Can a multistage approach improve individual tree mortality predictions across the complex mixed-species and managed forests of eastern North America?

Tree mortality plays a fundamental role in the dynamics of forest ecosystems,yet it is one of the most difficult phenomena to accurately predict.Various modeling strategies have been developed to improve individual tree mortality predictions.One less explored strategy is the use of a multistage modeling approach.Potential improvements from this approach have remained largely unknown.In this study,we developed a novel multistage approach and compared its performance in individual tree mortality predictions with a more conventional approach using an identical individual tree mortality model formulation.Extensive permanent plot data(n=9442)covering the Acadian Region of North America and over multiple decades(1965–2014)were used in this study.Our results indicated that the model behavior with the multistage approach better depicted the observed mortality and showed a notable improvement over the conventional approach.The difference between the observed and predicted numbers of dead trees using the multistage approach was much smaller when compared with the conventional approach.In addition,tree survival probabilities predicted by the multistage approach generally were not significantly different from the observations,whereas the conventional approach consistently underestimated mortality across species and overestimated tree survival probabilities over the large range of DBH in the data.The new multistage approach also predictions of zero mortality in individual plots,a result not possible in conventional models.Finally,the new approach was more tolerant of modeling errors because it based estimates on ranked tree mortality rather than error-prone predicted values.Overall,this new multistage approach deserves to be considered and tested in future studies.

Modelling tree mortality across diameter classes using mixedeffects zero-inflated models

The mortality of trees across diameter class model is a useful tool for predicting changes in stand structure.Mortality data commonly contain a large fraction of zeros and general discrete models thus show more errors.Based on the traditional Poisson model and the negative binomial model,different forms of zero-inflated and hurdle models were applied to spruce-fir mixed forests data to simulate the number of dead trees.By comparing the residuals and Vuong test statistics,the zero-inflated negative binomial model performed best.A random effect was added to improve the model accuracy;however,the mixed-effects zero-inflated model did not show increased advantages.According to the model principle,the zeroinflated negative binomial model was the most suitable,indicating that the"0"events in this study,mainly from the sample"0",i.e.,the zero mortality data,are largely due to the limitations of the experimental design and sample selection.These results also show that the number of dead trees in the diameter class is positively correlated with the number of trees in that class and the mean stand diameter,and inversely related to class size,and slope and aspect of the site.

Tree mortality and regeneration of Euphrates poplar riparian forests along the Tarim River,Northwest China

Background:Tree mortality and regeneration(seedling and sapling recruitment)are essential components of forest dynamics in arid regions,especially where subjected to serious eco-hydrological problems.In recent decades,the mortality of the Euphrates poplar(Populus euphratica)along the Tarim River in Northwest China has increased.However,few studies have quantified the causes of mortality and regeneration in this azonal riparian forest type.Methods:The present study describes the annual hydrological response of tree mortality and regeneration in forest gaps.A total of 60 canopy gaps were investigated in six replicate grid plots(50m×50 m)and the annual runoff and water consumption data during the period of 1955–2016 were collected from hydrological stations in the middle reaches of the Tarim River.We compared the regeneration density of seedlings and saplings within the canopy gap areas(CGAs),undercanopy areas(UCAs),and uncovered riverbank areas(RBAs)through detailed field investigation.Results:Our study found that the mortality of young and middle-aged gap makers has increased remarkably over recent decades,particularly since the year 1996.The main results indicated that regional water scarcity was the primary limiting factor for long-term changes in tree mortality,as shown by a significant correlation between the diameter at breast height(DBH)of dead trees and the annual surface water.The average density(or regeneration rate)of seedlings and saplings was highest in the RBAs,intermediate in the CGAs,and lowest in the UCAs.Compared with the UCAs,the CGAs promote tree regeneration to some extent by providing favorable conditions for the survival and growth of seedlings and saplings,which would otherwise be suppressed in the understory.Furthermore,although the density of seedlings and saplings in the CGAs was not as high as in the RBAs,the survival rate was higher in the CGAs than in the RBAs.Conclusion:Forest canopy gaps in floodplain areas can play a decisive role in the long-term germination and regeneration of plant species.However,as a typical phreatophyte in this hyper-arid region,the ecosystem structure,functions and services of this fragile P.euphratica floodplain forests are threatened by a continuous decrease of water resources,due to excessive water use for agricultural irrigation,which has resulted in a severe reduction of intact poplar forests.Furthermore,the survival of seedlings and saplings is influenced by light availability and soil water at the regional scale.Our findings suggest that policymakers may need to reconsider the restoration and regeneration measures implemented in riparian P.euphratica forests to improve flood water efficiency and create canopy gaps.Our results provide with valuable reference information for the conservation and sustainable development of floodplain forest ecosystems.

