Bat Use of Remnant Old-Growth Redwood Stands

Most of the old-growth redwood ( Sequoia sempervirens ) in California has been cut; regenerating forests will probably never resemble those that were harvested, and what old growth remains on private land occurs in small, isolated remnant patches. The landscapes in which these stands occur differ so markedly from their original condition that their value as habitat to many species of wildlife, including bats, is unknown. Previous research in unfragmented redwood forests demonstrated that bats use basal hollows in old-growth redwoods as roosts. We sought to determine whether bats use similar old-growth trees as roosts when they occur in small, remnant patches of isolated old growth on commercial forest land. We compared bat occurrence in remnant and contiguous stands by collecting guano in traps suspended in hollows and by monitoring flight activity with ultrasonic bat detectors. Hollows in trees within the remnant stands had significantly more guano deposited per tree than the trees within the contiguous forest. The mean numbers of bat passes per night were statistically indistinguishable between the two treatments, although mean flight activity in the remnant stands was greater than in the contiguous forest. Bats frequently used basal hollows in small (<5 ha) stands of remnant old growth, which may be due to the closer proximity of remnants to stream courses, to their greater interface with edge where foraging success may be greater, or to the fact that the lower density of hollow-bearing trees in remnants than in contiguous forest favored greater use per tree. Significant use of small, residual old-growth redwood provides reason to maintain these remnants in managed landscapes as potentially important habitat for forest bats.     

Potential site productivity influences the rate of forest structural development

Development and maintenance of structurally complex forests in landscapes formerly managed for timber production is an increasingly common management objective. It has been postulated that the rate of forest structural development increases with site productivity. We tested this hypothesis for Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forests using a network of permanent study plots established following complete timber harvest of the original old-growth forests. Forest structural development was assessed by comparing empirical measures of live tree structure to published values for Douglas-fir forests spanning a range of ages and structural conditions. The rate of forest structural development--resilience--exhibited a positive relationship with site index, a measure of potential site productivity. Density of shade-intolerant conifers declined in all study stands from an initial range of 336-4068 trees/ha to a range of 168-642 trees/ha at the most recent measurement. Angiosperm tree species declined from an initial range of 40-371 trees/ha to zero in seven of the nine plots in which they were present. Trends in shade-tolerant tree density were complex: density ranged from 0 to 575 trees/ha at the first measurement and was still highly variable (25-389 trees/ha) at the most recent measurement. Multivariate analysis identified the abundance of hardwood tree species as the strongest compositional trend apparent over the study period. However, structural variables showed a strong positive association with increasing shade-tolerant basal area and little or no association with abundance of hardwood species. Thus, while tree species succession and forest structural development occur contemporaneously, they are not equivalent processes, and their respective rates are not necessarily linearly related. The results of this study support the idea that silvicultural treatments to accelerate forest structural development should be concentrated on lower productivity sites when the management objective is reserve-wide coverage of structurally complex forests. Alternatively, high-productivity sites should be prioritized for restoration treatments when the management objective is to develop structurally complex forests on a portion of the landscape.     

Spatially explicit modeling of overstory manipulations in young forests: Effects on stand structure and light

Young forests can be manipulated in diverse ways to enhance their ecological values. We used stem maps from two dense, second-growth stands in western Washington and a spatially explicit light model (tRAYci) to simulate effects of five silvicultural manipulations on diameter distribution, species composition, spatial patterning, and light availability. Each treatment removed 30% of the basal area, but differed in how trees were selected for removal. Three primary treatments were thin from below (removing the smallest trees), random thin (removing trees randomly), and gap creation (removing all trees in circles ∼1 tree height in diameter). Two additional treatments combined elements of these approaches: random ecological thin (a mixture of thin from below and random thin) and structured ecological thin (a mixture of thin from below and gap creation).     

