• ABBREVIATION
• SYNONYMS
• NRCS PLANT CODE
• COMMON NAMES
• TAXONOMY
• LIFE FORM
• FEDERAL LEGAL STATUS
• OTHER STATUS
• AUTHORSHIP AND CITATION

ABBREVIATION:

SYNONYMS:

NRCS PLANT CODE [191]:

COMMON NAMES:

TAXONOMY:

LIFE FORM:

FEDERAL LEGAL STATUS:

OTHER STATUS:

Pennsylvania: The state rank of inkberry is SX and the state status is PX. These designations indicate inkberry is believed to be extinct within Pennsylvania [149].

AUTHORSHIP AND CITATION:

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

GENERAL DISTRIBUTION:

ECOSYSTEMS [58]:

STATES:

BLM PHYSIOGRAPHIC REGIONS [10]:

KUCHLER [109] PLANT ASSOCIATIONS:

SAF COVER TYPES [51]:

SRM (RANGELAND) COVER TYPES [179]:

HABITAT TYPES AND PLANT COMMUNITIES:

Inkberry is associated with pitch pine (Pinus rigida) on the Atlantic coastal plain [6,12,49,120,124,182]. Other associated tree species in these communities include white oak (Quercus alba) [6,12], bear oak (Quercus ilicifolia) [12,24,32,124], blackjack oak (Quercus marilandica) [124,182], dwarf chinkapin oak (Quercus prinoides) [24,32], post oak (Quercus stellata) [124,182], black oak (Quercus velutina) [6,12,182], shortleaf pine (Pinus echinata) [12,32,182], and sassafras (Sassafrass albinum) [124]. Associated shrub species include downy serviceberry (Amelanchier arborea) [24], coastal sweetpepperbush (Clethra alnifolia) [32,182],  black huckleberry [6,12,24,32,124,182], dwarf huckleberry (Gaylussacia dumosa) [32], blue huckleberry (Gaylussacia frondosa) [24,32,49,124,182], wintergreen [12,24,182], sheep-laurel [24,49,124,182], mountain-laurel (Kalmia latifolia) [24,32,124], swamp doghobble (Leucothoe racemosa) [182], maleberry (Lyonia ligustrina) [12], piedmont staggerbush (Lyonia mariana) [182], bayberry (Morella spp.) [6,182], black chokecherry (Photinia melanocarpa) [24], black cherry (Prunus serotina) [11], flameleaf sumac (Rhus copallinum) [24], cat greenbrier (Smilax glauca) [24,182], roundleaf greenbrier (Smilax rotundifolia) [182], Virginia tephrosia (Tephrosia virginiana) [12], lowbush blueberry [24], and Blue Ridge blueberry (Vaccinium pallidum) [12,24,124,182]. Additional understory associates include Pennsylvania sedge (Carex pensylvanica) [12,24], leatherleaf (Chamaedaphne calyculata) [49], sweetfern (Comptonia peregrina) [12,24,32,124,182], western brackenfern (Pteridium aquilinum) [12,24], and anisescented goldenrod (Solidago odora) [12].

In Atlantic white-cedar (Chamaecyparis thyoides) forests, inkberry occurs with red maple, pitch pine [138], and redbay (Persea borbonia) [127]. Understory species occurring in these communities include sweetpepperbush (Clethra spp.) [135,138], blue huckleberry [135], large gallberry (Ilex coriacea) [127,135], American holly (Ilex opaca) [127], common winterberry (Ilex verticellata) [138], maleberry [135], fetterbush lyonia (Lyonia lucida) [127,135], partridgeberry (Mitchella repens) [127], southern bayberry (Morella caroliniensis) [135], cinnamon fern (Osmunda cinnamomea) [138], Virginia creeper (Parthenocissus quinquefolia) [127], swamp azalea (Rhododendron viscosum) [138], poison-ivy (Toxicodendron radicans) [127], highbush blueberry (Vaccinium corymbosum) [138], black highbush blueberry (Vaccinium fuscatum) [135], possumhaw,  Virginia chainfern (Woodwardia virginica), and mosses (Sphagnum spp.) [138].

