Galax urceolata



INTRODUCTORY


 

  William S. Justice @ USDA-NRCS PLANTS Database
AUTHORSHIP AND CITATION:
League, Kevin R. 2006. Galax urceolata. 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/ [].

FEIS ABBREVIATION:
GALURC

SYNONYMS:
Galax aphylla L. [28,52]

NRCS PLANT CODE [67]:
GAUR2

COMMON NAMES:
galax
beetleweed
wandflower

TAXONOMY:
The scientific name of galax is Galax urceolata (Poir.) Brummitt (Diapensiaceae). Galax is a monotypic genus [41,63,73].

LIFE FORM:
Forb-shrub

FEDERAL LEGAL STATUS:
None

OTHER STATUS:
Galax is ranked as globally secure (G5). It is state-listed as vulnerable (S2) in West Virginia [47].

DISTRIBUTION AND OCCURRENCE

SPECIES: Galax urceolata
GENERAL DISTRIBUTION:
Galax occurs in the Appalachian Mountain, Piedmont, and Coastal Plain regions of the eastern United States. It is most common at the center of its distribution in the southern Appalachian Mountains. Its northernmost occurrence is in New York [30,41,51,63] and northeastern Massachusetts [47,51]. Galax distribution apparently skirts Pennsylvania, where it had not been collected (as of 2005). Its distribution continues south from Ohio to northern Alabama and Georgia [41,51,63]. Galax may be nonnative in New York, Massachusetts, Washington, DC., and Ohio (review by [51]). Plants database provides a state distributional map of galax.

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

ECOSYSTEMS [27]:
FRES10 White-red-jack pine
FRES11 Spruce-fir
FRES12 Longleaf-slash pine
FRES13 Loblolly-shortleaf pine
FRES14 Oak-pine
FRES15 Oak-hickory
FRES16 Oak-gum-cypress
FRES18 Maple-beech-birch

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

AL GA KY MA NY
NC OH SC TN VA
WV DC

KUCHLER [42] PLANT ASSOCIATIONS:
K095 Great Lakes pine forest
K096 Northeastern spruce-fir forest
K097 Southeastern spruce-fir forest
K098 Northern floodplain forest
K099 Maple-basswood forest
K100 Oak-hickory forest
K101 Elm-ash forest
K102 Beech-maple forest
K103 Mixed mesophytic forest
K104 Appalachian oak forest
K106 Northern hardwoods
K107 Northern hardwoods-fir forest
K108 Northern hardwoods-spruce forest
K109 Transition between K104 and K106
K110 Northeastern oak-pine forest
K111 Oak-hickory-pine
K112 Southern mixed forest
K113 Southern floodplain forest
K114 Pocosin

SAF COVER TYPES [24]:
5 Balsam fir
12 Black spruce
14 Northern pin oak
17 Pin cherry
18 Paper birch
19 Gray birch-red maple
20 White pine-northern red oak-red maple
21 Eastern white pine
22 White pine-hemlock
23 Eastern hemlock
24 Hemlock-yellow birch
25 Sugar maple-beech-yellow birch
26 Sugar maple-basswood
27 Sugar maple
28 Black cherry-maple
30 Red spruce-yellow birch
31 Red spruce-sugar maple-beech
32 Red spruce
33 Red spruce-balsam fir
34 Red spruce-Fraser fir
35 Paper birch-red spruce-balsam fir
39 Black ash-American elm-red maple
40 Post oak-blackjack oak
42 Bur oak
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
60 Beech-sugar maple
62 Silver maple-American elm
64 Sassafras-persimmon
65 Pin oak-sweetgum
70 Longleaf pine
71 Longleaf pine-scrub oak
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 (laurel) oak
89 Live oak
91 Swamp chestnut oak-cherrybark oak
92 Sweetgum-willow oak
93 Sugarberry-American elm-green ash
94 Sycamore-sweetgum-American elm
96 Overcup oak-water hickory
97 Atlantic white-cedar
107 White spruce
108 Red maple
109 Hawthorn
110 Black oak

SRM (RANGELAND) COVER TYPES [60]:
421 Chokecherry-serviceberry-rose
809 Mixed hardwood and pine
810 Longleaf pine-turkey oak hills
812 North Florida flatwoods

HABITAT TYPES AND PLANT COMMUNITIES:
Galax occurs in the understory of a variety of plant communities in the northeastern, Mid-Atlantic, and southern United States. It is most common in oak-pine (Quercus-Pinus spp.) forests that support an understory of mountain-laurel (Kalmia latifolia) and/or rosebay (Rhododendron maximum). Both mountain-laurel and rosebay are strongly associated with galax throughout galax's distribution [2,43,45,68]. Galax also occurs in oak-pine, hardwood, mixed conifer-hardwood, conifer, and shrubland communities.

