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Introductory

SPECIES: Smilax bona-nox | Saw Greenbrier
ABBREVIATION : SMIBON SYNONYMS : NO-ENTRY SCS PLANT CODE : SMBO2 COMMON NAMES : saw greenbrier greenbrier catbrier bullbrier chinabrier tramp's trouble TAXONOMY : The currently accepted scientific name for saw greenbrier is Smilax bona-nox L. (Smilacaceae, formerly Liliaceae) [12,16,19,30,41]. The following varieties are recognized by various authors: S. b. var. bona-nox S. b. var. hastata (Willd.) DC [12,30,41] S. b. var. exaruiculata Fern. [12,41] S. b. var. hederaefolia (Beyrich) Fern. [12,30,34,41] S. b. var. littoralis Coker [41] Throughout this write-up, the term 'greenbrier' will be used to refer to cases where other Smilax species are treated with saw greenbrier, or where Smilax species are undifferentiated. LIFE FORM : Vine FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : Janet Sullivan, March 1994 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Sullivan, Janet. 1994. Smilax bona-nox. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Smilax bona-nox | Saw Greenbrier
GENERAL DISTRIBUTION : The range of saw greenbrier extends from Florida to Texas and eastern Mexico, north to Maryland, Kentucky, southern Indiana, southern Illinois, Missouri, and southeastern Kansas [12,16,17,34]. ECOSYSTEMS : FRES12 Longleaf - slash pine FRES14 Oak - pine FRES15 Oak - hickory FRES17 Elm - ash - cottonwood STATES : AL AR FL GA IL IN KS KY LA MD MS MO NC OK SC TN TX VA MEXICO ADMINISTRATIVE UNITS : ASIS BICY BITH BLRI CAHA CALO CHCH COLO COSW CUGA CUIS EVER FOCA FODO GWCA GRSM GUIS HOBE HOSP JELA MACA NATR OBRI OZAR RICH SHIL WICR BLM PHYSIOGRAPHIC REGIONS : 14 Great Plains KUCHLER PLANT ASSOCIATIONS : K081 Oak savanna K082 Mosaic of K074 and K100 K084 Cross Timbers K089 Black Belt K092 Everglades K100 Oak - hickory forest K104 Appalachian oak forest K105 Mangrove K111 Oak - hickory - pine forest K112 Southern mixed forest K114 Pocosin K115 Sand pine scrub K116 Subtropical pine forest SAF COVER TYPES : 52 White oak - black oak - northern red oak 53 White oak 57 Yellow-poplar 63 Cottonwood 62 Silver maple - American elm 70 Longleaf pine 71 Longleaf pine - scrub oak 73 Southern redcedar 75 Shortleaf pine 76 Shortleaf pine - oak 78 Virginia pine - oak 81 Loblolly pine 82 Loblolly pine - hardwood 83 Longleaf pine - slash pine 87 Sweetgum - yellow-poplar 88 Willow oak - water oak - diamondleaf oak 89 Live oak 92 Sweetgum - willow oak 93 Sugarberry - American elm - green ash 94 Sycamore - sweetgum - American elm 97 Atlantic white-cedar 102 Baldcypress - tupelo 103 Water tupelo - swamp tupelo 104 Sweetbay - swamp tupelo - redbay 110 Black oak SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Saw greenbrier occurs in a wide variety of habitat and community types, and is not characteristic of any particular conditions. Its most common understory associates include muscadine grape (Vitus rotundifolia), Virginia creeper (Parthenocissus quinquefolia), trumpetcreeper (Campsis radicans), peppervine (Ampelopsis arborea), Alabama supplejack (Berchemia scandens), and eastern poison-ivy (Toxicodendron radicans) [2,6,11,25,39].

