Fire Effects Information System (FEIS)

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Galax urceolata

• INTRODUCTORY
• DISTRIBUTION AND OCCURRENCE
• BOTANICAL AND ECOLOGICAL CHARACTERISTICS
• FIRE ECOLOGY
• FIRE EFFECTS
• MANAGEMENT CONSIDERATIONS
• REFERENCES

INTRODUCTORY

DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• ECOSYSTEMS
• STATES/PROVINCES
• BLM PHYSIOGRAPHIC REGIONS
• KUCHLER PLANT ASSOCIATIONS
• SAF COVER TYPES
• SRM (RANGELAND) COVER TYPES
• HABITAT TYPES AND PLANT COMMUNITIES

There are 2 races of beetleweed. Their distributions mostly overlap. See General Botanical Characteristics and Site Characteristics for details on morphologies and habitats, respectively, distinguishing beetleweed races.

Beetleweed is a common understory species of oak-hickory forests in the east-central and southern United States. It most commonly associates with chestnut oak and scarlet oak (Q. coccinea), respectively [1,59]. Chestnut oak (Quercus prinus) is beetleweed's most frequent overstory dominant across beetleweed's distribution [1]. In a mixed-oak forest on the Jefferson National Forest of Virginia, overstory dominants of beetleweed are chestnut oak, scarlet oak, and red maple (Acer rubrum). Ericaceous shrubs including black huckleberry (G. baccata) and Blue Ridge blueberry (Vaccinium pallidum) dominate the shrub layer. American alumroot (Heuchera americana) is a common forb (review by [59]). Beetleweed is an important component of the chestnut oak/mountain-laurel community in South Carolina's Jocassee Gorges. Bear huckleberry (Gaylussacia ursina), Piedmont rhododendron (Rhododendron minus), and fairywand (Chamaelirium luteum) are other important species [1]. Besides chestnut and scarlet oak, other oak-hickory forest dominants sometimes associated with beetleweed include blackjack oak (Quercus marilandica), post oak (Q. stellata), northern red oak (Quercus rubra), white oak (Q. alba), black oak (Q. velutina), scarlet oak, southern red oak (Q. falcata), turkey oak (Q. laevis), pignut hickory (C. glabra), black hickory (C. texana), and mockernut hickory (C. tomentosa). Understory tree and shrub associates include flowering dogwood (Cornus florida), blueberries (Vaccinium spp.), huckleberries (Gaylussacia spp.), and sumacs (Rhus spp.). Herbaceous plant associates include bluestems (Andropogon spp.), little bluestem (Schizachyrium scoparium), and sedges (Carex spp.) [21,29,31,72].

Beetleweed was important in preblight American chestnut (Castanea dentata) stands. In a stump survey reconstructing early 20th century American chestnut riparian forests of the Blue Ridge Mountains, rosebay dominated former American chestnut stands disturbed by blight and harvest of infected American chestnuts in the 1920s and 1930s. Beetleweed and partridgeberry (Mitchella repens) were abundant in the rosebay thickets, and other herbaceous species were nearly absent [68]. Postblight American chestnut forests have mostly succeeded to rosebay thickets, mountain-laurel thickets, or oak-hickory forest [19,45,68].

Beetleweed is a common herbaceous associate of xeric oak-pine forest. Percent cover of beetleweed ranges from 5% to 20% in this forest type. Oak-pine forests are common on south-facing slopes in the central and southern Appalachian, Piedmont, and Coastal Plain regions. A 1935 publication noted that beetleweed was common in a shortleaf pine (Pinus echinata)-chestnut oak community on Pine Mountain, Kentucky. Understory shrubs included black huckleberry, Blue Ridge blueberry, and mountain-laurel. Narrowleaf silkgrass (Pityopsis graminifolia), downy danthonia (Danthonia sericea), and partridgeberry (Mitchella repens) were common herbaceous associates [6].

