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.     

Forest dynamics in Oregon landscapes: evaluation and application of an individual-based model.

The FORCLIM model of forest dynamics was tested against field survey data for its ability to simulate basal area and composition of old forests across broad climatic gradients in western Oregon, USA. The model was also tested for its ability to capture successional trends in ecoregions of the west Cascade Range. It was then applied to simulate present and future (1990-2050) forest landscape dynamics of a watershed in the west Cascades. Various regimes of climate change and harvesting in the watershed were considered in the landscape application. The model was able to capture much of the variation in forest basal area and composition in western Oregon even though temperature and precipitation were the only inputs that were varied among simulated sites. The measured decline in total basal area from tall coastal forests eastward to interior steppe was matched by simulations. Changes in simulated forest dominants also approximated those in the actual data. Simulated abundances of a few minor species did not match actual abundances, however. Subsequent projections of climate change and harvest effects in a west Cascades landscape indicated no change in forest dominance as of 2050. Yet, climate-driven shifts in the distributions of some species were projected. The simulation of both stand-replacing and partial-stand disturbances across western Oregon improved agreement between simulated and actual data. Simulations with fire as an agent of partial disturbance suggested that frequent fires of low severity can alter forest composition and structure as much or more than severe fires at historic frequencies.     

Modeling Dynamics of Habitat-Interior Bird Populations in Fragmented Landscapes

Abstract: A stochastic computer model was used to examine the effects of varying degrees of habitat fragmentation on the dynamics of a hypothetical population of forest-interior bid. The primary demographic parameter that influenced the population's dynamics was fecundity, which varied as a function of how far a birds territory was from an ecological edge. As our model landscape became more fragmented the proportion of forest habitat that was near edges increased geometrically, and the population's overall fecundity dropped as a result. The model demonstrates that impaired reproduction in a fragmented landscape is, by itself a sufficient disruption of the population's dynamics to generate population declines and shifts in distribution similar to those observed in the fragmented forests of southern Wisconsin. Without immigration of recruits from other regions where reproduction is better, habitat-interior populations in a severely fragmented landscape can become locally extinct.     

Effects of Livestock Grazing on Stand Dynamics and Soils in Upland Forests of the Interior West

Many ponderosa pine and mixed-conifer forests of the western, interior United States have undergone substantial structural and compositional changes since settlement of the West by Euro-Americans. Historically, these forests consisted of widely spaced, fire-tolerant trees underlain by dense grass swards. Over the last 100 years they have developed into dense stands consisting of more fire-sensitive and disease-susceptible species. These changes, sometimes referred to as a decline in "forest health," have been attributed primarily to two factors: active suppression of low-intensity fires (which formerly reduced tree recruitment, especially of fire-sensitive, shade-tolerant species), and selective logging of larger, more fire-tolerant trees. A third factor, livestock grazing, is seldom discussed, although it may be as important as the other two factors. Livestock alter forest dynamics by (1) reducing the biomass and density of understory grasses and sedges, which otherwise outcompete conifer seedlings and prevent dense tree recruitment, and (2) reducing the abundance of fine fuels, which formerly carried low-intensity fires through forests. Grazing by domestic livestock has thereby contributed to increasingly dense western forests and to changes in tree species composition. In addition, exclosure studies have shown that livestock alter ecosystem processes by reducing the cover of herbaceous plants and litter, disturbing and compacting soils, reducing water infiltration rates, and increasing soil erosion.     

Simulation of interannual responses of trembling aspen stands to climatic variation and insect defoliation in western Canada

A carbon-based model has been developed to simulate responses of trembling aspen (Populus tremuloides Michx.) stands to interannual climatic variation and insect defoliation. The model is designed for medium time scale (10–100 years) simulations and requires only daily maximum and minimum temperature and precipitation as meteorological inputs. The modelling approach is similar to FOREST-BGC but includes additional processes known to be important in deciduous forests. These include removal of leaf area during outbreaks of forest tent caterpillar (Malacosoma disstria Hbn.), phenological changes in leaf area index, storage and allocation of non-structural carbohydrate and the contribution of understorey vegetation to evapotranspiration. The model was used for simulations of growth and mortality of biomass carbon in two mature aspen forests located in the climatically dry transition zone between the boreal forest and prairie grassland regions of Saskatchewan, Canada. Model inputs of annual defoliation intensity were based on historic records of insect defoliation and the incidence of light-coloured tree rings in disks or cores collected from aspen at each of the two sites. At both sites, moderately good correlations (r²=0.47–0.54) were obtained between modelled interannual changes in stem carbon growth and observed interannual changes in stem basal area increment obtained from tree-ring analysis. Model outputs of stem biomass carbon were found to be highly sensitive to parameters describing seasonal leaf area duration, insect defoliation intensity, photosynthesis and root respiration and carbohydrate allocation to growth versus storage.     

