Comparison of the Susceptibility of Quercus petraea, Q. robur and Q. rubra to Collybia fusipes

Collybia fusipes is the cause of a root rot of Quercus petraea (sessile oak), Q. robur (pedunculate oak) and Q. rubra (red oak). This parasite is often reported to cause problems in stands of red oaks and field evidence suggests that this North American oak species is more susceptible than the two European oak species. Young saplings of the three oak species and of chestnut, Castanea sativa, in the glasshouse, and also mature sessile and red oaks growing in the same stand were inoculated with C. fusipes to compare their susceptibility. Red oak, both as young seedlings and mature trees, was more susceptible to C. fusipes than sessile oak. Chestnut seedlings were as susceptible as sessile oak. Susceptibility of pedunculate oak seedlings was intermediate between red oak and sessile oak. In one experiment this species was significantly less susceptible than red oak, and in the other it was as susceptible.     

Sudden Oak Death, A New Forest Disease in California

A forest disease caused by a newly described pathogen, Phytophthora ramorum, is presently affecting a number of woody plant species in central and northern coastal California and southern Oregon, U.S.A. Among the most severely affected tree species are coast live oak, Quercus agrifolia, and tanoak, Lithocarpus densiflorus. In March 2000, we established plots in Marin County, CA, to assess symptom progression and the association of other organisms with diseased trees. Symptomatic trees exhibit 'bleeding' of viscous sap from apparently intact bark, typically within approximately 2 m of the soil. At least three species of bark and ambrosia beetles colonize bleeding oaks. These beetles colonized every bleeding tree that died during the first year of the study (N = 23) while the foliage was still green. By March 2001, numbers of symptomatic and dead trees increased for both coast live oak and tanoak. Symptomatic coast live oaks totaled 26% in March 2000 and 27% in March 2001. During the same period, mortality increased from 6% to 10%. For tanoaks, 41% were symptomatic in 2000, rising to 49% in 2001. Mortality was 11% in 2000 and 13% in 2001. The values for coast live oak are broadly consistent with independently acquired infection and mortality estimates derived from an unbiased transect method.     

Relationships between health of Quercus robur, occurrence of Phytophthora species and site conditions in southern Sweden

The effect of Phytophthora species, soil chemistry, precipitation and temperature on the vitality of oak was evaluated in 32 oak stands in southern Sweden. In addition, the relationship between the occurrence of Phytophthora species and soil conditions was determined. The results showed that there was a weak association between the presence of P. quercina , the most frequently recovered Phytophthora species in southern Sweden, and the vitality of the oak stands (determined from estimates of crown defoliation of individual trees). The pathogens occurred more frequently in clayey and loamy soils that were less acidic and which had higher base saturation. However, they were found in all but the most acidic soils (pH < 3·5). In stands where Phytophthora species were not present, positive correlations between the average crown defoliation and proportion of damaged trees with average summer precipitation and average annual precipitation were found. There were no significant differences in soil chemistry between healthy and declining stands included in this study, and no significant correlations were found between any soil parameter and crown vitality. Based on the results from these 32 oak stands, it is likely that the decline of oaks in southern Sweden can be attributed to several different site-specific factors, such as infection by P. quercina or unusual weather events, which interact with a number of biotic and abiotic factors, leading to oak decline.     

Root infection by Phytophthora cinnamomi in seedlings of three oak species

The combination of soil infestation with Phytophthora cinnamomi and repetitive flooding was studied on 1-year-old plants of Quercus ilex (holm oak) and Q. suber (cork oak). In a second experiment, using 2-year-old plants of the same species and of red oak ( Q. rubra ), the soil infestation was followed by two drought-rewatering cycles. Oak predawn leaf water potential (PLWP) and stomatal conductance (gs) were monitored during both experiments. Root infection, root loss, wilting and mortality were assessed at the end of the experiments. Q. ilex exhibited the highest susceptibility to P. cinnamomi , and Q. rubra the lowest. Root infections caused by P. cinnamomi were more severe in the flooding than in the drought experiment. The most noticeable effect of the infection on plant water relations was a decrease in stomatal conductance. This occurred at different times after inoculation, varying with species susceptibility and experiment. Inoculation with P. cinnamomi induced a decrease of PLWP in Q. ilex plants, and in some Q. suber plants exhibiting a severe root loss. The results further showed that the relationship between PLWP and gs was modified by infection with P. cinnamomi . The combination of flooding and infection with P. cinnamomi acted synergistically on the water relations of Q. ilex . By contrast, there was no significant increase in disease severity due to the postinoculation water stress imposed on the oaks.     