Within oak-pine forests, chestnut oak is the most common overstory associate of beetleweed; scarlet oak is 2nd. Dominant pines include pitch pine (P. rigida), Table Mountain pine (P. pungens), and/or Virginia pine (P. virginiana). In a review, Murphy and Nowacki [46] noted beetleweed as important in high-elevation, old-growth Table Mountain pine communities in the Great Smoky Mountains. Pitch pine, scarlet oak, chestnut oak, and black tupelo (Nyssa sylvatica) were common overstory associates; mountain-laurel and blueberries dominated the understory. With 5% to 20% coverage, beetleweed, trailing arbutus (Epigaea repens), and eastern teaberry (Gaultheria procumbens) dominated groundlayer vegetation [46].

Beetleweed is a common understory component of northern hardwood forests. These forests are generally found at mid- to high elevations in the central and northern Appalachian Mountains, often transitioning to spruce-fir or mixed hardwood forest at higher and lower elevations, respectively [45]. Common overstory tree species include sugar maple (Acer saccharum), basswood (Tilia americana), yellow birch (B. alleghaniensis), black cherry (Prunus serotina), red spruce (Picea rubens), white spruce (P. glauca), American beech (Fagus grandifolia), eastern white pine (Pinus strobus), eastern hemlock (Tsuga canadensis), northern red oak, white oak, and yellow-poplar (Liriodendron tulipifera). Understory associates include beaked hazel (Corylus cornuta), eastern leatherwood (Dirca palustris), red elderberry (Sambucus racemosa var. racemosa), alternate-leaf dogwood (Cornus alternifolia), bush-honeysuckle (Diervilla lonicera), Canada yew (Taxus canadensis), red raspberry (Rubus idaeus), and blackberries. Herbaceous species include Carolina springbeauty (Claytonia caroliniana), snow trillium (Trillium grandiflorum), anemones (Anemone spp.), marsh blue violet (Viola cucullata), downy yellow violet (V. pubescens), hairy Solomon's seal (Polygonatum pubescens), starry Solomon's-seal (Maianthemum stellatum), hairy sweet-cicely (Osmorhiza claytonii), adderstongues (Ophioglossum spp.), Jack-in-the pulpit (Arisaema triphyllum), bigleaf aster (Eurybia macrophylla), and clubmosses (Lycopodiaceae) [45,58,72].

Beetleweed is a groundlayer species in mixed hardwood forests. These forests often support a high level of plant diversity [29]. Overstory hardwood and conifer associates of beetleweed are numerous within the type. They include northern red oak, white oak, black oak, scarlet oak, southern red oak, post oak, yellow-poplar, eastern white pine, American beech, sugar maple, red maple, black cherry, American basswood, sweetgum (Liquidambar styraciflua), white ash (Fraxinus americana), green ash (F. pennsylvanica), quaking aspen (Populus tremuloides), hickories, black tupelo, black walnut (Juglans nigra), jack pine (Pinus banksiana), eastern hemlock, and elms (Ulmus spp.). Common mid-canopy tree associates include flowering dogwood, hollies (Ilex spp.), eastern hophornbeam (Ostrya virginiana), sassafras (Sassafras albidum), American bladdernut (Staphylea trifolia), eastern redbud (Cercis canadensis), common persimmon (Diospyros virginiana), and serviceberries (Amelanchier spp.). Common understory shrubs and vines include greenbriers (Smilax spp.), blueberries, rosebay, eastern leatherwood, witch-hazel (Hamamelis virginiana), beaked hazel, spicebush (Lindera benzoin), poison-ivy (Toxicodendron radicans), and grapes (Vitis spp.) [9,45,58,72].

Beetleweed frequents openings or open stands of spruce-fir (Picea-Abies spp.) forest in the central and southern Appalachian arboreal highlands, mountain tops, and "balds" [9,58]. Southern spruce-fir forests are dominated by red spruce, which mixes with hardwoods on mid-elevation slopes. Common overstory associates include Fraser fir (A. fraseri), yellow buckeye (Aesculus flava), sweet birch (Betula lenta), and black cherry. Rhododendrons (Rhododendron spp.), American mountain-ash (Sorbus americana), and possumhaw (Viburnum nudum var. cassinoides) are common understory dominants. Other shrub associates include highbush cranberry (V. edule), mountain holly (I. montana), speckled alder (Alnus rugosa), pin cherry (Prunus pensylvanica), serviceberries, raspberries (Rubus spp.), blueberries, and huckleberries [58].