Galax 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 galax's most frequent overstory dominant across galax's distribution [1]. In a mixed-oak forest on the Jefferson National Forest of Virginia, overstory dominants of galax 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]). Galax 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 galax 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].

Galax 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. Galax 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].

Galax is a common herbaceous associate of xeric oak-pine forest. Percent cover of galax 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 galax 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 galax; 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 galax 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, galax, trailing arbutus (Epigaea repens), and eastern teaberry (Gaultheria procumbens) dominated groundlayer vegetation [46].

Galax 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].

Galax 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 galax 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].

Galax 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].

Galax 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 galax, Virginia heartleaf (Hexastylis virginica), and Carolina silverbell (Halesia carolina) [57,58,72].

Galax 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 galax'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 galax'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, galax 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 galax is a dominant or important component of the ground layer.

KY: oak-hickory forests [5]
NC: early seral mixed conifer-hardwood (white pine-oak/eastern hemlock-yellow-poplar) [36]
        mixed hardwood-eastern hemlock acidic cove forest [58]
        chestnut oak/mountain-laurel/galax xeric ridge forest [13,58]
        mesic longleaf pine-blackjack oak/mountain-laurel/galax woodland [50]
SC:  mountain-laurel balds [1]
WI:  chestnut oak forest [17]
VA: mixed pine-hardwood acidic cove forest
       southern Appalachian chestnut oak/mountain-laurel/galax xeric ridge forest
       heath balds
       high-elevation red oak/galax forest [25]
       mixed-hardwood acidic cove forest [25,26]
       eastern hemlock/galax forest
       Carolina hemlock bluffs
       mixed pine-oak/heath
       chestnut oak/galax mesic forest
       montane, piedmont, and coastal plain acidic cliffs
       piedmont/coastal plain heath bluffs
       montane palustrine spray cliffs [25]
       southern Appalachian mixed oak-hickory forest
WV: southern Appalachian mixed oak-hickory forest [31]

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Galax urceolata
GENERAL BOTANICAL CHARACTERISTICS:
This description provides characteristics that may be relevant to fire ecology, and is not meant for identification. Keys for identification are available (e.g. [41,63,73]).

Galax 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. Galax is rhizomatous. Roots are fibrous [52,63].

Galax 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 galax are 4 inches (10 cm) across, while tetraploid galax'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].

RAUNKIAER [55] LIFE FORM:
Hemicryptophyte

REGENERATION PROCESSES:
Galax reproduces sexually and asexually. Information pertaining to the reproductive biology of galax is sparse, and the degree to which galax relies on reproduction from seed vs. cloning is unknown.

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

Polyploidy may give galax 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: Galax's small, light seed is "easily transported" [9]. Mechanisms for transport are not described in current (2006) literature.

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

Asexual regeneration: Galax 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 galax was lacking. Further research is needed on galax's life history.

SITE CHARACTERISTICS:
Galax occurs in mesic and xeric forested sites in the Appalachian mountains, piedmont, and coastal plains [28,31,52,59,63]. Baldwin [2] described an ideal galax site as "a shaded, mesic habitat in the mountains with a soil that is rich, acid, and humus-covered."

Elevation: Precise elevational data are sparse for galax. However, galax 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]. Galax 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 galax occur throughout the distribution of diploid galax. They also occur on the Virginia coastal plain, where diploid populations do not occur. Additionally, tetraploid galax are more common at high elevations than diploid galax [2].

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

Climate: Considerable climatic diversity is found in galax'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 galax'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 galax's southern range. Mean summer temperatures are less variable, ranging from 70 to 72 F (21-22 C) [14,29].

SUCCESSIONAL STATUS:
Galax occurs in early [1,19,36], mid- [20], and late-successional forests [1,13,19,35]. It is shade tolerant [68]. Partial shade provides best light conditions for galax (review by [51]).

Early succession: A study of forest succession in mixed conifer-hardwood riparian forests of the southern Appalachians noted galax's rarely documented importance in early succession. With 69% frequency, galax 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].