Interaction between root rot basidiomycetes and Phytophthora species on pedunculate oak

Forest declines are usually complex multifactorial phenomena that involve interactions between different factors. The possible interaction between different types of mycelial pathogens was investigated through artificial inoculation of oak seedlings, involving two root rot basidiomycetes, Collybia fusipes and Armillaria mellea, and two Phytophthora species, P. cinnamomi and P. cambivora. These pathogens were inoculated onto young Quercus robur saplings in greenhouse conditions, either alone or combining a root rot basidiomycete with a Phytophthora species. Three out of the four Phytophthora spp.*root rot basidiomycete combinations tested resulted in significantly greater damage to the oak host than the sum of the damages induced by the individual pathogens. This positive interaction could be significant in oak decline syndrome.     

Effects of pear tree physiology on fire blight progression in perennial branches and on expression of pathogenicity genes in Erwinia amylovora

The interaction between Erwinia amylovora (the causal agent of fire blight) and the physiological status of pear trees was examined under orchard conditions. The physiological status of the trees was defined qualitatively, using host phenology and vigour as measures, and quantitatively, using the sorbitol content in annual shoots as a measure. Qualitatively, tree response to fire blight was governed by phenological stage at the time of infection and vigour: low vigour trees inoculated in the autumn (just before entering dormancy) and high vigour trees inoculated in the spring (soon after bloom) were more susceptible than high vigour trees inoculated in the autumn and low vigour trees inoculated in the spring. Quantitatively, the rate of symptom progression in perennial branches (SPR) was significantly (P ≤ 0.001) correlated to the absolute value of the rate of sorbitol content change (|SCR|). The relationship between hrp genes expression of transformed E. amylovora (estimated according to hrpE and hrpJ expression) and |SCR| was determined on 1 year-old trees. Expression of hrp genes was significantly correlated with |SCR| (P = 0.004) and 63.5% of the variability in the hrp genes expression was attributed to |SCR| values. The expression of hrp genes increased gradually and asymptotically with increasing |SCR| values; further increase in |SCR| did not affect the expression.     

Red Oak Borer, Enaphalodes rufulus (Coleoptera: Cerambycidae), in the Ozark Mountains of Arkansas, U.S.A.: An Unexpected and Remarkable Forest Disturbance

A complex interaction of multiple factors has resulted in an oak decline event in oak-hickory forests of the Ozark Mountains of Arkansas and Missouri, U.S.A. The most striking feature of this situation is an unprecedented population explosion of red oak borer, a species of cerambycid beetle, Enaphalodes rufulus (Haldeman), which appears to be causing extensive mortality to mature red oaks (Quercus, subgenus Erythrobalanus). The insect is a native species, historically a minor pest of oaks, found throughout the eastern United States. Beetles normally reproduce in living oaks, as larvae initially feed in phloem tissue and subsequently bore into xylem where pupation occurs. The life cycle is two years in length and synchronous adult emergence occurs in odd-numbered years. Data from previous research indicate average attack densities of less than four per tree with a high of 71 on a single tree. Historical emergence densities are similarly low and the highest reported was 15 adults from one tree. Our research is concerned with understanding factors contributing to this outbreak; developing sampling methods for red oak borer; assessing oak mortality; and evaluating site and stand conditions associated with the current outbreak. Results of our initial sampling reveal dramatically higher average attack densities of 244 per m2 and emergence densities of 18 per m2 of bark surface area. We confirm a three-week period of adult emergence and activity during mid-June to early July. We also report on possible management responses by federal and state agencies to this remarkable epidemic and oak mortality crisis.     

