Blue‐stain infections in roots,stems and branches of declining Pinus sylvestris trees in a dry inner alpine valley in Switzerland

Increased mortality rates in Scots pine (Pinus sylvestris) forests have recently been observed in the inner alpine Swiss Rhone valley. Drought, in combination with stand competition, mistletoe infections as well as nematode and insect infestations, appears to be the main factor for the decline. In focus of this study was the occurrence and role of fungal pathogens in the decline dynamics. Branches, stems and roots of 208 trees in five different crown transparency classes were collected and inspected for blue stain and fungal infections. Neither Armillaria species nor Heterobasidon annosum s. str. were detected, but blue stain was commonly observed. Visible blue stain increased with increasing crown transparency. Among the recently dead trees, 80% showed visible blue stain in the branches, 90% in the roots and 100% in the stems. In the crown transparency classes 2 and 3 (25–60% crown transparency), five of the 103 trees showed visible blue stain in the roots, one of 130 trees in the stem but none in the branches. Blue‐stain fungi were isolated from all parts of the trees and from all crown transparency classes. Overall incidence of blue stain was highest in the roots and lowest in the branches. In class 2, roots of 60% of the trees were visibly blue‐stained or developed blue stain in culture, but stems of only 24% and branches of 14% of the trees. In the roots Leptographium species, mostly L. serpens, dominated. From stems and branches, mainly Ophiostoma species were isolated. The positive relationship between the incidence of blue stain and crown transparency, in combination with the high infection levels of roots of fairly vigorous Scots pines, indicates the pathogenic potential of the blue‐stain fungi. Hence, these fungi together with their insect vectors may well act as an important contributing factor involved in pine decline.     

Quantifying the effect of pine mistletoe on the growth of Scots pine

Mistletoe infection results in substantial growth losses in mistletoe‐infected forests. This study reports and evaluates the results of retrospective analyses of radial growth of Scots pine (Pinus sylvestris) in relation to the level of infection of pine mistletoe (Viscum album ssp. austriacum). A total of 43 Scots pine trees were destructively sampled from different sites. Of these trees, 14 were uninfected and 29 were infected. Infection classes were determined using six‐class dwarf mistletoe rating system (DMRS). All needle and mistletoe biomass were removed completely and weighed for each sampled tree. Subsamples from needles and all mistletoe biomass were taken to the laboratory for oven‐dried weight determinations. Five‐cm‐thick wood discs were cut from the stem at the breast height (1.3 m) to determine annual basal area increment for the last 25 years. In addition to DMRS, new infection classes were created using mistletoe‐to‐needle biomass (MB/NB) ratio. The results showed that the radial growth losses could be as much as 41% to 64% at different infection levels. The rate of growth loss in relation to DMRS and MB/NB ratio was similar, but with a larger variability in DMRS values. The results showed that both DMRS rating and MB/NB ratio seem to be important for quantifying growth loss on Scots pine trees infected with mistletoe. The results of this study can also be invaluable in modelling the effects of mistletoe on the growth of Scots pine trees.     

Transformation of western hemlock (Tsuga heterophylla) tree crowns by dwarf mistletoe (Arceuthobium tsugense,Viscaceae)

Dwarf mistletoes (Arceuthobium species) are arboreal, hemiparasitic plants of conifers that can change the structure and function of the tree crown. Hemlock dwarf mistletoe (Arceuthobium tsugense subsp. tsugense) principally parasitizes western hemlock (Tsuga heterophylla) and effects 10.8% of all western hemlock trees in Oregon, USA. In this study, we climbed 16 western hemlock trees (age 97–321 years, height 33–54.7 m) across a gradient of infection (0%–100% of branches infected) and measured occurrence of all dwarf mistletoe infections, dwarf mistletoe caused deformities, foliage, branch and crown metrics, and sapwood area. We then modelled over 25 different response variables using linear and generalized linear models with three metrics of severity as explanatory variables: total infection incidence, proportion of all live branches infected, and proportion of all live, infected branches with 33 per cent or more foliage distal to infection. A strong effect of dwarf mistletoe intensification was the reduction of branch foliage and an increase in the proportional amount of foliage distal to infections, with severely infected trees having the majority of foliage distal to infections. Increasing severity led to an apparent crown compaction as crown volumes decreased and became increasingly comprised of deformities. Sapwood area was unrelated to infection severity. Branch length and diameters were unrelated to increasing infection severity despite severely infected branches supporting 1–70 infections. The most severely infected tree had 3,615 individual plants in the crown. Our results suggested that shifts in crown structure and branch deformation, foliage amount, and foliage distal to infection, reflected a likely reduction of capacity for tree growth that coincided with a hypothesized increase in resource demand by dwarf mistletoe plants as infection severity intensified.     

