Allotropa virgata - Candystick

Family: Ericaceae (Crowberry) ^{[E-flora](POWO,2026)}

"This unusual looking species is native to the western US (CA, ID, MT, NV, OR, WA) and Canada (BC). In British Columbia, it is primarily found in the southwestern corner of the province, where it has been collected most frequently on eastern Vancouver Island. There are reports from the area north of Kamloops, and the Sicamous area. It may be overlooked in some areas. Candystick is an erect perennial rhizomatous clonal species 10-50 cm tall with scale-like leaves and single, often multiple, unbranched stems. Stems arise from "a diffuse, deep-seated root system bearing branch roots and adventitious buds at intervals along its length" (Lichthardt and Mancuso 1991). Seeds are tiny (1 mm), with more than 100 per seed capsule, and are wind-dispersed. Plants persist after seed dispersal. Deer have been observed browsing this species, and may play a role in seed dispersal (Bureau of Land Management 2010)." ^[E-flora]

"Candystick is a member of the Monotropoideae--the subfamily of plants within the Ericaceae that lack chlorophyll. It is mycotrophic, and obtains its food from mycorrhizal fungi that are associated with nearby coniferous trees (sometimes hardwood trees (Lichthardt and Mancuso 1991)). In BC, it has been found in mesic to dry Douglas-fir forests in the lowland and montane zones at elevations of 30 to 900 m on nutrient poor sites. In the US, it is commonly found in lodgepole pine-beargrass communities on granitic soils but has also been found under ponderosa pine [Pinus ponderosa], subalpine fir [Abies lasiocarpa] and western red cedar [Thuja plicata] (Lichthardt and Mancuso 1991)." ^[E-flora]

"Although not documented, it is possible that this species may exhibit dormancy and flowering stems may not appear every year--like other members of the Monotropoideae which can remain dormant for up to thirteen years (Bureau of Land Management 2010)." ^[E-flora]

Habitat / Range : "Mesic to dry forests in the montane and lowland zones; infrequent in extreme SW BC; S to MT, ID and CA." ^{[IFBC-E-flora]} "The native range of this species is W. Canada to W. U.S.A. It is a holomycotrophic perennial and grows primarily in the temperate biome. " ^(POWO,2026)

Status: Native. ^[E-flora]

• "General:Perennial herb from a rhizome; parasites on mycorrhizal fungi associated with trees; "stems" erect, simple, 10-50 cm tall, 4-10 mm thick, glabrous, white with red or maroon stripes; persistent after seed dispersal; lacking chlorophyll." ^{[IFBC-E-flora]}
  
• "Leaves:Linear-lanceolate, scale-like, 20-35 mm long." ^{[IFBC-E-flora]}
  
• "Flowers:Inflorescence a spike-like, densely-flowered, terminal raceme, 5-20 cm long (about half the length of the stem); flower stalks erect; corollas cup-shaped, the petals 5, white or pinkish to brownish, about 5 mm long; calyces generally absent, rarely 2-4 sepals; stamens 10, exserted, purplish, anthers without awns." ^{[IFBC-E-flora]}
  
• "Fruits:Capsules, nearly globe-shaped, up to 5 mm wide." ^{[IFBC-E-flora]}
  
• "Notes:Petals are often incorrectly described as sepals. Stem and leaves can be considered part of the inflorescence.^{" [IFBC-E-flora]}
  

Similar Species:

Lichthardt and Mancuso (1991) provide the following write up on similar species: "Allotropa may be confused with other non-green members of the heath family which often share the same habitat. Pindrops (Pterospora andromedea) has sticky, brown flowering stems that also dry and persist a long time after dying. Unlike Allotropa, its flowers face downward, born at the ends of delicate, reflexed pedicels. It is generally much taller than Allotropa (3 dm to over a meter). Pinesap (Hypopitys monotropa) is similar in height to Allotropa, but is uniformly pinkish to straw-colored, drying to black. In flower the main stem bends over (nods) distinctly, but straightens again as fruits develop. Some non-green orchids, especially coralroot (Corallorhiza spp.), are sometimes mistaken for Allotropa. Coralroot has slender stems, rather variable in color from purplish to reddish-brown to albino. However, stems do not bear the small, pointed leaf blades always present on Allotropa." ^[E-flora]