Importance of conserving large and old trees to continuity of tree-related microhabitats

Protecting structural features, such as tree-related microhabitats (TreMs), is a cost-effective tool crucial for biodiversity conservation applicable to large forested landscapes. Although the development of TreMs is influenced by tree diameter, species, and vitality, the relationships between tree age and TreM profile remain poorly understood. Using a tree-ring-based approach and a large data set of 8038 trees, we modeled the effects of tree age, diameter, and site characteristics on TreM richness and occurrence across some of the most intact primary temperate forests in Europe, including mixed beech and spruce forests. We observed an overall increase in TreM richness on old and large trees in both forest types. The occurrence of specific TreM groups was variably related to tree age and diameter, but some TreM groups (e.g., epiphytes) had a stronger positive relationship with tree species and elevation. Although many TreM groups were positively associated with tree age and diameter, only two TreM groups in spruce stands reacted exclusively to tree age (insect galleries and exposed sapwood) without responding to diameter. Thus, the retention of trees for conservation purposes based on tree diameter appears to be a generally feasible approach with a rather low risk of underrepresentation of TreMs. Because greater tree age and diameter positively affected TreM development, placing a greater emphasis on conserving large trees and allowing them to reach older ages, for example, through the establishment of conservation reserves, would better maintain the continuity of TreM resource and associated biodiversity. However, this approach may be difficult due to the widespread intensification of forest management and global climate change.     

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.     

Quantifying Habitat Requirements of Tree-Living Species in Fragmented Boreal Forests with Bayesian Methods

Abstract:  Quantitative conservation objectives require detailed consideration of the habitat requirements of target species. Tree-living bryophytes, lichens, and fungi are a critical and declining biodiversity component of boreal forests. To understand their requirements, Bayesian methods were used to analyze the relationships between the occurrence of individual species and habitat factors at the tree and the stand scale in a naturally fragmented boreal forest landscape. The importance of unexplained between-stand variation in occurrence of species was estimated, and the ability of derived models to predict species' occurrence was tested. The occurrence of species was affected by quality of individual trees. Furthermore, the relationships between occurrence of species at the tree level and size and shape of stands indicated edge effects, implying that some species were restricted to interior habitats of large, regular stands. Yet for the habitat factors studied, requirements of many species appeared similar. Species occurrence also varied between stands; most of the seemingly suitable trees in some stands were unoccupied. The models captured most variation in species occurrence at tree level. They also successfully accounted for between-stand variation in species occurrence, thus providing realistic simulations of stand-level occupancy of species. Important unexplained between-stand variation in species occurrence warns against a simplified view that only local habitat factors influence species' occurrence. Apparently, similar stands will host populations of different sizes due to historical, spatial, and stochastic factors. Thus, habitat suitability cannot be assessed simply by population sizes, and stands lacking a species may still provide suitable habitat and merit protection.     

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.     

Seeing the forest for the heterogeneous trees: stand-scale resource distributions emerge from tree-scale structure

Forest ecosystem processes depend on local interactions that are modified by the spatial pattern of trees and resources. Effects of resource supplies on processes such as regeneration are increasingly well understood, yet we have few tools to compare resource heterogeneity among forests that differ in structural complexity. We used a neighborhood approach to examine understory light and nutrient availability in a well-replicated and large-scale variable-retention harvesting experiment in a red pine forest in Minnesota, USA. The experiment included an unharvested control and three harvesting treatments with similar tree abundance but different patterns of retention (evenly dispersed as well as aggregated retention achieved by cutting 0.1- or 0.3-ha gaps). We measured light and soil nutrients across all treatments and mapped trees around each sample point to develop an index of neighborhood effects (NI). Field data and simulation modeling were used to test hypotheses that the mean and heterogeneity of resource availability would increase with patchiness because of greater variation in competitive environments. Our treatments dramatically altered the types and abundances of competitive neighborhoods (NI) in each stand and resulted in significantly nonlinear relationships of light, nitrogen and phosphorus availability to NI. Hence, the distribution of neighborhoods in each treatment had a significant impact on resource availability and heterogeneity. In dense control stands, neighborhood variation had little impact on resource availability, whereas in more open stands (retention treatments), it had large effects on light and modest effects on soil nutrients. Our results demonstrate that tree spatial pattern can affect resource availability and heterogeneity in explainable and predictable ways, and that neighborhood models provide a useful tool for scaling heterogeneity from the individual tree to the stand. These insights are needed to anticipate the outcomes of silvicultural manipulations and should become more holistically integrated into both basic ecological and management science.     