The effects of pine mistletoe (Viscum album subsp. austriacum) on the growth of Scots pine and Crimean pine in Turkey

In this study, the effect of pine mistletoe (Viscum album subsp. austriacum) on basal area increment of Crimean pine and Scots pine was investigated. Dendrochronological data were collected from 223 (71 uninfected and 152 infected) Crimean pines and 195 (77 uninfected and 118 infected) Scots pines located in Kastamonu province of Turkey in 2014. Infected sample trees were classified as light, moderate or severe infection levels. Growth trends and basal area increment loses were compared between uninfected and infected trees for the periods of the last 10, 20 and 30 years. In addition, infection status of forest stands was investigated using temporary sample plots; 27 plots in Crimean pine stands and 26 plots in Scots pine. Results demonstrated that basal area increments were negatively affected by pine mistletoe for both species. Mean basal area increment losses of infected trees for the last decade were determined as 24% for Scots pine and 26% for Crimean pine. Basal area increment losses varied by infection levels (light, moderate and severe) as follows: 25%, 20% and 28% for Scots pines and 20%, 32% and 9% for Crimean pines. Scots pine stands were more severely infected by pine mistletoe than Crimean pine stands. There were negative correlations between number of infected trees and stand density for both species, while positive correlation was detected between the number of infected trees and mean diameter for Scots pine. The results of this study indicate that the pine mistletoe infection has negative effect on radial growth of Scots pine and Crimean pine trees. The results can be an important contribution to the forest management and protection activities in mistletoe-infected stands.     

Synergistic effect of drought and chestnut blight (Cryphonectria parasitica) on growth decline of European chestnut (Castanea sativa)

The chestnut blight fungus [Cryphonectria parasitica (Murill) Barr] has threatened European chestnut stands (Castanea sativa Mill.) in the 20th century, but infected trees recovered because of the appearance of hypovirulent strains. However, within a dry inner Alpine valley (Italy), blight‐infected C. sativa showing various degrees of crown dieback and dead trees were found. We conducted a dendroecological analysis to retrospectively evaluate a possible synergistic effect of blight infection in the early 1970s and climate stress on the growth decline of C. sativa. In the Eisack Valley (Italy), where annual precipitation is <700 mm, increment cores were taken from blight‐infected C. sativa (n = 103) showing different levels of decline symptoms, i.e. extent of crown dieback (healthy, moderate, severe), and from dead trees. Ring width and basal area increment (BAI) chronologies were developed based on dendroecological methods. Growth–climate relationships were explored using response function analysis and Pearson correlation coefficients. Major findings of our study were: (i) C. sativa growth is limited by low precipitation from December to February and high temperatures in June, (ii) BAI of all vitality classes except healthy trees shows a decreasing trend since 1980; and (iii) a severe drought in 1996 accelerated growth decline and caused death of infected C. sativa individuals. Because of the strong influence of climate on radial tree growth within the study area and observed divergent growth trends in selected vitality classes after infestation by hypovirulent C. parasitica strains in the early 1970s, we conclude that although blight infection is a prerequisite for observed growth declines, soil water availability in the years of drought strongly affects susceptibility to tree death.     

Factors influencing the formation of heartwood discolouration in sycamore (<Emphasis Type="Italic">Acer pseudoplatanus</Emphasis> L.)

