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Historic, Archive Document Do not assume content reflects current scientific knowledge, policies, or practices. Forest Insect sae DYW er:Ws Leaflet 49' Revised September 2003 U.S. Department of Agriculture ¢ Forest Service Dwarf Mistletoes of Ponderosa Pine __ Jerome S. Beatty’ and Robert L. Mathiasen* The dwarf mistletoes (Arceuthobium Southwestern dwarf mistletoe (A. Spp.) are important parasites found vaginatum subsp. cryptopodum) throughout most of the range of pon- occurs on the Rocky Mountain vari- derosa pine (Pinus ponderosa). ety of ponderosa pine (Pinus pon- derosa var. scopulorum) from Ponderosa pine 1s parasitized by Arizona and New Mexico into central two widespread species. Utah and northern Colorado. Western dwarf mistletoe (A. campylopo- dum) affects the Pacific coast variety of ponderosa pine (P ponderosa var. pon- derosa) from southern California to northern Washington and western Idaho (Figure 1). Although both dwarf mistletoes occur primarily on ponderosa pine, they | also commonly infect some related pines (Table 1). Southwestern dwarf mistletoe is equally damag- | ing to Arizona pine (Pinus arizonica) and Apache pine (Pinus engelmannii) in the Figure 1. Distribution of OY southwestem ) southwestern and western bm dwarf mistletoe ; dwarf mistletoes (shaded CAA Pliers: Ss areas) and ponderosa pine Ld Meiovaa' = (thick blue line). 'This Insect and Disease Leaflet combines Leaflet #19 “Dwarf mistletoes of Ponderosa Pine in the Southwest” and #40, “Western dwarf mistletoe on Ponderosa Pine.” * Jerome Beatty is Deputy Director, USDA Forest Service, Forest Health Protection, Washington, DC; * Robert Mathiasen is Associate Professor, School of Forestry, Northern Arizona University, Flagstaff, AZ. I Southwest. Similarly, western dwarf Dwarf mistletoes do not have mistletoe frequently infects Jeffrey large leaves; instead their leaves are pine (Pinus jeffreyi) and Coulter pine small squamate scales found at the (Pinus coulteri) in California. nodes of shoots. The average height of aerial shoots for these mistletoes is Both mistletoes will also attack about 4 to 6 inches (10-15 cm). some introduced pines; those hosts are listed in Table 1. Aerial shoots arise from a network Table 1. Hosts ofs outhwestern and western dwarf mistletoe. Susceptibility Class | Southwestern Dwarf Mistletoe | Western Dwarf Mistletoe (Arceuthobium vaginatum) (Arceuthobium campylopodum) Principal hosts Rocky Mountain ponderosa pine | Pacific coast ponderosa pine Arizona pine Jeffrey pine Apache pine Secondary hosts’ Coulter pine Knobcone pi Occasional hosts? Rocky Mountain bristlecone pine | Grey pine Rocky Mountain lodgepole pine Rocky Mountain lodgepole pine Sierra lodgepole pine Rare hosts’ Limber pine Sugar pine Southwestern white pine ‘Trees frequently attacked when they are associated with infected principal hosts *Trees occasionally attacked when associated with infected principal host *Trees rarely attacked, even when associated with infected principal hosts Life History of root-like absorbing strands a imbedded in host tissues. This net- The external (aerial) shoots of west- work, called the endophytic system, ern dwarf mistletoe are olive green to consists of cortical strands growing brown or yellow-brown (Figure 2). within the bark and sinkers within the The aerial shoots of southwestern wood. The endophytic system dwarf mistletoe are orange to reddish- obtains nutrients and water from the brown to almost black (Figure 3). host tree. Figure 2. Female shoots of western dwarf Table 2. Flowering and seed dispersal periods of the two ponderosa pine dwarf mistletoes. Characteristic Southwestern Dwarf Mistletoe | Western Dwarf Mistletoe (Arceuthobium vaginatum) (Arceuthobium campylopodum) Flowering period August - October Seed dispersal period | July - August September - November Germination Seeds germinate immediately Seeds overwinter and germinate after seed dispersal fhe following April - May The endophytic system usually in length. Seed dispersal is one of lives as long as adjacent host tissues the most interesting characteristics of are alive. The mistletoe is dependent dwarf mistletoes. Seeds are dis- upon its host trees for water and charged explosively from ripe fruits nutrients, and most of its carbohy- in late summer for southwestern drates. Although the aerial shoots dwarf mistletoe and early fall for contain chlorophyll that allows them western dwarf mistletoe (Table 2). to produce small amounts of carbohy- Seeds may travel 10-13 m, but drates, the major function of aerial most land within 3-5 m of the dis- shoots 1s reproduction. Male and female flowers are seminating shoot. A sticky seed coat- small and produced on separate ing called viscin enables seeds to plants. Southwestern dwarf mistletoe stick to most objects they strike. flowers in the spring and western Foliage is the most common receiv- dwarf mistletoe in the fall (Table 2). ing surface. Viscin acts as a lubricant Insects and wind pollinate female when moistened by rain. Seeds slide flowers. down and either