Effects of Atmospheric Nitrogen Deposition on the Herbaceous Layer of a Central Appalachian Hardwood Forest

Additions of nitrogen (N) have been shown to alter species diversity of plant communities, with most experimental studies having been carried out in communities dominated by herbaceous species. We examined seasonal and inter-annual patterns of change in the herbaceous layer of two watersheds of a central Appalachian hardwood forest that differed in experimental treatment. This study was carried out at the Fernow Experimental Forest, West Virginia, using two adjacent watersheds: WS4 (mature, second-growth hardwood stand, untreated reference), and WS3. Seven circular 0.04-ha sample plots were established in eachwatershed to represent its full range of elevation and slope aspect. The herbaceous layer was sampled by identifying and visually estimating cover (%) of all vascular plants. Sampling was carried out in mid-July of 1991 and repeated at approximately the same time in 1992. In 1994, these same plots were sampled each month fromMay to October. Seasonal patterns of herb layer dynamics were assessed for the complete 1994 data set, whereasinter-annual variability was based on plot data from 1991, 1992, and the July sample of 1994. There were nosignificant differences between watersheds for any sample year for any of the other herb layer characteristics measured, including herb layer cover, species richness, evenness, and diversity. Cover on WS4 decreased significantly from 1991 to 1992, followed by no change to 1994. By contrast, herb layer cover did not varysignificantly across years on WS3. Cover of the herbaceous layer of both watersheds increased from early in the growing season to the middle of the growing season, decreasing thereafter, with no significant differencesbetween WS3 and WS4 for any of the monthly cover means in 1994. Similar seasonal patterns found for herblayer cover—and lack of significant differences between watersheds—were also evident for species diversityand richness. By contrast, there was little seasonal change in herb layer species evenness, which was nearlyidentical between watersheds for all months except October. Seasonal patterns for individual species/speciesgroups were closely similar between watersheds, especially for Viola rotundifolia and Viola spp. Species richnessand species diversity were linearly related to herb layer cover for both WS3 and WS4, suggesting that spatialand temporal increases in cover were more related to recruitment of herb layer species than to growth of existingspecies. Results of this study indicate that there have been negligible responses of the herb layer to 6 yr of additions to WS3.

Effects of Silvicultural Practices on Soil Carbon and Nitrogen in a Nitrogen Saturated Central Appalachian (USA) Hardwood Forest Ecosystem

Silvicultural treatments represent disturbances to forest ecosystems often resulting in transient increases in net nitrification and leaching of nitrate and base cations from the soil. Response of soil carbon (C) is more complex, decreasing from enhanced soil respiration and increasing from enhanced postharvest inputs of detritus. Because nitrogen (N) saturation can have similar effects on cation mobility, timber harvesting in N-saturated forests may contribute to a decline in both soil C and base cation fertility, decreasing tree growth. Although studies have addressed effects of either forest harvesting or N saturation separately, few data exist on their combined effects. Our study examined the responses of soil C and N to several commercially used silvicultural treatments within the Fernow Experimental Forest, West Virginia, USA, a site with N-saturated soils. Soil analyses included soil organic matter (SOM), C, N, C/N ratios, pH, and net nitrification. We hypothesized the following gradient of disturbance intensity among silvicultural practices (from most to least intense): even-age with intensive harvesting (EA-I), even-age with extensive harvesting, even-age with commercial harvesting, diameter limit, and single-tree harvesting (ST). We anticipated that effects on soil C and N would be greatest for EA-I and least with ST. Tree species exhibited a response to the gradient of disturbance intensity, with early successional species more predominant in high-intensity treatments and late successional species more predominant in low-intensity treatments. Results for soil variables, however, generally did not support our predictions, with few significant differences among treatments and between treatments and their paired controls for any of the measured soil variables. Multiple regression indicated that the best predictors for net nitrification among samples were SOM (positive relationship) and pH (negative relationship). This finding confirms the challenge of sustainable management of N-saturated forests.

