Difference between revisions of "Buchnera americana"

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| range_map_caption = Natural range of ''Buchnera americana'' from USDA NRCS [http://www.plants.usda.gov/core/profile?symbol=BUAM Plants Database].
 
| range_map_caption = Natural range of ''Buchnera americana'' from USDA NRCS [http://www.plants.usda.gov/core/profile?symbol=BUAM Plants Database].
 
}}
 
}}
Common name: American bluehearts
+
Common names: American Bluehearts; Prairie Bluehearts; Plains Bluehearts
 
==Taxonomic notes==
 
==Taxonomic notes==
 +
Synonyms: none <ref name=weakley>Weakley, A.S. 2020. Flora of the Southeastern United States. Edition of 20 October 2020. University of North Carolina at Chapel Hill, Chapel Hill, North Carolina.</ref>
 +
 +
Varieties: none<ref name=weakley/>
 +
 
==Description==  
 
==Description==  
 
<!-- Basic life history facts such as annual/perrenial, monoecious/dioecious, root morphology, seed type, etc. -->
 
<!-- Basic life history facts such as annual/perrenial, monoecious/dioecious, root morphology, seed type, etc. -->
''Buchnera americana'' is a hispid to pilose perennial with erect, simple stems growing up to 4-8 dm tall. The entire plant turns black when dried. The leaves are opposite, elliptic to ovate-lanceolate in shape. The leaves are entire or irregularly serrate or crenulate, sessile. The inflorescence is an open spike, the flower are in the axils of opposite bracts and subtended by 2 bractlets. The calyx tube is cylindrical, grows up to 3.5-5 mm long, the lobes 5, lanceolate in shape, slightly unequal, up to 1 mm or less long. The corolla is purple or white in color, subsalverform, weakly zygomorphic, lobes 5, shorter than the calyx tube. Up to 4 fertile stamens are present with anthers with only a single sac. The capsule is ovoid or pyriform, grows  up to 5-6 mm long and is loculicidal in shape.<ref name="Radford">Radford, Albert E., Harry E. Ahles, and C. Ritchie Bell. Manual of the Vascular Flora of the Carolinas. 1964, 1968. The University of North Carolina Press. 954-5. Print.</ref>
+
''Buchnera americana'' is a hairy perennial plant with erect, simple stems growing between 40 - 80 cm tall. The leaves are oppositely arranged and elliptic to ovate-lanceolate in shape with entire or irregularly serrate margins and are noticeably 3-veined. The inflorescence is an open spike with the flowers in the axils of opposite bracts and supported by 2 bractlets. The calyx tube is cylindrical and grows up to 10 - 12 mm long with 5 lobes that are lanceolate in shape and roughly 5 - 8 mm long. The bilaterally symmetrical flowers are purple or white in color with 5 petals that form a narrow tube and abruptly bend out at right angles. There are up to 4 fertile stamens present with anthers with a single sac. The capsule seeds are ovoid or pyriform in shape and 5 - 6 mm long.<ref name="Radford">Radford, Albert E., Harry E. Ahles, and C. Ritchie Bell. Manual of the Vascular Flora of the Carolinas. 1964, 1968. The University of North Carolina Press. 954-5. Print.</ref>
 
 
Specifically, for ''Buchnera americana'', they have noticeable 3-veined leaves that are ovate-lanceolate in shape. The corolla is purple in color. The calyx tube is 10-12 mm long, the lobes are 5-8 mm long. Flowers from July to September.<ref name="Radford"></ref>
 
  
 
==Distribution==
 
==Distribution==
Critically imperiled in Ontario, Georgia, Virginia, North Carolina, and Indiana.<ref name="natureserve"/> It is the most widely distributed North American ''Buchnera'' species, and is found historically in 24 states.<ref name="wildflower">[[http://www.wildflower.org/plants/result.php?id_plant=BUAM]]Lady Bird Johnson Wildflower Center. Accessed: April 4, 2016</ref>
+
It is the most widely distributed North American ''Buchnera'' species, and is found historically in 24 states.<ref name="wildflower">[[https://www.wildflower.org/plants/search.php?search_field=&newsearch=true]]Lady Bird Johnson Wildflower Center. Accessed: April 4, 2016</ref>
  
