Simulation and analysis of outbreaks of bark beetle infestations and their management at the stand level

Outbreaks of bark beetles in forests can result in substantial economic losses. Understanding the factors that influence the development and spread of bark beetle outbreaks is crucial for forest management and for predicting outbreak risks, especially with the expected global warming. Although much research has been done on the ecology and phenology of bark beetles, the complex interplay between beetles, host trees, beetle antagonists and forest management makes predicting beetle population development especially difficult. Using the recent infestations of the European Spruce Bark Beetle (Ips typographus L. Col. Scol.) in the Bavarian Forest National Park (Germany) as a case study, we developed a spatially explicit agent-based simulation model (SAMBIA) that takes into account individual trees and beetles. This model primarily provides a tool for analysing and understanding the spatial and temporal aspects of bark beetles outbreaks at the stand scale. Furthermore, the model should allow an estimation of the effectiveness of concurrent impacts of both antagonists and management to confine outbreak dynamics in practice. We also used the model to predict outbreak probabilities in various settings. The simulation results indicated a distinct threshold behaviour of the system in response to pressure by antagonists or management of the bark beetle population. Despite the different scenarios considered, we were able to extract from the simulations a simple rule of thumb for the successful control of an outbreak: if roughly 80% of individual beetles are killed by antagonists or foresters, outbreaks will rarely take place. Our model allows the core dynamics of this complex system to be reduced to this inherent common denominator.     

Antennal responses of four species of tree-killing bark beetles (Coleoptera: Scolytidae) to volatiles collected from beetles,and their host and nonhost conifers

Summary. Host selection in tree-killing bark beetles (Coleoptera: Scolytidae) is mediated by a complex of semiochemical cues. Using gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectrometric analyses, we conducted a comparative study of the electrophysiological responses of four species of tree-killing bark beetles, the Douglas-fir beetle, Dendroctonus pseudotsugae, Hopkins, the mountain pine beetle, D. ponderosae Hopkins, the spruce beetle, D. rufipennis Kirby, and the western balsam bark beetle, Dryocoetes confusus Swaine, to volatiles captured by aeration of 1) bole and foliage of four sympatric species of conifers, Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, lodgepole pine, Pinus contorta var. latifolia Engelm., interior spruce, Picea engelmannii x glauca, and interior fir, Abies lasiocarpa x bifolia, and 2) con- and heterospecific beetles at three stages of attack. We identified 13 monoterpenes in the conifers and nine compounds in the volatiles of beetles that elicited antennal responses. There was no qualitative difference in the terpene constitution of the four species of conifers and very little difference across beetle species in their antennal response to compounds from conifers or beetles. The lack of species-specific major or minor components in conifers suggests that beetles would need to detect differences in the ratios of different compounds in conifers to discriminate among them. Attraction to hosts and avoidance of nonhost conifers may be accentuated by perception of compounds emitted by con- and heterospecific beetles, respectively. The 22 compounds identified are candidate semiochemicals with potential behavioural roles in host location and discrimination.     

Impacts of changing frost regimes on Swedish forests: Incorporating cold hardiness in a regional ecosystem model

