Occurrence,incidence and associations among fungal pathogens and Agrilus auroguttatus,and their roles in Quercus agrifolia decline in California

Synchronous decline of oak (Quercus spp.) trees in woodlands has been described in Europe and eastern North America as a complex interaction of stressors that predispose, incite or contribute to tree death. This study presents a 2‐year (2010–2011) assessment of the role of pathogens in coast live oak (Quercus agrifolia) woodlands in southern California where oak mortality occurs in locations that are infested and uninfested by the goldspotted oak borer (GSOB, Agrilus auroguttatus). Cumulative coast live oak mortality was not significantly different between sites and was weakly correlated with Diplodia corticola and GSOB incidence and negatively correlated with annual relative humidity. Multiple logistic regression models explained the presence of individual fungi or GSOB at the tree level. Fisher's exact test analysis determined that the presence of D. corticola, Fusarium solani, Dothiorella iberica, Cryptosporiopsis querciphila and Diatrypella verrucaeformis were each related to origin of sample location on tree, and C. querciphila was additionally related to symptom type on the bole. Multiple linear regression models showed high correlation between environmental variables and plot‐level incidence of both GSOB and D. corticola. Disease incidence (DI) for D. corticola was highest in GSOB‐uninfested locations. Jaccard index of association (J) showed that D. corticola was negatively associated with the presence of GSOB, F. solani and C. querciphila. Results suggest that oak decline in California is an example of a complex syndrome involving strong regional differences in factors that are associated with the problem.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

Carbon and nitrogen cycling immediately following bark beetle outbreaks in southwestern ponderosa pine forests

Bark beetle infestation is a well-known cause of historical low-level disturbance in southwestern ponderosa pine forests, but recent fire exclusion and increased tree densities have enabled large-scale bark beetle outbreaks with unknown consequences for ecosystem function. Uninfested and beetle-infested plots (n = 10 pairs of plots on two aspects) of ponderosa pine were compared over one growing season in the Sierra Ancha Experimental Forest, AZ to determine whether infestation was correlated with differences in carbon (C) and nitrogen (N) pools and fluxes in aboveground biomass and soils. Infested plots had at least 80% of the overstory ponderosa pine trees attacked by bark beetles within 2 years of our measurements. Both uninfested and infested plots stored ∼9 kg C m⁻² in aboveground tree biomass, but infested plots held 60% of this aboveground tree biomass in dead trees, compared to 5% in uninfested plots. We hypothesized that decreased belowground C allocation following beetle-induced tree mortality would alter soil respiration rates, but this hypothesis was not supported; throughout the growing season, soil respiration in infested plots was similar to uninfested plots. In contrast, several results supported the hypothesis that premature needlefall from infested trees provided a pulse of low C:N needlefall that altered soil N cycling. The C:N mass ratio of pine needlefall in infested plots (∼45) was lower than uninfested plots (∼95) throughout the growing season. Mineral soils from infested plots had greater laboratory net nitrification rates and field resin bag ammonium accumulation than uninfested plots. As bark beetle outbreaks become increasingly prevalent in western landscapes, longer-term biogeochemical studies on interactions with other disturbances (e.g. fire, harvesting, etc.) will be required to predict changes in ecosystem structure and function.     

Amino nitrogen and phenolic constituents of bark of American beech,Fagus grandifolia,and infestation by beech scale,Cryptococcus fagisuga

Concentrations of amino acids, total amino nitrogen, and phenols, ratio of phenol to amino nitrogen, and pH in the bark of American beech, Fagus grandifolia, were determined to see if these chemical constituents were correlated with susceptibility of trees to infestation by the beech scale, Cryptococcus fagisuga. The relationship of these bark constituents with scale infestation levels (none, light, and moderate), tree size (large vs. small trees), and bark shading was determined. Levels of some individual amino acids and total amino acid content in moderately infested trees were significantly higher than in uninfested trees. Large uninfested beech trees, which tend to be more readily infested by scale, had significantly higher concentrations of aspartic acid. Concentrations of phenol in the outer bark of both infested and uninfested trees were significantly higher than in the inner bark. However in bark infected by Nectria, phenol levels in the inner bark were significantly higher than in the outer bark.     

