Individual-tree diameter growth models for black spruce and jack pine plantations in northern Ontario

Individual-tree distance independent diameter growth models were developed for black spruce and jack pine plantations. Data used in this study came via stem analysis on 1170 black spruce (Picea mariana [Mill.] B.S.P.) and 800 jack pine (Pinus banksiana Lamb.) trees sampled from 75 stands of 25 even-aged monospecific plantations for each species in the Canadian boreal forest region of northern Ontario. Of the 75 stands, 50 were randomly selected for each species and all trees from these stands were used for model development. Trees from the remaining stands were used for model evaluation.A nonlinear mixed-effects approach was applied in fitting the diameter growth models. The predictive accuracy of the models was improved by including random effects coefficients. Four selection criteria - random, dominant or codominant, tree size close to quadratic mean diameter, and small sized - were evaluated for accuracy in predicting random effects for a new stand using the developed models. Random effects predicted based on trees selected using the random selection criterion provided more accurate diameter predictions than those using trees obtained via other selection criteria for both species. The models developed here are very important to forest managers as the diameters predicted by these models or, their stand-level summaries (i.e., basal area, average diameter), are used as inputs in any forest growth and yield models. In addition, individual-tree diameter growth models can be used to directly forecast changes in diameter distribution of stands.     

Forecasting tree mortality using change metrics derived from MODIS satellite data

Insect-induced tree mortality can cause substantial timber and carbon losses in many regions of the world. There is a critical need to forecast tree mortality to guide forest management decisions. Moderate Resolution Imaging Spectroradiometer (MODIS) satellite imagery provides inexpensive and frequent coverage over large areas, facilitating forest health monitoring. This study examined time series of MODIS satellite images to forecast tree mortality for a Pinus radiata plantation in southern New South Wales, Australia. Dead tree density derived from ADS40 aerial imagery was used to evaluate the performance of change metrics derived from time series of MODIS-based vegetation indices. Continuous subset selection by LASSO regression and model assessment using a variant of the bootstrap were used to select the best performing change metrics out of a large amount of predictor variables to account for over-fitting. The results suggest that 250 m 16-daily MODIS images are effective for forecasting tree mortality. Seasonal change metrics derived from the Normalized Difference Vegetation Index (NDVI) outperformed the Enhanced Vegetation Index (EVI) and the Normalized Difference Infrared Index (NDII). Temporal analysis illustrated that optimal forecasting power was obtained using change metrics based on three years of satellite data for this population. The forecast could be used to optimise the scheduling of detailed forest health surveys and silvicultural operations which currently are planned based on stratified, annual assessments. This coarse-scale, spatio-temporal analysis represents a potentially cost-effective early warning approach to forecasting tree mortality in pine plantations by identifying compartments that require more detailed investigation.     

The contribution of competition to tree mortality in old-growth coniferous forests

Competition is a well-documented contributor to tree mortality in temperate forests, with numerous studies documenting a relationship between tree death and the competitive environment. Models frequently rely on competition as the only non-random mechanism affecting tree mortality. However, for mature forests, competition may cease to be the primary driver of mortality.We use a large, long-term dataset to study the importance of competition in determining tree mortality in old-growth forests on the western slope of the Sierra Nevada of California, U.S.A. We make use of the comparative spatial configuration of dead and live trees, changes in tree spatial pattern through time, and field assessments of contributors to an individual tree's death to quantify competitive effects.Competition was apparently a significant contributor to tree mortality in these forests. Trees that died tended to be in more competitive environments than trees that survived, and suppression frequently appeared as a factor contributing to mortality. On the other hand, based on spatial pattern analyses, only three of 14 plots demonstrated compelling evidence that competition was dominating mortality. Most of the rest of the plots fell within the expectation for random mortality, and three fit neither the random nor the competition model. These results suggest that while competition is often playing a significant role in tree mortality processes in these forests it only infrequently governs those processes. In addition, the field assessments indicated a substantial presence of biotic mortality agents in trees that died.While competition is almost certainly important, demographics in these forests cannot accurately be characterized without a better grasp of other mortality processes. In particular, we likely need a better understanding of biotic agents and their interactions with one another and with competition.     