Understanding the effects of climate change on boreal forests which hold about 7% of the global terrestrial biomass carbon is a major issue. An important mechanism in boreal tree species is acclimatization to seasonal variations in temperature (cold hardiness) to withstand low temperatures during winter. Temperature drops below the hardiness level may cause frost damage. Increased climate variability under global and regional warming might lead to more severe frost damage events, with consequences for tree individuals, populations and ecosystems. We assessed the potential future impacts of changing frost regimes on Norway spruce (Picea abies L. Karst.) in Sweden. A cold hardiness and frost damage model were incorporated within a dynamic ecosystem model, LPJ-GUESS. The frost tolerance of Norway spruce was calculated based on daily mean temperature fluctuations, corresponding to time and temperature dependent chemical reactions and cellular adjustments. The severity of frost damage was calculated as a growth-reducing factor when the minimum temperature was below the frost tolerance. The hardiness model was linked to the ecosystem model by reducing needle biomass and thereby growth according to the calculated severity of frost damage. A sensitivity analysis of the hardiness model revealed that the severity of frost events was significantly altered by variations in the hardening rate and dehardening rate during current climate conditions. The modelled occurrence and intensity of frost events was related to observed crown defoliation, indicating that 6-12% of the needle loss could be attributed to frost damage. When driving the combined ecosystem-hardiness model with future climate from a regional climate model (RCM), the results suggest a decreasing number and strength of extreme frost events particularly in northern Sweden and strongly increasing productivity for Norway spruce by the end of the 21st century as a result of longer growing seasons and increasing atmospheric CO₂ concentrations. However, according to the model, frost damage might decrease the potential productivity by as much as 25% early in the century.     

Modelling habitat overlap among sympatric mesocarnivores in southern Illinois,USA

Few researchers have developed large-scale habitat models for sympatric carnivore species. We created habitat models for red foxes (Vulpes vulpes), coyotes (Canis latrans) and bobcats (Lynx rufus) in southern Illinois, USA, using the Penrose distance statistic, remotely sensed landscape data, and sighting location data within a GIS. Our objectives were to quantify and spatially model potential habitat differences among species. Habitat variables were quantified for 1-km² buffered areas around mesocarnivore sighting locations. Following variable reduction procedures, five habitat variables (percentage of grassland patches, interspersion–juxtaposition of forest patches, mean fractal dimension of wetland patches and the landscape, and road density) were used for analysis. Only one variable differed (P < 0.05) between red fox and coyote sighting areas (road density) and bobcat and coyote sighting areas (mean fractal dimension of the landscape). However, all five variables differed between red fox and bobcat sighting areas, indicating considerable differences in habitat affiliation between this pair-group. Compared to bobcats, red fox sightings were affiliated with more grassland cover and larger grassland patches, higher road densities, lower interspersion and juxtaposition of forest patches, and lower mean fractal dimension of wetland patches. These differences can be explained by different life history requirements relative to specific cover types. We then used the Penrose distance statistic to create habitat models for red foxes and bobcats, respectively, based on the five-variable dataset. An independent set of sighting locations were used to validate these models; model fit was good with 65% of mesocarnivore locations within the top 50% of Penrose distance values. In general, red foxes were affiliated with mixtures of agricultural and grassland cover, whereas bobcats were associated with a combination of grassland, wetland, and forest cover. The greatest habitat overlap between red foxes and bobcats was found at the interface between forested areas and more open cover types. Our study provides insight into habitat overlap among sympatric mesocarnivores, and the distance-based modelling approach we used has numerous applications for modelling wildlife–habitat relationships over large scales.     

A process-based population dynamics model to explore target and non-target impacts of a biological control agent

The risks and benefits associated with efforts to control invasive alien species using classical biological control are being subjected to increasing scrutiny. A process-based population dynamics model was developed to explore the interactions between a folivorous biological control agent, Cleopus japonicus, and its plant host Buddleja davidii. The model revealed that climate could have a significant impact upon the interactions between B. davidii and C. japonicus. At the coolest sites, the impact of C. japonicus on B. davidii was slowed, but it was still eventually capable of controlling populations of B. davidii. At the warmer sites where both B. davidii and C. japonicus grew faster, B. davidii succumbed rapidly to weevil damage. We hypothesise that barring an encounter with a natural enemy, C. japonicus will eventually be able to provide sustained control B. davidii throughout the North Island of New Zealand. The model scenarios illustrate the potential for the C. japonicus population to attain high densities rapidly, and to defoliate patches of B. davidii, creating the potential for spill-over feeding on non-target plants. The potential magnitude of this threat will depend partly on the climate suitability for C. japonicus, the pattern by which it migrates in response to a reduction in the available leaf resource, and the suitability of non-target plants as hosts. In all migration scenarios considered, the pattern of population growth and resource consumption by C. japonicus was exponential, with a strong tendency toward complete utilisation of resource patches more quickly at the warmer compared to colder sites. In addition to providing some useful hypotheses about the effects of climate on the biological control system, and the non-target risks, it also provides some insight into the mechanisms by which climate affects the system.     

