Black Stain Root Disease of Conifers Paul F. Hessburg, Donald J. Goheen, and Robert V. Bega The black stain fungus—Lep- the disease was often mistakenly at- tographium wageneri (Kendrick) tributed to other, more easily identi- Wingfield*—infects and kills several fied root diseases or to bark beetles, species of western conifers. The fun- which are commonly associated with gus colonizes water-conducting tis- the rapid decline and death of black sues of the host's roots, root collars, stain-infected trees. and lower stems, ultimately blocking the movement of water to foliage. Black stain occurs in many locations Severely infected trees exhibit wilting throughout the western United States. symptoms characteristic of vascular At present, the greatest development of wilt diseases. Black stain kills young the disease occurs in southeastern and trees within a year or two of infection. northwestern California, southwestern Older infected trees decline more and east-central Oregon, the central slowly (over 2 to 8 years) and are often Sierra Nevada, and southern Colorado. predisposed to bark beetle infestation. In recent years, reports of black stain in young, intensively managed stands Distribution have increased dramatically, espe- cially in Oregon and California. The Black stain root disease is thought to disease affects trees in high-use recre- be native to western coniferous ation areas and areas important for forests. Although the disease was first wildlife management as well as those discovered in 1938, further spread on lands dedicated to timber produc- went virtually undetected until the tion. 1970s. Tree mortality associated with ————————————— Paul Hessberg is research plant pathologist at the USDA Forest Service's Pacific Northwest Research Sta- tion, Forestry Sciences Laboratory, Wenatchee, Washington; Donald Goheen is plant pathologist/entomolo- gist at the USDA Forest Service's Southwest Oregon Forest Insect and Disease Technical Center, Rogue River National Forest, Medford, Oregon; and Robert Bega retired as a research plant pathologist from the USDA Forest Service's Pacific Southwest Forest and Range Experiment Station, Berkeley, California. *Other names have been given to the fungus that causes black stain root disease: Ophiostonia wagneri (Goheen and Cobb) Harrington is the name of the sexual stage; Ceratocystis wageneri, and Verticicladiella wageneri are no longer considered taxonomically correct. 1 Hosts ponderosum) attacks hard pines and The principal hosts of L. wa- occasionally hemlocks. Although lim- generi are Douglas-fir, ponderosa ited cross-over has been shown experi- pine, Jeffrey pine, pinyon, and sin- mentally with the Douglas-fir and hard gleleaf pinyon. Seldom-damaged pine variants, natural occurrences have hosts in the United States include not been detected. For purposes of lodgepole pine, western hemlock, resource management, each of the mountain hemlock, sugar pine, west- variants can be considered host spe- ern white pine, and knob-cone pine. cific. Tree species highly resistant or im- mune to infection are "cedars" (Alaska Black stain root disease occurs over a yellow-cedar, Port-Orford-cedar, wide range of environmental condi- western redcedar, and incense-cedar), tions, from the hot and semi-arid spruce, larch, junipers, and true firs. Southwest to cool Pacific coastal ar- eas. Soil type does not appear to be a Recent research indicates that there major factor in the distribution of the are three distinct variants of L. wa- disease. Soil moisture and tempera- generi, each with specific host prefer- ture, however, may influence disease ences. One (var. wageneri) attacks distribution significantly. Cool, moist pinyon, another (var. pseudotsugae) soil conditions in spring and early attacks Douglas-fir, and the third (var. summer are ideal for infection, Figure 1 — Crown symptoms of black stain Figure 2 — Faded, sparse crown of pon- root on young Douglas-fir. derosa pine with black stain root disease. 2 growth, and tree-to-tree spread of the pathogen. Symptoms and Damage Trees infected by the black stain fungus usually exhibit symptoms of gradual de- cline before they die (figures 1 and 2). In early stages of decline, terminal growth is reduced and older needles become chlorotic. As the disease progresses, older needles are shed prematurely, new needles are somewhat stunted and yellow, and re- duced internodal growth is evident on lat- eral branches. In advanced stages, new growth is compact and chlorotic, with a tufted appearance, and tree crowns exhibit very sparse foliage. They also bear "distress" cone crops. Very small trees, or those affected by other significant stress fac- tors, may succumb quickly without exhibit- Figure 3 — Typical dark sapwood stain as- sociated with black stain root disease. Figure 4 — Cross section through affected root showing arc-shaped pattern of black stain. 