fall off needles or become lodged on bark near the base Fruits complete their develop- of needles. Seeds are fastened in ment 12 tol5 months after pollina- place when the viscin dries. tion. The mature fruits contain one seed averaging about ().2 inches (3 mm) Seeds are often destroyed by visible. When shoots fall off of insects and fungi or dislodged by rain infected branches they leave behind or snow, so only a small proportion small cup-like structures embedded in of the seeds dispersed actually sur- the bark. These are called basal cups. vive and cause new infections. As time passes and the endophyt- Seeds of southwestern dwarf ic system of the parasite becomes mistletoe germinate immediately after more extensive, the branching habit dispersal, while those of western of the host may become distorted and dwarf mistletoe over-winter and ger- witches’ brooms are formed. minate in early spring. Witches' brooms are variously shaped masses of abnormal branch and twig A root-like structure called a growth. Broomed branches usually radicle emerges from the seed and outlive their uninfected neighbors by grows along the bark surface until an many years. They divert food from obstruction, usually a needle base, 1s uninfected parts of the tree and thus encountered. The radicle then forms are instrumental in reducing vigor a mound of tissue called a holdfast. and causing premature death of The holdfast develops an infection severely infected trees. peg, which penetrates into host tissue. The mistletoe's endophytic system Long-lived witches' brooms also then develops in the outer bark and result in excessively large knots; it is wood of the infected part of the host. not uncommon for the base of broomed branches on otherwise clear Infection occurs most readily in trunks to become over 10 inches (25 twigs less than five years old because cm) in diameter. Resinous cankers their bark is more easily penetrated may result from mistletoe infection of than older twigs. the main stem, especially on smaller Aerial shoots typically appear 2 trees. to 3 years after initial infection. Severely infested pine stands typ- Infections that have not yet produced ically have many trees with stunted aerial shoots are called latent infec- growth, witches' brooms, dying or tions. The typical length of time dead tops and branches, and dead needed for female plants to complete trees. Dieback occurs as nutrients their life cycle from initial establish- and water needed by growing tree ment to dissemination of the first tops are diverted to infections that are seed crop is 4 to 5 years for both of usually concentrated in the lower or these dwarf mistletoes. mid crowns. Symptoms and Signs of Foliage near infections becomes sparse and off-color and gradually the Infection upper branches die. Eventually height growth slows, and such trees The first symptom of dwarf mistletoe may subsequently be attacked and infection is the appearance of slight killed by bark beetles. These stands swellings at infection sites. Swellings eventually contain numerous dying become visible | to 2 years after and dead trees, usually bearing rem- infection occurs. Aerial portions of nants of witches’ brooms. male and female plants appear after a year or two after the swellings are Spread and Intensification crown third have infections). The three ratings are then added to obtain Several interrelated factors influence a tree rating ranging from 0 (healthy tree-to-tree spread of ponderosa pine trees) to 6 (severely infected trees). dwarf mistletoes. These include tree The tree ratings of all live trees in a size, tree spacing, stand structure, stand or plot (including uninfected species composition of stands, and ones) are then summed and the total infection position. divided by the number of live trees to obtain an average rating for the stand. In single-storied stands, spread is Average dwarf mistletoe ratings for a estimated to be 2-3 feet (0.6-1.0 m) stand can then be used to help deter- per year. Spread in multi-storied mine what management options can stands is more rapid because under- be considered for the infested area. story trees are exposed to dwarf mistletoe seeds from infected over- Dwarf mistletoe ratings can also story trees. Presence of non-host tree be used to help decide which trees species can slow the spread of pon- should be removed or retained if a derosa pine dwarf mistletoe. Spread decision is made to conduct harvest- rates in very dense stands are less ing or thinning treatments in the than in more open stands because infested area. dwarf mistletoe seed production is As a rough rule-of-thumb, inten- usually reduced due to limited light sification of pine dwarf mistletoe and lower host vigor, and many seeds averages about one DMR class per are trapped before they travel far. decade for individual trees, but varies Dwarf mistletoe seeds from plants with tree size, stand position, and high in tree crowns tend to travel far- overstory infection. Infection intensi- ther than those in lower portions of fies most rapidly in sapling or pole- crowns. size trees