Sycamore (Acer pseudoplatanus L.) is a tree species with highly valuable wood. Similar to European beech (Fagus sylvatica L.), the stemwood of sycamore can be devalued due to the discolouration i.e. brown heart (hearwood discolouration, hereafter). This paper aims to establish the influence of tree traits and site characteristics on the formation of heartwood discolouration and the possibility of predicting the occurrence of heartwood discolouration. For this purpose, 351 sycamore trees from 38 sites in Slovenia, most of them on carbonate bedrock and in the mountain vegetation belt, were analysed. Stem analysis was carried out on all the trees and the age, diameter at breast height (dbh), height of crown base, forking and crown size, were established. On the stump and upper fronts of the first and second logs, the maximum heartwood discolouration diameter was measured. Using logistic regression, the positive influence of age, relative crown length and average diameter increment on the formation of heartwood discolouration was ascertained. Conversely, the probability of heartwood discolouration was diminished by diameter increment in the last 20 years. In the less productive sites, the probability of formation of heartwood discolouration was relatively high in small diameter trees, but it increased slowly with diameter. Tree forking also contributed to a large extent of the heartwood discolouration. After the diameter at breast height achieves 45 cm, the formation of heartwood discolouration on the first log is highly probable. As in beech, the heartwood discolouration increases along the stem axis up to a height of 6–8 m, and decreases in the higher parts of the tree.     

Growth characteristics and reproductive output of dwarf mistletoe-infected Juniperus polycarpos in Iran

Dwarf mistletoes are parasitic flowering plants that infect conifers, resulting in substantial loss of growth and mortality. Recently, forest managers in Iran are contemplating whether infection of Juniperus polycarpos C. Koch forests by dwarf mistletoe, Arceuthobium oxycedri （DC.） M. Bieb, influences tree vigor and contributes to insuffieient natural regeneration. The present study aimed at assessing the severity of infection and its impact on growth and reproductive output of./., polycar- pos. Infected and uninfected trees （n =20 each） were selected for assess- ment of diameter, height, crown area, and crown volume as well as quantity and quality of cones and seeds. The severity of infection of trees was determined by Hawksworth＇s 6-class dwarf mistletoe rating （DMR） system. The DMR system revealed that 40% of the infected sample trees were lightly infected （DMR =1-2） and 60% were moderately infected （DMR =3--4）. Growth characteristics did not differ significantly （p 〉 0.05） between infected and uninfected trees. However, moderate infec- tion affected the reproductive output of./. polycarpos by significantly （p 〈0.05） reducing the mean number of cones per unit area of the crown, increasing the number of damaged seeds, and reducing seed size and seedgermination capacity. We conclude that reproductive output of J.. poly- carpos is more sensitive than growth characters to moderate infection by juniper dwarf mistletoe, and this might partly account for poor natural regeneration.     

Mistletoe effects on Scots pine decline following drought events: insights from within-tree spatial patterns, growth and carbohydrates

Forest decline has been attributed to the interaction of several stressors including biotic factors such as mistletoes and climate-induced drought stress. However, few data exist on how mistletoes are spatially arranged within trees and how this spatial pattern is related to changes in radial growth, responses to drought stress and carbon use. We used dendrochronology to quantify how mistletoe (Viscum album L.) infestation and drought stress affected long-term growth patterns in Pinus sylvestris L. at different heights. Basal area increment (BAI) trends and comparisons between trees of three different infestation degrees (without mistletoe, ID1; moderately infested trees, ID2; and severely infested trees, ID3) were performed using linear mixed-effects models. To identify the main climatic drivers of tree growth tree-ring widths were converted into indexed chronologies and related to climate data using correlation functions. We performed spatial analyses of the 3D distribution of mistletoe individuals and their ages within the crowns of three severely infested pines to describe their patterns. Lastly, we quantified carbohydrate and nitrogen concentrations in needles and sapwood of branches from severely infested trees and from trees without mistletoe. Mistletoe individuals formed strongly clustered groups of similar age within tree crowns and their age increased towards the crown apex. Mistletoe infestation negatively impacted growth but this effect was stronger near the tree apex than in the rest of sampled heights, causing an average loss of 64% in BAI (loss of BAI was ～51% at 1.3 m or near the tree base). We found that BAI of severely infested trees and moderately or non-infested trees diverged since 2001 and such divergence was magnified by drought. Infested trees had lower concentrations of soluble sugars in their needles than non-infested ones. We conclude that mistletoe infestation causes growth decline and increases the sensitivity of trees to drought stress.     

