UNIVERSITY OF CALIFORNIA COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION BERKELEY, CALIFORNIA CIRCULAR No. 289 May, 1925 OAK FUNGUS IN ORCHARD TREES BY A. H. HENDEICKSON' ^^ Oak fungus (Armillaria mellea) is a root disease of widespread economic importance in California. It attacks most of the common deciduous fruit trees and is also found on many other plants including annuals and perennials. As its common name indicates, the native oak trees are generally considered the source of infection. The disease spreads underground and often kills orchard trees over a considerable area. Trees replanted in infected orchards from which diseased trees have been removed usually die within a few years unless resistant species are used. APPEARANCE AND BEHAVIOE The characteristic behavior of trees infected with this disease makes it comparatively easy to identify the fungus. Diseased trees may die at any time during the year. Their roots show flat white fan-shaped mycelium (a thin felted sheet of fungous threads) between the bark and the wood. This mycelium is usually confined to the roots, and only in exceptional cases does it extend into the trunk above ground. The odor of a freshly dug root from a diseased tree is distinctive and is described by W. T. Home as having a ''sharp, rather agreeable mush- room smell." The presence of the flat white mycelium and peculiar mushroom odor serves as an excellent means of identifying this disease. Roots of fruit trees attacked by it decay rather quickly. The advanc- ing edge of the fungus is usually shown by a darkening of the wood. Slender brown or purple thread-like cords are sometimes found on the surface of the roots. These root-like organs are the rhizomorphs by which the fungus spreads underground from tree to tree. It is believed that the rhizomorphs are the principal means of spreading the disease. They are not always present in abundance on fruit tree roots and are often passed unnoticed. Whenever one of these rhizomorphs * Assistant Professor of Pomology, University of California. Z UNIVERSITY OF CALIFORNIA^ EXPERIMENT STATION comes in contact with a tree root, it quickly enters and in time infects the entire root system. During the rainy season clumps of toadstool sporophores often appear near the crown of the diseased trees. These toadstools are of a light tan color on the top, usually showing light brown scales. The loAver surface is white and is made up of fine delicate structures known as gills, which bear the spores or reproductive bodies. The size of these clumps varies considerably. They may consist of comparatively few toadstools, or they may be as much as two feet in diameter. The spores, which are produced in great numbers, apparently do not attack living plants, and hence there is nothing to be gained by destroying the toadstools. Ordinarily, the whole clump turns dark within a few days, becomes infested with the larvae of certain fungous gnats, and melts down to a slimy mass. While the spores are not thought to be important in disseminating the disease, the presence of one of these clumps indicates that the fungus is established in the tree and that the tree is doomed to die within a comparatively short time. Usually, the whole tree dies suddenly (fig. 1) but occasionally one side may die and leave the remainder apparently healthy. This condition seems due to the fact that the roots on one side are killed before the disease has become established throughout the root system. The remainder of the tree generally dies within a year or two. When a fruit tree has once become infected, it is inevitably killed. After the disease reaches the crown, treatments with disinfectants or by surgery have not proved effective. There are but few, if any, authentic cases where an infected tree has survived more than one or two seasons after treatment. One of the chief characteristics of the disease is its manner of spreading. From the original source of infection, the disease spreads outward from tree to tree in all directions. The infected area is irregularly circular in outline, and the border usually contains one or more trees showing various stages of injury. SOUECES OF INFECTION In the treatment of any plant disease, a knowledge of the source of infection is important in order that the disease may be stopped, if possible, before it attacks plants under cultivation. Armillaria mellea is found on many native species, including the oak, willow and madrono. The indigenous oak trees probably form the chief source of infection, but it is not known whether all oaks carry the disease. It is not difficult to find, in orchards which have been long owned by one