Effects of changing climate on water and nitrogen availability with implications on the productivity of Norway spruce stands in Southern Finland

An integrated process-based model was used to study how the changing climate affects the availability of water and nitrogen, and consequently the dynamics of productivity of Norway spruce (Picea abies) on sites with different initial soil water conditions in southern Finland over a 100-year period. The sensitivity of the total stem volume growth in relation to short-term availability of water and nitrogen was also analyzed. We found that a high proportion (about 88–92%) of the total precipitation was lost in total evapotranspiration (incl. canopy evaporation (E_c), transpiration (E_t) and ground surface evaporation (E_g)), under both current and changing climate. Furthermore, under the changing climate the cumulative amount of E_c and E_g were significantly higher, while E_t was largely lower than under the current climate. Additionally, the elevated temperature and increased expansion of needle area index (L) enhanced E_c. Under the changing climate, the increasing soil water deficit (W_d) reduced the canopy stomatal conductance (g_cs), the E_t, humus yield (H, available nitrogen source) and nitrogen uptake (N_up) of the trees. During the latter phases of the simulation period, the canopy net photosynthesis (P_nc) was lower due to the reduced N_up and soil water availability. This also reduced the total stem volume production (V_s) on the site with the lower initial soil moisture content. The growth was slightly more sensitive to the change in precipitation than to the change in nitrogen content of the needles, when the elevated temperature was assumed. According to our findings, drought stress episodes may become more frequent under the changing climate. Thus, adaptive management strategies should be developed to sustain the productivity of Norway spruce in these conditions, and thus, to mitigate the adverse impacts of climate change.     

Modeling amphibian energetics, habitat suitability, and movements of western toads, Anaxyrus (=Bufo) boreas, across present and future landscapes

Effective conservation of amphibian populations requires the prediction of how amphibians use and move through a landscape. Amphibians are closely coupled to their physical environment. Thus an approach that uses the physiological attributes of amphibians, together with knowledge of their natural history, should be helpful. We used Niche Mapper™ to model the known movements and habitat use patterns of a population of Western toads (Anaxyrus (=Bufo) boreas) occupying forested habitats in southeastern Idaho. Niche Mapper uses first principles of environmental biophysics to combine features of topography, climate, land cover, and animal features to model microclimates and animal physiology and behavior across landscapes. Niche Mapper reproduced core body temperatures (T_c) and evaporation rates of live toads with average errors of 1.6 ± 0.4 °C and 0.8 ± 0.2 g/h, respectively. For four different habitat types, it reproduced similar mid-summer daily temperature patterns as those measured in the field and calculated evaporation rates (g/h) with an average error rate of 7.2 ± 5.5%. Sensitivity analyses indicate these errors do not significantly affect estimates of food consumption or activity. Using Niche Mapper we predicted the daily habitats used by free-ranging toads; our accuracy for female toads was greater than for male toads (74.2 ± 6.8% and 53.6 ± 15.8%, respectively), reflecting the stronger patterns of habitat selection among females. Using these changing to construct a cost surface, we also reconstructed movement paths that were consistent with field observations. The effect of climate warming on toads depends on the interaction of temperature and atmospheric moisture. If climate change occurs as predicted, results from Niche Mapper suggests that climate warming will increase the physiological cost of landscapes thereby limiting the activity for toads in different habitats.     