under severely infected Nearly all spread is local and larger trees. results from explosive discharge of Impacts seeds. Wind exerts a minor influence on distance and direction of seed travel. Birds and other animals are Infection of ponderosa pine by dwarf responsible for some long-distance mistletoe causes increased mortality, spread when seeds stick to them and reduced growth rates and loss of later are rubbed off onto susceptible vigor, lowered timber quality, TEES reduced cone and seed production, and increased susceptibility to other The 6-class dwarf mistletoe rat- damaging agents. These damaging ing (DMR) system is useful for quan- effects result from the dwarf mistle- tifying intensity of infection in trees toe plants taking food and water from and stands. For this system, the live the host, thus reducing the amount crown of the tree is visually divided available for the tree's normal growth, into thirds and each third rated as: 0 protective, and reproductive processes. = no visible infection, | = light infec- tion (less than half of the branches in The effect of dwarf mistletoe on the crown third have dwarf mistletoe growth increases with severity of infections), or 2 = heavy infection infection and is especially acute in (more than half of the branches in the severely infected trees (DMR = 5 or 6). Tree volume growth can be reduced by as much as 50% in Management of ponderosa pine dwarf severely infected trees. Severely mistletoes must recognize their value infected trees (DMR = 6) usually live as functional components of forest less than 10 years in the Southwest. ecosystems. In areas where timber production or developed recreation is Quality of lumber in infected the primary goal, direct control of trees can be reduced because dwarf dwarf mistletoe may be warranted. In mistletoe sometimes infects the boles other areas, where wildlife or esthetic of trees. Knots often are associated values are more important, maintain- with branches supporting witches' ing or even increasing dwarf mistle- brooms. Severely infected trees typi- toe populations may be appropriate. cally produce few cones, and those that are produced are smaller than Cultural Controls normal and contain fewer seeds. Dwarf mistletoes can adversely The only practical control of pon- affect other forest values. Witches' derosa pine dwarf mistletoes over brooms and dead branches can large forested areas is through cultur- increase the hazard potential in recre- al treatments. No chemical or biolog- ation sites because they may cause ical controls are available for treating trees and branches to break and fall. stands. Dead and dying trees detract from Profitable production of timber in visual quality. Potential for wildfires many local areas often depends on is increased because of resin soaked management of dwarf mistletoe. live branches, dead branches, Because the parasite can cause large increased tree mortality, and the accu- reductions in yield, management mulation of dead, resinous branches should be considered in all timber- around the bases of infected trees. producing stands where dwarf mistle- However, dwarf mistletoe infec- toes occur. tion can also have beneficial effects. Successful timber volume pro- Flowers, shoots, and fruits are food duction is almost impossible in multi- for insects, birds, and mammals. storied, infested stands. Silvicultural Mortality caused by dwarf mistletoe, treatments designed to achieve sin- either directly or by predisposing gle-storied stands offer the best trees to other agents, provides snags prospects of preventing unacceptable as habitat for cavity-nesting birds losses to dwarf mistletoes. and, eventually, coarse-woody debris on the forest floor. Dwarf mistletoe The most effective method for brooms create additional structure eliminating dwarf mistletoes from and various species of wildlife use timber-producing forests is complete the witches’ brooms as nesting sites. harvest of infested stands by clearcut- ting. After usable trees are harvested, Management all remaining infected trees should be killed. In forest ecosystems, dwarf mistle- To minimize invasion of young toes have value as individual, biologi- pine stands by dwarf mistletoe from cal species and act as disturbance infected border trees, the ratio of agents, influencing both the structure perimeter to area of clearcuts should and function of forest communities. be as low as possible. That is, cut decline rapidly about 10 years after openings should be roughly circular they are exposed to full sunlight by and not long, narrow strips. thinning. Advantage should be taken of Because of their rapid decline, any potential barriers to dwarf mistle- moderately infected trees should not toe spread, such as roads, ridgetops, be left when stands are being sani- natural openings, and changes in tim- tized unless they can be expected to ber types when laying out the bound- reach merchantable size within 15 ary of a clearcut. years. When clearcutting of dwarf Thinning priorities should be mistletoe-infested stands is not appro- based on both dwarf mistletoe