The Ecological Significance of the Herbaceous Layer in Temperate Forest Ecosystems

Despite a growing awareness that the herbaceous layer serves a special role in maintaining the structure and function of forests, this stratum remainsan underappreciated aspect of forest ecosystems. In this article I review and synthesize information concerning the herb layer’s structure,composition, and dynamics to emphasize its role as an integral component of forest ecosystems. Because species diversity is highest in the herb layeramong all forest strata, forest biodiversity is largely a function of the herb-layer community. Competitive interactions within the herb layer candetermine the initial success of plants occupying higher strata, including the regeneration of dominant overstory tree species. Furthermore, the herblayer and the overstory can become linked through parallel responses to similar environmental gradients. These relationships between strata varyboth spatially and temporally. Because the herb layer responds sensitively to disturbance across broad spatial and temporal scales, its dynamics canprovide important information regarding the site characteristics of forests, including patterns of past land-use practices. Thus, the herb layer has asignificance that belies its diminutive stature.

Intra-annual radial growth and water relations of trees - implications towards a growth mechanism

There is a missing link between tree physiological and wood-anatomical knowledge which makes it impossible mechanistically to explain and predict the radial growth of individual trees from climate data. Empirical data of microclimatic factors, intra-annual growth rates, and tree-specific ratios between actual and potential transpiration (T PET−1) of trees of three species (Quercus pubescens, Pinus sylvestris, and Picea abies) at two dry sites in the central Wallis, Switzerland, were recorded from 2002 to 2004 at a 10 min resolution. This included the exceptionally hot and dry summer of 2003. These data were analysed in terms of direct (current conditions) and indirect impacts (predispositions of the past year) on growth. Rain was found to be the only factor which, to a large extent, consistently explained the radial increment for all three tree species at both sites and in the short term as well. Other factors had some explanatory power on the seasonal time-scale only. Quercus pubescens built up much of its tree ring before bud break. Pinus sylvestris and Picea abies started radial growth 1–2 weeks after Quercus pubescens and this was despite the fact that they had a high T PET−1 before budburst and radial growth started. A high T PET−1 was assumed to be related to open stomata, a very high net CO2 assimilation rate, and thus a potential carbon (C)-income for the tree. The main period of radial growth covered about 30–70% of the productive days of a year. In terms of C-allocation, these results mean that Quercus pubescens depended entirely on internal C-stores in the early phase of radial growth and that for all three species there was a long time period of C-assimilation which was not used for radial growth in above-ground wood. The results further suggest a strong dependence of radial growth on the current tree water relations and only secondarily on the C-balance. A concept is discussed which links radial growth over a feedback loop to actual tree water-relations and long-term affected C-storage to microclimate.

Effects of elevated atmospheric CO2 and O3 on wood density, antomical proerties and decomposition of Northern Hardwoods

Anthropogenic activities continue to drive atmospheric CO2 and O3 concentrations to levels higher than during the pre-industrial era. Accumulating evidence indicates that both elevated CO2 and elevated O3 could modify the quantity and biochemistry of woody plant biomass. Anatomical properties of woody plants are largely influenced by the activity of the cambium and the growth characteristics of wood cells, which are in turn influenced by a range of environmental factors. Hence, alterations in the concentrations of atmospheric CO2 and / or O3 could also impact wood anatomical properties. Many fungi derive their metabolic resources for growth from plant litter, including woody tissue, and therefore modifications in the quantity, biochemistry and anatomical properties of woody plants in response to elevated CO2 and / or O3 could impact the community of wood-decaying fungi and rates of wood decomposition. Consequently carbon and nutrient cycling and productivity of terrestrial ecosystem could also be impacted. Alterations in wood structure and biochemistry of woody plants could also impact wood density and subsequently impact wood quality. This dissertation examined the long term effects of elevated CO2 and / or O3 on wood anatomical properties, wood density, wood-decaying fungi and wood decomposition of northern hardwood tree species at the Aspen Free-Air CO2 and O3 Enrichment (Aspen FACE) project, near Rhinelander, WI, USA. Anatomical properties of wood varied significantly with species and aspen genotypes and radial position within the stem. Elevated CO2 did not have significant effects on wood anatomical properties in trembling aspen, paper birch or sugar maple, except for marginally increasing (P < 0.1) the number of vessels per square millimeter. Elevated O3 marginally or significantly altered vessel lumen diameter, cell wall area and vessel lumen area proportions depending on species and radial position. In line with the modifications in the anatomical properties, elevated CO2 and O3, alone, significantly modified wood density but effects were species and / or genotype specific. However, the effects of elevated CO2 and O3, alone, on wood anatomical properties and density were ameliorated when in combination. Wood species had a much greater impact on the wood-decaying fungal community and initial wood decomposition rate than did growth or decomposition of wood in elevated CO2 and / or O3. Polyporales, Agaricales, and Russulales were the dominant orders of fungi isolated. Based on the current results, future higher levels of CO2 and O3 may have moderate effects on wood quality of northern hardwoods, but for utilization purposes these may not be considered significant. However, wood-decaying fungal community composition and decomposition of northern hardwoods may be altered via shifts in species and / or genotype composition under future higher levels of CO2 and O3.

