Difference between revisions of "Opuntia humifusa"

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Common names: Devil's-tongue; Eastern prickly pear
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Common names: Devil's-tongue; Eastern prickly pear<ref name="weakley">Weakley, A.S. 2015. Flora of the southern and mid-atlantic states. Working Draft of 21 May 2015. University of North Carolina at Chapel Hill, Chapel Hill, North Carolina.</ref>
 
==Taxonomic notes==
 
==Taxonomic notes==
Synonyms: ''Opuntia humifusa'' (Rafinesque) Rafinesque var. ''humifusa''; ''O. compressa'' (Salisbury) J.F. Macbride var. ''compressa''; ''O. compressa''; ''O. impedita'' Small; ''O. macrarthra'' Gibbes; ''O. opuntia'' (Linnaeus) Karten; ''O. calcicola'' Wherry; ''O. rafinesquei'' Engelmann
+
Synonyms: ''Opuntia compressa'' (Salisbury) J.F. Macbride var. ''compressa''; ''O. impedita'' Small; ''O. macrarthra'' Gibbes; ''O. opuntia'' (Linnaeus) Karten; ''O. calcicola'' Wherry; ''O. rafinesquei'' Engelmann.<ref name="weakley">Weakley, A.S. 2015. Flora of the southern and mid-atlantic states. Working Draft of 21 May 2015. University of North Carolina at Chapel Hill, Chapel Hill, North Carolina.</ref>
 +
 
 +
Varieties: none.<ref name="weakley">Weakley, A.S. 2015. Flora of the southern and mid-atlantic states. Working Draft of 21 May 2015. University of North Carolina at Chapel Hill, Chapel Hill, North Carolina.</ref>
  
 
==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. -->
 
A description of ''Opuntia humifusa'' is provided in [http://efloras.org/florataxon.aspx?flora_id=1&taxon_id=242415184 The Flora of North America].
 
A description of ''Opuntia humifusa'' is provided in [http://efloras.org/florataxon.aspx?flora_id=1&taxon_id=242415184 The Flora of North America].
 +
 +
The leaves of ''Opuntia humifusa'' are 1.5-6 cm long, with the longest leaves having an acute to acuminate tip. The hairs of the leaf surfaces are similarly long and the leaf sheaths and culm axis are glabrate to pilose with hairs shorter than 1.5 mm. The awns are glabrous and the lemma is 9-11 veined.<ref name="weakley">Weakley, A.S. 2015. Flora of the southern and mid-atlantic states. Working Draft of 21 May 2015. University of North Carolina at Chapel Hill, Chapel Hill, North Carolina.</ref>
 +
 +
The root system of ''Opuntia humifusa'' includes stem tubers which store non-structural carbohydrates (NSC) important for both resprouting following fire and persisting during long periods of fire exclusion.<ref name="Diaz"> Diaz-Toribio, M.H. and F. E. Putz 2021. Underground carbohydrate stores and storage organs in fire-maintained longleaf pine savannas in Florida, USA. American Journal of Botany 108: 432-442.</ref> Diaz-Toribio and Putz (2021) recorded this species to have an NSC concentration of 159.7 mg/g (ranking 32 out of 100 species studied) and water content of 47.2% (ranking 3 out of 100 species studied).<ref name = "Diaz"/>
 +
 +
According to Diaz-Torbio and Putz (2021), ''Opuntia humifusa'' has stem tubers with a below-ground to above-ground biomass ratio of 0.606 and nonstructural carbohydrate concentration of 159.7 mg g<sup>-1</sup>.<ref>Diaz‐Toribio, M. H. and F. E. Putz. 2021. Underground carbohydrate stores and storage organs in fire‐maintained longleaf pine savannas in Florida, USA. American Journal of Botany 108(3):432-442.</ref>
  
