Multiple drivers of large-scale lichen decline in boreal forest canopies.

Thin, hair-like lichens (Alectoria, Bryoria, Usnea) form conspicuous epiphyte communities across the boreal biome. These poikilohydric organisms provide important ecosystem functions and are useful indicators of global change. We analyse how environmental drivers influence changes in occurrence and length of these lichens on Norway spruce (Picea abies) over 10 years in managed forests in Sweden using data from >6000 trees. Alectoria and Usnea showed strong declines in southern-central regions, whereas Bryoria declined in northern regions. Overall, relative loss rates across the country ranged from 1.7% per year in Alectoria to 0.5% in Bryoria. These losses contrasted with increased length of Bryoria and Usnea in some regions. Occurrence trajectories (extinction, colonization, presence, absence) on remeasured trees correlated best with temperature, rain, nitrogen deposition, and stand age in multinomial logistic regression models. Our analysis strongly suggests that industrial forestry, in combination with nitrogen, is the main driver of lichen declines. Logging of forests with long continuity of tree cover, short rotation cycles, substrate limitation and low light in dense forests are harmful for lichens. Nitrogen deposition has decreased but is apparently still sufficiently high to prevent recovery. Warming correlated with occurrence trajectories of Alectoria and Bryoria, likely by altering hydration regimes and increasing respiration during autumn/winter. The large-scale lichen decline on an important host has cascading effects on biodiversity and function of boreal forest canopies. Forest management must apply a broad spectrum of methods, including uneven-aged continuous cover forestry and retention of large patches, to secure the ecosystem functions of these important canopy components under future climates. Our findings highlight interactions among drivers of lichen decline (forestry, nitrogen, climate), functional traits (dispersal, lichen colour, sensitivity to nitrogen, water storage), and population processes (extinction/colonization).

Plant functional traits have globally consistent effects on competition.

Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions, but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear. Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits--wood density, specific leaf area and maximum height--consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies. Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our trait-based approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.

Improving the precision of sample-based forest damage inventories through two-phase sampling and post-stratification using remotely sensed auxiliary information.

Many countries have a national forest inventory (NFI) designed to produce statistically sound estimates of forest parameters. However, this type of inventory may not provide reliable results for forest damage which usually affects only small parts of the forest in a country. For this reason, specially designed forest damage inventories are performed in many countries, sometimes in coordination with the NFIs. In this study, we evaluated a new approach for damage inventory where existing NFI data form the basis for two-phase sampling for stratification and remotely sensed auxiliary data are applied for further improvement of precision through post-stratification. We applied Monte Carlo sampling simulation to evaluate different sampling strategies linked to different damage scenarios. The use of existing NFI data in a two-phase sampling for stratification design resulted in a relative efficiency of 50 % or lower, i.e., the variance was at least halved compared to a simple random sample of the same size. With post-stratification based on simulated remotely sensed auxiliary data, there was additional improvement, which depended on the accuracy of the auxiliary data and the properties of the forest damage. In many cases, the relative efficiency was further reduced by as much as one-half. In conclusion, the results show that substantial gains in precision can be obtained by utilizing auxiliary information in forest damage surveys, through two-phase sampling, through post-stratification, and through the combination of these two approaches, i.e., post-stratified two-phase sampling for stratification.

Monitoring trees outside forests: a review.

Trees outside forests (TOFs) are an important natural resource that contributes substantially to national biomass and carbon stocks and to the livelihood of people in many regions. Over the last decades, decision makers have become increasingly aware of the importance of TOF, and as a consequence, this tree resource is nowadays often considered in forest monitoring systems. Our review shows that in many cases, TOF are included in national forest inventories, applying traditional methodologies with relatively sparse networks of field sample plots. Only in some countries, such as India, the design of the inventories has considered the special features of how TOFs occur in the landscape. Several research studies utilising remote sensing for monitoring TOF have been conducted lately, but very few studies include comparative studies to optimise sampling strategies for TOF. Our review indicates that methods combining remote sensing and field surveys appear to be very promising, especially when remote sensing techniques that assess both the horizontal and vertical structures of tree resources are applied. For example, two-phase sampling strategies with laser scanning in the first phase and a field survey in the second phase appear to be effective for assessing TOF resources. However, TOFs often exhibit different characteristics than forest trees. Thus, to improve TOF monitoring, there is often a need to develop models, e.g. for biomass assessment, that are specifically adapted to this tree resource. Alternatively, field-based remote sensing methods that provide structural information about individual trees, notably terrestrial laser scanning, could be further developed for TOF monitoring applications. This also would have a potential to reduce the problem of accessing TOF during field surveys, which is a problem, for example, in countries where TOF are present on intensively utilised private grounds like gardens and agricultural fields.

