Difference between revisions of "Centrosema virginianum"

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| range_map_caption = Natural range of ''Centrosema virginianum'' from USDA NRCS [http://www.plants.usda.gov/core/profile?symbol=CEVI2 Plants Database].
 
| range_map_caption = Natural range of ''Centrosema virginianum'' from USDA NRCS [http://www.plants.usda.gov/core/profile?symbol=CEVI2 Plants Database].
 
}}
 
}}
Common name: Spurred Butterfly Pea
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Common name: Spurred butterfly pea
 
==Taxonomic notes==
 
==Taxonomic notes==
Synonyms: ''Centrosema virginianum'' var. ''virginianum''; ''C. virginianum'' var. ''ellipticum'' Fernald; ''Bradburya virginiana'' (Linnaeus) Kuntze; ''C. virginianum'' var. ''angustifolium'' (A.P. de Candolle) Grisebach
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Synonyms: ''Bradburya virginiana'' (Linnaeus) Kuntze.<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>
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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==<!-- Basic life history facts such as annual/perrenial, monoecious/dioecious, root morphology, seed type, etc. -->  
 
==Description==<!-- Basic life history facts such as annual/perrenial, monoecious/dioecious, root morphology, seed type, etc. -->  
''Centrosema virginianum'' is a perennial herbaceous vine. It grows as a twining vine, climbing on Smilax vines and low vegetation, frequently appearing among grasses<ref name="fsu">Florida State University Robert K. Godfrey Herbarium database. URL: http://herbarium.bio.fsu.edu. Last accessed: June 2014. Collectors: Loran C. Anderson, John C. Ogden, Gwynn W. Ramsey, R.K. Godfrey, R. S. Mitchell; R. C. Phillips, K. Craddock Burks, Gary R. Knight, D. W. Mather, C. Jackson, D. B. Ward, Mary Margaret Williams, O. Lakela, Brenda Herring, Jame Amoroso, Gwynn W. Ramsey, Richard Mitchell, Gail A. Steverson, Grady W. Reinert, George R. Cooley, R. J. Eaton, R. Kral, Cecil R Slaughter, Andre F. Clewell, R. Komarek, R. F. Doren, Kevin Oakes, Richard Gaskalla, Lisa Keppner, Clarke Hudson, Wilbur H Duncan, Jean Wooten, H. R. Totten, R. L. Wilbur, C. Ritchie Bell, Delzie Demaree, F. S. Earle, A. B. Seymour, Samuel B. Jones, Jr., H. R. Reed, A. B. Seymour, Michael B. Brooks, Sidney McDaniel, D. C. Bain, D. S. Correll, H. B. Correll, Lloyd H. Shinners, Geo M. Merrill, and H J Hamby. States and Counties: Alabama: Baldwin. Arkansas: Little Rock. Florida: Bay, Citrus, Collier, Duval, Escambia, Franklin, Gadsden, Gulf, Hillsborough, Jackson, Jefferson, Leon, Liberty, Manatee, Marion, Okaloosa, Polk, St Johns, St. Lucie, Suwannee, Wakulla, and Washington. Georgia: Bartow, Grady, Madison, and Thomas. Mississippi: Forrest, Harrison, Jackson, Pearl River, and Pike. North Carolina: Alamance, Orange, and Wilkes. Texas: Angelina, Bastrop, Freestone, Harris, Morris, Tarrant, and Van Zandt.</ref>.
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''Centrosema virginianum'' is a perennial herbaceous vine. It grows as a twining vine, climbing on other vegetation.<ref name="fsu">Florida State University Robert K. Godfrey Herbarium database. URL: http://herbarium.bio.fsu.edu. Last accessed: June 2014. Collectors: Loran C. Anderson, John C. Ogden, Gwynn W. Ramsey, R.K. Godfrey, R. S. Mitchell; R. C. Phillips, K. Craddock Burks, Gary R. Knight, D. W. Mather, C. Jackson, D. B. Ward, Mary Margaret Williams, O. Lakela, Brenda Herring, Jame Amoroso, Gwynn W. Ramsey, Richard Mitchell, Gail A. Steverson, Grady W. Reinert, George R. Cooley, R. J. Eaton, R. Kral, Cecil R Slaughter, Andre F. Clewell, R. Komarek, R. F. Doren, Kevin Oakes, Richard Gaskalla, Lisa Keppner, Clarke Hudson, Wilbur H Duncan, Jean Wooten, H. R. Totten, R. L. Wilbur, C. Ritchie Bell, Delzie Demaree, F. S. Earle, A. B. Seymour, Samuel B. Jones, Jr., H. R. Reed, A. B. Seymour, Michael B. Brooks, Sidney McDaniel, D. C. Bain, D. S. Correll, H. B. Correll, Lloyd H. Shinners, Geo M. Merrill, and H J Hamby. States and Counties: Alabama: Baldwin. Arkansas: Little Rock. Florida: Bay, Citrus, Collier, Duval, Escambia, Franklin, Gadsden, Gulf, Hillsborough, Jackson, Jefferson, Leon, Liberty, Manatee, Marion, Okaloosa, Polk, St Johns, St. Lucie, Suwannee, Wakulla, and Washington. Georgia: Bartow, Grady, Madison, and Thomas. Mississippi: Forrest, Harrison, Jackson, Pearl River, and Pike. North Carolina: Alamance, Orange, and Wilkes. Texas: Angelina, Bastrop, Freestone, Harris, Morris, Tarrant, and Van Zandt.</ref> The leaflet langth to width ratio is very variable.<ref name= "Weakley">Weakley, A. S. (2015). Flora of the Southern and Mid-Atlantic States. Chapel Hill, NC, University of North Carolina Herbarium.</ref>
  