Inkberry may dominate pocosin or shrub swamp vegetation [7,148,157], commonly occurring with southern bayberry [201], swamp titi (Cyrilla racemiflora) [47,94,148,156,157,176,201], cane (Arundinaria gigantea) [90,148,201], red chokecherry (Photinia pyrifolia) [157], cinnamon fern [201], coastal sweetpepperbush [90], greenbrier (Smilax spp.) [148,156,157,176,201], swamp doghobble [47], fetterbush lyonia [47,90,94,157], honeycup (Zenobia pulverulenta) [94,148,156,176], dwarf huckleberry [41], highbush blueberry [47,157], wax myrtle (Morella cerifera) [90,157,176,201,206], northern bayberry [41], and moss (Sphagnum spp.) [41,157,201]. Also occurring in pocosin vegetation is scattered red maple [156,201], Atlantic white-cedar [156], loblolly bay (Gordonia lasianthus) [94,156,157,176], sweetgum (Liquidambar styraciflua) [201], sweetbay (Magnolia virginiana) [47,94,148,156,157,176,201], redbay [94,156,157,176], pond pine (Pinus serotina) [90,94,157,176,206], and bald cypress (Taxodium distichum) [156].

In pond pine forests, inkberry occurs with bluestem (Andropogon spp.) [92], cane [16,92], sweetpepperbush [16,72,92], swamp cyrilla [16,72], huckleberry (Gaylussacia spp.) [72,92], large gallberry [16], St. Johnswort (Hypericum spp.) [92], maleberry [72], fetterbush lyonia [44,72,92], wax myrtle [16,44,57], cinnamon fern, sumac (Rhus spp.) [92], saw-palmetto (Serenoa repens) [16,44,72], laureleaf greenbrier (Smilax laurifolia) [16,72], honeycup [16], and blueberry [44,92]. Other tree species include red maple [16,72], Atlantic white-cedar [72], loblolly bay], sweetgum [16], sweetbay [72,92], redbay [44], and loblolly pine (Pinus taeda) [72].

Inkberry is found in bay forests with loblolly bay [5,18,126,176], sweetbay [5,126,152,176], and redbay [5,18,126,176]. Understory species in these communities include coastal sweetpepperbush, cane [126], buckwheat tree (Cliftonia monophylla) [18,126], swamp titi [126,176], huckleberry [176], dahoon (Ilex cassine) [5,18,126,176], large gallberry [126,176], deciduous holly (Ilex decidua) [126], smooth winterberry (Ilex laevigata) [176], American holly [126], yaupon (Ilex vomitoria) [18], Virginia sweetspire (Itea virginica), sheep-laurel, swamp doghobble [126], maleberry [5,126], fetterbush lyonia [5,18,126,176], bayberry [5,18,126,176], swamp azalea [176], greenbrier [126,176], poison sumac (Toxicodendron vernix) [18], highbush blueberry, black highbush blueberry [5,176], Virginia chainfern [176], and honeycup [126,176].

Inkberry occurs in bald cypress forests with red maple, sweetbay, blackgum (Nyssa sylvatica), redbay [170,171], and slash pine (Pinus elliottii) [54]. Understory species include chalky bluestem (Andropogon capillipes), baccharis (Baccharis spp.), toothed midsorus fern (Blechnum serrulatum), smallspike false nettle (Boehmeria cylindrica), spadeleaf (Centella asiatica), Jamaica swamp sawgrass (Cladium mariscus) [54], coastal sweetpepperbush [170,171], flatsedge (Cyperus spp.) [54], swamp titi [170,171], southern umbrella-sedge (Fuirena scirpoidea), peelbark St. Johnswort (Hypericum fasciculatum), clustered bushmint (Hyptis alata) [54], dahoon [54,170,171], large gallberry, Virginia sweetspire, swamp doghobble [170,171], primrose-willow (Ludwigia spp.), rusty staggerbush (Lyonia ferruginea) [54], fetterbush lyonia [170,171], wax myrtle, wild pennyroyal (Piloblephis rigida), camphorweed (Pluchea spp.), cabbage palmetto (Sabal palmetto), licorice weed (Scoparia dulcis) [54], laurel greenbrier, coral greenbrier (Smilax walteri), and blueberry [170,171]. 

Inkberry is associated with pond cypress (Taxodium distichum var. nutans) [134,202], occurring as an understory species with swamp cyrilla, eastern swamp privet (Forestiera acuminata), yaupon [202], wax myrtle [134,202], and viburnum (Viburnum spp.) [202]. Tree species in these communities include red maple, blackgum, and slash pine [134].