Galax 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 galax; 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 galax, dominated the undisturbed understory. Although method of postdisturbance establishment was not part of the study, galax establishment after 1967 was probably from seed because several successive herbicide treatments in the 1960s would have killed most mature plants. Galax distribution on undisturbed and disturbed plots was [19]:

  Frequency (%) Density (plants/m) Relative density
Undisturbed 38 0.8 5.1
Disturbed 3 0.1 <0.1

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

Postharvest year 25 27 32 41
Relative biomass (%) 0.0 2.2 6.5 14.8

In vegetation surveys of reclaimed surface coal mines of southwestern Virginia, galax 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 galax's successional position on coal mine sites.

Some heath balds where galax 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 galax exposure on balds could vary greatly. Galax's successional role in heath balds is not described in current (2006) literature.

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

SEASONAL DEVELOPMENT:
Galax germinates in early spring [51]. Other phenological events are listed below.
State Leaves emerge Flowers Disperses seed
West Virginia ---- June-July [63] ----
Carolinas ---- May-July Aug.-Oct. [52]
Appalachians 1 May- 15 June [25] ---- ----
Blue Ridge Mts. ---- May-July [73] ----
Northeast ---- June-July [28] ----

In the reclamation study discussed above, the authors noted that galax was wilted and dormant during August surveys [39].

FIRE ECOLOGY

SPECIES: Galax urceolata
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: As of this writing (2006), no literature specifically addressed galax fire adaptations. It is likely that galax's rhizomes survive fire, as galax has been found on burned sites in the 1st postfire growing season [16].

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 galax tetraploidy.

Fire regimes: Fire was historically important in maintaining oak-pine and high-elevation, xeric pine forests where galax occurs [46,58,71]. In the Appalachian Mountains, where galax 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 galax 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 galax 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/galax 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 galax 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 galax 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)
maple-beech-birch Acer-Fagus-Betula spp. >1,000
silver maple-American elm Acer saccharinum-Ulmus americana <5 to 200
sugar maple Acer saccharum >1,000
sugar maple-basswood Acer saccharum-Tilia americana >1,000
beech-sugar maple Fagus spp.-Acer saccharum >1,000
black ash Fraxinus nigra <35 to 200
yellow-poplar Liriodendron tulipifera <35 [70]
northeastern spruce-fir Picea-Abies spp. 35-200 [18]
southeastern spruce-fir Picea-Abies spp. 35 to >200 [70]
red spruce* Picea rubens 35-200 [18]
shortleaf pine Pinus echinata 2-15
shortleaf pine-oak Pinus echinata-Quercus spp. <10
longleaf pine-scrub oak Pinus palustris-Quercus spp. 6-10
Table Mountain pine Pinus pungens <35 to 200 [70]
red-white-jack pine* Pinus resinosa-P. strobus-P. banksiana 10-300 [18,37]
pitch pine Pinus rigida 6-25 [8,38]
pocosin Pinus serotina 3-8
eastern white pine Pinus strobus 35-200
eastern white pine-eastern hemlock Pinus strobus-Tsuga canadensis 35-200
eastern white pine-northern red oak-red maple Pinus strobus-Quercus rubra-Acer rubrum 35-200
loblolly-shortleaf pine Pinus taeda-P. echinata 10 to <35
Virginia pine Pinus virginiana 10 to <35
Virginia pine-oak Pinus virginiana-Quercus spp. 10 to <35
sycamore-sweetgum-American elm Platanus occidentalis-Liquidambar styraciflua-Ulmus americana <35 to 200 [70]
aspen-birch Populus tremuloides-Betula papyrifera 35-200 [18,70]
black cherry-sugar maple Prunus serotina-Acer saccharum >1,000
oak-hickory Quercus-Carya spp. <35 [70]
oak-juniper woodland (Southwest) Quercus-Juniperus spp. <35 to <200 [49]
northeastern oak-pine Quercus-Pinus spp. 10 to <35 [70]
southeastern oak-pine Quercus-Pinus spp. <10
white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra <35
northern pin oak Quercus ellipsoidalis <35
bear oak Quercus ilicifolia <35
chestnut oak Quercus prinus 3-8
northern red oak Quercus rubra 10 to <35
post oak-blackjack oak Quercus stellata-Q. marilandica <10
black oak Quercus velutina <35
eastern hemlock-yellow birch Tsuga canadensis-Betula alleghaniensis >200 [70]
eastern hemlock-white pine Tsuga canadensis-Pinus strobus x = 47 [15]
elm-ash-cottonwood Ulmus-Fraxinus-Populus spp. <35 to 200 [18,70]
*fire return interval varies widely; trends in variation are noted in the species review