Invasion patterns of emerald ash borer and European earthworms in forested ecosystems

Invasive and exotic species present a serious threat to the health and sustainability of natural ecosystems. These species often benefit from anthropogenic activities that aid their introduction and dispersal. This dissertation focuses on invasion dynamics of the emerald ash borer, native to Asia, and European earthworms. These species have shown detrimental impacts in invaded forest ecosystems across the Great Lakes region, and continue to spread via human-assisted long distance dispersal and by natural modes of dispersal into interior forests from areas of introduction. Successful forest management requires that the impact and effect of invasive species be considered and incorporated into management plans. Understanding patterns and constraints of introduction, establishment, and spread will aid in this effort. To assist in efforts to locate introduction points of emerald ash borer, a multicriteria risk model was developed to predict the highest risk areas. Important parameters in the model were road proximity, land cover type, and campground proximity. The model correctly predicted 85% of known emerald ash borer invasion sites to be at high risk. The model’s predictions across northern Michigan can be used to focus and guide future monitoring efforts. Similar modeling efforts were applied to the prediction of European earthworm invasion in northern Michigan forests. Field sampling provided a means to improve upon modeling efforts for earthworms to create current and future predictions of earthworm invasion. Those sites with high soil pH and high basal area of earthworm preferred overstory species (such as basswood and maples) had the highest likelihood of European earthworm invasion. Expanding beyond Michigan into the Upper Great Lakes region, earthworm populations were sampled across six National Wildlife Refuges to identify potential correlates and deduce specific drivers and constraints of earthworm invasion. Earthworm communities across all refuges were influenced by patterns of anthropogenic activity both within refuges and in surrounding ecoregions of study. Forest composition, soil pH, soil organic matter, anthropogenic cover, and agriculture proximity also proved to be important drivers of earthworm abundance and community composition. While there are few management options to remove either emerald ash borer or European earthworms from forests after they have become well established, prevention and early detection are important and can be beneficial. An improved understanding the factors controlling the distribution and invasion patterns of exotic species across the landscape will aid efforts to determine their consequences and generate appropriate forest management solutions to sustain ecosystem health in the presence of these invaders.

Evaluation of sugar maple dieback in the Upper Great Lakes region and development of a forest health youth education program

Acer saccharum Marsh., is one of the most valuable trees in the northern hardwood forests. Severe dieback was recently reported by area foresters in the western Upper Great Lakes Region. Sugar Maple has had a history of dieback over the last 100 years throughout its range and different variables have been identified as being the predisposing and inciting factors in different regions at different times. Some of the most common factors attributed to previous maple dieback episodes were insect defoliation outbreaks, inadequate precipitation, poor soils, atmospheric deposition, fungal pathogens, poor management, or a combination of these. The current sugar maple dieback was evaluated to determine the etiology, severity, and change in dieback on both industry and public lands. A network of 120 sugar maple health evaluation plots was established in the Upper Peninsula, Michigan, northern Wisconsin, and eastern Minnesota and evaluated annually from 2009-2012. Mean sugar maple crown dieback between 2009-2012 was 12.4% (ranging from 0.8-75.5%) across the region. Overall, during the sampling period, mean dieback decreased by 5% but individual plots and trees continued to decline. Relationships were examined between sugar maple dieback and growth, habitat conditions, ownership, climate, soil, foliage nutrients, and the maple pathogen sapstreak. The only statistically significant factor was found to be a high level of forest floor impacts due to exotic earthworm activity. Sugar maple on soils with lower pH had less earthworm impacts, less dieback, and higher growth rates than those on soils more favorable to earthworms. Nutritional status of foliage and soil was correlated with dieback and growth suggesting perturbation of nutrient cycling may be predisposing or contributing to dieback. The previous winter's snowfall totals, length of stay on the ground, and number of days with freezing temperatures had a significant positive relationship to sugar maple growth rates. Sapstreak disease, Ceratocystis virescens, may be contributing to dieback in some stands but was not related to the amount of dieback in the region. The ultimate goal of this research is to help forest managers in the Great Lakes Region prevent, anticipate, reduce, and/or salvage stands with dieback and loss in the future. An improved understanding of the complex etiology associated with sugar maple dieback in the Upper Great Lakes Region is necessary to make appropriate silvicultural decisions. Forest Health education helps increase awareness and proactive forest management in the face of changing forest ecosystems. Lessons are included to assist educators in incorporating forest health into standard biological disciplines at the secondary school curricula.