Beetleweed occurs in the groundlayer of Carolina hemlock (Tsuga caroliniana) forest. Overstory associates include red maple, chestnut oak, and sweet birch. Understory vegetation is not diverse in the type [57]. Mountain-laurel and rosebay are common shrubs. Groundlayer vegetation includes vines such as Virginia creeper (Parthenocissus quinquefolia) and partridgeberry, and forbs including beetleweed, Virginia heartleaf (Hexastylis virginica), and Carolina silverbell (Halesia carolina) [57,58,72].

Beetleweed is occasional on upland and mesic sites within longleaf pine (P. palustris) forests and savannas in and along the Atlantic and Gulf coastal plains and lower Piedmont regions of Georgia and Alabama. Associated species on mesic coastal plains include southern red oak, blackjack oak, water oak, flowering dogwood, black tupelo, sweetgum, common persimmon, and sassafras. Associated species on xeric sandhill sites include turkey oak, bluejack oak (Q. incana), and live oak (Q. virginiana). Associated shrubs include inkberry (I. glabra), yaupon (I. vomitoria), large gallberry (I. coriacea), wax-myrtle (Myrica cerifera), blueberries, huckleberries, blackberries, saw-palmetto (Serenoa repens), sweetbay (Magnolia virginiana), cyrilla (Cyrilla racemiflora), and buckwheat tree (Cliftonia monophylla). Pineland threeawn (Aristida stricta) is the primary groundcover on longleaf pine sites within beetleweed's distribution [14,50,58,72].

Shrub balds often occupy the highest (>4,000 feet (1,200 m)) mountain peaks in the central and southern Appalachian Mountains. Dense thickets of usually ericaceous shrubs dominate. Bald dominants and associates across beetleweed's range include mountain-laurel, Catawba rosebay (Rhododendron catawbiense), highbush blueberry (Vaccinium corymbosum), black chokeberry (Photinia melanocarpa), mountain sweetpepperbush (Clethra acuminata), mountain holly, possumhaw, blackberries, and American mountain-ash [9,58,72]. In South Carolina, beetleweed is common to codominant on high-elevation mountain-laurel balds [1]. In North Carolina, it forms a shrub/forb community with silvery nailwort (Paronychia argyrocoma) on the summit of King's Pinnacle, just above bear oak-dominated shrubland [4].

The following vegetation typings describe plant communities where beetleweed is a dominant or important component of the ground layer.

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

• GENERAL BOTANICAL CHARACTERISTICS
• RAUNKIAER LIFE FORM
• REGENERATION PROCESSES
• SITE CHARACTERISTICS
• SUCCESSIONAL STATUS
• SEASONAL DEVELOPMENT

Beetleweed is a perennial forb or subshrub that reaches 2 to 12 inches (6-30 cm) in height [41,73]. Its evergreen leaves are heart-shaped, thick, and glossy. They are singular, arising from a 1- to 10-inch (3-25 cm)- long petiole [52,63]. Leaf life span is 18 months [56]. Numerous small perfect flowers are 3 to 4 mm wide, and are arranged in a spike-shaped raceme on an 8- to 16-inch (20-40 cm) flower stem. Seeds are <1 mm long. They are contained in a small capsule. Each capsule contains "several to numerous" seeds. Beetleweed is rhizomatous. Roots are fibrous [52,63].

Beetleweed has 2 races, diploid and tetraploid, that are morphologically distinct [2,3,64]. Leaf morphology differs geographically and genetically among populations, with mountainous and southern populations (which are mostly tetraploid) having large leaves, and northern populations (mostly diploid) having small leaves [2,30]. Under best growing conditions (see Site Characteristics), leaves of diploid beetleweed are 4 inches (10 cm) across, while tetraploid beetleweed's leaves may reach 6 inches (20 cm) across. On poor sites, the 2 races may both have small leaves and be indistinguishable in the field [2].

Breeding system: Beetleweed is monoecious [41]. In a 1992 to 1999 survey, Greller and Clemants [30] noted beetleweed establishment and population expansion by seed spread on Long Island, New York, where beetleweed was previously uncollected and is probably not native.