family, authentic cases where the disease has developed in areas from which oak trees had been removed many years before. Oak roots CiRC. 289] OAK FUNGUS IN ORCHARD TREES decay very slowly and may remain in the soil twenty or thirty years after the tree has been cut down. It seems as if the disease attacks the roots of the fruit trees only when they come into contact with the oak root fragments. Thus, the fruit trees may not be killed for many years after the orchard is planted. Apparently the pieces of oak roots are a potential source of infection until after they are completely decayed. It has also been observed that many Armillaria spots develop in orchards planted along streams or old water courses. The source of Fig, 1. — Mature apricot tree killed by oak fungus during the growing season. infection in these cases may be the native vegetation along the banks or the fragments of roots carried down by the flood waters and deposited on the adjoining land and covered with sediment. Nursery trees may also form a source of infection, as the disease has been found, in a few cases, on nursery trees. It is probable that only a very small proportion of trees are diseased as they come from the nursery. In Tasmania,* Armillaria has been reported to have spread from underground drains made of wood. When tile was substituted for the wood, the rate of spread of the disease was appreciably diminished. * Bulletin 57. Root-Eot of Fruit Trees, by H. M. Xieholls Stock Department, Tasmania. Agricultural and 4 UNIVERSITY OF CALIFORNIA EXPERIMENT STATION Relative Susceptihility of Species. — Nearly all of the deciduous fruit trees grown in California are susceptible to the disease. Only three exceptions have thus far been discovered. The three species w^hose resistance seems to be vv^ell established, are the French pear, Pyrus communis; Northern California Black Walnut, Juglans cali- fornica hindsii; and the Fig, Ficus carica. Oak fungus also attacks raspberries, blackberries, and many ornamental shrubs, and has been found on rhubarb and potato plants. English or Persian walnuts (fig. 2) and many hybrids between the English and other species are not resistant. Prunus davidiana, recently introduced as a stock for peaches, and pear trees on Japanese stock, which may include a mixture of several species, show more or less susceptibility. Some of the pear root stocks used recently for their resistance to pear blight, have not been grown long enough in districts infected with Armillaria to determine their degree of resistance. Of the three principal rootstocks used for stone fruits, observation seems to indicate that myrobalan is the most resistant, surpassing either the peach or the almond root. Some growers are replanting their infected areas with myrobalan seedlings, to be topworked later with the desired varieties. None of these plantings are as yet old enough to show decisively whether or not this method is better than the usual practice of planting budded nursery trees. In some districts the apricot root on heavy soil seems to show a comparatively high degree of resistance, but on sandy soil, this root is killed very quickly. Peaches and cherries are known to be susceptible. Factors in the Bate of Spread of the Fungus. — The kind of soil iti which the fruit trees are growing usually seems to exert little influence on the rate of spread of the fungus. The disease has been found in soils of many types, ranging from light sand to heavy clay and adobe, but, dry, gravelly soils and the sandy sediment soils often found along creek beds seems to show a somewhat more rapid spread of the disease than do the heavy clay types. In some parts of the state, there is an impression among growers that irrigation tends to increase the rate of spread of the disease, but this belief has not, as yet, been substantiated. Irrigation may be a means of causing the disease to die out by hastening the decay of infected roots. This, if true, would be a means of quickly making an infected piece of soil safe for planting. The rate of spread of the disease shows great variation in different regions. The loss may range from one or two to forty or fifty trees a year. Where the annual losses are large it appears that several diseased spots have coalesced to form CiRC. OAK FUNGUS IN ORCHARD TREES a single large area, covering sometimes several acres. Limited observa- tions seem to show that with prunes and apricots in adobe soil, the rate of spread is considerably slower than where these fruits are on gravelly or loamy soil. The vigor of the individual tree does not seem to have any effect on its resistance to the disease. Young and vigorous trees