 
==Distribution==
 
==Distribution==
It is the only cactus to be widespread in the eastern United States. <ref name="wildflower">[[http://www.wildflower.org/plants/result.php?id_plant=OPHU]] Lady Bird Johnson Wildflower Center. Accessed: February 12, 2016</ref> It is found in southern Canada as well as the eastern United States. <ref name=gold> Goldstein, G. and P. S. Nobel (1994). "Water Relations and Low-Temperature Acclimation for Cactus Species Varying in Freezing Tolerance." Plant Physiology 104(2): 675-681.</ref>
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It is the only cactus to be widespread in the eastern United States.<ref name="wildflower">[[http://www.wildflower.org/plants/result.php?id_plant=OPHU]] Lady Bird Johnson Wildflower Center. Accessed: February 12, 2016</ref> It is mainly restricted to the Appalachian Mountains, in states like Deleware, Maryland, New Jersey, Virginia, and West Virginia, but also occurs in central and northcentral Mississippi. Further surveying could reveal populations in Alabama, northwestern Georgia, western South Carolina, western North Carolina, and northeastern Tennessee.<ref name="weakley">Weakley, A.S. 2015. Flora of the southern and mid-atlantic states. Working Draft of 21 May 2015. University of North Carolina at Chapel Hill, Chapel Hill, North Carolina.</ref>
  
 
==Ecology==
 
==Ecology==
Line 33: Line 41:
 
''Opuntia humifusa'' can occur in upland longleaf pines, sand dunes, ''Pinus clausa'' scrubs, open sand flats, sandhills, and pine/oak scrubs. It has been found in disturbed areas such as loblolly tree farms and roadside depressions. Associated species include ''Lyonia ferruginea, Lyonia lucida, Serenoa repens, Quercus geminata, Q. chapmanii, Persea humilis, Ceratiola, Osmanthus megacarpus, Rhynchospora megalocarpa, Galactia elliottii'', and ''Smilax auriculata.''<ref name="FSU Herbarium">Florida State University Robert K. Godfrey Herbarium database. URL: [http://herbarium.bio.fsu.edu http://herbarium.bio.fsu.edu]. Last accessed: October 2015. Collectors: Loran C. Anderson, Michael Blaker, M. Borgman, James R. Burkhaulter, George R. Cooley, R.J. Eaton,  Patricia Elliott, J. Kevin England, Robert K. Godfrey, Darren Jackson, Ed Keppner, Lisa Keppner, Andrew McAllister, Sidney McDaniel, K.M. Meyer, James D. Ray Jr., Erik Robinson, L. Rosen, C.E. Smith, A. Townesmith, Kenneth A. Wilson, Carroll E. Wood Jr.. States and Counties: Alabama: Dale. Florida: Alabama, Escambia, Franklin, Gadsden, Hernando, Jackson, Lafayette, Leon, Liberty, Orange, Putnam, Seminole, Walton, Wakulla, Washington. Compiled by Tall Timbers Research Station and Land Conservancy.</ref>
 
''Opuntia humifusa'' can occur in upland longleaf pines, sand dunes, ''Pinus clausa'' scrubs, open sand flats, sandhills, and pine/oak scrubs. It has been found in disturbed areas such as loblolly tree farms and roadside depressions. Associated species include ''Lyonia ferruginea, Lyonia lucida, Serenoa repens, Quercus geminata, Q. chapmanii, Persea humilis, Ceratiola, Osmanthus megacarpus, Rhynchospora megalocarpa, Galactia elliottii'', and ''Smilax auriculata.''<ref name="FSU Herbarium">Florida State University Robert K. Godfrey Herbarium database. URL: [http://herbarium.bio.fsu.edu http://herbarium.bio.fsu.edu]. Last accessed: October 2015. Collectors: Loran C. Anderson, Michael Blaker, M. Borgman, James R. Burkhaulter, George R. Cooley, R.J. Eaton,  Patricia Elliott, J. Kevin England, Robert K. Godfrey, Darren Jackson, Ed Keppner, Lisa Keppner, Andrew McAllister, Sidney McDaniel, K.M. Meyer, James D. Ray Jr., Erik Robinson, L. Rosen, C.E. Smith, A. Townesmith, Kenneth A. Wilson, Carroll E. Wood Jr.. States and Counties: Alabama: Dale. Florida: Alabama, Escambia, Franklin, Gadsden, Hernando, Jackson, Lafayette, Leon, Liberty, Orange, Putnam, Seminole, Walton, Wakulla, Washington. Compiled by Tall Timbers Research Station and Land Conservancy.</ref>
  