Spatio-temporal modelling of ground vegetation development in mountain spruce forests

More complex models of forest ecosystems are required to understand how land-cover changes can impact vegetation dynamics and spatial pattern. In order to document spatio-temporal modelling abilities, the observations conducted in the declined climax mountain Norway spruce forest during the recovery period (1995-2006) are used for simulation and spatial analysis in the GIS environment. The developed spatio-temporal model is used for simulation of forest vegetation dynamics in a mountain spruce forest in the framework of regeneration processes after stress from air pollution. In order to explore the spatial and temporal phenomena of regeneration processes, the spatio-temporal model is based on a large set of ordinary differential equations that solve dynamic processes in sets of microsites arranged in grids for each ground vegetation species and each age group of Norway spruce seedlings. The spatial extent of the explored site is composed of a set of 50 × 50 microsites. Each microsite is represented by a square with dimensions of 1 m × 1 m. The presented simulation studies are mainly focused on seedlings from the seed year 1992, in order to explore the longest monitored time series of survival. It is based on exponential growth models that are related to the environmental conditions for each microsite. The canopy gaps based on estimates of the local crown projected area, the soil type layer, and the dominant grass density are used to provide case simulation studies. The first case study simulates the influence of microsite positions in relation to the local tree crown projections on the survival of spruce seedlings. It is assumed that the density of the trees is the main factor that determines the light and heat supply to the ground level of the Norway spruce seedlings. The second case study extends the previous study to include terms that determine the growth ratio in dependence on the crown projection area. The third case study provides further extensions in order to simulate growth ratio relations to the local soil type. The fourth case study demonstrates the local influence of the dominant grasses, such as Avenella flexuosa and Calamagrostis villosa, on the natural regeneration of Norway spruce. Starting from the conditions at the sites before the recovery period, the case simulation studies are able to project the short-term succession for a regeneration decade and the approximate long-term development. In addition to the standard simulation procedures based on solution of ordinary differential equations, spatio-temporal modelling in the GIS environment is able to provide spatial data management, analysis and visualization of the data.     

Non-linear ecological processes, fires, environmental heterogeneity and shrub invasion in northwestern Patagonia

The emergent behaviors of nature are not only the sum of interactions among ecosystem parts but also depend on the organization of these interactions. Fire, climate and vegetation patterns produce non-linear fire propagation across the landscape. Environmental heterogeneity, like outcrop presence and hare density, increases landscape patchiness and makes possible the occupation of fire refuges by plants, like Fabiana imbricata shrubs. We monitored shrub recruitment and cover during nine postfire years in northwestern Patagonia grasslands and we studied the long-term population dynamics under different environmental conditions through a matrix model, exploring different fire frequencies and spring precipitation regimes. Both, the field monitoring and the model seem to confirm the relationships between shrub invasion and fire. The climate change forecast predicts an increase in the frequency of El Niño Southern Oscillation phenomena that could causes more coupled fires—rainy springs in northwestern Patagonia, and consequently, more recruitment windows for shrubs, like F. imbricata. The matrix model also indicates that this scenario would be the most favourable for shrub invasion. Our results contribute to the knowledge of the ecosystem properties and processes, providing useful information to improve the grasslands sustainable use.     

GIS-based modeling of the geographic distribution of Quercus emoryi Torr. (Fagaceae) in México and identification of significant environmental factors influencing the species’ distribution

