Signs and symptoms associated with Heterobasidion annosum and Armillaria ostoyae infection in dead and dying mountain pine (Pinus mugo ssp. uncinata)

Symptoms and signs associated with root rot caused by Heterobasidion annosum or Armillaria ostoyae in mountain pines (Pinus mugo ssp. uncinata) were investigated in the Swiss Alps. A sample of dying or recently dead mountain pine trees (≥12 cm d.b.h.) and saplings (<1.3 m height) was assessed for root pathogen infection by taking root samples followed by isolations in the laboratory. From a subsample, an additional core was taken from the butt of each tree and evaluated in the same fashion. A total of 157 dying or recently dead mountain pine trees and 184 saplings with roots infected by either of the two pathogens or which lacked infection were analyzed using logistic regression models. The main objectives were to determine the most prominent symptoms induced by the fungi (resinosis), signs of the fungi (mycelia, fruiting bodies and rhizomorphs), and tree characteristics (d.b.h./height and evidence of wounds) that would allow an easy and reliable determination of H. annosum and/or A. ostoyae infection of mountain pines in the field. Heterobasidion annosum caused both root and butt rot on mountain pine, whereas A. ostoyae was mostly restricted to the root systems of the trees sampled. The most discriminating sign for the presence of A. ostoyae infection was the presence of characteristic mycelial fans, and for H. annosum root rot the presence of H. annosum mycelia (sheets of paper‐thin mycelium and mycelial pustules). In addition, resinosis was a powerful predictor for A. ostoyae in trees. Symptoms and signs indicating A. ostoyae or H. annosum infections were more reliable for saplings than for mature trees. Armillaria rhizomorphs were not useful in detecting A. ostoyae infection and, if present, were often formed by saprophytic Armillaria species. Heterobasidion annosum fruiting bodies were rarely observed and poorly reflected the widespread occurrence of this pathogen in the mountain pine forests.     

Tree vigour and the susceptibility of Douglas fir to Armillaria root disease

In this study the effects of thinning, fertilization and pruning on the vigour of Douglas fir (Pseudotsuga menziesii) and its susceptibility to Armillaria root disease were explored. Tree vigour was defined as the relative capacity for tree growth, expressed as the above-ground biomass increment per unit of photosynthetic tissue, or growth efficiency (GE). It has been hypothesized that trees with higher GE can better resist pathogen attack, and that GE can be used as a predictor of tree susceptibility to disease. In a previous study, four Douglas fir plantations were thinned, fertilized and pruned in all combinations, and the effects of these treatments on tree vigour were measured after 10 years. Root disease was not a factor in the initial study design, and mortality was ignored until 8 years after the treatments were applied. The results of an earlier study were utilized and the correlation between Armillaria root disease incidence and the effects of earlier stand treatments on tree growth was tested. Armillaria ostoyae was the primary mortality agent in the study area. Disease incidence on infested subplots ranged from 2 to 20%. Armillaria ostoyae incidence was the highest at medium tree density (6.1%), slightly lower on the low density (5.6%), and lowest on the unthinned plots (3.8%). There were no significant correlations between disease incidence and previous tree growth. The vigour of trees that became symptomatic or died by 1993 was not significantly different in 1983–85 from the vigour of trees that remained asymptomatic. On these sites, in areas of infestation, A. ostoyae is killing the largest, fastest growing trees, as well as less vigorous trees. Armillaria continues to cause mortality, regardless of the growth efficiency or growth rate of the host.     

Pathogenicity and virulence of Armillaria sinapina and host response to infection in Douglas‐fir,western hemlock and western redcedar in the southern Interior of British Columbia

The frequency of infection, lesion characteristics and anatomical changes in phloem and cambial tissues caused by Armillaria sinapina were studied on inoculated trees of Douglas‐fir, western hemlock and western redcedar and compared with results of A. ostoyae inoculations previously reported on the same host species. Similar percentages of inoculations resulting in infection of roots on the three hosts indicate that A. sinapina and A. ostoyae are equally pathogenic. Armillaria ostoyae was more virulent than A. sinapina as demonstrated by fungal exudates from A. ostoyae inoculum blocks, which appeared to cause lesions on roots; the higher frequency at which lignified impervious tissue (IT) and necrophylactic periderm (NP) developed in bark and following cambial invasion, compartmentalization; the large proportion of roots that showed no visible host response; the large zones of IT formed under continuous stimulation by A. ostoyae advancing in inner bark; and the high frequency of breaching of NP barriers. Spread of A. sinapina mycelium in host species appeared slower than that of A. ostoyae, particularly in Douglas‐fir and western hemlock. In western redcedar, A. ostoyae induced stronger host responses than those following invasion by A. sinapina, which included further expansion of the induced rhytidome response, traumatic phloem resin duct formation and higher numbers of polyphenolic parenchyma comprising its barrier zone. Where damage by A. sinapina ensued, it was always associated with high inoculum potential. The ecology of virulent and less virulent species of Armillaria in natural forests is discussed.     