''Centrosema virginianum'' is seen growing in a tailing, climbing, and twining fashion. The plant is a perennial herbaceous vine that grows 0.5-1.5 m long, and is more or less minutely pubescent throughout. The leaves are 3-foliolate; the leaflets are widely to narrowly ovate, ovate-lanceolate or oblong to elliptic, are conspicously reticulate, mostly growing 2-7 cm long, and are stipellate. The stipules are ovate-lanceolate to lanceolate, striate, persistent, 1.5-4 mm long. The racemes are with peduncles usually growing 1-5 cm long, the zig-zag rachis bears 1-4 nodes each with an ovate bract growing up to 1.5-3 cm long subtending a pedicel (growing 2-10 cm long) surmounted by 2 ovate bractlets growing 0.8-1.2 cm long. The calyx is somewhat hidden by the braclets, the tube is broadly hemispheric, growing 4-5 mm long, the lobes are linear-subulate, growing 0.6-1.4 cm long, the lowermost longest. The petals are pale blue-violet to lavender in color, growing 2.5-3.5 cm long, is spurred near the base, the wigs and keel are nearly equal in size, ca. 2 cm long. The stamens are diadelphous, 9 and 1. The legume is linear, flattened, and grows 7-14 cm long and ca. 4 mm broad, is subsessile, many seeded with an elongate, persistent, beak-like style, valves longitudinally twisting after dehiscence<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. 635-6. Print</ref>.
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''Centrosema virginianum'' is seen growing in a tailing, climbing, and twining fashion. The plant is a perennial herbaceous vine that grows 0.5-1.5 m long, and is more or less minutely pubescent throughout. The leaves are 3-foliolate; the leaflets are widely to narrowly ovate, ovate-lanceolate or oblong to elliptic, are conspicously reticulate, mostly growing 2-7 cm long, and are stipellate. The stipules are ovate-lanceolate to lanceolate, striate, persistent, 1.5-4 mm long. The racemes are with peduncles usually growing 1-5 cm long, the zig-zag rachis bears 1-4 nodes each with an ovate bract growing up to 1.5-3 cm long subtending a pedicel (growing 2-10 cm long) surmounted by 2 ovate bractlets growing 0.8-1.2 cm long. The calyx is somewhat hidden by the braclets, the tube is broadly hemispheric, growing 4-5 mm long, the lobes are linear-subulate, growing 0.6-1.4 cm long, the lowermost longest. The petals are pale blue-violet to lavender in color, growing 2.5-3.5 cm long, is spurred near the base, the wigs and keel are nearly equal in size, ca. 2 cm long. The stamens are diadelphous, 9 and 1. The legume is linear, flattened, and grows 7-14 cm long and ca. 4 mm broad, is subsessile, many seeded with an elongate, persistent, beak-like style, valves longitudinally twisting after dehiscence.<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. 635-6. Print</ref>
  
 
==Distribution==
 
==Distribution==
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''C. virginianum'' is native to the southeast United States, north up to Illinois and New Jersey, along the east coast and south to Florida, and west to Texas and Oklahoma. It is also native to Puerto Rico as well as the Virgin Islands.<ref name= "USDA">USDA, NRCS. (2016). The PLANTS Database (http://plants.usda.gov, 4 April 2019). National Plant Data Team, Greensboro, NC 27401-4901 USA.</ref>
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==Ecology==
 
==Ecology==
It is a legume that has one of the highest nitrogen-fixing potentials<ref name="cathey">Cathey, S. E., L. R. Boring, et al. (2010). "Assessment of N2 fixation capability of native legumes from the longleaf pine-wiregrass ecosystem." Environmental and Experimental Botany 67: 444-450.</ref>. Because of this, it may be able to help restore N lost from fire<ref name="hainds">Hainds, M. J., R. J. Mitchell, et al. (1999). "Distribution of native legumes (Leguminoseae) in frequently burned longleaf pine (Pinaceae)-wiregrass (Poaceae) ecosystems." American Journal of Botany 86: 1606-1614.</ref>. By mid-season in June and July, a maximum nitrogen-fixing rate was observed<ref name="cathey"/>.
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It is a legume that has one of the highest nitrogen-fixing potentials.<ref name="cathey">Cathey, S. E., L. R. Boring, et al. (2010). "Assessment of N2 fixation capability of native legumes from the longleaf pine-wiregrass ecosystem." Environmental and Experimental Botany 67: 444-450.</ref> Because of this, it may be able to help restore N lost from fire.<ref name="hainds">Hainds, M. J., R. J. Mitchell, et al. (1999). "Distribution of native legumes (Leguminoseae) in frequently burned longleaf pine (Pinaceae)-wiregrass (Poaceae) ecosystems." American Journal of Botany 86: 1606-1614.</ref> By mid-season in June and July, a maximum nitrogen-fixing rate was observed.<ref name="cathey"/>
 