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.     

Chronic cork oak decline and water status: new insights

Chronic decline and Sudden death are two syndromes of cork oak (Quercus suber) dieback. Mortality is associated with water stress, but underlying physiological mechanisms are poorly understood. Here, we investigated the physiological performance of declining trees during the summer drought. Leaf water potential, gas-exchange, fluorescence of photosystem II and leaf and root starch concentration were compared in healthy (asymptomatic) and declining trees. Low annual cork increment in declining trees indicated tree decline for several years. All trees showed similar water status in spring. In summer, declining trees showed lower predawn leaf water potential (?2.0 vs. ?0.8 MPa), but unexpectedly higher midday leaf water potential than healthy trees (?2.8 vs. ?3.3 MPa). The higher midday water potential was linked to by means of strongly reduced stomatal conductance and, consequently, transpiration. This study is pioneer showing that declining trees had high midday water potential. A tendency for lower sap flow driving force (the difference between predawn and midday water potential) in declining trees was also associated with reduced photosynthesis, suggesting that chronic dieback may be associated with low carbon uptake. However, starch in roots and leaves was very low and not correlated to the health status of trees. Declining trees showed lower water-use efficiency and non-photochemical quenching in summer, indicating less resistance to drought. Contrarily to chronic decline, one tree that underwent sudden death presented predawn leaf water potential below the cavitation threshold.     

Tree recovery during the aftermath of an outbreak episode of the Hungarian spruce scale in southern Sweden

In 2010, the first, and so far only, infestation of the Hungarian spruce scale (Physokermes inopinatus) and accompanying sooty mould occurred in Scania, southernmost Sweden. About 1000?ha of Norway spruce (Picea abies) were affected, and the trees suffered from the sucking of the insects as well as from the dense sooty mould that covered the needles. Salvage cuttings were carried out in many of the massively attacked forest stands, both in response to the fear that the trees otherwise would die, e.g. from secondary bark beetle attacks and to prevent spreading of the infestation. The aim of this study was to provide basic, quantitative knowledge on the aftermath response of trees that were heavily infested, but not exposed to salvage cutting. Growth characteristics, in terms of needle weight, shoot length and tree-ring size were measured on infested and uninfested trees to compare and contrast the spruce growth before, during and after the scale outbreak. The infestation resulted in dwarf annual shoots, stunted needles and thin tree rings. The needle weight returned to normal the following year, whereas shoot length and tree rings required one growing season before full recovery.     

Radial growth changes following hemlock woolly adelgid infestation of eastern hemlock

? Context

Hemlock woolly adelgid (Adelges tsugae) is an invasive insect that is defoliating and killing eastern hemlock (Tsuga canadensis) in the USA.

? Aims

We quantified changes in tree-ring growth rates and wood anatomy for living trees infested with hemlock woolly adelgid across six sites from Massachusetts (42°41′N) to Georgia (34°53′N) to identify growth responses of eastern hemlock that had survived infestation.

? Methods

Annual ring widths from infested eastern hemlocks were cross-dated and measured. Growth rates before and after infestation were compared. Two infested trees from Virginia were cut, and thin sections were prepared to identify changes in cell properties.

? Results

At three sites, trees experienced a significant decrease in radial growth after hemlock woolly adelgid arrival; however, the other three sites showed no change or increase in growth. Latewood produced after hemlock woolly adelgid infestation had significantly smaller cells with reduced cell wall thickness compared to latewood prior to infestation.

? Conclusion

At half the sites where hemlock woolly adelgid infested eastern hemlock trees were sampled, radial growth increased or remained unchanged. This unexpected response may be due to reduced competition due to mortality of other eastern hemlocks or physiological compensatory responses of increased photosynthetic rate and increased water use efficiency experienced by eastern hemlock infested with hemlock woolly adelgid.     