Height–diameter equations for boreal tree species in Ontario using a mixed-effects modeling approach

Height–diameter relationships based on stand characteristics (trees/ha, basal area, and dominant stand height) were investigated for balsam fir, balsam poplar, black spruce, jack pine, red pine, trembling aspen, white birch, and white spruce using data from permanent growth study plots in northern Ontario, Canada. Approximately half the data were used to estimate model parameters with the rest used for model evaluation. Multiple Chapman–Richards functions with parameters expressed in terms of various stand characteristics were fit to determine the best models for predicting height.     

A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests

Greenhouse gas emissions have significantly altered global climate, and will continue to do so in the future. Increases in the frequency, duration, and/or severity of drought and heat stress associated with climate change could fundamentally alter the composition, structure, and biogeography of forests in many regions. Of particular concern are potential increases in tree mortality associated with climate-induced physiological stress and interactions with other climate-mediated processes such as insect outbreaks and wildfire. Despite this risk, existing projections of tree mortality are based on models that lack functionally realistic mortality mechanisms, and there has been no attempt to track observations of climate-driven tree mortality globally. Here we present the first global assessment of recent tree mortality attributed to drought and heat stress. Although episodic mortality occurs in the absence of climate change, studies compiled here suggest that at least some of the world's forested ecosystems already may be responding to climate change and raise concern that forests may become increasingly vulnerable to higher background tree mortality rates and die-off in response to future warming and drought, even in environments that are not normally considered water-limited. This further suggests risks to ecosystem services, including the loss of sequestered forest carbon and associated atmospheric feedbacks. Our review also identifies key information gaps and scientific uncertainties that currently hinder our ability to predict tree mortality in response to climate change and emphasizes the need for a globally coordinated observation system. Overall, our review reveals the potential for amplified tree mortality due to drought and heat in forests worldwide.     

Pinus pinaster Ait. tree mortality following wildfire in Spain

Maritime pine (Pinus pinaster Ait.) is the tree species most affected by wildfire in the Iberian Peninsula. Prediction of the probability of fire-injured tree mortality is critical for management of burned areas, evaluation of the ecological and economic impact of wildfire and prescribed fire planning and application. Pine bark beetles (Scolytidae) frequently attack burned maritime pine stands and cause extensive post-fire mortality throughout the Iberian Peninsula. In the present study, maritime pine trees were monitored for three years following 14 wildfires in four ecotypes in Spain (11 fires in Galicia (Galician ecotype - NW Spain), one fire in Portillo (Meseta-Castellana ecotype - Central Spain), one fire in Rodenal (Rodenal ecotype - Central Spain), and one fire in Genalguacil (Sierra Bermeja ecotype - SW Spain)). Data on tree attributes, crown and bole injury, ground fire severity, Ips sp. presence and tree survival were obtained by examining 3085 trees. Logistic regression models for predicting the probability of delayed maritime pine mortality were developed by use of generalized estimated equations (GEE). An ample range of response to fire damage in mortality was evident among the four ecotypes and different models were fitted for each. The most important variables for predicting tree mortality were total crown volume damaged, presence of Ips sp. attack and cambium kill rating. The results highlight the extensive presence of Ips sp. in burned maritime pine forests and its importance in tree mortality process, the ample range of response of P. pinaster, in terms of post-fire mortality, as well as the need to develop site specific mortality models for the different ecotypes of this species following fire.     

Modelling growth for two-storied stands: Incorporating an effect of shading on the tree growth into growth and yield models

The objective of this paper is to review research into growth and yield modelling for two-storied plantation forests. Difficulty with modelling growth for this kind of forest, compared with pure even-aged forests, is mentioned. Then research in this field is divided into three parts and described. First, growth models and equations which were driven theoretically from biological assumptions and may be available to develop growth models for two-storied stands are reviewed. Secondly, research into methods of controling light levels and the relationship between the growth of trees and light level beneath a canopy are reviewed. Thirdly, growth models developed for two-storied stands, especially two-storied plantation forests, are reviewed. In conclusion, some problems which are still left in this field are pointed out.     