===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.-->  
It is found in a wide range of natural and disturbed conditions, including frequently burned sandhills, upland longleaf-wiregrass and old-field pinelands<ref name="hainds"/><ref name="cushwa">Cushwa, C. T. (1966). The response of herbaceous vegetation to prescribed burning. Asheville, USDA Forest Service.</ref> and flatwoods, coastal island dunes and shorelines, open areas within mangrove swamps, wooded floodplains and edges of hardwood forests, and bogs .  It can be found in loblolly pine communities<ref name="cushwa"/>. It can also be found in longleaf pine-wiregrass communities<ref name="hainds"/>.
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It is found in a wide range of natural and disturbed conditions, including frequently burned sandhills, upland longleaf-wiregrass and old-field pinelands<ref name="hainds"/><ref name="cushwa">Cushwa, C. T. (1966). The response of herbaceous vegetation to prescribed burning. Asheville, USDA Forest Service.</ref> and flatwoods, coastal island dunes and shorelines, open areas within mangrove swamps, wooded floodplains and edges of hardwood forests, and bogs .  It can be found in loblolly pine communities.<ref name="cushwa"/> It can also be found in longleaf pine-wiregrass communities.<ref name="hainds"/> As well, ''C. virginianum'' is a characteristic species of the shortleaf pine-oak-hickory community.<ref name= "Clewell">Clewell, A. F. (2013). "Prior prevalence of shortleaf pine-oak-hickory woodlands in the Tallahassee red hills." Castanea 78(4): 266-276.</ref> It is tolerant of overstory canopies that decrease the light level to about half the ambient (i.e., it can live in partially shaded areas and its nitrogen-fixing capability won't be significantly affected).<ref name="cathey"/> It grows in highly disturbed areas, but it is also ubiquitous in high quality native longleaf pine uplands and sandhills. It occurs in soils ranging from deep sands (Entisols) to sandy loams (Ultisols). ''C. virginianum'' responds positively to soil disturbance in coastal plains of South Carolina,<ref>Brudvig, L.A. and E.I. Damchen. (2011). Land-use history, historical connectivity, and land management interact to determine longleaf pine woodland understory richness and composition. Ecography 34: 257-266.</ref> including agriculture-based disturbance, marking it as an indicator species of post-agricultural woodlands.<ref>Brudvig, L.A., E Grman, C.W. Habeck, and J.A. Ledvina. (2013). Strong legacy of agricultural land use on soils and understory plant communities in longleaf pine woodlands. Forest Ecology and Management 310: 944-955.</ref> It does not respond to soil disturbance by clearcutting and chopping in North Florida flatwoods forests.<ref>Moore, W.H., B.F. Swindel, and W.S. Terry. (1982). Vegetative Response to Clearcutting and Chopping in a North Florida Flatwoods Forest. Journal of Range Management 35(2):214-218.</ref>
It is tolerant of overstory canopies that decrease the light level to about half the ambient (i.e., it can live in partially shaded areas and its nitrogen-fixing capability won't be significantly affected)<ref name="cathey"/>. It grows in highly disturbed areas, but it is also ubiquitous in high quality native longleaf pine uplands and sandhills.
 
It occurs in soils ranging from deep sands (Entisols) to sandy loams (Ultisols).
 
  
Associated species includes Blackberry Bramble, turkey oak, longleaf pine, ''Galactia, Strophostyles, Smilax, Penstemon, Lechea, Chrysopsis, Brumelia, Centrosema, Euphorbia, Cassia, Serenoa repens, Quercus incana, Quercus chapmanii, Diospyros, Aristida, Andropogon,'' bahia grass, ''Rubus,'' cloverleaf, ''Pinus taeda, Liquidambar styraciflua'', and others<ref name="fsu"/>.
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Associated species includes Blackberry Bramble, turkey oak, longleaf pine, ''Galactia, Strophostyles, Smilax, Penstemon, Lechea, Chrysopsis, Brumelia, Centrosema, Euphorbia, Cassia, Serenoa repens, Quercus incana, Quercus chapmanii, Diospyros, Aristida, Andropogon,'' bahia grass, ''Rubus,'' cloverleaf, ''Pinus taeda, Liquidambar styraciflua'', 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/ -->  
 
===Phenology===<!--Timing off flowering, fruiting, seed dispersal, and environmental triggers.  Cite PanFlora website if appropriate: http://www.gilnelson.com/PanFlora/ -->  
It flowers in April through October and fruits primarily in June thorugh September<ref name="fsu"/>.  
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''C. virginianum'' has been observed flowering between April and October, with peak  inflorescence in June and July, and fruits primarily in June thorugh September.<ref name="fsu"/><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>
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===Seed dispersal===
 
===Seed dispersal===
According to Kay Kirkman, a plant ecologist, this species disperses by explosion mechanisms or by ants. <ref name="KK"> Kay Kirkman, unpublished data, 2015. </ref>
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This species is thought to be dispersed by ants and/or explosive dehiscence. <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>
  
