SPECIES: Ilex glabra


Ilex glabra: INTRODUCTORY

INTRODUCTORY

SPECIES: Ilex glabra
AUTHORSHIP AND CITATION:
Anderson, Michelle D. 2001. Ilex glabra. 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/ [].

ABBREVIATION:
ILEGLA

SYNONYMS:
No entry

NRCS PLANT CODE [191]:
ILGL

COMMON NAMES:
inkberry
gallberry
bitter gallberry

TAXONOMY:
The currently accepted name of inkberry is Ilex glabra (L.) Gray (Aquifoliaceae) [101].

LIFE FORM:
Shrub

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
Maine: The state legal status of inkberry is threatened, and the state rarity status is S1 (critically imperiled) [122].

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].


DISTRIBUTION AND OCCURRENCE

SPECIES: Ilex glabra
GENERAL DISTRIBUTION:
Inkberry occurs on the Atlantic coastal plain from Nova Scotia and the New England states south to southern Florida, and west on the Gulf coastal plain to eastern Texas [50,64,65,101,153,165,174]. The PLANTS database provides a distributional map of inkberry.

ECOSYSTEMS [60]:
FRES11 Spruce-fir
FRES12 Longleaf-slash pine
FRES13 Loblolly-shortleaf pine
FRES14 Oak-pine
FRES15 Oak-hickory
FRES16 Oak-gum-cypress
FRES41 Wet grasslands

STATES:
AL AR CT DE
FL GA ME MD
MA MS NH NJ
NY NC RI SC
TX VT VA WV
DC NS

BLM PHYSIOGRAPHIC REGIONS [12]:
None

KUCHLER [109] PLANT ASSOCIATIONS:
K073 Northern cordgrass prairie
K076 Blackland prairie
K077 Bluestem-sacahuista prairie
K078 Southern cordgrass prairie
K079 Palmetto prairie
K080 Marl everglades
K083 Cedar glades
K088 Fayette prairie
K089 Black Belt
K090 Live oak-sea oats
K091 Cypress savanna
K092 Everglades
K096 Northeastern spruce-fir forest
K100 Oak-hickory forest
K103 Mixed mesophytic forest
K105 Mangrove
K106 Northern hardwoods
K108 Northern hardwoods-spruce forest
K110 Northeastern oak-pine forest
K111 Oak-hickory-pine
K112 Southern mixed forest
K113 Southern floodplain forest
K114 Pocosin
K115 Sand pine scrub
K116 Subtropical pine forest

SAF COVER TYPES [53]:
18 Paper birch
19 Gray birch-red maple
20 White pine-northern red oak-red maple
21 Eastern white pine
22 White pine-hemlock
39 Black ash-American elm-red maple
40 Post oak-blackjack oak
43 Bear oak
44 Chestnut oak
45 Pitch pine
51 White pine-chestnut oak
52 White oak-black oak-northern red oak
53 White oak
57 Yellow-poplar
59 Yellow-poplar-white oak-northern red oak
64 Sassafras-persimmon
65 Pin oak-sweetgum
69 Sand pine
70 Longleaf pine
71 Longleaf pine-scrub oak
72 Southern scrub oak
73 Southern redcedar
74 Cabbage palmetto
75 Shortleaf pine
76 Shortleaf pine-oak
78 Virginia pine-oak
79 Virginia pine
80 Loblolly pine-shortleaf pine
81 Loblolly pine
82 Loblolly pine-hardwood
83 Longleaf pine-slash pine
84 Slash pine
85 Slash pine-hardwood
87 Sweetgum-yellow-poplar
88 Willow oak-water oak-diamondleaf (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
95 Black willow
96 Overcup oak-water hickory
97 Atlantic white-cedar
98 Pond pine
100 Pondcypress
101 Baldcypress
102 Baldcypress-tupelo
103 Water tupelo-swamp tupelo
104 Sweetbay-swamp tupelo-redbay
105 Tropical hardwoods
106 Mangrove
108 Red maple
109 Hawthorn
110 Black oak
111 South Florida slash pine

SRM (RANGELAND) COVER TYPES [179]:
726 Cordgrass
806 Gulf Coast salt marsh
807 Gulf Coast fresh marsh
808 Sand pine scrub
809 Mixed hardwood and pine
810 Longleaf pine-turkey oak hills
811 South Florida flatwoods
812 North Florida flatwoods
813 Cutthroat seeps
814 Cabbage palm flatwoods
815 Upland hardwood hammocks
816 Cabbage palm hammocks
817 Oak hammocks
818 Florida salt marsh
819 Freshwater marsh and ponds
820 Everglades flatwoods
821 Pitcher plant bogs
822 Slough