The oak gallwasp <Emphasis Type="Italic">Aphelonyx persica:</Emphasis> a new record from Turkey,with some new host records

A new record is reported of the oak gallwasp Aphelonyx persica on oaks (Quercus cerris, Q. ithaburensis, Q. trojana) in the Inland Aegean Region of Turkey. This species is recorded for the first time on Q. cerris and Q. trojana. Details are given of the geographical distribution, hosts and phenology.     

Phytophthora species in oak ecosystems in Turkey and their association with declining oak trees

From 1999 to 2001, a survey on the occurrence of Phytophthora spp. in the rhizosphere soil of healthy and declining oak trees was conducted in 51 oak stands in Turkey. Seven Phytophthora spp. were recovered from six out of the nine oak species sampled: P .  cinnamomi , P .  citricola , P .  cryptogea , P .  gonapodyides , P .  quercina , Phytophthora sp. 1 and Phytophthora sp. 2. The most frequently isolated species, P .  quercina , was very common on slopes susceptible to drought. It occurred in four different climatic zones and on six Quercus spp., suggesting that it is native to oaks. The second most common species, P .  citricola , was separated into three subgroups: type C was recovered only in Anatolia, whereas A and B occurred only in the European part of Turkey. Phytophthora cinnamomi was recovered at one site only, and may not be involved in oak decline in Turkey. The other four species were recovered sporadically. On affected sites there was a significant association between deteriorating crown status and the presence of Phytophthora spp., particularly P .  quercina . The occurrence of Phytophthora species was significantly influenced by soil pH. Stem inoculation tests on oak seedlings revealed that Q .  petraea was the most susceptible species.     

Efficacy of phosphonic acid, metalaxyl-M and copper hydroxide against Phytophthora ramorum in vitro and in planta

The ability of metalaxyl-M, phosphonic acid in the form of phosphonate, and copper hydroxide to inhibit different stages in the life cycle of Phytophthora ramorum , the causal agent of sudden oak death (SOD), was tested in vitro using 12 isolates from the North American forest lineage. In addition, experiments were conducted in planta to study the ability of phosphonic acid injections and metalaxyl-M drenches to control pathogen growth on saplings of California coast live oak ( Quercus agrifolia ), and of copper hydroxide foliar sprays to control infection of California bay laurel ( Umbellularia californica ) leaves. Phytophthora ramorum was only moderately sensitive to phosphonic acid in vitro , but was highly sensitive to copper hydroxide. In planta experiments indicated the broad efficacy of phosphonic acid injections and of copper hydroxide sprays in preventing growth of P. ramorum in oaks and bay laurels, respectively. Finally, although metalaxyl-M was effective in vitro , drenches of potted oak trees using this active ingredient were largely ineffective in reducing the growth rate of the pathogen in planta .     

Impact of <Emphasis Type="Italic">Erysiphe alphitoides</Emphasis> on transpiration and photosynthesis in <Emphasis Type="Italic">Quercus robur</Emphasis> leaves

Oak powdery mildew, (Erysiphe alphitoides) causes one of the most common diseases of oaks. We assessed the impact of this pathogen on photosynthesis and water relations of infected leaves using greenhouse-grown oak seedlings. Transpiration of seedlings infected by oak powdery mildew was also investigated. Altogether, E. alphitoides had a low impact on host gas exchange whether at the leaf or whole plant scale. Maximal stomatal conductance of infected leaves was reduced by 20–30% compared to healthy controls. Severely infected seedlings did not experience any detectable change of whole plant transpiration. The reduction in net CO₂ assimilation, A_n, was less than proportional to the fraction of leaf area infected. Powdery mildew reduced both the maximal light-driven electron flux (J_max) and the apparent maximal carboxylation velocity (Vc_max) although Vc_max was slightly more impacted than J_max. No compensation for the infection occurred in healthy leaves of partly infected seedlings as the reduced photosynthesis in the infected leaves was not paralleled by increased A_n levels in the healthy leaves of the seedlings. However, E. alphitoides had a strong impact on the leaf life-span of infected leaves. It is concluded that the moderate effect of E. alphitoides on oak might be related to the small impact on net CO₂ assimilation rates and on tree transpiration; nevertheless, the severe reduction in leaf life-span of heavily infected leaves may lead to decreased carbon uptake over the growth season.     