The contribution of trees outside forests to national tree biomass and carbon stocks--a comparative study across three continents.

In contrast to forest trees, trees outside forests (TOF) often are not included in the national monitoring of tree resources. Consequently, data about this particular resource is rare, and available information is typically fragmented across the different institutions and stakeholders that deal with one or more of the various TOF types. Thus, even if information is available, it is difficult to aggregate data into overall national statistics. However, the National Forest Monitoring and Assessment (NFMA) programme of FAO offers a unique possibility to study TOF resources because TOF are integrated by default into the NFMA inventory design. We have analysed NFMA data from 11 countries across three continents. For six countries, we found that more than 10% of the national above-ground tree biomass was actually accumulated outside forests. The highest value (73%) was observed for Bangladesh (total forest cover 8.1%, average biomass per hectare in forest 33.4 t ha(-1)) and the lowest (3%) was observed for Zambia (total forest cover 63.9%, average biomass per hectare in forest 32 t ha(-1)). Average TOF biomass stocks were estimated to be smaller than 10 t ha(-1). However, given the large extent of non-forest areas, these stocks sum up to considerable quantities in many countries. There are good reasons to overcome sectoral boundaries and to extend national forest monitoring programmes on a more systematic basis that includes TOF. Such an approach, for example, would generate a more complete picture of the national tree biomass. In the context of climate change mitigation and adaptation, international climate mitigation programmes (e.g. Clean Development Mechanism and Reduced Emission from Deforestation and Degradation) focus on forest trees without considering the impact of TOF, a consideration this study finds crucial if accurate measurements of national tree biomass and carbon pools are required.

Three-phase hierarchical model-based and hybrid inference.

Global commitments to mitigating climate change and halting biodiversity loss require reliable information about Earth's ecosystems. Increasingly, such information is obtained from multiple sources of remotely sensed data combined with data acquired in the field. This new wealth of data poses challenges regarding the combination of different data sources to derive the required information and assess uncertainties. In this article, we show how predictors and their variances can be derived when hierarchically nested models are applied. Previous studies have developed methods for cases involving two modeling steps, such as biomass prediction relying on tree-level allometric models and models linking plot-level field data with remotely sensed data. This study extends the analysis to cases involving three modeling steps to cover new important applications. The additional step might involve an intermediate model, linking field and remotely sensed data available from a small sample, for making predictions that are subsequently used for training a final prediction model based on remotely sensed data:•In cases where the data in the final step are available wall-to-wall, we denote the approach three-phase hierarchical model-based inference (3pHMB),•In cases where the data in the final step are available as a probability sample, we denote the approach three-phase hierarchical hybrid inference (3pHHY).

Adapting forest health assessments to changing perspectives on threats--a case example from Sweden.

A revised Swedish forest health assessment system is presented. The assessment system is composed of several interacting components which target information needs for strategic and operational decision making and accommodate a continuously expanding knowledge base. The main motivation for separating information for strategic and operational decision making is that major damage outbreaks are often scattered throughout the landscape. Generally, large-scale inventories (such as national forest inventories) cannot provide adequate information for mitigation measures. In addition to broad monitoring programs that provide time-series information on known damaging agents and their effects, there is also a need for local and regional inventories adapted to specific damage events. While information for decision making is the major focus of the health assessment system, the system also contributes to expanding the knowledge base of forest conditions. For example, the integrated monitoring programs provide a better understanding of ecological processes linked to forest health. The new health assessment system should be able to respond to the need for quick and reliable information and thus will be an important part of the future monitoring of Swedish forests.

National Inventory of Landscapes in Sweden (NILS)--scope, design, and experiences from establishing a multiscale biodiversity monitoring system.

