Arceuthobium tsugense (hemlock dwarf mistletoe)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Air Temperature
- Rainfall Regime
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Plant Trade
- Impact Summary
- Risk and Impact Factors
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Arceuthobium tsugense (Rosendahl) G.N.Jones
Preferred Common Name
- hemlock dwarf mistletoe
Other Scientific Names
- Arceuthobium douglasii var. tsugense (Rosendahl) M.E.Jones
- Razoumofskya tsugensis Rosendahl
International Common Names
- English: dwarf mistletoe: hemlock; mountain hemlock dwarf mistletoe; western hemlock dwarf mistletoe
- ARETS (Arceuthobium tsugense)
Summary of InvasivenessTop of page
Their potential to establish in other areas is limited by the need for the living parasite to survive on the pathway and reproduce after entry. Nevertheless, the risk of economic impact is considerable if host species are available. The conifers at greatest risk would be species, known to be hosts, planted as exotics in other continents, but there is also a certain possibility of spread to related species, not known to be hosts.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Dicotyledonae
- Order: Santalales
- Family: Viscaceae
- Genus: Arceuthobium
- Species: Arceuthobium tsugense
Notes on Taxonomy and NomenclatureTop of page
A. tsugense is in subgenus Vaginata, section Campylopoda, series Campylopoda.
Subsp. mertensianae (mountain hemlock dwarf mistletoe) is distinguished on Tsuga mertensiana, in the western USA, while the nominate subspecies (western hemlock dwarf mistletoe) occurs in the same area on T. heterophylla through British Columbia, Canada up to the southern tip of Alaska, USA. The two subspecies can be differentiated by electrophoretic methods (Nickrent and Stell, 1990), but the so-called shore pine race does not appear distinct.
DescriptionTop of page
Plant TypeTop of page
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 30 Jun 2021
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|United States||Present, Localized|
Risk of IntroductionTop of page
The risk of accidental introduction is already well recognized and trade in conifer plants is correspondingly controlled in many countries. Exotic Arceuthobium species are also specifically listed as prohibited imports in the European Union, other European countries, Australia, New Zealand, Turkey, Tanzania and no doubt many others. North American countries similarly restrict import of conifers.
HabitatTop of page
Habitat ListTop of page
|Terrestrial||Managed||Managed forests, plantations and orchards||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Natural forests||Present, no further details||Harmful (pest or invasive)|
Hosts/Species AffectedTop of page
Host Plants and Other Plants AffectedTop of page
|Abies amabilis (Pacific silver fir)||Pinaceae||Other|
|Abies lasiocarpa (rocky mountain fir)||Pinaceae||Other|
|Abies procera (noble fir)||Pinaceae||Other|
|Pinus contorta (lodgepole pine)||Pinaceae||Other|
|Pinus lambertiana (big pine)||Pinaceae||Other|
|Tsuga heterophylla (western hemlock)||Pinaceae||Main|
|Tsuga mertensiana (Patton's hemlock)||Pinaceae||Main|
Growth StagesTop of page
List of Symptoms/SignsTop of page
|Stems / distortion|
|Stems / witches broom|
Biology and EcologyTop of page
Chromosome number 2n = 28 (Hawksworth and Wiens, 1996).
Physiology and Phenology
Arceuthobium spp. are obligate parasites, depending on an endophytic system within the host branch to draw water and nutrients from their hosts. Germination usually occurs in the spring following autumn dispersal and is favoured by light. The seedling shows negative phototropism, causing it to bore directly into the host shoot, even from below. Seedlings of most Arceuthobium spp. can only penetrate young branches less than 5 years old. Most Arceuthobium spp. have no phloem tissue. Transfer of nutrients, including sugars, may depend on close association of host and parasite parenchyma cells, and apoplastic movement via the walls of these cells. Graniferous tracheary elements could also be involved (see Hawksworth and Wiens (1996), for detailed discussion on this topic).
Photosynthesis is apparently important in supporting the seedling as it germinates and attaches, but for the next 2-7 years (usually 3-4) of its life, the parasite persists only as the endophyte inside the host tissue without any aerial shoot. Even after emergence of the aerial shoots, the established parasite has a relatively low photosynthetic capacity, usually much less than 50% of 'normal'.
Once emerged, the parasite shoots of Arceuthobium spp. produce fruits annually, for at least 2 years, and often for 5 years or more (Hawksworth and Wiens, 1996).
