SPECIES: Gaylussacia ursina

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Gaylussacia ursina: INTRODUCTORY


SPECIES: Gaylussacia ursina



2005 Linda Lee, University of South Carolina Herbarium

Gucker, Corey L. 2005. Gaylussacia ursina. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].




bear huckleberry

The currently accepted scientific name of bear huckleberry is Gaylussacia ursina (Ericaceae) (M.A. Curtis) Torr. & Gray [21,30,45].

In this review, the common name bear huckleberry is used to refer to literature specific to Gaylussacia ursina. The generic term "huckleberries" is used when citing literature that did not specify species and instead referred to either huckleberries or Gaylussacia species.





SPECIES: Gaylussacia ursina
Bear huckleberry occupies a narrow range. It occurs in the mountainous regions of North Carolina and adjacent parts of South Carolina, Georgia, Tennessee [30,45], and southwestern Virginia [40]. Plants database provides a distributional map of bear huckleberry.

FRES10 White-red-jack pine
FRES13 Loblolly-shortleaf pine
FRES14 Oak-pine
FRES15 Oak-hickory

STATES/PROVINCES: (key to state/province abbreviations)


K104 Appalachian oak forest
K111 Oak-hickory-pine
K112 Southern mixed forest

21 Eastern white pine
22 White pine-hemlock
23 Eastern hemlock
43 Bear oak
44 Chestnut oak
45 Pitch pine
50 Black locust
51 White pine-chestnut oak
52 White oak-black oak-northern red oak
53 White oak
55 Northern red oak
57 Yellow-poplar
58 Yellow-poplar-eastern hemlock
59 Yellow-poplar-white oak-northern red oak
75 Shortleaf pine
76 Shortleaf pine-oak
78 Virginia pine-oak
79 Virginia pine


Bear huckleberry is common in many forest types occurring predominantly in the Blue Ridge and Great Smoky mountains of the southern Appalachian Mountain Range. Elliott and Swank [9] found bear huckleberry density was 10 times greater in more xeric oak-pine (Quercus-Pinus spp.) forests than in more mesic mixed oak forests of the southern Appalachians.

Deciduous forests:
In oak-hickory (Carya spp.) forests of the southern Appalachians, chestnut oak (Q. prinus), scarlet oak (Q. coccinea), northern red oak (Q. rubra), white oak (Q. alba), and  black oak (Q. velutina) dominate. While American chestnut (Castanea dentata) trees dominated these forests in the past, American chestnut sprouts are now restricted to the understory where bear huckleberry is also common [3,33]. Bear huckleberry is also present in the understory of subxeric, oak/heath (Ericaceae) forests below 3,500 feet (1,067 m) in the Great Smoky Mountains. In what have formerly been described as chestnut oak forests of the Great Smoky Mountains, shrub coverage can be as high as 80% on mid-elevation sites. Important shrub species include bear huckleberry, Rhododendron spp., and mountain-laurel (Kalmia latifolia) [43].

White oak, northern red oak, chestnut oak, mockernut hickory (C. alba), and pignut hickory (C. glabra) characterize the montane oak-hickory forests of the southern Appalachians where bear huckleberry is common. Flame azalea (R. calendulaceum) and red maple (Acer rubrum) are also common associates. The white oak/bear huckleberry-flame azalea community is associated with the rare Ultramafic Outcrop Barren forest type that occurs on xeric sites of the Blue Ridge Mountains [33].

Bear huckleberry is also found in the relatively high-elevation northern red oak (Q. rubra var. ambigua) and montane white oak forest types. A dense shrub layer of bear huckleberry, mountain-laurel, flame azalea, and highbush blueberry (Vaccinium corymbosum) is common in montane white oak forests. Mountain-laurel, black huckleberry (Gaylussacia baccata), hillside blueberry (V. pallidum), deerberry (V. stamineum), catawba rhododendron (R. catawbiense), and bear huckleberry also form a dense shrub layer in pine-oak/heath forests [28,33].

Coniferous forests:
In pitch pine-Table Mountain pine (P. rigida-P. pungens) forests, canopy species can include shortleaf pine (P. echinata), Virginia pine (P. virginiana), black locust (Robinia pseudoacacia), and several oak species. Mountain-laurel, mountain fetterbush (Pieris floribunda), blueberries (Vaccinium spp.), and huckleberries dominate the shrub layer in these forests [4,44].