Armillaria species distribution on symptomatic hosts in northern hardwood and mixed oak forests in western Massachusetts

Armillaria spp. are some of the most important forest pathogens in mixed hardwood forests of southern New England, yet their role as prominent disturbance agents is still not fully appreciated. We investigated the distribution of Armillaria species across eight separate stands of northern hardwood and mixed oak forests in western Massachusetts. We were specifically interested in the Armillaria species distribution from live, symptomatic hosts and not in determining overall incidence in the forest. From 32 plots (16 within each forest type), 320 isolates were collected. Armillaria was routinely encountered causing disease of live trees. In total, 89% (286/320) of all isolations came from live hosts exhibiting symptoms of root and butt rot. Overall, A. gallica was the dominant species in each forest type, making up 88/160 (55%) isolates from northern hardwood and 153/160 (96%) of all isolations from mixed oak stands. However, northern hardwood forests showed much greater species diversity, as A. calvescens, A. gemina, A. ostoyae, and A. sinapina were all found. At one site, a northern hardwood forest surrounding a high elevation spruce-fir forest, A. ostoyae was the most abundant species encountered. All five Armillaria species were found causing disease of live hosts, including A. gemina, a species considered by some as weakly virulent. Armillaria gallica was found on 22/23 tree species’ sampled, and was found most often causing butt rot.     

Temporal evolution of collar lesions associated with ash dieback and the occurrence of Armillaria in Belgian forests

Ash dieback, caused by the fungus Hymenoscyphus fraxineus, has been observed in Europe for several years. In Belgium, the disease was first reported in 2010. Besides crown defoliation and dieback, collar lesions have sometimes been reported. To evaluate the prevalence and the progression of collar lesions and crown defoliation in ash dieback‐affected stands of various ages, a survey was conducted in 2013 and 2014 on 268 ash trees (Fraxinus excelsior) originating from 17 Walloon forest stands. The results showed that the proportion of trees with collar lesions greatly increased between June 2013 and September 2014 and that there appeared to be no significant link between a tree's diameter‐at‐breast height (DBH) and collar lesion occurrence. The mean percentage of defoliation increased in each forest stand across time, with observations conducted in September 2013 and 2014 showing a positive correlation with the mean percentage of trees with collar lesions. Molecular tests were carried out on 103 additional trees originating from 12 of the 17 stands to evaluate the occurrence of H. fraxineus and Armillaria spp. at the collar level. Most of the trees (98%) were infected by H. fraxineus. In contrast, only 41% of the samples were infected with Armillaria spp., most commonly A. gallica and A. cepistipes. This study discusses the role of Armillaria spp. and the rapid increase in the number of trees with collar lesions within the context of the evolution of ash dieback in Europe.     

Stump infection by Armillaria in first-rotation conifers

Information about the entry of Armillaria into first-rotation pine and spruce stands was obtained by searching for infected stumps, rhizomorph systems or trees that had been killed. In pines Armillaria foci were very rare. In pure Norway spruce Armillaria lutea and A. mellea were detected in stumps but rhizomorphs did not extend into the soil; in Norway spruce mixed with oak, by contrast, A. lutea sometimes produced extensive rhizomorph systems. In Sitka spruce small groups of trees had been killed by A. ostoyae. All foci investigated in conifers contained different genotypes of Armillaria and probably originated from spore infection of stumps created by thinning. Some implications of these findings are discussed.     