VALUE AND USE

SPECIES: Smilax bona-nox | Saw Greenbrier
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : The fruits of saw greenbrier are eaten by wood ducks, ruffed grouse, wild turkeys, fish crows, black bears, opossums, raccoons, squirrels, and many species of songbirds [7]. White-tailed deer browse the foliage [5,19,43]. The commonly low and straggling growth form of saw greenbrier tends to form an impenetrable mass of prickly branches, which creates good cover for small mammals and birds [34]. In Kansas, an increase in the numbers of white-footed mice was associated with an increase in woody and weedy species (including saw greenbrier); the increase in mice was attributed to increased low cover. The mice were rare on the study site prior to the loss of an American elm overstory due to Dutch elm disease [13]. PALATABILITY : Saw greenbrier is palatable browse for white-tailed deer [6]. NUTRITIONAL VALUE : NO-ENTRY COVER VALUE : NO-ENTRY VALUE FOR REHABILITATION OF DISTURBED SITES : Saw greenbrier occurred on unreclaimed lignite surface mine sites in east-central Texas. It was recorded on 15-, 30-, and 50-year-old sites and in adjacent undisturbed forest. Its highest frequency occurred in undisturbed forest sites [33]. OTHER USES AND VALUES : NO-ENTRY MANAGEMENT CONSIDERATIONS : Saw greenbrier productivity in the understory of a loblolly pine plantation was highest under medium thinning intensities [5]. Closed canopy plantations produced little browse [6]. Saw greenbrier is considered a pest species in some areas; it is difficult to eradicate due to its persistent woody rhizome [7]. It is resistant to most herbicides, but can be controlled with karbutilate [42].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Smilax bona-nox | Saw Greenbrier
GENERAL BOTANICAL CHARACTERISTICS : Saw greenbrier is a native, evergreen [8] to semievergreen [16] or deciduous [7], rhizomatous vine up to 26 feet (8 m) in length [12]. The quadrate stems and branches have scattered to numerous stiff prickles [7,12]. There are two forms of rhizomes: ligneous, thickened, knotty tubers 0.8 to 2.4 inches (2-6 cm) thick in clusters up to 7.9 inches (20 cm) across [23], and more slender rhizomes which give rise to the erect stems [7,12,16,23]. The inflorescence is an umbel borne on an axillary peduncle. The fruit is a one-seeded drupe [30]. RAUNKIAER LIFE FORM : Phanerophyte Geophyte REGENERATION PROCESSES : Saw greenbrier reproduces by seed and by rhizomes. The seeds are animal dispersed and can be carried long distances by birds [8]. SITE CHARACTERISTICS : Saw greenbrier occurs in a variety of habitats, including dry and wet woods, thickets, and hammocks, and disturbed sites such as clearings, roadsides, fencerows, and old fields. It tolerates a wide variety of soils, including dry to moist sands, rocky soils, rich loams, and saturated swamp soils high in organic matter [7,10,12,16,34,41]. SUCCESSIONAL STATUS : Facultative Seral Species Saw greenbrier is often found in early seres and disturbed sites. It is listed with other plants characterized as early arrivals following disturbance [8]. In sand dune succession of barrier islands off North Carolina, several vines successively colonize inland dunes: Virginia creeper, eastern poison-ivy, and saw greenbrier [2]. Saw greenbrier was reported from years 0 to 10 in oldfield succession in Georgia, but was not discussed for later stages and the authors implied that the early colonizers were crowded out by Japanese honeysuckle (Lonicera japonica) and other species [27]. Saw greenbrier was the most widely distributed and abundant vine on a 3-year-old gravel pit in eastern Texas. It was present in successively lower numbers on a 5-year-old gravel pit, a 47-year-old gravel pit, and the adjacent unexcavated forest [44]. In Kansas, loss of American elms (Ulmus americana) to Dutch elm disease further opened an already open canopy and created conditions where cover values of woody and weedy species increased, including that of saw greenbrier [13]. In Florida, saw greenbrier occurred in a stand composed of large, old trees (mostly laurel oak [Quercus laurifolia], pignut hickory [Carya glabra], and magnolia [Magnolia grandiflora]) with no evidence of past fire, logging, or grazing [8]. SEASONAL DEVELOPMENT : Saw greenbrier flowers from April to May [17,19,30], April to June [41], or May to July [7] depending on latitude. The fruit ripens from September to October, persisting on the vine through the winter [7,19,34].