Comparing modern and presettlement forest dynamics of a subboreal wilderness: does spruce budworm enhance fire risk?

Insect disturbance is often thought to increase fire risk through enhanced fuel loadings, particularly in coniferous forest ecosystems. Yet insect disturbances also affect successional pathways and landscape structure that interact with fire disturbances (and vice-versa) over longer time scales. We applied a landscape succession and disturbance model (LANDIS-II) to evaluate the relative strength of interactions between spruce budworm (Choristoneura fumiferana) outbreaks and fire disturbances in the Boundary Waters Canoe Area (BWCA) in northern Minnesota (USA). Disturbance interactions were evaluated for two different scenarios: presettlement forests and fire regimes vs. contemporary forests and fire regimes. Forest composition under the contemporary scenario trended toward mixtures of deciduous species (primarily Betula papyrifera and Populus spp.) and shade-tolerant conifers (Picea mariana, Abies balsamea, Thuja occidentalis), with disturbances dominated by a combination of budworm defoliation and high-severity fires. The presettlement scenario retained comparatively more "big pines" (i.e., Pinus strobus, P. resinosa) and tamarack (L. laricina), and experienced less budworm disturbance and a comparatively less-severe fire regime. Spruce budworm disturbance decreased area burned and fire severity under both scenarios when averaged across the entire 300-year simulations. Contrary to past research, area burned and fire severity during outbreak decades were each similar to that observed in non-outbreak decades. Our analyses suggest budworm disturbances within forests of the BWCA have a comparatively weak effect on long-term forest composition due to a combination of characteristics. These include strict host specificity, fine-scaled patchiness created by defoliation damage, and advance regeneration of its primary host, balsam fir (A. balsamea) that allows its host to persist despite repeated disturbances. Understanding the nature of the three-way interaction between budworm, fire, and composition has important ramifications for both fire mitigation strategies and ecosystem restoration initiatives. We conclude that budworm disturbance can partially mitigate long-term future fire risk by periodically reducing live ladder fuel within the mixed forest types of the BWCA but will do little to reverse the compositional trends caused in part by reduced fire rotations.     

Modelling the spatial population dynamics of the green oak leaf roller (Tortrix viridana) using density dependent competitive interactions: Effects of herbivore mortality and varying host-plant quality

Cyclic population dynamics of forest insects with periods of more than two generations have been discussed in relation to a variety of extrinsic and intrinsic forces. In the present study, we employed the selection pressure of density dependent competitive interactions according to Witting's equations (Witting, 2000) as driver for a discrete spatiotemporal model of the green oak leaf roller (Tortrix viridana). The model was successfully parameterised to rebuild the cyclic population dynamics of an empirical data set of a 30-year leaf roller monitoring in Russia. Our analysis focussed on the role of herbivore mortality and host plant food quality, which have a significant effect on T. viridana population dynamics. An additional egg or larvae mortality lowers population density and can lead to selection pressures that favour individuals with higher growth rate. This increased population growth rate can not only compensate the additional mortality, but also can lead to higher average moth abundances in subsequent generations. Furthermore, we analysed the effect of inter- and intraspecific variation in host plant quality on herbivore population dynamics and the spatial distribution of abundance and defoliation patterns. We found significant effects of the qualitative composition of a trees neighbourhood on the herbivore population of the respective tree. Also, the patchy damage patterns observable in reality have been reproduced by the present model. The applicability of the model approach and the putative genetic processes underlying Witting's model are discussed.     