Effects of forest management on the diversity of deadwood-inhabiting fungi in Central European forests

Land use and land use change affect deadwood amount, quality and associated biodiversity in forest ecosystems. Old growth or virgin forests, which are exceptionally rare in temperate Europe harbor more deadwood and associated fungal species than managed forests. Whether and how more recent abandonment of management, to reestablish more natural forests, affects deadwood amount and fungal diversity on deadwood is unknown. Our main aim was to compare deadwood amount, characteristics and deadwood inhabiting fungi in differently managed forest types typical for large areas of Central Europe. We sampled deadwood inhabiting fungi on 27 forest plots of 400 m2 each in three geographically distant regions in Germany. Three forest management types, namely managed coniferous, managed deciduous and unmanaged deciduous forests, were represented by nine plots each. In autumn 2008 we collected all fungal fruiting bodies on deadwood >7 cm of diameter. We found deadwood amounts and fungal species numbers in unmanaged forests to be lower than in managed forests, which we attributed to the lack of natural tree death during the short time since management abandonment of usually 10–30 years. However, rarefaction analysis among deadwood items in forest plots indicated a slightly higher species density in unmanaged forests, which may be the first signal of a positive effect on fungal species richness on deadwood after management was abandoned. Although the three study regions span a large geographical gradient, we did not detect differences in the fungal species composition or in deadwood amounts and patterns, which reflects the wide distribution of this group of organisms and points to consistent management procedures among study regions. A very clear composition difference however occurred between deciduous and coniferous wood showing species substrate specialization. We conclude that the amount of deadwood is the main driver of deadwood fungal species richness, and substrate diversity in terms of various decay degrees, deadwood tree species and deadwood size are also important. Thus, to promote species richness of deadwood fungi it is vital to enhance deadwood amounts and diversity

Designing forest biodiversity experiments: general considerations illustrated by a new large experiment in subtropical China

1. Biodiversity-ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25.8x25.8m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial ecoscape' to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achieving specific research goals and synergy with previous experiments involves trade-offs between different designs and requires manifold design decisions.

Central European hardwood trees in a high-CO 2 future: synthesis of an 8-year forest canopy CO 2 enrichment project

Rapidly increasing atmospheric CO2 is not only changing the climate system but may also affect the biosphere directly through stimulation of plant growth and ecosystem carbon and nutrient cycling. Although forest ecosystems play a critical role in the global carbon cycle, experimental information on forest responses to rising CO2 is scarce, due to the sheer size of trees. Here, we present a synthesis of the only study world-wide where a diverse set of mature broadleaved trees growing in a natural forest has been exposed to future atmospheric CO2 levels (c. 550ppm) by free-air CO2 enrichment (FACE). We show that litter production, leaf traits and radial growth across the studied hardwood species remained unaffected by elevated CO2 over 8years. CO2 enrichment reduced tree water consumption resulting in detectable soil moisture savings. Soil air CO2 and dissolved inorganic carbon both increased suggesting enhanced below-ground activity. Carbon release to the rhizosphere and/or higher soil moisture primed nitrification and nitrate leaching under elevated CO2; however, the export of dissolved organic carbon remained unaltered.Synthesis. Our findings provide no evidence for carbon-limitation in five central European hardwood trees at current ambient CO2 concentrations. The results of this long-term study challenge the idea of a universal CO2 fertilization effect on forests, as commonly assumed in climate-carbon cycle models.

Assessing traditional knowledge on forest uses to understand forest ecosystem dynamics

Forest ecosystems worldwide are heavily influenced by human activities. Information on these human activities are key for understanding ecosystem dynamics, especially as some of these human activities have long-term consequences, i.e. legacy effects. We assessed the diversity of forest uses across the Swiss Alps and the respective traditional forest-related knowledge (TFRK) by conducting 56 oral history interviews in five regions. As TFRK tends to be underrepresented in written records, oral history proves to be a very valuable approach. We classified the information gathered in 61 specific forest use practices, characterized by activity, product and use, in order to gain a comprehensive picture of the regional variability but also cross-regional ubiquity of certain forest uses. Based on these results we are able to name the ecologically most relevant forest uses which were most likely, or potentially occurring in Central European forests. We suggest a simple, systematic historical assessment of these relevant forest uses to be applied in all studies on forest ecosystems. This procedure will enable scientists to better evaluate to what degree human activities had an impact on forest ecosystem dynamics in their study area.