Bear huckleberry is also common in the eastern hemlock (Tsuga canadensis) forests of North Carolina and is typically found in eastern hemlock/rosebay (R. maximum) communities [33].


SPECIES: Gaylussacia ursina
This description provides characteristics that may be relevant to fire ecology and is not meant for identification. Keys for identification are available [30,45].

Bear huckleberry is a deciduous native shrub that generally grows 3 feet (0.9 m) or greater in height [30]. The aboveground biomass can be great [40]. However, the rhizomes are likely confined to the 1st 3 to 6 inches (7.6-15.2 cm) of soil or duff [22],(Weakley, personal communication [42]). Bear huckleberry has alternate leaves that are elliptic, 2 to 4 inches (5-10 cm) long, and 0.79 to 1.8 inches (2- 4.5 cm) wide. Flowers are produced on 2-year-old wood [30]. The fruit is a 10-locule drupe with 1 seed per locule [17].


Bear huckleberry reproduces sexually and asexually.

Breeding system: Information pertaining to the reproductive biology of bear huckleberry is lacking. Bear huckleberry produces perfect flowers [29], and Renfro [32] successfully grew bear huckleberry from seed when performing tests of its susceptibility to ozone damage.

The degree to which bear huckleberry relies on sexual versus asexual reproduction is unknown. The following discussion of another huckleberry species suggests that reliance may be on one or the other regeneration strategy and may be disturbance dependent. Soil collected from mature oak forests of Montgomery County, Virginia, where black huckleberry dominated, was not found to contain any black huckleberry seed [34]. Likewise, research in the Atlantic Coastal Plains found vegetative reproduction was the only recruitment for black huckleberry over 3 years [26]. Matlack and others [26] suggest that an increased fire frequency in the Coastal Plains study area may explain the lack of sexual regeneration.

Pollination: Bear huckleberry flowers have a nectar disk on the surface of the ovary to reward insect visitors [29]. Bees and small butterflies are considered the chief huckleberry pollinators [23].

Seed dispersal: Animals may disperse bear huckleberry. Some suggest that the series of color changes that occur as huckleberry fruits ripen is a seed dispersal adaptation. It is thought that the color cues ensure that seed-dispersing wildlife remain in the area until fruits are ripe, increasing the probability of successful seed dispersal.  Huckleberry seeds typically remain viable after passage through animal seed dispersers [23].

Asexual regeneration: Bear huckleberry is strongly rhizomatous. New shouts are produced at approximately  1-foot (30.5 cm) intervals along the rhizome [42].

No literature addressed seed production, seed banking, germination, or seedling establishment and growth for bear huckleberry. Further research is needed.

In its range, bear huckleberry is common in mid- to high-elevation deciduous, coniferous, and mixed-woodland habitats. Elevational ranges are provided below by forest type.

Table Mountain-pitch pine 2,297 to 3,215 feet (700-980 m) [44]
Pine-oak/heath 2,500 to 5,000 feet (762-1,524 m) [33]
Pine/heath 3,400 feet (1,036 m)
Oak-hickory 4,000 feet (1,219 m) [4]
Mixed deciduous 2,297 to 2,953 feet (700-900 m) [9,10]

Soils: Forests soils where bear huckleberry is common are shallow, nutrient poor, acidic, and quick draining [4,33,44]. The pH of pine/heath forests of the Great Smoky Mountains of North Carolina and Tennessee range from 4.1 to 5.8 [4].

Climate: Temperatures associated with the mountainous regions of Tennessee, Georgia, and North and South Carolina are relatively mild. In a review of the environmental characteristics associated with Table Mountain pine forests, Della-Bianca [8] reported 170 to 180 frost-free days. Precipitation in this forest type ranges from a low of 27 inches (690 mm) to a high of 80 inches (2,030 mm) annually. In the Table Mountain pine-pitch pine forests, summer droughts are common [5,44]. In southwestern North Carolina mixed-deciduous forests, precipitation averages 71 inches (1,800 mm) annually. Growing season temperatures in this area average 65.3 F (18.5 C), and dormant-season temperatures average 44 F (6.7 C) [9,10].