Effects of metals and pH on in vitro growth of Armillaria ostoyae and other root and butt rot fungi of red spruce

Armillaria ostoyae, Perenniporia subacida, Resinicium bicolor and Scytinostroma galactinum, root and butt rot fungi found on red spruce, Picea rubens, were tested, in vitro, for their sensitivity to metals typically found in high elevation forest soils where red spruce grows. Rhizomorph production by A. ostoyae from woody inocula in soils from red spruce stands at three elevations at each of five mountainous sites in the eastern United States was inhibited completely in the mineral soil from all elevations at all sites, and was also reduced significantly in the organic horizon from the upper two elevations at three of the sites. Inhibition was correlated with concentrations of metal ions in the soil. Growth of rhizomorphs into an agar medium containing lead and other heavy metals was inhibited for isolates of A. ostoyae from red spruce, but not for an isolate of Armillaria gallica from sugar maple; aluminium inhibited rhizomorph growth of isolates of both species. Mycelial growth of all four root and butt rot fungi was inhibited by lead, aluminium and other heavy metals depending on the solubility and concentration of metal and pH of the medium; growth inhibition was usually greater at an initial pH of 3.5 than at pH 4.5. Metal ions inhibited radial growth of Armillaria species more than that of the other three fungi. Rhizomorph growth of Armillaria was inhibited more than radial growth. Because local spread of A. ostoyae occurs frequently by means of rhizomorph growth between near roots, increases in lead, aluminium and other metals in the forest floor may contribute to this fungus' scarcity in high elevation soils and reduced incidence of infection at these sites in the eastern United States.     

Nachweis der GattungArmillaria (Fr.:Fr.) Staude undHeterobasidion annosum (Fr.) Bref. in Fichte (Picea abies [L.] Karst.) und Erfassung der klonalen Ausbreitung vonA. ostoyae-Genotypen unter Verwendung molekularer Methoden

Zusammenfassung Methoden zur Entwicklung molekularer Marker mit Hilfe der Polymerase-Kettenreaktion (PCR) werden beispielhaft an den beiden forstlichen PathogenenArmillaria spp. undH. annosum erl?utert. Unter Verwendung der aus „internal transcribed spacer“ (ITS)-DNA-Sequenzen abgeleiteten Primerpaare ARM-1/ARM-2 bzw. HET-7/HET-8 konntenArmillaria spp.- bzw.H. annosum-spezifische DNA-Abschnitte mittels PCR amplifiziert werden. Diese PCR-Diagnosemethode erlaubt einen schnellen und effizienten Frühnachweis dieser beiden forstlich relevanten Wurzel- und Stockf?uleerreger in verschiedenen Substraten bzw. Pflanzengeweben. Die genetische Variabilit?t von 20A. ostoyae-Isolaten aus verschiedenen geographischen Herkunftsgebieten wurde untersucht. Die UPGMA-Clusteranalyse der unter Verwendung von 10 Decamer-Zufallsprimern (OPA 01-10) erhaltenen „random amplified polymorphic DNAs“ (RAPDs)-Muster gruppierte die Isolate in Untergruppen mit 40–96% ?hnlichkeit, was auf eine hohe intraspezifische genetische Variation hindeutet. Die potentielle Rolle der historischen und gegenw?rtigen Ausbreitung von Fichtenpflanzen für die genetische Variation vonA. ostoyae in Europa wird diskutiert. Die aufgefundenen polymorphen DNA-Marker wurden für die Erfassung der Populationsstruktur bzw. -dynamik sowie der r?umlichen Ausbreitung vonA. ostoyae-„genets“ in ausgew?hlten Befallsgebieten genutzt. Erste Untersuchungen haben unterschiedliche Ausbreitungsstrategien vonA. ostoyae in Abh?ngigkeit von Klima-, Standort- und Immissionsfaktoren aufgezeigt.      

Ecology of Armillaria species on silver fir (Abies alba) in the Spanish Pyrenees

? We describe the distribution and the ecology of three Armillaria species observed in silver fir (Abies alba) forests of the Pyrenees.

? We surveyed the presence and abundance of Armillaria above and belowground in 29 stands. Isolates were identified by the PCR-RFLP pattern of the IGS-1 region of their ribosomal DNA. We measured several ecological and management parameters of each stand in order to describe Armillaria infected sites.

? Armillaria cepistipes was the most abundant of three species observed. Armillaria gallica was dominant in soils with a higher pH and at lower elevations. Armillaria ostoyae seemed to be more frequent in stands where A. alba recently increased its dominance relative to other forest tree species. Thinning activities correlated with an increased abundance of Armillaria belowground. In 83% of the stands the same Armillaria species was observed above and belowground.

? It seems that in a conifer forest, A. cepistipes can be more frequent than A. ostoyae, a virulent conifer pathogen. Since logging is related to a higher abundance of Armillaria in the soil, the particular Armillaria species present in a given stand could be considered an additional site factor when making management decisions.