Ecological Indicator Information

"A shade-intolerant, submontane to montane, Western North American parasite distributed more in the Pacific than the Cordilleran re­gion. Occurs in cool temperate and cool meso­thermal climates on very dry to moderately dry, nitrogen-poor soils (Mor humus forms). Its occurrence decreases with increasing latitude, precipitation, and continentality. Sporadic in open-canopy Douglas-fir forests on water-shedding sites. Commonly associated with Hylocomium splendens and Pleurozium schreberi. An oxylophytic species characteristic of moisture-deficient sites." - Indicator Plants of Coastal British Columbia (Information applies to coastal locations only)^[E-flora]

Biochemicals

Pigments

• Antheraxanthin 51.71 nanograms/gram fresh wt. ^{(Cummings,1998)}
• Lutein/zeaxanthin 46.81 nanograms/gram fresh wt. ^{(Cummings,1998)}
• Violaxanthin 4.54 nanograms/gram fresh wt. ^{(Cummings,1998)}

"The presence of carotenoid pigments in these heterotrophs is not surprising, because carotenoid pigments are known to have non-photosynthetic functions, such as photoprotection (GOODWlN 1980). Even within fully photosynthetic plants carotenoid pigments are distributed in many non-photosyntheic tissues (GOODWIN 1980)." ^{(Cummings,1998)}

Cultivation & Propagation

"Because Allotropa spreads by underground, perennating buds on lateral roots, 0.5 to 2 ft (15-61 cm) deep, it can survive a ground fire that does not kill the host tree." ^{(Massicotte et al.,2010)}

"Allotropa is a clonal species that spreads by extensive lateral roots, up to 4 ft (12 dm) long, bearing adventitious buds (Luoma 1987, Wallace 1975). The lateral roots occur as deep as 2 ft (61 cm). At the point of bud initiation a mass of fibrous roots is produced. It is these fibrous roots that form the mycorrhizal relationship. Each new individual propagated in this way is genetically identical to the one before and is therefore technically a ramet (Harper 1977) as opposed to an individual propagated by seed, or genet. Successive ramets may be as far as 1 m (3.2 ft) apart (Castellano, pers. comm.)." ^{(Massicotte et al.,2010)}

"The small seeds of Allotropa lack nutrient reserves and must therefore establish a mycorrhizal association immediately upon germinating. Although this sounds like a rare event, seeds are released in late summer and autumn, at which time some mycorrhizal fungi are particularly active and supplies of carbohydrates in litterfall and plant matter are at their peak (Leake 1994). Thus, seeds may not need to lie dormant for long. Seeds of Pterospora have been observed to retain viability for 9 weeks after maturation (Bakshi 1959, in Leake 1994). Because of their mycotrophic nature it is suspected that the period from seed germination to flowering may extend over many years (Leake 1994). All attempts at germinating Allotropa seeds in the lab, in the presence of a suitable fungus, have failed (Castellano, pers. comm.)." ^{(Massicotte et al.,2010)}

"Extensive, stand-replacing fires appear to have played an important role in the past and current distribution of Allotropa." ^{(Massicotte et al.,2010)}

Mycorrhizal

"This is the mycorrhizal association formed by the achlorophyllous plants that used to be placed in the Family Montropaceae, though they are currently placed in the Family Ericaceae; it’s convenient to talk about them collectively so we will call them monotropes. The group includes the genera Monotropa, Monotropsis, Allotropa, Hemitomes, Pityopus, Pleuricospora, Pterospora and Sarcodes.
All these monotropes are highly unusual plants in that they are all entirely lacking in chlorophyll, and so are unable to photosynthesise. Instead, they parasitise their mycorrhizas (they are called mycoheterotrophs), using the fungi not only to obtain minerals and nutrients from the soil, but also to tap into the carbohydrate supplies of nearby photosynthesising plants via their root connections to the mycorrhizal fungus." ^{[Moore 21stFungi]}