Hydraulic properties and drought response of a tropical bamboo (Cephalostachyum pergracile)

Bamboo plants are an essential component of tropical ecosystems,yet their vulnerability to climate extremes,such as drought,is poorly understood due to limited knowledge of their hydraulic properties.Cephalostachyum pergracile,a commonly used tropical bamboo species,exhibited a substantially higher mortality rate than other co-occurring bamboos during a severe drought event in 2019,but the underlying mechanisms remain unclear.This study investigated the leaf and stem hydraulic traits related to drought responses,including leaf-stem embolism resistance(P50leaf;P50stem) estimated using optical and X-ray microtomography methods,leaf pressure-volume and water-releasing curves.Additionally,we investigated the seasonal water potentials,native embolism level(PLC) and xylem water source using stable isotope.We found that C.pergracile exhibited strong resistance to embolism,showing low P50leaf,P50stem,and turgor loss point,despite its rapid leaf water loss.Interestingly,its leaves displayed greater resistance to embolism than its stem,suggesting a lack of effective hydraulic vulnerability segmentation(HVS) to protect the stem from excessive xylem tension.During the dry season,approximately 49% of the water was absorbed from the upper 20-cm-deep soil layer.Consequently,significant diurnal variation in leaf water potentials and an increase in midday PLC from 5.87±2.33% in the wet season to 12.87±4.09%in the dry season were observed.In summary,this study demonstrated that the rapid leaf water loss,high reliance on surface water,and a lack of effective HVS in C.pergracile accelerated water depletion and increased xylem embolism even in the typical dry season,which may explain its high mortality rate during extreme drought events in 2019.

Dead standing pine trees in a boreal forest landscape in the Kalevala National Park,northern Fennoscandia: amount,population characteristics and spatial pattern

Background: After their death, Scots pine trees can remain standing for decades and sometimes up to 200 years,forming long-lasting and ecologically important structures in boreal forest landscapes. Standing dead pines decay very slowly and with time develop into ‘kelo' trees, which are characterized by hard wood with silvery-colored appearance. These kelo trees represent an ecologically important, long lasting and visually striking element of the structure of natural pine-dominated forests in boreal Fennoscandia that is nowadays virtually absent from managed forest landscapes.Methods: We examined and mapped the amount, structural features, site characteristics and spatial distribution of dead standing pine trees over a ten hectare area in an unmanaged boreal forest landscape in the Kalevala National Park in Russian Viena Karelia.Results: The mean basal area of dead standing pine trees in the forested part of the landscape was 1.7 m^2?ha^(-1) and the estimated volume 12.7 m^3?ha^(-1). From the total number of standing dead pine trees 65% were kelo trees, with a basal area of 1.1 m^2?ha^(-1) and volume of 8.0 m^3?ha^(-1), the remainder consisting of standing dead pines along the continuum between a recently dead tree and a kelo tree. Overall, standing dead pines were distributed throughout the study area, but there was a tendency towards spatial clustering up to <100 m distances. Standing dead pines were most commonly situated on flat ground or in the mid slope in the local topography.In addition, standing dead pines contributed to substrate diversity also by commonly having charred wood and broken tops. Based on the presence of dead pine snags in different stage of transition from a recently dead pine to a kelo with silvery surface, it seems evident that the process of kelo recruitment was continuously in action in the studied landscape.Conclusions: Kelo trees are an omnipresent feature in natural pine-dominated forest landscapes with important contribution to forest structural and substrate diversity. Because of their longevity and extremely slow turnover dynamics and importance for biodiversity, protection of vulnerable kelo tree populations, and ensuring their continuous recruitment, should be of high priority in forest restoration and sustainable management.