It occurs in southern Canada, where average nighttime temperatures can reach -4 degrees Celsius. In order to prevent intracellular freeze dehydration and ice formation, individuals have an accumulation of sugars and mannitol in their cells. <ref name=gold/>
+
''O. humifusa'' was unaffected by soil disturbance by backe seeding, high-severity burning, and salvage logging in central Florida sand pine scrub.<ref name=Greenberg> Greenberg, C.H., D.G. Neary, L.D. Harris, and S.P. Linda. (1995). Vegetation Recovery Following High-intensity Wildfire and Silvicultural Treatments in Sand Pine Scrub. American Midland Naturalist 133(1):149-163.</ref>
 +
 
 +
It had mixed responses to clearcutting in central Florida. In some cases the plant increased its cover and in other areas it was unaffected by the practice. It has shown regrowth and been unaffected in reestablished native sand pine scrub that was disturbed by clearcutting.<ref name=Greenberg/>
 +
 
 +
This species increased its cover in response to soil disturbance by single roller chopping and broadcast seeding in central Florida pine scrub habitat. It has shown regrowth in reestablished native habitat that was disturbed by these practices.<ref name=Greenberg/>
 +
 
 +
Additionally, ''O. humifusa'' reduced its frequency and density in response to soil disturbance by roller chopping in northwest Florida sandhills. It has shown resistance to regrowth and been unaffected by reestablished native sandhill habitat that was disturbed by this practice.<ref>Hebb, E.A. (1971). Site Preparation Decreases Game Food Plants in Florida Sandhills. The Journal of Wildlife Management 35(1):155-162.</ref>
 +
 
 +
It occurs in southern Canada, where average nighttime temperatures can reach -4 degrees Celsius. In order to prevent intracellular freeze dehydration and ice formation, individuals have an accumulation of sugars and mannitol in their cells.<ref name=gold/>
  
 
===Phenology=== <!--Timing off flowering, fruiting, seed dispersal, and environmental triggers.  Cite PanFlora website if appropriate: http://www.gilnelson.com/PanFlora/ -->
 
===Phenology=== <!--Timing off flowering, fruiting, seed dispersal, and environmental triggers.  Cite PanFlora website if appropriate: http://www.gilnelson.com/PanFlora/ -->
It has been observed flowering April through July and fruiting January, May and December.<ref name="FSU Herbarium"/>
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''O. humifusa'' has been observed flowering April through July with peak inflorescence in May and fruiting January, May, and December.<ref name="FSU Herbarium"/><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: 12 DEC 2016</ref>
  
Root stocks and detached cladodes can propagate vegetatively for up to 12 months after detachment. <ref name=gold/>
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Root stocks and detached cladodes can propagate vegetatively for up to 12 months after detachment.<ref name=gold/>
  
 
===Seed dispersal===
 
===Seed dispersal===
The fruit is eaten and dispersed by birds, rabbits, woodrats, prairie-dogs, mice, ground squirrels, and white-tailed deer. <ref name="fs">[[http://www.fs.fed.us/database/feis/plants/cactus/opuhum/all.html]] Accessed: February 13, 2016</ref> According to Kay Kirkman, a plant ecologist, this species disperses by being consumed by vertebrates (being assumed). <ref name="KK"> Kay Kirkman, unpublished data, 2015. </ref>
+
This species is thought to be dispersed by consumption by vertebrates.<ref> Kirkman, L. Katherine. Unpublished database of seed dispersal mode of plants found in Coastal Plain longleaf pine-grasslands of the Jones Ecological Research Center, Georgia.</ref> The fruit is eaten and dispersed by birds, rabbits, woodrats, prairie-dogs, mice, ground squirrels, and white-tailed deer.<ref name="fs">[[http://www.fs.fed.us/database/feis/plants/cactus/opuhum/all.html]] Accessed: February 13, 2016</ref>
  
