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Introductory

SPECIES: Ilex opaca | American Holly
ABBREVIATION : ILEOPA SYNONYMS : NO-ENTRY SCS PLANT CODE : ILOP ILOPA COMMON NAMES : American holly dune holly hummock holly scrub holly TAXONOMY : The currently accepted scientific name of American holly is Ilex opaca Ait. The Ilex genus consists of 13 species belonging to the holly family (Aquifoliacea). There are no forms or subspecies of American holly. Recognized varieties include [21,34]: I. opaca var. opaca American holly I. opaca var. arenicola (Ashe) Ashe scrub or hummock holly I. opaca hybridizes with I. cassine to produce I. x attenata Ashe [47]. LIFE FORM : Tree, Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : Milo Coladonato, September 1991 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Coladoanto, Milo. 1991. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Ilex opaca | American Holly
GENERAL DISTRIBUTION : From the maritime forests of Massachusetts, American holly is scattered along the coast to Delaware. It grows inland to several Pennsylvania counties and to extreme southeastern Ohio. It occurs abundantly southward throughout the Coastal Plain, Piedmont, and Appalachians. Its range extends south to mid-peninsular Florida and west to eastern Texas and southern Missouri [23,46]. It is cultivated in Hawaii [49] ECOSYSTEMS : FRES10 White - red - jack pine FRES12 Longleaf - slash pine FRES13 Loblolly - shortleaf pine FRES14 Oak - pine FRES15 Oak - hickory FRES16 Oak - gum - cypress STATES : AL AR CT DE FL GA HI KY LA MA MD MS MO NC OH OK PA RI SC TN TX VA WV ADMINISTRATIVE UNITS : GWMP GRSM GUIS HOBE HOSP MACA MANA NATR NERI OBRI PRWI RICH ROCR SHEN SHIL BLM PHYSIOGRAPHIC REGIONS : NO-ENTRY KUCHLER PLANT ASSOCIATIONS : K089 Black Belt K090 Live oak - sea oats K095 Great Lakes pine forest K100 Oak - hickory forest K104 Appalachian oak forest K110 Northeastern oak - pine forest K111 Oak - hickory - pine forest K112 Southern mixed forest K113 Southern floodplain forest K114 Pocosin K115 Sand pine - scrub SAF COVER TYPES : 21 Eastern white pine 40 Post oak - blackjack oak 43 Bear oak 44 Chestnut oak 45 Pitch pine 46 Eastern redcedar 51 White pine - chestnut oak 52 White oak - black oak - northern red oak 53 White oak 57 Yellow poplar 58 Yellow poplar - eastern hemlock 59 Yellow poplar - white oak - northern red oak 61 River birch - sycamore 64 Sassafras - persimmon 65 Pin oak - sweetgum 69 Sand pine 70 Longleaf pine 71 Longleaf pine - scrub oak 72 Southern scrub oak 73 Southern redcedar 74 Cabbage palmetto 75 Shortleaf pine 76 Shortleaf pine - oak 78 Virginia pine - oak 79 Virginia pine 80 Loblolly pine - shortleaf pine 81 Loblolly pine 82 Loblolly pine - hardwood 83 Longleaf pine - slash pine 84 Slash pine 85 Slash pine - hardwood 87 Sweetgum - yellow poplar 88 Willow oak - water oak - diamondleaf oak 89 Live oak 91 Swamp chestnut oak - cherrybark oak 92 Sweetgum - willow oak 96 Overcup oak - water hickory 97 Atlantic white cedar 98 Pond pine 100 Pondcypress 101 Baldcypress 103 Water tupelo - swamp tupelo 104 Sweetbay - swamp tupelo - red bay 110 Black oak SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : NO-ENTRY