The greatest concentration of oak species in the world is believed to be found in Mexico. These species are potentially useful for reforestation because of their capacity to adapt to diverse environments. Knowledge of their geographic distribution and of species–environment relations is essential for decision-making in the management and conservation of natural resources. The objectives of this study were to develop a model of the distribution of Quercus emoryi Torr. in Mexico, using geographic information systems and data layers of climatic and other variables, and to determine the variables that significantly influence the distribution of the species. The study consisted of the following steps: (A) selection of the target species from a botanical scientific collection, (B) characterization of the collecting sites using images with values or categories of the variables, (C) model building with the overlay of images that meet the habitat conditions determined from the characterization of sites, (D) model validation with independent data in order to determine the precision of the model, (E) model calibration through adjustment of the intervals of some variables, and (F) sensitivity analysis using precision and concordance non-parametric statistics applied to pairs of images. Results show that the intervals of the variables that best describe the species’ habitat are the following: altitude from 1650 to 2750 amsl, slope from 0 to 66°; average minimum temperature of January from −12 to −3 °C; mean temperature of June from 11 to 25 °C; mean annual precipitation from 218 to 1225 mm; soil units: lithosol, eutric cambisol, haplic phaeozem, chromic luvisol, rendzina, luvic xerosol, mollic planosol, pellic vertisol, eutric regosol; type of vegetation: oak forest, oak–pine forest, pine forest, pine–oak forest, juniperus forest, low open forest, natural grassland and chaparral. The resulting model of the geographic distribution of Quercus emoryi in Mexico had the following values for non-parametric statistics of precision and agreement: Kappa index of 0.613 and 0.788, overall accuracy of 0.806 and 0.894, sensitivity of 0.650 and 0.825, specificity of 0.963, positive predictive value of 0.945 and 0.957 and negative predictive value of 0.733 and 0.846. Results indicate that the variable average minimum temperature of January, with a maximum value of −3 °C, is an important factor in limiting the species’ distribution.     

A multi-agent simulation to assess the risk of malaria re-emergence in southern France

A multi-agent simulation (MAS) was developed to assess the risk of malaria re-emergence in the Camargue in southern France, a non-endemic area where mosquitoes of the genus Anopheles (Culicidae) live. The contact rate between people and potential malaria vectors, or the human biting rate, is one of the key factor to predict the risk of re-emergence of malaria, would the parasite be introduced in the region. Our model (called MALCAM) represents the different agents that could influence malaria transmission in the Camargue – people, mosquitoes, animal hosts and the landscape – in a spatially explicit environment. The model simulates spatial and temporal variations in human biting rate at the landscape scale. These variations depend on the distribution of people and potential vectors, their behaviour and their interactions. A land use/cover map was used as a cellular-spatial support for the movements of and interactions between mobile agents. The model was tested for its sensitivity to variations in parameter values, and for the agreement between field observations and model predictions. The MALCAM model provides a tool to better understand the interactions between the multiple agents of the disease transmission system, and the land use and land cover factors that control the spatial heterogeneity in these interactions. It allows testing hypotheses and scenarios related to disease dynamics by varying the value of exogenous biological, geographical, or human factors. This application of agent-based modelling to a human vector-borne disease can be adapted to different diseases and regions.     

Volatile attractants for three Pteromalid parasitoids attacking concealed spruce bark beetles

Summary. The odour perceptive abilities, and preferences, of three bark beetle parasitoid species; Rhopalicus tutela (Walker), Roptrocerus mirus (Walker), and Roptrocerus xylophagorum (Ratzeburg) (Hymenoptera: Pteromalidae), were investigated to isolate and identify the essential compounds involved in host location. These parasitoids attack several economically important bark beetle species and oviposit preferentially on late larval stages concealed under the bark of conifers. Odours were collected from Norway spruce logs (Picea abies L. Karst.) containing Ips typographus L. (Coleoptera: Scolytidae) larvae. Biologically active compounds were isolated by coupled gas chromatographic-electroantennographic detection (GC-EAD), and identified by GC-mass spectrometry (GC-MS). Based on these analyses, four different synthetic baits were prepared and tested in a Y-tube walking bioassay. In the complex odour samples from spruce logs, only 16 compounds were EAD-active. The tested R. tutela and R. mirus females displayed similar trends in antennal activity to EAD-active compounds, responding mainly to oxygenated monoterpenes that indicate damaged conifers. Consequently, the synthetic baits were exclusively prepared with oxygenated monoterpenes. Parasitoid females (R. tutela and R. mirus) preferred spruce logs containing susceptible hosts over fresh logs, while male parasitoids (R. mirus) did not show any preference. However, when odours from fresh logs were mixed with synthetic baits (mimicking the odour composition of logs containing susceptible hosts), these combinations attracted female parasitoids (R. tutela, R. mirus, and R. xylophagorum). All synthetic baits seemed to be equally attractive to female parasitoids. Received 12 October 2000; accepted 18 January 2001     