     

Armillaria ostoyae associated with dying 60‐year‐old Scots pines in northern Turkey

We investigated an Armillaria root disease centre in a 60‐year‐old, naturally regenerated Pinus sylvestris stand located at 1250 m altitude in Sinop province in northern Turkey. Based on sequence analysis of the internal transcribed spacer region of the rDNA, the mycelial fans sampled from the infected trees were Armillaria ostoyae. Multilocus genotyping indicated that the fans belonged to a single genet, at least 0.2 ha in size. Drought as a predisposing factor is discussed. The study area received 50–70 and 25–50% of the normal precipitation in May, June and August in 2009 and in August and September in 2010, respectively. Moreover, earlier excavation work to prepare a forest road had probably disrupted the water table in the affected stand.     

Comparison of the virulence of Armillaria cepistipes and Armillaria ostoyae on four Norway spruce provenances

The basidiomycetes Armillaria cepistipes and Armillaria ostoyae frequently occur in the same forest stand. In this study, we determined the virulence of 20 isolates of A. cepistipes and 16 isolates of A. ostoyae on four different provenances of 2‐year‐old Norway spruce (Picea abies). Within 30 months after inoculation, 1.1 and 19.1% of the seedlings inoculated with A. cepistipes and A. ostoyae, respectively, had died or were dying. The incidence of dead and dying seedlings varied between 3 and 49% among the A. ostoyae isolates. The virulence of an isolate was positively correlated to its ability to produce rhizomorphs. One Norway spruce provenance showed significantly lower susceptibility to A. ostoyae than the other three. Rhizomorphs of both Armillaria species were attached to the root surface. The attached rhizomorphs of A. ostoyae, however, were associated with significantly more lesions. The virulence of the isolates was not correlated with their wood‐degrading capability for either of the Armillaria species.     

Ecology of Armillaria species on conifers in Japan

Distribution, host preference and pathogenicity of Japanese Armillaria species on conifers were investigated on the basis of field collections of 65 isolates. We identified seven Armillaria species from 19 conifer species including six major Japanese plantation conifers using mating tests and sequences of the translation elongation‐1 α gene. Armillaria mellea, Armillaria ostoyae, Armillaria cepistipes and Armillaria sinapina were frequently collected, whereas Armillaria nabsnona, Armillaria tabescens and a biological species Nagasawa’s E were rare. On the basis of host condition when the isolates were collected, A. mellea, A. ostoyae, A. cepistipes and A. tabescens are considered as moderate to aggressive pathogens of conifers in Japan.     

Identification and partial characterization of an extracellular manganese-dependent peroxidase in Armillaria ostoyae and Armillaria mellea

Laccase and manganese-dependent peroxidase (Mn peroxidase) activities were detected in the culture media of Armillaria ostoyae and A. mellea. Mn peroxidase was produced in significantly higher quantity by the A. ostoyae isolates and was purified by chromatography from one isolate of this species. Some properties of the purified enzyme were examined (absorption spectrum, H₂O₂ and MnSO₄ optimal concentrations, pH optimum and lactate stimulation). Enzymes of potential importance in the lignin degradation (especially Mn peroxidase) by Armillaria sp. are compared to those of other root-rotting fungi. The possible role of Mn peroxidase in modulating the pathogenicity of Armillaria sp. is discussed.     

Untersuchungen zur Pathogenität des BaumholznematodenBursaphelenchus fraudulentus Rühm