The landscape-level and multiscale biodiversity monitoring program National Inventory of Landscapes in Sweden (NILS) was launched in 2003. NILS is conducted as a sample-based stratified inventory that acquires data across several spatial scales, which is accomplished by combining aerial photo interpretation with field inventory. A total of 631 sample units are distributed across the land base of Sweden, of which 20% are surveyed each year. By 2007 NILS completed the first 5-year inventory phase. As the reinventory in the second 5-year phase (2008-2012) proceeds, experiences and insights accumulate and reflections are made on the setup and accomplishment of the monitoring scheme. In this article, the emphasis is placed on background, scope, objectives, design, and experiences of the NILS program. The main objective to collect data for and perform analyses of natural landscape changes, degree of anthropogenic impact, prerequisites for natural biological diversity and ecological processes at landscape scale. Different environmental conditions that can have direct or indirect effects on biological diversity are monitored. The program provides data for national and international policy and offers an infrastructure for other monitoring program and research projects. NILS has attracted significant national and international interest during its relatively short time of existence; the number of stakeholders and cooperation partners steadily increases. This is constructive and strengthens the incentive for the multiscale monitoring approach.

Monitoring landscape metrics by point sampling: accuracy in estimating Shannon's diversity and edge density.

Environmental monitoring of landscapes is of increasing interest. To quantify landscape patterns, a number of metrics are used, of which Shannon's diversity, edge length, and density are studied here. As an alternative to complete mapping, point sampling was applied to estimate the metrics for already mapped landscapes selected from the National Inventory of Landscapes in Sweden (NILS). Monte-Carlo simulation was applied to study the performance of different designs. Random and systematic samplings were applied for four sample sizes and five buffer widths. The latter feature was relevant for edge length, since length was estimated through the number of points falling in buffer areas around edges. In addition, two landscape complexities were tested by applying two classification schemes with seven or 20 land cover classes to the NILS data. As expected, the root mean square error (RMSE) of the estimators decreased with increasing sample size. The estimators of both metrics were slightly biased, but the bias of Shannon's diversity estimator was shown to decrease when sample size increased. In the edge length case, an increasing buffer width resulted in larger bias due to the increased impact of boundary conditions; this effect was shown to be independent of sample size. However, we also developed adjusted estimators that eliminate the bias of the edge length estimator. The rates of decrease of RMSE with increasing sample size and buffer width were quantified by a regression model. Finally, indicative cost-accuracy relationships were derived showing that point sampling could be a competitive alternative to complete wall-to-wall mapping.

Estimating dead wood during national forest inventories: a review of inventory methodologies and suggestions for harmonization.

Efforts to assess forest ecosystem carbon stocks, biodiversity, and fire hazards have spurred the need for comprehensive assessments of forest ecosystem dead wood (DW) components around the world. Currently, information regarding the prevalence, status, and methods of DW inventories occurring in the world's forested landscapes is scattered. The goal of this study is to describe the status, DW components measured, sample methods employed, and DW component thresholds used by national forest inventories that currently inventory DW around the world. Study results indicate that most countries do not inventory forest DW. Globally, we estimate that about 13% of countries inventory DW using a diversity of sample methods and DW component definitions. A common feature among DW inventories was that most countries had only just begun DW inventories and employ very low sample intensities. There are major hurdles to harmonizing national forest inventories of DW: differences in population definitions, lack of clarity on sample protocols/estimation procedures, and sparse availability of inventory data/reports. Increasing database/estimation flexibility, developing common dimensional thresholds of DW components, publishing inventory procedures/protocols, releasing inventory data/reports to international peer review, and increasing communication (e.g., workshops) among countries inventorying DW are suggestions forwarded by this study to increase DW inventory harmonization.

Increased spruce tree growth in Central Europe since 1960s.