"Although often referred to as a saprophyte, Allotropa, like other non-green members of the Ericaceae, is actually mycotrophic–it obtains carbohydrates from a mycorrhizal fungus associated with its roots. The fungal mycelium is shared with a photosynthesizing plant that indirectly supplies carbohydrates to the mycotroph via the fungus. The mycotroph appears to be parasitic on the fungus, but there could exist a much more complex interaction in which all three partners benefit." ^{(Massicotte et al.,2010)}

"Although inland populations of Allotropa are strongly tied to lodgepole pine stands, it has been demonstrated that its roots can be colonized by fungal associates of various conifers as well as several hardwood species (Castellano and Trappe 1985). Species of fungi that have been identified from the roots of Allotropa include Rhizopogon vinicolor, Cenococcum geophilum (Castellano and Trappe 1985), and Tricholoma magnivelare (Castellano, pers. comm.). The mushroom producing fungus T. magnivelare commonly occurs with Allotropa in its coastal range. Tricholoma magnivelare has been observed at several Allotropa sites on the Beaverhead National Forest and (Carver 1993) and has been identified from Allotropa roots collected on the Nez Perce National Forest." ^{(Massicotte et al.,2010)}

Personal Note: (Bidartondo, Martin I. ,2005) & (Massicotte et al.,2010) reference Cenococcum geophilum as a mycorrhizal partner of A. virgata, citing (Castellano and Trappe 1985). The direct quote from the original source, as noted below, would seem to indicate otherwise.

Truffles: "Typical Rhizopogon vinicolor Smith mycorrhizae (Zak, 1971) were observed on roots of A. virgata, H. congestum, and P. fimbriolata. Mycorrhizae of Cenococcum geophilum Fr. (Mikola, 1948) were a minor component of P. fimbriolata, P. andro? medea, and H. congestum root systems." ^{(castellano1985)}

Matsutake Indicator Species: In China, particularly in the north-east, matsutake always occurs in pure stands of Pinus spp. mixed with a few Quercus magnolica and with bush and herb layers containing fewer than 20 species, including Rhododendron spp., Lespedeza bicolor Turez., Atractilis japonoca Koidz. ex Kitam. and Carex callitrichos V. Krecz., which are considered indicator plants for matsutake-producing areas. The indicator plant in Japan and Korea is Juniperus rigida Sieb. et Zucc. and in the USA Allotropa virgata T. et G. ex Gray.^(Yun&Hall)

"Many pickers I spoke to said that John Getz was “legendary” in the picking scene in general, and the matsutake scene in particular, a fact he credits to having noticed a plant that he said “seeks out” the matsutake mycelium, Allotropa virgata, known as the candy-stick. Since then it has been established that A. virgata does indeed have a parasitic relationship with matsutake mycelium that in turn has a mycorrhizal relationship with the pine tree." ^{[Myco Bone]}

• Tricholoma magnivelare ^{(Bidartondo, Bruns, 2001)}
• Tritholoma pardinum ^{[Heijden ME]}
• Tricholoma flavovirens ^{[Heijden ME]}
• Rhizopogon vinicolor ^{(castellano1985)}

"Lefevre et aL (1998) used molecular characterisation and morphology to show that Allotropa virgata associates exclusively with Tricholoma magnivelare. Kretzer et aL (2000) re-examined the associates of Sarcodes sanguinea from a much larger sample (76 plants versus 12) with a combination of ITS-RFLP and ITS sequence data obtained directly from roots; cultures were also iso- lated from the Sarcodes roots and used to synthesise mycorrhizae with pine. All sequences, RFLP patterns and cultures belonged to the Rhizopogon ellenae species complex. Additional sampling throughout most of the geographic range of Sarcodes sanguinea confirms this result (M. Bidartondo, pers. comm.). Interestingly, the R. ellenae complex is closely related to, but distinct from, the Rhizopogon species that associate with P. andromedea." ^{[Heijden ME]}

