The survival of Ceratocystis fagacearum in South Carolina

This research, which involved observing and sampling artificially inoculated oak-wilted turkey oak trees, substantiates previous field observations that C. fagacearum does not quickly kill oak trees in South Carolina. C. fagacearum seems to quickly lose virulence during the first growing season following inoculation. Rather than being quickly killed, as occurs with infected red oaks in more temperate climates, oak trees in South Carolina either survive infection or die much more slowly. Moisture, total carbohydrate, and reducing sugar contents of infected trees indicate that these trees suffer an initial physiologic trauma. Branches die first, followed by upper stems, and in few cases lower stems. Some trees, however, have survived inoculation, and initial partial wilting, but have since recovered.     

Identification and distribution of Armillaria species associated with an oak decline event in the Arkansas Ozarks

Forests in the Ozark Mountains of northern Arkansas recently experienced a widespread oak decline event. Armillaria, a root rot fungus, has been associated with other oak decline events and may have been an important contributing factor to tree mortality in this event. Although Armillaria has been identified from the Ozark Mountains in Missouri, it has never been investigated in the Arkansas Ozarks. Molecular diagnostic techniques were used in this study to identify species of Armillaria present on roots removed from dead trees of two common oak species, northern red oak, Quercus rubra L., and white oak, Q. alba L., from three geographic areas and on three topographic positions – ridges, south‐ and west‐facing benches. Armillaria(A. mellea, A. gallica or A. tabescens) was identified from 31% of root samples taken from 102 trees in seven of nine sample plots. Armillaria mellea, occurred most often (20 samples, both oak species on seven plots) followed by A. gallica (10 samples, northern red oak only on four plots), and A. tabescens occurred twice (on northern red oak in a single plot). Thus, all three Armillaria species occurred on northern red oaks while A. mellea was the only species recovered from white oaks. Results varied by topographic position with samples from tree roots on ridges having the fewest positive identifications, one of 29. West‐facing benches had the highest positive samples with 20 of 41 testing positive and trees on south‐facing benches were intermediate with 11 of 32 samples from infected trees. This study documents the occurrence of three species of Armillaria in the Arkansas Ozarks and their association with oak mortality resulting from an oak decline event coupled with a red oak borer, Enaphalodes rufulus, outbreak. Further, it documents some potential variation in host/pathogen combinations and forest site conditions.     

Fine root dynamics of oak saplings in response to Phytophthora cinnamomi infection under different temperatures and durations

The belowground effects of Phytophthora cinnamomi on 1‐year‐old saplings of two common oak species in mid‐Atlantic US forests, white (Quercus alba) and red oak (Q. rubra), were examined after incubation in pathogen‐infested soilless potting mix. Fine root lengths (0–1.5 mm in diameter) of both oak species were quantified after incubation at successive 30‐day intervals up to 300 days, for a total of 10 incubation periods. In addition, colony‐forming units (CFU) of P. cinnamomi were quantified after white oak saplings were incubated in infested soilless potting mix at different temperature/duration combinations that reflect soil conditions present in the mid‐Atlantic United States. Impact of P. cinnamomi on fine root lengths of red and white oak saplings varied considerably over time. Significant periods of fine root loss occurred primarily during spring, when bud break and leaf flush began for both oak species. Red oaks had 17% fine root loss on average, while white oaks appeared more resistant to P. cinnamomi infection with a 2% decrease in fine roots over the course of the experiment. Phytophthora cinnamomi CFU declined significantly with exposure to all incubation temperatures except 8°C. This was in contrast to in vitro experiments, where the optimum temperature for mycelial growth was determined to be 21°C and above. Significant fine root loss caused by P. cinnamomi depended on plant phenology and the oak species tested. Extreme soil temperatures have a significant adverse impact on temporal changes of P. cinnamomi population.     