POSTFIRE REGENERATION STRATEGY [62]:
Surface rhizome/chamaephytic root crown in organic mantle or on soil surface
Rhizomatous herb, rhizome in soil

FIRE EFFECTS

SPECIES: Galax urceolata
IMMEDIATE FIRE EFFECT ON PLANT:
Galax is highly flammable under dry conditions [66], and fire probably top-kills galax [16].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
Fire effects on galax likely vary with season and severity, ranging from partial to complete consumption of the aboveground plant.

PLANT RESPONSE TO FIRE:
Galax establishes after fire [9,23], although documentation of the method (rhizome sprouts and/or seedling establishment) is lacking as of this writing (2006). It is likely that if plants are healthy before fire and there is sufficient rainfall afterwards, top-killed galax rhizomes sprout readily after fire. However, galax's postfire recovery rate varies, and current research (as of 2006) does not explain the variation. Research is needed on galax's fire ecology.

Several studies note galax presence on burns [9,16,22,23]. For example, galax 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].

Galax 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." Galax's postfire frequency and height were low compared to 27 and 31 other herbs present on 2 study plots. Galax 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]:

  Frequency
(%)
Biomass
(g/plot)
Height
(cm)
Density
(stems/plot)
Jacob Branch East Site, 1991 3 0.40 3.7 1.0
Jacob Branch West Site, 1992 3 3.48 7.3 1.3

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Galax postfire growth rate is not always slow. Galax recovered quickly after prescribed burning in a pitch pine-chestnut-oak forest on the Nantahala National Forest, North Carolina. Burning was conducted in the spring and varied from "light" to "heavy" in severity. Galax percent cover at postfire year 2 was double that of prefire coverage, though still less than 1% [22].

After a felling and burning treatment in a chestnut oak-pitch pine/mountain-laurel forest on the Nantahala National Forest, galax 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 galax abundance decreased substantially, presumably due to interference from other plant species. Because initial posttreatment recovery was rapid, the authors presumed that galax's postfire growth was from rhizome sprouts. Density and percent cover of galax were as follows [16]:

  Postfire year Density (stems/m) Cover
(%)
Pretreatment 0 4.0 1.1
Posttreatment 1 4.5 1.3
2 4.8 1.2
4 1.0 0.1

Galax 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. Galax declined on 2- and 4-burn plots, but increased on 3-burn plots. Importance value of galax on each treatment was [53]:

Number of burns

0 1 2 3 4
29.34 29.56 0.85 40.25 6.77

Galax is noted as occurring in deeply shaded, late-successional forest [13,19,35,74], although whether galax 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 galax 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. Galax 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, galax was the most abundant herbaceous species present on the oldest burns. Mean percent cover of galax on permanent plots was [35]:

  Study Years Cover (%)
Burned before 1940 1977-1978 1.17
1995 1.08
Burned 1976-1977 1977 0.30
1978 0.33
1979 0.27
1980 0.35
1984 0.15
1995 0.17

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

FIRE MANAGEMENT CONSIDERATIONS:
Galax evolved under a wide variety of fire regimes, and information of its rate of postfire recovery is sparse for all plant communities in which it occurs (as of 2006). How quickly galax recovers from fire will vary with fire regime, plant community, fuels, and fire season. Harvest history also undoubtedly affects galax's ability to recover after fire. Because harvesting (see Other Management Considerations) reduces galax's photosynthetic capacity and biomass, populations that have been heavily harvested probably have reduced rhizome carbohydrate reserve. Therefore, they have reduced ability to sprout and produce seed compared to unharvested populations. Data are need on 1) sustainable levels of galax harvest and 2) interactive effects of harvesting and fire.

Galax leaves may help generate severe fires that provide important ecological benefits. Turrill and others [66] report that in Table Mountain/mountain-laurel/galax communities on the Chattahoochee National Forest of Georgia, groundlayer galax 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 galax and blueberry-fueled surface fires up to the serotinous cones of Table Mountain pine, aiding in the pine's postfire regeneration [66].

Elliot and Clinton [21] provide allometric equations for predicting aboveground dry weight of galax in the southern Appalachian Mountains.

