Sanitary felling of Norway spruce due to spruce bark beetles in Slovenia: A model and projections for various climate change scenarios

A model is presented to predict sanitary felling of Norway spruce (Picea abies) due to spruce bark beetles (Ips typographus, Pityogenes chalcographus) in Slovenia according to different climate change scenarios. The model incorporates 21 variables that are directly or indirectly related to the dependent variable, and that can be arranged into five groups: climate, forest, landscape, topography, and soil. The soil properties are represented by 8 variables, 4 variables define the topography, 4 describe the climate, 4 define the landscape, and one additional variable provides the quantity of Norway spruce present in the model cell. The model was developed using the M5′ model tree. The basic spatial unit of the model is 1 km², and the time resolution is 1 year. The model evaluation was performed by three different measures: (1) the correlation coefficient (51.9%), (2) the Theil's inequality coefficient (0.49) and (3) the modelling efficiency (0.32). Validation of the model was carried out by 10-fold cross-validation. The model tree consists of 28 linear models, and model was calculated for three different climate change scenarios extending over a period until 2100, in 10-year intervals. The model is valid for the entire area of Slovenia; however, climate change projections were made only for the Maribor region (596 km²). The model assumes that relationships among the incorporated factors will remain unchanged under climate change, and the influence of humans was not taken into account. The structure of the model reveals the great importance of landscape variables, which proved to be positively correlated with the dependent variable. Variables that describe the water regime in the model cell were also highly correlated with the dependent variable, with evapotranspiration and parent material being of particular importance. The results of the model support the hypothesis that bark beetles do greater damage to Norway spruce artificially planted out of its native range in Slovenia, i.e., lowlands and soils rich in N, P, and K. The model calculation for climate change scenarios in the Maribor region shows an increase in sanitary felling of Norway spruce due to spruce bark beetles, for all scenarios. The model provides a path towards better understanding of the complex ecological interactions involved in bark beetle outbreaks. Potential application of the results in forest management and planning is discussed.     

Simulating forest succession after blowdown events: The crucial role of space for a realistic management

Ecological patterns vary in space and time. Therefore, when using dynamic models in ecology, the spatial aspect should not be neglected prematurely since it could possibly change the model outcomes to a considerable extent. In view of this problem, we describe here a method how to construct a non-spatial version from a spatially explicit simulation model. The principle idea is to suppress the spatial correlations of cells in a grid in time by continuously re-assigning a random neighbourhood for each cell on the grid. Since this procedure actually eliminates the spatial dimensions, it allows to quantify the unadulterated impact of spatial processes on the model results. To illustrate an important application of this approach in the context of forest management we use a grid-based model that simulates succession of Norway spruce (Picea abies (L.) Karst.) at mountainous sites after blowdown events. The output of this model is compared with the results of the deduced non-spatial version of this model regarding the predicted amount of re-growing trees. The non-spatial version dramatically overestimates the number of spruce trees on different microsites. Thus, the uncritical use of the non-spatial model might give reason to wrong management decisions that are based on too optimistic predictions. In practice, this may lead to dangerous situations, especially in mountain forests serving as protection against avalanches and landslides. This example demonstrates the successful applicability of our approach. Our method can be interpreted as a contribution to an extended sensitivity analysis: it analyses the sensitivity of the results due to structural changes of the model. This sensitivity allows one to estimate the redundancy or the necessity of spatially explicit processes in a model with regard to the parsimony principle of modelling. Since our approach is not dependent on special features of the simulation model used here, it is assumed to be applicable for other spatial models, too, and can thus be considered of general interest for a diligent model analysis.     