Involvement of soilborne Phytophthora species in Central European oak decline and the effect of site factors on the disease

A survey was made on the occurrence of soilborne Phytophthora species in 35 oak stands on a range of geologically different sites in Bavaria. The most widespread species were P. quercina , P. cambivora and P. citricola . Seven other Phytophthora species were isolated infrequently. The fine root systems of 106 healthy and 111 declining mature trees of Quercus robur and Q. petraea were intensively investigated. The results indicate that, depending on the site conditions, at least two different complex diseases are referred to under the name 'oak decline'. On sites with a mean soil pH (CaCl₂)  3·5 and sandy-loamy to clayey soil texture Phytophthora spp. were commonly isolated from rhizosphere soil, and highly significant correlations existed between crown transparency and various root parameters. Oaks with P. quercina or other Phytophthora spp. in their rhizosphere had markedly higher levels of fine root damage than oaks without Phytophthora spp., and were subject to a relative risk of severe crown symptoms of 2·1 and 2·8, respectively. In contrast, in stands with sandy to sandy-loamy soils and a mean soil pH  3·9, Phytophthora spp. were not found. In these stands, correlations between crown transparency and various root parameters were either less significant or not significant. It is concluded that Phytophthora species are strongly involved in oak decline on sandy-loamy to clayey sites with a mean soil-pH (CaCl₂)  3·5.     

Massenvermehrung von phyllophagen Schmetterlingen in Laubw?ldern des Rheinlands

Zusammenfassung  In den Jahren 1996 und 1997 wurde in ausgew?hlten Gebieten des Rheinlands die Populationsdichte von Schmetterlingsraupen erfa?t, die an Eichen (Quercas robur undQ. petraea) und an Hainbuchen (Carpinus betulus) Kahlfra?, an Rotbuchen (Fagus sylvatica) erheblichen Fra?schaden verursachten. An dem Blattfra? waren insgesamt 63 Schmetterlingsraupen beteiligt. An Eichen erfolgte Kahlfra? gleichzeitig durch mehrere Arten (Operophtera brumala, O. fagata, Erannis defoliaria undTortrix viridana), w?hrend an Hainbuche und RotbucheOperophtera fagata für den hohen Fra?schaden verantwortlich war. Standorte, an denen Massenvermehrungen nachgewiesen wurden, waren gut mit N?hrstoffen versorgt und best?tigen so die Hypothese zur Ressourcenverfügbarkeit. Die Populationsdichten der Schmetterlingsraupen waren an Eichen so gro?, da? es zu inter- und intraspezifischer Konkurrenz (scramble competition) kam. In Eichenbest?nden, die mit Rotbuchen unterbaut waren, wurde die Konkurrenzsituation unter den Schmetterlingsraupen vermindert. Somit k?nnen waldbauliche Ma?nahmen, besonders bei einer Massenvermehrung vonOperophtera fagata, zu einer Verl?ngerung der Gradation beitragen.

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Mass occurrence of phyllophagous caterpillars in the deciduous forests of the Rhineland

We investigated species assemblages of Lepidoptera which fed onQuercus petraea, Carpinus betulus andFagus sylvatica trees growing in commercial forests in the Rhineland (Germany). In total we found 63 different species of caterpillars. Several species (Operophtera fagata, O. brumata, Tortrix viridana, Agriopis leucophaearia, A. aurantiaria andErannis defoliaria) appeared in high abundances on oak trees, which they together defoliated. Defoliation ofCarpinus betulus as well as high feeding damages onFagus sylvatica were caused mainly by a single species,Operophtera fagata. From our observations of individual caterpillars and their extraordinarily high feeding damages (=45%) onFagus sylvatica trees which were growing in the shade of oak trees, in comparison to individualF. sylvatica trees not growing beneath oak trees (feeding damage= 7%), we made the following conclusions: Firstly, in the case of mass occurrence a severe influence of scramble competition occurs between caterpillars. Secondly, a prolongation of gradation of polyphagous caterpillars is caused when stratification byFagus sylvatica living as undergrowth of oak trees occurs. Defoliated trees occurred in sites with enhanced nutrient availability. This supports the resource availability hypothesis. Although there was a surplus of nutrients in the soil and within the roots we found a deficiency of P in the leaves.