 
==Ecology==
 
==Ecology==
''B. americana'' is a hemiparasitic species, meaning that it gets nourishment from the roots of other plants, but also produces its own nourishment through photosynthesis, and can survive with out a host. Hemiparasitic plants have modified roots called haustoria, that allow it to attach to a host plant and obtain nutrients. Host for this species include: ''Pinus clausa, P. echinata, P. elliottii, P. palustris, P. strobes, P. taeda, Celtis laevigata, Fraxinus pensylvanica, Liquidambar styraciflua, Liriodendron tulipifera, Nyssa aquatica, N. sylvatica, Platanus occidentalis, Populus deltoids, Quercus alba, Q. falcata'', and ''Q. shumardii''<ref name="natureserve">[[http://explorer.natureserve.org/servlet/NatureServe?searchName=Buchnera+americana]]NatureServe. Accessed: April 4, 2016</ref>.
+
''B. americana'' is a hemiparasitic species, meaning that it gets nourishment from the roots of other plants, but also produces its own nourishment through photosynthesis, and can survive with out a host. Hemiparasitic plants have modified roots called haustoria, that allow it to attach to a host plant and obtain nutrients. Host for this species include: ''Pinus clausa, [[Pinus echinata]], [[Pinus elliottii]], [[Pinus palustris]], Pinus strobes, [[Pinus taeda]], Celtis laevigata, Fraxinus pensylvanica, [[Liquidambar styraciflua]], Liriodendron tulipifera, Nyssa aquatica, [[Nyssa sylvatica]], Platanus occidentalis, Populus deltoids, Quercus alba, [[Quercus falcata]]'', and ''Quercus shumardii''<ref name="natureserve">[[http://explorer.natureserve.org/servlet/NatureServe?searchName=Buchnera+americana]]NatureServe. Accessed: April 4, 2016</ref>. However, one study found that ''B. americana'' can mature without ever attaching to a host plant.<ref name= "Musselman2">Musselman, L. J. and W. F. Mann, Jr (1978). Root parasites of southern forests. , USDA Forest Service, Southern For. Exp. Station, New Orleans, LA. Gen. Tech. Rpt. SO-20. : 76.</ref>
 
===Habitat=== <!--Natural communities, human disturbed habitats, topography, hydrology, soils, light, fire regime requirements for removal of competition, etc.-->
 
===Habitat=== <!--Natural communities, human disturbed habitats, topography, hydrology, soils, light, fire regime requirements for removal of competition, etc.-->
 
''B. americana'' is found in moist environments near ponds and low brackish marshes, at the edges of hammocks, and in hillside bogs. However, it can also appear in drier situations like savannas, pine flatwoods, dry open woods, hillside prairies, open prairies, coastal dunes, and shallow, grassy depressions. In addition, it can be found in disturbed areas such as roadsides, ditches, sandy fields, open pastures, and clearings that have become weedy. This species prefers open, sunny conditions, but occurs in a range of soil types, from dry loamy sand, to moist black sandy peat, seasonally damp areas, acidic sandy soils, and clay soils.<ref name="fsu">Florida State University Robert K. Godfrey Herbarium database. URL: http://herbarium.bio.fsu.edu. Last accessed: June 2014. Collectors: George R. Cooley, R. J. Eaton, O. Lakela, R. Kral, Elmer C. Prichard, Loran C. Anderson, Norlan C. Henderson, Mabel Kral, F. H. Sargent, Carroll E. Wood, Jr., Mary Atkinson, William P. Adams, C. Jackson, R.K. Godfrey, William Lindsey, W. P. Adams, Gary R. Knight, Jane Brockmann, Jackie Patman, James D. Ray, Gary R. Knight, Jean W. Wooten, Raymond Athey, Bruce Hansen, U. T. Waterfall, Paul L. Redfearn, Jr., Charles T. Bryson, A.F. Clewell, S.W. Leonard, Samuel B. Jones, Jr., John W. Thieret, Mary Margaret Williams, D. S. Correll, Chas. C. Deam, R. B. Channel, Josephine Skehan, Delzie Demaree, D.B. Ward, S.S. Ward, H.L. Blomquist, Baltzell, Kenneth A. Wilson, Grady W. Reinert, Julie Neel, R. A. Norris, R. Komarek, Thomas E. Miller, and James R. Burkhalter. States and Counties: Alabama: Wilcox. Arkansas: Prairie. Florida: Calhoun, Citrus, Charlotte, Collier, De Soto, Dixie, Escambia, Franklin, Hamilton, Hardee, Hernando, Highlands, Hillsborough, Jackson, Jasper, Lee, Leon, Levy, Liberty, Martin, Monroe, Osceola, Okaloosa, Pasco, Polk, Sarasota, St Johns, Taylor, Volusia, and Wakulla. Georgia: Charlton, Grady, McIntosh, and Thomas. Kentucky: Coffee and Lyon. Louisiana: Jefferson-Davis. Mississippi: Clay, Forrest, Jackson, and Newton. Missouri: Cambria, Cedar, Polk, and Vernon. North Carolina: Brunswick and Carteret. Oklahoma: Payne. South Carolina: Lee. Texas: Galveston.</ref>
 