Occurrence of Diplodia corticola,including new oak host records,in Wisconsin,USA

In the United States, diseased oaks (Quercus species) exhibit tip blight, branch and stem cankers, and dieback often attributed to Diplodia species or related fungi. Emergence of Diplodia corticola as a pathogen of European oaks, and reports of this fungus in the eastern and western United States, prompted re‐examination of strains from Wisconsin. These had been obtained in the late 1990s and early 2000s and previously identified only as Diplodia species. Nuclear ribosomal ITS sequences were obtained from the Wisconsin strains and analysed with other sequences from GenBank. Wisconsin strains confirmed as D. corticola were from northern red oak (Q. rubra), black oak (Q. velutina), white oak (Q. alba) and bur oak (Q. macrocarpa). Other strains from oaks in Wisconsin were D. mutila and D. seriata. Wound inoculation of northern red, white and bur oak seedlings with D. corticola in a greenhouse resulted in shoot death and stem lesions, from which the pathogen was reisolated. We conclude that D. corticola has been present in the northcentral United States for at least two decades and report two previously unrecognized hosts of this pathogen: white oak and bur oak. The roles of D. corticola, related fungi and influences of other environmental factors in deterioration of oak health in North America merit additional investigation.     

Natural regeneration in oak wilt centers

Openings, 0.2 ha and less, and 3–12 years old, caused by the oak wilt fungus Ceratocystis fagacearum had abundant oak reproduction, averaging 4700 stems per ha. Species of other woody reproduction made up over 27 000 stems per ha. Seedling sprouts comprised 54%, seedlings 30%, and stumps sprouts the remainder of the reproduction. No infections were found except a few on stump sprouts of diseased oak stumps. Oakhickory reproduction was abundant on all site qualities, but especially so on the poorer sites with over 8000 stems per ha. Sugar maple reproduction was much more abundant on high quality sites, also with over 8000 stems per ha. The reproduction in the oak wilt openings was judged to be similar to that of selection cuttings in disease-free but otherwise similar stands, and should respond in like manner.     

Genetic diversity and population structure of Raffaelea quercus‐mongolicae,a fungus associated with oak mortality in South Korea

Raffaelea quercus‐mongolicae is a fungus associated with oak wilt and deemed to cause extensive oak mortality in South Korea. Since the discovery of this fungus on a dead Mongolian oak (Quercus mongolica) in 2004, the mortality continued to spread southwards in South Korea. Despite continued expansion of the disease and associated significant impacts on forest ecosystems, information is lacking about the origin and genetic diversity of R. quercus‐mongolicae. Restriction‐site‐associated DNA (RAD) sequencing was used to assess genetic diversity and population structure among five populations (provinces) of R. quercus‐mongolicae in South Korea. In total, 179 single nucleotide polymorphisms (SNPs) were identified among 2,639 RAD loci across the nuclear genome of the 54 R. quercus‐mongolicae isolates (0.0012 SNPs per bp), which displayed an overall low expected heterozygosity and no apparent population structure. The low genetic diversity and no apparent population structure among South Korean populations of this ambrosia beetle‐vectored fungus support the hypothesis that this fungus was introduced to South Korea.     

Performance of northern red oak enrichment plantings in naturally regenerating Southern Appalachian hardwood stands

The limited success of methods to naturally regenerate northern red oak (Quercus rubra L.) has increased the use of artificial techniques to improve overall oak composition. Enrichment plantings are often recommended as a means to supplement species composition within the existing natural reproduction. Previous enrichment efforts have often resulted in low survival and poor growth, generally due to poor planting stock quality and a lack of competition control. In this study, high quality northern red oak seedlings were established on four recently harvested sites in western North Carolina using one of four competition control treatments (untreated, year 1 control, year 2 control or a weed mat) in a 1.2 × 1.2 m area around each seedling with or without a soil fertility amendment. The 3 year results show that competition control and fertilization treatments had few positive impacts on survival and growth of the planted seedlings. These results suggest that either the treatment area was insufficient to adequately release the seedlings, or other factors like belowground competition from existing advance reproduction and/or stump sprouts may be important determinants of the overall success of northern red oak enrichment plantings on recently harvested sites.     