Evaluation of discrete host–parasitoid models for diamondback moth and Diadegma semiclausum field time population density series

The performance of discrete mathematical models to describe the population dynamics of diamondback moth (DBM) (Plutella xylostella L.) and its parasitoid Diadegma semiclausum was investigated. The parameter values for several well-known models (Nicholson–Bailey, Hassell and Varley, Beddington, Free and Lawton, May, Holling type 2, 3 and Getz and Mills functional responses) were estimated. The models were tested on 20 consecutive sets of time series data collected at 14 days interval for pest and parasitoid populations obtained from a highland cabbage growing area in eastern Kenya. Model parameters were estimated from minimized squared difference between the numerical solution of the model equations and the empirical data using Powell's method. Maximum calculated DBM growth rates varied between 0.02 and 0.07. The carrying capacity determined at 16.5 DBM/plant by the Beddington et al. model was within the range of field data. However, all the estimated parameter values relating to the parasitoid, including the instantaneous searching rate (0.07–0.28), per capita searching efficiency (0.20–0.27), search time (5.20–5.33), handling time (0.77–0.90), and parasitism aggregation index (0.33), were well outside the range encountered empirically. All models evaluated for DBM under Durbin–Watson criteria, except the May model, were not autocorrelated with respect to residuals. In contrast, the criteria applied to the parasitoid residuals showed strong autocorrelations. Thus, these models failed to estimate parasitoid dynamics. We conclude that the interactions of the DBM with its parasitoid cannot be explained by any of the models tested. Two factors may be associated with this failure. First, the parasitoid in this integrated biological control system may not be playing a major role in regulating DBM population. Second, and perhaps more likely, poor correlations reflect gross inadequacies in the theoretical assumptions that underlie the existing models.     

Effects of Forest Fragmentation and Habitat Degradation on West African Leaf-Litter Frogs

Abstract:  Habitat degradation alters the dynamics and composition of anuran assemblages in tropical forests. The effects of forest fragmentation on the composition of anuran assemblages are so far poorly known. We studied the joint influence of forest fragmentation and degradation on leaf-litter frogs. We specifically asked whether the processes structuring leaf-litter anuran assemblages in fragmented forests are the same as those in continuous forests. We analyzed anuran assemblages with respect to habitat characteristics, including fragmentation and degradation parameters. In comparison with continuous forests, species richness and diversity were lower and assemblage composition was altered in forest fragments. These changes seemed to be mainly caused by habitat degradation rather than forest fragmentation. Availability of aquatic sites for breeding, vegetation structure (including those variables indicating degradation), and leaf-litter cover had the most influence on the presence of single species. The comparatively small impact of fragmentation on anurans might be due to the location of the study area; it still possessed large tracts of continuous forest. These forest blocks may stabilize the regional rainforest climate and thus weaken the effects of fragmentation .     

桂西北喀斯特区原生林与次生林凋落物及养分归还特征比较

森林凋落物是森林生态系统的重要组成成分,其养分归还量在一定程度上决定着土壤养分有效性的高低。在土层浅薄且土被很不连续的我国喀斯特区域进行凋落物生物量及养分归还研究对我们更深刻地了解该区养分循环具有至关重要的意义。本文比较分析了桂西北喀斯特区3种原生林与3种次生林的全年凋落物量、组成、月凋落物量动态及养分归还量与动态。结果表明,圆果化香(Platycarya longipes Wu)、大叶蚊母树(Distylium Sieb.et Zucc.)与青檀(Pteroceltis tatarinowii Maxim.)3种原生林的年凋落物总量分别为2342.16,4057.99和1834.36kg·hm-2,而圆叶乌桕(Sapium rotundifolium Hemsl.)、八角枫(Alangium chinense(Lour.) Harms)和黄荆(Vitex negundo L.)3种次生林的年凋落物总量分别为3192.82,3284.26,2469.90kg·hm-2,除大叶蚊母树外,次生林年凋落物总量大于原生林。凋落物的组成中,叶凋落生物量均占总凋落物量的80%左右,甚至更高,而圆叶乌桕、八角枫和黄荆3种次生林群落的叶凋落物量占总凋落物量的百分比大于圆果化香、大叶蚊母树以及青檀3种原生林。凋落物的养分归还量的月动态与凋落物量的月动态一致,原生林呈"U"形曲线,而次生林则呈"W"形曲线。原生林和次生林凋落物的年养分归还量均为C﹥N﹥K﹥P,且次生林的C、N、P养分的归还量大于原生林。     