Tree Mortality,Foliage Recovery and Top-kill in Stands of Scots Pine (Pinus sylvestris) Subsequent to Defoliation by the Pine Looper (Bupalus piniaria)

In 1996, the pine looper (Bupalus piniaria) (L.) defoliated 7000 ha of Scots pine (Pinus sylvestris L.) forest at Hökensås, in southern Sweden. To study tree mortality, foliage recovery and top-kill, plots were laid out in stands with varying levels of defoliation in autumn 1997. Tree mortality peaked 2 yrs after defoliation, and amounted to 25% in stands suffering from 90–100% defoliation. Suppressed trees suffered higher mortality than intermediate and dominant trees. In stands suffering <90% defoliation, tree mortality did not exceed 8%. Foliage recovery in moderately and severely defoliated stands was not complete at the end of the 4 yr study period, whereas slightly defoliated stands had regained full foliage in 1998. Top-kill was most frequent in severely defoliated stands, and 50% of all trees in these stands suffered from top-kill at the end of the study period in spring 2001.     

Towards standardised crown condition assessment in poplar plantations

? This work aims at developing new tools for rapid assessment of forest health indicators in poplar plantations.

? Crown transparency and discoloration were visually evaluated in all trees of four 15 m-radius sub-plots in 32 poplar clonal plantations, which were chosen according to a factorial scheme with three factors: tree age, site quality and understorey vegetation management. A subset of trees was assessed using digital photos processed with a semi-automatic image analysis system (the CROCO software) in order to compare visual and digital crown transparency estimates.

? Poplar crown conditions were better in young stands and rich sites. Harrowing understorey vegetation improved tree health in poor sites. Samples of 20 trees per stand provided the same information about crown transparency and discoloration as 60 trees. Calibration curves of digital crown transparency estimates were successfully fitted against visual crown transparency estimates. The same effects of stand age and site quality could be detected with digital crown transparency as response variable.

? The use of digital photos processed with CROCO in ca. twenty trees per stand is therefore recommended to accurately and objectively monitor crown condition in clonal poplar plantations.

     

Tree growth as indicator of tree vitality and of tree reaction to environmental stress: a review

The intensive monitoring plots (Level II) of ICP Forests serve to examine the effects of air pollution and other stress factors on forest condition, including tree vitality. However, tree vitality cannot be measured directly. Indicators, such as tree growth or crown transparency, may instead be used. Tree growth processes can be ranked by order of importance in foliage growth, root growth, bud growth, storage tissue growth, stem growth, growth of defence compounds and reproductive growth. Under stress photosynthesis is reduced and carbon allocation is altered. Stem growth may be reduced early on as it is not directly vital to the tree. Actual growth must be compared against a reference growth, such as the growth of trees without the presumed stress, the growth of presumed healthy trees, the growth in a presumed stress-free period or the expected growth derived from models. Several examples from intensive monitoring plots in Switzerland illustrate how tree-growth reactions to environmental stresses may serve as vitality indicator. Crown transparency and growth can complement each other. For example, defoliation by insects becomes first visible in crown transparency while stem growth reaction occurs with delay. On the other hand, extreme summer drought as observed in large parts of Europe in 2003 affects stem growth almost immediately, while foliage reduction becomes only visible months later. Residuals of tree growth models may also serve as indicators of changed environmental conditions. Certain stresses, such as drought or insect defoliation cause immediate reactions and are not detectable in five-year growth intervals. Therefore, annual or inter-annual stem growth should be assessed in long-term monitoring plots. An erratum to this article can be found at     

The influence of prescribed crown fire on lodgepole pine dwarf mistletoe (Arceuthobium americanum) populations 33 years post‐fire

Dwarf mistletoes (Arceuthobium spp.) are a group of obligate, hemiparasitic plants that infect numerous species in the Pinaceae in North America. Wildland fire is considered to be the primary natural agent influencing the population and distribution of dwarf mistletoes across landscapes. Based on this understanding, prescribed fire has been suggested as a potential method for dwarf mistletoe sanitation and control; however, experimental work has primarily focused on prescribed surface fire. In this study, we report long‐term impacts of three experimental crown fires on dwarf mistletoe severity in infested lodgepole pine stands in Colorado 33 years post‐fire. The three fires achieved tree mortality rates ranging from 20% to 100%. Our results suggested a significant negative relationship between the amount of fire‐caused tree mortality and future dwarf mistletoe severity. These findings supported the presumed natural role of fire in altering dwarf mistletoe populations, which perhaps exhibits a linear relationship between fire‐caused host tree mortality and future dwarf mistletoe severity.     