Polyploidy may give beetleweed greater ecological amplitude than a simple diploid state [9,47]. Although geographic distributions of the 2 races overlap, they apparently do not cross-breed [47].

Seed dispersal: Beetleweed's small, light seed is "easily transported" [9]. Mechanisms for transport are not described in current (2006) literature.

Seed bank: The degree to which beetleweed relies on a seed bank is unknown. In seed bank study conducted in a chestnut oak-scarlet oak forest on the Jefferson National Forest, beetleweed seed was neither visually apparent in soil samples, nor did it germinate from soil samples in the greenhouse. However, beetleweed was a dominant forb in the study area [59]. Methodologies can affect seed bank trials [61], and absence of beetleweed emergence in seed bank studies does not mean that beetleweed does not form a seed bank. Further studies are needed on seed ecology of beetleweed.

Asexual regeneration: Beetleweed reproduces asexually [48] from rhizome sprouts [51,52,55,63].

As of 2006, published literature describing pollination, seed dispersal, seed production, seed banking, germination, seedling establishment, or growth for beetleweed was lacking. Further research is needed on beetleweed's life history.

Elevation: Precise elevational data are sparse for beetleweed. However, beetleweed is reported on sites with a wide elevational range, from low-elevation coastal plains and highest-elevation mountain peaks. In Virginia, for example, it is reported on low-elevation coastal plains (~0-70 feet (20 m)) and at 1,940 feet (591 m) in a Carolina hemlock forest [2,57]. Beetleweed has been collected on the highest peaks of the Appalachian Mountains: 5,964 feet (1,818 m) on Grandfather Mountain in North Carolina and 5,200 feet (1,600 m) on Mt. LeConte in Tennessee [2].

Tetraploid populations of beetleweed occur throughout the distribution of diploid beetleweed. They also occur on the Virginia coastal plain, where diploid populations do not occur. Additionally, tetraploid beetleweed are more common at high elevations than diploid beetleweed [2].

Soils: Beetleweed commonly occurs on rocky or sandy acidic soils on slopes, ridges, and mountain hillsides [74]. Best growth occurs on moist, acidic soils, although beetleweed occurs on dry soils in chestnut oak and a few other habitats (review by [25]). In Kentucky, beetleweed has been noted on soils with a pH as low as 3.9 [6].

Climate: Considerable climatic diversity is found in beetleweed's range. Climate ranges from subtropical along the southeastern coastal plains to temperate further inland. In general, temperature, precipitation, and length of growing season increase to the south. However, a wide variety of local microclimatic conditions exist in the complex topography of the Appalachian mountain region. Seasonal weather patterns are driven by alternating cold/dry continental air masses from Canada and warm/moist air from the Gulf of Mexico. Precipitation is generally distributed uniformly throughout the year, mostly as rain. Snow and ice are common in the winter months in beetleweed's northern range and high-elevation mountainous terrain [14,29]. Mean annual precipitation ranges from 39 to 80 or more inches (990-2,000 mm) in the Great Smoky Mountains [9]. Depending on location, annual snow accumulations range from 8 to 48 inches (200-1,220 mm). Tropical cyclones are possible in summer and fall months. Seasonal variations in temperature increase away from the coast. Mean winter temperatures vary from -18 °F (-28 °C) on high-elevation sites and in the north [9] to 64 °F (18 °C) in beetleweed's southern range. Mean summer temperatures are less variable, ranging from 70 to 72 °F (21-22 °C) [14,29].

Early succession: A study of forest succession in mixed conifer-hardwood riparian forests of the southern Appalachians noted beetleweed's rarely documented importance in early succession. With 69% frequency, beetleweed was the herbaceous dominant in early seral yellow-poplar-red maple-mountain magnolia (Magnolia fraseri) stands on the Thompson River corridor of North Carolina. It was also present but less frequent on 2 other North Carolina sites in late succession: an eastern white pine-white oak forest and an eastern hemlock-white oak forest (22% and 12% frequency, respectively) [1].