 
===Seed bank and germination===
 
===Seed bank and germination===
Germination rate is low.<ref name="fs"/>
+
The germination rate is low.<ref name="fs"/> ''O. humifusa'' is known to occur in above-ground vegetation and in the fall surveyed seed bank in the southern ridge sandhill regions in south-central Florida.<ref>Lang, N. L. 2011. Soil seed bank dynamics of southern Lake Wales Ridge sandhill communities. Archbold Biological Station, Venus, Florida,15 pg. </ref>
  
<!--===Fire ecology===--> <!--Fire tolerance, fire dependence, adaptive fire responses-->
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===Fire ecology=== <!--Fire tolerance, fire dependence, adaptive fire responses-->
 +
O. humifusa will resprout and recruit seedlings during post-fire recovery.<ref>Lang, N. L. 2011. Soil seed bank dynamics of southern Lake Wales Ridge sandhill communities. Archbold Biological Station, Venus, Florida,15 pg. </ref>
  
 
===Pollination===
 
===Pollination===
The following Hymenoptera families and species were observed visiting flowers of ''Opuntia humifusa'' at Archbold Biological Station: <ref name="Deyrup 2015">Deyrup, M.A. and N.D. 2015. Database of observations of Hymenoptera visitations to flowers of plants on Archbold Biological Station, Florida, USA.</ref>
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''Opuntia humifusa'' has been observed at the Archbold Biological Station to host bees from the Apidae family such as  ''Apis mellifera, Bombus impatiens, B. pennsylvanicus'' and ''Mellisodes communis'', sweat bees from the Halictidae family such as ''Agapostemon splendens, Augochlorella aurata, Augochloropsis sumptuosa, Halictus poeyi, Lasioglossum nymphalis,'' and ''L. puteulanum'', and leafcutting bees from the family Megachilidae such as ''Dianthidium floridiense, Lithurgus gibbosus, Megachile brevis pseudobrevis'' and ''M. policaris''.<ref name="Deyrup 2015">Deyrup, M.A. and N.D. 2015. Database of observations of Hymenoptera visitations to flowers of plants on Archbold Biological Station, Florida, USA.</ref> Additionally, ''O. humifusa'' has been observed to host sweat bees from the Halictidae family such as ''Agapostemon virescens, Lasioglossum leucocomum, L. pruinosum'' and ''L. semicaeruleum'', true bugs such as ''Myodocha serripes'' (family Lygaeidae), and leafcutting bugs from the Megachilidae family such as ''Megachile addenda, Megachile centuncularis'' and ''Osmia texana''.<ref>Discoverlife.org [https://www.discoverlife.org/20/q?search=Bidens+albaDiscoverlife.org|Discoverlife.org]</ref>
 
 
Apidae:  ''Apis mellifera, Bombus impatiens, B. pennsylvanicus, Mellisodes communis''
 
 
 
Halictidae''Agapostemon splendens, Augochlorella aurata, Augochloropsis sumptuosa, Halictus poeyi, Lasioglossum nymphalis, L. puteulanum''
 
 
 
Megachilidae:  ''Dianthidium floridiense, Lithurgus gibbosus, Megachile brevis pseudobrevis, M. policaris''
 