Patch selection by wintering white-tailed deer : consequences for eastern hemlock regeneration at local and landscape scales

Tsuga canadensis (eastern hemlock) is a highly shade-tolerant, late-successional, and long-lived conifer species found throughout eastern North America. It is most often found in pure or nearly pure stands, because highly acidic and nutrient poor forest floor conditions are thought to favor T. canadensis regeneration while simultaneously limiting the establishment of some hardwood species with greater nutrient requirements. Once a common species, T. canadensis is currently experiencing widescale declines across its range. The hemlock woolly adelgid (Adelges tsugae) is decimating the population across its eastern distribution. Across the Upper Great Lakes region, where the adelgid is currently being held at bay by cold winter temperatures, T. canadensis has been experiencing failures in regeneration attributed, in part, to herbivory by white-tailed deer (Odocoileus virginianus). Deer utilize T. canadensis stands as winter habitat in areas of high snow depth. Tsuga canadensis, once a major component of these forests, currently exists at just a fraction of its pre-settlement abundance due to historic logging and contemporary forest management practices, and what remains is found in small remnant patches surrounded by second- and third-growth deciduous forests. The deer population across the region, however, is likely double that of pre-European settlement times. In this dissertation I explore the relationship between white-tailed deer use of T. canadensis as winter habitat and the effect this use is having on regeneration and forest succession. For this research I quantified stand composition and structure and abiotic variables of elevation and snow depth in 39 randomly selected T. canadensis stands from across the western Upper Peninsula of Michigan. I also quantified composition and the configuration of the landscapes surrounding these stands. I measured relative deer use of T. canadensis stands as pellet group piles deposited in each stand during each of three consecutive winters, 2005-06, 2006-07, and 2007-08. The results of this research suggest that deer use of T. canadensis stands as winter habitat is influenced primarily by snow depth, elevation, and the composition and configuration of the greater landscapes surrounding these stands. Specifically, stands with more heterogeneous landscapes surrounding them (i.e., a patchy mosaic of conifer, deciduous, and open cover) had higher relative deer use than stands surrounded by homogenous deciduous forest cover. Additionally, the intensity of use and the number of stands used was greater in years with higher average snow depth. Tsuga canadensis regeneration in these stands was negatively associated with deer use and Acer saccharum (sugar maple) basal area. Of the 39 stands, 17 and 22 stands had no T. canadensis regeneration in small and large sapling categories, respectively. Acer saccharum was the most common understory tree species, and the importance of A. saccharum in the understory (stems < 10 cm dbh) of the stands was positively associated with overstory A. saccharum dominance. Tsuga canadensis establishment was associated with high-decay coarse woody debris and moss, and deciduous leaf litter inputs in these stands may be limiting access to these important microsites. Furthermore, A. saccharum is more tolerant to the effects of deer herbivory than T. canadensis, giving A. saccharum a competitive advantage in stands being utilized as winter habitat by deer. My research suggests that limited microsite availability, in conjunction with deer herbivory, may be leading to an erosion in T. canadensis patch stability and an altered successional trajectory toward one of A. saccharum dominance, an alternately stable climax species.