MANAGEMENT CONSIDERATIONS

SPECIES: Galax urceolata
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Wildlife graze galax. It is preferred forage for white-tailed deer and wild turkeys in Virginia [12]. Heaviest white-tailed deer use is in fall and winter [40]; heaviest year-round deer use is in years of poor acorn production [32,40]. To date (2006), information on galax's nutritional content and its use as cover by small animals is lacking. Further research is needed on plant-animal relationships for galax.

VALUE FOR REHABILITATION OF DISTURBED SITES:
Because of its spreading rhizomes, galax is a good choice for planting on sites subject to erosion. It is propagated from rhizome cuttings and seed, and can be mass-cultivated. Predny and Chamberlain [51] review sowing and planting guidelines for galax seed and rhizomes.

OTHER USES:
Galax is harvested for use in the floral industry. Its durable, shiny evergreen leaves, which are brightly colored in fall, are highly prized as background foliage in floral arrangements. It is also used for landscaping [51].

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

OTHER MANAGEMENT CONSIDERATIONS:
High demand by the floral industry is threatening galax population stability on some sites. An experienced harvester can pull approximately 5,000 leaves/day. Pulling and uprooting galax is a common harvest method and is faster than breaking off individual leaves, but pulling removes rhizomes and kills plants so harvested [51]. The larger-leaved, tetraploid galax is harvested most often. High elevations of the Appalachian Mountains, where galax is mostly tetraploid, contain "all the territory worth while for 'picking' the Galax" (Woodruff 1940, personal communication in [2]). As of 2005, galax harvesting was restricted from 1 May through 15 June, when new leaves emerge. Research to determine sustainable harvestable levels, and fine-tuning the harvest season for galax, is ongoing (as of 2006) [51].