Forest dynamics after successive spruce budworm outbreaks in mixedwood forests

In order to assess the long-term spatiotemporal influence of the spruce budworm in sub-boreal mixedwood forests, we studied the effect of three successive outbreaks in a region of western Quebec, Canada. We used dendrochronology to detect past outbreaks in three areas (111-185 ha), based on the recruitment age of balsam fir (Abies balsamea) and on growth patterns of white spruce (Picea glauca), the two main host species of this defoliating insect. We also used a series of aerial photographs taken between 1935 and 2003 to evaluate overstory mortality and post-outbreak succession patterns in these same areas. Individual outbreaks had a spatially homogenous impact on host species throughout the region, but successive outbreaks differed in intensity: the two outbreaks around 1910 and 1980 caused widespread mortality in the overstory, but an outbreak around 1945 had little impact, probably because the forest mosaic had not yet recuperated from the 1910 outbreak. No clear outbreak was detected in the later part of the 19th century. In portions of the study areas where the 1910 outbreak had a major impact, between 36% and 50% of the stands were reoccupied by balsam fir stands in the period up to the 1980 outbreak (cyclic succession), the rest being at least partly replaced by nonhost species such as Betula spp. Changes in forest composition after the 1910 outbreak were mostly associated with upper-slope positions in all study areas. The 1980 outbreak also had a higher impact than earlier outbreaks in lower-slope positions dominated by black spruce (Picea mariana)-balsam fir mixtures. These results suggest that, at the regional scale, the abundance of mature or over-mature balsam fir stands does not determine the outbreak cycle. When an outbreak occurs, however, its impact will be strongly constrained by forest characteristics such as stand composition and structure, which are themselves influenced by previous disturbances and slope position.     

Ecology of Pacific Yew (Taxus brevifolia) in Western Oregon and Washington

Taxus brevifolia , a subcanopy tree or shrub in forests of the Pacific Northwest, has been harvested intensively in recent years. With management concerns as an impetus, we examined the distribution and population dynamics of Taxus based on data from the mountains of western Oregon and Washington. Surveys of natural forests, long-term studies of forest recovery following logging, and census data on marked trees in forest stands support the hypothesis that Taxus is a widespread but predominantly late-successional species. Sensitive to fire and slow to recover from disturbance on many sites, Taxus attains maximal basal area and adult stem density in old forests. Colonization of Taxus is often slow in potentially suitable habitats. Conservation of Taxus at the landscape level may require large, unmanaged reserves and maintenance of patches of old forest within managed forests. Long rotations (centuries) between harvest events will enhance the long-term viability of the species. Practices designed to accelerate the development of old-growth forest structure will not benefit Taxus and other species requiring long disturbance-free intervals for recovery.     

酸雨对外来植物入侵的影响

酸雨和外来种入侵都是全球关注的问题。结合外来入侵植物的生态适应特性以及酸雨的危害特征,系统分析了酸雨对外来植物入侵产生的影响。酸雨对外来植物入侵的影响是复杂多样的。酸雨导致群落冠层稀疏,群落透光率增加,加之氮沉降后土壤、水体氮素的增加,有利于生长力强的外来喜阳植物入侵;酸雨加速土壤酸化,促使基本离子淋失以及A1毒等危害植物的生长发育,植物的内源激素以及化感作用发生改变,适应力和耐受力强的外来植物在与本地植物竞争中处于相对优势而成为入侵种;酸雨以及外来植物入侵改变了土壤微生物群落结构,影响本地植物的生长而促使外来植物的入侵。     

Simulating the potential for ecological restoration of dryland forests in Mexico under different disturbance regimes

Examining the potential for ecological restoration is important in areas where anthropogenic disturbance has degraded forest landscapes. However, the conditions under which restoration of degraded tropical dry forests (TDF) might be achieved in practice have not been determined in detail. In this study, we used LANDIS-II, a spatially explicit model of forest dynamics, to assess the potential for passive restoration of TDF through natural regeneration. The model was applied to two Mexican landscapes under six different disturbance regimes, focusing on the impact of fire and cattle grazing on forest cover, structure and composition. Model results identified two main findings. First, tropical dry forests are more resilient to anthropogenic disturbance than expected. Results suggested that under both a scenario of small, infrequent fires and a scenario of large, frequent fires, forest area can increase relatively rapidly. However, forest structure and composition differed markedly between these scenarios. After 400 years, the landscape becomes increasingly occupied by relatively shade-tolerant species under small, infrequent fires, but only species with both relatively high shade tolerance and high fire tolerance can thrive under conditions with large, frequent fires. Second, we demonstrated that different forms of disturbance can interact in unexpected ways. Our projections revealed that when grazing acts in combination with fire, forest cover, structure and composition vary dramatically depending on the frequency and extent of the fires. Results indicated that grazing and fire have a synergistic effect causing a reduction in forest cover greater than the sum of their individual effects. This suggests that passive landscape-scale restoration of TDF is achievable in both Mexican study areas only if grazing is reduced, and fires are carefully managed to reduce their frequency and intensity.     