Inkberry is commonly found in southeastern pine flatwoods communities, dominated by sand pine (Pinus elliottii var. elliottii) [65,116], slash pine [4,5,32,38,43,79,95,96,125,128,129,130,133,144,146,162,184], longleaf pine (Pinus palustris) [4,14,15,30,31,39,43,59,77,78,79,125,128,129,133,146,162,167,176,184], pond pine [4,133,184], and loblolly pine [31,59,60,118,131,159,167]. Several oak species occur in flatwoods communities, including Chapman oak (Quercus chapmanii) [39], southern red oak (Quercus falcata) [39,118,121,159], sand live oak (Quercus geminata) [184], bluejack oak (Quercus incana) [39], turkey oak (Quercus laevis) [39], laurel oak (Quercus laurifolia) [11,39,79], runner oak (Quercus margarettiae) [125], blackjack oak [118,121], dwarf live oak (Quercus minima) [4,39,56,77,184], myrtle oak (Quercus myrtifolia) [56], water oak (Quercus nigra) [4,11,60,79,159], willow oak (Quercus phellos) [118,121,159], running oak (Quercus pumila) [133,162,184], post oak [60,118,121], and live oak [4,11,30,43]. Other tree species occurring in these communities include red maple [4,39,60,79], persimmon (Diospyros spp.) [15], ash (Fraxinus spp.) [4], sweetgum [4,11,59,60,118,121,131,159], loblolly bay [39], sweetbay [39,72,79], blackgum [59,60,72,118,121,131,159], redbay [11,61,72], swamp bay (Persea palustris) [79], shortleaf pine [59,60,131], spruce pine (Pinus glauca) [108], and pond cypress [43]. 

Inkberry occurs as an understory dominant in the flatwoods/inkberry plant association in Florida [5] and in the holly-bayberry-titi plant association in North Carolina [201]. It is also an understory dominant in the longleaf pine/blue huckleberry-inkberry community type in North Carolina [104]. Understory associates of inkberry in southeastern pine flatwoods include cane [55,59], pawpaw (Asimina spp.) [38], buckwheat tree [72], tarflower [1,2,4,39], sedges (Carex spp.) [55], sweetpepperbush [72,118,121,131], flowering dogwood (Cornus florida) [15,99], black titi [190], swamp titi [72,103,167], huckleberry [4,5,14,15,38], St. Johnswort [56,72], large gallberry [30,103,146,176,190], yaupon [14,98,99,146], mountain-laurel [103], hairy-laurel (Kalmia hirsuta) [84,184], sand myrtle (Leiophyllum buxifolium) [103], fetterbush lyonia [4,5,39,43,61,103,125,130,144,184,190], rusty staggerbush [116], coastalplain staggerbush (Lyonia fruticosa) [20,116], piedmont staggerbush [103,176], wax myrtle [4,14,30,38,39,43,59,79,144,150,159,176,184], southern bayberry [121], red chokecherry [56], azalea (Rhododendron spp.) [56], beakrushes (Rhynchospora spp.) [55], sumac [14,43], blackberry (Rubus spp.) [14,15,39,112,146], cabbage palm (Sabal palmetto) [43,196], saw-palmetto [4,5,14,38,39,43,77,78,79,125,128,129,144,150], greenbrier [72,190], blueberry [5,14,15,59,72,79,99,125,162,176,184], and viburnum [39]. Grasses occurring in these communities include threeawn (Aristida spp.) [4,39,77,84,97,125,130,133,146,162,167], common carpet grass (Axonopus fissifolius) [20,68], toothache grass (Ctenium aromaticum) [20,115], panic grass (Panicum spp.) [3,35,36,55,72,115,118,146], muhly (Muhlenbergia spp.) [55], bluestem (Schizachyrium and Andropogon spp.) [35,39,55,59,68,98,99,115,118,125,146,146], Indian grass (Sorghastrum spp.) [20,55], Curtis' dropseed (Sporobolus curtissii) [20,36,68,118], and Florida dropseed (Sporobolus floridanus) [115].

In hammock vegetation of Florida, inkberry occurs with various hickories (Carya spp.) and other hardwoods, southern magnolia (Magnolia grandiflora), wax myrtle, lancewood (Nectandria coriacea), West Indian cherry (Prunus myrtifolia), evergreen oaks (Quercus spp.), and coontie (Zamia pumila) [57].

Inkberry is also an important component of saw palmetto-dominated scrub vegetation, occurring with pineland threeawn (Aristida stricta), tarflower, rusty staggerbush, coastalplain staggerbush, fetterbush lyonia, wax myrtle, redbay, western brackenfern, Chapman oak, myrtle oak, sand live oak, greenbrier, shiny blueberry (Vaccinium myrsinites), deerberry (Vaccinium stamineum), and tallow wood (Ximenia americana) [173].