Development of tree hollows in pedunculate oak (Quercus robur)

Many invertebrates, birds and mammals are dependent on hollow trees. For landscape planning that aims at persistence of species inhabiting hollow trees it is crucial to understand the development of such trees. In this study we constructed an individual-based simulation model to predict diameter distribution and formation of hollows in oak tree populations. Based on tree ring data from individual trees, we estimated the ages when hollow formation commences for pedunculate oak (Quercus robur) in southeast Sweden. At ages of about 200–300 years, 50% of the trees had hollows. Among trees <100 years old, less than 1% had hollows, while all >400-year-old trees had hollows. Hollows formed at earlier ages in fast-growing trees than in slow-growing trees, which may be because hollows are formed when big branches shed, and branches are thicker on fast-growing trees in comparison to slow-growing trees of the same age. The simulation model was evaluated by predicting the frequency of presence of hollows in relation to tree size in seven oak stands in the study area. The evaluation suggested that future studies should focus on tree mortality at different conditions. Tree ring methods on individual trees are useful in studies on development of hollow trees as they allow analysis of the variability in time for hollow formation among trees.     

Effects of past growth trends and current water use strategies on Scots pine and pubescent oak drought sensitivity

Drought-induced decline is affecting Pinus sylvestris populations in southern Europe, with very little impact on the more drought-tolerant Quercus pubescens. Although multiple studies have investigated interspecific differences in water use and growth strategies, the link between these two processes and how they vary within drought-exposed populations remains poorly understood. Here, we analysed tree ring and sap flow data from P. sylvestris and Q. pubescens stands in the Pyrenees in order to (1) evaluate differences in climate–growth responses among species, (2) disentangle the role of past growth trends and water use strategies in individual trees drought sensitivity and (3) assess whether such intraspecific patterns vary between species. Both species have suffered recent climatic constraints related to increased aridity. However, the effects of past growth trends and current water use traits on drought sensitivity varied among them. Initially, fast-growing ‘drought-sensitive’ pines displayed a higher gas exchange potential but were more sensitive to evaporative demand and soil moisture. They also showed lower water use efficiency for growth (WUE_BAI) and current growth decline. In contrast, initially, slow-growing ‘drought-tolerant’ pines showed the opposite water use traits and currently maintain the highest growth rates. In comparison, neither current WUE_BAI nor recent growth trends varied across Q. pubescens climate–growth groups. Nonetheless, ‘drought-sensitive’ oaks showed the lowest gas exchange potential and the highest growth rates under milder conditions. Our results show a strong effect of past growth trends and current water use strategies on tree resilience to increased aridity, which is more evident in P. sylvestris.     

Effects of climate on radial growth of Norway spruce and interactions with attacks by the bark beetle Dendroctonus micans (Kug., Coleoptera: Scolytidae): a dendroecological study in the French Massif Central

Samples of Norway spruce (Picea abies [L.] Karst) were dendrochronologically investigated in order to detect infestations by Dendroctonus micans (Kug.), the great spruce bark beetle (Col. Scolytidae), a relatively recent introduction to France. Uninfested natural forests located in the north-eastern French Alps and heavily infested plantations in the Ardèche region (Massif Central) were compared. The penetration holes bored in trunks by the bark beetle induced visible marks on wood, such as extreme ring width reductions, locally missing rings and crescent-shaped resin patches between consecutive rings that make possible a post-infestation dating.The outbreak began in 1979, 5 years prior to first insect visual detection by foresters. In the infested forest, tree basal area growth was not as sustained as in uninfested natural stands, but showed an inflection point at an unusually young tree age (from 30 to 40 years). Ring widths showing extreme synchronous radial growth reductions were caused either by excessively cold periods (e.g. in 1948, 1980, 1984, 1992) or by summer drought (as in 1986). Most of these weak growth years were shared with uninfested sites. In healthy forests, the consequences of extremely cold years were usually recorded only in high elevation stands, especially near the timberline, whereas summer drought effects were mostly visible in low altitude forests. By contrast, both phenomena were recorded in the infected Ardèche plantation. An analysis of tree-rings and monthly climate confirmed that Norway spruce growth in Ardèche plantations was reduced by excessively low minimum temperature during most parts of the year prior to ring formation, by higher than average maximum temperature during current spring and summer, and by drought in winter, spring and summer. Thus, the regional Ardèche climate with both cold winters and dry summers (especially in July) seems to weaken spruce trees planted there. Moreover, tree sensitivity to climate was found to be greatly enhanced by insect infestation. Such interactions between climatic stress and insect outbreak led to forest dieback in a 15–20-year period, when trees were still young (less than 70 years), and without any tree recovery. Therefore, in that region spruce plantations should be replaced by non-host species of Dendroctonus micans, especially where soil conditions may exacerbate drought effects.     