Beetleweed occurred relatively soon after extreme disturbances in the Nantahala Mountains of southwestern North Carolina pushed a forest into early succession. On the Wine Spring Creek Watershed of what is now the Coweeta Hydrologic Laboratory, a former American chestnut forest was "heavily" logged from 1912 to 1923. The forest succeeded to mixed oak after chestnut blight infestations in the early 1920s. Further logging and a type conversion followed decades later. A riparian corridor was partially logged in 1941; after that, treatments involved the entire watershed. The watershed was clearcut and pile burned in 1958; planted to sixweeks grass (Vulpia octoflora, a native annual) in 1959; repeatedly treated with 2,4-D from 1960 to 1965 to suppress woody shrubs including beetleweed; fertilized in 1965; treated with atrazine (to kill grass) and paraquat and 2,4-D (to kill shrubs) in 1967; then left undisturbed. Plots (0.02-ha) were sampled in 1995 to determine understory recovery, using adjacent undisturbed plots (70 years since last harvest) as controls. Forbs dominated the disturbed understory in 1995; red maple and shrubs, including beetleweed, dominated the undisturbed understory. Although method of postdisturbance establishment was not part of the study, beetleweed establishment after 1967 was probably from seed because several successive herbicide treatments in the 1960s would have killed most mature plants. Beetleweed distribution on undisturbed and disturbed plots was [19]:

Mid-succession: In another Coweeta Hydrologic Laboratory study, Elliott and others [20] found beetleweed recovered gradually after a chestnut oak-scarlet oak-pitch pine stand was clearcut in 1952. Beetleweed was the 2nd most common herbaceous species in the study area before clearcutting, representing 19% of total understory species composition. Forty-one years later, beetleweed showed 3rd greatest percent relative biomass compared to other understory species. Pre- and postharvest measurements were in different units. Beetleweed's relative biomass was [20]:

In vegetation surveys of reclaimed surface coal mines of southwestern Virginia, beetleweed occurred only on the oldest (>35 years) reclamation sites. The sites were planted with nonnative grasses ands legumes, native black locust (Robinia pseudoacacia), and eastern white pine, which is native to the general region but not to the reclamation sites [39]. Nonnative plantings probably altered the successional trajectory. Without comparative studies, it is difficult to access beetleweed's successional position on coal mine sites.

Some heath balds where beetleweed is common to dominant in the understory are successional to chestnut oak, conifer, or oak-pine forest without fire or other disturbance. Depending upon time-since-fire or other disturbance, the heathlands vary from open to very dense (review by [25]), so degree of beetleweed exposure on balds could vary greatly. Beetleweed's successional role in heath balds is not described in current (2006) literature.

Late succession: Beetleweed presence has been noted in heavily shaded, old-growth mixed- and northern hardwood forests [45,58,72].

FIRE ECOLOGY

• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

Laboratory and field experiments suggest that extreme heat or cold shock during flowering can induce polyploidy [54,64]. Baldwin [2] suggested that repeated fire disturbances and/or glaciation may have played an evolutionary role in the development of beetleweed tetraploidy.

Fire regimes: Fire was historically important in maintaining oak-pine and high-elevation, xeric pine forests where beetleweed occurs [46,58,71]. In the Appalachian Mountains, where beetleweed is most common, such communities experienced both stand-replacing and mixed-severity fires. In Great Smoky Mountain National Park, occurrence of both anthropogenic and natural fires increased with elevation, and lightning fires were most frequent on xeric, high-elevation sites [5,33] where beetleweed is common [13,58]. Whittaker [72] suggested that xeric, even-aged pine forests of the Great Smoky Mountains were mostly maintained by severe fire that replaced the existing stand; however, as the even-aged stand matured, pine mortality created small openings where gap succession occurred. He also stated the "fire alone does not produce these pine stands" [72]. Hurricanes, tornadoes, and less severe wind storms also maintained seral pine and oak-pine stands [46,65]. Time between stand-replacing fires varied between sites and among pine species.

