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Forest Insect & Disease Management Guide for the Northern and Central

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Forest Insect & Disease Management Guide for the Northern and Central 2009 Forest Health Protection and State Forestry Organizations Management Guide for Brennan A. Ferguson Ferguson Forest Black Stain Root Disease Pathology Consulting, Inc.; Missoula, MT Leptographium wageneri (Kendr.) Wingf. Hosts: This disease is common and damaging in Topics Douglas-fir and west of the Cascade range of Oregon, Ponderosa pine Washington and northeast California, and Introduction 1 Pinyon pine in parts of eastern Oregon. Nomenclature 2 Occasionally, other It rarely causes significant damage in Host infection 2 pine species. inland or Rocky Mountain forests. Pathological 2 anatomy Pathogenesis 3 Introduction Black stain root disease BSRD was detected in 18.6% of Symptoms 4 (BSRD) is a wilt-like disease of 500, 10- to 30-year-old Douglas-fir Pathogen survival 5 conifers caused by the native, plantations, compared to 1.2% of Insect vectors 5 insect-vectored, fungal pathogen same with Armillaria root disease Leptographium wageneri (Kendr.) and 7.0% with laminated root rot Soil disturbance 6 Wingf. (Harrington and Cobb (Hessburg et al. 2001). In a subset Precommercia 7 1983). It is considered one of the of eighty of the 500 plantations l thinning five most-damaging root diseases that underwent intensive BSRD in Western forests (Hadfield et al. surveys the percentage of crop Disease 8 management 1986). Black stain root disease is trees affected was: <0.1% in 93% widespread across much of the of stands; 0.1-2% in 3% of stands; References cited 9 range of its hosts, but incidence 2.1-5% in 3% of stands; and >5% and severity, and thus the in 1% of the stands (Hessburg et al. importance to forest management, 2001). In the most heavily vary greatly. impacted Douglas-fir plantations in Incidence and severity of southwest Oregon and northwest Key Points BSRD in Douglas-fir and California, stocking levels were ponderosa pine is greatest west of found to be reduced by as much as A wilt disease the Cascade Range in Oregon, 50% (DeNitto 1982, Goheen et al. Vectored by Washington, and northern 1983, Goheen et al. 1984, Goheen insects, and then California. In southwest Oregon et al. 1985). spreads root-to- root Rarely damaging OVERVIEW OF in the northern BLACK STAIN ROOT DISEASE MANAGEMENT Rockies. 1. Favor mixed species. Plant mixtures that include resistant species May be serious in and favor resistant species during precommercial thinning. off-site plantations 2. Minimize soil disturbance. Avoid tractor-logging which may Avoid attracting compact soil and attract the insect vectors. the insect vectors 3. Avoid tree injury or remove injured trees of host species. Injured trees attract vectors. Minimize injuries during skidding, falling, road building, and brushing operations, especially near young trees. Page 2 Black Stain Root Disease Hansen and Goheen (1988) Hunt 1988), Phellinus weirii (Filip reported that 30-40 cm diameter and Goheen 1982, Goheen and PATHOGEN Douglas-fir continued to die on Filip 1980, Morrison and Hunt NOMENCLATURE, long-term study plots across 1988, Witcosky 1989), and TAXONOMY, AND western Washington and Oregon. Heterobasidion annosum (Kelsey POPULATION Losses in ponderosa pine can be et al. 1998b). GENETICS locally severe as well (Byler et al. In Idaho and Montana it is rare 1979, Owen 2000). for BSRD to be found acting as a There are three varieties As an insect-vectored pathogen, primary pathogen of Douglas-fir of L. wageneri. They, and with the vectors generally or pines, and the disease is not and the hosts they primarily affect, are: attracted to weakened trees such as perceived as a concern in forest those infected by other root management. Where it has been L. wageneri var. pathogens, L. wageneri often occurs observed it is in conjunction with pseudotsugae - in complexes with other root other root pathogens considered to Douglas-fir (Harrington and disease fungi (Filip and Goheen be the primary agent of mortality Cobb 1987); 1982, Goheen and Filip 1980, (Byler et al. 1983), or in off-site Morrison and Hunt 1988). It has ponderosa pine plantations. Other L. wageneri var. been found colonizing the same wood-staining, root-infecting ponderosum - ponderosa, Jeffrey, roots of Douglas-fir as Armillaria fungi in the genus Leptographium and lodgepole pines ostoyae (Byler et al. 1983, Goheen can also be found in Idaho and (Harrington and and Hansen 1978, Morrison and Montana (Wingfield et al. 1994). Cobb 1986); L. wageneri var. wageneri - Host infection singleleaf pinyon and Leptographium wageneri pinyon pines grafts like other root disease fungi, (Harrington and cannot decay wood or penetrate but can also grow through the soil Cobb 1986). non-wounded host roots. Smith for a few centimeters (Goheen (1969), using an isolate collected 1976, Hessburg and Hansen 1986a, ponderosa pine, found the fungus Landis and Helburg 1976). had no ability to break down Hessburg and Hansen (2000) cellulose. Hessburg (1984) demonstrated that minute wounds concluded that L. wageneri cannot and natural openings in seedling penetrate to the xylem