Retention of canopy trees as biological legacies for balancing woody biomass production and biodiversity in managed aspen forests of the Great Lakes Region

Green-tree retention under the conceptual framework of ecological forestry has the potential to provide both biomass feedstock for industry and maintain quality wildlife habitat. I examined the effects of retained canopy trees as biological legacies (“legacy trees”) in aspen (Populus spp.) forests on above-ground live woody biomass, understory plant floristic quality, and bird diversity. Additionally, I evaluated habitat quality for a high conservation priority species, the Golden-winged Warbler (Vermivora chrysoptera). I selected 27 aspen-dominated forest stands in northern Wisconsin with nine stands in each of three legacy tree retention treatments (conifer retention, hardwood retention, and clearcuts or no retention) across a chronosequence (4-36 years post-harvest). Conifer retention stands had greater legacy tree and all tree species biomass but lower regenerating tree biomass than clearcuts. Coniferous but not hardwood legacy trees appeared to suppress regenerating tree biomass. I evaluated the floristic quality of the understory plant assemblage by estimating the mean coefficient of conservatism (C). Mean C was lower in young stands than in middle-age or old stands; there was a marginally significant (p=0.058) interaction effect between legacy tree retention treatment and stand age. Late-seral plant species were positively associated with stand age and legacy tree diameter or age revealing an important relationship between legacy tree retention and stand development. Bird species richness was greatest in stands with hardwood retention particularly early in stand development. Six conservation priority bird species were indicators of legacy tree retention or clearcuts. Retention of legacy trees in aspen stands provided higher quality nest habitat for the Golden-winged Warbler than clearcuts based on high pairing success and nesting activity. Retention of hardwoods, particularly northern red oak (Quercus rubra), yielded the most consistent positive effects in this study with the highest bird species richness and the highest quality habitat for the Golden-winged Warbler. This treatment maintained stand biomass comparable to clearcuts and did not suppress regenerating tree biomass. In conclusion, legacy tree retention can enhance even-aged management techniques to produce a win-win scenario for the conservation of declining bird species and late-seral understory plants and for production of woody biomass feedstock from naturally regenerating aspen forests.

Assessing success of forest restoration efforts in degraded montane cloud forests in southern Mexico

Montane cloud forests are home to great biodiversity. However, non-sustainable anthropogenic activities have led to the loss of forest cover in southern Mexico. Increasing conservation, restoration and sustainable use of forest resources prevents the loss of cloud forests. In this study, success of forest restoration was evaluated in a degraded forest of Highlands Chiapas. The goal of this study was to assess the structure and composition of native tree species. We evaluated vegetation composition at three sites that had undergone enrichment plantings. Floristic composition and structure of the herbaceous, seedling, sapling, and overstory layers were measured. A total of sixty-six native tree species were recorded. Enrichment planting was found to have increased tree diversity. Moreover, 54% of the planted species were found in the understory, indicating that they were successfully recruiting. In conclusion, enrichment planting can aid in the conservation of forest cover in degraded areas.

Small-scale timber stand management techniques : a case study of woodlots in Isangati, Tanzania

Small-scale village woodlots of less than 0.5ha are the preferred use of land for local farmers with extra land in the village of Isangati, a small community located in the southern highlands of Tanzania. Farmers view woodlots as lucrative investments that do not involve intensive labor or time. The climate is ideal for the types of trees grown and the risks are minimal with no serious threats from insects, fires, thieves, or grazing livestock. It was hypothesized that small-scale village woodlot owners were not maximizing timber outputs with their current timber stand management and harvesting techniques. Personal interviews were conducted over a five month period and field data was collected at each farmer’s woodlots over a seven month period. Woodlot field data included woodlot size, number of trees, tree species, tree height, dbh, age, and spacing. The results indicated that the lack of proper woodlot management techniques results in failure to fully capitalize on the investment of woodlots. While farmers should continue with their current harvesting rotations, some of the reasons for not maximizing tree growth include close spacing (2m x 2m), no tree thinning, extreme pruning (60% of tree), and little to no weeding. Through education and hands-on woodlot management workshops, the farmers could increase their timber output and value of woodlots.