Do evergreen and deciduous trees have different effects on net N mineralization in soil?

Evergreen and deciduous plants are widely expected to have different impacts on soil nitrogen (N) availability because of differences in leaf litter chemistry and ensuing effects on net N mineralization (N(min)). We evaluated this hypothesis by compiling published data on net N(min) rates beneath co-occurring stands of evergreen and deciduous trees. The compiled data included 35 sets of co-occurring stands in temperate and boreal forests. Evergreen and deciduous stands did not have consistently divergent effects on net N(min) rates; net N(min) beneath deciduous trees was higher when comparing natural stands (19 contrasts), but equivalent to evergreens in plantations (16 contrasts). We also compared net N(min) rates beneath pairs of co-occurring genera. Most pairs of genera did not differ consistently, i.e., tree species from one genus had higher net N(min) at some sites and lower net N(min) at other sites. Moreover, several common deciduous genera (Acer, Betula, Populus) and deciduous Quercus spp. did not typically have higher net N(min) rates than common evergreen genera (Pinus, Picea). There are several reasons why tree effects on net N(min) are poorly predicted by leaf habit and phylogeny. For example, the amount of N mineralized from decomposing leaves might be less than the amount of N mineralized from organic matter pools that are less affected by leaf litter traits, such as dead roots and soil organic matter. Also, effects of plant traits and plant groups on net N(min) probably depend on site-specific factors such as stand age and soil type.     

Bridging the gap between ecophysiological and genetic knowledge to assess the adaptive potential of European beech

In this study we aimed to combine knowledge of the ecophysiology and genetics of European beech to assess the potential of this species to adapt to environmental change. Therefore, we performed field and experimental studies on the genetic and ecophysiological functioning of beech. This information was integrated through a coupled genetic–ecophysiological model for individual trees that was parameterized with information derived from our own studies or from the literature. Using the model, we evaluated the adaptive response of beech stands in two ways: firstly, through sensitivity analyses (of initial genetic diversity, pollen dispersal distance, heritability of selected phenotypic traits, and forest management, representing disturbances) and secondly, through the evaluation of the responses of phenotypic traits and their genetic diversity to four management regimes applied to 10 study plots distributed over Western Europe. The model results indicate that the interval between recruitment events strongly affects the rate of adaptive response, because selection is most severe during the early stages of forest development. Forest management regimes largely determine recruitment intervals and thereby the potential for adaptive responses. Forest management regimes also determine the number of mother trees that contribute to the next generation and thereby the genetic variation that is maintained. Consequently, undisturbed forests maintain the largest amount of genetic variation, as recruitment intervals approach the longevity of trees and many mother trees contribute to the next generation. However, undisturbed forests have the slowest adaptive response, for the same reasons.Gene flow through pollen dispersal may compensate for the loss in genetic diversity brought about by selection. The sensitivity analysis showed that the total genetic diversity of a 2 ha stand is not affected by gene flow if the pollen distance distribution is varied from highly left-skewed to almost flat. However, a stand with a prevailing short-distance gene flow has a more pronounced spatial genetic structure than stands with equal short- and long-distance gene flows. The build-up of a spatial genetic structure is also strongly determined by the recruitment interval. Overall, the modelling results indicate that European beech has high adaptive potential to environmental change if recruitment intervals are short and many mother trees contribute to the next generation.The findings have two implications for modelling studies on the impacts of climate change on forests. Firstly: it cannot be taken for granted that parameter values remain constant over a time horizon of even a few generations – this is particularly important for threshold values subject to strong selection, like budburst, frost hardiness, drought tolerance, as used in species area models. Secondly: forest management should be taken into account in future assessments, as management affects the rate of adaptive response and thereby the response on trees and forests to environmental change, and because few forests are unmanaged. We conclude that a coupled ecophysiological and quantitative genetic tree model is a useful tool for such studies.     