severi- priate, shelterwood and seed tree har- ty and live crown ratios. First priori- vests can be good alternative even- ty for removal should be all trees age management methods. Trees with a live crown ratio < 40% or selected to provide shelter or seeds trees with live crown ratios < 60% should be uninfected or only lightly with a DMR of 5 or 6. Last trees to infected (DMR < 3). be removed should be those with a live crown ratio > 80% and a DMR <3. Moderately and severely infected trees, in addition to being a source of In stands where management of dwarf mistletoe seeds, produce poorer hiding and thermal cover for wildlife crops of tree seeds. Infected shelter- is a primary consideration, infected wood or seed trees should be trees with large witches' broom removed as soon as susceptible repro- should be treated by girdling rather duction becomes established. As a than felling. general rule for most dwarf mistle- Reexamination of stands 5 years toes it is desirable to remove the after sanitation is desirable to deter- infected overstory before the young mine if additional sanitation 1s need- stand is 3 feet (1 m) tall or 10 years ed. Moderately and severely infested old, whichever occurs first. stands should not be sanitized In mixed-species stands that con- because many trees with latent infec- tain pines infected by dwarf mistle- tions will be missed, and once these toe, silvicultural treatments should trees are released from competition, favor other tree species. Non-hosts the number of dwarf mistletoe plants left between infected and non-infect- will increase rapidly within their ed pines prevent or slow spread and crowns. Such stands should either be intensification of the parasite. clearcut and regenerated or left unthinned to avoid their rapid Thinning or sanitation by decline. removal of infected trees can be an effective treatment in lightly infested Latent dwarf mistletoe infections in stands. Lightly infested is defined as trees in densely stocked stands and in those stands in which there are shaded reproduction are very difficult acceptable numbers of desirable to detect. It should be assumed that dwarf mistletoe-free or lightly infect- understory trees more than 3 feet (1 ed (DMR <3) trees. Trees with one- m) tall that have been overtopped by half or more of their crowns infected infected trees for at least 10 years are by dwarf mistletoe (DMR >3) may probably infected. Recreation Management Assistance In recreation areas, sanitation treat- Resource managers can get more ments that favor non-susceptible information about the identification hosts or remove infected overstory and management of dwarf mistletoes trees are appropriate. Pruning, in of ponderosa pine by contacting a conjunction with removal of severely County Cooperative Extension agent, infected trees, can prolong the life of their local state forestry office, or individual trees. their regional USDA Forest Service, Forest Health Protection office. For high value trees, removal of live witches' brooms that exert a large References drain on infected trees can improve their vigor and prolong their life. Hawksworth, F.G. 1961. Trees may have to be repruned occa- Dwarfmistletoe of ponderosa pine in sionally to remove developing witch- the Southwest. USDA Forest Service es' brooms. Candidates for pruning Technical Bulletin No. 1246, 112 p. are trees with infections only in the lower half of their crown. Hawksworth, F.G. 1977. The 6-class In addition to prolonging tree dwarf mistletoe rating system. life, pruning can reduce the danger of USDA Forest Service Research Note trees or branches breaking and pre- RM-48, 7 p. vent damage and subsequent liability from falling branches. Trees with Hawksworth, F.G., and D. Wiens. bole infections may contain decayed 1996. Dwarf mistletoes: biology, wood and therefore may have a high- pathology, and systematics. USDA er potential to fail. Forest Service Agriculture Handbook 709, 410 p. Pesticides used improperly can be injurious to humans, animals, and plants. Follow direc- tions and read all precautions on the labels. Consult your local forest pathologist, county agricultural agent, or State extension agent about restrictions and registered uses of par- ticular pesticides. The United States Department of Agriculture (USDA) prohibits discrimination in its programs and activities on the basis of race, color, national origin, gender, religion, age, disability, political beliefs, sexual orientation, and marital or family status. (Not all prohibited bases apply to all programs.) Persons with disabilities who require alternative means of communication of program information (Braille, large print, audiotape, etc.) should contact USDA’s TARGET Center at (202) 720-2600 (voice and TDD). To file a complaint of discrimination, write USDA, Director, Office of Civil Rights, Room 326-W, Whitten Building, 14th and Independence Avenue, SW, Washington, DC 20250-9410 or call (202) 720- 5964 (voice or TDD). USDA is an equal opportunity provider and employer. R6-NR-FID-PR-01-03

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