Observations on the colonization of oak wilt mats (Ceratocystis fagacearum) by Pesotum piceae

Subcortical mats of Ceratocystis fagacearum were found to become readily overgrown by Pesotum piceae. Thus by reducing the chance of contamination of insect vectors with spores of the pathogen P. piceae is likely to contribute to the low efficiency of oak wilt transmissions.     

Perceptions of Private Forest Owners in West Virginia on the Use of Prescribed Fire in Forestry

Prescribed fire is the controlled application of fire to accomplish planned land management objectives. Public acceptance of prescribed fire varies in different parts of the world, and is little known in the state of West Virginia, USA. West Virginia is about 80% forested, and most of the forest land is in private ownership. Further, the region is home to oak (Quercus spp.), a possibly fire-dependent and highly valuable genus. Due to real and perceived risks associated with prescribed fires, it was hypothesized that forest owners are unwilling to accept the use of prescribed fire as a forest management option, even for the regeneration of oak. Non-industrial private forest owners in West Virginia were surveyed to shed light on their attitudes and opinions regarding the use of prescribed fire as a forest management tool. Contrary to the hypothesis, 64% of the responding owners were supportive of the use of prescribed fire. Acceptance was related to knowledge of prescribed fires or of firefighters. Major concerns included personal and property safety and being informed prior to burning. Therefore, while acceptance of prescribed fires among forest owners is high, timely notification can ensure further and, likely, increased, cooperation from the landowners.     

Effects of Various Silvicultural Systems on Regeneration in Mixed Hardwood Stands of the Southern Appalachians

This study investigated the effects of clear-cutting and several other commonly used silvicultural systems on regeneration at seven sites in the Appalachian Mountains of Virginia and West Virginia. These even-aged oak dominated stands ranged in age from 63 to 100 yr and were located on medium quality oak sites (site indices from 18 to 23 m, base age 50). The treatments evaluated included a clear-cut, commercial harvest, shelterwood, leave-tree, group selection, preharvest herbicide, and control.

Mixed model ANOVA was used to analyze treatment response in five species groups: (a) oak (Quercus spp.); (b) maple (Acer spp.); (c) black cherry (Prunus serotina Ehrh.), and yellow-poplar (Liriodendron tulipifera L.); (d) miscellaneous overstory hardwoods; and (e) midstory species that typically do not occupy main canopy positions in mature stands. Response variables included stem density, importance value, average height for all regeneration, and the tallest 365 stems ha^?1. Comparisons were made among treatments, species groups, and between regeneration of sprout and seedling origin.

Alternative systems with residual trees reduced the regenerations overall mean height growth compared to the clear-cut by 0.34 to 0.74 m. Current conditions indicate oak will be a lesser component of the future stand, represented mostly through stump sprouting, and maple will likely increase in proportion among all treatments.     

Dendroecology and species co-existence in an old-growth Quercus–Acer–Tilia talus slope forest in the central Appalachians,USA

Dendroecological techniques were used to examine the disturbance history and patterns of species recruitment in an old-growth Quercus rubra L. (northern red oak)–Acer saccharum Marsh. (sugar maple)–Tilia americana L. (basswood) forest on a steep, talus slope in eastern West Virginia. The forest was uneven-aged as were the populations of red oak. Sugar maple dominated the sapling layer, which comprised little or no basswood and red oak. A compilation of major and moderate releases (indicative of disturbance) in 25 cores revealed single or multiple release events in every decade from 1870–1990. The high elevation of the forest coupled with a fertile sub-soil beneath the talus ameliorated the outwardly harsh conditions of the site, allowing for the domination of typically mesophytic, nutrient demanding tree species. We observed several fire scarred trees as well as extensive small-scale blow-down throughout the forest. Frequent disturbance events were probably crucial to the co-existence and continuous canopy recruitment of the relatively light demanding red oak with highly shade tolerant sugar maple and basswood. The strong successional replacement tendencies of red oak by northern hardwoods noted elsewhere in the eastern US may be less apparent on high elevation, rocky sites in the central Appalachians. Thus, this is a unique case study of long-term red oak domination with later successional species in an old-growth forest.     