Influence of overstory removal on growth of epiphytic mosses and lichens in western Oregon.

Will old-growth-associated epiphytes survive if the forest canopy is opened around them by thinning or partial harvest? If old-growth association is due to a species' environmental tolerances, it may not survive in the relatively open stands that result from such treatments. If, however, old-growth association is due to dispersal limitations rather than environmental tolerances, retention of host trees as refugia and sources of inoculum might carry populations of old-growth-associated epiphytes into young stands. We studied growth rates of lichen and moss transplants in a Douglas-fir (Pseudotsuga menziesii) forest (tree ages approximately 55 yr) in western Oregon for nine months before and 27 months after moderate thinning, creation of 0.4-ha patch cuts, and in control areas. We also assessed moss sporophyte production. We contrasted responses of one moss species, Isothecium myosuroides sensu lato, which is ubiquitous in forests of varying ages, with those of another moss, Antitrichia curtipendula, and a lichen, Lobaria oregana, which are both associated with old-growth forests. Both old-growth associates grew faster in thinned areas and patch cuts than in controls, while Isothecuim grew most slowly and produced fewest sporophytes in patch cuts. These species are likely to survive in remnants, assuming they can remain attached, and may be successful in young stands if they can disperse and establish there. Our results suggest that logging with green-tree retention and other silvicultural practices that preserve trees or shrubs hosting the species studied here are likely to encourage these species' development in managed forests.     

The importance of age-related decline in forest NPP for modeling regional carbon balances.

We show the implications of the commonly observed age-related decline in aboveground productivity of forests, and hence forest age structure, on the carbon dynamics of European forests in response to historical changes in environmental conditions. Size-dependent carbon allocation in trees to counteract increasing hydraulic resistance with tree height has been hypothesized to be responsible for this decline. Incorporated into a global terrestrial biosphere model (the Lund-Potsdam-Jena model, LPJ), this hypothesis improves the simulated increase in biomass with stand age. Application of the advanced model, including a generic representation of forest management in even-aged stands, for 77 European provinces shows that model-based estimates of biomass development with age compare favorably with inventory-based estimates for different tree species. Model estimates of biomass densities on province and country levels, and trends in growth increment along an annual mean temperature gradient are in broad agreement with inventory data. However, the level of agreement between modeled and inventory-based estimates varies markedly between countries and provinces. The model is able to reproduce the present-day age structure of forests and the ratio of biomass removals to increment on a European scale based on observed changes in climate, atmospheric CO2 concentration, forest area, and wood demand between 1948 and 2000. Vegetation in European forests is modeled to sequester carbon at a rate of 100 Tg C/yr, which corresponds well to forest inventory-based estimates.     

Contrasted dynamics of northern Labrador tree lines caused by climate change and migrational lag

The northern Québec-Labrador tree lines are the most climatically stressed tree ecosystems of eastern North America. In particular, white spruce (Picea glauca) tree line populations distributed between 56 degrees N and 58 degrees N and 61 degrees W and 66 degrees W show contrasted responses to recent changes in climate according to their geographic position relative to the Labrador Sea. Along the coast, the northernmost latitudinal and altitudinal tree lines responded positively to warming over the last 50 years with invading spruce several tens of meters above the current tree line. In contrast, white spruce tree lines across the wind-exposed Labrador plateau are located much higher in altitude and have receded a few tens of meters beginning around AD 1740-1750 and have not yet recovered. Whereas no field evidence of recent fire and insect damage was found, all inland tree line stands were progressively damaged likely due to mechanical defoliation of wind-exposed trees. Massive tree death in the 19th century caused a reduction in the number of seed-bearing trees, and declining tree lines were not replenished by seedlings. Recent warming reported for northern latitudes has not been strong enough to change the regressive tree line trajectory in interior Labrador. However, white spruce expansion above coastal tree line in the northernmost forest site in Labrador is in line with current climatic trends. It is hypothesized that the species is still advancing toward its potential tree line higher in altitude due to delayed postglacial migration. The slow advance of white spruce in northernmost coastal Labrador is likely caused by the rugged topography of the Torngat-Kaumajet-Kiglapait mountains.     