are as liable to attack as old ones. With old trees, the presence of the disease may Fig. 2. — Walnut tree on English walnut root killed by oak fungus. be suspected for a year or more before death occurs, but young trees die very quickly after the first indications are given by the drying and falling off of the leaves. The belief has long been held among growers that so long as its native host is alive, the fungus does not leave it to attack fruit trees. While this contention may be difficult to prove, it is an interesting fact that many examples have been found of prune and apricot trees grow- ing for years in close proximity to oak trees without contracting the b UNIVERSITY OF CALIFORNIA EXPERIMENT STATION disease. There is some evidence to show that Armillaria exists in an oak tree but does not kill it. While the oak tree is living, the fungus is apparently unable to penetrate to the cambium layer. After the death of the host the disease spreads throughout the dead or dying roots. In the case of fruit trees, the fungus is apparently able to penetrate to the cambium immediately. History of a Typical Orcliard Infected with Armillaria. — A strik- ing example of how oak fungus spreads is shown in the accompanying map of an orchard in Santa Clara County (fig. 3). The part of the orchard shown on the map originally contained 1297 prune and apricot trees planted in 1894 on the square system, 22 feet apart. French prunes, part of which were grafted on myrobalan and part on almond root, made up the majority of the planting. The apricots were top- worked to Sugar prunes when seven years old. It is interesting to note that these trees, as shown in the northeast corner of the map, have withstood the advance of the disease fully as well as, if not better than, the French variety on Myrobalan and on almond root. Losses first occurred when the trees were five or six years old, and have been continuous since that time. Where trees died the vacancies were refilled with prune trees, but these second plantings did not long survive. Next, apples were tried, and although somewhat more resist- ant to the fungus, they were removed because this fruit did not pay. Then, for a time the vacant spaces were devoted to garden vegetables and alfalfa. Finally, walnuts were planted. Part of the first planting of walnuts was on English root. These trees were killed after a few years. The remaining walnut trees which were grafted on the Cali- fornia black w^alnut stock survived and at the present time are about 12 or 14 years old and are bearing satisfactory crops. Judging from the apparent age of the replanted walnut trees the area now infected with oak fungus started from four or five points or sources. These original spots gr?vv in size and finally coalesced to form the irregular shaped area shown within the dotted lines on the map (p. 7). It seems that one area began near the lower southeast corner, where the orchard approaches the banks of a creek. It is probable that infection started from the native trees along the bank. Other infected areas developed in various places and gradually spread until they came together to form one large diseased patch. The owner of the orchard still remembers the approximate location of the oak trees which were cut down more than 30 years ago. In some cases the location of these oaks apparently coincides with what seem to have been the beginning points of the disease in the orchard. Giro. 289" OAK FUNGUS IN ORCHARD TREES VYV ScxvOcoc CVo^t(L Co. • •\) • •/ X X vO ooJ%. is % K. X X X X X X ;t y X X X X X X X o X X o X X X XX X XXXXX XX X XXXXXXX XX xxxxxx xx>t y< X X X. XXX X X XXXXX X X X XOXX XXXXX X X. X X X X X xxxxxxxx O X X X X X X % o a X X X X xoxK X X xxxxxox XX XX XXXXX xxxxxxxx X X X X X XX XP XXXXX .'O '\, X X X X X X X X *''^9,o \_x XXXXX x''\o "o o X X X X x» XX x\o O X X / X X X X X X', o X X y- X XXOAXX X;0 X X X X xxA x;o X X X X X X XXX x" XX X X>txAX X X X X X X XXXXX X X X X XXXX X X X X. XXXXXXX X Fig. 3. — A typical example of an Armillaria-infected orchard, and how the disease slowly but surely spreads over the whole area if not checked during its early stages. This part of the orchard originally contained 1297 prune and apricot trees planted in 1894. Missing places are now principally planted to walnut trees on resistant stocks. 