3 hibiting gradual decline symptoms; fo- liage may change rapidly from green to yellow or reddish brown. Crown symptoms of trees affected by black stain root disease are very simi- lar usually a lighter color and typically are wedge-shaped in cross-section or they discolor the entire sapwood ra- dius. Black stain root disease affects groups of trees in distinct infection centers (figures 5-7). Typical infection centers have trees in various stages of decline near the perimeter and dead trees in the interior nearer the origin of initial infection. Infection centers usually oc- cur in well-stocked stands where a preferred host predominates or occurs Figure 5 — Black stain root disease center in in unmixed clumps. In stands where a second-growth ponderosa pine plantation. species composition is well mixed, Figure 6 — Typical black stain root disease center in a Douglas-fir plantation. 4 Figure 7 — Black stain root disease center in singleleaf pinyon. Figure 8 — Insect vectors of the black stain root disease fungus on Douglas-fir: Hylastes nigri- nus, Steremnius carinatus, and Pissodes fasciatus. 5 infection and mortality are less com- roots of recently dead or dying host mon, but isolated trees or tree clusters trees including those infected by L. can be infected. wageneri. Several vector species—H. nigrinus, P. fasciatus, and S. carina- Disease Cycle tus—also breed in Douglas-fir stumps, Long-distance spread of the black which are susceptible to infection by stain fungus involves insect vectors L. wageneri for up to 7 months after (figure 8). The root-feeding barkbee- stems have been severed. tles Hylastes macer LeConte and H. nigrinus (Mannerheim) are believed to Leptographium wageneri sporulates be the primary vectors of the fungus readily inside insect galleries in in- on ponderosa pine and Douglas-fir, fected roots and root collars. Stalked, respectively. Two weevils—Sterem- microscopic fruiting bodies nius carinatus (Boheman) and Pis- (conidiophores) form on gallery sodes fasciatus LeConte—have also walls, each bearing a sticky spore been implicated as vectors in Douglas- droplet that protrudes into the gallery. fir, but their involvement in local and These sticky spore droplets are well long-distance spread of the pathogen suited to insect dispersal (figure 9). is still poorly understood. During emergence, some young adult beetles are contaminated with spores Vector insects commonly breed in as they brush against spore droplets in galleries or pupal chambers. Contami- Figure 9 — Enlargement (25x) of fruiting bodies of the black stain root disease fungus in an insect gallery, showing sticky spore masses. 6 -nated beetles fly (or walk) from brood the pathogen through soil. Small trees, burrow through the duff and soil, rootlets (< 5 mm., or .2 in diame- visit roots of healthy, recently dead, or ter) serve as infection courts where dying trees, and deposit spores on root roots are not grafted. Infection sapwood exposed during feeding. centers in ponderosa pines and Wounds that expose sapwood xylem pinyons enlarge on the margin at are required for infection, because L. an average annual rate of 1.0 m/yr wageneri hyphae are unable to pene- (3.2 ft/yr). Infection centers in trate live bark and cambial tissues. Douglas-fir enlarge at a rate of 0.8 to 1.5 m/yr (31.5 to 59.1 in/yr), but Once established, the black stain fun- enlargement of infection centers gus colonizes root and stem sapwood with this host often decreases xylem, reducing water uptake and the markedly when stands reach 30 to vertical ascent of xylem sap by clog- 35 years of age. The rate of en- ging water conducting vessels with hy- largement of black stain infection phae and host reaction compounds. In centers is 3 to 5 times faster than live trees, the fungus is confined to that of other common root dis- tracheids; parenchymatous tissues are eases. not invaded. Infected trees die (usually in summer, when soil moisture is de- Leptographium wageneri is rela- pleted) because of inability to take up tively nonpersistent in infected water. root systems, remaining active usually no more than 1 year after Black stain infection centers are most its host dies. prevalent in areas where substantial tree damage or site disturbance has Management and Cultural occurred, especially along roads and Controls skid trails, in areas having a history of In recent years, black stain root disease tractor logging and resultant soil com- has been detected in many new areas, paction, and in areas that have been often causing locally severe damage. precommercially thinned. The appear- Incidence appears to be steadily in- ance of infection centers in areas of creasing.