A. tsugense, like other Arceuthobium spp., is dioecious. Pollination appears to be predominantly due to insects (especially ants and flies) but may also occur by wind (Hawksworth and Wiens, 1996). Anthesis occurs between July and September in A. tsugense subsp. tsugense, which is about 1-2 weeks earlier than in subsp. mertensianae. However, the seed dispersal period for subsp. tsugense (late September to early November) is about 2-4 weeks later than for subsp. mertensianae. No true seed is formed, as there is no testa, but the embryo is embedded in chlorophyllous endosperm, surrounded by viscin. This will be referred to as a seed for convenience. The embryo is green, a few millimetres long, and has a meristematic radicular apex without a root cap. Dispersal of the seed is exceptional, involving a hydrostatic, explosive process which expels the seed for some distance from the parent plant. Most dispersal occurs as temperatures rise and humidity declines in the morning. The viscin ensures that it is retained by any host shoot that is hit, but if this is a needle, it may slide down with gravity to the base of the needle and germinate there. Although this is the main means of dispersal over a short range, long-distance dispersal also occurs as a result of seeds sticking to birds or mammals. However, any seeds that are ingested by animals are destroyed. Seeds of Arceuthobium spp. do not generally show dormancy and germination normally occurs in the first season after dispersal, though seeds may retain dormancy for 1-4 years when stored in ideal conditions.
The main environmental constraint on an Arceuthobium sp. is the presence of its host, which is in turn determined by multiple environmental requirements. The different North American species most obviously differ in the latitudinal limits of their range, from those that occur in Canada and northern US states, to those which are confined to Mexico, with all intermediates. Species also differ from those with an essentially coastal distribution to those with a continental distribution. The relevant factors further interact to determine an altitudinal range, reflecting the fact that conifers form a distinctive element of montane vegetation. Soil conditions have practically no importance.
Air TemperatureTop of page
|Parameter||Lower limit||Upper limit|
|Mean annual temperature (ºC)||1||15|
|Mean maximum temperature of hottest month (ºC)||18||32|
|Mean minimum temperature of coldest month (ºC)||-26||-2|
RainfallTop of page
|Parameter||Lower limit||Upper limit||Description|
|Dry season duration||2||12||number of consecutive months with <40 mm rainfall|
|Mean annual rainfall||200||1150||mm; lower/upper limits|
Rainfall RegimeTop of page
Natural enemiesTop of page
Notes on Natural EnemiesTop of page
Means of Movement and DispersalTop of page
Natural dispersal of Arceuthobium spp. is by the explosive fruits which can expel the seeds at speeds of 2.6 m/s up to a 15-m distance (Hinds and Hawksworth, 1965). In spite of this, the natural spread may not exceed about 1.5 m/annum (Hawksworth, 1958).
Vector Transmission (Biotic)
Seeds of Arceuthobium spp. falling onto the plumage of birds, or the fur of animals, tend to stick and may be dispersed for long distances. About 7% of birds and mammals trapped carried seeds, rising to 22% during the 2-week period of maximum seed release (Hawksworth and Johnson, 1989).
Logging and movement of timber which has not been completely de-barked, can result in movement of complete plants of Arceuthobium and possible transfer of seeds and establishment of new infestations.
Accidental introduction of Arceuthobium spp. into new areas or continents does not appear very likely. Seeds are short-lived, and unlikely to reach a host tree under circumstances in which they could develop. Conifer plants could carry living mistletoe plants, especially in the prolonged endophytic stage before the external plant develops, but young plants, as normally traded, are not very likely to be infected. Mistletoes could be carried on cut branches, including Christmas trees, and possibly on logs with bark (though mistletoes normally occur on the branches of trees, not on trunks). It is unlikely that mistletoes borne on cut, dead plants present any risk of transmission. Accordingly, introduction can be prevented relatively easily. The prohibition of import of plants for planting of the main host genera (as established, for example, in the phytosanitary regulations of the European Union) blocks the only really dangerous pathway.
Intentional introduction seems extremely unlikely, other than for research.
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Stems (above ground)/Shoots/Trunks/Branches||whole plants|
|Plant parts not known to carry the pest in trade/transport|
|Fruits (inc. pods)|
|Growing medium accompanying plants|
|True seeds (inc. grain)|
Impact SummaryTop of page
|Fisheries / aquaculture||None|
ImpactTop of page
A. tsugense is a severe pathogen of Tsuga mertensiana and T. heterophylla in western North America.
Risk and Impact FactorsTop of page
- Invasive in its native range
- Highly mobile locally
- Has high reproductive potential
- Negatively impacts agriculture
- Competition - monopolizing resources
- Difficult/costly to control
Similarities to Other Species/ConditionsTop of page
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.The following methods for controlling Arceuthobium spp. may have utility for the management of A. tsugense.
In the absence of any simple direct means of control of dwarf mistletoes, and the vast areas of forest involved, cultural management is virtually the only approach to the problem, the techniques varying according to the type of stand in which the problem occurs. Management options listed by Hawksworth and Johnson (1989) include:
- Use RMYLD model to predict yields (Edminster, 1978; Hawksworth, 1978)
- Favour or plant resistant tree species
- Prune infected branches and witches' brooms
- Destroy the whole stand (including the use of fire) and regenerate
- Fell non-merchantable infected trees
- Sanitation thin
- Harvest and regenerate the stand
- Do nothing.