8 UNIVERSITY OF CALIFORNIA EXPERIMENT STATION At the present time approximately 860 trees of the original plant- ing have been killed. This amounts to a loss of nearly 66 per cent in 23 years. In other words, the average loss was about 37 trees a year on an area of about 14 acres. The economic loss resulting from the death of 37 mature prune trees each year, together with the non- productive period which must elapse before the replanted walnuts come into bearing amounts to a considerable sum. A study of this orchard, which is no different from many others in various parts of the state, is of peculiar interest, because of the very evident lessons it imparts. It shows first, the inexorable progress of the fungus when once established in a prune or apricot orchard ; second, the economic loss resulting from the death of bearing trees; and third, the best solution as yet found for this problem : viz., replant- ing with disease-resistant trees such as walnuts on California black Avalnut root. METHODS OF CONTEOL Up to the present time no method of treatment intended to control this disease, after it is discovered, has been effective. The solution of the problem seems to lie in lessening the losses by checking the spread of the fungus. Isolating the diseased trees by digging trenches around the diseased area or by replanting with resistant stocks after the orig- inal trees have died will accomplish this. Trenching around the infected area at a safe distance beyond the visible limits of the fungus may be expected to give a reasonable amount of protection for the remainder of the orchard. This method, however, is practicable only when the diseased area is relatively small. It is also beset with danger because diseased roots may be left outside the barrier. In the case of many plant diseases, one of the principal methods of control is by pruning off diseased parts and by burning or plowing under infected material. This method is not applicable to controlling a root disease of a large tree. When a diseased tree is dug out of the soil, it is not practicable to remove all of the roots. The idea of disinfecting the soil presents itself in this connection. Various chemicals have been tested for this purpose. Up to the present time, the tests have not been particularly successful. Disin- fecting the holes from which diseased trees have been removed with such materials as copper sulfate, formalin, sodium cyanide and carbon bisulfide has not proved to be a reliable method of permanently pre- venting infection of the replanted trees. Neither has the application of hydrated lime to the soil around trees adjoining diseased areas proved to be effective in preventing the spread of the fungus (fig. 4). CiRC. 289] OAK FUNGUS IN ORCHARD TREES In Tasmania, excellent results are reported from the application of permanganate of potash or iron sulfate on the main roots of trees infected with Armillaria. Similar treatments (fig. 4) applied to trees in California using Weed Killer (proprietary arsenic compound).* Fig. 4. — Partly diseased prune tree with main roots uncovered for treatment with disinfecting materials. No useful results obtained. permanganate of potash/* iron sulfate,* formaldehyde and Bordeaux, all failed to control the disease. Likewise, treatments w^ith various * Tliese materials have been used by Col. G. E. Stockle of Mountain View on a c-onimercial scale for several vcars without success. 10 UNIVERSITY OF CALIFORNIA EXPERIMENT STATION chemicals by so-called tree doctors have not checked the spread of the fungus. Barriers. — Armillaria spreads underground from tree to tree by means of root-like fungous strands known as rhizomorphs. Healthy »^.r Fig. 5. — Portion of a narrow concrete barrier built around an oak fungus area to prevent spread of the disease. trees are thus infected from adjacent diseased ones. Theoretically, the disease can be checked by preventing the underground spread of these rhizomorphs. Various methods to accomplish this purpose have been suggested and carried out to a limited extent. The chief idea CiRC. 289] OAK FUNGUS IN ORCHARD TREES 11 in this method of control is to dig around the diseased area a trench of sufficient depth to extend below the area occupied by roots. This trench may simply be refilled after cutting off all crossing roots and later reopened for the same purpose ; or it may be lined with material J Fig. 6.^J'.c(-;iusr tills uiii' hUiali infected iv-; - .,.,-.., ...- wall, the barrier, faintly shown on the right of the photograph, failed to prevent the spread of the fungus. The barrier should have been placed one tree row farther away from the diseased area. like tar-paper; or it may have constructed in it a narrow concrete (fig. 5) wall extending from slightly below plowing depth to the bottom of the trench. 