Pterospora

Pterospora

Secure  (NatureServe)[1]
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Ericales
Family: Ericaceae
Subfamily: Monotropoideae
Tribe: Pterosporeae
Genus: Pterospora
Nutt. 1818
Species:
P. andromedea
Binomial name
Pterospora andromedea
Nutt. 1818

Pterospora, commonly known as pinedrops, woodland pinedrops, Albany beechdrops, or giant bird's nest, is a North American genus in the subfamily Monotropoideae of the heath family, and includes only the species Pterospora andromedea. It grows as a mycoheterotroph (relying on fungi rather than photosynthesis for nutrients) in coniferous or mixed forests. It is widespread across much of Canada as well as the western and northeastern United States to Mexico. Along with Monotropa it is one of the more frequently encountered genera of the Monotropoideae.[2][3]

Description

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The visible portion of Pterospora andromedea is a fleshy, unbranched, reddish to yellowish flower spike (raceme) 15–170 cm (6–67 in) in height,[4] though it has been reported to occasionally attain a height of 2 meters (6.6 feet).[5] The above-ground stalks (inflorescences) resemble an emerging asparagus spear in shape and are usually found in small clusters between June and August.[6][7] The inflorescences are hairy and noticeably sticky to the touch. This is caused by the presence of hairs which exude a sticky substance (glandular hairs). The inflorescences are covered by leaves that have been reduced to scale-like structures known as bracts that are 2–3.5 centimeters long and 0.5–1 centimeter wide.[4] The lower portion of the flowering stem is tightly covered in bracts and they become more widely spaced higher on the stalk.[8]

Pterospora flowers

The upper portion of the inflorescence has a series of white, yellowish, or rusty urn-shaped flowers that face downward.[9] Each flower may have five sepals surrounding the base of the flower,[6] which measure 4–6 millimeters long and are 2—3 millimeters wide.[4] The fused petals (the corolla) measure 6–9 millimeters in length.[6]

The fruit is a capsule with five lobes and 7–10 millimeters wide. Inside the capsules there are large numbers of very small seeds with, thin papery wings that are much wider than the seeds themselves.[6] Plants exist for most of their life as a mass of branching roots that form round root balls. These root balls may be penetrated by the roots of other plants, but there are no vascular connections between them.[10] At most, the total volume of a root ball is no more than 150–200 cubic centimeters. The inflorescence will grow from buds on roots which grow horizontally.[8]

Once fruiting is complete the flowering stems dry out and persist in the forest for a significant length of time.[11]

The main plants that may be confused with Pterospora andromedea are the mycoheterotrophic orchids in genus Corallorhiza, commonly called the coral roots. However, their flowers are bilaterally symmetrical, not regular like those of pinedrops.[11]

Fungal interactions

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P. andromedea, like all members of Monotropoideae, is a mycoheterotroph. This is a form of carbon acquisition that is parasitic on fungal organisms and epiparastic of photosynthetic plants which are symbionts to the fungal host. Because P. andromedea is achlorophyllous[12] this relationship is an obligate symbiosis for it, but is not ubiquitous in the fungal host. All monotropoideae are host specific to a select few fungal counterparts which in turn makes them specific to the photosynthetic organism associated with their fungal host. In the case of P. andromedea fungal host specificity leans heavily towards Rhizopogon salebrosus[13] in the western distribution and Rhizopogon kretzerae[14][15] in the eastern distribution but broadly seems to be ubiquitous symbionts with Rhizopogon subgenus Amylopogon.[3][16] Rhizopogon species also exhibit high host specificity and sub-genus Amylopogon is primarily associated with the Pinus genus. Fungal exoenzymatic activity has been shown to be required for seed germination of P. andromedea however the requisite enzymes are not exclusively produced within subgenus Amylopogon indicating that seed colonization by fungi outside of the observed host specificity is possible however ecologically restricted by some currently unknown mechanism.[17] Though often described as a parasitic relationship, there is some evidence that it may be a mutualistic partnership.[10]

Taxonomy

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Illustration from John Lindley's Collectanea botanica

Pterospora andromedea was first scientifically described by Thomas Nuttall in 1818. He based his work on specimens collected by Charles Whitlow in Canada near Niagara Falls. Amos Eaton published a description of the plant as Monotropa procera at almost the same time, but available evidence shows that Nuttall's publication was before Eaton's.[18] As of 2024 Pterospora andromedea is the only species in the genus, making it a monotypic taxon.[19]