Modelling the impacts of various thinning intensities on tree growth and survival in a mixed species eucalypt forest in central Gippsland,Victoria, Australia

The response of tree survival and diameter growth to thinning treatments was examined over 29 years, in various thinning treatments established in a 21-year-old even-aged mixed species regenerating forest in Victoria, Australia. The treatments were control, crown release, strip thinning and three different intensities of thinning from below (light, moderate, and heavy). Each treatment was replicated three times in a complete randomised design. Logistic and multilevel regression analyses showed that tree survival, growth and thinning response (change of tree growth due to a thinning treatment) were functions of tree species, size, age, removed and remaining competition, as well as time since the treatment. Mean annual tree diameter growth in unthinned stands was highest for Eucalyptus sieberi L. Johnson (1.9 mm) followed by Eucalyptus baxteri (Benth.) Maiden & Blakely ex J. Black (1.6 mm), and lowest for both Eucalyptus consideniana (Maiden) and Eucalyptus radiata (Sieber ex DC) combined (0.7 mm). Diameter growth increased with tree size for both E. sieberi and E. baxteri, but not for E. consideniana and E. radiata. Smaller trees were more likely to die due to shading and suppression than their larger counterparts. A mortality model suggested, however, that both shading and suppression had very little effect on trees in both E. consideniana and E. radiata species, which were less likely to die compared to trees in the other species. This result indicates that both E. consideniana and E. radiata species may be relatively shade tolerant compared with the other species. Total thinning response was a sum of positive (increased growing space) and negative (thinning stress) effects. Following thinning, smaller trees showed signs of thinning stress for the first one or two years, after which the highest percentage thinning response was observed. While larger trees were initially less responsive to thinning, the rate of decrease in the response for subsequent years was greater in smaller trees than larger ones. The average amount of thinning response showed similar trends to diameter growth increasing from E. sieberi (1.7 mm) through E. baxteri (0.6 mm) to both E. consideniana and E. radiata (0.5 mm). This translates into low average percentage thinning response in E. baxteri (34%), twice as much in both E. consideniana and E. radiata (69%) and highest overall percentage response in E. sieberi (87%). Thinning response and the duration of this response appeared to increase with thinning intensity and was still evident 29 years after thinning. Heavy thinning did, however, reduce the number of trees to a severely under-stocked condition, which prohibited optimum site occupancy, requiring 29 years of post-thinning development for the heavily thinned stands to regain their pre-thinning stand basal area.     

Modelling root rot incidence in Sweden using tree,site and stand variables

Decay and root rot caused by Heterobasidion annosum (Fr.) Bref. s. lato is the most serious disease of Norway spruce [Picea abies (L.) Karst.]. Mathematical models of disease development have recently been developed for forestry planning purposes. Functions for predicting the probability of decay were developed from two data sets, one comprising trees and another comprising stumps. From the years 1983–2001, 45,587 Norway spruce trees from the Swedish National Forest Inventory (NFI) were analysed for decay incidence at 1.3 m height and correlation with environmental conditions. The decay frequency increased in all studied regions from the first to the second half of the period for trees with comparable tree and environmental attributes. In a stepwise logistic regression, sets of functions were developed showing significance regarding stand age, site index class, temperature sum, height above sea level, diameter at 1.3 m, soil moisture and texture, proportion of spruce and eastern coordinates. The functions were calibrated and validated with a data set from the Swedish NFI from the years 1993–2002 comprising 7,893 stumps. The calibration of decay incidence at breast height to stump height doubled the decay incidence (R ²=0.85). The developed functions could be used to establish initial conditions for dynamic modelling of disease and in strategic planning.     

Individual tree diameter increment model for managed even-aged stands of ponderosa pine throughout the western United States using a multilevel linear mixed effects model

A diameter increment model is developed and evaluated for individual trees of ponderosa pine throughout the species range in the United States using a multilevel linear mixed model. Stochastic variability is broken down among period, locale, plot, tree and within-tree components. Covariates acting at tree and stand level, as breast height diameter, density, site index, and competition indices are included in the model as fixed effects in order to explain residual variability. The data set used in this study came from long-term permanent research plots in even-aged, pure stands both planted and of natural origin. The data base consists of six levels-of-growing stock studies supplemented by initial spacing and other permanent-plot thinning studies for a total of 310 plots, 34,263 trees and 153,854 observations. Regression analysis is the preferred technique used in growth and yield modeling in forestry. We choose the mixed effects models instead of the regression analysis approach because it allows for proper treatment of error terms in a repeated measures analysis framework. Regional growth and yield models exist for ponderosa pine. However, data collection and analysis procedures differ. As a result, comparisons of growth responses that may be due to geographic variation of the species are not possible. Our goal is to present a single distance-independent diameter increment model applicable throughout the geographic range of ponderosa pine in the United States and by using only data from long-term permanent plots on sites capable of the productivity estimated by Meyer [Meyer, W.H., 1938. Yield of Even-Aged Stands of Ponderosa Pine. US Department of Agriculture Technical Bulletin 630].     