HABITAT TYPES AND PLANT COMMUNITIES:
In Nova Scotia rocky heathlands, inkberry is common in black huckleberry (Gaylussacia baccata) communities [186] and the mixed shrub vegetation type. It occurs with serviceberry (Amelanchier spp.) [185], wild sarsaparilla (Aralia nudicaulis), sheep-laurel (Kalmia angustifolia), wintergreen (Gaultheria procumbens) [186], sweetgale (Myrica gale) [185], northern bayberry (Morella pensylvanica) [185,186], lowbush blueberry (Vaccinium angustifolium) [186], and possumhaw (Viburnum nudum). Tree species include red maple (Acer rubrum) [185,186], American green alder (Alnus viridis ssp. crispa) [186], gray birch (Betula populifolia), black spruce (Picea mariana) [185,186], eastern white pine (Pinus strobus) [186], and quaking aspen (Populus tremuloides) [185]. Inkberry also occurs in dense spruce (Picea spp.) woods of Nova Scotia [165].

Inkberry is associated with pitch pine (Pinus rigida) on the Atlantic coastal plain [6,14,51,120,124,182]. Other associated tree species in these communities include white oak (Quercus alba) [6,14], bear oak (Quercus ilicifolia) [14,26,34,124], blackjack oak (Quercus marilandica) [124,182], dwarf chinkapin oak (Quercus prinoides) [26,34], post oak (Quercus stellata) [124,182], black oak (Quercus velutina) [6,14,182], shortleaf pine (Pinus echinata) [14,34,182], and sassafras (Sassafrass albinum) [124]. Associated shrub species include downy serviceberry (Amelanchier arborea) [26], coastal sweetpepperbush (Clethra alnifolia) [34,182],  black huckleberry [6,14,26,34,124,182], dwarf huckleberry (Gaylussacia dumosa) [34], blue huckleberry (Gaylussacia frondosa) [26,34,51,124,182], wintergreen [14,26,182], sheep-laurel [26,51,124,182], mountain-laurel (Kalmia latifolia) [26,34,124], swamp doghobble (Leucothoe racemosa) [182], maleberry (Lyonia ligustrina) [14], piedmont staggerbush (Lyonia mariana) [182], bayberry (Morella spp.) [6,182], black chokecherry (Photinia melanocarpa) [26], black cherry (Prunus serotina) [13], flameleaf sumac (Rhus copallinum) [26], cat greenbrier (Smilax glauca) [26,182], roundleaf greenbrier (Smilax rotundifolia) [182], Virginia tephrosia (Tephrosia virginiana) [14], lowbush blueberry [26], and Blue Ridge blueberry (Vaccinium pallidum) [14,26,124,182]. Additional understory associates include Pennsylvania sedge (Carex pensylvanica) [14,26], leatherleaf (Chamaedaphne calyculata) [51], sweetfern (Comptonia peregrina) [14,26,34,124,182], western brackenfern (Pteridium aquilinum) [14,26], and anisescented goldenrod (Solidago odora) [14].

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 [8,148,157], commonly occurring with southern bayberry [201], swamp titi (Cyrilla racemiflora) [11,49,148,156,157,176,201], cane (Arundinaria gigantea) [92,148,201], red chokecherry (Photinia pyrifolia) [157], cinnamon fern [201], coastal sweetpepperbush [92], greenbrier (Smilax spp.) [148,156,157,176,201], swamp doghobble [49], fetterbush lyonia [11,49,92,157], honeycup (Zenobia pulverulenta) [11,148,156,176], dwarf huckleberry [43], highbush blueberry [49,157], wax myrtle (Morella cerifera) [92,157,176,201,206], northern bayberry [43], and moss (Sphagnum spp.) [43,157,201]. Also occurring in pocosin vegetation is scattered red maple [156,201], Atlantic white-cedar [156], loblolly bay (Gordonia lasianthus) [11,156,157,176], sweetgum (Liquidambar styraciflua) [201], sweetbay (Magnolia virginiana) [11,49,148,156,157,176,201], redbay [11,156,157,176], pond pine (Pinus serotina) [11,92,157,176,206], and bald cypress (Taxodium distichum) [156].

In pond pine forests, inkberry occurs with bluestem (Andropogon spp.) [94], cane [18,94], sweetpepperbush [18,74,94], swamp cyrilla [18,74], huckleberry (Gaylussacia spp.) [74,94], large gallberry [18], St. Johnswort (Hypericum spp.) [94], maleberry [74], fetterbush lyonia [46,74,94], wax myrtle [18,46,59], cinnamon fern, sumac (Rhus spp.) [94], saw-palmetto (Serenoa repens) [18,46,74], laureleaf greenbrier (Smilax laurifolia) [18,74], honeycup [18], and blueberry [46,94]. Other tree species include red maple [18,74], Atlantic white-cedar [74], loblolly bay], sweetgum [18], sweetbay [74,94], redbay [46], and loblolly pine (Pinus taeda) [74].