Holzmassenverluste in Zerreichenbeständen durch Inonotus nidus-pici Pilát

Volume loss in Turkey oak coppice due to Inonotus nidus-pici Pilat. Inonotus nidus-pici Pilat (order Aphyllophorales, family Mucronoporaceae) causes serious damage to stands of Turkey oak (Quercus cerris L.) in South West Europe. The decay it causes in the heartwood gives rise to considerable losses in volume.     

Modified exponential nitrogen loading to promote morphological quality and nutrient storage of bareroot-cultured Quercus rubra and Quercus alba seedlings

Abstract

Exponential nutrient loading has been used to improve nursery fertilizer uptake efficiency of conifer seedlings, but the technique has received little attention in the culture of temperate deciduous hardwoods. This study examined responses of northern red oak (Quercus rubra L.) and white oak (Q. alba L.) seedlings to modified exponential nitrogen loading during bareroot nursery culture using a broad range of nutrient supply from 0 to 3.35 g nitrogen (N) per plant per season for 18 weeks in Indiana, USA. Seedling growth and nutritional parameters followed a curvilinear pattern that ranged from deficiency to toxicity with increased fertilization consistent with trends depicted in the proposed model for nutrient loading. Fertilization increased plant dry mass by 113–260% for red oak and 49–144% for white oak. Severe nutrient deficiency occurred under indigenous soil fertility, and limited phosphorus and potassium uptake were found to inhibit seedling growth at higher N supply. The sufficiency and optimum rates were determined to be 0.84 and 1.68 g N per seedling per season, respectively, under the current cultural conditions. Fertilization at 1.68 g N per plant increased N content by 40% in red oak and 35% in white oak. This approach may be used to help refine nursery fertilization practices in hardwood culture to produce high-quality seedlings for field planting.     

Effects of prescribed fire on canopy foliar chemistry and suitability for an insect herbivore

We assessed the effects of a prescribed fire on the phytochemical characteristics and vigor of overstory chestnut oak, scarlet oak, and red maple, and measured the impact of potential changes on herbivore fitness. We compared foliar carbohydrates, tannins, nutrients, and fiber concentrations in foliage from burned and non-burned forest canopies. There were significant differences in most foliar characteristics between tree species. Total non-structural carbohydrate concentrations in scarlet oak and red maple foliage, and calcium levels in red maple foliage, were significantly lower in burned plots, but other phytochemical characteristics were largely unaffected by burning. Tree growth also varied with species. Burning increased chestnut oak relative growth, decreased scarlet oak growth, and had no affect on red maple growth. Scarlet oak and red maple foliage from burned and non-burned forest canopies were assayed for gypsy moth performance. Caterpillars fed foliage from scarlet oak, the preferred host, grew larger and developed more rapidly than did those fed red maple foliage. There were no significant burn treatment differences in caterpillar development within either tree species, suggesting that managers using prescribed fire to promote oak regeneration need not worry about enhancing forest stand susceptibility to gypsy moth. However, the fire in this study was of low to moderate intensity; more intense fires may alter foliar palatability.     

Survival of Ceratocystis fagacearum in branches of trees killed by oak wilt in Minnesota

The survival of Ceratocystis fagacearum in the branches of Quercus ellipsoidalis killed by oak wilt in Minnesota depends on the time of year when the trees die. In trees, which wilt in May or June, the pathogen can be isolated from the xylem for only 1–2 months. In trees which die later in the summer it may persist through to the following spring. Observations on oak bark beetles (Pseudopityophthorus spp.) breeding in the dead trees do not suggest that these beetles are important vectors of the disease in Minnesota.     

Oak and red maple seedling survival and growth following periodic prescribed fire on xeric ridgetops on the Cumberland Plateau

To test the hypothesis that repeated prescribed fires alone can improve the status of oak regeneration, a long-term seedling population study was established to follow permanently tagged chestnut oak (Q. prinus L.), scarlet oak (Q. coccinea Muenchh.), and red maple (Acer rubrum L.) seedlings over 8 years in sites where fire was excluded, and where fire was applied either three (3×) or four (4×) times.     