     

Apple proliferation resistance of <Emphasis Type="Italic">Malus sieboldii</Emphasis>-based rootstocks in comparison to rootstocks derived from other <Emphasis Type="Italic">Malus</Emphasis> species

In three trials carried out over a period of 24 years, open-pollinated seedlings of Malus sieboldii and M. sargentii and 22 apomictic rootstock selections with either M. sieboldii, M. sargentii or M. hupehensis in their parentage were examined for apple proliferation (AP) resistance in comparison to clonal M. x domestica-based rootstocks M 9, M 11, M 13, stocks of the B (Budagovski) and the Polish P series and M. robusta seedlings. Following experimental inoculation or natural infection the Golden Delicious-grafted trees on most of the M. sieboldii-derived progenies showed a high level of AP resistance expressed by low cumulative disease indices, a high percentage of non or little affected trees, low incidence of the small fruit symptom and non or little effect on vigour. Trees on M 9 and M 11, B 118 and M. robusta seedlings were moderately susceptible while trees on progenies with M. sargentii and M. hupehensis parentage, rootstocks of the P series, B 9, B 490 and M 13 proved highly susceptible. The screening also showed that rootstocks with M. sieboldii and M. sargentii parentage are often highly susceptible to latent apple viruses. Trees on most of the M. sieboldii-based progenies were more vigorous than trees on standard stock M 9, whereas the vigour of some progenies from selections with M. sargentii parentage was in the range of M 9 or even lower. Productivity was often correlated with the vigour.     

Effect of competition on spatial patterns of oak forests on the Chinese Loess Plateau

Reforestation or natural forest regeneration is an alternative measure for controlling soil erosion in degraded land on the Chinese Loess Plateau(CLP). However, our understanding of the temporal dynamics and the spatial patterns of forest regeneration remains inadequate. Two oak forests at different development stages were investigated to determine the spatial patterns of competitions(intraspecies and interspecies) during different successional stages. The intraspecies and interspecies spatial relationships among different tree diameters at breast height were analyzed at multiple scales by Kriging interpolation method and univariate and bivariate O-ring statistics. Our analytical results indicated that self-correlation and competition intensity were relatively high between oak and pine trees in the early development stage of oak forests due to their clumped distributions of heavy seeds. Birch trees had a lower competition in comparison to oak trees although birch was the dominant species. Therefore, asymmetric competition of oak trees was most likely to have led to their edge dispersal and their success in replacing the pioneer species. Asymmetric competition means that larger individuals obtained a disproportionately large share of the resources and suppressed the growth of smaller individuals. Kriging interpolation analysis showed a tendency towards homogenization caused by interspecies competition during the succession of oak forests. Our results demonstrated that the competition was the driving factor in the spatial distribution of oak forests on the CLP.     

Aerial Application of Bacillus thuringiensis (Dipel) against Colotois (Himera) pennaria L. in Comparison with ULV-Application of Various Insecticides 1)

In spring 1972 experiments were carried out in Lower Austria with the Bacillus thuringiensis isolate Dipel in order to assess its effectiveness in controlling Colotois pennaria L. infesting oak and other broad-leaved trees. The results obtained compared favourably with those produced by the insecticides Nexion²), malathion and Dipterex³), although both malathion and Dipterex yielded higher mortality. Similar results were obtained when B. thuringiensis was used to control Stilpnotia salicis L., feeding on poplar trees. Neodiprion sertifer Geoffr. and Pris-tiphora abietina Christ., however, were not affected.     

Prediction and Mapping of the Impact of Winter Temperature on the Development of Phytophthora cinnamomi-Induced Cankers on Red and Pedunculate Oak in France