In sand scrub vegetation, inkberry occurs with  pawpaw (Asimina spp.), tarflower [38], sand heath (Ceratiola ericoides), garberia (Garberia heterophylla) [139], huckleberry [38], St. Andrew's cross (Hypericum hypericoides), Carolina holly (Ilex ambigua) [139], dahoon [38], staggerbush (Lyonia spp.), wax myrtle [38,139], Chapman oak [139], sand live oak [38,139], laurel oak [38], myrtle oak, redbay, silk bay, devilwood (Osmanthus americanus), oak mistletoe (Phoradendron leucarpum) [139], sand pine (Pinus clausa) [38,139], scrub plum (Prunus geniculata) [139], hog plum (Prunus umbellata) [38], tallow wood, western brackenfern, flameleaf sumac (Rhus copallinum), scrub palmetto (Sabal etonia) [139], saw palmetto [38,139], greenbrier, shiny blueberry, and deerberry [139].

Coastal sand dune associates of inkberry include live oak, hog plum, prickly-pear (Opuntia spp.), staggerbush, periwinkle (Catharanthus spp.), saltbush (Atriplex spp.), avocado (Persea spp.), banana (Musa spp.), and scrub oak [38].

In dry prairie vegetation, inkberry occurs with pawpaw (Asimina spp.) [38,43], tarflower [136,158], sedge (Cyperus spp.) [28,38], coastalplain St. Johnswort (Hypericum brachyphyllum) [136], whitehead bogbutton (Lachnocaulon anceps) [89], rusty staggerbush [158], coastalplain staggerbush [53,89,136], fetterbush lyonia [43,53,136], wax myrtle [53], eastern prickly-pear (Opuntia humifusa) [28], sand live oak [89], dwarf live oak [53,101,136], running oak [43,53], plumed beaksedge (Rhynchospora plumosa) [89], saw-palmetto [4,38,53,89,101,136,158], shiny blueberry [53,136], Darrow's blueberry (Vaccinium darrowii) [89], common carpetgrass [101], Curtis' dropseed [80], threeawn [4,80,89,136,158], sawgrass (Cladium spp.) [38], cutthroat grass (Panicum abscissum) [209], little bluestem (Schizachyrium scoparium), lopsided Indian grass (Sorghastrum secundum), narrowleaf silkgrass (Pityopsis graminifolia), and vanillaleaf (Carphephorus odoritissimus) [89].

Classifications identifying inkberry as a plant community dominant are as follows:

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

GENERAL BOTANICAL CHARACTERISTICS:

Inkberry is a stoloniferous or rhizomatous shrub forming extensive, dense colonies [45,48,63,69,110,116,153]. Inkberry may comprise 67 to 75% of shrub cover on coastal plain sites [116]. Several aerial stems arise singly or in groups from a rhizome that ranges in diameter from 0.2 to 0.5 inch (0.5-1.3 cm). A few small fibrous or woody roots occur along the entire rhizome, and longer roots (up to 48 inches or 120 cm) occur at infrequent intervals.  These longer roots may extend to the water table if it is near the surface; the rhizome itself grows in the upper 2 inches (5 cm) of soil [114]. In coastal plain loblolly forests, inkberry was estimated to have 660 to 1,459 rootstocks per acre (1,650-3,648 rootstocks/ha) [132].

Male plants have flowers occurring in clumps of 3 on long peduncles, while female plants have flowers occurring singly [45,69] or in clumps of up to 3 flowers [45]. Inkberry fruits grow individually or in small clusters, with 3 to 9 seeds [69].

RAUNKIAER [154] LIFE FORM:

REGENERATION PROCESSES:

Breeding system: Inkberry is dioecious [45,62,63,69,153].

Pollination: Inkberry is pollinated by insects [123].

Seed production: In general, inkberry produces good seed crops annually, though individual plants or clumps may bear heavily one year and be barren the next [69]. Immediately following fire, inkberry fruit production drops off, but within 4 years high fruit yields occur [187]. The pruning of older inkberry growth may cause inkberry to flower with renewed vigor, though 2 seasons are required [116]. Inkberry does not flower or produce fruit until the 2nd year after injury, at which point it often bears profusely [69,80,116]. In Georgia slash pine plantations, inkberry fruited heavily in all successional stages except recently burned stands, and produced maximum yields in 4-year-old plantations. Fruit yields were generally higher in young stands than in older stands [97]. After the 3rd or 4th season, the yield of inkberry fruits gradually declines, though fruit production occurs for many years [116].

Seed dispersal: Inkberry seeds are likely dispersed by birds [19].