Stand scale variability of topsoil organic matter composition in a high-elevation Norway spruce forest ecosystem

Our knowledge about the effect of single-tree influence areas on the physicochemical properties of the underlying mineral soil in forest ecosystems is still limited. This restricts our ability to adequately estimate future changes in soil functioning due to forest management practices. We studied the stand scale spatial variation of different soil organic matter species investigated by 13C NMR spectroscopy, lignin phenol and neutral sugar analysis under an unmanaged mountainous high-elevation Norway spruce (Picea abies L.) forest in central Europe. Multivariate geostatistical approaches were applied to relate the spatial patterns of the different soil organic matter species to topographic parameters, bulk density, oxalate- and dithionite-extractable iron, pH, and the impact of tree distribution. Soil samples were taken from the mineral top soil. Generally, the stand scale distribution patterns of different soil organic matter compounds could be divided into two groups: Those compounds, which were significantly spatially correlated with topography/altitude and those with small scale spatial pattern (range ≤ 10 m) that was closely related to tree distribution. The concentration of plant-derived soil organic matter components, such as lignin, at a given sampling point was significantly spatially related to the distance of the nearest tree (p ≤ 0.05). In contrast, the spatial distribution of mainly microbial-derived compounds (e.g. galactose and mannose) could be attributed to the dominating impact of small-scale topography and the contribution of poorly crystalline iron oxides that were significantly larger in the central depression of the study site compared to crest and slope positions. Our results demonstrate that topographic parameters dominate the distribution of overall topsoil organic carbon (OC) stocks at temperate high-elevation forest ecosystems, particularly in sloped terrain. However, trees superimpose topography-controlled OC biogeochemistry beneath their crown by releasing litter and changing soil conditions in comparison to open areas. This may lead to distinct zones with different mechanisms of soil organic matter degradation and also stabilization in forest stands.

Flujos de gases de efecto invernadero en suelos forestales y agroforestales del centro de la Península Ibérica=Greenhouse gas flows in forest and agroforestry soils in the center of the Iberian Peninsula