Some pine and oak-pine communities dominated by serotinous pines have moderate to long fire-return intervals. Table Mountain pine historically experienced medium-return interval, stand-replacement fires [66]. Barden [3] documented a Table Mountain pine stand on Glass Mountain, North Carolina, that established after a fire 87 years previously. Fire may be very infrequent on some sites: Zobel [74] suggests that Table Mountain pine stands on rocky outcrops or shale slopes, where hardwood regeneration is poor, are self-sustaining, and rarely if ever experience fire. On sites where succession proceeds to understory hardwoods, understory burning probably helped keep the hardwoods in check [46].

In a 1935 survey of a shortleaf pine-chestnut oak community in the Cumberland Mountains of Kentucky, Braun [6] noted and 0.5- to 1-inch (cm) duff layer of pine, oak, and mountain-laurel leaves.

Mesic pine forests: Not all pine forests where beetleweed occurs have stand-replacement fire regimes. Longleaf and other nonserotinous pines historically experienced frequent surface fires [70]. Peet and Allard [50] state that longleaf pine-blackjack oak/mountain-laurel/beetleweed communities of North Carolina require "exceptionally high fire frequency." Occurring on mesic, north-facing slopes, these communities are successionally replaced relatively quickly when fire is excluded [50].

Heath balds where beetleweed is common to dominant in the understory were apparently maintained by stand-replacing fire, landslides, extreme weather conditions, and/or windthrow in presettlement times. Logging in the 20th Century stopped tree invasion onto the balds. Possibly due to highly acidic soils, some balds appear stable despite lack of disturbance such as fire [10,25,72]. On other and possibly most balds, however, repeated fires may have been the primary disturbance that maintained the shrubland [7,25]. Cain [10] found that all of the heath balds he sampled in the Great Smoky Mountains had burned multiple times. For heath bald types, fires in balds undergoing successional replacement by red spruce are probably most severe [25].

The following table provides fire return intervals for plant communities and ecosystems where beetleweed is important. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".

FIRE EFFECTS

• IMMEDIATE FIRE EFFECT ON PLANT
• DISCUSSION AND QUALIFICATION OF FIRE EFFECT
• PLANT RESPONSE TO FIRE
• DISCUSSION AND QUALIFICATION OF PLANT RESPONSE
• FIRE MANAGEMENT CONSIDERATIONS

Several studies note beetleweed presence on burns [9,16,22,23]. For example, beetleweed occurred with 1% to 2% frequency and 32% cover in an "old" and a "recent" burn in the Great Smoky Mountains of North Carolina and Tennessee. Time-since-fire was not determined for the old burn; the recent burn occurred "several years" before the 1929 survey [9].

Beetleweed can be slow to establish after fire relative to associated species. To increase the pine component and overall diversity of a mixed pitch pine-scarlet oak-chestnut oak/mountain-laurel forest, broadcast prescribed burning and white pine plantings were done following clearcutting on the Nantahala National Forest of North Carolina. Clearcutting occurred in 1990 and was finished by late July; prescribed burning was conducted about 2 months later on 18 and 19 September 1990. The fire consumed the forest litter and fine woody material. Large woody debris in the burn's interior was "consumed or reduced." Beetleweed's postfire frequency and height were low compared to 27 and 31 other herbs present on 2 study plots. Beetleweed mean biomass, height, and density on 2 study sites are given below. Plots were 0.05 ha (n=5); plots on the 2 sites were measured in successive years [23]:

After a felling and burning treatment in a chestnut oak-pitch pine/mountain-laurel forest on the Nantahala National Forest, beetleweed exceeded pretreatment density and percent cover levels 1 year following treatments. The site was clearcut in summer 1990; vegetation left to cure for 44 to 89 postharvest days; then burned under prescription in September 1990. Four years after treatment beetleweed abundance decreased substantially, presumably due to interference from other plant species. Because initial posttreatment recovery was rapid, the authors presumed that beetleweed's postfire growth was from rhizome sprouts. Density and percent cover of beetleweed were as follows [16]:

Beetleweed showed mixed response after short-interval, repeat prescribed burning near the Green River of North Carolina. Prescribed burning was conducted in a Table Mountain-pitch pine/mountain-laurel stand. There were 5 treatments: an unburned control, a single burn, and 2-, 3-, and 4-repeat burns. Repeat-burn plots were subjected to fire every 3 to 4 years. Beetleweed declined on 2- and 4-burn plots, but increased on 3-burn plots. Importance value of beetleweed on each treatment was [53]:

Beetleweed is noted as occurring in deeply shaded, late-successional forest [13,19,35,74], although whether beetleweed is most favored in long-unburned or otherwise undisturbed forest is unclear. In a postfire succession study in Great Smoky Mountains National Park, Harrod and others [35] found that beetleweed was more abundant in xeric mixed Virginia pine-pitch pine-scarlet oak stands that had not burned for over 60 years compared to stands that had burned more recently. Beetleweed density was 3 to 4 times greater on the oldest pine-oak burn plots (>60 years since fire) compared to younger pine-oak burn plots. With slightly more than 1% cover, beetleweed was the most abundant herbaceous species present on the oldest burns. Mean percent cover of beetleweed on permanent plots was [35]:

The following Research Project Summaries provide further information on prescribed fire use and postfire response of plant community species including beetleweed:

• Early postfire effects of a prescribed fire in the southern Appalachians of North Carolina
• Early postfire response of southern Appalachian Table Mountain-pitch pine stands to prescribed fires in North Carolina and Virginia

Beetleweed leaves may help generate severe fires that provide important ecological benefits. Turrill and others [66] report that in Table Mountain/mountain-laurel/beetleweed communities on the Chattahoochee National Forest of Georgia, groundlayer beetleweed and blueberries (Vaccinium spp.) promote "hot" surface fires that remove even deep build-ups of litter and organic matter under dry conditions. In turn, tall mountain-laurel shrubs can become ladder fuels that carry beetleweed and blueberry-fueled surface fires up to the serotinous cones of Table Mountain pine, aiding in the pine's postfire regeneration [66].

MANAGEMENT CONSIDERATIONS

• IMPORTANCE TO LIVESTOCK AND WILDLIFE
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER USES
• OTHER MANAGEMENT CONSIDERATIONS

Native Americans traditionally used beetleweed for treatment of kidney ailments [41].

Galax urceolata: REFERENCES

1. Abella, Scott R.; Shelburne, Victor B. 2004. Ecological species groups of South Carolina's Jocassee Gorges, southern Appalachian Mountains. Journal of the Torrey Botanical Society. 131(3): 220-231. [54882]

2. Baldwin, J. T., Jr. 1941. Galax: the genus and its chromosomes. Journal of Heredity. 32: 249-254. [60214]

3. Barden, L. S. 1977. Self-maintaining populations of Pinus pungens Lam. in the southern Appalachian Mountains. Castanea. 42: 316-323. [17652]

4. Barden, Lawrence S. 2000. A common species at the edge of its range: conservation of bear oak (Quercus ilicifolia) and its low elevation rocky summit community in North Carolina. Natural Areas Journal. 20(1): 85-89. [34902]

5. Barden, Lawrence S.; Woods, Frank W. 1974. Characteristics of lightning fires in southern Appalachian forests. In: Proceedings, annual Tall Timbers fire ecology conference; 1973 March 22-23; Tallahassee, FL. No. 13. Tallahassee, FL: Tall Timbers Research Station: 345-361. [19012]

6. Braun, E. Lucy. 1935. The vegetation of Pine Mountain, Kentucky: an analysis of the influence of soils and slope exposure as determined by geological structure. The American Midland Naturalist. 16(4): 517-565. [54879]

7. Brown, Dalton Milford. 1941. Vegetation of Roan Mountain: a phytosociological and successional study. Ecological Monographs. 11: 61-97. [23349]

8. 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]

9. 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]

10. 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]

11. Cain, Stanley A. 1944. Foundations of plant geography. New York: Harper & Brothers. 556 p. [56074]

12. Carlile, D. W.; Tipton, A. R.; Whelan, J. B.; Sharik, T. L. 1978. Changes in productivity of food for white-tailed deer and wild turkey following a forest thinning operation in the Ridge and Valley Province of Virginia. Virginia Journal of Science. 29(2): 58. [10119]

13. 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]

14. Christensen, Norman L. 1988. Vegetation of the southeastern Coastal Plain. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. Cambridge: Cambridge University Press: 317-363. [17414]

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