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Proceedings of the Symposium on Dwarf Misletoe Control Through Forest Management

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Proceedings of the Symposium on Dwarf Misletoe Control Through Forest Management Bases for Control BIOLOGICAL FACTORS OF DWARF MISTLETOE IN RELATION TO CONTROL-I/ 2 Frank G. Hawkswortb/ Abstract: Dwarf mistletoes have been present in North American coniferous forests for essentially as long as their hosts. Most of the world's dwarf mistletoes occur in North America where they attack all six genera of the Pinaceae: Pinus, Picea, Abie?, Larix, T:uga, and Pseudotsuga. Dwarf mistletoes have many effects including: growth loss, mortality, reduction in seed crops, lowered wood quality, predisposition of infected trees to insects and fungi, and ecologic effects. Features that make the dwarf mistletoes amenable to silvicul- tural control measures are: they are obligate parasites so will live only as long as the host does; they are generally host specific; they have long life cycles, frequently 4 to 6 years; the rate of spread through stands is slow, generally on the order of 1 to 2 feet per year; and they are readily detected as they are non-microscopic organisms and cause distinctive symptoms. INTRODUCTION HISTORY Dick Parmeter and I have been asked to The dwarf mistletoes are no newcomers to help "set the stage" for this symposium by North America. We know from fossil evidence outlining some basics of dwarf mistletoe biol- that these parasites have been here at least ogy, stand dynamics, and ecological factors since the Miocene period, or about 25 million relating to control. I'll discuss the follow- years. They presumably migrated across the ing items relating to biology and let Dick Bering area from eastern Asia, along with their tackle the difficult topics of stand dynamics coniferous hosts. Thus, the dwarf mistletoes and ecological factors. and Western conifers have co-evolved over the millenia. We must keep this in mind when we History tend to regard mistletoes and other "pests" as The genus Arceuthobium something foreign to, and not an integral part Host relationships of, forest ecosystems (Hawksworth 1975). Dwarf mistletoe effects Dwarf mistletoe life cycles The earliest written record of dwarf Bases for control mistletoe that I have found is from southern Europe (Clusius 1601). The first North American collection was by the famous explorer and natur- Ñ'presente at Symposium on Dwarf Mistletoe alist Alexander von Humbolt, in Veracruz, Mexico, Control through Forest Management, Berkeley, in 1804. David Douglas, the botanical explorer, was apoarently the first to discover dwarf Calif. April 11-13, 1978. mistletoes in what is now the United States ;Supervisory Forest Pathologist, Rocky Mountain (Douglas 1914). In 1826 he found two dwarf Forest and Range Experiment Station, Forest mistletoes in eastern Washington, which we can Service, U.S. Department of Agriculture, Fort now determine were Arceuthobium americanum on lodgepole pine, and campylopodum on ponderosa Collins, Colo. 4. pine (although he thought the latter was red The aerial parts of the dwarf mistletoe (the pine). shoots) are essentially reproductive structures. The shoots range from as small as 1/4 inch high Several early western botanists in the in Arceuthobium minutissimum of the Himalayas 1850's collected dwarf mistletoes which were to as high as nearly 3 feet for A. globosum sent to Dr. George Engelmann of the Missouri in Guatemala, but most species have shoots trom Botanical Garden, who described most of the 3 to 5 inches high. Dwarf mistletoe color species in the West. Dr. James R. Wier, ranges from black to red, purple, green, gray, pioneer forest pathologist in the Inland Empire brown, to yellow (Hawksworth and Wiens 1973). from 1910 to 1923, was the first to point out The root system consists of two main parts: the seriousness of the dwarf mistletoe problem (1) the cortical strands which occur primarily in the coniferous forests of the West. Recog- longitudinally in the inner bark, and (2) the nition of them as a serious problem in the sinkers, which extend from the cortical West, however, was slow to develop. It took strands radially into ray tissues of the host several early western Forest Pathologists-- xylem. The sinkers, which become imbedded in notably Lake S. Gill (who prepared a dwarf the xylem as the host tissues grow around them, mistletoe monograph (Gill 1935)), Toby Childs, provide the mechanism for water conduction from Willis W. Wagener, and J. S. Boyce--to bring host to parasite tissues (Kuijt 1960). about this awareness. These missionary path- ologists helped show that not only were these The dwarf mistletoes contain chlorophyll parasites causing severe losses in the West, so they can manufacture some of their carbo- but more importantly, that the dwarf mistletoes hydrates, but they obtain most of their