Critical evaluation of diameter increment modelling in the Great Lakes region

Simulations of forest stand dynamics in a modelling framework including Forest Vegetation Simulator (FVS) are diameter driven, thus the diameter or basal area increment model needs a special attention. This dissertation critically evaluates diameter or basal area increment models and modelling approaches in the context of the Great Lakes region of the United States and Canada. A set of related studies are presented that critically evaluate the sub-model for change in individual tree basal diameter used in the Forest Vegetation Simulator (FVS), a dominant forestry model in the Great Lakes region. Various historical implementations of the STEMS (Stand and Tree Evaluation and Modeling System) family of diameter increment models, including the current public release of the Lake States variant of FVS (LS-FVS), were tested for the 30 most common tree species using data from the Michigan Forest Inventory and Analysis (FIA) program. The results showed that current public release of the LS-FVS diameter increment model over-predicts 10-year diameter increment by 17% on average. Also the study affirms that a simple adjustment factor as a function of a single predictor, dbh (diameter at breast height) used in the past versions, provides an inadequate correction of model prediction bias. In order to re-engineer the basal diameter increment model, the historical, conceptual and philosophical differences among the individual tree increment model families and their modelling approaches were analyzed and discussed. Two underlying conceptual approaches toward diameter or basal area increment modelling have been often used: the potential-modifier (POTMOD) and composite (COMP) approaches, which are exemplified by the STEMS/TWIGS and Prognosis models, respectively. It is argued that both approaches essentially use a similar base function and neither is conceptually different from a biological perspective, even though they look different in their model forms. No matter what modelling approach is used, the base function is the foundation of an increment model. Two base functions – gamma and Box-Lucas – were identified as candidate base functions for forestry applications. The results of a comparative analysis of empirical fits showed that quality of fit is essentially similar, and both are sufficiently detailed and flexible for forestry applications. The choice of either base function in order to model diameter or basal area increment is dependent upon personal preference; however, the gamma base function may be preferred over the Box-Lucas, as it fits the periodic increment data in both a linear and nonlinear composite model form. Finally, the utility of site index as a predictor variable has been criticized, as it has been widely used in models for complex, mixed species forest stands though not well suited for this purpose. An alternative to site index in an increment model was explored, using site index and a combination of climate variables and Forest Ecosystem Classification (FEC) ecosites and data from the Province of Ontario, Canada. The results showed that a combination of climate and FEC ecosites variables can replace site index in the diameter increment model.

Early season ozone uptake is important for determining ozone tolerance in two trembling aspen clones

There is substantial genetic variability in response to ozone amongst and within tree species. Aspen is a highly variable species with a wide range of responses to ozone. Aspen response to elevated O3 levels is being investigated at the Aspen FACE site near Rhinelander, WI where five aspen clones of varying O3 tolerance have been fumigated with elevated O3 over the past decade. In this study, we examined the physiological differences in two of the aspen clones that differed significantly in their O3 tolerance with 8L being tolerant and 42E being sensitive. Throughout the 2007 and 2008 growing seasons we periodically estimated instantaneous photosynthetic rates, ACi responses and light response curves. The results of our study suggest that aspen clone 8L’s tolerance is due in part to decreased stomatal conductance early in the season, which lowered ozone uptake. Later during the season O3 uptake was comparable for the two clones. Our results also suggest the response of Vcmax, TPU, Rd, Gm, light compensation point and quantum flux to elevated O3 did not differ significantly between the two clones. Ozone uptake is important for ozone tolerance in clone 8L early in the season but cannot explain late season tolerance. Photosynthetic parameters for the two clones were similar, so clone 8L’s ozone tolerance is not due to a more efficient photosynthetic system.

INSTANTANEOUS PHOTOSYNTHETIC RESPONSE TO TEMPERATURE OF MATURE FOREST CANOPIES AND EXPERIMENTALLY WARMED SEEDLINGS

Tropical trees have been shown to be more susceptible to warming compared to temperate species, and have shown growth and photosynthetic declines at elevated temperatures as little as 3oC above ambient. However, regional and global vegetation models lack the data needed to accurately represent physiological response to increased temperatures in tropical forests. We compared the instantaneous photosynthetic responses to elevated temperatures of four mature tropical rainforest tree species in Puerto Rico and the temperate broadleaf species sugar maple (Acer saccharum) in Michigan. Contrary to expectations, leaves in the upper canopy of both temperate and tropical forests had temperature optima that are already exceeded by mean daily leaf temperatures. This indicates that tropical and temperate forests are already seeing photosynthesis decline at mid-day temperature. This decline may worsen as air temperatures rise with climate change if trees are unable to acclimate, increasing the likelihood that forests may shift from carbon sinks to sources. A secondary study was conducted on experimentally warmed sugar maple seedlings to determine if photosynthesis had been able to acclimate to +5oC air temperature over four years. Species abundance models had predicted a decline of sugar maple within the Upper Peninsula of Michigan over the next 100 years, due to elevated temperature and altered precipitation. Instantaneous photosynthetic temperature response curves on both control and heated seedlings showed that the differences between treatments were not statistically significant, though there was a 16% increase in temperature optima and a 3% increase in maximum rates of photosynthesis in warmed plots. Though evidence of acclimation was not significant, the seedlings did not fare poorly as the models suggest.