Zusammenfassung Der LaubbaumnematodeBursaphelenchus frandulentus ist eng mit dem QuarantäneschädlingB. xylophilus verwandt und kann in hoher Anzahl in toten oder absterbenden Laubbäumen gefunden werden. Um zu klären, obB. frandulentus pathogen für Laub- oder Nadelbaumarten sein kann, wurden mikroskopische Untersuchungen am Holz von mitB. frandulentus befallenen Eichen durchgeführt und Infektionsversuche im Gewächshaus mit dreijährigen Bäumen angelegt.Die Auswertung von Holzschnitten ergab, daßB. fraudulentus nur in den Tracheen vorhanden war. Es wurden keine direkten Zell- oder Gewebezerstörungen beobachtet, die auf eine Saugtätigkeit des Nematoden zurückzuführen wären. In den Tracheen waren ebenfalls Pilzhyphen vorhanden, die die Nematoden möglicherweise als Nahrungsquelle nutzen.AnQuercus robus, Fagus sylvatica undPinus sylvestris wurden Inokulationen mit 15 000B. fraudulentus/Baum durchgeführt, oder auch kombinierte Pilz-Nematoden-Inokulationen mitTrichoderma sp. oderVerticillium leptobactrum und zusätzlich jeweils 12 000B. fraudulentus/Baum. Während der elfmonatigen Versuchsdauer wurden weder durchB. fraudulentus alleine noch in Kombination mit den Pilzen Krankheitssymptome hervorgerufen. Die Bäume zeigten keine Chlorosen oder Welkesymptome und trichen im Folgejahr wieder gut aus. Aus den Bäumen der untersuchten Arten konnten nach 11 Monaten vitaleB. fraudulentus in unterschiedlicher Anzahl aus dem Holz reisoliert werden, insgesamt jedoch deutlich weniger als inokuliert wurde. Die beiden inokulierten PilzartenV. leptobactrum undTrichoderma sp. hatten keinen fördernden Einfluß auf die Vermehrung und Ausbreitung vonB. fraudulentus inQ. robur undF. sylvatica.Die Ergebnisse zeigen deutlich, daßB. fraudulentus apathogen fürQ. robur, F. sylvatica undP. sylvestris ist.

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Studies on the pathogenicity of the deciduous tree nematodeBursaphelenchus fraudulentus

Bursaphelenchus fraudulentus is closely related to the quarantine pestB. xylophilus and can be found in high numbers in dead or severely damaged deciduous tree species. To find out ifB. fraudulentus can be pathogenic for deciduous or coniferous trees, inoculation tests with three year old trees were conducted in the greenhouse. In addition to this, wood of infested oak trees was microscopically investigated.The microscopical evaluation of thin wood sections showed thatB. fraudulentus occurred only in the tracheae. Besides the nematodes fungal hyphae were present which could serve as food source for the nematodes. No cell or tissue destruction—which could have been attributed to a possible feeding activity of the nematodes—was visible.In a first test seriesB. fraudulentus was inoculated intoQuercus robur, Fagus sylvatica andPinus sylvestris (15,000 nematodes/tree). In a second test series combined nematodefungus-inoculations withTrichoderma sp. orVerticillium leptobactrum together withB. fraudulentus (12,000 nematodes/tree) were carried out. None of the treatments showed any chloroses or wilting symptoms during the test period (11 months) and all the trees developed new shoots in the following spring. After 11 months viableB. fraudulentus could be reisolated from the wood of all inoculated trees in varying numbers, but in general the number of nematodes represented only a small percentage of the inoculum density. The two inoculated fungiV. leptobactrum andTrichoderma sp. had no promotive effect on multiplying and spreading ofB. fraudulentus inQ. robur andF. sylvatica.These results demonstrate clearly thatB. fraudulentus is apathogenic forQ. robur, F. sylvatica andP. sylvestris.

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Mit einer Abbildung und 5 Tabellen     

Mycelial fan formation of three sympatric Armillaria species on excised stem segments of Picea abies

Mycelial fan formation was studied in five Armillaria cepistipes, ten A. borealis and ten diploid and six haploid A. ostoyae strains on excised stem segments of Picea abies. Stem segments were either non‐autoclaved or autoclaved, representing dying and dead wood, respectively. To confirm the identity of mycelial fans on non‐autoclaved stem segments, re‐isolations were made and isolates characterized with microsatellite markers. Mycelial fan formation on autoclaved stem segments was fast and reliable for most of the tested Armillaria strains. On non‐autoclaved stem segments, mycelial fan formation was slower, more erratic and less predictable. Mycelial fan formation was fastest in A. cepistipes closely followed by A. borealis and was slowest in A. ostoyae. For two A. cepistipes and four A. ostoyae strains (all diploid), growth rates of mycelial fans were estimated in a time course experiment. They ranged between 5.1 and 8.7 mm/day for autoclaved and between 1.4 and 4.7 mm/day for non‐autoclaved stem segments. The haploid A. ostoyae strains also formed mycelial fans on autoclaved stem segments, but typically slower and less reliably than the diploid strains. Whether haploid strains are able to produce mycelial fans on non‐autoclaved stem segments remains unknown because of accidental diploidization of the original haploid strains which was likely caused by basidiospores introduced into the study system on the non‐autoclaved stems. Overall, the method developed in this study may be useful for further investigations into the genetic, physiological and biochemical nature of mycelial fan formation in the genus Armillaria.     