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]
15. Cleland, David T.; Crow, Thomas R.; Saunders, Sari C.; Dickmann, Donald I.; Maclean, Ann L.; Jordan, James K.; Watson, Richard L.; Sloan, Alyssa M.; Brosofske, Kimberley D. 2004. Characterizing historical and modern fire regimes in Michigan (USA): a landscape ecosystem approach. Landscape Ecology. 19: 311-325. [54326]
16. 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]
17. Curtis, John T. 1959. The vegetation of Wisconsin. Madison, WI: The University of Wisconsin Press. 657 p. [7116]
18. Duchesne, Luc C.; Hawkes, Brad C. 2000. Fire in northern 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: 35-51. [36982]
19. 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]
20. Elliott, Katherine J.; Boring, Lindsay R.; Swank, Wayne T.; Haines, Bruce R. 1997. Successional changes in plant species diversity and composition after clearcutting a southern Appalachian watershed. Forest Ecology and Management. 92: 67-85. [27629]
21. Elliott, Katherine J.; Clinton, Barton D. 1993. Equations for estimating biomass of herbaceous and woody vegetation in early-successional southern Appalachian pine-hardwood forests. Res. Note SE-365. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 7 p. [22273]
22. 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]
23. Elliott, Katherine J.; Vose, James M. 1995. Evaluation of the competitive environment for white pine (Pinus strobus L.) seedlings planted on prescribed burn sites in the s. Appalachians. Forest Science. 41(3): 513-530. [26758]
24. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]
25. Fleming, G. P.; Coulling, P. P.; Patterson, K. D. 2005. The natural communities of Virginia: classification of ecological community groups. Second approximation. Version 2.1. Richmond, VA: Virginia Department of Conservation and Recreation, Division of Natural Heritage. Available: http://www.dcr.virginia.gov/dnh/ncintro.htm [2005, November 3]. [54983]
26. Fleming, Gary P. 2003. Ecological communities of the Northern Virginia Blue Ridge [Online]. Virginia Department of Conservation and Recreation, Division of Natural Heritage (Producer). Available: http://www.dcr.virginia.gov/dnh/Noblueridge.pdf [2005, November 21]. [55271]
27. 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]
28. 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]
29. Greller, Andrew M. 1977. A classification of mature forests on Long Island, New York. Bulletin of the Torrey Botanical Club. 104(4): 376-382. [22020]
30. Greller, Andrew M.; Clemants, Steven E. 2001. Flora of West Hills Park, Suffolk County, New York, with considerations of provenance of some long-distance disjuncts. Journal of the Torrey Botanical Society. 128(1): 76-89. [55625]
31. Hammond, Daniel N. 1997. Characterization of vascular plant species composition and relative abundance in southern Appalachian mixed-oak forests. Blacksburg, VA: Virginia Polytechnic Institute and State University. 113 p. Thesis. [54881]
32. Harlow, Richard F.; Whelan, James B.; Crawford, Hewlette S.; Skeen, John E. 1975. Deer foods during years of oak mast abundance and scarcity. Journal of Wildlife Management. 39(2): 330-336. [10088]
33. Harmon, M. E.; Bratton, S. P.; White, P. S. 1983. Disturbance and vegetation response in relation to environmental gradients in the Great Smoky Mountains. Vegetatio. 55: 129-139. [21309]
34. Harmon, Mark. 1982. Fire history of the westernmost portion of Great Smoky Mountains National Park. Bulletin of the Torrey Botanical Club. 109(1): 74-79. [9754]
35. Harrod, J. C.; Harmon, M. E.; White, P. S. 2000. Post-fire succession and 20th century reduction in fire frequency on xeric southern Appalachian sites. Journal of Vegetation Science. 11(4): 465-472. [38753]
36. Hedman, Criag W.; Van Lear, David H. 1995. Vegetative structure and composition of southern Appalachian forests. Bulletin of the Torrey Botanical Club. 122(2): 134-144. [55023]
37. Heinselman, Miron L. 1970. The natural role of fire in northern conifer forests. In: The role of fire in the Intermountain West: Symposium proceedings; 1970 October 27-29; Missoula, MT. Missoula, MT: Intermountain Fire Research Council: 30-41. In cooperation with: University of Montana, School of Forestry. [15735]
38. 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]
39. Holl, Karen D. 2002. Long-term vegetation recovery on reclaimed coal surface mines in the eastern USA. Journal of Applied Ecology. 39(6): 960-970. [47075]
40. Johnson, A. Sydney; Hale, Philip E.; Ford, William M.; [and others]. 1995. White-tailed deer foraging in relation to successional stage, overstory type and management of southern Appalachian forests. The American Midland Naturalist. 133(1): 18-35. [25953]
41. Kartesz, John T.; Meacham, Christopher A. 1999. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Available: North Carolina Botanical Garden. In cooperation with: The Nature Conservancy, Natural Resources Conservation Service, and U.S. Fish and Wildlife Service [2001, January 16]. [36715]
42. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. [3455]
43. Little, Parker L. 2002. An exploration and study of White Rock Mountain, WV for unusual forms of Rhododendron species. American Rhododendron Society Journal. 56(3): 122-131. [55138]
44. McLeod, D. E. 1988. Vegetation patterns, floristics, and environmental relationships in the Black and Craggy mountains of North Carolina. University of North Carolina, Chapel Hill. 222 p. Ph.D. dissertation. [60570]
45. McNab, W. Henry; Browning, Sara A.; Simon, Steven A.; Fouts, Penelope F. 1999. An unconventional approach to ecosystem unit classification in western North Carolina, USA. Forest Ecology and Management. 114: 405-420. [54831]