FIRE ECOLOGY

SPECIES: Smilax bona-nox | Saw Greenbrier
FIRE ECOLOGY OR ADAPTATIONS : Saw greenbrier is tolerant of periodic fire because it will sprout from the rhizomes when top-killed. It is not dependent on fire for regeneration; it occurs in both fire-tolerant communities and communities which infrequently experience fire. Saw greenbrier occurs in the pine flatwoods of the lower Atlantic Coastal Plain, which were historically maintained in open condition by periodic fire, and are now managed with prescribed fires [15]. Similarly, it is often found in longleaf pine (Pinus palustris) communities which were also historically maintained by fire. Longleaf pine communities have largely been replaced by other communities, mostly loblolly pine, which can also be managed with prescribed fire [43]. Saw greenbrier is a member of Florida bay swamps, which experience fire on the average of once per century [10]. Where saw greenbrier occurs on Cumberland Island, Georgia, its distribution is probably only partly affected by fire. The scrub and marsh communities on Cumberland Island historically experienced wildfires approximately every 20 to 27 years. Oak (Quercus spp.)/saw-palmetto (Serenoa repens) communities are vulnerable to fires burning into them from adjacent scrub or marsh. Greenbriers occurred on forested sites, decreased at forest/marsh and forest/scrub interfaces, and were not present in interior marsh and scrub sites. The authors concluded that the marsh/forest and scrub/forest boundaries are controlled by fluctuation in the water table and not by fire [26]. POSTFIRE REGENERATION STRATEGY : Rhizomatous shrub, rhizome in soil Secondary colonizer - off-site seed

FIRE EFFECTS

SPECIES: Smilax bona-nox | Saw Greenbrier
IMMEDIATE FIRE EFFECT ON PLANT : Saw greenbrier is probably top-killed by most fires and subsequently sprouts from the rhizomes. Mortality due to a winter prescribed fire in Texas ranged from 11 percent to 31 percent for most understory plants, including saw greenbrier [36]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : In Oklahoma, a post oak (Quercus stellata)-blackjack oak (Quercus marilandica) and tallgrass prairie mosaic was subjected to prescribed fire to determine the response of understory species to fire and timing of fire. The groundlayer vegetation was dominated by little bluestem (Schizachyrium scoparium). Saw greenbrier was present only on sites that were selected to receive prescribed fire in summer (July 1979), and showed very little difference in cover following the fire [1]. In Texas, a prescribed fire in March, 1974, consumed 80 to 90 percent of the previous year's needle and leaf cast and 50 percent of old litter under a loblolly pine-shortleaf pine (Pinus echinata) plantation. Average saw greenbrier height decreased from 11.8 feet (3.60 m) to 4.79 feet (1.46 m) and the average number of stems per plant increased from 1.47 to 1.62 (measured July 1975) [36]. Also in Texas, prescribed surface fires were conducted in February, 1982 to assess the response of vegetation under either Texas live oak (Quercus virginiana var. fusiformis) or post oak. By July, saw greenbrier had increased in relative dominance and frequency on both site types [20]. In Florida, frequent prescribed fires in longleaf pine-slash pine (Pinus elliotii) communities have prevented the formation of a hardwood midstory. Saw greenbrier was the most common vine in these fire-maintained stands [38]. Also in Florida, longleaf pine-turkey oak (Quercus laevis) stands were subjected to periodic prescribed fire. Greenbriers were present in low numbers (15 percent occurrence) and were subjected to heavy spring browsing (90 percent of twigs browsed=90 percent utilization) on 1-year-old burns. Greenbriers were not reported from study plots that represented postfire years 2, 3, and 4 [18]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : Since herbicides alone do not control saw greenbrier, prescribed burning has been suggested to help hold it to desired densities for wildlife habitat and to improve its browse value [32]. However, in the Cross Timbers of Oklahoma, herbicides plus annual spring fires had no effect on saw greenbrier cover [37].