Seed banking: No information

Germination: In general, members of the genus Ilex have immature embryos at time of ripeness, necessitating a period of after-ripening before germination will occur [45].

Seedling establishment/growth: Natural inkberry seedlings are rare; virtually all new growth originates from sprouting [69]. A study of a slash pine savannah in Mississippi found a greater frequency of established inkberry adults and a greater density of inkberry seedlings near the base of trees (<6.5 ft or 2 m). This trend is attributed to a higher seed rain underneath trees, presumably due to the dispersion of seeds by birds [19].

Asexual regeneration: Inkberry regenerates from readily sprouting rhizomes [62,63,116,119]. Mechanical damage to inkberry plants or simply exposure of a rhizome by removing soil may stimulate a new shoot to grow [116].

SITE CHARACTERISTICS:

Elevation and topography: Inkberry commonly occurs on the lower elevations of the coastal plain [203]. It occurs at elevations from 14 to 249 feet (4.3-76 m) [8,112,121,131,152]. Inkberry grows on sites with flat to slightly undulating topography [1,48,65,97,118,131,144,194,202].

Site types: Inkberry is found in wetlands [100,103,157,174], bogs [33,45,62,63,103,157,208], seeps [45,62,63], pocosins [7,153,156,157,176,206], swamps [72,134,135,148,170,171,174], lake and pond borders [4,72,164], and creek bottoms [45,55,62,63]. It also occurs in upland areas [55,103] including pine flatwoods [33,45,48,62,63,72,153,207,208], pine barrens [153], prairies [62,63,196], terraces [184], and savannahs [38,45,48,62,63,72,153]. Inkberry is also found on coastal sand dunes [38,196] and swales [33,208].

In Nova Scotia, inkberry is commonly found on rocky barrens and dry hillsides, in addition to swampy or mesic areas [165,185].

Soils: Soils supporting inkberry include coarse till [185,186], fine sand [4,21,38,54,65,68,69,70,72,77,78,121,125,131,150,196], loam [11,68,70,131], clay [54,70,125], marl [38,54,196], and peat [41,69,72,126,148,176]. These soils may have an underlying, impervious clay hardpan [4,114]. Soils often have low organic matter [4,54,72] and nutrient content [4,38,121,126,131,184,185], and are often acidic [1,4,45,62,63,69,121,126,131,134,152,176,184,186]. Soils may be calcareous [43,196], though Halls [69] reports that inkberry grows poorly on soils with high lime content.

Though it occurs on soils with low levels of extractable phosphorus (<2.0 ppm) [70], inkberry growth may be enhanced by pine needle fall due to increased phosphorus concentrations [21].

Inkberry is common on both well-drained [1,38,185,188] and poorly drained sites [1,20,38,56,57,65,68,72,77,97,176] with moist or wet soils [4,11,17,32,56,77,102,133,144,151] and a shallow or fluctuating water table [17,57,72,148,176,206]. The soils are often seasonally wet [4,72,152] or periodically flooded [1,134,184,196], saturated in winter and droughty during the growing season [54,72,148]. In New Jersey, inkberry may be restricted to areas where the roots can extend to the water table, and inkberry seldom occurs in abundance on upland sites [114].

SUCCESSIONAL STATUS:

SEASONAL DEVELOPMENT:

• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

FIRE ECOLOGY OR ADAPTATIONS:

Fuels: Though green year-round, inkberry is highly flammable [105,106,168] due to the volatile and easily ignited substances in the plant tissues [26,94,107,163]. However, flammability is also affected by fuel moisture content, fuel load, and live moisture content. Higher leaf moisture content corresponds to a decrease in the amount of volatiles and a decreased flammability of leaves [26]. The heat value of inkberry leaves is greater than 5000 calories per gram [86]. The volatile oils in inkberry foliage ignite quickly under certain weather conditions, and when the temperature is high enough may suddenly flame to create an "intense" fire [106].

Once ignition has occurred, the intensity of a fire is strongly influenced by the structure of the fuels in allowing heat transfer and by the energy content of the fuel [166]. Over the temperature range of 200 to 900 degrees Fahrenheit (100-500 ^oC), the majority of inkberry combustion gases are derived from ether and benzene-ethanol extractives [166,175], both of which play an important role in the initiation of combustion at low temperatures (below 572 ^oF or 300 ^oC). The benzene-ethanol extractives also increase the intensity of the fire at higher temperatures (above 572 ^oF or 300 ^oC) [175]. The total extractive content, or combustible gases, of inkberry is 44.6% of foliage dry weight. A summary of inkberry foliage extractive content by dry weight is presented below [166]:

Though foliage comprises 30 to 45% of inkberry's total dry weight, inkberry stems present a greater contribution to fuels composition, comprising 55 to 69% of dry weight [129]. 