While bear huckleberry can be present in early and late-seral stages of community development, it is considered dominant in mid-seral communities [5,16]. Bear huckleberry, northern red oak, and white oak typified mid-seral communities in the high rainfall region of the Nantahala National Forest of North Carolina [5]. Similarly, Gattis and others [16] report that bear huckleberry, oaks, and hickories dominate mid-seral communities in the same area. In both studies, these communities were at mid-elevations (2,600-3,300 feet (792-1,006 m)) and were intermediate along a xeric to mesic moisture scale. These findings, however, do not imply that bear huckleberry is not present in early seral and/or late-seral communities as well. Von Holle and Simberloff [40] report finding bear huckleberry in a closed-canopy forest of southwestern Virginia, where Big Stony Creek often floods the area, suggesting bear huckleberry may tolerate early seral conditions.

In southern Appalachian mixed-deciduous forests of North Carolina, sites were severely disturbed by clear-cutting vegetation, burning slash piles, planting grass seed, and treating emerging woody vegetation with herbicide. Twenty-eight years following these treatments, bear huckleberry was present at a frequency of 12% and at a density of 0.8 plant/m. On nearby undisturbed sites, aged as 70-year-old forests, bear huckleberry frequency was 38% and density was 6.0 plants/m [10].

The southern pine beetle creates gaps in southern Appalachian forests by killing pitch pine during stressful drought conditions. In the absence of fire, these historically pitch pine-dominated forests convert to oak-dominated forests. The density of huckleberries was recorded in undisturbed early seral pitch pine forests and in 3-, 8-, 11-, and 16-year-old forest gaps. The density of huckleberries in undisturbed forests and in the gaps is given below [36]:

Gap age

0 (Undisturbed)





Overstory density (trees/ha)




Huckleberry density (plants/ha)






Bear huckleberry typically flowers in May or June [30,45], and fruit ripens sometime from July to September [30].


SPECIES: Gaylussacia ursina
Fire adaptations: As of the writing of this review (2005), no literature specifically addressed bear huckleberry fire adaptations. It is likely that underground plant structures survive fire as bear huckleberry is often found on burned sites the first postfire growing season [7,18].

Fire regimes: Southern oak and pine forests are fire-adapted ecosystems and are highly impacted by changes in fire management. In these forests, lack of fire is often a chief explanation for poor regeneration of dominant pine and oak species. Historically, in montane oak-hickory and chestnut oak forests, low- to moderate-severity surface fires were common [33]. In mid-elevation (3,500-5,000 feet (1,067-1,524 m) northern red oak forests of the Blue Ridge Mountains, canopy removal by fire was uncommon, and the highest-elevation sites rarely burned. However, the typical xeric conditions and exposed landscapes occupied by northern red oak forests suggest the potential for large surface fires.

Literature cited by Elliott and others [10] indicates that prior to 1842, native Cherokee tribes burned southern Appalachian forests semiannually, a practice European settlers continued through the early 1900s. Fire exclusion in these forests, together with excessive shrub growth (including bear huckleberry), are suggested reasons for poor oak establishment. Without the creation of gaps in the forest canopy, oaks rarely progress from sapling to pole stage [9].

In presettlement time, Table Mountain pine-pitch pine forests of the southern Appalachians were likely confined to xeric sites but expanded during severe droughts that fueled high-severity fires. Expansion likely continued during the settlement of these forest areas, as Native Americans and European settlers burned the forests often. However, the absence of fire and changes in land use in the southern Appalachians have resulted in poor Table Mountain and pitch pine regeneration [44]. Some of the driest pine-, oak-, and heath-dominated communities experience frequent lightning. Fires in these forests allowed for shade-intolerant pines to regenerate [33]. Successful fire suppression in xeric southern Appalachian pine and pine-oak forests has resulted in mature, closed-canopy forests with decreased grass and forb cover. With fire, these forests have more open canopies, increased herbaceous cover, and increased species richness. Fire exclusion will probably change the way fires behave in these areas today. Fire sizes may be smaller and successful ignitions fewer in fire-suppressed communities [18]. After an extended period without fire, however, fuel loads build and eventually may result in larger, more severe fires.