Tree growth response to recent environmental changes is of key interest for forest ecology. This study addressed the following questions with respect to Norway spruce (Picea abies, L. Karst.) in Central Europe: Has tree growth accelerated during the last five decades? What are the main environmental drivers of the observed tree radial stem growth and how much variability can be explained by them? Using a nationwide dendrochronological sampling of Norway spruce in the Czech Republic (1246 trees, 266 plots), novel regional tree-ring width chronologies for 40(±10)- and 60(±10)-year old trees were assembled, averaged across three elevation zones (break points at 500 and 700m). Correspondingly averaged drivers, including temperature, precipitation, nitrogen (N) deposition and ambient CO2 concentration, were used in a general linear model (GLM) to analyze the contribution of these in explaining tree ring width variability for the period from 1961 to 2013. Spruce tree radial stem growth responded strongly to the changing environment in Central Europe during the period, with a mean tree ring width increase of 24 and 32% for the 40- and 60-year old trees, respectively. The indicative General Linear Model analysis identified CO2, precipitation during the vegetation season, spring air temperature (March-May) and N-deposition as the significant covariates of growth, with the latter including interactions with elevation zones. The regression models explained 57% and 55% of the variability in the two tree ring width chronologies, respectively. Growth response to N-deposition showed the highest variability along the elevation gradient with growth stimulation/limitation at sites below/above 700m. A strong sensitivity of stem growth to CO2 was also indicated, suggesting that the effect of rising ambient CO2 concentration (direct or indirect by increased water use efficiency) should be considered in analyses of long-term growth together with climatic factors and N-deposition.

Discerning environmental factors affecting current tree growth in Central Europe.

We examined the effect of individual environmental factors on the current spruce tree growth assessed from a repeated country-level statistical landscape (incl. forest) survey in the Czech Republic. An extensive set of variables related to tree size, competition, site characteristics including soil texture, chemistry, N deposition and climate was tested within a random-effect model to explain growth in the conditions of dominantly managed forest ecosystems. The current spruce basal area increment was assessed from two consecutive landscape surveys conducted in 2008/2009 and six years later in 2014/2015. Tree size, age and competition within forest stands were found to be the dominant explanatory variables, whereas the expression of site characteristics, environmental and climatic drives was weaker. The significant site variables affecting growth included soil C/N ratio and soil exchangeable acidity (pH KCl; positive response) reflecting soil chemistry, long-term N-deposition (averaged since 1975) in combination with soil texture (clay content) and Standardized Precipitation Index (SPI), a drought index expressing moisture conditions. Sensitivity of growth to N-deposition was positive, although weak. SPI was positively related to and significant in explaining tree growth when expressed for the growth season. Except SPI, no significant relation of growth was determined to altitude-related variables (temperature, growth season length). We identified the current spruce growth optimum at elevations about 800ma.s.l. or higher in the conditions of the country. This suggests that at lower elevations, limitation by a more pronounced water deficit dominates, whereas direct temperature limitation may concern the less frequent higher elevations. The mixed linear model of spruce tree growth explained 55 and 65% of the variability with fixed and random effects included, respectively, and provided new insights on the current spruce tree growth and factors affecting it within the environmental gradients of the country.

Merging national forest and national forest health inventories to obtain an integrated forest resource inventory--experiences from Bavaria, Slovenia and Sweden.

BACKGROUNDS, MATERIAL AND METHODS: To meet the demands of sustainable forest management and international commitments, European nations have designed a variety of forest-monitoring systems for specific needs. While the majority of countries are committed to independent, single-purpose inventorying, a minority of countries have merged their single-purpose forest inventory systems into integrated forest resource inventories. The statistical efficiencies of the Bavarian, Slovene and Swedish integrated forest resource inventory designs are investigated with the various statistical parameters of the variables of growing stock volume, shares of damaged trees, and deadwood volume. The parameters are derived by using the estimators for the given inventory designs. The required sample sizes are derived via the general formula for non-stratified independent samples and via statistical power analyses. The cost effectiveness of the designs is compared via two simple cost effectiveness ratios. RESULTS: In terms of precision, the most illustrative parameters of the variables are relative standard errors; their values range between 1% and 3% if the variables' variations are low (s%<80%) and are higher in the case of higher variations. A comparison of the actual and required sample sizes shows that the actual sample sizes were deliberately set high to provide precise estimates for the majority of variables and strata. In turn, the successive inventories are statistically efficient, because they allow detecting the mean changes of variables with powers higher than 90%; the highest precision is attained for the changes of growing stock volume and the lowest for the changes of the shares of damaged trees. Two indicators of cost effectiveness also show that the time input spent for measuring one variable decreases with the complexity of inventories. CONCLUSION: There is an increasing need for credible information on forest resources to be used for decision making and national and international policy making. Such information can be cost-efficiently provided through integrated forest resource inventories.