Natural tree collectives of pure oriental spruce [Picea orientalis (L.) Link] on mountain forests in Turkey

Distribution area of oriental spruce [Picea orientalis (L.) Link.] in the world is only in the north-east of Turkey and Caucasian. Because of being the semi monopoly tree with respect to its distribution and representing the upper forest line, it is necessary to analyse, evaluate and model the stand structures of oriental spruce forests in Turkey. In this research, some sampling plots were selected in timberline and treeline in the subalpine forest zone in Turkey. In these sampling plots some information about occurrence and development of the tree collectives was obtained. A total of 12 sampling plots (6 in timberline and 6 of them in treeline) were studied and horizontal and vertical stand profiles were obtained, while number of trees ranges between 2-86 in the tree collectives in treeline and in timberline 3-12. According to this, area per tree in treeline and in timberline is determined as 1.02 m2 and 3.75 m2 on an average respectively. Mean age of trees to reach breast height is 43 years in treeline sampling plots and 22 years in timberline sampling plots. According to the ratio of h (mean height) / d1.30 (diameter at breast height), stand stability values were calculated and it was determined if the stands were stable on the basis of the sampling plots. Stability values of the sampling plots changed between 33 and 75.     

The influence of successional processes and disturbance on the structure of Tsuga canadensis forests

Old-growth forests are valuable sources of ecological, conservation, and management information, yet these ecosystems have received little study in New England, due in large part to their regional scarcity. To increase our understanding of the structures and processes common in these rare forests, we studied the abundance of downed coarse woody debris (CWD) and snags and live-tree size-class distributions in 16 old-growth hemlock forests in western Massachusetts. Old-growth stands were compared with eight adjacent second-growth hemlock forests to gain a better understanding of the structural differences between these two classes of forests resulting from contrasting histories. In addition, we used stand-level dendroecological reconstructions to investigate the linkages between disturbance history and old-growth forest structure using an information-theoretic model selection framework. Old-growth stands exhibit a much higher degree of structural complexity than second-growth forests. In particular, old-growth stands had larger overstory trees and greater volumes of downed coarse woody debris (135.2 vs. 33.2 m3/ha) and snags (21.2 vs. 10.7 m3/ha). Second-growth stands were characterized by either skewed unimodal or reverse-J shaped diameter distributions, while old-growth forests contained bell-shaped, skewed unimodal, rotated sigmoid, and reverse J-shaped distributions. The variation in structural attributes among old-growth stands, particularly the abundance of downed CWD, was closely related to disturbance history. In particular, old-growth stands experiencing moderate levels of canopy disturbance during the last century (1930s and 1980s) had greater accumulations of CWD, highlighting the importance of gap-scale disturbances in shaping the long-term development and structural characteristics of old-growth forests. These findings are important for the development of natural disturbance-based silvicultural systems that may be used to restore important forest characteristics lacking in New England second-growth stands by integrating structural legacies of disturbance (e.g., downed CWD) and resultant tree-size distribution patterns. This silvicultural approach would emulate the often episodic nature of CWD recruitment within old-growth forests.     