Defoliation levels of oriental spruce by Ips typographus (L.) in relation to elevation and exposure

A study concerning the effects of elevation and exposure of the spruce forests on defoliation levels of oriental spruce (Picea orientalis (L.) Link.) by Ips typographus L. was carried out during 2005 and 2006 in Artvin-Hatila National Park, Turkey Nine spruce stands were selected at 3 zones of elevations (1000-1350 m, 1350-1700 m and 1700-2000 m) and at different aspects to assess the role of elevation and exposure in the crown defoliation level and body length of beetles. Influence of bark thickness and trunk diameter at 1.3 m on the damage caused by the pest was investigated as well. The results of the study were as follows: (1) The mean defoliation level was highest at 1700-2000 m following by 1350-1700 m and 1000-1350 m. (2) The highest defoliation levels occurred on southern slopes following by eastern and northern slopes at 1700-2000 m. (3) No statistical differences were found in the mean bark thickness between tree defoliation levels 1, 2, 3 and 4. (4) Mean trunk diameters of dead trees (level 4) were significantly greater than those with defoliation levels 0, 1 and 2. (5) Mean body length of I. typographus at upper zones was significantly higher than those at middle and lower zones.     

Monoterpene chemodiversity of ponderosa pine in relation to herbivory and bark beetle colonization

Summary. Ponderosa pine, Pinus ponderosa Laws. (Pinaceae), forests in Arizona have suffered from a nine-year period of drought and bark beetle, Ips lecontei Swaine (Coleoptera: Scolytidae), outbreaks. Abiotic and biotic stress in ponderosa pine results in the induced synthesis of certain monoterpenes that may in turn affect bark beetle behavior and survival. In this study, we investigate whether induced monoterpene production could result in a different monoterpene composition that remains stored in the needles or the trunk resin of the tree. Needle and resin samples in addition to trunk cores were collected from ponderosa pines at three locations in Arizona. Ungulate browsing induced a significant increase in limonene (P=0.010) and in chemodiversity (P=0.009), a measure of the evenness of distribution among the monoterpenes present in needles. We compared the level of ‘stress’ of the trees by measuring the thickness of annual rings in living trees and those that were killed by bark beetles. Where drought occurred, the spacing of annual rings from the last 10 years of trees killed by bark beetles was significantly smaller (P=0.020) compared to living trees. There was no difference in the monoterpene composition between the core sections of closest spacing of annual rings (stressed years) compared to the sections of widest spacing, which indicates that monoterpenes are distributed evenly throughout the extended resin system. In the area where the degree of drought was less overall, none of the individual monoterpenes present in the resin was related to bark beetle killed trees. However, about half the living pines had resin in which one of the major monoterpenes (α-pinene, Δ³-carene, and limonene) was absent, and these trees had a lower monoterpene chemodiversity compared to trees killed by bark beetles. Trees with these three major monoterpenes, corresponding to the average relative proportion in living pines at that location, may sustain higher selection and colonization by bark beetles.     

