Arceuthobium campylopodum (western dwarf misletoe)
- 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
- 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 campylopodum Engelmann
Preferred Common Name
- western dwarf misletoe
Other Scientific Names
- Razoumofskya campylopoda (Engelmann) Kuntze
- ARECA (Arceuthobium campylodum)
- ARECP (Arceuthobium campylopodum)
Summary of InvasivenessTop of page Arceuthobium spp. do not spread rapidly and cannot be considered highly invasive. They do, however, constitute a serious threat as a result of their ability to build up gradually over the life of a forest and cause severe damaging effects on a number of important forest species.
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 campylopodum
Notes on Taxonomy and NomenclatureTop of page A detailed discussion of the taxonomy and taxonomic history of the genus Arceuthobium is provided by Hawksworth and Wiens (1996). The genus Arceuthobium is a member of the plant family Viscaceae and is a clearly defined group of small (generally less than 20 cm high), variously coloured flowering plants that are aerial parasites on conifers of the families Pinaceae and Cupressaceae. They are considered to be the most evolutionarily specialized genus of the family Viscaceae. Arceuthobium was previously included in the subfamily Viscoideae of the family Loranthaceae, but the subfamilies Loranthoideae and Viscoideae are now generally considered to have family status (Loranthaceae and Viscaceae).
A. campylopodum is a member of subgenus Vaginata, section Campylopoda, series Campylopoda.
This species was considered by some early authors to include A. occidentale, but the two species are now recognized to be distinct in morphology, geographical distribution and hosts; the latter having a more southerly range in California and mainly attacking Pinus sabiniana, P. radiata, P. muricata and P. contorta.
DescriptionTop of page A. campylopodum, like other Arceuthobium spp., is an obligate parasite with an endophytic 'root' system ramifying within the host branch. This endophyte expands within the cortex and becomes embedded in the xylem for some years before aerial shoots are produced, encircling the infected branch and growing along it. A. campylopodum plants are 8 to 13 cm tall, olive green to yellow in pistillate (female) plants and brownish in staminate (male) plants, loosely branched; internodes 6 times as long as wide; basal diameter of shoots more than 3 mm; staminate flowers 3.0 mm across, perianth 3-merous (occasioinally 4-merous); mature fruits 5.0 x 3.0 mm.
Plant TypeTop of page Parasitic
DistributionTop of page Mainly limited to forested areas in the western seaboard states of the USA, with a wide altitudinal range. A USDA website on North American plants (USDA, 2004) gives a record in Montana, but it is possible that this refers back to a 1935 report attributed to A. laricis by Hawksworth and Wiens (1996). Outside the USA, A. campylopodum extends just into the north of Baja California State (Mexico). Connell (1967) gives a record in the south of British Columbia (Canada), communicated by the Canadian NPPO. However, authors from the USA do not consider that A. campylopodum occurs in Canada.
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.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Canada||Restricted distribution||EPPO, 2014|
|-British Columbia||Present||Native||Invasive||EPPO, 2014|
|Mexico||Restricted distribution||Native||Invasive||Hawksworth and Wiens, 1996; EPPO, 2014|
|USA||Restricted distribution||EPPO, 2014|
|-California||Present||Native||Invasive||Hawksworth and Wiens, 1996; EPPO, 2014|
|-Idaho||Present||Native||Invasive||Hawksworth and Wiens, 1996; EPPO, 2014|
|-Montana||Indigenous, localized||Native||Invasive||USDA, 2004|
|-Nevada||Indigenous, localized||Native||Invasive||Hawksworth and Wiens, 1996|
|-Oregon||Present||Native||Not invasive||Hawksworth and Wiens, 1996; EPPO, 2014|
|-Washington||Present||Native||Not invasive||Hawksworth and Wiens, 1996; EPPO, 2014|
Risk of IntroductionTop of page The risk presented by Arceuthobium spp. introductions into other areas of the world is related to the availability of their hosts. The most obvious risk arises from the fact that several North American hosts (for example, Pinus contorta, P. ponderosa, Tsuga spp., and Pseudotsuga menziesii) have been more or less widely planted in other continents, in the absence of these mistletoes (curiously, P. radiata, one of the North American pines most widely planted around the world, is hardly reported as an Arceuthobium host, nor is Picea sitchensis, planted in parts of Europe). Conversely, the European or Asian hosts of Arceuthobium have not been substantially planted outside their natural range. A secondary risk is that, although in their natural range Arceuthobium spp. occur rather rarely on species other than their main hosts, there is limited data suggesting that they may readily infect some exotic species. There is accordingly a certain risk that Arceuthobium spp. may spread to and affect such exotic hosts if they are introduced into other continents, e.g. P. sylvestris in Europe, Juniperus virginiana in North America.
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 A. campylopodum is limited to forests in which its specific hosts are present in substantial numbers.