An innovative computer design for modeling forest landscape change in very large spatial extents with fine resolutions

Although forest landscape models (FLMs) have benefited greatly from ongoing advances of computer technology and software engineering, computing capacity remains a bottleneck in the design and development of FLMs. Computer memory overhead and run time efficiency are primary limiting factors when applying forest landscape models to simulate large landscapes with fine spatial resolutions and great vegetation detail. We introduce LANDIS PRO 6.0, a landscape model that simulates forest succession and disturbances on a wide range of spatial and temporal scales. LANDIS PRO 6.0 improves on existing forest landscape models with two new data structures and algorithms (hash table and run-length compression). The innovative computer design enables LANDIS PRO 6.0 to simulate very large (>10⁸ ha) landscapes with a 30-m spatial resolution, which to our knowledge no other raster forest landscape models can do. We demonstrate model behavior and performance through application to five nested forest landscapes with varying sizes (from 1 million to 100 million 0.09-ha cells) in the southern Missouri Ozarks. The simulation results showed significant and variable effects of changing spatial extent on simulated forest succession patterns. Results highlighted the utility of a model like LANDIS PRO 6.0 that is capable of efficiently simulating large landscapes and scaling up forest landscape processes to a common regional scale of analysis. The programming methodology presented here may significantly advance the development of next generation of forest landscape models.     

The impact of fragmentation and density regulation on forest succession in the Atlantic rain forest

The Atlantic Rain Forest, an important biodiversity hot spot, has faced severe habitat loss since the last century which has resulted in a highly fragmented landscape with a large number of small forest patches (<100 ha). For conservation planning it is essential to understand how current and future forest regeneration depends on ecological processes, fragment size and the connection to the regional seed pool. We have investigated the following questions by applying the forest growth simulation model FORMIND to the situation of the Atlantic Forest in the state of São Paulo, SE Brazil: (1) which set of parameters describing the local regeneration and level of density regulation can reproduce the biomass distribution and stem density of an old growth forest in a reserve? (2) Which additional processes apart from those describing the dynamics of an old growth forest, drive forest succession of small isolated fragments? (3) Which role does external seed input play during succession? Therefore, more than 300 tree species have been classified into nine plant functional types (PFTs), which are characterized by maximum potential height and shade tolerance. We differentiate between two seed dispersal modes: (i) local dispersal, i.e. all seedlings originated from fertile trees within the simulated area and (ii) external seed rain. Local seed dispersal has been parameterized following the pattern oriented approach, using biomass estimates of old growth forest. We have found that moderate density regulation is essential to achieve coexistence for a broad range of regeneration parameters. Considering the expected uncertainty and variability in the regeneration processes it is important that the forest dynamics are robust to variations in the regeneration parameters. Furthermore, edge effects such as increased mortality at the border and external seed rain have been necessary to reproduce the patterns for small isolated fragments. Overall, simulated biomass is much lower in the fragments compared to the continuous forest, whereas shade tolerant species are affected most strongly by fragmentation. Our simulations can supplement empirical studies by extrapolating local knowledge on edge effects of fragments to larger temporal and spatial scales. In particular our results show the importance of external seed rain and therefore highlight the importance of structural connectivity between regenerating fragments and mature forest stands.     

Modelling forest management within a global vegetation model—Part 1: Model structure and general behaviour

This article describes a new forest management module (FMM) that explicitly simulates forest stand growth and management within a process-based global vegetation model (GVM) called ORCHIDEE. The net primary productivity simulated by ORCHIDEE is used as an input to the FMM. The FMM then calculates stand and management characteristics such as stand density, tree size distribution, tree growth, the timing and intensity of thinnings and clear-cuts, wood extraction and litter generated after thinning. Some of these variables are then fed back to ORCHIDEE. These computations are made possible with a distribution-based modelling of individual tree size. The model derives natural mortality from the relative density index (rdi), a competition index based on tree size and stand density. Based on the common forestry management principle of avoiding natural mortality, a set of rules is defined to calculate the recurrent intensity and frequency of forestry operations during the stand lifetime. The new-coupled model is called ORCHIDEE-FM (forest management).The general behaviour of ORCHIDEE-FM is analysed for a broadleaf forest in north-eastern France. Flux simulation throughout a forest rotation compare well with the literature values, both in absolute values and dynamics.Results from ORCHIDEE-FM highlight the impact of forest management on ecosystem C-cycling, both in terms of carbon fluxes and stocks. In particular, the average net ecosystem productivity (NEP) of 225 gC m⁻² year⁻¹ is close to the biome average of 311 gC m⁻² year⁻¹. The NEP of the “unmanaged” case is 40% lower, leading us to conclude that management explains 40% of the cumulated carbon sink over 150 years. A sensitivity analysis reveals 4 major avenues for improvement: a better determination of initial conditions, an improved allocation scheme to explain age-related decline in productivity, and an increased specificity of both the self-thinning curve and the biomass-diameter allometry.     