ABSTRACT Phytophthora cinnamomi is the causal agent of a perennial canker that develops on the lower bole on northern red oak and pedunculate oak. The disease has a limited range in Europe, being reported only in southwest France. This limited distribution is probably linked to the susceptibility of P. cinnamomi to frost. A model was developed in previous work to estimate the impact of temperatures of <0 degrees C on the winter survival of P. cinnamomi in trunk cortical tissues and on the subsequent development of cankers. In this article, we report the use of this model to simulate canker development in 503 locations across France during a 30-year period. The predicted canker extension decreased sharply when the median P. cinnamomi winter survival index decreased from 0.95 to 0.65, with cankers that poorly developed when the median survival index was lower than 0.5 to 0.6. The actual incidence of the disease in 192 stands located across southwest France was compared with that of the model outputs. Both presence of disease in stands and frequency of cankered trees in infected stands, but not canker size on infected trees, were strongly related to the median P. cinnamomi survival index. No disease was present in stands with median survival index lower than 0.65, and the frequency of cankered trees in infected stands remained very low in stands with a median survival index between 0.65 and 0.70. Aspect was an additional factor explaining disease incidence, while the effect of elevation was likely due to its effect on winter temperatures. Maps of winter suitability to P. cinnamomi-induced cankers on oaks in France are presented.     

Early survival of Quercus ilex subspecies from different populations after infections and co‐infections by multiple Phytophthora species

Forests in Europe are threatened by increased diversity of Phytophthora species, but effects on trees of simultaneous infections by Phytophthora and ecological consequences of their coexistence are unknown. This study explored variation in early survival of Quercus ilex to Phytophthora infections and assessed interactions between Phytophthora species when trees were co‐infected. Three Phytophthora species (P. cinnamomi, P. gonapodyides and P. quercina), seeds from 16 populations of Q. ilex (ballota and ilex subspecies) and two infection times were used as sources of variation in two experiments. The influence of Phytophthora species, Q. ilex subspecies and populations on plant germination and survival were analysed using generalized linear mixed models and survival analysis techniques. Germination rates were not influenced by Phytophthora spp. (P = 0.194) but by the subspecies and populations of Q. ilex (P < 0.001). In Phytophthora‐infested soils, Q. ilex subsp. ilex germinated at higher rates than Q. ilex subsp. ballota. Plant survival was strongly influenced by Phytophthora species (P < 0.001), not by the subspecies and populations of Q. ilex. Seedling mortality was reduced and delayed if a less virulent Phytophthora species infected plants prior to infection by a more virulent Phytophthora species. The results help to explain oak decline syndrome and the lack of natural and artificial regeneration of Q. ilex forests. Lack of interspecific variability of early survival to Phytophthora spp. discourages direct sowing for artificial reforestation programmes. Large, thick seeds, giving plants rapid growth, are advantageous traits when soils are infested with Phytophthora spp.     

Ceratocystis species on Quercus robur with oak decline symptoms in southern Poland1

In southern Poland during 1986–1989, six species of Ceratocystis were found on sick and dead Quercus robur. The frequency of their occurrence in above-ground portions and roots of trees is given. The possibility is discussed whether oak decline, caused by a complex of abiotic and biotic factors, is intensified when trees are colonized by Ceratocystis.     

A multiplex real-time PCR assay enables simultaneous rapid detection and quantification of bacteria associated with acute oak decline

Acute oak decline (AOD) is a syndrome affecting mature oak trees and is characterized by stem bleeds from vertical fissures on trunks, and inner bark necrosis caused by a polybacterial consortium, in which Gibbsiella quercinecans and Brenneria goodwinii, and to a lesser extent Rahnella victoriana and Lonsdalea britannica, play key roles. Here we report a novel multiplex real-time PCR assay that enables simultaneous and rapid detection and quantification of these four bacterial species from stem bleed swabs. Experiments with axenic cultures were performed to determine specificity and sensitivity of the multiplex quantitative PCR (qPCR). Whilst the primer/probe set for B. goodwinii was species-specific, primer/probe sets for the other three species were able to identify other members of their respective genera. There was no cross detection of genera within the multiplex qPCR, and non-target bacteria were not detected. The multiplex AOD assay had differential sensitivity for each bacterial species. The assay was evaluated on swab samples collected from stem bleeds of declining oak trees at a site in south-east England and was able to detect all four bacterial species. Absolute quantification of the bacteria from swab samples was possible through the inclusion of a standard curve prepared from dilutions of gene copy standards. This diagnostic tool will facilitate rapid detection of AOD-associated bacteria from samples that can easily be taken by non-specialists without specific training, and will also find application in other experimental work such as pathogenicity and control trials.