  
===Use by animals=== <!--Herbivory, granivory, insect hosting, etc.-->
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===Herbivory and toxicology===
It is an important food for pocket gophers. <ref name=myers> Myers, G. T. and T. A. Vaughan (1964). "Food Habits of the Plains Pocket Gopher in Eastern Colorado." Journal of Mammalogy 45(4): 588-598.</ref> It is a host to the invasive cactus moth (''Cactoblastis cactorum''). <ref name=jez> Jezorek, H. and P. Stiling (2012). "LACK OF ASSOCIATIONAL EFFECTS BETWEEN TWO HOSTS OF AN INVASIVE HERBIVORE: OPUNTIA SPP. AND CACTOBLASTIS CACTORUM (LEPIDOPTERA: PYRALIDAE)." The Florida Entomologist 95(4): 1048-1057. </ref> The large pads provides nesting to bobwhite quail. <ref name="illinois">[[http://www.illinoiswildflowers.info/prairie/plantx/prickly_pearx.htm]]Illinois Wildflowers. Accessed: February 15, 2016</ref>
+
It is an important food for pocket gophers.<ref name=myers> Myers, G. T. and T. A. Vaughan (1964). "Food Habits of the Plains Pocket Gopher in Eastern Colorado." Journal of Mammalogy 45(4): 588-598.</ref> It is a host to the invasive cactus moth (''Cactoblastis cactorum'').<ref name=jez> Jezorek, H. and P. Stiling (2012). "LACK OF ASSOCIATIONAL EFFECTS BETWEEN TWO HOSTS OF AN INVASIVE HERBIVORE: OPUNTIA SPP. AND CACTOBLASTIS CACTORUM (LEPIDOPTERA: PYRALIDAE)." The Florida Entomologist 95(4): 1048-1057. </ref> The large pads provides nesting to bobwhite quail.<ref name="illinois">[[http://www.illinoiswildflowers.info/prairie/plantx/prickly_pearx.htm]]Illinois Wildflowers. Accessed: February 15, 2016</ref>
 
<!--===Diseases and parasites===-->
 
<!--===Diseases and parasites===-->
  
==Conservation and management==
+
==Conservation, cultivation, and restoration==
  
==Cultivation and restoration==
+
==Cultural use==
Native Americans used the pads to heal wounds, warts, and drank pad tea for respiratory issues.<ref name="wildflower"/>
+
Native Americans used the pads to heal wounds, warts, and drank pad tea for respiratory issues.<ref name="wildflower"/> It was also used as a poultice to stimulate milk supply in nursing mothers. The roots were used to stimulate hair growth and boiled with milk to treat dysentery, and a tea of the flowers could be used for urinary issues. A tea from the stem could be used in treating headaches, eyes problems, and insomnia.<ref> Korchmal, Arnold & Connie. 1973. A Guide to the Medicinal Plants of the United States. The New York Times Book Company, New York.</ref>
  
 
==Photo Gallery==
 
==Photo Gallery==

Latest revision as of 09:13, 3 July 2024

Opuntia humifusa
Opun humi.jpg
Photo by Wayne Matchett, SpaceCoastWildflowers.com
Scientific classification
Kingdom: Plantae
Division: Magnoliophyta - Flowering plants
Class: Magnoliopsida – Dicotyledons
Order: Caryophyllales
Family: Cactaceae
Genus: Opuntia
Species: O. humifusa
Binomial name
Opuntia humifusa
(Raf.) Raf.
Opun humi dist.jpg
Natural range of Opuntia humifusa from USDA NRCS Plants Database.

Common names: Devil's-tongue; Eastern prickly pear[1]

Taxonomic notes

Synonyms: Opuntia compressa (Salisbury) J.F. Macbride var. compressa; O. impedita Small; O. macrarthra Gibbes; O. opuntia (Linnaeus) Karten; O. calcicola Wherry; O. rafinesquei Engelmann.[1]

Varieties: none.[1]

Description

A description of Opuntia humifusa is provided in The Flora of North America.

The leaves of Opuntia humifusa are 1.5-6 cm long, with the longest leaves having an acute to acuminate tip. The hairs of the leaf surfaces are similarly long and the leaf sheaths and culm axis are glabrate to pilose with hairs shorter than 1.5 mm. The awns are glabrous and the lemma is 9-11 veined.[1]

The root system of Opuntia humifusa includes stem tubers which store non-structural carbohydrates (NSC) important for both resprouting following fire and persisting during long periods of fire exclusion.[2] Diaz-Toribio and Putz (2021) recorded this species to have an NSC concentration of 159.7 mg/g (ranking 32 out of 100 species studied) and water content of 47.2% (ranking 3 out of 100 species studied).[2]

According to Diaz-Torbio and Putz (2021), Opuntia humifusa has stem tubers with a below-ground to above-ground biomass ratio of 0.606 and nonstructural carbohydrate concentration of 159.7 mg g-1.[3]