Hawksworth and Johnson (1989) also refer to mechanisms to help prevent infection, including the use of natural or man-made barriers (roads, streams, strips of non-susceptible forest) to reduce (re)invasion from adjacent infested stands; and removing infected trees before re-planting/regeneration.
Detailed surveys are an essential ingredient of successful control programmes and the 6-class rating system (Hawksworth, 1977) is widely accepted as a standard. This involves a 0-6 score based on 0, 1 or 2 for each third (lower, middle, upper) of the tree; 0 for no infection, 1 for light infection (less than half the branches affected) or 2 for heavy infection (more than half infected).
In recently harvested, regenerating stands, the emphasis is on the complete removal of any infected trees over 2 m, regardless of commercial value, both within the stand, and along borders to a distance of 18 m, before the regeneration is 1 m high.
In pre-commercial stands in which surveys show less than 40% infected trees, it should be economic to practice selective thinning to remove all those infected. Above 40% this is unlikely to be economic. Severely infested stands may best be harvested early and regenerated, but decisions may require use of available models to help devise the most economic option. Some of the available models are described by Muir and Geils (2002).
Dwarf mistletoes may contribute in various ways to biodiversity - by creating openings in the forest following tree death, by providing nesting sites in the 'brooms' and by providing food for a range of vertebrates and invertebrates. There can therefore be some conflict between the requirements of forest exploitation, and environmental concerns.
Pruning may be appropriate as a means of reducing damage to individual trees, but more generally to reduce the source of infection for surrounding trees. The practicality, however, is that it will only be feasible in particular amenity and recreation areas.
Clear-felling (with or without fire) is appropriate where a stand is so severely infested that it needs to be abandoned and regenerated or re-planted.
The only chemical approved for use against dwarf mistletoes is the ethylene-releasing growth regulator, ethephon, which can cause abscission of the shoots and delay fresh seeding for 2-4 years, but there is eventual re-growth from the endophyte. It is difficult to achieve good coverage in larger trees from the ground, whereas applications from the air fail to penetrate the canopy adequately. The treatment is therefore of interest mainly for high-value amenity trees.
Glomerella cingulata is being developed as a biocontrol agent for use on A. americanum and A. tsugense (Geils et al., 2002) and has shown promise in field trials. Work is also in progress on two other pathogens, Caliciopsis arceuthobii and Nectria neomacrospora. There is no known method for biological control of A. tsugense.
Hawksworth and Johnson (1989) emphasise the importance of integrating dwarf mistletoe control with measures to reduce damage from the mountain pine beetle (Dendroctonus ponderosae).
ReferencesTop of page
Edminster CB, 1978. RMYLD: computation of yield tables for even-aged and two-storied stands. Research Paper RM-199, Fort Collins, USA: United States Department of Agriculture Forest Service.
EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm
Geils BW; Tovar JC; Moody B; (technical coordinators), 2002. Mistletoes of North American Conifers. General Technical Report RMRS-GTR-98. Ogden, USA: United States Department of Agriculture Forest Service.
Hawksworth FG, 1958. Rate of spread and intensification of dwarf mistletoe in young Lodgepole Pine stands. Journal of Forestry, 56:404-407.
Hawksworth FG, 1978. Intermediate cuttings in mistletoe-infested lodgepole pine and southwestern ponderosa pine stands. General Technical Report, Pacific Southwest Forest and Range Experiment Station, No. PSW-31:86-92
Hawksworth FG; Johnson DW, 1989. Biology and management of dwarf mistletoe in lodgepole pine in the Rocky Mountains. General Technical Report - Rocky Mountain Forest and Range Experiment Station, USDA Forest Service, No. RM-169:ii + 38 pp.
Hawksworth FG; Wiens D, 1996. Dwarf Mistletoes: Biology, Pathology, and Systematics. Agriculture Handbook 709. Washington DC, USA: United States Department of Agriculture Forest Service.
Hinds TE; Hawksworth FG, 1965. Seed dispersal velocity in four dwarf mistletoes. Science, 148:517-519.
Nickrent DL; Stell AL, 1990. Electrophoretic evidence for genetic differentiation in two host races of hemlock dwarf mistletoe (Arceuthobium tsugense). Biochemical Systematics and Ecology, 18(4):267-280
Rediske JH; Shea KR, 1961. The production and translocation of photosynthate in dwarf mistletoe and Lodgepole Pine. American Journal of Botany, 48:447-452.
CABI, Undated. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI
Hawksworth FG, Wiens D, 1996. Dwarf Mistletoes: Biology, Pathology, and Systematics. In: Agriculture Handbook 709, Washington DC, USA: United States Department of Agriculture Forest Service.
Mathiasen R, Daugherty C, 2009. First report of mountain hemlock dwarf mistletoe (Arceuthobium tsugense subsp. mertensianae) on sugar pine (Pinus lambertiana) from Oregon. Plant Disease. 93 (3), 321. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-93-3-0321A
Distribution MapsTop of page
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