The following table provides fire return intervals for plant communities and ecosystems where bear huckleberry is important. For further information, see the FEIS review of the dominant species listed below.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
yellow-poplar Liriodendron tulipifera < 35
shortleaf pine Pinus echinata 2-15
shortleaf pine-oak P. echinata-Quercus spp. < 10
Table Mountain pine P. pungens < 35 to 200 [41]
pitch pine P. rigida 6-25 [2,19]
eastern white pine P. strobus 35-200
eastern white pine-eastern hemlock P. strobus-Tsuga canadensis 35-200
eastern white pine-northern red oak-red maple P. strobus-Quercus rubra-Acer rubrum 35-200
Virginia pine P. virginiana 10 to < 35
Virginia pine-oak P. virginiana-Quercus spp. 10 to < 35
oak-hickory Quercus-Carya spp. < 35
southeastern oak-pine Quercus-Pinus spp. < 10
white oak-black oak-northern red oak Q. alba-Q. velutina-Q. rubra < 35
chestnut oak Q. prinus 3-8
northern red oak Q. rubra 10 to < 35
black oak Q. velutina < 35 [41]

Rhizomatous shrub, rhizome in soil
Ground residual colonizer (on-site, initial community)


SPECIES: Gaylussacia ursina
Clinton (personal communication [6]) researched fire effects in the southern Appalachian Mountains and indicated that bear huckleberry is top-killed by fire.

No additional information is available on this topic.

Bear huckleberry likely establishes on burned sites through rhizome and root crown sprouting. Research regarding its capacity to establish from seed following fire is unknown as of this writing, 2005. Bear huckleberry's presence in plant communities by the 1st postfire growing season suggests that sprouting or rapid seedling establishment of bear huckleberry occurs following fire. A fire researcher in North Carolina predicts that regeneration occurs through crown sprouting and considers postfire sprouting to be prolific in bear huckleberry (Clinton, personal communication [6]).

Information regarding the response of other huckleberries to fire may prove useful, as bear huckleberry fire research is limited. Research on the postfire response of black huckleberry indicates sprouting occurs from rhizomatous buds near the base of top-killed shoots [25]. Several huckleberries (black huckleberry, G. dumosa, and G. frondosa) showed increased fruit production in postfire growing seasons [11,20].

Postfire succession was studied in pine and pine-oak forests in the western portion of Great Smoky Mountains National Park. A mix of prescribed fires occurred in summer, fall, and winter, and the percentage of bear huckleberry basal area killed varied. Changes in bear huckleberry coverage were periodically evaluated for 22 years following the fires. The cover of bear huckleberry was much greater on unburned sites than on burned sites. Presented below is the percent cover of bear huckleberry following burning of multiple plots in 1976 to 1977 (only 1 plot burned in 1977). The coverage of bear huckleberry, averaged over all burned plots, was [18]:


Unburned sites

Burned sites


1977-78 1995 1977 1978 1979 1980 1984 1995

Bear huckleberry  cover (%) 

12.34 8.37 0.04 0.05 0.18 0.13 0.41 0.65

In the Nantahala National Forest, "degraded" 80-year-old stands of pine-hardwood forests were prescribed burned. All trees were cut in the summer of 1990 and cured for 44 to 89 days before fires were set in September. The average fuel moistures were between 28% and 33% at the time of burning. Forest floor and soil temperatures ranged between 113 F (45 C) and 140 F (60 C). Peak flame temperatures were between 1,157 F (625 C) and 1,477 F (803 C).  By the 3rd postfire growing season, burned sites had bear huckleberry cover similar to that of prefire coverage. However, bear huckleberry density was double that of the prefire community. The difference in pre- and postfire coverages and densities were not analyzed statistically. The postfire response of bear huckleberry is given below [7]:

  Prefire 1991 1992 1994
Density (stems/m) 0.5 0.6 0.6 1.3
Cover (%) 0.4 0.5 0.4 0.4

The information regarding fire effects on bear huckleberry is sparse. The use of fire for the management of bear huckleberry requires more information than is available at this time. If and when available, pre- and postfire vegetation responses for this species should be assessed and results be made available to managers.


SPECIES: Gaylussacia ursina
Several eastern forest mammals eat the sweet huckleberry fruits produced in mid- to late summer. Huckleberries are considered an important food source for grouse and white-tailed deer [38]. The low stature of bear huckleberry suggests that several small mammals may feed on its fruits. Common gray foxes, red foxes, raccoons, black bears, eastern fox squirrels, and white-footed mice typically feed on huckleberry fruits. Bird species also likely rely on bear huckleberry fruits. Since insects normally hatch in spring and early summer, the mid- to late-summer fruit production provides a valuable food source for birds [23]. Research in Alabama indicates huckleberries were a major component of wild turkey summer diets [12]. Although this study was conducted outside bear huckleberry's range, wild turkeys are found within the range of bear huckleberry and likely feed on its fruits.