Ectomycorrhizal development in a Pinus thunbergii stand in relation to location on a slope and effect on tree mortality from pine wilt disease

The relationship between ectomycorrhizal development and mortality from pine wilt disease was studied in an artificial Pinus thunbergii Parl. stand on a slope. The development of ectomycorrhizae and the survival of the trees showed the same tendency, which suggests a correlation between mycorrhizal development and resistance to pine wilt disease. The development of pine roots and mycorrhizae was greater in the upper part of the slope. The ratio of mycorrhizae to the total of mycorrhizae and fine taproots was also higher in the upper part of the slope. Tree mortality was clearly biased and more trees survived in the upper part of the slope than in the middle and the lower parts. There was no significant difference between the upper and the lower part of the slope in the number of feeding wounds made by the pine sawyer beetle, which demonstrates the opportunity of infection with this disease. There was no clear correlation between the development of mycorrhizae and the composition of the soil substrate such as total carbon, nitrogen, and phosphorus. The abundant mycorrhizae in the upper part of the slope, which mitigate drought stress, may also have decreased the rate of tree mortality.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.     

Modelling dominant height growth from national forest inventory individual tree data with short time series and large age errors

We developed dominant height growth models for Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) in Norway using national forest inventory (NFI) data. The data were collected for a different purpose which potentially causes problems for dominant height growth modelling due to short time series and large age errors. We used the generalized algebraic difference approach and fitted 15 different models using nested regression techniques. Despite the potential problems of NFI data the models fitted to these data were unbiased for most of the age and site index range covered by the NFI data when tested against independent data from long-term experiments (LTE). Biased predictions for young stands and better site indices that are better represented in the LTE data, led us to fit models to a combined data set for unbiased predictions across the total data range. The models fitted to the combined data that were unbiased with little residual variation when tested against an independent data set based on stem analysis of 73 sample trees from southeastern Norway. No indications of regional differences in dominant height growth across Norway were detected. We tested whether the better growing conditions during the short time series (22 years) of the NFI data had affected our dominant height growth models relative to long-term growing conditions, but found only minor bias. The combination with LTE data that have been collected during a longer period (91 years) reduced this potential bias. The dominant height growth models presented here can be used as potential height growth models in individual tree-based forest growth models or as site index models.     

Effects of flooding on the recruitment, damage and mortality of riparian tree species: A field and simulation study on the Rhine floodplain

The extensive flooding by the river Rhine on May 12 1999 provided an opportunity to investigate the impact of such an extreme event in terms of damage and mortality of adult trees in floodplains. Such data is highly valuable for determining the potential impact of climate change on the zonation of tree species along rivers. We analysed an extensive dataset of the damage and mortality suffered by groups of adult trees of the following species as a consequence of this flood: the hardwoods Acer campestre L., Acer platanoides L., Acer pseudoplatanus L., Alnus glutinosa (L.) Gaertn., Carpinus betulus L., Fagus sylvatica L., Fraxinus excelsior L., Juglans nigra L., Prunus avium (L.) L., Quercus robur L., Tilia cordata Mill., Ulmus laevis Pall. and Ulmus minor Miller, and the softwoods Salix spp. L. and Populus spp. L.A logistic survivorship curve revealed that mortality of A. platanoides, A. pseudoplatanus and T. cordata increased significantly with increasing duration of flooding; C. betulus and F. excelsior showed a significant increase of damage and mortality with increasing flooding depth. There was no mortality of Salix spp. and Populus spp. in either the flooded or unflooded areas. No statistically significant relationships were found for the other tree species. Multivariate analysis revealed that flooding duration, flooding depth and flooding velocity explain 19%, 11% and 8%, respectively, of the variation in damage and mortality of trees.The survivorship curves of adult trees obtained in this study were combined with similar curves of saplings based on an earlier study and applied in an individual-tree, process-based simulation model. The simulated effects of flooding on an initial random distribution of trees species on a hypothetical floodplain resulted in a realistic zonation of tree species along the river. When extreme events were simulated, the zonation shifted upward. This demonstrates the model's usefulness in assessment and planning studies of the impacts of climate change on tree species composition in river floodplains in north-west Europe.