Pterospora has consistently been shown to be more closely related to Sarcodes than any other member of the Monotropoideae.[20]

Names

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The genus name is derived from the morphology of the seeds which have narrow flaps of tissue on the side and therefore appear winged: pteron (Gr.) = wing, spora (Gr.) = seed. The specific name andromedea derives from the resemblance of the flowers to those of another genus in the Ericaceae, Andromeda.[2][3]

In English it is known by the common name "pinedrops" and the similar "woodland pinedrops" and "giant pinedrops".[21][22][1] Because Edwin James found the plant near Albany, New York it is also known as "Albany beechdrops".[23] Another name is "giant bird's-nest".[24]

In the Salish–Spokane–Kalispel language this species is called "senchelep tapemis" which can be translated as "coyote's arrow".[25] In the Jemez language its name translates as "Elk Girl tree".[26]

Range and habitat

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The fruits are five-celled woody capsules (Wenatchee Mountains, Washington).

Pterospora andromedea is native to North America from Alaska to central Mexico.[27] It has two populations, the larger in western North America and the other in the east, separated by the Great Plains.[10] In Canada it is found in the western provinces of British Columbia, Alberta, and Saskatchewan and in the eastern provinces of New Brunswick, Ontario, Prince Edward Island, and Québec.[27] In the western United States it is found widely in Arizona, California, Colorado, Idaho, Montana, Nevada, New Mexico, Oregon, South Dakota, Utah, Washington state, and Wyoming. It has also been reported in Sioux County, Nebraska and without county level distributions in Texas. America's eastern population is found in Michigan, New Hampshire, New York, Vermont, Wisconsin, and possibly in Pennsylvania.[28] It grows or has grown in many Mexican states including Chihuahua, Coahuila, Durango, Hidalgo, Mexico City, Mexico State, Morelos, Nuevo Leon, Oaxaca, Puebla, Sonora, Tlaxcala, and Veracruz.[29] It is also very rarely found in the highest elevations of Sierra de San Pedro Mártir in Baja California.[30]

In the western United States it is mainly found under conifer trees in areas with deep accumulations of needles and dry to moderately moist soils,[11] but may also be found under some oak trees.[6] More specifically they are found in Rocky Mountain Douglas-fir forests,[21] ponderosa pine forests,[7] the Sierra Nevada upper montane forest,[31] and under eastern white pines and balsam firs around the Great Lakes (especially on dunes that have become forested).[32]

Ecology

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In addition to not photosynthesizing, pinedrops are also a protocarnivorous plant. The sticky stems trap small insects and enzymes break them down for nitrogen.[7]

Though not a typical hummingbird plant, the broad-tailed hummingbird (Selasphorus platycercus) has been seen drinking nectar from pinedrop flowers.[33]

Conservation

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In 2019 NatureServe evaluated the conservation status species and as "globally secure" (G5), a species that is not significantly declining over the whole of its range. At the state and provincial level it also found it to be "secure" S5 in British Columbia and "apparently secure" (S4) in Montana. However, it is "vulnerable" (S3) in Wyoming, Nevada, and Alberta. In the rest of the west they evaluated it as "imperiled" (S2) in Nebraska and "critically imperiled" (S1) in Texas and Saskatchewan. In the eastern part of its range it imperiled or critically imperiled in every state and province with the exceptions of Massachusetts, where the species was not evaluated, and Prince Edward Island, where it is possibly extirpated. It is imperiled in Michigan, Ontario, Quebec and critically imperiled in New Brunswick, New Hampshire, New York, Vermont, and Wisconsin.[1]

Uses

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The seed stalks are occasionally collected to be used as long lasting ornaments in dry floral arrangements.[11] Though the flowers are considered desirable by plant collectors,[34] the very specific requirements of its fungal host make this plant practically impossible to transplant.[35]