Design and validation of a spatially explicit simulation model for bottomland hardwood forests

Species composition in forests depends on the interaction of species traits and species availability. Yet many forest simulation models focus only on interactions of adult trees and saplings, ignoring how species become members of the community. We modify a published forest model for bottomland hardwood forests (program SWAMP [Phipps, R.L., 1979. Simulation of wetlands forest vegetation dynamics. Ecol. Modell. 7, 257–288]) to make it spatially explicit and incorporate explicit seed production and dispersal algorithms. The resulting individual-based, spatially explicit forest simulator (YAFSIM) combines mechanistic seed dispersal with growth and mortality of trees to track forest dynamics over time. We describe the structure of the model and test its validity for dynamics in small bottomland hardwood patches in the Mississippi Alluvial Valley. Dynamics of species composition and basal areas of trees predicted by Yazoo Forest Simulator (YAFSIM) were similar to those of natural second- and old-growth bottomland forests. However, diversity of simulated forest patches declined over time largely because of random dynamics acting on small, isolated populations.     

Biodiversity Differences between Managed and Unmanaged Forests: Meta-Analysis of Species Richness in Europe

Abstract: Past and present pressures on forest resources have led to a drastic decrease in the surface area of unmanaged forests in Europe. Changes in forest structure, composition, and dynamics inevitably lead to changes in the biodiversity of forest‐dwelling species. The possible biodiversity gains and losses due to forest management (i.e., anthropogenic pressures related to direct forest resource use), however, have never been assessed at a pan‐European scale. We used meta‐analysis to review 49 published papers containing 120 individual comparisons of species richness between unmanaged and managed forests throughout Europe. We explored the response of different taxonomic groups and the variability of their response with respect to time since abandonment and intensity of forest management. Species richness was slightly higher in unmanaged than in managed forests. Species dependent on forest cover continuity, deadwood, and large trees (bryophytes, lichens, fungi, saproxylic beetles) and carabids were negatively affected by forest management. In contrast, vascular plant species were favored. The response for birds was heterogeneous and probably depended more on factors such as landscape patterns. The global difference in species richness between unmanaged and managed forests increased with time since abandonment and indicated a gradual recovery of biodiversity. Clearcut forests in which the composition of tree species changed had the strongest effect on species richness, but the effects of different types of management on taxa could not be assessed in a robust way because of low numbers of replications in the management‐intensity classes. Our results show that some taxa are more affected by forestry than others, but there is a need for research into poorly studied species groups in Europe and in particular locations. Our meta‐analysis supports the need for a coordinated European research network to study and monitor the biodiversity of different taxa in managed and unmanaged forests.     

Microsite-limited recruitment controls fern colonization of post-agricultural forests

Assessing the relative roles of dispersal limitation and environmental effects in population dynamics and community assembly is fundamental to understanding patterns of species distribution and diversity. In forests growing on abandoned agricultural lands, both legacies of vegetation disturbance and changes in the abiotic environment shape the diversity and composition of recovering communities. Here I specify how interactions among historical, environmental, and biological factors influence species distributions, focusing on three fern species with contrasting distributions across forests of different history in central New York, USA: Dryopteris carthusiana, Dryopteris intermedia, and Polystichum acrostichoides. Using population surveys, spore-trap and spore-bank studies, and a three-year field experiment, I compare demographic rates among species and between forest types to determine which life history stages limit colonization and which traits explain species distributions. Adult plants of all three species were larger and more likely to produce spores in post-agricultural forests than in adjacent, uncleared stands. Though lower population densities led to fewer spores in post-agricultural soils, spore availability still exceeded recruitment by four to five orders of magnitude. Sowing additional spores had relatively little effect, while microhabitat conditions had the greatest impact on establishment rates. Given similar microsites, the two forest types had equal rates of establishment, but some forest-floor features preferentially occupied by juvenile plants were less frequent in post-agricultural stands. The availability of suitable sites for establishment, created by small-scale heterogeneity on forest floors, thus limits both the growth of fern populations and the colonization of new habitats. In fact, reduced microtopographic variation in post-agricultural forests may represent a greater hindrance to plant establishment than changes in mean environmental conditions. Among the three fern species, establishment rates differed as species distributions would predict, with the strongest colonizer consistently having the highest rates and the slowest colonizer the lowest. Rather than random or trait-mediated dispersal, the different distributions of these species reflect life history traits that determine establishment rates and thus colonization ability. This case study demonstrates that ecological interactions based on the unique life histories of individual species can override dispersal in determining species distributions.     