''B. americana'' is found in moist environments near ponds and low brackish marshes, at the edges of hammocks, and in hillside bogs. However, it can also appear in drier situations like savannas, pine flatwoods, dry open woods, hillside prairies, open prairies, coastal dunes, and shallow, grassy depressions. In addition, it can be found in disturbed areas such as roadsides, ditches, sandy fields, open pastures, and clearings that have become weedy. This species prefers open, sunny conditions, but occurs in a range of soil types, from dry loamy sand, to moist black sandy peat, seasonally damp areas, acidic sandy soils, and clay soils.<ref name="fsu">Florida State University Robert K. Godfrey Herbarium database. URL: http://herbarium.bio.fsu.edu. Last accessed: June 2014. Collectors: George R. Cooley, R. J. Eaton, O. Lakela, R. Kral, Elmer C. Prichard, Loran C. Anderson, Norlan C. Henderson, Mabel Kral, F. H. Sargent, Carroll E. Wood, Jr., Mary Atkinson, William P. Adams, C. Jackson, R.K. Godfrey, William Lindsey, W. P. Adams, Gary R. Knight, Jane Brockmann, Jackie Patman, James D. Ray, Gary R. Knight, Jean W. Wooten, Raymond Athey, Bruce Hansen, U. T. Waterfall, Paul L. Redfearn, Jr., Charles T. Bryson, A.F. Clewell, S.W. Leonard, Samuel B. Jones, Jr., John W. Thieret, Mary Margaret Williams, D. S. Correll, Chas. C. Deam, R. B. Channel, Josephine Skehan, Delzie Demaree, D.B. Ward, S.S. Ward, H.L. Blomquist, Baltzell, Kenneth A. Wilson, Grady W. Reinert, Julie Neel, R. A. Norris, R. Komarek, Thomas E. Miller, and James R. Burkhalter. States and Counties: Alabama: Wilcox. Arkansas: Prairie. Florida: Calhoun, Citrus, Charlotte, Collier, De Soto, Dixie, Escambia, Franklin, Hamilton, Hardee, Hernando, Highlands, Hillsborough, Jackson, Jasper, Lee, Leon, Levy, Liberty, Martin, Monroe, Osceola, Okaloosa, Pasco, Polk, Sarasota, St Johns, Taylor, Volusia, and Wakulla. Georgia: Charlton, Grady, McIntosh, and Thomas. Kentucky: Coffee and Lyon. Louisiana: Jefferson-Davis. Mississippi: Clay, Forrest, Jackson, and Newton. Missouri: Cambria, Cedar, Polk, and Vernon. North Carolina: Brunswick and Carteret. Oklahoma: Payne. South Carolina: Lee. Texas: Galveston.</ref>
  
Associated species includes slash pine, longleaf pine, saw palmetto, cypress, ''Hypericum opacum, Eryngium integrifolium, Xyris elliottii, Habenaria integra, Polygala ramosa, Polygala lutea, Ctenium aromaticus, Hibiscus aculeatus, Lobelia, Baccharis, Myrica, Erigeron strigosus, Polypremum procumbens'', and others.<ref name="fsu"/>
+
Associated species includes slash pine, longleaf pine, saw palmetto, cypress, ''Hypericum opacum, [[Eryngium integrifolium]], [[Xyris elliottii]], Habenaria integra, Polygala ramosa, [[Polygala lutea]], Ctenium aromaticus, [[Hibiscus aculeatus]], Lobelia, Baccharis, Myrica, [[Erigeron strigosus]], [[Polypremum procumbens]]'', and others.<ref name="fsu"/>
  