Phylogeography and host range of Armillaria gallica in riparian forests of the northern Great Plains,USA

Root disease pathogens, including Armillaria, are a leading cause of growth loss and tree mortality in forest ecosystems of North America. Armillaria spp. have a wide host range and can cause significant reductions in tree growth that may lead to mortality. DNA sequence comparisons and phylogenetic studies have allowed a better understanding of Armillaria spp. taxonomic diversity. Genetic sequencing has facilitated the mapping of species distributions and host associations, providing insights into Armillaria ecology. These studies can help to inform forest management and are essential in the development of disease risk maps, leading to more effective management strategies for Armillaria root disease. Armillaria surveys were conducted on publicly owned lands in North Dakota, South Dakota, and Nebraska, U.S.A. Surveyed stands consisted of riparian forests ≥0.4 hectares in area. Armillaria was found at 78 of 101 sites. A total of 57 Armillaria isolates—associated with 12 host tree species—were used for DNA sequencing of the translation elongation factor‐1 alpha (tef1) gene. Armillaria gallica was the only species identified within the study sites. Results suggest that A. gallica is a common root pathogen of hardwood trees in riparian forests of the northern Great Plains with a wider host range and geographic distribution than previously recognized.     

Brenneria spp. and Rahnella victoriana associated with acute oak decline symptoms on oak and hornbeam in Iran

Decline diseases of forest trees are complex syndromes not attributable to single causal factors. In Iran, symptoms of decline disease have been observed in a number of native forest species including Quercus castaneifolia (chestnut‐leaved oak), Q. brantii (Persian oak) and Carpinus betulus (hornbeam). The symptoms are prevalent in the northern forests and the Zagros mountain forests. There are parallels between the disease in Iran and acute oak decline (AOD) reported in the UK, specifically the presence of weeping cankers, which have been attributed to a polybacterial complex wherein Brenneria goodwinii is considered a key necrogen. Based on the AOD symptomatology, and as a first step towards discovering potential causal agents of the stem weeping symptoms of affected trees in Iran, necrotic tissues were tested primarily for the presence of B. goodwinii. Symptomatic Q. castaneifolia and C. betulus from the Mazandaran Province and symptomatic Q. brantii from Kohgiluyeh and Boyerahmad Province were sampled. Isolation and culture on a selective medium yielded uniform bacterial colonies. Isolates were characterized using phenotypic and genotypic (DNA sequencing) tests. The isolates were phenotypically identical to members of Pectobacteriaceae and Yersiniaceae, specifically Brenneria and Rahnella spp. The nucleotide sequences of the 16S rRNA and housekeeping genes gyrB, infB and atpD (MLSA) amplified via PCR demonstrated that the isolates from the trees in Iran were indeed B. goodwinii, B. roseae subsp. roseae, Rahnella victoriana and an unknown species of Brenneria. Most bacteria isolated from non‐symptomatic trees were Gram‐positive, and Pseudomonas spp. were dominant, but bacterial species isolated from the diseased trees were not detected in healthy trees. Hypersensitivity response tests were positive, but inoculation on saplings was more variable with internal necrosis developing only once in the test period. Therefore, further testing is required. This is the first report of the incidence of B. goodwinii, B. roseae subsp. roseae, R. victoriana and Brenneria sp. associated with acute oak decline‐like symptoms on Q. castaneifolia, Q. brantii and C. betulus across the western forests of Iran and in the world.     

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.     

Armillaria ostoyae in managed coniferous forests in Kastamonu in Turkey

Although several Armillaria species have been reported in Turkey, there is little information about their ecology in Turkish forests. In this study, we investigated five forest stands, approximately 5–74 ha in size, in Kastamonu province in the Black Sea Region of Turkey for the presence of Armillaria species in stumps and logs. The stands were mixed Abies nordmanniana ssp. bornmülleriana and Pinus sylvestris forests managed using a selective cuttings system; the proportion of fir in the total number of stems and stumps ranged from 36 to 98%. Based on sequence analysis of the internal transcribed spacer and intergenic spacer regions of the rDNA, all rhizomorphs sampled from the stumps and logs were of Armillaria ostoyae. The size of the genets was estimated with random amplified microsatellites analysis of the isolates and ranged from single stumps to approximately 450 m². One to seven genets were found in each stand. These results indicate that the genets had arisen from spores and vegetative spread was limited on most sites.