Inkberry is found in bay forests with loblolly bay [5,20,126,176], sweetbay [5,126,152,176], and redbay [5,20,126,176]. Understory species in these communities include coastal sweetpepperbush, cane [126], buckwheat tree (Cliftonia monophylla) [20,126], swamp titi [126,176], huckleberry [176], dahoon (Ilex cassine) [5,20,126,176], large gallberry [126,176], deciduous holly (Ilex decidua) [126], smooth winterberry (Ilex laevigata) [176], American holly [126], yaupon (Ilex vomitoria) [20], Virginia sweetspire (Itea virginica), sheep-laurel, swamp doghobble [126], maleberry [5,126], fetterbush lyonia [5,20,126,176], bayberry [5,20,126,176], swamp azalea [176], greenbrier [126,176], poison sumac (Toxicodendron vernix) [20], 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) [56]. 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) [56], coastal sweetpepperbush [170,171], flatsedge (Cyperus spp.) [56], swamp titi [170,171], southern umbrella-sedge (Fuirena scirpoidea), peelbark St. Johnswort (Hypericum fasciculatum), clustered bushmint (Hyptis alata) [56], dahoon [56,170,171], large gallberry, Virginia sweetspire, swamp doghobble [170,171], primrose-willow (Ludwigia spp.), rusty staggerbush (Lyonia ferruginea) [56], fetterbush lyonia [170,171], wax myrtle, wild pennyroyal (Piloblephis rigida), camphorweed (Pluchea spp.), cabbage palmetto (Sabal palmetto), licorice weed (Scoparia dulcis) [56], 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) [67,116], slash pine [4,5,34,40,45,81,96,97,125,128,129,130,133,144,146,162,184], longleaf pine (Pinus palustris) [4,16,17,32,33,41,45,61,79,80,81,125,128,129,133,146,162,167,176,184], pond pine [4,133,184], and loblolly pine [33,61,62,118,131,159,167]. Several oak species occur in flatwoods communities, including Chapman oak (Quercus chapmanii) [41], southern red oak (Quercus falcata) [41,118,121,159], sand live oak (Quercus geminata) [184], bluejack oak (Quercus incana) [41], turkey oak (Quercus laevis) [41], laurel oak (Quercus laurifolia) [13,41,81], runner oak (Quercus margarettiae) [125], blackjack oak [118,121], dwarf live oak (Quercus minima) [4,41,58,79,184], myrtle oak (Quercus myrtifolia) [58], water oak (Quercus nigra) [4,13,62,81,159], willow oak (Quercus phellos) [118,121,159], running oak (Quercus pumila) [133,162,184], post oak [62,118,121], and live oak [4,13,32,45]. Other tree species occurring in these communities include red maple [4,41,62,81], persimmon (Diospyros spp.) [17], ash (Fraxinus spp.) [4], sweetgum [4,13,61,62,118,121,131,159], loblolly bay [41], sweetbay [41,74,81], blackgum [61,62,74,118,121,131,159], redbay [13,63,74], swamp bay (Persea palustris) [81], shortleaf pine [61,62,131], spruce pine (Pinus glauca) [108], and pond cypress [45]. 

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 [105]. Understory associates of inkberry in southeastern pine flatwoods include cane [57,61], pawpaw (Asimina spp.) [40], buckwheat tree [74], tarflower [1,2,4,41], sedges (Carex spp.) [57], sweetpepperbush [74,118,121,131], flowering dogwood (Cornus florida) [17,100], black titi [190], swamp titi [74,104,167], huckleberry [4,5,16,17,40], St. Johnswort [58,74], large gallberry [32,104,146,176,190], yaupon [16,99,100,146], mountain-laurel [104], hairy-laurel (Kalmia hirsuta) [86,184], sand myrtle (Leiophyllum buxifolium) [104], fetterbush lyonia [4,5,41,45,63,104,125,130,144,184,190], rusty staggerbush [116], coastalplain staggerbush (Lyonia fruticosa) [22,116], piedmont staggerbush [104,176], wax myrtle [4,16,32,40,41,45,61,81,144,150,159,176,184], southern bayberry [121], red chokecherry [58], azalea (Rhododendron spp.) [58], beakrushes (Rhynchospora spp.) [57], sumac [16,45], blackberry (Rubus spp.) [16,17,41,112,146], cabbage palm (Sabal palmetto) [45,196], saw-palmetto [4,5,16,40,41,45,79,80,81,125,128,129,144,150], greenbrier [74,190], blueberry [5,16,17,61,74,81,100,125,162,176,184], and viburnum [41]. Grasses occurring in these communities include threeawn (Aristida spp.) [4,41,79,86,98,125,130,133,146,162,167], common carpet grass (Axonopus fissifolius) [22,70], toothache grass (Ctenium aromaticum) [22,115], panic grass (Panicum spp.) [3,37,38,57,74,115,118,146], muhly (Muhlenbergia spp.) [57], bluestem (Schizachyrium and Andropogon spp.) [37,41,57,61,70,99,100,115,118,125,146,146], Indian grass (Sorghastrum spp.) [22,57], Curtis' dropseed (Sporobolus curtissii) [22,38,70,118], 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) [59].