on its own root systems, where a direct path to and requires an existing opening to the xylem was exposed, serve as initiate an infection. infection points for L. wageneri Leptographium wageneri spreads hyphae growing in soil. tree-to-tree at root contacts and Pathological anatomy The pathological anatomy of pattern compared to the wedge- BSRD has been studied in pinyon shaped staining commonly seen pine (Wagener and Mielke 1961), with blue stain fungi. Smith ponderosa pine (Smith 1967), and (1967) reported that mature host Figure 1. Cross-section of Douglas-fir (Hessburg and Hansen tracheids were the sole habitat for infected Douglas-fir root showing typical staining 1987). Wagener and Mielke the fungus in infected ponderosa pattern. (Photo courtesy of (1961) were the first to note that pine, and that hyphal movement E. Hansen, Oregon State pathogen growth followed the between tracheids was via University) annual rings (Figure 1), a unique bordered-pit pairs. Black Stain Root Disease Page 3 Hessburg and Hansen (1987) found the pigmented hyphae of L. wageneri exclusively in the sapwood xylem tracheids of Douglas-fir, moving between tracheids via bordered-pit pairs (Figures 2-3). The most noticeable host response to L. wageneri var. Figure 2. Tangential view of amber- pseudotsugae infection is partial to colored L. wageneri hyphae within complete plugging of tracheids by Douglas-fir tracheids. gums; this and plugging of tracheids by pathogen hyphae distances up the stem from the root were proposed as the primary collar (Hessburg and Hansen, 1987) Figure 3. L. wageneri hyphae passing through pit pairs of cause of water blockage and (Figure 4). Once a stain column Douglas-fir tracheids. subsequent wilting (Hessburg and extends up to the root collar it can Hansen 1987). Non-pathogen- descend down a non-infected root. colonized tracheids adjacent to Resinosus is often present on the exterior of infected roots or outside colonized tissue become plugged as The most noticeable well, and stain columns sometimes stain columns above the root collar. host response is consist of 20% hyphae-clogged Earlywood is typically colonized partial to complete tracheids and 80% gum-clogged prior to latewood within the same plugging of tracheids tracheids. Colonization of tissue by growth ring, so that in cross- by gums.. the the amber-colored hyphae results in sectional view the stain columns primary cause of columns of black-stained tissue appear as concentric rings in the water blockage and earlywood of successive growth subsequent wilting extending along infected roots and (Hessburg and rootlets of all sizes, and for short rings (Figure 1). Hansen 1987) Pathogenesis (disease development) The wilt-inducing nature of and like normal heartwood, BSRD is seen in host responses to pathological heartwood does not advancing colonization. Hessburg function in water transport. In 20- (1984) reported a critical water year-old Douglas-fir, the mean shortage occurred in Douglas-fir basal, cross-sectional area of seedlings 30 to 40 days after normal heartwood in non-infected inoculation with L. wageneri. trees was 23%; the basal, normal- Lawson (1988) found colonization plus-pathological-heartwood in of Douglas-fir by L. wageneri non-symptomatic, diseased trees corresponded to reduced water was 44%; and the basal, normal- uptake and transpiration, and that plus-pathological-heartwood in an increase in colonization symptomatic, diseased trees was correlated with a 50% reduction in 77% (Lawson 1988) (Figure 5). water transport to needles At breast height the percentages of compared to non-infected heartwood in these same trees Figure 4. Staining caused by L. wageneri var. controls. Douglas-fir forms were 18%, 32%, and 64%, pseudotsugae in Douglas-fir. pathological heartwood in reaction respectively. (Photo courtesy of E. Hansen, to colonization by L. wageneri, Oregon State University) Page 4 Black Stain Root Disease Impacts on water transport are similar in ponderosa pine. Infection by L. wageneri had a profound effect on hydraulic conductivity in ponderosa pine roots examined by Joseph et al. (1998). Nearly 100% of the xylem in apparently non-infected, healthy roots was capable of conducting water, compared to just 12% in roots infected with L. wageneri. The specific conductivity of infected roots was 5% that of healthy roots, while the conductivity of functional xylem Figure 5. Dye-uptake (green) in BSRD-infected (left) and non-infected (right) basal cross-sections of Douglas-fir (See Lawson 1988). in diseased roots was only one Pathological heartwood in infected cross-section has extended to third that of functional xylem in encompass L. wageneri stain columns. (Photo by T.T. Lawson; courtesy healthy roots. of E. Hansen, Oregon State University) Symptoms Black stain root disease in reduction prior to death (Morrison Douglas-fir causes progressive and Hunt 1988), while severe crown symptoms similar to other leader reduction in young root diseases (Figure 6). Leader, Douglas-fir plantations in branch, and needle length is southwest Oregon preceded death reduced, followed by loss of older in most cases by three and four needles and a resultant thinning years (Witcosky 1981).
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