The growth and production modeling of individual trees of Eucalyptus urophylla plantations

Individual tree models(ITMs)are classified as growth and production models for projecting current and future forest stands.ITMs are more complex than other growth and production models,show a higher level of detail and,consequently,produce a better modeling resolution.However,the accuracy and efficiency of ITMs have not been properly assessed to date.In this study,we estimated the growth in height,diameter,and individual tree volume of a Eucalyptus urophylla plantation by applying an ITM.We used a continuing forest inventory dataset in which 1554 individual trees within 29 permanent plots were measured in the field over a 6-year period(24 to 72 months).Each individual tree volume was estimated for future tree age.To achieve this,we adjusted the model to predict the height and diameter growth,and the probability of mortality as a function of the competition index.The ITM accuracy was assessed based on the analysis of variance results and,subsequently,the multiple mean comparison test at the 5%significance level.The tree volumes predicted by the ITM for the forest stand aged 72 months,beginning at ages 24,36,48,and 60 months,were compared to the field measured tree volume acquired from the 72-month forest inventory that was used as the reference age.Estimated and observed tree volumes were similar when the estimation was based on the 48-month forest plots.These results might help to reduce financial costs of forest inventory because the ITM produces accurate future predictions of forest stand stocks.Our estimated ITM for Eucalyptus plantations using measurement intervals up to 2 years is recommended because it significantly reduced the projected volume discrepancy compared to the field measurements.

Spatial Distribution of Hemlock Woolly Adelgid Induced Hemlock Mortality in the Southern Appalachians

Hemlock woolly adelgid (Adelges Tsugae Annand, HWA) outbreaks are posing a major threat to eastern hemlock (Tsuga canadensis L. Carr.) and Carolina hemlock (Tsuga caroliniana Engelm.) forest landscapes in the eastern USA. As foundation species, hemlocks play a variety of functional roles in forest landscapes. These species usually occur as isolated canopies and mixed species in landscapes where variation in topography is extreme. Spatially explicit inventory information on HWA induced hemlock mortality at landscape scale does not exist. High resolution aerial imageries enable landscape scale assessment even at the individual tree level. Accordingly, our goal was to investigate spatial pattern and distribution of HWA induced hemlock mortality using a high resolution aerial image mosaic in the Linville River Gorge, Southern Appalachians, western North Carolina. Our study objectives were: 1) to detect dead trees within the Lower Linville River watershed;2) to estimate the area occupied by dead trees in the forest canopy surface;3) to investigate the relationship of dead hemlocks and topography;and 4) to define the spatial pattern of the dead trees. We found ca. 10,000 dead trees within the study area, occupying over 7 ha of the canopy surface with an average area of 36 m2 per dead tree. The density of the dead trees was higher in proximity to the Linville River, at higher elevations, and on northern and northwestern aspects. Spatial pattern of the dead trees was generally clustered at all spatial scales. We suggest that although the reduction in plant biomass resulting from herbivory within the landscapes is modest, impact of the clustered distribution of hemlock mortality, especially in the riparian zones, is noteworthy. Our analysis of the pattern of hemlock decline provides new means for projecting future impacts of HWA on the range of hemlock distribution in eastern North America.

Mangrove forests as a nature-based solution for coastal flood protection:Biophysical and ecological considerations

Nature-based coastal protection is increasingly recognised as a potentially sustainable and cost-effective solution to reduce coastal flood risk.It uses coastal ecosystems such as mangrove forests to create resilient designs for coastal flood protection.However,to use mangroves effectively as a nature-based measure for flood risk reduction,we must understand the biophysical processes that govern risk reduction capacity through mangrove ecosystem size and structure.In this perspective,we evaluate the current state of knowledge on local physical drivers and ecological processes that determine mangrove functioning as part of a nature-based flood defence.We show that the forest properties that comprise coastal flood protection are well-known,but models cannot yet pinpoint how spatial heterogeneity of the forest structure affects the capacity for wave or surge attenuation.Overall,there is relatively good understanding of the ecological processes that drive forest structure and size,but there is a lack of knowledge on how daily bed-level dynamics link to long-term biogeomorphic forest dynamics,and on the role of combined stressors influencing forest retreat.Integrating simulation models of forest structure under changing physical(e.g.due to sea-level change)and ecological drivers with hydrodynamic attenuation models will allow for better projections of long-term natural coastal protection.