VALUE AND USE

SPECIES: Ilex opaca | American Holly
WOOD PRODUCTS VALUE : The wood of American holly is heavy, tough, and close-grained. It shrinks considerably, checks or warps badly unless properly seasoned, and is not durable under exposure [47]. The wood is used for veneer and to a limited extent as pulpwood and lumber. The greates use of the wood is for cabinets, interior finish, novelties, handles, fixtures, and scientific instruments. When dyed black to resemble ebony, it is used for piano keys, violin pegs, and fingerboards [23]. IMPORTANCE TO LIVESTOCK AND WILDLIFE : Birds are the principal consumers of American holly fruit, although deer, squirrels, and other small animals eat them. At least 18 species of birds, including songbirds, mourning doves, wild turkeys, and northern bobwhite, are known to eat the fruit [42,46]. Cattle and deer sometimes browse the foliage [9,18]. PALATABILITY : The palatability of American holly to white-tailed deer and cattle is considered poor. Deer and cattle generally consume American holly only when more preferred browse is unavailable [33]. NUTRITIONAL VALUE : Lay [32] listed nutrient percentage values for American holly browse collected in winter and summer on a pine-hardwood forest in Newton County, Texas: N-Free Phosphoric Protein Fat Fiber extract Ash acid Calcium Su 5.50 --- --- --- --- 0.14 --- W 6.73 3.16 26.10 46.17 2.84 0.14 0.70 These levels are low for protein and deficient for phosphoric acid, but high for calcium [32]. COVER VALUE : Cavities in American holly provide nesting habitat for the endangered red-cockaded woodpecker [28]. VALUE FOR REHABILITATION OF DISTURBED SITES : American holly is useful for rehabilitating areas that have been damaged by salt spray. It is more resistant to damage from salt spray than any associated woody species in the maritime forest of New England [23,25]. OTHER USES AND VALUES : The attractiveness of American holly's foliage is its principal value, whether as a forest tree, planted ornamental, or Christmas decoration. It can be used for yard, street, park planting, or for hedges [6,47]. MANAGEMENT CONSIDERATIONS : American holly is considered an undesirable shrub that competes with pines and desirable hardwoods for light, moisture, and nutrients. Streamline basal application of the herbicide Garlon 4 is an effective means of controlling American holly [38]. The greatest damage to American holly trees is the indiscriminate harvesting of foliage with berries for Christmas decorating. Before laws were passed in Maryland and Delaware to protect the holly, there was a "roadside" market for holly collected from trees that did not belong to the harvesters. Trees were left mutilated and many died [23].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Ilex opaca | American Holly
GENERAL BOTANICAL CHARACTERISTICS : American holly is a native evergreen tree that grows to 50 feet (16 m) [12,17]. Its evergreen leaves are leathery, with sharp pointed tips and spiny toothed margins. The branches are short and crooked, and the crown is rounded or pyramidal. The greenish white flowers are unisexual, dioecious, and borne on short-stalked, axillary cymose clusters. The fruit is a round, bright red, orange, or occasionally yellow, four-seeded drupe or pyrene. The bark is thin, gray, and often warty [21,40] RAUNKIAER LIFE FORM : Undisturbed State: Panerophyte (mesophanerophyte) Undisturbed State: Panerophyte (microphanerophyte) Undisturbed State: Phanerophyte (nanophanerophyte) Burned or Clipped State: Cryptophyte (geophyte) REGENERATION PROCESSES : Sexual: American holly is pollinated by insects, including bees, ants, wasps, and night-flying moths [23]. Seed germination is slow, requiring 16 months to 3 years in nature [46]. Seeds are mainly dispersed by birds and small mammals [10,13]. Seed production may be low in years of heavy spring rain, as rain can diminish the wide dissemination of pollen. A frost can kill the spring flowers, eliminating the fruit crop. Frequent prescribed burning will also reduce fruit production [20]. Vegetative: American holly sprouts from basal dormant buds [23,44]. Propagation: Transplanting of young holly trees should be done during the dormant season, usually November through March. Small plants may be dug bare-rooted if roots are kept moist, but larger plants should be balled and burlapped. When wild hollies are transplanted from the woods, tops should be severely pruned and most of the remaining leaves removed. Small trees should be allowed to flower before transplanting to ensure the selection of fruit-bearing individuals. Root pruning to a depth of 2 to 3 feet (0.6-0.9 m) a year before lifting improves transplanting success. Holly can be produced from cuttings taken in August or September and December. Cuttings should be taken from the current season's ripened wood, with a small section of 2-year-old wood including several leaves. Cuttings should be set slanting in about 6 inches (15 cm) of moist peatmoss, with the leaves lying flat on the surface [23]. SITE CHARACTERISTICS : American holly is primarily a plant of the humid Southeast. It occupies a wide variety of soils, from nearly sterile Inceptisols of the Atlantic sandy beaches to fertile but thin mountain Ultisols to an elevation of approximately 3,000 feet (915 m) [12,23]. The largest trees are found in the rich bottomlands and swamps of the Coastal Plain. Growth is best in moist, slightly acidic, well-drained sites such as upland pine sites and hammocks [8,11]. Trees will not survive flooding or saturated soils for more than 17 percent of the growing season. In the northeastern portion of its range, holly is found on sandy soils on the Coast and on dry gravely soils farther inland [2,12]. American holly is a common understory component in the longleaf pine (Pinus palustris)-slash pine (P. elliotti) forests of the Coastal Plain. Other common associates include sweetgum (Liquidambar styraciflua), flowering dogwood (Cornus florida), American beech (Fagus grandifolia), red maple (Acer rubrum), white oak (Quercus alba), water oak (Q. nigra), hickory (Carya spp.), white ash (Fraxinus americana), yellow poplar (Liriodendron tulipifera), black tupelo (Nysaa sylvatica), southern red oak (Quercus falcata), and post oak (Q. stellata) [23]. SUCCESSIONAL STATUS : American holly is seldom dominant because of its slow growth and relatively short stature [3,16]. It is very shade tolerant and can survive in the understory of most forest canopies [20]. American holly's slow growth allows faster growing species to overtop it. Shade and root competition in natural stands reduces the average height of hollies compared with those growing in full sunlight. Crown area is reduced by more than one-third under medium shade and by more than one-half under heavy shade [23,30]. SEASONAL DEVELOPMENT : American holly begins flowering in April in the southern parts of its range and in June at its northern limits. The fruit ripen from September through December and remain on the tree through most of the winter [46,47].