In the pine flatwoods of the southeastern United States, the presence of inkberry, especially inkberry thickets, increases the frequency, hazard, and severity of forest fires [69,82,147,192]. Average fuel weights of inkberry (tons/acre) in inkberry-dominated roughs of the longleaf-slash pine forest type in the Osceola National Forest, are presented below [22]. Average fuel weights were based on air-dry weight of 2 samples of all material less than 1 inch in diameter that usually burn in headfires under dry conditions.

Fire regimes: Inkberry is an important understory component in longleaf-slash pine and loblolly-shortleaf pine ecosystems. Longleaf-slash pine communities experience frequent surface fire, historically occurring at 1- to 8-year intervals. Surface fire in loblolly-shortleaf pine communities also occurs frequently, at 2- to 15-year intervals [195].

Fire regimes for plant communities and ecosystems in which inkberry occurs are summarized below. For further information regarding fire regimes and fire ecology of communities and ecosystems where inkberry is found, see the 'Fire Ecology and Adaptations' section of the FEIS species summary for the plant community or ecosystem dominants listed below.

POSTFIRE REGENERATION STRATEGY [183]:

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

IMMEDIATE FIRE EFFECT ON PLANT:

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:

PLANT RESPONSE TO FIRE:

Inkberry response to prescribed fire: Inkberry sprouts quickly and prolifically following fire. Following a February prescribed burn in a Georgia longleaf pine/slash pine community, inkberry responded with a rapid increase in biomass from the end of April to early June. From June to early August, the growth slowed. From August to the end of October, the growth curve flattened, with little biomass increase recorded.  Growth ceased in November [82].

On the southeastern coastal plain, inkberry may act as a major inhibitor of herbaceous plants in longleaf pine/wiregrass understories. A temporary reduction in the percent cover of inkberry by fire may provide an opportunity for expansion and colonization by herbaceous species, increasing understory plant species richness and diversity as well as the biomass productivity of grasses and shrubs. In a Georgia longleaf pine/wiregrass forest, percent cover of inkberry was significantly reduced (p<0.05) by annual, biennial, and triennial prescribed fire treatments, from 50% cover on unburned plots to 35% on burned plots [20]. Prescribed burns at 3- to 4- year intervals restrict the presence and cover of inkberry, though it resprouts quickly [66,192]. Lewis and Hart [119] observed that where inkberry stems were burned to the ground, resprouts were half as tall as unburned stems within 9 months. On 1 site in Georgia, the number of inkberry stems was significantly greater (p<0.05), though percent cover was significantly less (p<0.05), on burned versus unburned plots 9 months postfire; on a 2nd plot, no significant differences were reported [119]. During the 2nd postfire year on sites in Florida, Georgia, and South Carolina, Hilman [81] recorded number of inkberry stems per acre:

In Florida longleaf pine/slash pine stands, prescribed fire treatments significantly reduced (p<0.01) inkberry canopy coverage below 4.9 feet (1.5 m), from 14.7% to 11.1% approximately 1 postfire year [137]. In saw-palmetto-dominated shrub vegetation of Florida, inkberry cover values reached or exceeded preburn values within 3 years. Information on mean inkberry percent cover is summarized in the following table [172]:

Another study conducted on the lower coastal plain of Georgia found that slash pine-harvested sites were dominated by inkberry 6-years postharvest and remained dominated by inkberry following prescribed fire. No difference was found in the frequency of inkberry pre- and postfire. Inkberry on these sites strongly competed with slash pine seedlings, and 6-year-old slash pine seedlings competing with inkberry were significantly smaller in diameter (p<0.025) than those not subject to the same competition [197].

Even without burning, inkberry persists and often increases [69,199,209], reducing forage production [199]. In Georgia flatwoods, inkberry foliage cover increased 3-fold during 20 years of continuous fire exclusion. In 1 series of plots unburned for 30 years, inkberry occurred as a dominant shrub [69]. On plots in Georgia last burned in 1941, inkberry percent cover increased from 7% in 1942 to 14% in 1951 and 39% in 1963 [80].