 
===Seed bank and germination===
 
===Seed bank and germination===
In Hier's study, changes in flowering did not seem to affect seed germination<ref name="hiers2000">Hiers, J. K., R. Wyatt, et al. (2000). "The effects of fire regime on legume reproduction in longleaf pine savannas: is a season selective?" Oecologia 125: 521-530.</ref>. It spreads clonally by production of rhizomes<ref name="hiers2007">Hiers, J. K. and R. J. Mitchell (2007). "The influence of burning and light availability on N-2-fixation of native legumes in longleaf pine woodlands." Journal of the Torrey Botanical Society 134: 398-409.</ref>.
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It spreads clonally by production of rhizomes.<ref name="hiers2007">Hiers, J. K. and R. J. Mitchell (2007). "The influence of burning and light availability on N-2-fixation of native legumes in longleaf pine woodlands." Journal of the Torrey Botanical Society 134: 398-409.</ref> Seed coats are hard and and seeds and remain viable in the seed bank for at least two years.<ref> Coffey, K. L. and L. K. Kirkman (2006). "Seed germination strategies of species with restoration potential in a fire-maintained pine savanna." Natural Areas Journal 26: 289-299. </ref> For optimal germination, ''C. virginianum'' needs about 2 seconds of scarification time. It was also found to have greatest dry heat germination at a heat index between 128.76 and 191.83, and steam duration of 10 seconds.<ref name= "Wiggers">Wiggers, M. S., et al. (2017). "Seed heat tolerance and germination of six legume species native to a fire-prone longleaf pine forest." Plant Ecology 218: 151-171.</ref> With a study on the effects of variation in fine fuel loads on post-burn germination, it was found to have greater germination in low fuel than high fuel with a lower mortality rate as well, although the greatest rate of germination and lowest mortality was with the control of no burn regiment.<ref name= "Wiggers2">Wiggers, M. S., et al. (2013). "Fine-scale variation in surface fire environment and legume germination in the longleaf pine ecosystem." Forest Ecology and Management 310: 54-63.</ref>
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===Fire ecology===<!--Fire tolerance, fire dependence, adaptive fire responses-->  
 
===Fire ecology===<!--Fire tolerance, fire dependence, adaptive fire responses-->  
It thrives under fire<ref name="cushwa"/>. Hendricks observed that the Piedmont National Wildlife Refuge plots, which had been under a 4-year burning regime since 1966, each contained more than 10 times more ''C. virginianum'' individuals per ha than the Oconee National Forest plots, which had no burning history<ref name="hendricks">Hendricks, J. J. and L. R. Boring (1999). "N2-fixation by native herbaceous legumes in burned pine ecosystems of the southeastern United States." Forest Ecology and Management 113: 167-177.</ref>. Seasonal burning does not seem to negatively affect nitrogen fixation<ref name="hiers2003">Hiers, J. K., R. J. Mitchell, et al. (2003). "Legumes native to longleaf pine savannas exhibit capacity for high N2-fixation rates and negligible impacts due to timing of fire." New Phytologist 157: 327-338</ref>. ''C. virginianum'' showed increased flowering synchrony in response to lightning-season burns<ref name="hiers2000"/>. It responded the best to March burns with respect to annual tissue inputs as well as nitrogen contribution<ref name="hiers2003"/>. ''C. virginianum'' showed robust flowering response to late winter/ early spring burns, supporting the response to March burns noted earlier. It has a mid-summer flowering peak<ref name="hiers2000"/>. Also, Hiers found no evidence that increased flowering affects nitrogen-fixing capability<ref name="hiers2003"/>.
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It thrives under fire.<ref name="cushwa"/> Hendricks observed that the Piedmont National Wildlife Refuge plots, which had been under a 4-year burning regime since 1966, each contained more than 10 times more ''C. virginianum'' individuals per ha than the Oconee National Forest plots, which had no burning history.<ref name="hendricks">Hendricks, J. J. and L. R. Boring (1999). "N2-fixation by native herbaceous legumes in burned pine ecosystems of the southeastern United States." Forest Ecology and Management 113: 167-177.</ref> Seasonal burning does not seem to negatively affect nitrogen fixation.<ref name="hiers2003">Hiers, J. K., R. J. Mitchell, et al. (2003). "Legumes native to longleaf pine savannas exhibit capacity for high N2-fixation rates and negligible impacts due to timing of fire." New Phytologist 157: 327-338</ref> ''C. virginianum'' showed increased flowering synchrony in response to lightning-season burns.<ref name="hiers2003"/> It responded the best to March burns with respect to annual tissue inputs as well as nitrogen contribution.<ref name="hiers2003"/> ''C. virginianum'' showed robust flowering response to late winter/ early spring burns, supporting the response to March burns noted earlier. It has a mid-summer flowering peak.<ref name="hiers2003"/> Also, one study found no evidence that increased flowering affects nitrogen-fixing capability.<ref name="hiers2003"/>
  