Palatability/nutritional value: Huckleberry fruits are palatable to many mammal species [23]. No specific literature was found regarding bear huckleberry use by wildlife or livestock.

Cover value: The dense shrub layer associated with montane white oak and pine-oak/heath forests [33] likely provides cover for wildlife.

A dense shrub community including huckleberries has successfully colonized power line right-of-ways in eastern abandoned fields. This suggests bear huckleberry may be valuable in revegetating disturbed sites. It is important to note, however, that these dense shrub communities may discourage the growth and establishment of canopy vegetation [23].

Bear huckleberry was an immediate and preserved food source utilized by Native Americans in the Cherokee tribe [27].

OTHER MANAGEMENT CONSIDERATIONS: More information regarding the specific use of bear huckleberry by eastern wildlife species is needed.

Gaylussacia ursina: References

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2. Buchholz, Kenneth; Good, Ralph E. 1982. Density, age structure, biomass and net annual aboveground productivity of dwarfed Pinus rigida Moll. from the New Jersey Pine Barren Plains. Bulletin of the Torrey Botanical Club. 109(1): 24-34. [8639]

3. Cain, Stanley A. 1930. An ecological study of the heath balds of the Great Smoky Mountains. Butler University Botanical Studies: Paper No. 13. Indianapolis, IN: Butler University. 1: 77-208. [22935]

4. 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]

5. Carter, Robert E., Jr.; Shelburne, Victor B. 1995. Landscape ecosystem classification of successional forest communities in the southern Appalachians. In: Edwards, M. Boyd, compiler. Proceedings, 8th biennial southern silvicultural research conference; 1994 November 1-3; Auburn, AL. Gen. Tech. Rep. SRS-1. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station: 46-50. [27256]

6. Clinton, Barry. 2005. [Email to Corey Gucker]. February 9. Bear huckleberry information request. Otto, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. On file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT: RWU 4403. [51539]

7. Clinton, Barton D.; Vose, James M. 2000. Plant succession and community restoration following felling and burning in the southern Appalachian Mountains. In: Moser, W. Keith; Moser, Cynthia F., eds. Fire and forest ecology: innovative silviculture and vegetation management: Proceedings of the 21st Tall Timbers fire ecology conference: an international symposium; 1998 April 14-16; Tallahassee, FL. No. 21. Tallahassee, FL: Tall Timbers Research, Inc: 22-29. [37602]

8. Della-Bianca, Lino. 1990. Pinus pungens Lamb. table mountain pine. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Volume 1. Conifers. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 425-432. [13400]

9. Elliott, K. J.; Swank, W. T. 1994. Impacts of drought on tree mortality and growth in a mixed hardwood forest. Journal of Vegetation Science. 5: 229-236. [23616]

10. Elliott, Katherine J.; Boring, Lindsay R.; Swank, Wayne T. 1998. Changes in vegetation structure and diversity after grass-to-forest succession in a southern Appalachian watershed. The American Midland Naturalist. 140(2): 219-232. [30080]

11. Elliott, Katherine J.; Hendrick, Ronald L.; Major, Amy E.; [and others]. 1999. Vegetation dynamics after a prescribed fire in the southern Appalachians. Forest Ecology and Management. 114(2-3): 199-213. [30079]

12. Exum, Jay H.; McGlincy, Joe A.; Speake, Dan W.; [and others]. 1987. Ecology of the eastern wild turkey in an intensively managed pine forest in southern Alabama. Bull. 23. Tallahassee, FL: Tall Timbers Research Station. 70 p. [17990]

13. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]

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15. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 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]

16. Gattis, Jane T.; Jones, Steven M.; McNab, W. Henry. 1993. Landscape ecosystem classification on the Highlands Ranger District, Nantahala National Forest, North Carolina. In: Brissette, John C., ed. Proceedings, 7th biennial southern silvicultural research conference; 1992 November 17-19; Mobile, AL. Gen. Tech. Rep. SO-93. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station: 223-228. [23267]

17. Gleason, Henry A.; Cronquist, Arthur. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York: New York Botanical Garden. 910 p. [20329]