12 UNIVERSITY OF CALIFORNIA EXPERIMENT STATION The chief objection to the use of the concrete barrier is its cost. The enclosed area is, therefore, usually made as small as possible. The danger in this practice lies in the fact that although the trees bordering the diseased area are apparently healthy, they may have one Fig. 7. — Stump of English walnut tree killed by oak fungus. The graft union was covered with soil and the disease entered the scion above the graft union. The black walnut root was not injured and sent up several sprouts after the top had died. or more infected roots. In this case, the disease soon makes its appear- ance outside of the barrier. This has actually happened in two instances known to the writer. In one case when a tree outside of the CiRC. 289] OAK FUNGUS IN ORCHARD TREES 13 barrier died, the source of infection was traced to a root which extended to the wall, but which was not observed to be infected when the barrier was built (fig. 6). For safety the barrier should be put not closer than between the second and third rows of healthy trees surrounding the diseased area. CONCLUSIONS Oak fungus is a root disease attacking many common deciduous fruit trees and ornamentals. All root stocks used at the present time for stone fruits are more or less susceptible to attack. Apples are usually considered highly resistant, especially if grown on Delicious stock, and the French pear root is practically immune. The fig and California black walnut are also highly resistant to this fungus. If the infected area is limited in extent, as in the case of young orchards when the disease is first noticed, it may be possible to check further progress of the fungus by enclosing the diseased spot with a concrete barrier as described in this paper. When the diseased area is extensive it should be replanted with one or more of the resistant species. In replanting with either pears on French root or with walnuts on black walnut root, care must be taken to see that the graft union (fig. 7) is not covered with soil, because cases have been observed w^here the susceptible scion was attacked by the disease, while the root Avas not affected. STATION PUBLICATIONS AVAILABI^E FOR FEEE DISTRIBUTION BULLETINS No. No. 253. Irrigation and Soil Conditions in the 352. Sierra Nevada Foothills, California. 261. Melaxiima of the Walnut, "Juglans 353. regia." 354. 262. Citrus Diseases of Florida and Cuba 357. Compared with Those of California. 263. Size Grades for Ripe Olives. 268. Growing and Grafting Olive Seedlings. 358. 273. Preliminary Report on Kearney Vine- yard Experimental Drain. 359. 275. The Cultivation of Belladonna in Cali- 361. fornia. 276. The Pomegranate. 362. 277. Sudan Grass 363. 278. Grain Sorghums. 279. Irrigation of Rice in California. 364. 280. Irrigation of Alfalfa in the Sacramento Vallev. 366. 283. The Olive Insects of California. 285. The Milk Goat in California. 367. 286. Commercial Fertilizers. 294. Bean Culture in California. 368. 304. A Study of the Effects of Freezes on Citrus in California. 369. 310. Plum Pollination. 370. 312. Mariout Barley. 371. 313. Pruning Young Deciduous Fruit Trees. 319. Caprifigs and Caprification. 372. 324. Storage of Perishable Fruit at Freezing Temperatures. 374. 325. Rice Irrigation Measurements and Ex- periments in Sacramento Valley, 1914-1919. 375. 328. Prune Growing in California. 331. Phylloxera-Resistant Stocks. 376. 334. Preliminary Volume Tables for Second- Growth Redwood. 377. 335. Cocoanut Meal as a Feed for Dairy 379. Cows and Other Livestock. 380. 339. The Relative Cost of Making Logs from Small and Large Timber. 381. 340. Control of the Pocket Gopher in Cali- fornia. 382. 343. Cheese Pests and Their Control. 344. Cold Storage as an Aid to the Market- 383. ing of Plums. 346. Almond Pollination. 384. 347. The Control of Red Spiders in Decidu- ous Orchards. 348. Pruning Young Olive Trees. 385. 349. A Study of Sidedraft and Tractor 386. Hitches. 350. Agriculture in Cut-over Redwood Lands. 287. Further Experiments in Plum Pollina- tion. Bovine Infectious Abortion. Results of Rice Experiments in 1922. A Self-mixing Dusting Mackine for Applying Dry Insecticides and Fungicides. Black Measles, Water Berries, and Related Vine Troubles. Fruit Beverage Investigations. Preliminary Yield Tables for Second Growth Redwood. Dust and the Tractor Engine. The Pruning of Citrus Trees in Cali- fornia. Fungicidal Dusts for the Control of Bunt. Turkish Tobacco Culture, Curing and Marketing. Methods of Harvesting and Irrigation in Relation to Mouldy Walnuts. Bacterial Decomposition of Olives dur- ing Pickling. Comparison of Woods for Butter Boxes. Browning of Yellow Newtown Apples. The Relative Cost of Yarding Small and Large Timber. The Cost of Producing Market Milk and Butterfat on 246 California Dairies. A Survey of Orchard Practices in the Citrus Industry of Southern Cali- fornia. Results of Rice Experiments at Cor- tena, 1923. Sun-Drying and Dehydration of Wal- nuts. The Cold Storage of Pears. Walnut Culture in California. Growth of Eucalyptus in California Plantations. Growing and Handling Asparagus Crowns. Pumping for Drainage in the San Joaquin Valley, California. Monilia Blossom Blight (Brown Rot) of Apricot. A Study of the Relative Values of Cer- tain Succulent Feeds and Alfalfa Meal as Sourses of Vitamin A for Poultry. Pollination of the Sweet Cherry. Pruning Bearing Deciduous Fruit Trees. Fig Smut. No. 87 113 117 Alfalfa. Correspondence Courses in Agriculture. The Selection and Cost of a Small Pumping Plant. House Fumigation. 