46. Murphy, Paul A.; Nowacki, Gregory J. 1997. An old-growth definition for xeric pine and pine-oak woodlands. Gen. Tech. Rep. SRS-7. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. 7 p. [28623]
47. NatureServe. 2005. Comprehensive report: Galax urceolata--beetle weed, [Online]. In: NatureServe Explorer: An online encyclopedia of life. Version 4.6. Arlington, VA: NatureServe (Producer). Available: http://www.natureserve.org/explorer/servlet/NatureServe?sourceTemplate=species_RptComprehensive.wmt&loadTemplate=species_RptStatus.wmt&paging=&elKey=158136&save=false&page=1#contributors [2005, November 22]. [55632]
48. Oosting, Henry J. 1942. An ecological analysis of the plant communities of the Piedmont, North Carolina. The American Midland Naturalist. 28: 1-126. [50588]
49. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; [and others]. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-volume 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. [36978]
50. Peet, Robert K.; Allard, Dorothy J. 1993. Longleaf pine vegetation of the southern Atlantic and Gulf Coast regions: a preliminary classification. In: Hermann, Sharon M., ed. The longleaf pine ecosystem: ecology, restoration and management: Proceedings, 18th Tall Timbers fire ecology conference; 1991 May 30 - June 2; Tallahassee, FL. Tallahassee, FL: Tall Timbers Research, Inc: 45-81. [28325]
51. Predny, Mary L.; Chamberlain, James L. 2005. Galax (Galax urceolata): an annotated bibliography. Gen. Tech. Report SRS-87. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. 33 p. Available online: http://www.srs.fs.usda.gov/pubs/21011 [2005, November 2]. [54979]
52. 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]
53. Randles, Russel B.; Van Lear, David H.; Waldrop, Thomas A. 2002. Periodic burning in table mountain-pitch pine stands. In: Outcalt, Kenneth W., ed. Proceedings, 11th biennial southern silvicultural research conference; 2001 March 20-22; Knoxville, TN. Gen. Tech. Rep. SRS-48. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station: 114-118. [41464]
54. Randolph, L. F. 1932. Some effects of high temperature on polyploidy and other variations in maize. Proceedings of the National Academy of Sciences of the United States of America. 18(3): 222-229. [60595]
55. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]
56. Reich, Peter B.; Walters, Michael B.; Ellsworth, David S.; Vose, James M.; Volin, John C.; Gresham, Charles; Bowman, William D. 1998. Relationships of leaf dark respiration to leaf nitrogen, specific leaf area and leaf life-span: a test across biomes and functional groups. Oecologia. 114: 471-482. [54832]
57. Rentch, James S.; Adams, Harold S.; Coxe, Robert B.; Stephenson, Steven L. 2000. An ecological study of a Carolina hemlock (Tsuga caroliniana) community in southwestern Virginia. Castanea. 65(1): 1-8. [39273]
58. 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 online: http://ils.unc.edu/parkproject/nhp/publications/class.pdf [2005, February 14]. [41937]
59. 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]
60. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]
61. Simpson, Robert L.; Leck, Mary Allessio; Parker, V. Thomas. 1989. Seed banks: general concepts and methodological issues. In: Leck, Mary Allessio; Parker, V. Thomas; Simpson, Robert L., eds. Ecology of soil seed banks. New York: Academic Press, Inc: 3-8. [60593]
62. 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]
63. Strausbaugh, P. D.; Core, Earl L. 1977. Flora of West Virginia. 2nd ed. Morgantown, WV: Seneca Books, Inc. 1079 p. [23213]
64. Tischler, G. 1935. Die bedeutung der polyploidie fur die verbreitung der angiospermen. Botanische Jahrbucher. 67: 1-36. [60597]
65. Turner, L. M. 1935. Catastrophes and pure stands of southern shortleaf pine. Ecology. 16(2): 213-215. [41631]
66. Turrill, Nicole L.; Buckner, Edward R.; Waldrop, Thomas A. 1997. Pinus pungens Lam. (Table Mountain pine): a threatened species without fire? In: Greenlee, Jason M., ed. Proceedings, 1st conference on fire effects on rare and endangered species and habitats; 1995 November 13-16; Coeur d'Alene, ID. Fairfield, WA: International Association of Wildland Fire: 301-306. [28153]
67. U.S. Department of Agriculture, Natural Resources Conservation Service. 2006. PLANTS database (2006), [Online]. Available: http://plants.usda.gov/. [34262]
68. Vandermast, D. B.; Van Lear, D. H. 2002. Riparian vegetation in the southern Appalachian mountains (USA) following chestnut blight. Forest Ecology and Management. 155: 97-106. [40782]
69. Vandermast, David B.; Van Lear, David H. 1999. Vegetative composition of riparian forest once dominated by American chestnut. In: Haywood, James D., ed. Proceedings, 10th biennial southern silvicultural research conference; 1999 February 16-18; Shreveport, LA. Gen. Tech. Rep. SRS-30. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station: 3-7. [34398]
70. 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]
71. Welch, Nicole Turrill. 1999. Occurrence of fire in southern Appalachian yellow pine forests as indicated by macroscopic charcoal in soil. Castanea. 64(4): 310-317. [39471]
72. Whittaker, R. H. 1956. Vegetation of the Great Smoky Mountains. Ecological Monographs. 26(1): 1-79. [11108]
73. Wofford, B. Eugene. 1989. Guide to the vascular plants of the Blue Ridge. Athens, GA: The University of Georgia Press. 384 p. [12908]
74. Zobel, Donald B. 1969. Factors affecting the distribution of Pinus pungens, an Appalachian endemic. Ecological Monographs. 39: 302-333. [17651]

FEIS Home Page