Shrub species affect distinctively the functioning of scattered Quercus ilex trees in Mediterranean open woodlands

A low tree stand density has been showed as necessary to thrive with summer drought in semiarid Mediterranean open woodlands. Shrub encroachment of these open woodlands is currently recommended to guarantee the persistence of the system, due to the nursery effect of shrubs on tree seedling. However, the increase in abundance and cover of a shrub understory in these water limited woodlands could bring consequences to tree overstory functioning. The present study analyzes the physiological status of scattered Quercus ilex L. trees in paired adjacent plots with and without the presence of a shrubby understory in CW Spain. Two contrasting shrub strategies were addressed in order to take into account possible species-specific effects: a dense-shallow rooting shrub (Cistus landanifer L.) and a sparse-deep rooting shrub (Retama sphaerocarpa (L.) Boiss). Leaf water potential (at predawn and midday), leaf gas exchange parameters (net photosynthetic rate and stomatal conductance), leaf nitrogen content and chlorophyll fluorescence transients (maximum photochemical efficiency and performance index, sensuStrasser et al., 2004) were measured during three consecutive summers. Trees growing with Cistus as understory showed significant lower leaf water potential, leaf gas exchange parameters, leaf nitrogen content and chlorophyll photochemical efficiency than trees growing without shrub competence. However, the presence of the legume Retama did not affect significantly the physiological state of Q. ilex. Thus, we conclude that the presence of a shrubby understory has the potential to modify the functioning of scattered trees, but these effects are species-specific.     

Susceptibility of ponderosa pine, Pinus ponderosa (Dougl. ex Laws.), to mountain pine beetle, Dendroctonus ponderosae Hopkins,attack in uneven-aged stands in the Black Hills of South Dakota and Wyoming USA

Mountain pine beetle, Dendroctonus ponderosae Hopkins can cause extensive tree mortality in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests in the Black Hills of South Dakota and Wyoming. Most studies that have examined stand susceptibility to mountain pine beetle have been conducted in even-aged stands. Land managers increasingly practice uneven-aged management. We established 84 clusters of four plots, one where bark beetle-caused mortality was present and three uninfested plots. For all plot trees we recorded species, tree diameter, and crown position and for ponderosa pine whether they were killed or infested by mountain pine beetle. Elevation, slope, and aspect were also recorded. We used classification trees to model the likelihood of bark beetle attack based on plot and site variables. The probability of individual tree attack within the infested plots was estimated using logistic regression. Basal area of ponderosa pine in trees ≥25.4 cm in diameter at breast height (dbh) and ponderosa pine stand density index were correlated with mountain pine beetle attack. Regression trees and linear regression indicated that the amount of observed tree mortality was associated with initial ponderosa pine basal area and ponderosa pine stand density index. Infested stands had higher total and ponderosa pine basal area, total and ponderosa pine stand density index, and ponderosa pine basal area in trees ≥25.4 cm dbh. The probability of individual tree attack within infested plots was positively correlated with tree diameter with ponderosa pine stand density index modifying the relationship. A tree of a given size was more likely to be attacked in a denser stand. We conclude that stands with higher ponderosa pine basal area in trees >25.4 cm and ponderosa pine stand density index are correlated with an increased likelihood of mountain pine beetle bark beetle attack. Information form this study will help forest managers in the identification of uneven-aged stands with a higher likelihood of bark beetle attack and expected levels of tree mortality.     