Seasonal patterns of carbohydrate levels in roots and stems of inkberry are unaffected by season of burning. Recovery of carbohydrates occurs within 1 year, regardless of the time of year inkberry is burned [69,80].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:

FIRE MANAGEMENT CONSIDERATIONS:

Contribution to fuels: Without frequent fire, the herbaceous understory of southeastern pine flatwoods is replaced by highly flammable species, including inkberry [34,169,198], which is dangerous to treat with fire due to its volatility. This may result in continued accumulation of wildland fuel loads and increasingly difficult and dangerous conditions for fire suppression [198]. A general increase in the size, number, and severity of fires accompanies increasing age of vegetation in the gallberry-saw-palmetto/wiregrass fuel type. This increase is attributed to fuel accumulation as the vegetation ages, and frequent burning may help manage these hazardous fuel levels [169]. Experimental burns in longleaf pine forests with an understory dominated by inkberry produced "very hot fires" [76]. On areas protected from fire in longleaf pine forests, inkberry attains much greater size than on frequently burned areas [79]. On sites in Georgia, prescribed fire reduced vegetative inkberry fuel 0 to 70% [83].

Following an initial burn on a longleaf pine site in Florida and longleaf pine and loblolly pine sites in South Carolina, fuel weights of inkberry were recorded. Total (total litter fuel plus total understory fuel) mean weight of inkberry is presented below in pounds per acre. On the Florida site, inkberry fuels increased for about  6 years before leveling off, while in South Carolina, fuel weights continued to increase. On these sites, inkberry represented 1.8 to 6.6% of the total fuels measured [168].

In uniform 4-year-old saw-palmetto-inkberry roughs, headfire spread rates can exceed 0.5 mph (0.8 km/hr) with flame lengths in excess of 20 feet (6 m) and fireline intensities of 2,000 Btu/ft/s under "good" burning conditions [195].

Inkberry forms specific fuel types with wax myrtle [210] and saw-palmetto [200]. A model for predicting particulate matter emissions from fires in the saw-palmetto-inkberry fuel type has been developed [200].

Inkberry control using fire: Campbell [29] states that annual winter burning in longleaf pine forests may alleviate inkberry competition with longleaf pine seedlings, and over a 20-year period, annual winter burning resulted in a significant decrease (p<0.05) in inkberry crown spread [117]. Following a winter burn in South Carolina, inkberry relative percent cover was recorded at 4.0%, contrasted with 12.1% on the control site [59]. However, Clewell [34] found that summer prescribed burning provides better control of inkberry than winter burning. Spring and summer prescribed burning reduces the frequency and size of inkberry plants, thinning out dense clumps that can shade out grasses and forbs [52]. Inkberry thrives with frequent winter burning [69,93]; though fire top-kills the plants, within a few months as many or more new sprouts grow [93]. Hilman and Hughes [80] found that resprouts of inkberry burned in January appear in late February, growing most rapidly from mid-April to mid-May and averaging 2 feet (0.6 m) in height by November.

Though fire will not eliminate inkberry, annual summer burning may be effective in retarding growth [80,93]. In 1 study, inkberry stands were burned for 3 consecutive years in each of the following months: January, April, June, August, and October. Regardless of the month of burning, the number of stems increased following the initial burn and remained fairly stable for the next 2 years, with sprouts growing approximately 2 feet (0.6 m) the 1st year after the initial burn. The effects of these consecutive burns are presented in the following table according to month treated [93].

However, in a study of longleaf pine communities in Alabama, inkberry contributed 60 to 78% of shrub biomass on winter, spring, and summer burn treatments, as well as on control plots. There was no difference between the effects of dormant season and growing season burns [110]. Repeated burning, even at the most vulnerable time, is not likely to eradicate inkberry [93,161].

• IMPORTANCE TO LIVESTOCK AND WILDLIFE
• PALATABILITY
• NUTRITIONAL VALUE
• COVER VALUE
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER MANAGEMENT CONSIDERATIONS

IMPORTANCE TO LIVESTOCK AND WILDLIFE:

The fruit, leaves, and twigs of inkberry are an important component of white-tailed deer [69,97,160,177,205] and Sambar deer [177] diets; inkberry is moderately preferred browse for white-tailed deer [37]. It is browsed by white-tailed deer primarily in  the fall and winter [69,177,205], and by Sambar deer in both the fall and spring [177].

Inkberry fruit may be important in the diets of small mammals [69,111], and it is an important component of the raccoon winter diet in Alabama [97]. The fruit and leaves provide food for black bears in the fall [42,69,71,73].

Inkberry fruit and leaves are a minor component of wild turkey [50,69,97,160], bobwhite quail [69,97,140,160], and songbird diets [9,69,97].