"We re-examined and expanded the sample size and geographical range of previous identifications to systematically evaluate mycorrhizal specificity in the Monotropoideae. To determine if different Monotropoideae lineages are specifically dependent on different lineages of fungi we identified the plant and fungal lineages involved in the monotropoid mycorrhizal symbiosis for 10 species over parts of their world-wide distribution. This sample represents all but two (Cheilotheca spp.) described species in the Monotropoideae....The 13 plant lineages and 31 fungal lineages involved were defined by nrITS sequence data. One-hundred and four populations, 331 plants, and 251 fungal basidiocarps were screened using molecular methods.... The Tricholomatoid group is poorly resolved, but the symbionts of Monotropa hypopithys, Pityopus californicus and Allotropa virgata shared shortest distances with its members (Fig. 1)." ^{(Bidartondo, Bruns, 2001)}

References

• (Bidartondo, Bruns, 2001) BIDARTONDO, MI, and TD BRUNS. "Extreme specificity in epiparasitic Monotropoideae (Ericaceae): widespread phylogenetic and geographical structure." Molecular Ecology 10 (2001): 2285-2295.
• (Bidartondo, Martin I. ,2005) Bidartondo, Martin I. "The evolutionary ecology of myco‐heterotrophy." New Phytologist 167.2 (2005): 335-352.
• (castellano1985) Castellano, Michael A., and James M. Trappe. "Mycorrhizal associations of five species of Monotropoideae in Oregon." Mycologia 77.3 (1985): 499-502.
• (Cummings,1998) Cummings, Michael P., and Nicholas A. Welschmeyer. "Pigment composition of putatively achlorophyllous angiosperms." Plant Systematics and Evolution 210.1 (1998): 105-111.
• [E-flora] Allotropa virgata [Accessed: 6/7/2014 7:25:41 AM ]
  • [IFBC] Illustrated Flora of B.C., 1998-2002 Douglas, G.W., G.B. Straley, D.V. Meidinger, and J. Pojar (Editors) Publication
• (Massicotte et al.,2010) Massicotte, Hugues B., et al. "Structural characteristics of root–fungus associations in two mycoheterotrophic species, Allotropa virgata and Pleuricospora fimbriolata (Monotropoideae), from southwest Oregon, USA." Mycorrhiza 20.6 (2010): 391-397.
• (POWO,2026) Allotropa virgata Torr. & A.Gray https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:582508-1, Accessed Feb 19, 2026
• Tricholoma Matsutake and Related Fungi, Wang Yun, Ian R. Hall, and Lynleya. Evans, Economic Botany 51(3) pp 311-327. 1997
• (Yun&Hall) Yun, Wang, Ian R. Hall, and Lynley A. Evans. "Ectomycorrhizal fungi with edible fruiting bodies 1. Tricholoma matsutake and related fungi." Economic botany 51.3 (1997): 311-327.

ALLOTROPA SUGAR STICK

1 sp. (Greek: different turned, from erect inflorescence) ^[Jepson]

Local Species;

1. Allotropa virgata - Candystick ^[E-flora]

"Copeland ( 1938 ) studied the morphology. Bidartondo and Bruns ( 2001, 2002 ) sequenced the ITS region of the fungal symbionts of a number of Allotropa plants from California and Oregon and found that all investigated specimens were associated with a narrow clade within the basidiomycete fungal genus Tricholoma." ^{[Merckx Myco]}

References

• [Jepson] Gary D. Wallace, 2012. Allotropa, in Jepson Flora Project (eds.) Jepson eFlora, http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?tid=12707, accessed on Mar 26 2014

Images

• 1 Mount Rainier National Park from Ashford, WA, United States, Public domain, via Wikimedia Commons
• 2 Gail, CC BY 4.0, via Wikimedia Commons

Journals of Interest

• Lefevre CK, Carter CM, Molina R (1998) Morphological and molecular evidence of specificity between Allotropa virgata and Tricholoma magnivelare. In: Second International Conference on Mycorrhiza, Poster Presentation, Program and Abstracts. Upp- sala, Sweden, p 107

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