Debido a la complejidad de los procesos que controlan el intercambio de gases de carbono (C) y nitrógeno (N) entre el suelo y la atmósfera, en los sistemas forestales y agroforestales, son comprensibles las incógnitas existentes respecto a la estimación de los flujos de los gases de efecto invernadero (GEI) y la capacidad como reservorios de carbono de los suelos, bajo diferentes formas de uso y regímenes de alteración a escala regional y global. Esta escasez de información justifica la necesidad de caracterizar la dinámica de intercambio de GEI en los ecosistemas Mediterráneos, en especial en el contexto actual de cambio climático, y el incremento asociado de temperatura y periodos de sequía, alteración de los patrones de precipitación, y el riesgo de incendios forestales; cuyas consecuencias afectarán tanto a los compartimentos de C y de N del suelo como a la capacidad de secuestro de C de estos ecosistemas. Dentro de este contexto se enmarca la presente tesis doctoral cuyo objetivo ha sido cuantificar y caracterizar los flujos de dióxido de carbono (CO2), de oxido nitroso (N2O) y de metano (CH4), junto con los stocks de C y N, en suelos forestales de Quercus ilex, Quercus pyrenaica y Pinus sylvestris afectados por incendios forestales; así como el estudiar el efecto de la gestión y la cubierta arbórea en la respiración del suelo y los stocks de C y N en una dehesa situada en el centro de la Península Ibérica. De manera que los flujos de CO2, N2O y CH4; y los parámetros físico-químicos y biológicos del suelo fueron estudiados en los diferentes tratamientos y ecosistemas a lo largo del trabajo que se presenta. Los resultados obtenidos muestran la existencia de variaciones temporales y espaciales de la respiración del suelo dentro de una escala geográfica pequeña, controladas principalmente por la temperatura y la humedad del suelo; y por los contenidos de C y N del suelo en un bosque de Pinus sylvestris en la vertiente norte de la Sierra de Guadarrama , en España. El análisis de los efectos de los incendios forestales a largo plazo (6-8 años) revela que las pérdidas anuales de C a través de la respiración del suelo en las zonas quemadas de Quercus ilex, Quercus pyrenaica y Pinus sylvestris fueron 450 gCm-2yr-1, 790 gCm-2yr-1 y 1220 gCm-2yr-1, respectivamente; lo que representa una reducción del 43%, 22% y 11% en comparación con las zonas no quemadas de dichas especies, debido a la destrucción de la masa arbórea. El efecto del fuego también alteró los flujos N2O y CH4 del suelo, de una forma diferente en los distintos ecosistemas y estacionalidades estudiadas. De tal modo, que los suelos quemados mostraron una mayor oxidación del CH4 en las masas de Q. ilex, y una menor oxidación en las de P. sylvestris; además de una disminución de los flujos de N2O en Q. pyrenaica. Los incendios también afectaron los parámetros microclimáticos de los suelos forestales, observándose un incremento de la temperatura del suelo y una disminución de la humedad en los emplazamientos quemados que en los no quemados. Los cationes intercambiables, el pH, el cociente C/N, el contenido en raicillas y la biomasa microbiana también disminuyeron en las zonas quemadas. Aunque el C orgánico del suelo no se alteró de manera significativa, si lo hizo la calidad de la materia orgánica, disminuyendo el carbono lábil y aumentando las formas recalcitrantes lo que se tradujo en menor sensibilidad de la respiración del suelo a la temperatura (valores de Q10) en las zonas quemadas. Los resultados del estudio realizado en la Dehesa muestran que las actividades silvopastorales estudiadas afectaron levemente y de forma no constante a la respiración del suelo y las condiciones microclimáticas del suelo. Se observó una reducción 12% de la respiración del suelo por efecto del pastoreo no intensivo. Sin embargo, se observaron incrementos de 3Mg/ha en los stocks de C y de 0.3 Mg/ha en los stocks de N en los suelos pastoreados en comparación con los no pastoreados. Aunque, no se observó un claro efecto de la labranza sobre la respiración del suelo en nuestro experimento, sin embargo si se observó una disminución de 3.5 Mg/ha en las reservas de C y de 0.3 Mg/ ha en las de N en los suelos labrados comparados con los no labrados. La copa del arbolado influyó de forma positiva tanto en la respiración del suelo, como en los stocks de C y N de los suelos. La humedad del suelo jugó un papel relevante en la sensibilidad de la respiración a la temperatura del suelo. Nuestros resultados ponen de manifiesto la sensibilidad de la respiración del suelo a cambios en la humedad y los parámetros edáficos, y sugieren que la aplicación de modelos estándar para estimar la respiración del suelo en áreas geográficas pequeñas puede no ser adecuada a menos que otros factores sean considerados en combinación con la temperatura del suelo. Además, las diferentes respuestas de los flujos de gases de efecto invernadero a los cambios, años después de la ocurrencia de incendios forestales, destaca la necesidad de incluir estos cambios en las futuras investigaciones de la dinámica del carbono en los ecosistemas mediterráneos. Por otra parte, las respuestas divergentes en los valores de respiración del suelo y en los contenidos de C y N del suelo observados en la dehesa, además de la contribución de la copa de los árboles en los nutrientes del suelo ilustran la importancia de mantener la gestión tradicional aplicada en beneficio de la capacidad de almacenar C en la dehesa estudiada. La información obtenida en este trabajo pretende contribuir a la mejora del conocimiento de la dinámica y el balance de C en los sistemas mediterráneos, además de ayudar a predecir el impacto del cambio climático en el intercambio de C entre los ecosistemas forestales y agroforestales y la atmósfera. ABSTRACT Due to the complexity of the processes that control the exchange of carbon (C) and nitrogen (N) gasses between soils and the atmosphere in forest and agroforestry ecosystems, understandable uncertainties exist as regards the estimation of greenhouse gas (GHG) fluxes and the soil sink capacity at regional and global scale under different forms of land use and disturbance regimes. These uncertainties justify the need to characterize the exchange dynamics of GHG between the atmosphere and soils in Mediterranean terrestrial ecosystems, particularly in the current context of climate change and the associated increase in temperature, drought periods, heavy rainfall events, and increased risk of wildfires, which affect not only the C and N pools but also the soil C sink capacity of these ecosystems. Within this context, the aims of the present thesis were, firstly, to quantify and characterize the fluxes of carbon dioxide (CO2), nitrous oxide (N2O) and methane (CH4) as well as the C and N stocks in Quercus ilex, Quercus pyrenaica and Pinus sylvestris stands affected by wildfires, and secondly, to study the effects of Quercus ilex canopy and management on both soil respiration and C and N pools in dehesa systems in the center of Iberian Peninsula. Soil CO2, N2O and CH4 fluxes, and soil physical-chemical and biological parameters were studied under the different treatments and ecosystems considered in this study. The results showed seasonal and spatial variations in soil respiration within small geographic areas, mainly controlled by soil temperature and moisture in addition to soil carbon and nitrogen stocks in mixed pine–oak forest ecosystems on the north facing slopes of the Sierra de Guadarrama in Spain. The analysis of long term effects of wildfires (6–8 years) revealed that annual carbon losses through soil respiration from burned sites in Quercus ilex, Quercus pyrenaica and Pinus sylvestris stands were 450 gCm-2yr-1, 790 gCm-2yr-1 and 1220 gCm-2yr-1, respectively; with burned sites emitting 43%, 22% and 11% less in burned as opposed to non-burned sites due the loss of trees. Fire may alter both N2O and CH4 fluxes although the magnitude of such variation depends on the site, soil characteristics and seasonal climatic conditions. The burned sites showed higher CH4 oxidation in Q.ilex stands, and lower oxidation rates in P. sylvestris stands. A reduction in N2O fluxes in Q. pyrenaica stands was detected at burned sites along with changes in soil microclimate; higher soil temperature and lower soil moisture content. Exchangeable cations, the C/N ratio, pH, fine root and microbial biomass were also found to decrease at burned sites. Although the soil organic carbon was not significantly altered, the quality of the organic matter changed, displaying a decrease in labile carbon and a relative increase in refractory forms, leading to lower sensitivity of soil respiration to temperature (Q10 values) at burned sites. The results from the dehesa study show that light grazing and superficial tilling practices used in the studied dehesa system in Spain had a slight but non-consistent impact on soil respiration and soil microclimate over the study period. The reduction in soil respiration in the dehesa system due to the effects of grazing was around 12 %. However, increments of 3Mg/ha in C stocks and 0.3 Mg/ha in N stocks in grazed soils were observed. Although no clear effect of tilling on soil respiration was found, a decrease of 3.5 Mg/ha in C stocks and 0.3 Mg/ha in N stocks was detected for tilled soils. The presence of a tree canopy induced increases in soil respiration, soil C and N stocks, while soil moisture was found to play an important role in soil respiration temperature response. Our results suggest that the use of standard models to estimate soil respiration in small geographical areas may not be adequate unless other factors are considered in addition to soil temperature. Furthermore, the different responses of GHG flux to climatic shifts, many years after the occurrence of wildfire, highlight the need to include these shifts in C dynamics in future research undertaken in Mediterranean ecosystems. Furthermore, divergent responses in soil respiration and soil C and N stocks to grazing or tilling practices in Dehesa systems, and the influence of tree canopy on soil respiration and soil nutrient content, illustrate the importance of maintaining beneficial management practices. Moreover, the carbon sequestration capacity of the Dehesa system studied may be enhanced through improvements in the management applied. It is hoped that the information obtained through this research will contribute towards improving our understanding of the dynamics and balance of C in Mediterranean systems, and help predict the impact of climate change on the exchange of C between forest and agroforestry ecosystems and the atmosphere.