FIRE ECOLOGY

SPECIES: Ilex opaca | American Holly
FIRE ECOLOGY OR ADAPTATIONS : American holly is very susceptible to fire. It may persist by sprouting from the root crown. [1,7,14]. POSTFIRE REGENERATION STRATEGY : survivor species; on-site surviving root crown or caudex off-site colonizer; seed carried by animals or water; postfire yr 1&2

FIRE EFFECTS

SPECIES: Ilex opaca | American Holly
IMMEDIATE FIRE EFFECT ON PLANT : Fire easily top-kills American holly [37]. Its thin bark is easily injured by fire [24,26]. The cambium layer is destroyed and the leaves and crown defoliated. Even large trees may be killed by light fires in the understory [27,48]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : American holly sprouts from surviving basal buds following fire [31]. Initial growth after fire is slow, averaging about 6 feet (1.8 m) in 16 years under medium shade [23]. Three annual fires in a southern pine forest reduced the number of fruit-producing holly trees by 95 percent [27]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : Fire is a very effective agent for controlling American holly. Seedlings and sprouts can usually be eliminated as a result of normal underburning regimes in most commercial pine stands [48].

REFERENCES

SPECIES: Ilex opaca | American Holly
REFERENCES : 1. Allen, Peter H. 1960. Scorch and mortality after a summer burn in loblolly pine. Res. Note No. 144. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 2 p. [12256] 2. Beaven, George Francis; Oosting, Henry J. 1939. Pocomoke Swamp: a study of a cypress swamp on the eastern shore of Maryland. Bulletin of the Torrey Botanical Club. 66: 376-389. [14507] 3. Billings, W. D. 1938. The structure and development of old field shortleaf pine stands and certain associated physical properties of the soil. Ecological Monographs. 8(3): 437-499. [10701] 4. Blair, Robert M.; Brunett, Louis E. 1976. Phytosociological changes after timber harvest in a southern pine ecosystem. Ecology. 57: 18-32. [9646] 5. Blinn, Charles R.; Buckner, Edward R. 1989. Normal foliar nutrient levels in North American forest trees: A summary. Station Bulletin 590-1989. St. Paul, MN: University of Minnesota, Minnesota Agricultural Experiment Station. 27 p. [15282] 6. Bonner, F. T. 1974. Ilex L. holly. In: Schopmeyer, C. S., ed. Seeds of woody plants in the United States. Agric. Handb. 450. Washington, DC: U.S. Department of Agriculture, Forest Service: 450-453. [7683] 7. Bruce, David. 1947. Thirty-two years of annual burning in longleaf pine. Journal of Forestry. 45(11): 809-814. [11001] 8. Cain, Stanley A. 1931. Ecological studies of the vegetation of the Great Smoky Mountains of North Carolina and Tennessee. Botanical Gazette. 91: 22-41. [10340] 9. Conover, M. R.; Kania, G. S. 1988. Browsing preference of white-tailed deer for different ornamental species. Wildlife Society Bulletin. 16: 175-179. [8933] 10. Daubenmire, Rexford. 1990. The Magnolia grandiflora-Quercus virginiana forest of Florida. American Midland Naturalist. 123: 331-347. [10871] 11. Delcourt, Hazel R.; Delcourt, Paul A. 1974. Primeval magnolia-holly-beech climax in Louisiana. Ecology. 55(3): 638-644. [11469] 12. Duncan, Wilbur H.; Duncan, Marion B. 1988. Trees of the southeastern United States. Athens, GA: The University of Georgia Press. 322 p. [12764] 13. Engstrom, R. Todd; Crawford, Robert L.; Baker, W. Wilson. 1984. Breeding bird populations in relation to changing forest structure following fire exclusion: a 15-year study. Wilson Bulletin. 96(3): 437-450. [9873] 14. Ewel, Katherine Carter; Mitsch, William J. 1978. The effects of fire on species composition in cypress dome ecosystems. Florida Scientist. 41(1): 25-31. [14634] 15. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 16. Feldman, Thomas D. 1987. Fire control and ecological succession in McCarty Woods, Hernando County , Florida. Florida Geographer. 21: 15-19. [8689] 17. 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] 18. Garrison, George A. 1972. Carbohydrate reserves and response to use. In: McKell, Cyrus M.; Blaisdell, James P.; Goodin, Joe R., eds. Wildland shrubs--their biology and utilization: Proceedings of a symposium; 1971 July; Logan, UT. Gen. Tech. Rep. INT-1. Ogden, UT; U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 271-278. [997] 19. 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] 20. Glitzenstein, Jeff S.; Harcombe, Paul A.; Streng, Donna R. 1986. Disturbance, succession, and maintenance of species diversity in an east Texas forest. Ecological Monographs. 56(3): 243-258. [9670] 21. 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] 22. Grelen, Harold E. 1983. Comparison of seasons and frequencies of burning in a young slash pine plantation. Res. Pap. SO-185. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. 5 p. [10996] 23. Grelen, H. E. 1990. Ilex opaca Ait. American holly. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Vol. 2. Hardwoods. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 379-385. [9131] 24. Hare, Robert C. 1965. Contribution of bark to fire resistance of southern trees. Journal of Forestry. 