===Phenology=== <!--Timing off flowering, fruiting, seed dispersal, and environmental triggers.  Cite PanFlora website if appropriate: http://www.gilnelson.com/PanFlora/ -->
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===Phenology=== <!--Timing off flowering, fruiting, and environmental triggers.  Cite PanFlora website if appropriate: http://www.gilnelson.com/PanFlora/ -->
 
''B. americana'' has been observed flowering as early as January, through October with peak inflorescence in April and May.<ref>Nelson, G.  [http://www.gilnelson.com/ PanFlora]: Plant data for the eastern United States with emphasis on the Southeastern Coastal Plains, Florida, and the Florida Panhandle. www.gilnelson.com/PanFlora/  Accessed: 7 DEC 2016</ref> Fruiting has been observed February through November.<ref name="fsu"/>
 
''B. americana'' has been observed flowering as early as January, through October with peak inflorescence in April and May.<ref>Nelson, G.  [http://www.gilnelson.com/ PanFlora]: Plant data for the eastern United States with emphasis on the Southeastern Coastal Plains, Florida, and the Florida Panhandle. www.gilnelson.com/PanFlora/  Accessed: 7 DEC 2016</ref> Fruiting has been observed February through November.<ref name="fsu"/>
 
  <!--===Seed dispersal===-->
 
  <!--===Seed dispersal===-->
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===Fire ecology===<!--Fire tolerance, fire dependence, adaptive fire responses-->  
 
===Fire ecology===<!--Fire tolerance, fire dependence, adaptive fire responses-->  
 
Fire kills plants that made shade out ''B. americana''.  
 
Fire kills plants that made shade out ''B. americana''.  
 +
 
===Pollination===
 
===Pollination===
Pollinated by butterflies.<ref name="regist"/>
+
''Buchnera americana'' is pollinated by butterflies.<ref name="regist"/>  
<!--===Use by animals===--><!--Herbivory, granivory, insect hosting, etc.-->  
+
 
 +
<!--===Herbivory and toxicology===-->
 
<!--===Diseases and parasites===-->
 
<!--===Diseases and parasites===-->
  
==Conservation and management==
+
==Conservation, cultivation, and restoration==
 
+
It is listed as endangered by the Committee on the Status of Endangered Wildlife in Canada for Ontario, the Indiana Department of Natural Resources, the Illinois Endangered Species Protection Board, the Maryland Department of Natural Resources, and the New York Division of Land and Forests.<ref name="natureserve"/><ref name= "Board">Illinois Endangered Species Protection Board. (2015). "Checklist of Illinois Endangered and Threatened Animals and Plants."</ref><ref name= "USDA">USDA, NRCS. (2016). The PLANTS Database (http://plants.usda.gov, 27 March 2019). National Plant Data Team, Greensboro, NC 27401-4901 USA.</ref> ''B. americana'' is also listed as threatened by the Ohio Department of Natural Resources, as possibly extirpated by the Michigan Department of Natural Resources, and as extirpated by the Maryland Department of Natural Resources and the Pennsylvania Department of Conservation and Natural Resources.<ref name= "USDA"/> While ''B. americana'' is considered to be an abundant weed on roadsides in central and southern Florida, it was noted that it does not seem to do any damage.<ref name= "Musselman">Musselman, L. J. (1996). "Parasitic weeds in the southern United States." Castanea 61: 271-292.</ref> This species is recommended for natural landscapes and habitat restorations, also recommended for butterfly gardens. Prefers moist to wet, well-drained to poorly drained sandy or limestone soils and does not tolerate long term flooding.<ref name="regional">[[http://www.regionalconservation.org/beta/nfyn/plantdetail.asp?tx=Buchamer]]Regional Conservation. Accessed: April 4, 2016</ref>
==Cultivation and restoration==
 
This species is recommended for natural landscapes and habitat restorations, also recommended for butterfly gardens. Prefers moist to wet, well-drained to poorly drained sandy or limestone soils and does not tolerate long term flooding.<ref name="regional">[[http://www.regionalconservation.org/beta/nfyn/plantdetail.asp?tx=Buchamer]]Regional Conservation. Accessed: April 4, 2016</ref>
 
