The
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''Arceuthobium'', commonly called dwarf
mistletoe
Mistletoe is the common name for obligate parasite, obligate parasitic plant, hemiparasitic plants in the Order (biology), order Santalales. They are attached to their host tree or shrub by a structure called the haustorium, through which they ...
s, is a genus of 42
species of
parasitic plant
A parasitic plant is a plant that derives some or all of its nutritional requirements from another living plant. They make up about 1% of angiosperms and are found in almost every biome. All Parasite, parasitic plants develop a specialized organ ...
s that parasitize members of
Pinaceae
The Pinaceae (), or pine family, are conifer trees or shrubs, including many of the well-known conifers of commercial importance such as Cedrus, cedars, firs, Tsuga, hemlocks, Pinyon_pine, piñons,
larches, pines and spruces. The family is incl ...
and
Cupressaceae in
North America
North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
,
Central America
Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
,
Asia
Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
,
Europe
Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
, and
Africa
Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
.
Of the 42 species that have been recognized, 39 and 21 of these are endemic to North America and the United States, respectively. They all have very reduced shoots and
leaves
A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
(mostly reduced to scales) with the bulk of the plant living under the host's bark. Recently the number of species within the genus has been reduced to 26 as a result of more detailed genetic analysis.
Description

They are
dioecious
Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
, individual plants being either male or female. The
fruit
In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering.
Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
is unusual in that it builds up hydrostatic pressure internally when ripe and shoots the single sticky
seed
In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
up to speeds nearly , an example of
rapid plant movement. The
lodgepole pine dwarf mistletoe,'' Arceuthobium americanum, has been found to explosively-disperse its seeds through
thermogenesis.
Rolena A.J. deBruyn, Mark Paetkau, Kelly A. Ross, David V. Godfrey & Cynthia Ross Friedman. Thermogenesis-triggered seed dispersal in dwarf mistletoe. Dwarf mistletoe seeds are enveloped in a
hygroscopic
Hygroscopy is the phenomenon of attracting and holding water molecules via either absorption (chemistry), absorption or adsorption from the surrounding Natural environment, environment, which is usually at normal or room temperature. If water mol ...
, glue-like substance called
viscin. Many fail to land on a suitable host's shoot, but some succeed, and in this way they are spread through the forests as a pest front. The spread of dwarf mistletoes in forest stands is greatest from the overstory to the understory, owing to gravity. Advantageous stand conditions for the spread of the parasite include an uneven-aged stand structure with severely infected hosts in dominant and codominant crown classes,
species composition dominated by the primary host, and tree densities between 175 and 500 trees/ha.
There are also several species from
Europe
Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
and
Asia
Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
including one of the smallest in the genus, ''A. minutissimum'' that lives on its host, ''
Pinus wallichiana'' in the
Himalayas
The Himalayas, or Himalaya ( ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the Earth's highest peaks, including the highest, Mount Everest. More than list of h ...
.
Effects of parasitism
In western forest ecosystems of
North America
North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, numerous dwarf mistletoe species are considered to be serious forest-borne disease agents. Severe dwarf mistletoe infection can result in a reduction in tree growth, premature tree mortality, reduced seed and cone development, and reduced wood quality, and increases the susceptibility of the host tree to pathogen and/or insect attack. Most of the commercially important conifers in western North America are parasitized by one or more dwarf mistletoes.
The interaction between dwarf mistletoes and their host can be generalized as a source-to-sink relationship. Dwarf mistletoes derive the majority of their nutrition from the host’s
vascular tissue
Vascular tissue is a complex transporting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. T ...
s. Dwarf mistletoes have a root-like
endophytic system, composed of primary and secondary
haustoria, which invade, but do not injure, both the
xylem
Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
and
phloem
Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
of the host. Because this root-like endophytic system is not
soil
Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...
borne, dwarf mistletoes are dependent solely on their host for water. Along a xylem-to-xylem link, dwarf mistletoes draw water from their host using the difference in water potential between parasite and host. The greater transpiration rate of the dwarf mistletoe produces a lower water potential, allowing water to flow from host to parasite. The water gradient or
transpiration
Transpiration is the process of water movement through a plant and its evaporation from aerial parts, such as leaves, stems and flowers. It is a passive process that requires no energy expense by the plant. Transpiration also cools plants, c ...
stream is consistently maintained, even when the host is under moderate water deficit.