Are trade-offs in plant resprouting manifested in community seed banks?

Trade-offs in allocation to resprouting vs. seedling regeneration in plants are predicted to occur along fire disturbance gradients. Increased resprouting ability should be generally favored in plant communities with a high probability of crown fire return. Hence, communities dominated by resprouters are predicted to have smaller seed banks than those dominated by species killed by fire. We tested whether there were trait shifts in resprouting ability among growth forms (short-lived herbaceous vs. ground-dwelling perennials vs. shrubs) and among communities (rocky outcrop vs. sclerophyll forest) with contrasting probabilities of crown fire return. Resprouting was more common in ground-dwelling perennials and in the sclerophyll forest community with a high probability of crown fire. Soil seed banks were sampled in rocky outcrop and sclerophyll forest communities in recently burned (18 months) and long-since-burned (12 years) locations at interspersed replicated sites. Collected seed banks were treated with orthogonal treatments of fire stimuli or no stimuli, and seedling emergence was measured in controlled conditions. Seed bank composition reflected the pattern of extant vegetation, with resprouting species being more common in the community with a higher probability of crown fire. Overall, however, resprouting species were poorly represented in the seed bank compared to those species killed by fire. Predicted shifts in allocation to seed production were strongly manifested in community seed banks across the disturbance gradient. Fewer species, seedlings, and seedlings per adult emerged from seed banks in the sclerophyll forest. This suggests that the dominance of resprouting species influences recruitment at the community scale. Community patterns in the seed bank also reflected predicted trade-offs with plant size and growth rate. Short-lived species that are killed by fire dominated the seed bank on rocky outcrops, while longer-lived resprouting species were found in low abundance. Life history trade-offs in persistence and regeneration strongly contribute to coexistence patterns between and within communities with contrasting probabilities of fire return.     

Application of a three-dimensional model for assessing effects of small clear-cuttings on radiation and soil temperature

A three-dimensional model Mixfor-3D of soil–vegetation–atmosphere transfer (SVAT) was developed and applied to estimate possible effects of tree clear-cutting on radiation and soil temperature regimes of a forest ecosystem. The Mixfor-3D model consists of several closely coupled 3D sub-models describing: forest stand structure; radiative transfer in a forest canopy; turbulent transfer of sensible heat, H₂O and CO₂ between ground surface and the atmospheric surface layer; evapotranspiration of ground surface vegetation and soil; heat and moisture transfer in soil. The model operates with the horizontal grid resolution, 2 m × 2 m; vertical resolution, 1 m and primary time step, 1 h.     

The importance of spatial autocorrelation,extent and resolution in predicting forest bird occurrence