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.     

Rapid and accurate detection of Ceratocystis fagacearum from stained wood and soil by nested and real‐time PCR

A nested and real‐time PCR assay was developed for the rapid and accurate detection of Ceratocystis fagacearum, which is the causal agent of oak wilt in stained wood and soil. Based on the differences of the internal transcribed spacer (ITS) sequences of Ceratocystis spp., one pair of species‐specific primers, CF01/CF02, was designed. Whereas a 280‐bp product was amplified using the purified DNA from three isolates of C. fagacearum as the template, no PCR product was obtained from template of other 18 fungi. The detection sensitivity was 10 pg genomic DNA per 25‐μl PCR reaction volume. To increase detection sensitivity, a nested PCR was developed by using ITS1/ITS4 as the first‐round primers and CF01/CF02 in the second round, as it can detect 1 pg genomic DNA per 25‐μl PCR reaction volume. More importantly, CF01/CF02 primers were successfully adapted to real‐time PCR with a detection limit of 0.1 pg genomic DNA per 20‐μl PCR reaction volume. Using these two methods, we could rapidly and accurately detect the pathogen in artificially infected wood and soil.     

Dendroclimatological analysis of major forest species of the central Appalachians

A dendroclimatological study was carried out in the 80-year-old Fernow Experimental Forest of the U.S. Forest Service near Parsons, West Virginia (39°20′N, 79°40′W). The relationship between radial and basal area increments and corresponding monthly, seasonal, and yearly mean air temperature and precipitation were obtained over 53 years for four species: black cherry (Prunus serotina Ehrh.), northern red oak (Quercus rubra L.), white ash (Fraxinus americana L.), and yellow poplar (Liriodendron tulipifera L.). These species show positive growth responses to rainfall of previous summer, autumn, and current summer. They show significantly inverse correlation to air temperature of the current growing season. The indices of radial increment show a close correlation to those of basal area increment. However, in our analyses, basal area increment indices are more sensitive to climatic variations than radial growth indices.     

Effects of Acidic Precipitation and Ectomycorrhizal Inoculation on Growth,Mineral Nutrition,and Xylem Water Potential of Juvenile Loblolly Pine and White Oak

Abstract

Individual and interactive effects of simulated acidic rainfall and mycorrhizal inoculation on growth and nutrient and water relations of loblolly pine (Pinus taedaL.) and white oak (Quercus albaL.) grown in a loam soil were examined. Seedlings of each species inoculated with basidiospores of the ectomycorrhizal fungus Pisolithus tinctorius(Pers.) Coker and Couch, a known my-cobiont of both loblolly pine and white oak, and uninoculated control seedlings received two simulated rains per week of either pH 3.6, 4.2, or 4.8 for 26 weeks. Higher acidity rainfall reduced the growth but increased mycorrhizal colonization of loblolly pine, while both loblolly pine and white oak exposed to these rains exhibited greater foliar injury. Inoculation with P. tinctoriusincreased growth and reduced foliar injury of both species. Foliar concentrations of P, S, and Cu in loblolly pine and white oak, Ca in loblolly pine, and Fe and Zn in white oak decreased with increasing rain acidity while the Al concentration of both species increased. Higher rainfall acidity also reduced soil pH and Ca and Mg concentrations while increasing soil AI. Foliage of inoculated seedlings of both species had higher N and P concentrations and lower Al concentrations than control seedlings. Following the final rain applications, a drought cycle was simulated by withholding irrigation for two weeks during which seedling xylem pressure potential and soil water potential were measured. One day after cessation of irrigation, xylem pressure potential of loblolly pine that had received pH 3.6 rains was lower than that of other treatments. Thereafter, xylem pressure potential and soil water potential of the inoculated treatment decreased below those of the control treatment in both species. These results suggest that acid deposition is detrimental to juvenile loblolly pine and white oak, but the magnitude of this effect is less than the positive response to ectomycorrhizal inoculation.