Drought tolerance and transplanting performance of holm oak (Quercus ilex) seedlings after drought hardening in the nursery

Drought stress is the main cause of mortality of holm oak (Quercus ilex L.) seedlings in forest plantations. We therefore assessed if drought hardening, applied in the nursery at the end of the growing season, enhanced the drought tolerance and transplanting performance of holm oak seedlings. Seedlings were subjected to three drought hardening intensities (low, moderate and severe) for 2.5 and 3.5 months, and compared with control seedlings. At the end of the hardening period, water relations, gas exchange and morphological attributes were determined, and survival and growth under mesic and xeric transplanting conditions were assessed. Drought hardening increased drought tolerance primarily by affecting physiological traits, with no effect on shoot/root ratio or specific leaf mass. Drought hardening reduced osmotic potential at saturation and at the turgor loss point, stomatal conductance, residual transpiration (RT) and new root growth capacity (RGC), but enhanced cell membrane stability. Among treated seedlings, the largest response occurred in seedlings subjected to moderate hardening. Severe hardening reduced shoot soluble sugar concentration and increased shoot starch concentration. Increasing the duration of hardening had no effect on water relations but reduced shoot mineral and starch concentrations. Variation in cell membrane stability, RT and RGC were negatively related to osmotic adjustment. Despite differences in drought tolerance, no differences in mortality and relative growth rate were observed between hardening treatments when the seedlings were transplanted under either mesic or xeric conditions.     

Crown transparency, tree mortality and stem growth of Pinus sylvestris, and colonization of Tomicus piniperda after an outbreak of Gremmeniella abietina

In the year 2000, large areas of forest in Sweden, mainly 30-50 year old Pinus sylvestris (L.) stands, were attacked by the fungus Gremmeniella abietina (Lagerb.) Morelet. The aims of this study were to investigate: (i) the relationship between G. abietina-induced tree crown transparency (CT) and P. sylvestris (L.) tree mortality; (ii) the influence of CT levels on stem growth; (iii) the recovery of the crown; and (iv) the association of CT and colonization by Tomicus piniperda (L.). Thirty-five permanent sample plots were established in five P. sylvestris stands (38-46 years old), infested by G. abietina, and 23 plots in four reference stands, not obviously infested.During the 5 years following the attack, the total mortality amounted to 454 trees ha⁻¹ and 7.8 m² ha⁻¹, on average, in the five infested stands, corresponding to 42% of the trees and 34% of the basal area at the time of the attack. Most of the mortality occurred within 2 years of the attack. The mortality of individual trees (2002-2005) was found to be related to the crown transparency (CT), the position of needle loss within the crown and the tree diameter at breast height. Based on our modeling, the probability of mortality was substantially increased if the initial CT-value was higher than 85%.Growth reductions were detected for individual trees with an initial CT of >c. 40%. In contrast, trees with a low initial CT (<c. 40%) were not affected and even exhibited increased growth. In the five infested stands, the reductions in basal area and volume increment were estimated to be 26-58%, and, 42-73%, respectively, during the five growing seasons after the attacks.The trees in the infested stands that were still alive in spring 2005 had started to recover in terms of CT. Breeding of T. piniperda on the P. sylvestris (L.) stems occurred almost exclusively on stems with a CT > 90%.The data from this study suggest that when a P. sylvestris (L.) stand has been attacked by G. abietina, trees with a CT above 80% should be felled; the remaining trees will have a high probability of survival and resistance to successful breeding by the T. piniperda.     