REFERENCES

SPECIES: Smilax bona-nox | Saw Greenbrier
REFERENCES : 1. Adams, Dwight E.; Anderson, Roger C.; Collins, Scott L. 1982. Differential response of woody and herbaceous species to summer and winter burning in an Oklahoma grassland. Southwestern Naturalist. 27: 55-61. [6282] 2. Au, Shu-fun. 1974. Vegetation and ecological processes on Shackleford Bank, North Carolina. Scientific Monograph Series No 6: NPS 113. Washington, DC: U.S. Department of the Interior, National Park Service. 86 p. [16101] 3. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434] 4. Best, G. Ronnie; Segal, Debra S.; Wolfe, Charlotte. 1990. Soil-vegetation correlations in selected wetlands and uplands of north-central Florida. Biol. Rep. 90(9). Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 51 p. [18161] 5. Blair, Robert M. 1960. Deer forage increased by thinnings in a Louisiana loblolly pine plantation. Journal of Wildlife Management. 24(4): 401-405. [16891] 6. Blair, Robert M.; Short, Henry L.; Epps, E. A., Jr. 1977. Seasonal nutrient yield and digestibility of deer forage from a young pine plantation. Journal of Wildlife Management. 41(4): 667-676. [16963] 7. Brown, Russell G.; Brown, Melvin L. 1972. Woody plants of Maryland. Baltimore, MD: Port City Press. 347 p. [21844] 8. Daubenmire, Rexford. 1990. The Magnolia grandiflora-Quercus virginiana forest of Florida. American Midland Naturalist. 123: 331-347. [10871] 9. Egler, Frank E. 1952. Southeast saline Everglades vegetation, Florida, and its management. Vegetatio. 3: 213-265. [11479] 10. Ewel, Katherine C. 1990. Swamps. In: Myers, Ronald L.; Ewel, John J., eds. Ecosystems of Florida. Orlando, FL: University of Central Florida Press: 281-322. [17392] 11. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 12. Fernald, Merritt Lyndon. 1950. Gray's manual of botany. [Corrections supplied by R. C. Rollins]. Portland, OR: Dioscorides Press. 1632 p. (Dudley, Theodore R., gen. ed.; Biosystematics, Floristic & Phylogeny Series; vol. 2). [14935] 13. Fitch, Henry S.; Kettle, W. Dean. 1983. Ecological succession in vegetation and small mammal populations on a natural area of northeastern Kansas. In: Kucera, Clair L., ed. Proceedings, 7th North American prairie conference; 1980 August 4-6; Springfield, MO. Columbia, MO: University of Missouri: 117-121. [3211] 14. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998] 15. Gilliam, Frank S.; Christensen, Norman L. 1986. Herb-layer response to burning in pine flatwoods of the lower Coastal Plain of South Carolina. Bulletin of the Torrey Botanical Club. 113(1): 42-45. [4419] 16. Godfrey, Robert K. 1988. Trees, shrubs, and woody vines of northern Florida and adjacent Georgia and Alabama. Athens, GA: The University of Georgia Press. 734 p. [10239] 17. Great Plains Flora Association. 1986. Flora of the Great Plains. Lawrence, KS: University Press of Kansas. 1392 p. [1603] 18. Harlow, Richard F.; Bielling, Paul. 1961. Controlled burning studies in longleaf pine-turkey oak association on the Ocala National Forest. Proceeding, Annual Conference of Southeastern Association of Game and Fish. 15: 9-24. [9905] 19. Hunter, Carl G. 1989. Trees, shrubs, and vines of Arkansas. Little Rock, AR: The Ozark Society Foundation. 207 p. [21266] 20. Hutcheson, Ann-Marie; Baccus, John T.; McClean, Terry M.; Fonteyn, Paul J. 1989. Response of herbaceous vegetation to prescribed burning in the Hill Country of Texas. Texas Journal of Agriculture and Natural Resources. 3: 42-47. [17777] 21. Kartesz, John T.; Kartesz, Rosemarie. 1980. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Volume II: The biota of North America. Chapel Hill, NC: The University of North Carolina Press; in confederation with Anne H. Lindsey and C. Richie Bell, North Carolina Botanical Garden. 500 p. [6954] 22. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384] 23. Martin, Ben F.; Tucker, S. C. 1985. Developmental studies in Smilax (Liliaceae). I. Organography and the shoot apex. American Journal of Botany. 72(1): 66-74. [15086] 24. Masters, Ronald E. 1989. Nutrient response to overstory removal and winter prescribed fire versus clearcutting and summer site prep burns in Oklahoma Ouachita Mountains. In: Waldrop, Thomas A., ed. Proceedings of pine-hardwood mixtures: a symposium on management and ecology of the type; 1989 April 18-19; Atlanta, GA. Gen. Tech. Rep. SE-58. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 256. [10283] 25. McLendon, Terry. 1991. Preliminary description of the vegetation of south Texas exclusive of coastal saline zones. Texas Journal of Science. 43(1): 13-32. [14890] 26. McPherson, Guy R.; Bratton, Susan P. 1991. Effects of disturbance on community boundary dynamics on Cumberland Island, Georgia. In: Proceedings, 17th Tall Timbers fire ecology conference; 1989 May 18-21; Tallahassee, FL. Tallahassee, FL: Tall Timbers Research Station: 163-182. [17607] 27. Nicholson, Stuart A.; Monk, Carl D. 1974. Plant species diversity in old-field succession on the Georgia piedmont. Ecology. 55: 1075-1085. [17523] 28. Nixon, E. S.; Ward, J. R.; Fountain, E. A.; Neck, J. S. 1991. Woody vegetation of an old-growth creekbottom forest in north-central Texas. Texas Journal of Science. 43(2): 157-164. [15407] 29. Pessin, L. J. 1938. The effect of vegetation on the growth of longleaf pine seedlings. Ecological Monographs. 8(1): 119-149. [10329] 30. Radford, Albert E.; Ahles, Harry E.; Bell, C. Ritchie. 1968. Manual of the vascular flora of the Carolinas. Chapel Hill, NC: The University of North Carolina Press. 1183 p. [7606] 31. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843] 32. Scifres, C. J. 1980. Integration of prescribed burning with other practices in brush management systems. In: Hanselka, C. Wayne, ed. Prescribed range burning in the coastal prairie and eastern Rio Grande Plains of Texas: Proceedings of a symposium; 1980 October 16; Kingsville, TX. College Station, TX: The Texas A&M University System, Texas Agricultural Extension Service: 65-71. [11451] 33. Skousen, J. G.; Call, C. A.; Knight, R. W. 1990. Natural revegetation of an unreclaimed lignite surface mine in east-central Texas. Southwestern Naturalist. 35(4): 434-440. [21195] 34. Stephens, H. A. 1973. Woody plants of the North Central Plains. Lawrence, KS: The University Press of Kansas. 530 p. [3804] 35. Stickney, Peter F. 1989. Seral origin of species originating in northern Rocky Mountain forests. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT; RWU 4403 files. 7 p. [20090] 36. Stransky, John J.; Halls, Lowell K. 1979. Effect of a winter fire on fruit yields of woody plants. Journal of Wildlife Management. 43(4): 1007-1010. [9660] 37. Stritzke, Jimmy F.; Engle, David M.; McCollum, F. Ted. 1991. Vegetation management in the Cross Timbers: response of woody species to herbicides and burning. Weed Technology. 5(2): 400-405. [16395] 38. Tanner, George W. 1987. Soils and vegetation of the longleaf/slash pine forest type, Apalachicola National Forest, Florida. In: Pearson, Henry A.; Smeins, Fred E.; Thill, Ronald E., compilers. Ecological, physical, and socioeconomic relationships within southern National Forests; 1987 May 26-27; Long Beach, MS. Gen. Tech. Rep. SO-68. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station: 186-200. [10173] 39. Thieret, John W. 1971. Quadrat study of a bottomland forest in St. Martin Parish, Louisiana. Castanea. 36: 174-181. [9923] 40. U.S. Department of Agriculture, Soil Conservation Service. 1982. National list of scientific plant names. Vol. 1. List of plant names. SCS-TP-159. Washington, DC. 416 p. [11573] 41. Vines, Robert A. 1960. Trees, shrubs, and woody vines of the Southwest. Austin, TX: University of Texas Press. 1104 p. [7707] 42. Hamel, Dennis R. 1981. Forest management chemicals: A guide to use when considering pesticides for forest management. Agric. Handb. 585. Washington, DC: U.S. Department of Agriculture, Forest Service. 512 p. [7847] 43. Goodrum, Phil D.; Reid, Vincent H. 1958. Deer browsing in the longleaf pine belt. In: Proceedings, 58th annual meeting of the Society of American Foresters; [Date of meeting unknown]; [Place of meeeting unknown]. Washington, DC: [Society of American Foresters]: 139-143. [17023] 44. Nixon, Elray S. 1975. Successional stages in a hardwood bottomland forest near Dallas, Texas. Southwestern Naturalist. 20: 323-335. [12250]

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