 
===Pollination===  
 
===Pollination===  
Its flower is highly specialized for pollination by large Hymenoptera (Spears 1987 cited by Hiers et al 2007). It requires bees for pollination to "trip" the pollen delivery mechanism. Pollinator-plant relationships appear to be robust to alteration in flowering phenology resulting from variation in season of burn (Hiers et al 2000).
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Its flower is highly specialized for pollination by large Hymenoptera.<ref name="spears">Spears, Jr. E. E. 1987. Island and mainland pollination ecology of ''Centrosema virginianum'' and ''Opuntia stricta''. J. Ecol. 75: 351-362.</ref> It requires bees for pollination to "trip" the pollen delivery mechanism. Pollinator-plant relationships appear to be robust to alteration in flowering phenology resulting from variation in season of burn.<ref name="hiers2003"/>''Bombus pennsylvanicus'' was observed feeding and collecting pollen.<ref>Godts J.E. 1990 The Upside-Down Flower Palmetto 10(4):3</ref>
  
 
===Use by animals===<!--Herbivory, granivory, insect hosting, etc.-->  
 
===Use by animals===<!--Herbivory, granivory, insect hosting, etc.-->  
Because ''C. virginianum'' is a legume, and legumes are high in protein and mineral content, a number of herbivores including but not limited to ''Gopherus polyphemus'', white-tailed deer, and bob-white quail, consume it (Hainds et al 1999).
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Because ''C. virginianum'' is a legume, and legumes are high in protein and mineral content, a number of herbivores including but not limited to ''Gopherus polyphemus'', white-tailed deer, and bob-white quail, consume it.<ref name="hainds"/> One study found that it is a significantly important plant in 1-year and 2-years stands for the bobwhite quail diet.<ref name= "Sweeney">Sweeney, J. M., et al. (1981). Bobwhite quail food in young Arkansas loblolly pine plantations. Arkansas Experiment Station bulletin 852. Fayetteville, AR, University of Arkansas, Divisionn of Agriculture, Agricultural Experiment Station.</ref> It averages to be about 10-25% of the diet for large mammals and terrestrial birds.<ref name= "Miller">Miller, J.H., and K.V. Miller. 1999. Forest plants of the southeast and their wildlife uses. Southern Weed Science Society.</ref>
  
<!--===Diseases and parasites===-->
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===Diseases and parasites===
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''C. virginianum'' can be infected by the root-knot nematode species ''Meloidogyne arenaria'', ''M. incognita'', and ''M. javanica'', but it is moderately resistant.<ref name= "Quesenberry">Quesenberry, K. H., et al. (2008). "Response of native southeastern U.S. legumes to root-knot nematodes." Crop Science 48: 2274-2278.</ref>
 +
 
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==Conservation and management==
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It is listed as endangered by the New Jersey Department of Environmental Protection and Energy.<ref name= "USDA"/> ''C. virginianum'' is tolerant to the herbicide imazapyr.<ref name= "proceedings">(2000). The role of fire in nongame wildlife management and community restoration: Traditional uses and new directions, Nashville, TN, USDA Forest Service, Northeastern Research Station.</ref> Also for management, it benefits from the overstory canopy being thinned.<ref name= "Brockway">Brockway, D. G. and C. E. Lewis (2003). "Influence of deer, cattle grazing and timber harvest on plant species diversity in a longleaf pine bluestem ecosystem." Forest Ecology and Management 175: 49-69.</ref>
  
==Conservation and Management==
 
 
==Cultivation and restoration==
 
==Cultivation and restoration==
 
==Photo Gallery==
 
==Photo Gallery==
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==References and notes==
 
==References and notes==
Cathey, S. E., L. R. Boring, et al. (2010). "Assessment of N2 fixation capability of native legumes from the longleaf pine-wiregrass ecosystem." Environmental and Experimental Botany 67: 444-450.
 
 
Cushwa, C. T. (1966). The response of herbaceous vegetation to prescribed burning. Asheville, USDA Forest Service.
 
 
Florida State University Robert K. Godfrey Herbarium database. URL: [http://herbarium.bio.fsu.edu http://herbarium.bio.fsu.edu]. Last accessed: June 2014.  Collectors: Loran C. Anderson, John C. Ogden, Gwynn W. Ramsey, R.K. Godfrey, R. S. Mitchell; R. C. Phillips, K. Craddock Burks, Gary R. Knight, D. W. Mather, C. Jackson, D. B. Ward, Mary Margaret Williams, O. Lakela, Brenda Herring, Jame Amoroso, Gwynn W. Ramsey, Richard Mitchell, Gail A. Steverson, Grady W. Reinert, George R. Cooley, R. J. Eaton, R. Kral, Cecil R Slaughter, Andre F. Clewell, R. Komarek, R. F. Doren, Kevin Oakes, Richard Gaskalla, Lisa Keppner, Clarke Hudson, Wilbur H Duncan, Jean Wooten, H. R. Totten, R. L. Wilbur, C. Ritchie Bell, Delzie Demaree, F. S. Earle, A. B. Seymour, Samuel B. Jones, Jr., H. R. Reed, A. B. Seymour, Michael B. Brooks, Sidney McDaniel, D. C. Bain, D. S. Correll, H. B. Correll, Lloyd H. Shinners, Geo M. Merrill, and H J Hamby.  States and Counties: Alabama: Baldwin. Arkansas: Little Rock. Florida: Bay, Citrus, Collier, Duval, Escambia, Franklin, Gadsden, Gulf, Hillsborough, Jackson, Jefferson, Leon, Liberty, Manatee, Marion, Okaloosa, Polk, St Johns, St. Lucie, Suwannee, Wakulla, and Washington. Georgia: Bartow, Grady, Madison, and Thomas. Mississippi: Forrest, Harrison, Jackson, Pearl River, and Pike. North Carolina: Alamance, Orange, and Wilkes. Texas: Angelina, Bastrop, Freestone, Harris, Morris, Tarrant, and Van Zandt.
 