Signals of summer drought in crown condition data from the German Level I network

Crown transparency estimates of Scots pine, Norway spruce, common beech, pedunculate and sessile oak, annually surveyed between 1990 and 2004 within a grid over Germany, provide a suitable response variable to study drought effects on forest trees. Major climatic factors, available on a monthly basis as plot-specifically interpolated values and parameters of site and stand conditions, biotic and other relevant factors were used as predictors in different cross- and length-sectional, and longitudinal models. Stand age is a considerable and most constant driver of crown transparency in all species. Pine, spruce and beech responded—mainly with a delay of 1 year—with some foliar loss in areas where there was a surplus of temperature after the generally hot and dry summer of 2003. Parallel time-series analyses delivered species-specific geographic large-scale patterns with delayed or recent precipitation deficits or temperature surpluses. Even if beech is partly responding in current years with leaf loss towards precipitation surpluses, defoliation is especially high 1 year after hot summers, partly a result of high seed sets after such summers. Crown condition of oak responds in dry and warm areas according to the drought stress hypothesis, however, in cool and wet mountainous ranges oak responds after wet summers with higher defoliation. Longitudinal approaches revealed for all 4-tree species significant relationships between crown condition and deviations from the long-term means of temperature, precipitation but also global radiation and wind speed. Results do not always match the drought stress hypothesis, however, this is not to expect considering the heterogeneous site, stand and climatic conditions across Germany. Complex interactions of climatic and biotic factors also impede simple relationships. Soil-related clusters reveal higher sensitivity of spruce and beech towards climatic drought factors on more acid soils with thin humus layers. Also clusters constructed from plot-specific courses of defoliation reveal groups with rather closer relationships like a group of pine plots in the Oberpfalz, which seems to be especially sensitive to summer drought.     

Epidemiology of Heterobasidion abietinum and Viscum album on silver fir (Abies alba) stands of the Pyrenees

In the last two decades, stand decline and increased mortality has affected silver fir (Abies alba) forests in the Spanish Pyrenees. Simultaneously severe occurrences of the root rot fungus Heterobasidion annosum s.l. and of the mistletoe Viscum album have been reported. We aimed to improve the understanding of the epidemiology of both pathogens in our region. All H. annosum isolates found on silver fir were typed as H. abietinum. H. abietinum was more frequently observed where cuttings had targeted fir trees rather than other species. H. abietinum fruiting bodies were observed in the most recently cut stumps. V. album was more abundant on more dominant fir trees, and in southern aspect stands. The number of V. album colonies in the stand correlated (R² = 0.40) with silver fir mortality. Stands with a high level of V. album infection tended to have a smaller percentage of basal area in species other than silver fir, and they tended to be located on more south‐facing slopes. H. abietinum was widespread in silver fir forests of the Pyrenees. Our data suggest that, in the Pyrenees, the observed H. abietinum incidence may represent a combination of both primary and secondary spread of the pathogen. Favouring mixed forests should be tested as a potential control method for V. album. The correlation between silver fir mortality and V. album infection warrants further study, as the observed tree mortality could have occurred due to other factors than V. album, such as drought damage.     