63(4): 248-251. [9915] 25. Hartley, Jeanne J.; Arner, Dale H.; Hartley, Danny R. 1990. Survival of planted woody species on disposal areas of the Tennessee-Tombigbee Waterway. In: Hughes, H. Glenn; Bonnicksen, Thomas M., eds. Restoration '89: the new management challenge: Proceedings, 1st annual meeting of the Society for Ecological Restoration; 1989 January 16-20; Oakland, CA. Madison, WI: The University of Wisconsin Arboretum, Society for Ecological Restoration: 244-250. [12090] 26. Hodgkins, Earl J. 1958. Effects of fire on undergrowth vegetation in upland southern pine forests. Ecology. 39(1): 36-46. [7632] 27. Johnson, A. Sydney; Landers, J. Larry. 1978. Fruit production in slash pine plantations in Georgia. Journal of Wildlife Management. 42(3): 606-613. [9855] 28. Kalisz, Paul J.; Boettcher, Susan E. 1991. Active and abandoned red-cockaded woodpecker habitat in Kentucky. Journal of Wildlife Management. 55(1): 146-154. [13837] 29. 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] 30. Kurz, Herman. 1944. Secondary forest succession in the Tallahassee Red Hills. Proceedings, Florida Academy of Science. 7(1): 59-100. [10799] 31. Langdon, O. Gordon. 1981. Some effects of prescribed fire on understory vegetation in loblolly pine stands. In: Wood, Gene W., ed. Prescribed fire and wildlife in southern forests: Proceedings of a symposium; 1981 April 6-8; Myrtle Beach, SC. Georgetown, SC: Clemson University, Belle W. Baruch Forest Science Institute: 143-153. [14821] 32. Lay, Daniel W. 1957. Browse quality and the effects of prescribed burning in southern pine forests. Journal of Forestry. 55: 342-347. [7633] 33. Lay, Daniel W. 1967. Browse palatability and the effects of prescribed burning in southern pine forests. Journal of Forestry. 65(11): 826-828. [145] 34. Little, Elbert L., Jr. 1979. Checklist of United States trees (native and naturalized). Agric. Handb. 541. Washington, DC: U.S. Department of Agriculture, Forest Service. 375 p. [2952] 35. Lotti, Thomas; Klawitter, Ralph A.; LeGrande, W. P. 1960. Prescribed burning for understory control in loblolly pine stands of the coastal plain. Station Pap. No. 116. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 19 p. [15417] 36. Lyon, L. Jack; Stickney, Peter F. 1976. Early vegetal succession following large northern Rocky Mountain wildfires. In: Proceedings, Tall Timbers fire ecology conference and Intermountain Fire Research Council fire and land management symposium; 1974 October 8-10; Missoula, MT. No. 14. Tallahassee, FL: Tall Timbers Research Station: 355-373. [1496] 37. McKinley, Carol E.; Day, Frank P., Jr. 1979. Herb. prod. in cut-burned, uncut-burned & contl areas of a Chamaecyparis thyoides (L.) BSP (Cupressaceae) stand in the Great Dismal Swamp. Bulletin of the Torrey Botanical Club. 106(1): 20-28. [14089] 38. Miller, James H. 1990. Streamline basal application of herbicide for small-stem hardwood control. Southern Journal of Applied Forestry. 14(4): 161-165. [13538] 39. Petruncio, Mark; Lea, Russ. 1985. Natural hardwood regeneration in the southern Appalachians. In: Shoulders, Eugene, ed. Proceedings, 3rd biennial southern silvicultural research conference; 1984 November 7-8; Atlanta, GA. General Technical Report SO-54. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station: 178-182. [7389] 40. 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] 41. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843] 42. Stransky, J. J.; Halls, L. K.; Nixon, E. S. 1976. Plants following timber harvest: importance to songbirds. Texas Forestry Pap. No. 28. Nacogdoches, TX: Stephen F. Austin State University, School of Forestry. 13 p. [15292] 43. Stransky, John J.; Huntley, Jimmy C.; Risner, Wanda J. 1986. Net community production dynamics in the herb-shrub stratum of a loblolly pine-hardwood forest: effects of clearcutting and site prepar. Gen. Tech. Rep. SO-61. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. 11 p. [9835] 44. Trousdell, Kenneth B. 1970. Disking and prescribed burning: sixth-year residual effects on loblolly pine and competing vegetation. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 6 p. [10190] 45. 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] 46. Van Dersal, William R. 1938. Native woody plants of the United States, their erosion-control and wildlife values. Washington, DC: U.S. Department of Agriculture. 362 p. [4240] 47. Vines, Robert A. 1960. Trees, shrubs, and woody vines of the Southwest. Austin, TX: University of Texas Press. 1104 p. [7707] 48. Watson, Geraldine E. 1986. Influence of fire on the longleaf pine - bluestem range in the Big Thicket region. In: Kulhavy, D. L.; Conner, R. N., eds. Wilderness and natural areas in the eastern United States: a management challenge. Nacogdoches, TX: Stephen F. Austin University: 181-185. [10334] 49. St. John, Harold. 1973. List and summary of the flowering plants in the Hawaiian islands. Hong Kong: Cathay Press Limited. 519 p. [25354]

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