See also

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References

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  1. ^ a b c NatureServe (2024). "Pterospora andromedea". Arlington, Virginia. Retrieved 28 March 2024.
  2. ^ a b Wallace, G.D. (1975), Studies of the Monotropoidiae (Ericaceae): taxonomy and distribution, The Wassman Journal of Biology
  3. ^ a b c Wallace, Gary D. "Pterospora Nuttall". Flora of North America @ efloras.org. Retrieved 1 May 2024.
  4. ^ a b c Wallace, Gary D. (5 November 2020). "Pterospora andromedea - FNA". Flora of North America. Retrieved 28 March 2024.
  5. ^ Bidartondo, Martin I. (August 2005). "The evolutionary ecology of myco-heterotrophy". New Phytologist. 167 (2): 335–352. doi:10.1111/j.1469-8137.2005.01429.x. PMID 15998389.
  6. ^ a b c d e Heil, Kenneth D.; O’Kane, Jr., Steve L.; Reeves, Linda Mary; Clifford, Arnold (2013). Flora of the Four Corners Region : Vascular Plants of the San Juan River Drainage, Arizona, Colorado, New Mexico, and Utah. St. Louis, Missouri: Missouri Botanical Garden. pp. 494, 499. Retrieved 30 April 2024.
  7. ^ a b c Dodson, Carolyn; Dunmire, William W. (2007). Mountain Wildflowers of the Southern Rockies : Revealing Their Natural History. Albuquerque, New Mexico: University of New Mexico Press. pp. 44–45. ISBN 978-0-8263-4244-7. Retrieved 2 May 2024.
  8. ^ a b Henderson, Margaret W. (1919). "A Comparative Study of the Structure and Saprophytism of the Pyrolaceae and Monotropaceae with Reference to Their Derivation from the Ericaceae". Contributions from the Botanical Laboratory of the University of Pennsylvania. 5 (1). University of Pennsylvania Press: 52, 76. Retrieved 3 May 2024.
  9. ^ Ferguson, Mary; Saunders, Richard M. (1982). Canadian Wildflowers Through the Seasons. Toronto, Canada: Van Nostrand Reinhold. p. 90. ISBN 0-7706-0018-2. Retrieved 2 May 2024.
  10. ^ a b c Li, Jianhua; Corajod, Jeffrey; Vander Stel, Holly; Homkes, Austin (2011). "The Mycorrhizal System of Pterospora Andromedea (Pine-Drops) in West Michigan Inferred from DNA Sequence Data". The Michigan Botanist. 50 (3): 129–136. ISSN 2166-4374. Retrieved 1 May 2024.
  11. ^ a b c d Craighead, John Johnson; Craighead Jr., Frank D.; Davis, Ray J. (1963). A Field Guide to Rocky Mountain Wildflowers : From Northern Arizona and New Mexico to British Columbia (5th printing ed.). Boston, Massachusetts: Houghton Mifflin. p. 136. ISBN 978-0-395-18324-3. Retrieved 30 April 2024.
  12. ^ Cummings, Michael P.; Welschmeyer, Nicholas A. (1998), "Pigment composition of putatively achlorophyllous angiosperms", Plant Systematics and Evolution, 210 (1–2): 105, Bibcode:1998PSyEv.210..105C, doi:10.1007/BF00984730
  13. ^ Dowie, Nicholas J.; Grubisha, Lisa C.; Burton, Brent A.; Klooster, Matthew R.; Miller, Steven L. (2017-01-02). "Increased phylogenetic resolution within the ecologically important Rhizopogon subgenus Amylopogon using 10 anonymous nuclear loci". Mycologia. 109 (1): 35–45. doi:10.1080/00275514.2017.1285165. ISSN 0027-5514. PMID 28402794. S2CID 12476105.
  14. ^ Grubisha, Lisa C.; Dowie, Nicholas J.; Miller, Steven L.; Hazard, Christina; Trowbridge, Steven M.; Horton, Thomas R.; Klooster, Matthew R. (July 2014). "Rhizopogon kretzerae sp. nov.: the rare fungal symbiont in the tripartite system with Pterospora andromedea and Pinus strobus". Botany. 92 (7): 527–534. doi:10.1139/cjb-2013-0309. ISSN 1916-2790.
  15. ^ Hazard, Christina; Lilleskov, Erik A.; Horton, Thomas R. (2011-10-12). "Is rarity of pinedrops (Pterospora andromedea) in eastern North America linked to rarity of its unique fungal symbiont?". Mycorrhiza. 22 (5): 393–402. doi:10.1007/s00572-011-0414-y. ISSN 0940-6360. PMID 21989709. S2CID 3259872.