Effects of Mexican dwarf mistletoe (Arceuthobium vaginatum subsp. vaginatum) on the growth of Pinus cooperi in Durango,Mexico—A case study

Mexican dwarf mistletoe (Arceuthobium vaginatum subsp. vaginatum, Viscaceae) is the most widespread and damaging parasitic plant in Mexico. It parasitizes 10 species of pines (Pinus spp., Pinaceae) as principal hosts, including Pinus cooperi, one of the economically most important pines in the state of Durango. As a case study, we used stem analysis to estimate the effects of Mexican dwarf mistletoe on volume and height growth of dwarf mistletoe‐infected P. cooperi in western Durango (Ejido El Brillante, Municipality Pueblo Nuevo). This case study sampled a total of 48 trees, 12 in each of four infection classes estimated using the 6‐class dwarf mistletoe rating system (DMR): uninfected (DMR 0), lightly infected (DMR 1–2), moderately infected (DMR 3–4) and severely infected (DMR 5–6). Significant reductions in both volume and height growth were found for moderately and severely infected trees when compared to uninfected trees. On average, reductions in volume growth and height growth were as high as 50% and 17%, respectively. The largest growth reductions were for moderately infected trees, but large growth reductions also occurred for severely infected trees. Because of the reduced growth associated with moderate to severe infection, Mexican dwarf mistletoe‐infested pine forests in Durango, where timber production is a high priority, should be managed using harvesting practices that reduce dwarf mistletoe infection, and thereby, increase forest productivity.     

Oak mortality associated with crown dieback and oak borer attack in the Ozark Highlands

Oak decline and related mortality have periodically plagued upland oak–hickory forests, particularly oak species in the red oak group, across the Ozark Highlands of Missouri, Arkansas and Oklahoma since the late 1970s. Advanced tree age and periodic drought, as well as Armillaria root fungi and oak borer attack are believed to contribute to oak decline and mortality. Declining trees first show foliage wilt and browning, followed by progressive branch dieback in the middle and/or upper crown. Many trees eventually die if severe crown dieback continues. In 2002, more than 4000 living oak trees ≥11 cm dbh in the relatively undisturbed mature oak forests of the Missouri Ozark Forest Ecosystem Project (MOFEP) were randomly selected and inventoried for tree species, dbh, crown class, crown width, crown dieback condition (healthy: <5% crown dieback, slight: >5–33%, moderate: 33–66%, and severe: >66%) and number of emergence holes created by oak borers on the lower 2.4 m of the tree bole. The same trees were remeasured in 2006 to determine their status (live or dead). In 2002, about 10% of the red oak trees showed moderate or severe crown dieback; this was twice the percentage observed for white oak species. Over 70% of trees in the red oak group had evidence of oak borer damage compared to 35% of trees in the white oak group. There was significant positive correlation between crown dieback and the number of borer emergence holes (p < 0.01). Logistic regression showed oak mortality was mainly related to crown width and dieback, and failed to detect any significant link with the number of oak borer emergence holes. Declining red oak group trees had higher mortality (3 or 4 times) than white oaks. The odds ratios of mortality of slightly, moderately, and severely declining trees versus healthy trees were, respectively, 2.0, 6.5, and 29.7 for black oak; 1.8, 3.8, and 8.3 for scarlet oak; and 2.6, 6.5 and 7.1 for white oaks.     

Individual vulnerability factors of Silver fir (Abies alba Mill.) to parasitism by two contrasting biotic agents: mistletoe (Viscum album L. ssp. abietis) and bark beetles (Coleoptera: Curculionidae: Scolytinae) during a decline process

? Context

In recent decades, there have been increasing reports of forest decline, especially in Mediterranean forest ecosystems. Decline in tree vitality is usually due to complex interactions between abiotic factors and biotic agents that attack weakened trees.

? Aims and methods

Estimating dendrometrical characteristics [basal area increment (BAI), age at DBH from tree ring counting, social status, height, and diameter], tree health status, and a competition index, we investigated the individual vulnerability of a French declining silver fir forest to both mistletoe (Viscum album L. ssp. abietis) and bark beetles (Pityophthorus pityographus Ratz., Pityokteines vorontzovi Jac., and Pityokteines spinidens Reitt.).

? Results

BAI was negatively correlated with both mistletoe infection (via mistletoe biomass) and bark beetle attack (number of insects per square meter), but there was evidence of divergence in tree choice between two groups of parasites. Mistletoe preferentially infected isolated and dominant trees that showed higher past growth rates than non-infected ones. Conversely, bark beetles mainly attacked defoliated and preferably declining trees with diameter (DBH) lower than 44.5 cm and slower past growth.

? Conclusion

While successive severe drought periods are thought to greatly weaken southern silver fir populations, mistletoe and bark beetles may contribute actively to their decline processes as inciting and contributing factors, respectively.     