carbo- could be controlled by silvicultural means. hydrates and all of their water and minerals They, in fact, helped build the strong founda- directly from the host. In contrast to most tion on which this symposium was constructed. mistletoes, carbohydrates manufactured by dwarf mistletoes are not translocated to the host (Leonard and Hull 1965). THE GENUS ARCEUTHOBIUM Arceuthobium, the only genus that occurs The dwarf mistletoes are unique as the in the eastern and western hemispheres, com- only group of parasitic higher plants confined prises some 31 known species. A few of these to conifers. They occur on conifer branches have recognizable subspecies on host forms or stems, and their endophytic (or root) so the grand total is now 38 dwarf mistletoes system is embedded in bark and wood tissues (table 1). This is six more than we knew of of the host (fig. 1). when we published our monograph 6 years ago (Hawksworth and Wiens 1972). North America has the "lion's share" of the world's dwarf mistletoes~32of the known 38. Most of these occur in Mexico and in the Western United States. The taxonomic status of the dwarf mistletoes, as indeed many of their host trees, is not yet "finalized." Several unsettled questions remain, e.g.; (1) the taxonomic status and host relationship of A. campylopodum and A. occidentale in California; (2) the status of dwarf mistletoes on white pines in northern California and southern Oregon, and (3) the status of "races" of the hemlock dwarf mistletoe--&. tsugense. It now seems fairly sure that A. tsugense has at least two "races" (Smith and Wass 1976) --one on shore pine in coastal British Columbia, Vancouver Island and Orcas Island, Washington, and one on western hemlock from Alaska to California. Possibly, the populations on mountain hemlock also are distinct. Figure I--Dwarf mistletoe in a pine stem-- showing shoots, cortical strands and sinkers. Table I--Synopsis of the genus Arceuthobium, with distribution and principal hosts of each taxon. Dwarf Mistletoe Distribution Principal hosts NEW WORLD TAXA 1. Arceuthobium abietinum a. f . sp. concoloris Western U.S. Abies concolor b. f. sp. magnificae Calif. , S. Oreg. --Abies magnif ica Arceuthobium abietis-religiosae Mexico --Abies spp. Arceuthobium americanum West. U.S., West. Can. Pinus contorta, x. banksiana Arceuthobium apachecum sw U.S. Pinus strobiformis Arceuthobium aureum a. Subspecies aureum Guatemala, Belice Pinus spp. b. Subspecies petersonii S. Mexico Pinus spp . Arceuthobium bicarinatum Hispaniola Pinus occidentalis Arceuthobium blumeri Ariz., Mexico Pinus strobiformis Arceuthobium californicum Calif., Oreg. Pinus lambertiana Arceuthobium campylopodum W. U.S., Mexico Pinus ponderosa, P. jeffreyi Arceuthobium cyanocarpum w. U.S. Pinus f lexilis Arceuthobium divaricatum W. U.S., Mexico Pinyons Arceuthobium douglasii B.C., W. U.S., Mexico Pseudotsuga menziesii Arceuthobium gillii a. Subspecies gillii SW U.S., Mexico Pinus leiophylla b. Subspecies nigrum Mexico Pinus spp. Arceuthobium globosun a. Subspecies globosum Mexico Pinus spp. b. Subspecies grandicaule Mexico, Guatemala Pinus spp. Arceuthobium guatemalense Mexico, Guatemala Pinus ayacahuite Arceu thobium hondurense Honduras --Pinus oocarpa Arceuthobium laricis B.C., NW U.S. Larix occidentalis Arceuthobium microcarpum Ariz., N. Mex. Picea engelmannii, x. pungens Arceuthobium occidentale Calif. PinuS-~sabiniana,g. radiata Arceuthobium pusillum E. Can., E. U.S. Picea mariana, P. glauca Arceuthobium rubrum Mexico P inus spp . Arceuthobium strictum Mexico Pinus spp. 23. Arceuthobium tsugense a. Western hemlock race Alaska to Calif. Tsuga heterophylla b. Shore pine race B.C., NW Wash. Pinus contorta 24. Arceuthobium vaginatum a. Subspecies vaginatum Mexico Pinus spp. b. Subspecies Colo. , Utah to Mexico Pinus ponderosa c. Subspecies durangense Mexico Pinus spp. 25. Arceuthobium verticillif lorum Mexico Pinus- spp. OLD WORLD TAXA 26. Arceuthobium azoricum Azores Juniperus brevifolia 27. Arceuthobium chinense SW China Abies sp., Keteleeria sp. 28. Arceuthobium juniperi-procerae Central Africa Juniperus procera 29. Arceuthobium minutissimum India, Pakistan Pinus wallichiana 30. Arceuthobium oxycedri Spain to India Juniperus spp. 31. Arceuthobium & SW China Pinus tabulaeformis HOST RELATIONSHIPS I emphasize that we're talking about dwarf mistletoe "effects", which may or may The dwarf mistletoes parasitize all six not be causing an "impact" on the forest genera in the Pinaceae in North America: resource. This, too, will be discussed in Pinus, Picea, Abies, Larix, Pseudotsuga and more detail by the next speaker. Tsuga. Most dwarf mistletoes parasitize pines as their principal hosts: Principal Hosts Number of Dwarf Mistletoes Hard pines Growth Loss White pines True firs The most common result of dwarf mistletoe Spruce infection is reduction of growth rates of in- Hemlock fected trees.
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