Climate and groundwater effects on the establishment, growth and death of Prosopis caldenia trees in the Pampas (Argentina)

Semiarid woodlands dominated by Prosopis caldenia thrive at the dry edge of the Argentinean Pampas. Deforestation and increased precipitation have driven sustained water table level rise in the region that are likely to affect the dynamics of remnant woody vegetation patches. Here we analyze the effect of climate and groundwater level on the establishment, growth and death of P. caldenia located on lowland (current water table <0.5 m deep) and upland (current water table 8 m deep) positions within rolling sandy landscape. Standard dendrochronological techniques were applied on 98 cross sections and cores of P. caldenia trees. Results allowed us to estimated that trees in the lowland established a few years earlier than in the upland (1929 vs. 1936) and died between 1991 and 2002, while trees in higher positions are still alive. As a result of a faster growth in the lowland, maximum mean basal area increment took place earlier (1950s vs. 1990s) and achieved a higher mean value than in the upland (41.63 vs. 37.41 mm²). While mean annual chronologies were not associated across stands, an opposite highly significant association was found for the mean growth trends suggesting long-term effects of water table depths on growth. We found a different association between mean annual growth and climate, with lowland trees showing a negative response to precipitation before and during the growing season, and upland trees displaying a positive response to summer rainfall inputs. Temperature at the end of the growing season had a positive effect in the lowland whereas temperature during the growing season had a positive effect in the upland. These results show how groundwater can induce diverging sensitivity of forest growth and survival to climate variability, enhancing growth at optimum depths (2-8 m in our study) but depressing it or even killing trees when it approaches the surface (<2 m in our study). Groundwater dynamic should be considered in forest management and conservation strategies in semiarid woodlands in Central Argentina.     

Fire history and understorey vegetation: Water and nutrient relations of Eucalyptus gomphocephala and E. delegatensis overstorey trees

Fire regimes in temperate forests and woodlands have changed significantly in Australia since European settlement. We hypothesised that an absence of fire leads to the increased development of woody understorey/midstorey and that this may be correlated with decreased water and/or nutrient availability in overstorey temperate eucalypts currently declining in health. Sites with a history of being long unburnt or recently (and frequently in the case of Eucalyptus gomphocephala woodland) burnt (relative to median fire intervals for the vegetation type) were established in E. gomphocephala woodland in Western Australia and in Eucalyptus delegatensis forest in Tasmania. In long unburnt sites in both E. gomphocephala woodland and E. delegatensis forest, there was greater percent cover of understorey/midstorey and eucalypts had higher water use efficiency, indicative of greater soil water limitation, as estimated by foliar carbon isotope ratios. In E. gomphocephala woodland foliar Cu and Zn were significantly lower in eucalypts of long unburnt, relative to frequently burnt, sites. In E. gomphocephala woodland, understorey/midstorey (shrub) cover was positively correlated, and foliar copper and zinc levels were negatively correlated to health of overstorey trees. In E. delegatensis forest foliar phosphorus (P) was significantly lower in eucalypts of long unburnt, relative to recently burnt, sites. In E. delegatensis forest moss cover was positively correlated and foliar P was negatively correlated to health of overstorey trees. The understorey/midstorey that develops in the long absence of fire may alter ecological processes that lead to less favourable water- and nutrient-relations in E. gomphocephala woodland and E. delegatensis forest that are associated with decline in crown health. However this study does not definitively show a link between understorey/midstorey vegetation and overstorey tree water- and nutrient-relations. This link will be investigated in future research.     

Incidence of Phytophthora species in oak forests in Austria and their possible involvement in oak decline

A survey on the occurrence of Phytophthora species in oak ecosystems in Austria was conducted from April to May 1999 and in June 2000. The investigations were carried out at 35 study sites distributed throughout the zone of oak forests in eastern Austria. Four oak species, including Quercus robur, Q. petraea, Q. cerris and Q. pubescens were considered in the survey. Rhizosphere soil samples were taken from sample trees, which consisted of healthy and declining trees as indicated by their crown transparency. Young oak leaflets were used as baits to recover Phytophthora species. The assemblage of Phytophthora spp. detected in Austrian oak forests consisted of five species, including Phytophthora quercina, P. citricola, P. gonapodyides, P. europaea and P. syringae. P. quercina and P. citricola were isolated from 11 and seven sites, respectively, and were thus the most common and most widely distributed species. The three other species were recovered only sporadically. P. citricola could be separated into two morphologically and genetically well‐characterized types (A and B). Phytophthora species, in particular the common P. quercina and P. citricola occurred on sites showing a wide variety of soil types, soil textures and moisture classes. There was mild evidence for connection between deteriorating crown status and the presence of Phytophthora spp. Furthermore, significant differences in contents of magnesium, as well as calcium, aluminium, nitrogen and carbon at different soil depths (0–10, 10–20 and 20–40 cm) were detected between Phytophthora‐infested and Phytophthora‐free sites. The results of the present study provide circumstantial evidence that Phytophthora species are involved in oak decline at certain sites in Austria.     