PALATABILITY:

NUTRITIONAL VALUE:

Little difference was observed in the nitrogen and phosphorus content of inkberry on burned and unburned areas. Lignin content was higher on unburned areas during the early leaf stage, but higher on burned areas during anthesis. Ether content of inkberry was much lower on burned areas, while calcium content was higher on burned areas [116].

Nutrient composition of inkberry was also evaluated in slash pine stands with a saw-palmetto/inkberry understory in Florida and Georgia. Composition of live and dead inkberry stems by diameter is presented below. Data include average values and standard errors [88]:

A study conducted in slash pine plantations compared the nutrient content of inkberry current-year growth in spring and summer, and evaluated the nutrient content in response to the residual effect of fertilization. A summary of that information is presented in the table below [205].

IVOMD levels were not nutritionally adequate for white-tailed deer, and phosphorus concentrations were well below proposed maintenance levels (0.28-0.3% at time of study). For the most part, maintenance levels of calcium (0.25%) and nitrogen (1.2%) for white-tailed deer were met or exceeded by inkberry [205].

COVER VALUE:

VALUE FOR REHABILITATION OF DISTURBED SITES:

Artificial regeneration: Propagation of inkberry may be achieved by transplanting the suckering shoots that develop around the base of plants. Inkberry cuttings root year-round when provided with 1,000 parts per million indolebutyric acid quick dip (5 seconds) or a commercial rooting powder, though untreated cuttings will also root [45]. 

Inkberry seed germination may be achieved by sowing the seed in a warm greenhouse, and then waiting for germination to occur. In greenhouse tests, 57% of the freshly collected, cleaned seeds germinated within 8 months [45]. Good germination is achieved by planting cleaned, collected seeds in moist, fungus-free medium at 60 degrees Fahrenheit (15.5 ^oC) until seeds sprout [69,91]. Seeds held at 60 degrees Fahrenheit (15.5 ^oC) germinated more quickly and in greater numbers than seeds held at 40, 50, 70, 80, or 90 degrees Fahrenheit (4, 10, 21, 27, or 32 ^oC). Germinations of 24, 56, and 70% occurred in 49, 69, and 102 days, respectively, at 60 degrees Fahrenheit (15.5 ^oC). Inkberry will germinate at this temperature in complete darkness, under constant illumination, or completely submerged in water [91]. Germination capacities of 70 to 95% were achieved in laboratory tests with seeds planted immediately following collection; however, germination may not occur until the 2nd or 3rd spring, suggesting potential inkberry seed banking. Some benefit may be obtained by stratifying seeds at alternating temperatures of 68 degrees Fahrenheit (20 ^oC) at night and 86 degrees Fahrenheit (30 ^oC) during the day for 60 days, followed by 60 days at 41 degrees Fahrenheit (5 ^oC) [13].

OTHER MANAGEMENT CONSIDERATIONS:

Inkberry thrives under browsing pressure [64]. In Georgia longleaf-slash pine forests, inkberry increased greatly with grazing, possibly due to reduced grass competition. The greatest increases occurred on more heavily grazed sites [67]. Extensive inkberry cover contributes to reduced herbage yields of pineland threeawn, Curtiss dropseed, bluestem grasses, and a variety of forbs [119]. Inkberry thickets often reduce or exclude more desirable forage species [68,69], reducing the grazing capacity of these areas for domestic livestock [69]. 

On sites in Florida, Neary [142] found that the height and diameter of loblolly pine seedlings decreased with increasing inkberry densities. Mechanical treatments may be useful in controlling or reducing competition by inkberry with tree seedlings [25]; however, inkberry shrubs that are mechanically cut develop new shoots from rhizomes and from adventitious buds that develop low on the cut stem, approximately 2 to 3 inches (5-7.5 cm) from the ground [116]. Though inkberry cover, frequency, and biomass are reduced by harvesting activities, inkberry might dominate sites in Florida where the slash pine overstory has been harvested [36]. In longleaf pine communities of Alabama, inkberry contributed 64 to 73% of shrub biomass 23 years following no treatment as well as chemical and mechanical understory treatments [110]. On sites in Mississippi, inkberry crown cover was significantly reduced (p<0.05) by a combination of cultivation and fertilizer treatments. Plowing and disking operations reduced crown cover from 58.3% to 21.5% 12 years after treatment. This mechanical treatment combined with low, medium, and high fertilizer rates (nitrogen, phosphorus, potassium) reduced crown cover to 10.7%, 4.1%, and 5.4%, respectively [204].

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Related categories for SPECIES: Ilex glabra | Inkberry