The contribution of structural indices to the modelling of Sitka spruce (Picea sitchensis) and birch (Betula spp.) crowns

Crown dimensions are important for the quantification of tree interactions in some growth models. This study investigates the potential for structural indices and other spatial measures to improve the prediction of crown radius and crown length for birch (Betula spp.) and Sitka spruce (Picea sitchensis (Bong.) Carr.) in forests in Wales. Crown dimensions were measured for 125 birch and 154 spruce in six fully stem-mapped research plots. These data were used to test the performance of a crown radius model and a crown length model which estimated crown dimensions on the basis of allometric relationships with stem dimensions. Spatial data from the six plots were used to calculate the structural indices mean directional index, diameter correlation index, species mingling, dbh and height dominance, and dbh differentiation, as well as the Hegyi competition index, and basal area of neighbours and larger neighbours, for each crown measurement sample tree, using various numbers of nearest neighbours. Two non-spatial indices, BAL and BALMOD, were also calculated for all sample trees for comparison. These spatial and non-spatial variables were then incorporated into modified crown dimension models. Model performances, in terms of efficiency and relative bias, were compared to determine whether the inclusion of spatial or non-spatial variables resulted in any improvements over models using tree dimensions alone. Crown length and radius were found to be correlated with most of the spatial measures studied. Models incorporating spatial variables gave improvements in performance over allometric models for every data set, and performed more consistently than models containing non-spatial variables. The greatest improvements were achieved for suppressed birch in unthinned forests which had irregularly shaped and strongly displaced crowns. The spatial variable contributing to the most efficient model for each data set varied widely. This points to the complexity of tree spatial interactions and indicates that there is a great deal of scope for investigating other structural indices and crown dimension model forms.     

Spatial pattern dynamics of oak mortality and associated disease symptoms in a California hardwood forest affected by sudden oak death

Sudden oak death is a disease affecting coastal forests in California and southern Oregon. The spatial pattern of disease dynamics is important for forest and landscape pathology; in this work we investigated the interaction across landscape scales of disease symptomology in coast live oaks, Quercus agrifolia, (trunk bleeding, presence of beetles, and presence of the fungus Hypoxylon thouarsianum) and tree mortality through time. We used two-dimensional spatial analysis tools with data gathered in point-centered-quarter format in 2001 and 2004 to quantify the population density of the disease through time; to examine the spatial pattern of tree mortality across scales through time; and to examine the spatial co-occurrence of disease symptoms with crown mortality through time. Early in the study period dead trees were strongly clustered at smaller scales (~300 m) and after three years this clustering was less pronounced. Bleeding on trees occurred in clusters away from dead trees, particularly in 2004, likely indicating a new cohort of infected trees. The presence of H. thouarsianum was strongly related to overstory mortality through time. Beetle-infested trees co-occurred with mortality in 2001. By 2004, they occurred throughout the forest, and were less strongly correlated with overstory tree mortality, suggesting a future peak of tree mortality.     

Tree mortality risk of oak due to gypsy moth

We present prediction models for estimating tree mortality resulting from gypsy moth, Lymantria dispar, defoliation in mixed oak, Quercus sp., forests. These models differ from previous work by including defoliation as a factor in the analysis. Defoliation intensity, initial tree crown condition (crown vigour), crown position, and species grouping classes were highly significant in categorical analysis of variance for mortality. Heavy defoliation intensity was shown to have a strong, consistent influence in increasing the probability of tree mortality. Classification and Regression Tree (CART) analysis, a binomial decision tree procedure, was used to develop prediction models of mortality risk for use by forest managers. The best decision tree had 65 groups that correctly classified 75% of the live trees and 76% of the dead trees. Models were run separately by defoliation class and provided correct classifications between 63 and 78% of the trees. Forest land managers can use these models to assign probabilities of death for moderate and heavy defoliation intensity levels and compare predicted mortality to mortality of undefoliated trees to determine how gypsy moth defoliation will affect their stands. The probabilities can be used to develop marking guides Lased on projected defoliation levels for implementing silvicultural treatments to minimize gypsy moth effects in forest stands prior to infestation.