Soil bacterial functional diversity is associated with the decline of Eucalyptus gomphocephala

This study investigates whether tree decline in Eucalyptus gomphocephala (tuart) is associated with the functional diversity of soil bacterial communities. We selected 12 sites with different stages of decline and assessed crown health [Crown density (CD), Foliage transparency (FT), Uncompacted live crown ratio (ULCR), Crown dieback ratio (CDR) and Epicormic index (EI)] and soil bacterial functional diversity based on Biolog EcoPlates™ incubation [Average well colour development (AWCD), Shannon diversity (H′), richness (S) and Shannon evenness (E)]. Crown health indices differed between sites with EI being the most robust indicator of decline in crown health followed by CDR and CD (P < 0.05). Soil bacterial indices collected at 0–10 and 20–30 cm soil depth between December (summer, dry season) and May (autumn, start of wet season) differed between sites (P < 0.05), and significant relationships between crown health indices, except ULCR, and all soil bacterial indices were observed. Principle component analysis (PCA) showed that a decrease in the utilization of carbohydrates, carboxylic acids, amino acids and amines by the soil bacterial communities correlated to sites with poor crown health, indicating some changes in physiological responses of bacterial groups with declining tree health. Using stepwise regression analyses, in the 0–10 cm soil layer in December, itaconic acid had a 46% contribution to the EI. Carboxylic acids, including itaconic acid, have a strong ability to solubilize soil minerals in calcareous soil, and these possibly increased the availability of soil mineral nutrients in the healthier sites compared to the declining sites, particularly in the dry season. In addition, lack of soil water in the declining sites limited soil bacterial diversity and was positively correlated with EI in the 0–10 cm soil layer in December. In conclusion, soil bacterial functional diversity has a strong relationship with tuart decline and the importance of soil microbes in tuart ecosystem health must be considered in the future.     

Spatial estimation of the density and carbon content of host populations for Phytophthora ramorum in California and Oregon

Outbreak of the emerging infectious disease sudden oak death continues to threaten California and Oregon forests following introduction of the exotic plant pathogen Phytophthora ramorum. Identifying areas at risk and forecasting changes in forest carbon following disease outbreak requires an understanding of the geographical distribution of host populations, which is unknown. In this study, we quantify and map the population density and carbon contents of five key host species for P. ramorum in California and Oregon, including four hosts killed by the pathogen (Notholithocarpus densiflorus, Quercus agrifolia, Quercus kelloggii and Quercus chrysolepis) and the foliar host Umbellularia californica which supports high sporulation rates. We integrate multiple sources of vegetation data, assembled from sparsely distributed (regional-scale) forest inventory and analysis (FIA) plots and more densely distributed (landscape-scale) plots for monitoring sudden oak death, and develop spatial prediction models based on correlation with environmental variables and spatial dependencies in host abundance. We estimate that 1.8 billion N. densiflorus trees (68 Tg C) and 2.6 billion Quercus host trees (227 Tg C) occur across 3.9 and 17.7 million ha of their respective habitat. A total of 436 million U.californica trees (14 Tg C) occur across 4.2 million ha which frequently overlap with Quercus and N. densiflorus host populations. Combination of landscape-scale data with FIA data resulted in more accurate estimation of host populations and their carbon contents. Forests of northern California and southwest Oregon have the highest concentration of the most susceptible hosts along with climatic conditions that favor pathogen spread. This study represents the first spatially-explicit estimate of P. ramorum host populations and their carbon contents which exceed previously published estimates. Our results will inform landscape- to regional-scale models of disease dynamics and guide management decisions regarding ecosystem impacts including risk of C release following widespread tree mortality.