Concerns about declines in forest biodiversity underscore the need for accurate estimates of the distribution and abundance of organisms at large scales and at resolutions that are fine enough to be appropriate for management. This paper addresses three major objectives: (i) to determine whether the resolution of typical air photo-derived forest inventory is sufficient for the accurate prediction of site occupancy by forest birds. We compared prediction success of habitat models using air photo variables to models with variables derived from finer resolution, ground-sampled vegetation plots. (ii) To test whether incorporating spatial autocorrelation into habitat models via autologistic regression increases prediction success. (iii) To determine whether landscape structure is an important factor in predicting bird distribution in forest-dominated landscapes. Models were tested locally (Greater Fundy Ecosystem [GFE]) using cross-validation, and regionally using an independent data set from an area located ca. 250 km to the northwest (Riley Brook [RB]). We found significant positive spatial autocorrelation in the residuals of at least one habitat model for 76% (16/21) of species examined. In these cases, the logistic regression assumption of spatially independent errors was violated. Logistic models that ignored spatial autocorrelation tended to overestimate habitat effects. Though overall prediction success was higher for autologistic models than logistic models in the GFE, the difference was only significantly improved for one species. Further, the inclusion of spatial covariates did little to improve model performance in the geographically discrete study area. For 62% (13/21) of species examined, landscape variables were significant predictors of forest bird occurrence even after statistically controlling for stand-level variability. However, broad spatial extents explained less variation than local factors. In the GFE, 76% (16/21) of air photo and 81% (17/21) of ground plot models were accurate enough to be of practical utility (AUC > 0.7). When applied to RB, both model types performed effectively for 55% (11/20) of the species examined. We did not detect an overall difference in prediction success between air photo and ground plot models in either study area. We conclude that air photo data are as effective as fine resolution vegetation data for predicting site occupancy for the majority of species in this study. These models will be of use to forest managers who are interested in mapping species distributions under various timber harvest scenarios, and to protected areas planners attempting to optimize reserve function.     

A geospatial model of forest dynamics with controlled trend surface

This paper proposes a method of controlled trend surface to simultaneously account for large-scale spatial trends and non-spatial local effects. With this method, a geospatial model of forest dynamics was developed for the Alaska boreal forest from 446 constantly monitored permanent sample plots. The geospatial component of this model represented large-scale spatial trends in recruitment, diameter growth, and mortality. The model was tested on two sets of validation plots which represented temporal and spatial extensions of the current sample coverage. The results suggest that the controlled trend surface model was generally more accurate than both the non-spatial and conventional trend surface models. With this model, we mapped the forest dynamics of the entire Alaska boreal region by aggregating predicted stand states across the region. It was predicted that under current conditions of climate and natural disturbances, most of the Alaska boreal forest region may undergo a major shift from deciduous-dominant to conifer-dominant, with an average increase of 0.33 m² ha year⁻¹ in basal area over the Twenty-First Century.     

Vegetation competition model for water and light limitation. II: Spatial dynamics of groundwater and vegetation

In temperate climates groundwater can have a profound effect on vegetation, because it strongly influences the spatio-temporal distribution of soil moisture in the rootzone and therefore the occurrence of water and oxygen stress of vegetation. This article focuses on vegetation and groundwater dynamics along a hill slope by developing and evaluating a fully coupled hydrological-vegetation model for a temperate forest ecosystem. The vegetation model is described in part 1 of this series of two papers. To simulate the hydrology an extended version of the saturated-unsaturated hydrological model STARWARS has been used. The coupled model is used to investigate both the short and long-term dynamics for a system of two species. Both compete for light and water where one is adapted to wet conditions and the other to dry conditions. The daily dynamics show that the influence of groundwater is particularly strong in spring when waterlogging occurs due to decreased evapotranspiration in winter. Long simulation runs of 1000 years were performed to study the equilibrium state for the two species. Comparison of simulation results with observations of groundwater depth and vegetation types along a dry-wet gradient in a natural forest shows that a reductionist approach is able to capture these patterns well. Sensitivity analysis shows that the border between wet- and dry-adapted species moves upslope with increased rainfall, decreased slope angle and decreased aquifer thickness. These results are similar to previous findings which were based on global maximization of ecosystem evaporation or minimizing ecosystem stress. Comparison of runs with a fixed and a dynamic groundwater table shows that a dynamic groundwater table facilitates a wider transition zone between vegetation types along the hill slope. In this transition the biomass of vegetation is higher in the case of a dynamic groundwater than in case of a static groundwater table. This underlines the importance of incorporating spatial groundwater dynamics in models of groundwater influenced ecosystems.     