Assessing forest response-type diversity to natural disturbances in Peninsular Spain and the Balearic Islands = Evaluación de la diversidad de tipos de respuesta a perturbaciones naturales en la España peninsular e Islas Baleares

Disturbances shape forest ecosystems by influencing their composition, structure, and processes. In the Mediterranean Basin, changes in the disturbance regimes have been predicted to occur in the next future with a higher occurrence of extreme events of drought, wildfire, and – to a lesser extent – windstorm. Woody species are the main elements defining the structure and functioning of forest ecosystems. Recently, response-type diversity has been pointed out as an appropriate indicator of ecosystems resilience. For this, we have elaborated a complete response-trait database for the tree and shrubby species considered in the Third Spanish National Forest Inventory (3SNFI). In the database, the presence or absence of nine response traits associated to drought, fire, and wind were assigned to each species. The database reflected the lack of information about some important traits (in particular for shrubby species) and allowed to determine those traits most widely distributed. The information contained in the database was then used to assess a relative index of forest resilience to these disturbances calculated from the abundance of response traits and the species redundancy for each plot of the 3SNFI; considering both tree and shrubby species. In general, few plots showed high values of the resilience index, probably because some traits were scarcely presented in the species and also because most plots presented very few species. The cartographic representation of the index showed low values for the stands located in mountainous ranges, which are mostly composed by species typical from central Europe. In the other side, Eucalyptus plantations in Galicia appeared as one thee the most resilient ecosystems, due to its higher adaptive capacity to persist after the occurrence of drought, fire, and windstorm events. We conclude that the response traits database can constitute a useful tool for forest management and planning and for future research to enhance the forest resilience.