A resource-depletion model of forest insect outbreaks

Detailed analyses of thresholded ecological interactions can improve our understanding of the transition from aperiodic to periodic dynamics. We develop a threshold model of the population dynamics of outbreaking bark beetle populations that alternate between non-epidemic and epidemic behavior. The model involves accumulation of resources during low-density periods and depletion during outbreaks. The transition between the two regimes is caused by disturbance events in the form of major tree felling by wind. The model is analyzed with particular reference to the population dynamics of the spruce bark beetle (Ips typographus) in Scandinavia for which a comprehensive literature allows full parameterization. The fairly constant outbreak lengths and the highly variable waiting time between outbreaks that are seen in the historical records of this species agree well with the predictions of the model. The thresholded resource-depletion dynamics result in substantial variation in the degree of periodicity between stochastic realizations. The completely aperiodic tree colonizations are partly predictable when the timing of the irregular windfall events are known. However, the predictability of inter-outbreak periods is low due to the large variation of cases falling most frequently in the middle between the extremes of purely nonperiodic (erratic) and periodic (cyclic) fluctuations.     

Modelling the response of tree growth to temperature and CO2 elevation as related to the fertility and current temperature sum of a site

A methodology for simulating climate change impacts on tree growth was introduced into a statistical growth and yield model in relation to variations in site fertility and location implemented with current temperature sum. This was based on a procedure in which the relative enhancement in stem volume growth was calculated from short-term runs of a physiological simulation model for Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies (L.) Karst.) and silver birch (Betula pendula Roth.) stands. These simulations were made for a set of stands with species-specific variations in stand characteristics, location and fertility type first in current climatic conditions and then in different combinations of CO₂ and temperature elevations. Based on these simulations, the relative enhancement of volume growth induced by the climate change (relative scenario effect, RSEv) was calculated and modelled in relation to: (i) CO₂ and temperature elevation, stand density and the competition status of the tree in its stand, and (ii) variations in site fertility type and current temperature sum of a stand. Finally, these transfer functions for RSEv were applied to adapt the stem volume growth in the statistical growth and yield model to reflect the response to climate change.     

Evaluation of the forest growth simulator SILVA on dominant trees in mature mixed Silver fir–Norway spruce stands in South-West Germany

Forest growth simulators go beyond a mere tabulation of empirical measurements by employing biometric models that functionally describe the dependence of forest growth of the initial forest structure, growth conditions and management regime. This makes them very flexible and allows predicting growth reactions for unknown and/or complex forest growth scenarios. When simulation outcomes are to be used in silvicultural strategic planning, the results are of direct and delicate importance, and the correct simulator performance must be ascertained. This is especially so when the considered forest situation differs from the forest data used to parameterise the model (e.g. different geographical region).In this article, the forest growth simulator SILVA (version 2.2) was validated for 55 long-term experimental plots of mature mixed Silver fir–Norway spruce stands in southwest Germany (Picea abies, Abies alba). The evaluation was restricted to the upper canopy trees during the survey period 1989–2004. Following the general evaluation criteria for ecological models from [Vanclay, J.K., Skovsgaard, J.P., 1997. Evaluating forest growth models. Ecol. Mod. 98, 1–12], a specific methodology was developed to evaluate the simulated height and diameter growth on the basis of forest growth principles.The qualitative analysis proved the SILVA growth algorithms to be in accordance with physiologically based standard growth equations. The quantitative evaluation was limited by incomplete knowledge of the site conditions. To overcome this problem, the experimental plots were regarded as a “heterogeneous growth series” which allows analysing the growth behaviour in a more general way. It could be shown that for the given data set, the SILVA simulations produced an overestimation of height growth (median: +61% spruce, +12% fir), and an underestimation of diameter growth and competition sensitivity (median: ?16% spruce, ?70% fir). The errors partially compensated in the volume growth resulting in an overall over-/underestimation of +9% for spruce and ?58% for fir (median).The unbalanced height and diameter growth cannot be compensated by a change in the site conditions because this affects both height and diameter growth either positive or negative. Hence, an adjustment of selected parameterisation values appears to offer the best solution to adapt SILVA to the considered forest situation. This approach of adaptive parameterisation is discussed against a more general background of deductive vs. inductive forest growth modelling.     