Decline and dieback of cork oak(Quercus suber L.)forests in the Mediterranean basin:a case study of Kroumirie,Northwest Tunisia

Assessing the vulnerability of forest ecosystems in the climate change context is a challenging task as the mechanisms that determine this vulnerability cannot be directly observed.Based on the ecological interrelationships between forests and climate,the present review focused on providing current information about vulnerability assessments of cork oak(Quercus suber L.)forests in the Mediterranean basin,especially,in the Kroumirie region(northwest Tunisia),currently under historic extreme drought conditions.From comparing recent findings in this region,we synthesized data on cork oak decline and mortality collected during the historic drought years 1988–1995 period.Climate change impacts cork forest decline,with special interest shown in elevated temperatures and drought;cork oak forest regeneration,and the adaptation of the Kroumirie forest to climate change,are reviewed herein.The studied region has been influenced largely by frequent prolonged drought periods,especially from 1988 to 1995.Droughts were found to consistently have a more detrimental impact on the growth and mortality rates of cork oak populations.Cork oak mortality was recorded for up to 63,622 trees.In the future,more research studies and observational data will be needed,which could represent an important key to understand ecosystem processes,and to facilitate the development of better models that project climate change impacts and vulnerability.The study is useful for researchers and forestry decision makers to develop the appropriate strategies to restore and protect ecosystems,and to help anticipate potential future droughts and climate change.

Twenty years of drought-mediated change in snag populations in mixed-conifer and ponderosa pine forests in Northern Arizona

Background:Snags(standing dead trees)are important biological legacies in forest systems,providing numerous resources as well as a record of recent tree mortality.From 1997 to 2017,we monitored snag populations in drought-influenced mixed-conifer and ponderosa pine(Pinus ponderosa)forests in northern Arizona.Results:Snag density increased significantly in both forest types.This increase was driven largely by a pulse in snag recruitment that occurred between 2002 and 2007,fol owing an extreme drought year in 2002,with snag recruitment returning to pre-pulse levels in subsequent time periods.Some later years during the study also were warmer and/or drier than average,but these years were not as extreme as 2002 and did not trigger the same level of snag recruitment.Snag recruitment was not equal across tree species and size classes,resulting in significant changes in species composition and size-class distributions of snag populations in both forest types.Because trees were far more abundant than snags in these forests,the effect of this mortality pulse on tree populations was far smal er than its effect on snag populations.Snag loss rates increased over time during the study,even though many snags were newly recruited.This may reflect the increasing prevalence of white fir snags and/or snags in the smal er size classes,which general y decay faster than snags of other species or larger snags.Thus,although total numbers of snags increased,many of the newly recruited snags may not persist long enough to be valuable as nesting substrates for native wildlife.Conclusions:Increases in snag abundance appeared to be due to a short-term tree mortality"event"rather than a longerterm pattern of elevated tree mortality.This mortality event fol owed a dry and extremely warm year(2002)embedded within a longer-term megadrought.Climate models suggest that years like 2002 may occur with increasing frequency in the southwestern U.S.Such years may result in additional mortality pulses,which in turn may strongly affect trajectories in abundance,structure,and composition of snag populations.Relative effects on tree populations likely wil be smal er,but,over time,also could be significant.

Coarse woody debris features of a warm temperate deciduous broad-leaved forest,northern China

Stocking and structural composition of a deciduous broad-leaved forest were determined to predict coarse woody debris quantity by quantifying the empirical relationships between these two attributes.The most ecologically significant families by stem density were Salicaceae,Betulaceae,Fagaceae,and Aceraceae.P opulus davidiana was the most dominant species followed by B etula dahurica,Quercus mongolica,and Acer mono.The four species accounted for 69.5%of total stems.Numerous small-diameter species characterized the coarse woody debris showing a reversed J-shaped distribution.The coarse debris of P.davidiana,B.dahurica,and Q.mongolica mainly comprised the 10–20 cm size class,whereas A.mono debris was mainly in the 5–10 cm size class.The spatial patterns of different size classes of coarse woody debris were analyzed using the g-function to determine the size of the tree at its death.The results indicate that the spatial patterns at the 0–50 m scale shifted gradually from an aggregated to a random pattern.For some species,the larger coarse debris might change from an aggregated to a random distribution more easily.Given the importance of coarse woody debris in forest ecosystems,its composition and patterns can improve understanding of community structure and dynamics.The aggregation pattern might be due to density dependence and self-thinning effects,as well as by succession and mortality.The four dominant species across the different size classes showed distinct aggregated distribution features at different spatial scales.This suggests a correlation between the dominant species population,size class,and aggregated distribution of coarse woody debris.