Spatio-temporal analysis of alpine ecotones: A spatial explicit model targeting altitudinal vegetation shifts

There is general agreement in literature that Alpine vegetation belt ecotones have shown a trend of upward migration in the last few decades. Despite the potential of such shifts as indicators of global change effects in mountain ecosystems, there are relatively few works focused on their assessment in a systematic and spatially explicit way. In this work our aim is to quantify the altitudinal shifts and analyse the spatial pattern dynamics of mountain ecotones. We developed a novel procedure to delineate the current and former state of three characteristic mountain ecotones, which we formalised as forest, tree and tundra lines. Our approach is based on the recognition of altitudinal extreme outposts identified with ecotone locations at a slope scale. The integration of multi-temporal datasets allows the identification and quantification of altitudinal advances and retreats in the outpost locations for a given period. We tested the method in a section of the Italian Alps for the period 1957-2003. Results show a general trend of an increase in altitude for the three ecotones, despite the occurrence of occasional decreases. We estimate decadal altitude increments of 25 m for forest line, 13 m for treeline and 11 m for tundra line. We also identified changes in ecotone spatial morphology between the two dates, with significant implications in connectivity and colonisation dynamics.     

Mapping eucalypt forest susceptible to dieback associated with bell miners (Manorina melanophys) using laser scanning, SPOT 5 and ancillary topographical data

Mapping the location and extent of forest at risk from damaging agents or processes assists forest managers in prioritizing their planning and operational mitigation activities. In Australia, Bell Miner Associated Dieback (BMAD) refers to a form of canopy decline observed in eucalypt crowns occupied by colonies of bell miners (Manorina melanophrys). High densities of bell miners are associated with decreased avian abundance and diversity and an increase in psyllid abundance on crown foliage. BMAD has recently been nominated as a key threatening process in New South Wales (NSW). Consequently, a modelling system for predicting bell miner distribution in coastal eucalypt forests of NSW has been developed. The presence or absence of bell miners was recorded in 130 plots located within a 12,800 ha catchment study area containing a range of eucalypt forest types. The modelling system was produced by integrating a machine learning software suite (WEKA), and the statistical software R within the geographic resources analysis support system (GRASS) geographical information system (GIS). The variable modelled was the binary variable: presence or absence of bell minors. Six modelling techniques (Logistic regression; generalised additive models; two tree-based ensemble classification algorithms, random forest and Adaboost and Neural Networks) were integrated with airborne laser scanning; SPOT 5 and topographic derived variables. Model evaluation and parameter selection were measured by three threshold dependent measures (sensitivity, specificity and kappa) and the threshold independent Receiver Operator Curve (ROC) analysis. The final presence and absence maps were obtained through maximisation of the kappa statistic and applied at a resolution of 10 m across the entire catchment study area. For this data set, the most accurate algorithm for predicting the distribution of bell miner colonies was random forest (kappa = 0.84; ROC area under curve = 0.97). Variables most commonly selected in the six models were the laser scanning metrics; coefficient of variation, skewness, and the 10th and 90th percentiles derived from the shape of the height frequency distribution which, in turn, is directly influenced by vertical structure of the forest. An image textural statistic based on the shortwave infrared (SWIR) band of SPOT 5 was also commonly selected by the models. The SWIR band is sensitive to vegetation and soil moisture content. These models predicted that forest stands with a sparse eucalypt canopy over a moist, dense understorey were susceptible to being colonised by bell miners and hence BMAD.     