 
Hainds, M. J., R. J. Mitchell, et al. (1999). "Distribution of native legumes (Leguminoseae) in frequently burned longleaf pine (Pinaceae)-wiregrass (Poaceae) ecosystems." American Journal of Botany 86: 1606-1614.
 
 
Hendricks, J. J. and L. R. Boring (1999). "N2-fixation by native herbaceous legumes in burned pine ecosystems of the southeastern United States." Forest Ecology and Management 113: 167-177.
 
 
Hiers, J. K., R. Wyatt, et al. (2000). "The effects of fire regime on legume reproduction in longleaf pine savannas: is a season selective?" Oecologia 125: 521-530.
 
 
Hiers, J. K., R. J. Mitchell, et al. (2003). "Legumes native to longleaf pine savannas exhibit capacity for high N2-fixation rates and negligible impacts due to timing of fire." New Phytologist 157: 327-338.
 
 
Hiers, J. K. and R. J. Mitchell (2007). "The influence of burning and light availability on N-2-fixation of native legumes in longleaf pine woodlands." Journal of the Torrey Botanical Society 134: 398-409.
 
 
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. 635-6. Print.
 
 
Spears, Jr. E. E. 1987. Island and mainland pollination ecology of Centrosema virginianum and Opuntia s trie ta. J. Ecol. 75: 351-362.
 

Revision as of 08:51, 15 September 2020

Centrosema virginianum
Centrosema virginianum Gil.jpg
Photo by Gil Nelson
Scientific classification
Kingdom: Plantae
Division: Magnoliophyta - Flowering plants
Class: Magnoliopsida – Dicotyledons
Order: Fabales
Family: Fabaceae ⁄ Leguminosae
Genus: Centrosema
Species: C. virginianum
Binomial name
Centrosema virginianum
(L.) Benth.
CENT VIRG dist.jpg
Natural range of Centrosema virginianum from USDA NRCS Plants Database.

Common name: Spurred butterfly pea

Taxonomic notes

Synonyms: Bradburya virginiana (Linnaeus) Kuntze.[1]

Varieties: none.[1]

Description

Centrosema virginianum is a perennial herbaceous vine. It grows as a twining vine, climbing on other vegetation.[2] The leaflet langth to width ratio is very variable.[3]

Centrosema virginianum is seen growing in a tailing, climbing, and twining fashion. The plant is a perennial herbaceous vine that grows 0.5-1.5 m long, and is more or less minutely pubescent throughout. The leaves are 3-foliolate; the leaflets are widely to narrowly ovate, ovate-lanceolate or oblong to elliptic, are conspicously reticulate, mostly growing 2-7 cm long, and are stipellate. The stipules are ovate-lanceolate to lanceolate, striate, persistent, 1.5-4 mm long. The racemes are with peduncles usually growing 1-5 cm long, the zig-zag rachis bears 1-4 nodes each with an ovate bract growing up to 1.5-3 cm long subtending a pedicel (growing 2-10 cm long) surmounted by 2 ovate bractlets growing 0.8-1.2 cm long. The calyx is somewhat hidden by the braclets, the tube is broadly hemispheric, growing 4-5 mm long, the lobes are linear-subulate, growing 0.6-1.4 cm long, the lowermost longest. The petals are pale blue-violet to lavender in color, growing 2.5-3.5 cm long, is spurred near the base, the wigs and keel are nearly equal in size, ca. 2 cm long. The stamens are diadelphous, 9 and 1. The legume is linear, flattened, and grows 7-14 cm long and ca. 4 mm broad, is subsessile, many seeded with an elongate, persistent, beak-like style, valves longitudinally twisting after dehiscence.[4]

Distribution

C. virginianum is native to the southeast United States, north up to Illinois and New Jersey, along the east coast and south to Florida, and west to Texas and Oklahoma. It is also native to Puerto Rico as well as the Virgin Islands.[5]

Ecology

It is a legume that has one of the highest nitrogen-fixing potentials.[6] Because of this, it may be able to help restore N lost from fire.[7] By mid-season in June and July, a maximum nitrogen-fixing rate was observed.[6]