Changes in biodiversity and vegetation composition in the central Swiss Alps during the transition from pristine forest to first farming

Aim: We investigate the response of vegetation composition and plant diversity to increasing land clearance, burning and agriculture at the Mesolithic–Neolithic transition (c. 6400–5000 bc) when first farming was introduced. Location: The Valais, a dry alpine valley in Switzerland. Methods: We combine high-resolution pollen, microscopic charcoal and sedimentological data to reconstruct past vegetation, fire and land use. Pollen evenness, rarefaction-based and accumulation-based palynological richness analyses were used to reconstruct past trends in plant diversity. Results: Our results show that from c. 5500 cal. yr bc, slash-and-burn activities created a more open landscape for agriculture, at the expense of Pinus and Betula forests. Land clearance by slash-and-burn promoted diverse grassland ecosystems, while on the long term it reduced woodland and forest diversity, affecting important tree species such as Ulmus and Tilia. Main conclusions: Understanding the resilience of Alpine ecosystems to past disturbance variability is relevant for future nature conservation plans. Our study suggests that forecasted land abandonment in the Alps will lead to pre-Neolithic conditions, with significant biodiversity losses in abandoned grassland ecosystems. Thus, management measures for biodiversity, such as ecological compensation areas, are needed in agricultural landscapes with a millennial history of human impact, such as the non-boreal European lowlands. Our study supports the hypothesis that species coexistence is maximized at an intermediate level of disturbances. For instance, species richness decreased when fire exceeded the quasi-natural variability observed during the Mesolithic times. Under a more natural disturbance regime, rather closed Pinus sylvestris and mixed oak forests would prevail.

North-south palaeohydrological contrasts in the central Mediterranean during the Holocene: tentative synthesis and working hypotheses