  16. ^ Cullings, K. W.; Szaro, T. M.; Bruns, T. D. (January 1996). "Evolution of extreme specialization within a lineage of ectomycorrhizal epiparasites". Nature. 379 (6560): 63–66. Bibcode:1996Natur.379...63C. doi:10.1038/379063a0.
  17. ^ BRUNS, THOMAS D.; READ, DAVID J. (2000-11-27). "In vitro germination of nonphotosynthetic, myco-heterotrophic plants stimulated by fungi isolated from the adult plants". New Phytologist. 148 (2): 335–342. doi:10.1046/j.1469-8137.2000.00766.x. ISSN 0028-646X.
  18. ^ Wallace, Gary D. (1 January 2022). "A Question of Priority: Pterospora andromedea Nuttall vs. Monotropa procera Torrey ex Eaton (Monotropoideae, Ericaceae)". Aliso: A Journal of Systematic and Floristic Botany. 40 (1): 57–63. doi:10.5642/aliso.BWQH1814. ISSN 0065-6275. Retrieved 28 March 2024.
  19. ^ "Pterospora Nutt". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 1 May 2024.
  20. ^ Bidartondo, M. I.; Bruns, T. D. (September 2001). "Extreme specificity in epiparasitic Monotropoideae (Ericaceae): widespread phylogenetic and geographical structure". Molecular Ecology. 10 (9): 2285–2295. Bibcode:2001MolEc..10.2285B. doi:10.1046/j.1365-294X.2001.01358.x. PMID 11555270.
  21. ^ a b Roberts, Rhoda N.; Nelson, Ruth Ashton (1967). Mountain Wild Flowers of Colorado and Adjacent Areas. Denver, Colorado: Denver Museum of Natural History. pp. 6, 34. Retrieved 1 May 2024.
  22. ^ Shaw, Richard J.; On, Danny (1979). Plants of Waterton-Glacier National Parks, and the Northern Rockies. Missoula, Montana: Mountain Press Publishing Company. p. 146. ISBN 978-0-87842-114-5. Retrieved 1 May 2024.
  23. ^ Clarke, John M.; House, Homer D. (1925). "Report of the State Botanist for 1924". New York State Museum Bulletin. 266. Albany, N.Y. : University of the State of New York: 11. Retrieved 1 May 2024.
  24. ^ Armstrong, Margaret; Thornber, J. J. (John James) (1915). Field Book of Western Wild Flowers. New York: C. P. Putnam's Sons. p. 360. Retrieved 1 May 2024.
  25. ^ Hart, Jeffrey A. (1979). "The Ethnobotany of the Flathead Indians of Western Montana". Botanical Museum Leaflets. 27 (10). Cambridge, Massachusetts: Harvard University: 282. Retrieved 30 April 2024.
  26. ^ Cook, Sarah Louise (1930). The Ethnobotany of Jemez Indians (Masters thesis). University of New Mexico. p. 26. Retrieved 21 May 2024.
  27. ^ a b "Pterospora andromedea Nutt". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 1 May 2024.
  28. ^ NRCS (1 May 2024), "Pterospora andromedea", PLANTS Database, United States Department of Agriculture (USDA)
  29. ^ Hassler, Michael (17 April 2024). "World Plants". Synonymic Checklist and Distribution of the World Flora. Version 19.2. Retrieved 1 May 2024.
  30. ^ Rebman, J. P.; Gibson, J.; Rich, K. (2016). "Annotated checklist of the vascular plants of Baja California, Mexico" (PDF). San Diego Society of Natural History. 45: 139.
  31. ^ Horn, Elizabeth L. (1998). Sierra Nevada Wildflowers. Missoula, Montana: Mountain Press Publishing Company. p. 4. ISBN 978-0-87842-388-0. Retrieved 3 May 2024.
  32. ^ Weatherbee, Ellen Elliott (2006). Guide to Great Lakes Coastal Plants. Ann Arbor, Michigan: University of Michigan Press. pp. 122–123. ISBN 978-0-472-03015-6. Retrieved 3 May 2024.
  33. ^ Waser, Nickolas M.; CaraDonna, Paul J.; Price, Mary V. (November 2018). "Atypical Flowers Can Be as Profitable as Typical Hummingbird Flowers". The American Naturalist. 192 (5): 644–653. doi:10.1086/699836. hdl:10150/631593. PMID 30332579.
  34. ^ Lindley, John (1821). Collectanea Botanica, or, Figures and Botanical Illustrations of Rare and Curious Exotic Plants (in Latin and English). London: Richard and Arthur Taylor, Shoe-Lane. p. 5. Retrieved 3 May 2024.
  35. ^ Trull, Sue. "Woodland Pinedrops". Plant of the Week. U.S. Forest Service, United States Department of Agriculture. Retrieved 3 May 2024.
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