Habitat

It is found in a wide range of natural and disturbed conditions, including frequently burned sandhills, upland longleaf-wiregrass and old-field pinelands[7][8] and flatwoods, coastal island dunes and shorelines, open areas within mangrove swamps, wooded floodplains and edges of hardwood forests, and bogs . It can be found in loblolly pine communities.[8] It can also be found in longleaf pine-wiregrass communities.[7] As well, C. virginianum is a characteristic species of the shortleaf pine-oak-hickory community.[9] It is tolerant of overstory canopies that decrease the light level to about half the ambient (i.e., it can live in partially shaded areas and its nitrogen-fixing capability won't be significantly affected).[6] It grows in highly disturbed areas, but it is also ubiquitous in high quality native longleaf pine uplands and sandhills. It occurs in soils ranging from deep sands (Entisols) to sandy loams (Ultisols). C. virginianum responds positively to soil disturbance in coastal plains of South Carolina,[10] including agriculture-based disturbance, marking it as an indicator species of post-agricultural woodlands.[11] It does not respond to soil disturbance by clearcutting and chopping in North Florida flatwoods forests.[12]

Associated species includes Blackberry Bramble, turkey oak, longleaf pine, Galactia, Strophostyles, Smilax, Penstemon, Lechea, Chrysopsis, Brumelia, Centrosema, Euphorbia, Cassia, Serenoa repens, Quercus incana, Quercus chapmanii, Diospyros, Aristida, Andropogon, bahia grass, Rubus, cloverleaf, Pinus taeda, Liquidambar styraciflua, and others.[2]

Phenology

C. virginianum has been observed flowering between April and October, with peak inflorescence in June and July, and fruits primarily in June thorugh September.[2][13]

Seed dispersal

This species is thought to be dispersed by ants and/or explosive dehiscence. [14]

Seed bank and germination

It spreads clonally by production of rhizomes.[15] Seed coats are hard and and seeds and remain viable in the seed bank for at least two years.[16] For optimal germination, C. virginianum needs about 2 seconds of scarification time. It was also found to have greatest dry heat germination at a heat index between 128.76 and 191.83, and steam duration of 10 seconds.[17] With a study on the effects of variation in fine fuel loads on post-burn germination, it was found to have greater germination in low fuel than high fuel with a lower mortality rate as well, although the greatest rate of germination and lowest mortality was with the control of no burn regiment.[18]

Fire ecology

It thrives under fire.[8] Hendricks observed that the Piedmont National Wildlife Refuge plots, which had been under a 4-year burning regime since 1966, each contained more than 10 times more C. virginianum individuals per ha than the Oconee National Forest plots, which had no burning history.[19] Seasonal burning does not seem to negatively affect nitrogen fixation.[20] C. virginianum showed increased flowering synchrony in response to lightning-season burns.[20] It responded the best to March burns with respect to annual tissue inputs as well as nitrogen contribution.[20] C. virginianum showed robust flowering response to late winter/ early spring burns, supporting the response to March burns noted earlier. It has a mid-summer flowering peak.[20] Also, one study found no evidence that increased flowering affects nitrogen-fixing capability.[20]

Pollination

Its flower is highly specialized for pollination by large Hymenoptera.[21] It requires bees for pollination to "trip" the pollen delivery mechanism. Pollinator-plant relationships appear to be robust to alteration in flowering phenology resulting from variation in season of burn.[20]Bombus pennsylvanicus was observed feeding and collecting pollen.[22]

Use by animals

Because C. virginianum is a legume, and legumes are high in protein and mineral content, a number of herbivores including but not limited to Gopherus polyphemus, white-tailed deer, and bob-white quail, consume it.[7] One study found that it is a significantly important plant in 1-year and 2-years stands for the bobwhite quail diet.[23] It averages to be about 10-25% of the diet for large mammals and terrestrial birds.[24]

Diseases and parasites

C. virginianum can be infected by the root-knot nematode species Meloidogyne arenaria, M. incognita, and M. javanica, but it is moderately resistant.[25]

Conservation and management

It is listed as endangered by the New Jersey Department of Environmental Protection and Energy.[5] C. virginianum is tolerant to the herbicide imazapyr.[26] Also for management, it benefits from the overstory canopy being thinned.[27]