  
 +
==Cultural use==
 
==Photo Gallery==
 
==Photo Gallery==
 
<gallery widths=180px>
 
<gallery widths=180px>
 
</gallery>
 
</gallery>
 
==References and notes==
 
==References and notes==

Latest revision as of 13:40, 22 May 2023

Buchnera americana
Buchnera americana Gil.jpg
photo by Gil Nelson
Scientific classification
Kingdom: Plantae
Division: Magnoliophyta - Flowering plants
Class: Magnoliopsida - Dicotyledons
Order: Scrophulariales
Family: Scrophulariaceae
Genus: Buchnera
Species: B. americana
Binomial name
Buchnera americana
L.
BUCH AMER dist.jpg
Natural range of Buchnera americana from USDA NRCS Plants Database.

Common names: American Bluehearts; Prairie Bluehearts; Plains Bluehearts

Taxonomic notes

Synonyms: none [1]

Varieties: none[1]

Description

Buchnera americana is a hairy perennial plant with erect, simple stems growing between 40 - 80 cm tall. The leaves are oppositely arranged and elliptic to ovate-lanceolate in shape with entire or irregularly serrate margins and are noticeably 3-veined. The inflorescence is an open spike with the flowers in the axils of opposite bracts and supported by 2 bractlets. The calyx tube is cylindrical and grows up to 10 - 12 mm long with 5 lobes that are lanceolate in shape and roughly 5 - 8 mm long. The bilaterally symmetrical flowers are purple or white in color with 5 petals that form a narrow tube and abruptly bend out at right angles. There are up to 4 fertile stamens present with anthers with a single sac. The capsule seeds are ovoid or pyriform in shape and 5 - 6 mm long.[2]

Distribution

It is the most widely distributed North American Buchnera species, and is found historically in 24 states.[3]

Ecology

B. americana is a hemiparasitic species, meaning that it gets nourishment from the roots of other plants, but also produces its own nourishment through photosynthesis, and can survive with out a host. Hemiparasitic plants have modified roots called haustoria, that allow it to attach to a host plant and obtain nutrients. Host for this species include: Pinus clausa, Pinus echinata, Pinus elliottii, Pinus palustris, Pinus strobes, Pinus taeda, Celtis laevigata, Fraxinus pensylvanica, Liquidambar styraciflua, Liriodendron tulipifera, Nyssa aquatica, Nyssa sylvatica, Platanus occidentalis, Populus deltoids, Quercus alba, Quercus falcata, and Quercus shumardii[4]. However, one study found that B. americana can mature without ever attaching to a host plant.[5]

Habitat

B. americana is found in moist environments near ponds and low brackish marshes, at the edges of hammocks, and in hillside bogs. However, it can also appear in drier situations like savannas, pine flatwoods, dry open woods, hillside prairies, open prairies, coastal dunes, and shallow, grassy depressions. In addition, it can be found in disturbed areas such as roadsides, ditches, sandy fields, open pastures, and clearings that have become weedy. This species prefers open, sunny conditions, but occurs in a range of soil types, from dry loamy sand, to moist black sandy peat, seasonally damp areas, acidic sandy soils, and clay soils.[6]

Associated species includes slash pine, longleaf pine, saw palmetto, cypress, Hypericum opacum, Eryngium integrifolium, Xyris elliottii, Habenaria integra, Polygala ramosa, Polygala lutea, Ctenium aromaticus, Hibiscus aculeatus, Lobelia, Baccharis, Myrica, Erigeron strigosus, Polypremum procumbens, and others.[6]

Phenology

B. americana has been observed flowering as early as January, through October with peak inflorescence in April and May.[7] Fruiting has been observed February through November.[6]

Seed bank and germination

Seeds can remain viable in the soil for 3 years. Requires light for germination.[8]

Fire ecology

Fire kills plants that made shade out B. americana.