On the basis of a multi-proxy approach and a strategy combining lacustrine and marine records along a north–south transect, data collected in the central Mediterranean within the framework of a collaborative project have led to reconstruction of high-resolution and well-dated palaeohydrological records and to assessment of their spatial and temporal coherency. Contrasting patterns of palaeohydrological changes have been evidenced in the central Mediterranean: south (north) of around 40° N of latitude, the middle part of the Holocene was characterised by lake-level maxima (minima), during an interval dated to ca. 10 300–4500 cal BP to the south and 9000–4500 cal BP to the north. Available data suggest that these contrasting palaeohydrological patterns operated throughout the Holocene, both on millennial and centennial scales. Regarding precipitation seasonality, maximum humidity in the central Mediterranean during the middle part of the Holocene was characterised by humid winters and dry summers north of ca. 40° N, and humid winters and summers south of ca. 40° N. This may explain an apparent conflict between palaeoclimatic records depending on the proxies used for reconstruction as well as the synchronous expansion of tree species taxa with contrasting climatic requirements. In addition, south of ca. 40° N, the first millennium of the Holocene was characterised by very dry climatic conditions not only in the eastern, but also in the central- and the western Mediterranean zones as reflected by low lake levels and delayed reforestation. These results suggest that, in addition to the influence of the Nile discharge reinforced by the African monsoon, the deposition of Sapropel 1 has been favoured (1) by an increase in winter precipitation in the northern Mediterranean borderlands, and (2) by an increase in winter and summer precipitation in the southern Mediterranean area. The climate reversal following the Holocene climate optimum appears to have been punctuated by two major climate changes around 7500 and 4500 cal BP. In the central Mediterranean, the Holocene palaeohydrological changes developed in response to a combination of orbital, ice-sheet and solar forcing factors. The maximum humidity interval in the south-central Mediterranean started ca. 10 300 cal BP, in correlation with the decline (1) of the possible blocking effects of the North Atlantic anticyclone linked to maximum insolation, and/or (2) of the influence of the remnant ice sheets and fresh water forcing in the North Atlantic Ocean. In the north-central Mediterranean, the lake-level minimum interval began only around 9000 cal BP when the Fennoscandian ice sheet disappeared and a prevailing positive NAO-(North Atlantic Oscillation) type circulation developed in the North Atlantic area. The major palaeohydrological oscillation around 4500–4000 cal BP may be a non-linear response to the gradual decrease in insolation, with additional key seasonal and interhemispheric changes. On a centennial scale, the successive climatic events which punctuated the entire Holocene in the central Mediterranean coincided with cooling events associated with deglacial outbursts in the North Atlantic area and decreases in solar activity during the interval 11 700–7000 cal BP, and to a possible combination of NAO-type circulation and solar forcing since ca. 7000 cal BP onwards. Thus, regarding the centennial-scale climatic oscillations, the Mediterranean Basin appears to have been strongly linked to the North Atlantic area and affected by solar activity over the entire Holocene. In addition to model experiments, a better understanding of forcing factors and past atmospheric circulation patterns behind the Holocene palaeohydrological changes in the Mediterranean area will require further investigation to establish additional high-resolution and well-dated records in selected locations around the Mediterranean Basin and in adjacent regions. Special attention should be paid to greater precision in the reconstruction, on millennial and centennial timescales, of changes in the latitudinal location of the limit between the northern and southern palaeohydrological Mediterranean sectors, depending on (1) the intensity and/or characteristics of climatic periods/oscillations (e.g. Holocene thermal maximum versus Neoglacial, as well as, for instance, the 8.2 ka event versus the 4 ka event or the Little Ice Age); and (2) on varying geographical conditions from the western to the eastern Mediterranean areas (longitudinal gradients). Finally, on the basis of projects using strategically located study sites, there is a need to explore possible influences of other general atmospheric circulation patterns than NAO, such as the East Atlantic–West Russian or North Sea–Caspian patterns, in explaining the apparent complexity of palaeoclimatic (palaeohydrological) Holocene records from the Mediterranean area.

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.

Climate and vegetation changes during the Lateglacial and early–middle Holocene at Lake Ledro (southern Alps, Italy)

Adding to the on-going debate regarding vegetation recolonisation (more particularly the timing) in Europe and climate change since the Lateglacial, this study investigates a long sediment core (LL081) from Lake Ledro (652ma.s.l., southern Alps, Italy). Environmental changes were reconstructed using multiproxy analysis (pollen-based vegetation and climate reconstruction, lake levels, magnetic susceptibility and X-ray fluorescence (XRF) measurements) recorded climate and land-use changes during the Lateglacial and early-middle Holocene. The well-dated and high-resolution pollen record of Lake Ledro is compared with vegetation records from the southern and northern Alps to trace the history of tree species distribution. An altitudedependent progressive time delay of the first continuous occurrence of Abies (fir) and of the Larix (larch) development has been observed since the Lateglacial in the southern Alps. This pattern suggests that the mid-altitude Lake Ledro area was not a refuge and that trees originated from lowlands or hilly areas (e.g. Euganean Hills) in northern Italy. Preboreal oscillations (ca. 11 000 cal BP), Boreal oscillations (ca. 10 200, 9300 cal BP) and the 8.2 kyr cold event suggest a centennial-scale climate forcing in the studied area. Picea (spruce) expansion occurred preferentially around 10 200 and 8200 cal BP in the south-eastern Alps, and therefore reflects the long-lasting cumulative effects of successive boreal and the 8.2 kyr cold event. The extension of Abies is contemporaneous with the 8.2 kyr event, but its development in the southern Alps benefits from the wettest interval 8200-7300 cal BP evidenced in high lake levels, flood activity and pollen-based climate reconstructions. Since ca. 7500 cal BP, a weak signal of pollen-based anthropogenic activities suggest weak human impact. The period between ca. 5700 and ca. 4100 cal BP is considered as a transition period to colder and wetter conditions (particularly during summers) that favoured a dense beech (Fagus) forest development which in return caused a distinctive yew (Taxus) decline.We conclude that climate was the dominant factor controlling vegetation changes and erosion processes during the early and middle Holocene (up to ca. 4100 cal BP).