Cultivation and restoration

Photo Gallery

References and notes

  1. 1.0 1.1 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 2.2 Florida State University Robert K. Godfrey Herbarium database. URL: http://herbarium.bio.fsu.edu. Last accessed: June 2014. Collectors: Loran C. Anderson, John C. Ogden, Gwynn W. Ramsey, R.K. Godfrey, R. S. Mitchell; R. C. Phillips, K. Craddock Burks, Gary R. Knight, D. W. Mather, C. Jackson, D. B. Ward, Mary Margaret Williams, O. Lakela, Brenda Herring, Jame Amoroso, Gwynn W. Ramsey, Richard Mitchell, Gail A. Steverson, Grady W. Reinert, George R. Cooley, R. J. Eaton, R. Kral, Cecil R Slaughter, Andre F. Clewell, R. Komarek, R. F. Doren, Kevin Oakes, Richard Gaskalla, Lisa Keppner, Clarke Hudson, Wilbur H Duncan, Jean Wooten, H. R. Totten, R. L. Wilbur, C. Ritchie Bell, Delzie Demaree, F. S. Earle, A. B. Seymour, Samuel B. Jones, Jr., H. R. Reed, A. B. Seymour, Michael B. Brooks, Sidney McDaniel, D. C. Bain, D. S. Correll, H. B. Correll, Lloyd H. Shinners, Geo M. Merrill, and H J Hamby. States and Counties: Alabama: Baldwin. Arkansas: Little Rock. Florida: Bay, Citrus, Collier, Duval, Escambia, Franklin, Gadsden, Gulf, Hillsborough, Jackson, Jefferson, Leon, Liberty, Manatee, Marion, Okaloosa, Polk, St Johns, St. Lucie, Suwannee, Wakulla, and Washington. Georgia: Bartow, Grady, Madison, and Thomas. Mississippi: Forrest, Harrison, Jackson, Pearl River, and Pike. North Carolina: Alamance, Orange, and Wilkes. Texas: Angelina, Bastrop, Freestone, Harris, Morris, Tarrant, and Van Zandt.
  3. Weakley, A. S. (2015). Flora of the Southern and Mid-Atlantic States. Chapel Hill, NC, University of North Carolina Herbarium.
  4. 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. 635-6. Print
  5. 5.0 5.1 USDA, NRCS. (2016). The PLANTS Database (http://plants.usda.gov, 4 April 2019). National Plant Data Team, Greensboro, NC 27401-4901 USA.
  6. 6.0 6.1 6.2 Cathey, S. E., L. R. Boring, et al. (2010). "Assessment of N2 fixation capability of native legumes from the longleaf pine-wiregrass ecosystem." Environmental and Experimental Botany 67: 444-450.
  7. 7.0 7.1 7.2 7.3 Hainds, M. J., R. J. Mitchell, et al. (1999). "Distribution of native legumes (Leguminoseae) in frequently burned longleaf pine (Pinaceae)-wiregrass (Poaceae) ecosystems." American Journal of Botany 86: 1606-1614.
  8. 8.0 8.1 8.2 Cushwa, C. T. (1966). The response of herbaceous vegetation to prescribed burning. Asheville, USDA Forest Service.
  9. Clewell, A. F. (2013). "Prior prevalence of shortleaf pine-oak-hickory woodlands in the Tallahassee red hills." Castanea 78(4): 266-276.
  10. Brudvig, L.A. and E.I. Damchen. (2011). Land-use history, historical connectivity, and land management interact to determine longleaf pine woodland understory richness and composition. Ecography 34: 257-266.
  11. Brudvig, L.A., E Grman, C.W. Habeck, and J.A. Ledvina. (2013). Strong legacy of agricultural land use on soils and understory plant communities in longleaf pine woodlands. Forest Ecology and Management 310: 944-955.
  12. Moore, W.H., B.F. Swindel, and W.S. Terry. (1982). Vegetative Response to Clearcutting and Chopping in a North Florida Flatwoods Forest. Journal of Range Management 35(2):214-218.
  13. 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
  14. 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.
  15. Hiers, J. K. and R. J. Mitchell (2007). "The influence of burning and light availability on N-2-fixation of native legumes in longleaf pine woodlands." Journal of the Torrey Botanical Society 134: 398-409.
  16. Coffey, K. L. and L. K. Kirkman (2006). "Seed germination strategies of species with restoration potential in a fire-maintained pine savanna." Natural Areas Journal 26: 289-299.
  17. Wiggers, M. S., et al. (2017). "Seed heat tolerance and germination of six legume species native to a fire-prone longleaf pine forest." Plant Ecology 218: 151-171.
  18. Wiggers, M. S., et al. (2013). "Fine-scale variation in surface fire environment and legume germination in the longleaf pine ecosystem." Forest Ecology and Management 310: 54-63.
  19. Hendricks, J. J. and L. R. Boring (1999). "N2-fixation by native herbaceous legumes in burned pine ecosystems of the southeastern United States." Forest Ecology and Management 113: 167-177.
  20. 20.0 20.1 20.2 20.3 20.4 20.5 Hiers, J. K., R. J. Mitchell, et al. (2003). "Legumes native to longleaf pine savannas exhibit capacity for high N2-fixation rates and negligible impacts due to timing of fire." New Phytologist 157: 327-338
  21. Spears, Jr. E. E. 1987. Island and mainland pollination ecology of Centrosema virginianum and Opuntia stricta. J. Ecol. 75: 351-362.
  22. Godts J.E. 1990 The Upside-Down Flower Palmetto 10(4):3
  23. Sweeney, J. M., et al. (1981). Bobwhite quail food in young Arkansas loblolly pine plantations. Arkansas Experiment Station bulletin 852. Fayetteville, AR, University of Arkansas, Divisionn of Agriculture, Agricultural Experiment Station.
  24. Miller, J.H., and K.V. Miller. 1999. Forest plants of the southeast and their wildlife uses. Southern Weed Science Society.
  25. Quesenberry, K. H., et al. (2008). "Response of native southeastern U.S. legumes to root-knot nematodes." Crop Science 48: 2274-2278.
  26. (2000). The role of fire in nongame wildlife management and community restoration: Traditional uses and new directions, Nashville, TN, USDA Forest Service, Northeastern Research Station.
  27. Brockway, D. G. and C. E. Lewis (2003). "Influence of deer, cattle grazing and timber harvest on plant species diversity in a longleaf pine bluestem ecosystem." Forest Ecology and Management 175: 49-69.