Pollination

Buchnera americana is pollinated by butterflies.[8]


Conservation, cultivation, and restoration

It is listed as endangered by the Committee on the Status of Endangered Wildlife in Canada for Ontario, the Indiana Department of Natural Resources, the Illinois Endangered Species Protection Board, the Maryland Department of Natural Resources, and the New York Division of Land and Forests.[4][9][10] B. americana is also listed as threatened by the Ohio Department of Natural Resources, as possibly extirpated by the Michigan Department of Natural Resources, and as extirpated by the Maryland Department of Natural Resources and the Pennsylvania Department of Conservation and Natural Resources.[10] While B. americana is considered to be an abundant weed on roadsides in central and southern Florida, it was noted that it does not seem to do any damage.[11] This species is recommended for natural landscapes and habitat restorations, also recommended for butterfly gardens. Prefers moist to wet, well-drained to poorly drained sandy or limestone soils and does not tolerate long term flooding.[12]

Cultural use

Photo Gallery

References and notes

  1. 1.0 1.1 Weakley, A.S. 2020. Flora of the Southeastern United States. Edition of 20 October 2020. University of North Carolina at Chapel Hill, Chapel Hill, North Carolina.
  2. Radford, Albert E., Harry E. Ahles, and C. Ritchie Bell. Manual of the Vascular Flora of the Carolinas. 1964, 1968. The University of North Carolina Press. 954-5. Print.
  3. [[1]]Lady Bird Johnson Wildflower Center. Accessed: April 4, 2016
  4. 4.0 4.1 [[2]]NatureServe. Accessed: April 4, 2016
  5. Musselman, L. J. and W. F. Mann, Jr (1978). Root parasites of southern forests. , USDA Forest Service, Southern For. Exp. Station, New Orleans, LA. Gen. Tech. Rpt. SO-20. : 76.
  6. 6.0 6.1 6.2 Florida State University Robert K. Godfrey Herbarium database. URL: http://herbarium.bio.fsu.edu. Last accessed: June 2014. Collectors: George R. Cooley, R. J. Eaton, O. Lakela, R. Kral, Elmer C. Prichard, Loran C. Anderson, Norlan C. Henderson, Mabel Kral, F. H. Sargent, Carroll E. Wood, Jr., Mary Atkinson, William P. Adams, C. Jackson, R.K. Godfrey, William Lindsey, W. P. Adams, Gary R. Knight, Jane Brockmann, Jackie Patman, James D. Ray, Gary R. Knight, Jean W. Wooten, Raymond Athey, Bruce Hansen, U. T. Waterfall, Paul L. Redfearn, Jr., Charles T. Bryson, A.F. Clewell, S.W. Leonard, Samuel B. Jones, Jr., John W. Thieret, Mary Margaret Williams, D. S. Correll, Chas. C. Deam, R. B. Channel, Josephine Skehan, Delzie Demaree, D.B. Ward, S.S. Ward, H.L. Blomquist, Baltzell, Kenneth A. Wilson, Grady W. Reinert, Julie Neel, R. A. Norris, R. Komarek, Thomas E. Miller, and James R. Burkhalter. States and Counties: Alabama: Wilcox. Arkansas: Prairie. Florida: Calhoun, Citrus, Charlotte, Collier, De Soto, Dixie, Escambia, Franklin, Hamilton, Hardee, Hernando, Highlands, Hillsborough, Jackson, Jasper, Lee, Leon, Levy, Liberty, Martin, Monroe, Osceola, Okaloosa, Pasco, Polk, Sarasota, St Johns, Taylor, Volusia, and Wakulla. Georgia: Charlton, Grady, McIntosh, and Thomas. Kentucky: Coffee and Lyon. Louisiana: Jefferson-Davis. Mississippi: Clay, Forrest, Jackson, and Newton. Missouri: Cambria, Cedar, Polk, and Vernon. North Carolina: Brunswick and Carteret. Oklahoma: Payne. South Carolina: Lee. Texas: Galveston.
  7. Nelson, G. PanFlora: Plant data for the eastern United States with emphasis on the Southeastern Coastal Plains, Florida, and the Florida Panhandle. www.gilnelson.com/PanFlora/ Accessed: 7 DEC 2016
  8. 8.0 8.1 [[3]]Accessed: April 4, 2016
  9. Illinois Endangered Species Protection Board. (2015). "Checklist of Illinois Endangered and Threatened Animals and Plants."
  10. 10.0 10.1 USDA, NRCS. (2016). The PLANTS Database (http://plants.usda.gov, 27 March 2019). National Plant Data Team, Greensboro, NC 27401-4901 USA.
  11. Musselman, L. J. (1996). "Parasitic weeds in the southern United States." Castanea 61: 271-292.
  12. [[4]]Regional Conservation. Accessed: April 4, 2016