The importance of large-diameter trees to the creation of snag and deadwood biomass

Background:Baseline levels of tree mortality can,over time,contribute to high snag densities and high levels of deadwood(down woody debris)if fire is infrequent and decomposition is slow.Deadwood can be important for tree recruitment,and it plays a major role in terrestrial carbon cycling,but deadwood is rarely examined in a spatially explicit context.Methods:Between 2011 and 2019,we annually tracked all trees and snags≥1 cm in diameter and mapped all pieces of deadwood≥10 cm diameter and≥1 m in length in 25.6 ha of Tsuga heterophylla/Pseudotsuga menziesii forest.We analyzed the amount,biomass,and spatial distribution of deadwood,and we assessed how various causes of mortality that contributed uniquely to deadwood creation.Results:Compared to aboveground woody live biomass of 481 Mg ha^(−1)(from trees≥10 cm diameter),snag biomass was 74 Mg ha^(−1) and deadwood biomass was 109 Mg ha^(−1)(from boles≥10 cm diameter).Biomass from large-diameter trees(≥60 cm)accounted for 85%,88%,and 58%,of trees,snags,and deadwood,respectively.Total aboveground woody live and dead biomass was 668 Mg ha^(−1).The annual production of downed wood(≥10 cm diameter)from tree boles averaged 4 Mg ha^(−1) yr^(−1).Woody debris was spatially heterogeneous,varying more than two orders of magnitude from 4 to 587 Mg ha^(−1) at the scale of 20 m×20 m quadrats.Almost all causes of deadwood creation varied in importance between large-diameter trees and small-diameter trees.Biomass of standing stems and deadwood had weak inverse distributions,reflecting the long period of time required for trees to reach large diameters following antecedent tree mortalities and the centennial scale time required for deadwood decomposition.Conclusion:Old-growth forests contain large stores of biomass in living trees,as well as in snag and deadwood biomass pools that are stable long after tree death.Ignoring biomass(or carbon)in deadwood pools can lead to substantial underestimations of sequestration and stability.

Changes in forest structure, species diversity and spatial pattern following hurricane disturbance in a Piedmont North Carolina forest, USA

Aims Large hurricanes have profound impacts on temperate forests,but owing to their infrequent nature these effects have rarely been examined in detail.In 1996,Hurricane Fran significantly damaged many long-term tree census plots in the Duke Forest on the North Carolina Piedmont,thereby providing an exceptional opportunity to examine pre-and post-hurricane forest compositional trajectories.Our goal was to examine immediate,short-term(0–4 years)and longer term(;5 year)hurricane-induced structural,spatial and compositional changes in the tree population(stem d.b.h>1 cm)in the context of our detailed,long-term knowledge of the dynamics of these forests.Methods We surveyed stem damage and tree mortality in 34 long-term permanent plots(ca.70-year record;404–1012 m^(2))and 7 large mapped tree stands(ca.20-year record;5250–65000 m^(2))representing both transition-phase,even-aged pine stands and uneven-aged upland hardwood forests.We employed three types of damage measures to quantify stand-level damage severity:percentage of stems damaged,percentage of basal area lost and a‘stand-level damage index’.Second-order spatial analysis(Ripley’s K-function)was used to investigate patterns in tree mortality.Important findings Our study found hurricane effects on the structural attributes of Piedmont forests to be variable and patchy.Changes in tree species composition,however,were modest.Uprooting was the major damage type for the overstory trees[diameter at breast height(d.b.h.)>10 cm]apparently due to the exposure of the crowns to high wind combined with heavy rainfall prior to and during the storm.Saplings,juvenile trees and small trees(1–10 cm d.b.h.)of the understory and midstory were mainly damaged by being pinned or bent by their damaged large neighbors.Hurricane-induced tree mortality varied weakly among species,was positively correlated with pre-hurricane tree size and remained up to 2-fold higher than pre-hurricane background mortality 5 years after the hurricane.Spatial point pattern analysis revealed a patchy distribution of tree mortality during the hurricane sampling interval.Hurricane Fran resulted in a dramatic increase in average gap size from ca.400 m^(2) pre-hurricane to ca 1100 m^(2) after the hurricane,whereas maximum gap sizes reached 18–34 times larger than the pre-hurricane levels.