Distribution

It is the only cactus to be widespread in the eastern United States.[4] It is mainly restricted to the Appalachian Mountains, in states like Deleware, Maryland, New Jersey, Virginia, and West Virginia, but also occurs in central and northcentral Mississippi. Further surveying could reveal populations in Alabama, northwestern Georgia, western South Carolina, western North Carolina, and northeastern Tennessee.[1]

Ecology

Habitat

Opuntia humifusa can occur in upland longleaf pines, sand dunes, Pinus clausa scrubs, open sand flats, sandhills, and pine/oak scrubs. It has been found in disturbed areas such as loblolly tree farms and roadside depressions. Associated species include Lyonia ferruginea, Lyonia lucida, Serenoa repens, Quercus geminata, Q. chapmanii, Persea humilis, Ceratiola, Osmanthus megacarpus, Rhynchospora megalocarpa, Galactia elliottii, and Smilax auriculata.[5]

O. humifusa was unaffected by soil disturbance by backe seeding, high-severity burning, and salvage logging in central Florida sand pine scrub.[6]

It had mixed responses to clearcutting in central Florida. In some cases the plant increased its cover and in other areas it was unaffected by the practice. It has shown regrowth and been unaffected in reestablished native sand pine scrub that was disturbed by clearcutting.[6]

This species increased its cover in response to soil disturbance by single roller chopping and broadcast seeding in central Florida pine scrub habitat. It has shown regrowth in reestablished native habitat that was disturbed by these practices.[6]

Additionally, O. humifusa reduced its frequency and density in response to soil disturbance by roller chopping in northwest Florida sandhills. It has shown resistance to regrowth and been unaffected by reestablished native sandhill habitat that was disturbed by this practice.[7]

It occurs in southern Canada, where average nighttime temperatures can reach -4 degrees Celsius. In order to prevent intracellular freeze dehydration and ice formation, individuals have an accumulation of sugars and mannitol in their cells.[8]

Phenology

O. humifusa has been observed flowering April through July with peak inflorescence in May and fruiting January, May, and December.[5][9]

Root stocks and detached cladodes can propagate vegetatively for up to 12 months after detachment.[8]

Seed dispersal

This species is thought to be dispersed by consumption by vertebrates.[10] The fruit is eaten and dispersed by birds, rabbits, woodrats, prairie-dogs, mice, ground squirrels, and white-tailed deer.[11]

Seed bank and germination

The germination rate is low.[11] O. humifusa is known to occur in above-ground vegetation and in the fall surveyed seed bank in the southern ridge sandhill regions in south-central Florida.[12]

Fire ecology

O. humifusa will resprout and recruit seedlings during post-fire recovery.[13]

Pollination

Opuntia humifusa has been observed at the Archbold Biological Station to host bees from the Apidae family such as Apis mellifera, Bombus impatiens, B. pennsylvanicus and Mellisodes communis, sweat bees from the Halictidae family such as Agapostemon splendens, Augochlorella aurata, Augochloropsis sumptuosa, Halictus poeyi, Lasioglossum nymphalis, and L. puteulanum, and leafcutting bees from the family Megachilidae such as Dianthidium floridiense, Lithurgus gibbosus, Megachile brevis pseudobrevis and M. policaris.[14] Additionally, O. humifusa has been observed to host sweat bees from the Halictidae family such as Agapostemon virescens, Lasioglossum leucocomum, L. pruinosum and L. semicaeruleum, true bugs such as Myodocha serripes (family Lygaeidae), and leafcutting bugs from the Megachilidae family such as Megachile addenda, Megachile centuncularis and Osmia texana.[15]

Herbivory and toxicology

It is an important food for pocket gophers.[16] It is a host to the invasive cactus moth (Cactoblastis cactorum).[17] The large pads provides nesting to bobwhite quail.[18]

Conservation, cultivation, and restoration

Cultural use

Native Americans used the pads to heal wounds, warts, and drank pad tea for respiratory issues.[4] It was also used as a poultice to stimulate milk supply in nursing mothers. The roots were used to stimulate hair growth and boiled with milk to treat dysentery, and a tea of the flowers could be used for urinary issues. A tea from the stem could be used in treating headaches, eyes problems, and insomnia.[19]