Modelling soil carbon development in Swedish coniferous forest soils—An uncertainty analysis of parameters and model estimates using the GLUE method

Boreal forest soils such as those in Sweden contain a large active carbon stock. Hence, a relatively small change in this stock can have a major impact on the Swedish national CO₂ balance. Understanding of the uncertainties in the estimations of soil carbon pools is critical for accurately assessing changes in carbon stocks in the national reports to UNFCCC and the Kyoto Protocol. Our objective was to analyse the parameter uncertainties of simulated estimates of the soil organic carbon (SOC) development between 1994 and 2002 in Swedish coniferous forests with the Q model. Both the sensitivity of model parameters and the uncertainties in simulations were assessed. Data of forests with Norway spruce, Scots pine and Lodgepole pine, from the Swedish Forest Soil Inventory (SFSI) were used. Data of 12 Swedish counties were used to calibrate parameter settings; and data from another 11 counties to validate. The “limits of acceptability” within GLUE were set at the 95% confidence interval for the annual, mean measured SOC at county scale. The calibration procedure reduced the parameter uncertainties and reshaped the distributions of the parameters county-specific. The average measured and simulated SOC amounts varied from 60 t C ha⁻¹ in northern to 140 t C ha⁻¹ in the southern Sweden. The calibrated model simulated the soil carbon pool within the limits of acceptability for all calibration counties except for one county during one year. The efficiency of the calibrated model varied strongly; for five out of 12 counties the model estimates agreed well with measurements, for two counties agreement was moderate and for five counties the agreement was poor. The lack of agreement can be explained with the high inter-annual variability of the down-scaled measured SOC estimates and changes in forest areas over time. We conclude that, although we succeed in reducing the uncertainty in the model estimates, calibrating of a regional scale process-oriented model using a national scale dataset is a sensitive balance between introducing and reducing uncertainties. Parameter distributions showed to be scale sensitive and county specific. Further analysis of uncertainties in the methods used for reporting SOC changes to the UNFCCC and Kyoto protocol is recommended.     

Estimating the potential impact of vegetation on the water cycle requires accurate soil water parameter estimation

It is well known that vegetation dynamics at the catchment scale depends on the prevailing weather and soil moisture conditions. Soil moisture, however, is not equally distributed in space due to differences in topography, weather patterns, soil properties and the type and amount of vegetation cover. To elucidate the complex interaction between vegetation and soil moisture, the dynamic vegetation model LPJ-GUESS (Smith et al., 2001), which provides estimations of vegetation dynamics, but does not consider lateral water fluxes was coupled with the hydrological TOPMODEL (cf. Beven, 2001) in order to be able to evaluate the importance of these lateral fluxes. The new model LG-TM was calibrated and validated in two climatically different mountain catchments. The estimations of runoff were good, when monthly and weekly time scales were considered, although the low flow periods at winter time were somewhat underestimated. The uncertainty in the climate induced change vegetation carbon storage caused by the uncertainty in soil parameters was up to 3-5 kg C m⁻² (depending on elevation and catchment), compared to the total change in vegetation carbon storage of 5-9 kg C m⁻². Therefore accurate estimates of the parameters influencing the water holding capacity of the soil, for example depth and porosity, are necessary when estimating future changes in vegetation carbon storage. Similarly, changes in plant transpiration due to climatic changes could be almost double as high (88 mm m⁻²) in the not calibrated model compared to the new model version (ca 50 mm m⁻² transpiration change). The uncertainties in these soil properties were found to be more important than the lateral water exchange between grid cells, even in steep topography at least for the temporal and spatial resolution used here.     