Indirect Effects of Pandemic Deer Overabundance Inferred from Caterpillar‐Host Relations

Externally feeding phytophagous insect larvae (i.e., caterpillars, here, larval Lepidoptera and sawflies, Hymenoptera: Symphyta) are important canopy herbivores and prey resources in temperate deciduous forests. However, composition of forest trees has changed dramatically in the eastern United States since 1900. In particular, browsing by high densities of white‐tailed deer (Odocoileus virginianus) has resulted in forests dominated by browse‐tolerant species, such as black cherry (Prunus serotina), and greatly reduced relative abundance of other tree species, notably pin cherry (Prunus pensylvanica) and birches (Betula spp.). To quantify effects of these changes on caterpillars, we sampled caterpillars from 960 branch tips of the 8 tree species that comprise 95% of trees in Allegheny hardwood forests: red maple (Acer rubrum), striped maple (Acer pensylvanicum), sugar maple (Acer saccharum), sweet birch (Betula lenta), yellow birch (Betula allegheniensis), American beech (Fagus grandifolia), black cherry, and pin cherry. We collected 547 caterpillar specimens that belonged to 66 Lepidoptera and 10 Hymenoptera species. Caterpillar density, species richness, and community composition differed significantly among tree species sampled. Pin cherry, nearly eliminated at high deer density, had the highest density and diversity of caterpillars. Pin cherry shared a common caterpillar community with black cherry, which was distinct from those of other tree hosts. As high deer density continues to replace diverse forests of cherries, maples, birches, and beech with monodominant stands of black cherry, up to 66% of caterpillar species may be eliminated. Hence, deer‐induced changes in forest vegetation are likely to ricochet back up forest food webs and therefore negatively affect species that depend on caterpillars and moths for food and pollination. Efectos Indirectos de la Sobreabundancia de Venados Pandémicos Inferida de Relaciones Orugas‐Huéspedes     

Browsing lawns? Responses of Acacia nigrescens to ungulate browsing in an African savanna

We measured browsing-induced responses of Acacia trees to investigate "browsing lawns" as an analogy to grazing lawns in a semiarid eutrophic African savanna. During the two-year field study, we measured plant tolerance, resistance, and phenological traits, while comparing variation in leaf nitrogen and specific leaf area (SLA) across stands of Acacia nigrescens, Miller, that had experienced markedly different histories of attack from large herbivores. Trees in heavily browsed stands developed (1) tolerance traits such as high regrowth abilities in shoots and leaves, high annual branch growth rates, extensive tree branching and evidence of internal N translocation, and (2) resistance traits such as close thorn spacing. However, phenological "escape" responses were weak even in heavily browsed stands. Overall, browsing strongly affected plant morpho-functional traits and decreased both the number of trees carrying pods and the number of pods per tree in heavily browsed stands. Hence, there is experimental evidence that tolerance and resistance traits may occur simultaneously at heavily browsed sites, but this comes at the expense of reproductive success. Such tolerance and resistance traits may coexist if browsers trigger and maintain a positive feedback loop in which trees are continually investing in regrowth (tolerance), and if the plant's physical defenses (resistance) are not nutritionally costly and are long-lived. Our results confirm that chronic browsing by ungulates can maintain A. nigrescens trees in a hedged state that is analogous to a grazing lawn. Further research is needed to fully understand the long-term effects of chronic browsing on reproduction within such tree populations, as well as the overall effects on nutrient cycling at the ecosystem level.     

Evaluation of sampling methods for the estimation of structural indices in forest stands

The paper is about the accurate (i.e. unbiased and precise) and efficient estimation of structural indices in forest stands. We present SIAFOR, a computer programme for the calculation of four nearest-neighbour indices, which describe the spatial arrangement of tree positions, the distribution pattern of species, and the size differentiation between trees. The study uses SIAFOR as a sampling simulator in eight completely stem-mapped forest stands of varying area and structural complexity. We statistically evaluate two sample types (distance and plot sampling), comparing sampling error, bias and minimum sample size for index estimation. We introduce the concepts of measurement expansion factor (MEF) and design expansion factor (DEF) for the technical evaluation of sample type efficiency (optimal sample type). Results indicate that sampling error can reach high levels and that minimum sample sizes for index estimation often amply exceed the limit of 20% of tree density or 20 trees per species per hectare, that we set as the highest feasible sample size in normal situations. We found clear feasibility limits (in terms of minimal tree densities and reachable accuracy levels) for the estimation of all investigated indices. Generally, equal or higher sample sizes are needed for plot sampling than for distance sampling to reach equal accuracy levels. Nevertheless, plot sampling resulted more efficient for the estimation of tree size differentiation at low to medium accuracy levels. For all other investigated indices distance sampling resulted more efficient than plot sampling. Minimum sample size increases with accuracy and is negatively correlated with tree density. At a given accuracy level minimum sample size is highest for the estimation of relative mingling and lowest for tree size differentiation; furthermore it is generally lower in large stands than in small ones. Because of the consistency of our conclusions in all of the investigated stands, we think they apply in most stands of similar area (between 1 and 10 ha) and species diversity (not more than four species).     