Photo Gallery

References and notes

  1. 1.0 1.1 1.2 1.3 1.4 Weakley, A.S. 2015. Flora of the southern and mid-atlantic states. Working Draft of 21 May 2015. University of North Carolina at Chapel Hill, Chapel Hill, North Carolina.
  2. 2.0 2.1 Diaz-Toribio, M.H. and F. E. Putz 2021. Underground carbohydrate stores and storage organs in fire-maintained longleaf pine savannas in Florida, USA. American Journal of Botany 108: 432-442.
  3. Diaz‐Toribio, M. H. and F. E. Putz. 2021. Underground carbohydrate stores and storage organs in fire‐maintained longleaf pine savannas in Florida, USA. American Journal of Botany 108(3):432-442.
  4. 4.0 4.1 [[1]] Lady Bird Johnson Wildflower Center. Accessed: February 12, 2016
  5. 5.0 5.1 Florida State University Robert K. Godfrey Herbarium database. URL: http://herbarium.bio.fsu.edu. Last accessed: October 2015. Collectors: Loran C. Anderson, Michael Blaker, M. Borgman, James R. Burkhaulter, George R. Cooley, R.J. Eaton, Patricia Elliott, J. Kevin England, Robert K. Godfrey, Darren Jackson, Ed Keppner, Lisa Keppner, Andrew McAllister, Sidney McDaniel, K.M. Meyer, James D. Ray Jr., Erik Robinson, L. Rosen, C.E. Smith, A. Townesmith, Kenneth A. Wilson, Carroll E. Wood Jr.. States and Counties: Alabama: Dale. Florida: Alabama, Escambia, Franklin, Gadsden, Hernando, Jackson, Lafayette, Leon, Liberty, Orange, Putnam, Seminole, Walton, Wakulla, Washington. Compiled by Tall Timbers Research Station and Land Conservancy.
  6. 6.0 6.1 6.2 Greenberg, C.H., D.G. Neary, L.D. Harris, and S.P. Linda. (1995). Vegetation Recovery Following High-intensity Wildfire and Silvicultural Treatments in Sand Pine Scrub. American Midland Naturalist 133(1):149-163.
  7. Hebb, E.A. (1971). Site Preparation Decreases Game Food Plants in Florida Sandhills. The Journal of Wildlife Management 35(1):155-162.
  8. 8.0 8.1 Cite error: Invalid <ref> tag; no text was provided for refs named gold
  9. 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: 12 DEC 2016
  10. Kirkman, L. Katherine. Unpublished database of seed dispersal mode of plants found in Coastal Plain longleaf pine-grasslands of the Jones Ecological Research Center, Georgia.
  11. 11.0 11.1 [[2]] Accessed: February 13, 2016
  12. Lang, N. L. 2011. Soil seed bank dynamics of southern Lake Wales Ridge sandhill communities. Archbold Biological Station, Venus, Florida,15 pg.
  13. Lang, N. L. 2011. Soil seed bank dynamics of southern Lake Wales Ridge sandhill communities. Archbold Biological Station, Venus, Florida,15 pg.
  14. Deyrup, M.A. and N.D. 2015. Database of observations of Hymenoptera visitations to flowers of plants on Archbold Biological Station, Florida, USA.
  15. Discoverlife.org [3]
  16. Myers, G. T. and T. A. Vaughan (1964). "Food Habits of the Plains Pocket Gopher in Eastern Colorado." Journal of Mammalogy 45(4): 588-598.
  17. Jezorek, H. and P. Stiling (2012). "LACK OF ASSOCIATIONAL EFFECTS BETWEEN TWO HOSTS OF AN INVASIVE HERBIVORE: OPUNTIA SPP. AND CACTOBLASTIS CACTORUM (LEPIDOPTERA: PYRALIDAE)." The Florida Entomologist 95(4): 1048-1057.
  18. [[4]]Illinois Wildflowers. Accessed: February 15, 2016
  19. Korchmal, Arnold & Connie. 1973. A Guide to the Medicinal Plants of the United States. The New York Times Book Company, New York.