Simulating regeneration and vegetation dynamics in Mediterranean coniferous forests

This study aims to provide a quantitative framework to model the dynamics of Mediterranean coniferous forests by integrating existing ecological data within a generic mathematical simulator. We developed an individual-based vegetation dynamics model, constrained on long-term field regeneration data, analyses of tree-rings and seed germination experiments. The simulator implements an asymmetric competition algorithm which is based on the location and size of each individual. Growth is parameterized through the analysis of tree-rings from more than thirty individuals of each of the three species of interest. A super-individual approach is implemented to simulate regeneration dynamics, constrained with available regeneration data across time-since-disturbance and light-availability gradients. The study concerns an insular population of an endemic to Greece Mediterranean fir (Abies cephalonica Loudon) on the island of Cephalonia (Ionian Sea) and two interacting populations of a Mediterranean pine (Pinus brutia Ten.) and a more temperate-oriented pine (Pinus nigra Arn. ssp. pallasiana) on the island of Lesbos (NE Aegean Sea), Greece. The model was validated against plot-level observations in terms of species standing biomass and regeneration vigour and adequately captured regeneration patterns and overall vegetation dynamics in both study sites. The potential effects of changing climatic patterns on the regeneration dynamics of the three species of interest were subsequently explored. With the assumption that a warmer future would probably cause changes in the duration of cold days, we tested how this change would affect the overall dynamics of the study sites, by focusing on the process of cold stratification upon seed germination. Following scenarios of a warmer future and under the current model parameterization, changes in the overall regeneration vigour controlled by a reduction in the amount of cold days, did not alter the overall dynamics in all plant populations studied. No changes were identified in the relative dominance of the interacting pine populations on Lesbos, while the observed reduction in the amount of emerging seedlings of A. cephalonica on Cephalonia did not affect biomass yield at later stages of stand development.     

Assessing the performance of theoretical and empirical tree mortality models using tree-ring series of Norway spruce

Successional dynamics of forests under current and changed climate are often investigated using gap models, a subset of forest succession models that simulate establishment, growth, and mortality of trees. However, the mortality submodels of gap models are largely based on theoretical assumptions, and have not been tested in detail.In the present study, we compared the performance of a range of theoretical mortality functions (TMFs) that are commonly used in gap models with several empirical mortality functions (EMFs) that were derived using logistic regression from growth patterns of tree-ring series as predictor variables. Data from dead and living Norway spruce (Picea abies (L.) Karst.) trees from subalpine forests at three study sites in Switzerland were used to this end.Three of the four EMFs consistently performed better at all three sites, while three of the four TMFs performed worse than the remaining mortality functions. At one site, these three EMFs correctly classified 71–78% of the dead trees (48–72% for the three TMFs) and 73% (49–64%) of the living trees. 44–54% (21–25%) of the dead trees were predicted to die within 15 years prior to death. 0–2% (7–10%) of the dead trees and 5% (19–31%) of the living trees were predicted to die more than 60 years prior to the last measured year.We conclude that, unless the parameters of the TMFs are optimized for individual species, the TMFs are not appropriate to predict the time of tree death, in spite of their widespread use. A substantial change in simulated forest succession is to be expected if the currently implemented TMFs in gap models are replaced by species-specific EMFs.     

Future distribution modelling: A stitch in time is not enough

The last two decades have seen an increasing number of studies assessing the impact of climate change upon biodiversity. A central assumption underpinning research into the potential future habitat of terrestrial biota is that species are presently in equilibrium with their environments and that quantitative climate models adequately represent the distribution of species. Recently, many alarming predictions have emerged concerning the extinction and redistribution of species. Here, we show that even large-scale models of the climatic niche dimensions of species are temporally variable. Distributional models were developed for Salix (willow) species occurring in the province of Ontario, Canada, using three historical climate data sets. Although historical data very accurately represented the distributions of willows, the inherent variability within the models of species based on different periods greatly influenced the direction and magnitude of projected distributional change. We expose a fundamental uncertainty with respect to predicting the responses of species to climate change.