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19. Hendrickson, William H. 1972. Perspective on fire and ecosystems in the United States. In: Fire in the environment: Symposium proceedings; 1972 May 1-5; Denver, CO. FS-276. [Washington, DC]: U.S. Department of Agriculture, Forest Service: 29-33. In cooperation with: Fire Services of Canada, Mexico, and the United States; Members of the Fire Management Study Group; North American Forestry Commission; FAO. [17276]

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22. Klein, Rob. 2005. [Email to Corey Gucker]. January 20. Bear huckleberry information request. Gatlinburg, TN: U.S. Department of the Interior, Great Smoky Mountains National Park. On file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT: RWU 4403. [51540]

23. Kricher, John C.; Morrison, Gordon. 1988. A field guide to eastern forests: North America. Peterson Field Guide Series 37. Boston: Houghton Mifflin Company. 368 p. [22677]

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25. Matlack, Glenn R. 1997. Resource allocation among clonal shoots of the fire-tolerant shrub Gaylussacia baccata. Oikos. 80(3): 509-518. [29359]

26. Matlack, Glenn R.; Gibson, David J.; Good, Ralph E. 1993. Regeneration of the shrub Gaylussacia baccata and associated species after low-intensity fire in an Atlantic coastal plain. American Journal of Botany. 80(2): 119-126. [20726]

27. Moerman, Dan. 1999. Native American ethnobotany database: Foods, drugs, dyes, and fibers of native North American peoples, [Online]. Available: http://www.umd.umich.edu/cgi-bin/herb/ [2003, March 2]. [37492]

28. Nelson, John B. 1986. The natural communities of South Carolina. Columbia, SC: South Carolina Wildlife & Marine Resources Department. 54 p. [15578]

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31. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]

32. Renfro, James R. 1989. Evaluating the effects of ozone on the plants of Great Smoky Mountains National Park. Park Science. 9(4): 22-23. [9337]

33. Schafale, Michael P.; Weakley, Alan S. 1990. Classification of the natural communities of North Carolina: Third approximation. Raleigh, NC: Department of Environment, Health, and Natural Resources, Division of Parks and Recreation, North Carolina Natural Heritage Program. 325 p. Available: http://ils.unc.edu/parkproject/nhp/publications/class.pdf [2005, February 14]. [41937]

34. Schiffman, Paula M.; Johnson, W. Carter. 1992. Sparse buried seed bank in a southern Appalachian oak forest: implications for succession. The American Midland Naturalist. 127(2): 258-267. [18191]

35. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]

36. Smith, Robert Nolan. 1991. Species composition, stand structure, and woody detrital dynamics associated with pine mortality in the southern Appalachians. Athens, GA: University of Georgia. 163 p. Thesis. [51018]

37. Stickney, Peter F. 1989. FEIS postfire regeneration workshop--April 12: Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. 10 p. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. [20090]

38. Thackston, Reginald E.; Hale, Philip E.; Johnson, A. Sydney; Harris, Michael J. 1982. Chemical composition of mountain-laurel Kalmia leaves from burned and unburned sites. Journal of Wildlife Management. 46(2): 492-496. [9076]

39. U.S. Department of Agriculture, National Resource Conservation Service. 2005. PLANTS database (2004), [Online]. Available: http://plants.usda.gov/. [34262]

40. Von Holle, Betsy; Simberloff, Daniel. 2004. Testing Fox's assembly rule: does plant invasion depend on recipient community structure? Oikos. 105(3): 551-563. [48592]

41. Wade, Dale D.; Brock, Brent L.; Brose, Patrick H.; [and others]. 2000. Fire in eastern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 53-96. [36983]

42. Weakley, Alan. 2005. [Email to Corey Gucker]. February 23. Bear huckleberry information request. Chapel Hill, NC: University of North Carolina, North Carolina Botanical Garden, Herbarium. On file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT: RWU 4403. [51884]

43. Whittaker, R. H. 1956. Vegetation of the Great Smoky Mountains. Ecological Monographs. 26(1): 1-79. [11108]

44. Williams, Charles E. 1998. History and status of Table Mountain pine-pitch pine forests of the southern Appalachian Mountains (USA). Natural Areas Journal. 18(1): 81-90. [27900]

45. Wofford, B. Eugene. 1989. Guide to the vascular plants of the Blue Ridge. Athens, GA: The University of Georgia Press. 384 p. [12908]

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