The sensitivity of carbon sequestration to harvesting and climate conditions in a temperate cypress forest: Observations and modeling

The role of disturbance and climate factors in determining the forest carbon balance was investigated at a Japanese cypress forest in central Japan with eddy flux measurements, tree-ring analyses, and a terrestrial biosphere model. The forest was established as a plantation after intermittent harvesting and replanting between 1959 and 1977, and acted as a strong carbon sink of approximately 500 g C m⁻² year⁻¹ for the measurement years between 2001 and 2007. A terrestrial biosphere model, BIOME-BGC, was validated using the eddy flux data at daily to interannual timescales, and the tree-ring width data at interannual to decadal timescales. According to the model simulation, during the observation period 270 ± 55 g C m⁻² year⁻¹ was additionally sequestered due to the indirect effects of the harvesting and planting, whereas the increase of CO₂ concentration and the change in climate increased the sink of 110 ± 40 and 30 ± 80 g C m⁻² year⁻¹, respectively. The model simulation shows that the forest is now recovering from harvesting, and that harvesting is a more important determinant of the current carbon sink than either interannual climate anomalies or increased atmospheric CO₂ concentration. We found that harvesting with long rotation length could be effective management activity in order to increase carbon sequestration, if the harvested timber is converted into products with long lifecycles.     

九寨沟和黄龙自然保护区原始林与次生林土壤物理性质比较

森林土壤的结构和贮水保水能力是决定土壤水分及其时空分布的关键,九寨沟和黄龙自然保护区均是以水为灵魂,其水量的多少及时空分布格局对景区景点的视觉观赏效果具有十分重要的作用.通过对几寨沟和黄龙核心景区原始林与次生林土壤容重、持水性能和孔隙度等指标的监测,探讨了不同植被类型下土壤结构及持水能力状况.结果表明:(1)由于较早地得到保护,九寨沟景区原始林、桦槭次生林土壤物理性质均优于四川西部其它同类犁植被,与此相反,人工云杉纯林土壤物理性质明显劣于其它植被类型,可能主要与云杉平根系统对土壤膨胀挤压和单优势乔木群落所形成的不利微气候影响有机物分解归还土壤有关;(2)在黄龙核心景区的原始云、冷杉林,发育于坡积母质的土壤物理性质明显优于发育于钙华母质卜的土壤物理性质.图3表1参31     

Fragmentation drives tropical forest fragments to early successional states: A modelling study for Brazilian Atlantic forests

Land use leads to massive habitat destruction and fragmentation in tropical forests. Despite its global dimensions the effects of fragmentation on ecosystem dynamics are not well understood due to the complexity of the problem. We present a simulation analysis performed by the individual-based model FORMIND. The model was applied to the Brazilian Atlantic Forest, one of the world's biodiversity hot spots, at the Plateau of São Paulo. This study investigates the long-term effects of fragmentation processes on structure and dynamics of different sized remnant tropical forest fragments (1-100 ha) at community and plant functional type (PFT) level. We disentangle the interplay of single effects of different key fragmentation processes (edge mortality, increased mortality of large trees, local seed loss and external seed rain) using simulation experiments in a full factorial design.Our analysis reveals that particularly small forest fragments below 25 ha suffer substantial structural changes, biomass and biodiversity loss in the long term. At community level biomass is reduced up to 60%. Two thirds of the mid- and late-successional species groups, especially shade-tolerant (late successional climax) species groups are prone of extinction in small fragments. The shade-tolerant species groups were most strongly affected; its tree number was reduced more than 60% mainly by increased edge mortality. This process proved to be the most powerful of those investigated, explaining alone more than 80% of the changes observed for this group. External seed rain was able to compensate approximately 30% of the observed fragmentation effects for shade-tolerant species.Our results suggest that tropical forest fragments will suffer strong structural changes in the long term, leading to tree species impoverishment. They may reach a new equilibrium with a substantially reduced subset of the initial species pool, and are driven towards an earlier successional state. The natural regeneration potential of a landscape scattered with forest fragments appears to be limited, as external seed rain is not able to fully compensate for the observed fragmentation-induced changes. Our findings suggest basic recommendations for the management of fragmented tropical forest landscapes.     