Tree growth inference and prediction from diameter censuses and ring widths.

Estimation of tree growth is based on sparse observations of tree diameter, ring widths, or increments read from a dendrometer. From annual measurements on a few trees (e.g., increment cores) or sporadic measurements from many trees (e.g., diameter censuses on mapped plots), relationships with resources, tree size, and climate are extrapolated to whole stands. There has been no way to formally integrate different types of data and problems of estimation that result from (1) multiple sources of observation error, which frequently result in impossible estimates of negative growth, (2) the fact that data are typically sparse (a few trees or a few years), whereas inference is needed broadly (many trees over many years), (3) the fact that some unknown fraction of the variance is shared across the population, and (4) the fact that growth rates of trees within competing stands are not independent. We develop a hierarchical Bayes state space model for tree growth that addresses all of these challenges, allowing for formal inference that is consistent with the available data and the assumption that growth is nonnegative. Prediction follows directly, incorporating the full uncertainty from inference with scenarios for "filling the gaps" for past growth rates and for future conditions affecting growth. An example involving multiple species and multiple stands with tree-ring data and up to 14 years of tree census data illustrates how different levels of information at the tree and stand level contribute to inference and prediction.     

Effect of prior disturbances on the extent and severity of wildfire in Colorado subalpine forests

Disturbances are important in creating spatial heterogeneity of vegetation patterns that in turn may affect the spread and severity of subsequent disturbances. Between 1997 and 2002 extensive areas of subalpine forests in northwestern Colorado were affected by a blowdown of trees, bark beetle outbreaks, and salvage logging. Some of these stands were also affected by severe fires in the late 19th century. During a severe drought in 2002, fires affected extensive areas of these subalpine forests. We evaluated and modeled the extent and severity of the 2002 fires in relation to these disturbances that occurred over the five years prior to the fires and in relation to late 19th century stand-replacing fires. Occurrence of disturbances prior to 2002 was reconstructed using a combination of tree-ring methods, aerial photograph interpretation, field surveys, and geographic information systems (GIS). The extent and severity of the 2002 fires were based on the normalized difference burn ratio (NDBR) derived from satellite imagery. GIS and classification trees were used to analyze the effects of prefire conditions on the 2002 fires. Previous disturbance history had a significant influence on the severity of the 2002 fires. Stands that were severely blown down (> 66% trees down) in 1997 burned more severely than other stands, and young (approximately 120 year old) postfire stands burned less severely than older stands. In contrast, prefire disturbances were poor predictors of fire extent, except that young (approximately 120 years old) postfire stands were less extensively burned than older stands. Salvage logging and bark beetle outbreaks that followed the 1997 blowdown (within the blowdown as well as in adjacent forest that was not blown down) did not appear to affect fire extent or severity. Conclusions regarding the influence of the beetle outbreaks on fire extent and severity are limited, however, by spatial and temporal limitations associated with aerial detection surveys of beetle activity. Thus, fire extent in these forests is largely independent of prefire disturbance history and vegetation conditions. In contrast, fire severity, even during extreme fire weather and in conjunction with a multiyear drought, is influenced by prefire stand conditions, including the history of previous disturbances.