In addition to host-water dependence, dwarf mistletoes must acquire
carbohydrate
A carbohydrate () is a biomolecule composed of carbon (C), hydrogen (H), and oxygen (O) atoms. The typical hydrogen-to-oxygen atomic ratio is 2:1, analogous to that of water, and is represented by the empirical formula (where ''m'' and ''n'' ...
and mineral nutrition from their hosts. Dwarf mistletoes have both
chlorophyll
Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
a and chlorophyll b and the necessary mechanisms for photosynthesis, but chlorophyll concentrations in dwarf mistletoes are approximately 1/5 to 1/10 of those found in their host’s foliage, and dwarf mistletoes have low photosynthetic rates as measured by the rate of
carbon fixation
Biological carbon fixation, or сarbon assimilation, is the Biological process, process by which living organisms convert Total inorganic carbon, inorganic carbon (particularly carbon dioxide, ) to Organic compound, organic compounds. These o ...
. The principal carbohydrate transported from the host to dwarf mistletoe is
sucrose
Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula .
For human consumption, sucrose is extracted and refined ...
. Because dwarf mistletoes are phloem-deficient, they draw carbohydrates from their hosts by connections to the host phloem and ray
parenchyma
upright=1.6, Lung parenchyma showing damage due to large subpleural bullae.
Parenchyma () is the bulk of functional substance in an animal organ such as the brain or lungs, or a structure such as a tumour. In zoology, it is the tissue that ...
. The rate of carbohydrate transport varies by season, but dwarf mistletoes continuously draw carbohydrates from their hosts throughout the year.
Dwarf mistletoe rating system
A standardized system called the Hawksworth 6-class dwarf mistletoe rating (DMR) system has been devised to determine how much dwarf mistletoe has infected a tree. To use this system, the living part of the
tree crown needs to be broken up into 3 sections, (top, middle, bottom). Each section is then rated either 0, 1, or 2 with a score of 0 being assigned for a uninfected section, 1 for light infection with fewer than half of the branches infected, or 2 for heavy infection with over half of the branches infected. The 3 numbers are then added together to give the total rating for the tree.
Species

* ''
Arceuthobium abietinum''
Engelm. ex Munz
* ''
Arceuthobium americanum''
Nutt. ex Engelm.
* ''
Arceuthobium apachecum''
Hawksworth & Wiens
* ''
Arceuthobium azoricum''
Hawksw. & Wiens
* ''
Arceuthobium blumeri''
A. Nels.
* ''
Arceuthobium californicum''
Hawksworth & Wiens
* ''
Arceuthobium campylopodum''
Engelm.
* ''
Arceuthobium cyanocarpum''
(A. Nels. ex Rydb.) A. Nels.
* ''
Arceuthobium divaricatum''
Engelm.
* ''
Arceuthobium douglasii''
Engelm.
* ''
Arceuthobium gillii''
Hawksworth & Wiens
* ''
Arceuthobium globosum''
Hawksw. & Wiens
* ''
Arceuthobium laricis''
(Piper) St. John
* ''
Arceuthobium littorum''
Hawksworth, Wiens & Nickrent
* ''
Arceuthobium microcarpum''
(Engelm.) Hawksworth & Wiens
* ''
Arceuthobium minutissimum''
Hook. f.
* ''
Arceuthobium monticola''
Hawksworth, Wiens & Nickrent
* ''
Arceuthobium occidentale''
Engelm.
* ''
Arceuthobium oxycedri''
(DC.) M.Bieb.
* ''
Arceuthobium pusillum''
Peck
* ''
Arceuthobium siskiyouense''
Hawksworth, Wiens & Nickrent
* ''
Arceuthobium tsugense''
(Rosendahl) G.N. Jones
* ''
Arceuthobium vaginatum''
(Humb. & Bonpl. ex Willd.) J. Presl (Syn. ''Viscum vaginatum''
Humb. & Bonpl. ex Willd.)
References
Further reading
* Hawksworth, F. G., & Wiens, D. (1996). ''Dwarf Mistletoes: Biology, Pathology, and Systematics''. USDA Forest Service, Agriculture Handbook 709.
* Kenaley, S.C., R.L. Mathiasen, & C.M. Daugherty. 2006. Selection of dwarf mistletoe-infected ponderosa pines by ''Ips'' species (Coleoptera: Scolytidae) in northern Arizona. WNAN 66:279-284.
*Mathiasen, R.L. 1996. Dwarf mistletoes in forest canopies. Northwest. Sci. 70:61-71.
External links
Maleshoots of ''A. campylopodum'' in CaliforniaFruiting plant of ''A. campylopodum'' on Pine shootPLANTS profile for Arceuthobium (United States) including range maps.US forest service data about dwarf mistletoes
{{Taxonbar, from=Q244255
Dioecious plants
Epiphytes
Parasitic plants
Santalaceae genera