Long-term Responses of Mountain Ecosystems to Environmental Changes: Resilience, Adjustment, and Vulnerability

The steep environmental gradients of mountain ecosystems over short distances reflect large gradients of several climatic parameters and hence provide excellent possibilities for ecological research on the effects of environmental change. To gain a better understanding of the dynamics of abiotic and biotic parameters of mountain ecosystems, long-term records are required since permanent plots in mountain regions cover in the best case about 50 - 70 years. In order to extend investigations of ecological dynamics beyond these temporal limitations of permanent plots, paleoecological approaches can be used if the sampling resolution can be adapted to ecological research questions, e.g. a sample every 10 years. Paleoecological studies in mountain ecosystems can provide new ecological insights through the combination of different spatial and temporal scales. [f we thus improve our understanding of processes across both steep environmental gradients and different time scales, we may be able to better estimate ecosystem responses to current and future environmental change (Ammann et al. 1993; Lotter et al. 1997). The complexity of ecological interactions in mountain regions forces us to concentrate on a number of sub-systems - without losing sight of the wider context. Here, we summarize a few case studies on the effects of Holocene climate change and disturbance on the vegetation of the Western Alps. To categorize the main response modes of vegetation to climatic change and disturbance in the Alps we use three classes of ecological behaviour: "resilience", "adjustment", and "vulnerability", We assume a resilient (or elastic) behaviour if vegetation is able to recover to its former state, regaining important ecosystem characteristics, such as floristic composition, biodiversity, species abundances, and biomass (e.g. Küttel 1990; Aber and Melillo 199 1). Conversely, vegetation displacements may occur in response to climatic change and/or disturbance. In some cases, this may culminate in irreversible large-scale processes such as species and/or community extinctions. Such drastic developments indicate high ecosystem vulnerability (or inelasticity or instability, for detailed definitions see Küttel 1990; Aber and Melillo 199 1) to climatic change and/or disturbance. In this sense, the "vulnerability" (or instability) of an ecosystem is expressed by the degree of failure to recover to the original state before disturbance and/or climatic change. Between these two extremes (resilience vs. vulnerability), ecosystem adjustments to climatic change and/or disturbance may occur, including the appearance of new and/or the disappearance of old species. The term "adjustment" is hence used to indicate the response of vegetational communities, which adapted to new environmental conditions without losing their main character. For forest ecosystems, we assume vegetational adjustments (rather than vulnerability) if the dominant (or co-dominant) tree species are not outnumbered or replaced by formerly unimportant plant species or new invaders. Adaptation as a genetic process is not discussed here and will require additional pbylogeographical studies (that incorporate the analysis of ancient DNA) in order to fully understand the distributions of ecotypes.

Lake-effect snow as the dominant control of mesic-forest distribution in Michigan, USA

1Recent studies demonstrated the sensitivity of northern forest ecosystems to changes in the amount and duration of snow cover at annual to decadal time scales. However, the consequences of snowfall variability remain uncertain for ecological variables operating at longer time scales, especially the distributions of forest communities. 2The Great Lakes region of North America offers a unique setting to examine the long-term effects of variable snowfall on forest communities. Lake-effect snow produces a three-fold gradient in annual snowfall over tens of kilometres, and dramatic edaphic variations occur among landform types resulting from Quaternary glaciations. We tested the hypothesis that these factors interact to control the distributions of mesic (dominated by Acer saccharum, Tsuga canadensis and Fagus grandifolia) and xeric forests (dominated by Pinus and Quercus spp.) in northern Lower Michigan. 3We compiled pre-European-settlement vegetation data and overlaid these data with records of climate, water balance and soil, onto Landtype Association polygons in a geographical information system. We then used multivariate adaptive regression splines to model the abundance of mesic vegetation in relation to environmental controls. 4Snowfall is the most predictive among five variables retained by our model, and it affects model performance 29% more than soil texture, the second most important variable. The abundance of mesic trees is high on fine-textured soils regardless of snowfall, but it increases with snowfall on coarse-textured substrates. Lake-effect snowfall also determines the species composition within mesic forests. The weighted importance of A. saccharum is significantly greater than of T. canadensis or F. grandifolia within the lake-effect snowbelt, whereas T. canadensis is more plentiful outside the snowbelt. These patterns are probably driven by the influence of snowfall on soil moisture, nutrient availability and fire return intervals. 5Our results imply that a key factor dictating the spatio-temporal patterns of forest communities in the vast region around the Great Lakes is how the lake-effect snowfall regime responds to global change. Snowfall reductions will probably cause a major decrease in the abundance of ecologically and economically important species, such as A. saccharum.