Habitat ListTop of page
|Managed forests, plantations and orchards||Present, no further details||Harmful (pest or invasive)|
|Natural forests||Present, no further details||Harmful (pest or invasive)|
Hosts/Species AffectedTop of page Under natural conditions A. campylopodum is restricted to certain Pinus spp. In addition to the main recorded hosts, it has very occasionally been found on others (e.g. P. lambertiana). It has been found on the exotics P. pinaster and P. sylvestris in arboreta. It has been artificially inoculated to various species, including Abies and Larix spp. (but has never been found occurring naturally on these genera). Recently, A. campylopodum has been found infecting planted Picea pungens (from the Rocky Mountains, USA) and Picea abies (from Europe) in California, USA (Mathiasen et al., 1998).
Host Plants and Other Plants AffectedTop of page
|Picea abies (common spruce)||Pinaceae||Other|
|Picea pungens (blue spruce)||Pinaceae||Other|
|Pinus attenuata (knobcode pine)||Pinaceae||Other|
|Pinus contorta var. latifolia (Lodgepole pine)||Pinaceae||Other|
|Pinus contorta var. murrayana (sierra lodgepole pine)||Pinaceae||Other|
|Pinus coulteri (big-cone pine)||Pinaceae||Other|
|Pinus jeffreyi (Jeffrey pine)||Pinaceae||Main|
|Pinus ponderosa (ponderosa pine)||Pinaceae||Main|
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
In common with other Arceuthobium spp., A. campylopodum is an obligate parasite, 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 produce fruits annually, for at least 2 years, and often for 5 years or more (Hawksworth and Wiens, 1996).
A. campylopodum, 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). Meiosis occurs in July, followed by peak anthesis from mid-August to early October with extremes from early August to late October (Hawksworth and Wiens, 1996). Fruits usually mature from early September to mid-November, with extremes of late August to late November, and have a maturation period of about 13 months. 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.. 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.
Grasshoppers (Melanoplus devastator) have been reported (Scharpf and Koerber, 1986) to feed on the aerial portions of A. campylopodum growing on Pinus jeffreyi.
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||0||12||number of consecutive months with <40 mm rainfall|
|Mean annual rainfall||200||1150||mm; lower/upper limits|
Rainfall RegimeTop of page Uniform
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
Notes on Natural EnemiesTop of page Major fungal parasites of Arceuthobium spp. in North America that occur on A. campylopodum are Caliciopsis arceuthobii and Glomerella cingulata (Hawksworth and Wiens, 1996).
Means of Movement and DispersalTop of page Natural Dispersal (Non-Biotic)
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 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 of Arceuthobium spp., this 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 could occur on or in timber that is not de-barked.
Intentional introduction would seem extremely unlikely, other than for research.
Plant TradeTop of page
|Plant parts not known to carry the pest in trade/transport|
|Fruits (inc. pods)|
|Growing medium accompanying plants|
|Stems (above ground)/Shoots/Trunks/Branches|
|True seeds (inc. grain)|
Impact SummaryTop of page
|Fisheries / aquaculture||None|
ImpactTop of page Arceuthobium species as a whole are regarded as some of the most serious of all pests/diseases of North American forests. Dwarf mistletoes are much more damaging to their hosts than the 'green' mistletoes in both Loranthaceae and Viscaceae. Having little photosynthetic capacity, they draw more heavily on host carbohydrate, and furthermore interfere with photosynthate translocation to the roots. The mistletoe has a girdling effect, resulting in an accumulation of photosynthate above the site of infection. Apparently carbohydrates are withheld from the roots in quantities sufficient to cause the characteristic decline of the tree (Rediske and Shea, 1961; Hawksworth and Wiens, 1996). There are also severe growth-regulatory effects resulting from cytokinin production at the point of infection and the redirection of host photosynthate into the resulting witches broom growths. These distort and suppress growth of branches and even the main trunk. Wood quality is further affected as a result of swellings, witches' brooms and knots, and structural weakening associated with shortened, distorted tracheids.
A. campylopodum is a serious pathogen on Pinus jeffreyi and P. ponderosa in western North America, apparently more damaging in the southern parts of its range.
Risk and Impact FactorsTop of page Invasiveness
- 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 Arceuthobium occidentale (q.v.) is similar to A. campylopodum but has basal diameter of shoots less than 3 mm, and mature fruits 4 mm long. Its range and hosts are also distinct.
Prevention and ControlTop of page Cultural Control
Resistance to A. campylopodum is known in Pinus ponderosa and P. jeffreyi (Scharpf, 1984).
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. General management options for dwarf mistletoes 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. Preliminary tests of ethephon for use against A. campylopodum on P. jeffreyii in California has given promising results, yielding shoot abscission rates of 90-100% when coverage is thorough. However, rapid resprouting from the endophytic system in A. campylopodum may limit its effectiveness (Parks and Hoffman, 1991). It is difficult to achieve good spray coverage of ethephon 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.
Colletotrichum gloeosporioides [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.
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.
Parks CA; Hoffman JT, 1991. Control of western dwarf mistletoe with the plant-growth regulator ethephon. Research Note PNW-RN-506. Portland, Oregon, USA: Department of Agriculture, Forest Service, Pacific Nortwest Forest and Range Management Station.
Rediske JH; Shea KR, 1961. The production and translocation of photosynthate in dwarf mistletoe and Lodgepole Pine. American Journal of Botany, 48:447-452.
United States Department of Agriculture, 2004. Plants Database. http://plants.usda.gov/index.html.
Distribution MapsTop of page
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