Modelling the impact of defoliation and leaf damage on forest plantation function and production

After presenting a short review of process-based model requirements to capture the plant dynamic response to defoliation, this paper describes the development and testing of a model of crown damage and defoliation for Eucalyptus. A model that calculates light interception and photosynthetic production for canopies that vary spatially and temporally in leaf area and photosynthetic properties is linked to the forest growth model CABALA. The process of photosynthetic up-regulation following defoliation is modelled with a simple conditional switch that triggers up-regulation when foliar damage or removal causes the ratio of functional leaf area to living tissue in the tree to change.We show that the model predicts satisfactorily when validated with trees of Eucalyptus nitens and Eucalyptus globulus from a range of sites of different ages, subject to different types of stress and different types of defoliation events (R² = 0.96 across a range of sites). However, the complexity of particular situations can cause the model to fail (e.g. very heavy defoliation events where branch death occurs).It is concluded that while the model will not cope with all situations, an appropriate level of generality has been captured to represent many of the physiological processes and feedbacks that occur following defoliation or leaf damage. This makes the model useful for guiding management interventions following pest attack and allows the development of scenarios including climate change impact analyses and decision-making on the merits of post-defoliation fertilisation to expedite recovery.     

The Potential for Species Conservation in Tropical Secondary Forests

Abstract: In the wake of widespread loss of old‐growth forests throughout the tropics, secondary forests will likely play a growing role in the conservation of forest biodiversity. We considered a complex hierarchy of factors that interact in space and time to determine the conservation potential of tropical secondary forests. Beyond the characteristics of local forest patches, spatial and temporal landscape dynamics influence the establishment, species composition, and persistence of secondary forests. Prospects for conservation of old‐growth species in secondary forests are maximized in regions where the ratio of secondary to old‐growth forest area is relatively low, older secondary forests have persisted, anthropogenic disturbance after abandonment is relatively low, seed‐dispersing fauna are present, and old‐growth forests are close to abandoned sites. The conservation value of a secondary forest is expected to increase over time, as species arriving from remaining old‐growth forest patches accumulate. Many studies are poorly replicated, which limits robust assessments of the number and abundance of old‐growth species present in secondary forests. Older secondary forests are not often studied and few long‐term studies are conducted in secondary forests. Available data indicate that both old‐growth and second‐growth forests are important to the persistence of forest species in tropical, human‐modified landscapes.     

Carbon dynamics of forests in Washington, U.S.A.: 21st century projections based on climate-driven changes in fire regimes

During the 21st century, climate-driven changes in fire regimes will be a key agent of change in forests of the U.S. Pacific Northwest (PNW). Understanding the response of forest carbon (C) dynamics to increases in fire will help quantify limits on the contribution of forest C storage to climate change mitigation and prioritize forest types for monitoring C storage and fire management to minimize C loss. In this study, we used projections of 21st century area burned to explore the consequences of changes in fire regimes on C dynamics in forests of Washington State. We used a novel empirical approach that takes advantage of chronosequences of C pools and fluxes and statistical properties of fire regimes to explore the effects of shifting age class distributions on C dynamics. Forests of the western Cascades are projected to be more sensitive to climate-driven increases in fire, and thus projected changes in C dynamics, than forests of the eastern Cascades. In the western Cascades, mean live biomass C is projected to decrease by 24-37%, and coarse woody debris (CWD) biomass C by 15-25% for the 2040s. Loss of live biomass C is projected to be lower for forests of the eastern Cascades and Okanogan Highlands (17-26%), and CWD biomass is projected to increase. Landscape mean net primary productivity is projected to increase in wet low-elevation forests of the western Cascades, but decrease elsewhere. These forests, and moist forests of the Okanogan Highlands, are projected to have the greatest percentage increases in consumption of live biomass. Percentage increases in consumption of CWD biomass are greater than 50% for all regions and up to four times greater than increases in consumption of live biomass. Carbon sequestration in PNW forests will be highly sensitive to increases in fire, suggesting a cautious approach to managing these forests for C sequestration to mitigate anthropogenic CO2 emissions.