Anticipating climate-induced changes to forest cover in Ontario and the implications for future bioenergy development

Climate change is expected to have marked impacts on forest ecosystems. In Ontario forests, this includes changes in tree growth, stand composition and disturbance regimes, with expected impacts on many forest-dependent communities, the bioeconomy, and other environmental considerations. In response to climate change, renewable energy systems, such as forest bioenergy, are emerging as critical tools for carbon emissions reductions and climate change mitigation. However, these systems may also need to adapt to changing forest conditions. Therefore, the aim of this research was to estimate changes in forest growth and forest cover in response to anticipated climatic changes in the year 2100 in Ontario forests, to ultimately explore the sustainability of bioenergy in the future. Using the Haliburton Forest and Wildlife Reserve in Ontario as a case study, this research used a spatial climate analog approach to match modeled Haliburton temperature and precipitation (via Fourth Canadian Regional Climate Model) to regions currently exhibiting similar climate (climate analogs). From there, current forest cover and growth rates of core species in Haliburton were compared to forests plots in analog regions from the US Forest Service Forest Inventory and Analysis (FIA). This comparison used two different emission scenarios, corresponding to a high and a mid-range emission future. This research then explored how these changes in forests may influence bioenergy feasibility in the future. It examined possible volume availability and composition of bioenergy feedstock under future conditions. This research points to a potential decline of softwoods in the Haliburton region with a simultaneous expansion of pre-established hardwoods such as northern red oak and red maple, as well as a potential loss in sugar maple cover. From a bioenergy perspective, hardwood residues may be the most feasible feedstock in the future with minimal change in biomass availability for energy production; under these possible conditions, small scale combined heat and power (CHP) and residential pellet use may be the most viable and ecologically sustainable options. Ultimately, understanding the way in which forests may change is important in informing meaningful policy and management, allowing for improved forest bioenergy systems, now and in the future.

Assessing the influence of abiotic factors and leaf-level properties on the stability of growing-season canopy greenness in a deciduous forest

Maps depicting spatial pattern in the stability of summer greenness could advance understanding of how forest ecosystems will respond to global changes such as a longer growing season. Declining summer greenness, or “greendown”, is spectrally related to declining near-infrared reflectance and is observed in most remote sensing time series to begin shortly after peak greenness at the end of spring and extend until the beginning of leaf coloration in autumn,. Understanding spatial patterns in the strength of greendown has recently become possible with the advancement of Landsat phenology products, which show that greendown patterns vary at scales appropriate for linking these patterns to proposed environmental forcing factors. This study tested two non-mutually exclusive hypotheses for how leaf measurements and environmental factors correlate with greendown and decreasing NIR reflectance across sites. At the landscape scale, we used linear regression to test the effects of maximum greenness, elevation, slope, aspect, solar irradiance and canopy rugosity on greendown. Secondly, we used leaf chemical traits and reflectance observations to test the effect of nitrogen availability and intrinsic water use efficiency on leaf-level greendown, and landscape-level greendown measured from Landsat. The study was conducted using Quercus alba canopies across 21 sites of an eastern deciduous forest in North America between June and August 2014. Our linear model explained greendown variance with an R2=0.47 with maximum greenness as the greatest model effect. Subsequent models excluding one model effect revealed elevation and aspect were the two topographic factors that explained the greatest amount of greendown variance. Regression results also demonstrated important interactions between all three variables, with the greatest interaction showing that aspect had greater influence on greendown at sites with steeper slopes. Leaf-level reflectance was correlated with foliar δ13C (proxy for intrinsic water use efficiency), but foliar δ13C did not translate into correlations with landscape-level variation in greendown from Landsat. Therefore, we conclude that Landsat greendown is primarily indicative of landscape position, with a small effect of canopy structure, and no measureable effect of leaf reflectance. With this understanding of Landsat greendown we can better explain the effects of landscape factors on vegetation reflectance and perhaps on phenology, which would be very useful for studying phenology in the context of global climate change