Modelling the spatial distribution of montane and subalpine forests in the central Alps using digital elevation models

The analysis of complex interactions between spatial distribution patterns of site factors and vegetation types is crucial for understanding high mountain ecosystems, especially in the view of a changing climate. Therefore, in the present study, a GIS and remote sensing-based approach is followed to produce a vegetation map for a study area in the Western Alps (Switzerland). Two major forest alliances are chosen for analysis: subalpine coniferous forest Vaccinio-Piceion/Larici-Pinetum cembrae and montane oak forest Quercion pubescenti-petraeae. As spatial information on site factors is commonly lacking in mountain areas, the use of a digital elevation model (DEM) is a potential substitute for use in vegetation analyses: it highly correlates with temperature, moisture, geomorphological processes and disturbance factors. Thus, it is important to analyse the capabilities of a DEM for indicating habitat conditions in a landscape characterised by high topodiversity and a patchwork of microclimatic habitats.For the purpose of identifying the potential of landform parameters for the indication of forest habitat structures in the present study, 24 primary and secondary landform parameters have been derived, indicating temperature and moisture distribution, exposure towards wind, snow, etc. Quantitative analyses were performed using statistical means such as contingency correlation coefficients and principal components analysis. The results formed the basis for the development of parallel-epiped-vegetation models (PED) used to simulate the spatial distribution patterns of the subalpine coniferous and the montane oak forest. It can be shown that topographic variables derived from a DEM at a spatial resolution of 25 m are very useful for indicating habitats of large forest types. Additionally potential forest sites in the cultural landscape, removed by human logging, can be reconstructed.Inaccuracies within the simulation results can partly be attributed to the insufficient parameterisation of geomorphologic activity and to poor spatial resolution of the DEM as compared to the vegetation data. Although the lack of information on the human dimension leads to some uncertainties in the interpretation of spatial patterns of vegetation, the exclusive use of topographic variables in vegetation models for the indication of forest habitats is very promising.     

Foliage profiles of individual trees determine competition,self-thinning,biomass and NPP of a Cryptomeria japonica forest stand: A simulation study based on a stand-scale process-based forest model

A simulation study was carried out to investigate simultaneously the effects of eco-physiological parameters on competitive asymmetry, self-thinning, stand biomass and NPP in a temperate forest using an atmosphere–vegetation dynamics interactive model (MINoSGI). In this study, we selected three eco-physiological relevant parameters as foliage profiles (i.e. vertical distribution of leaf area density) of individual trees (distribution pattern is described by the parameter η), biomass allocation pattern in individual tree growth (χ) and the maximum carboxylation velocity (V_max). The position of the maximal leaf area density shifts upward in the canopy with increasing η. For scenarios with η < 4 (foliage concentrated in the lowest canopy layer) or η > 12 (foliage concentrated in the uppermost canopy layer), a low degree of competitive asymmetry was produced. These scenarios resulted in the survival of subordinate trees due to a brighter lower canopy environment when η < 4 or the generation of spatially separated foliage profiles between dominant and subordinate trees when η > 12. In contrast, competition between trees was most asymmetric when 4 ≤ η ≤ 12 (vertically widespread foliage profile in the canopy), especially when η = 8. In such cases, vertically widespread foliage of dominant trees lowered the opportunity of light acquisition for subordinate trees and reduced their carbon gain. The resulting reduction in carbon gain of subordinate trees yielded a higher degree of competitive asymmetry and ultimately higher mortality of subordinate trees. It was also shown that 4 ≤ η ≤ 12 generated higher self-thinning speed, smaller accumulated NPP, litter-fall and potential stand biomass as compared with the scenarios with η < 4 or η > 12. In contrast, our simulation revealed small effects of χ or V_max on the above-mentioned variables as compared with those of η. In particular, it is notable that greater V_max would not produce greater potential stand biomass and accumulated NPP although it has been thought that physiological parameters relevant to photosynthesis such as V_max influence dynamic changes in forest stand biomass and NPP (e.g. the greater the V_max, the greater the NPP). Overall, it is suggested that foliage profiles rather than biomass allocation or maximum carboxylation velocity greatly govern forest dynamics, stand biomass, NPP and litter-fall.