129. The Control of Citrus Insects. 136. Melilotus indica as a Green-Manure Crop for California. Oidium or Powdery Mildew of the Vine. Feeding and Management of Hogs. Some Observations on the Bulk Hand- ling of Grain in California. Irrigation Practice in Growing Small Fruit in California. 127. 144. 151. 152. 154. CIRCULARS No. 155. 157. 160. 161. 164. 165. Bovine Tuberculosis. Control of the Pear Scab. Lettuce Growing in California. Potatoes in California. Small Fruit Culture in California. Fundamentals of Sugar Beet Culture under California Conditions. 166. The County Farm Bureau. 167. Feeding Stuffs of Minor Importance. 170. Fertilizing California Soils for the 1918 Ci'op. 173. The Construction of the Wood-Hoop Silo. 178. The Packing of Apples in California. CmCVhARS— (Continued) No. 179. 184. 190. 199. 202. 203. 208. 209. 210. 212. 214. 215. 217. 220. 228. 231. 232. 233. 234. 235. 236. 237. 238. 239. 240. 241. 243. 244. 245. 247. 248. 249. 250. Factors of Importance in Producing Milk of Low Bacterial Count. A Flock of Sheep on the Farm. Agriculture Clubs in California. Onion Growing in California. County Organizations for Rural Fire Control. Peat as a Manure Substitute. Summary of the Annual Reports of the Farm Advisors of California. The Function of the Farm Bureau. Suggestions to the Settler in California. Salvaging Rain-Damaged Prunes. Seed Treatment for the Prevention of Cereal Smuts. Feeding Dairy Cows in California. Methods for Marketing Vegetables in California. Unfermented Fruit Juices. Vineyard Irrigation in Arid Climates. The Home Vineyard. Harvesting and Handling California Cherries for Eastern Shipment. Artificial Incubation. Winter Injury to Young Walnut Trees during 1921-22. Soil Analysis and Soil and Plant Inter- relations. The Common Hawks and Owls of Cali- fornia from the Standpoint of the Rancher. Directions for the Tanning and Dress- of Furs. The Apricot in California. Harvesting and Handling Apricots and Plums for Eastern Shipment. Harvesting and Handling Pears for Eastern Shipment. Harvesting and Handling Peaches for Eastern Shipment. Marmalade Juice and Jelly Juice from Citrus Fruits. Central Wire Bracing for Fruit Trees. Vine Pruning Systems. Colonization and Rural Development. Some Common Errors in Vine Pruning and Their Remedies. Replacing Missing Vines. Measurement of Irrigation Water on the Farm. No. 251. Recommendations Concerning the Com- mon Diseases and Parasites of Poultry in California. 252. Supports for Vines. 253. Vineyard Plans. 254. The Use of Artificial Light to Increase Winter Egg Production. 255. Leguminous Plants as Organic Fertil- izer in California Agriculture. 256. The Control of Wild Morning Glory. 257. The Small-Seeded Horse Bean. 258. Thinning Deciduous Fruits. 259. Pear By-products. 260. A Selected List of References Relating to Irrigation in California. 261. Sewing Grain Sacks. 262. Cabbage Growing in California. 263. Tomato Production in California. 264. Preliminary Essentials to Bovine Tuber- culosis Control. 265. Plant Disease and Pest Control. 266. Analyzing the Citrus Orchard by Means of Simple Tree Records. 267. The Tendency of Tractors to Rise in Front; Causes and Remedies. 268. Inexpensive Lavor-saving Poultry Ap- pliances. 269. An Orchard Brush Burner. 270. A Farm Septic Tank. 271. Brooding Chicks Artificially. 272. California Farm Tenancy and Methods of Leasing. 273. Saving the Gophered Citrus Tree. 275. Marketable California Decorative Greens. 276. Home Canning. 277. Head, Cane, and Cordon Pruning of Vines. 278. Olive Pickling in Mediterranean Coun- tries. 279. The Preparation and Refining of Olive Oil in Southern Europe. 281. The Results of a Survey to Determine the Cost of Producing Beef in Cali- fornia. 282. Prevention of Insect Attack on Stored Grain. 283. Fertilizing Citrus Trees in California. 284. The Almond in California. The publications listed above may be had by addressing College of Agriculture, University of California, Berkeley, California. 12m-5,'25