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 [40], sand heath (Ceratiola ericoides), garberia (Garberia heterophylla) [139], huckleberry [40], St. Andrew's cross (Hypericum hypericoides), Carolina holly (Ilex ambigua) [139], dahoon [40], staggerbush (Lyonia spp.), wax myrtle [40,139], Chapman oak [139], sand live oak [40,139], laurel oak [40], myrtle oak, redbay, silk bay, devilwood (Osmanthus americanus), oak mistletoe (Phoradendron leucarpum) [139], sand pine (Pinus clausa) [40,139], scrub plum (Prunus geniculata) [139], hog plum (Prunus umbellata) [40], tallow wood, western brackenfern, flameleaf sumac (Rhus copallinum), scrub palmetto (Sabal etonia) [139], saw palmetto [40,139], 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 [40].

In dry prairie vegetation, inkberry occurs with pawpaw (Asimina spp.) [40,45], tarflower [136,158], sedge (Cyperus spp.) [30,40], coastalplain St. Johnswort (Hypericum brachyphyllum) [136], whitehead bogbutton (Lachnocaulon anceps) [91], rusty staggerbush [158], coastalplain staggerbush [55,91,136], fetterbush lyonia [45,55,136], wax myrtle [55], eastern prickly-pear (Opuntia humifusa) [30], sand live oak [91], dwarf live oak [55,102,136], running oak [45,55], plumed beaksedge (Rhynchospora plumosa) [91], saw-palmetto [4,40,55,91,102,136,158], shiny blueberry [55,136], Darrow's blueberry (Vaccinium darrowii) [91], common carpetgrass [102], Curtis' dropseed [82], threeawn [4,82,91,136,158], sawgrass (Cladium spp.) [40], cutthroat grass (Panicum abscissum) [209], little bluestem (Schizachyrium scoparium), lopsided Indian grass (Sorghastrum secundum), narrowleaf silkgrass (Pityopsis graminifolia), and vanillaleaf (Carphephorus odoritissimus) [91].

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

Florida [5]
North Carolina [105,201]

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Ilex glabra
GENERAL BOTANICAL CHARACTERISTICS:
Inkberry is a native evergreen shrub [51,58,71,100,101,153,181,192] with dense foliage [100], growing 2 to 12 feet (0.5-4 m) tall [15,47,71,192] and 6.5 to 10 feet (2-3 m) wide [47]. The persistent leaves of inkberry are 1.2 to 1.8 inches (3-4.6 cm) long and 0.6 to 0.8 inch (1.5-2 cm) wide [71].

Inkberry is a stoloniferous or rhizomatous shrub forming extensive, dense colonies [47,50,65,71,110,116,153]. 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 [47,71] or in clumps of up to 3 flowers [47]. Inkberry fruits grow individually or in small clusters, with 3 to 9 seeds [71].

RAUNKIAER [154] LIFE FORM:
Phanerophyte
Geophyte

REGENERATION PROCESSES:
Inkberry regenerates by sprouting [64,65,116,119] and from seed [128].

Breeding system: Inkberry is dioecious [47,64,65,71,153].

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 [71]. 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 [71,82,116]. 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 [98]. 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 [21].

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 [47].

Seedling establishment/growth: Natural inkberry seedlings are rare; virtually all new growth originates from sprouting [71]. 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 [21].

Asexual regeneration: Inkberry regenerates from readily sprouting rhizomes [64,65,116,119]. Mechanical damage to inkberry plants or simply exposure of a rhizome by removing soil may stimulate a new shoot to grow [116].

SITE CHARACTERISTICS:
Climate: Inkberry occurs in areas subject to alternating seasons of flooding and droughty conditions [4], with a summer rainy season (80% of precipitation) from May through October and a winter dry season from November through April [56]. The climate is humid mesothermal with mild winters and warm summers [61,184], and precipitation typically exceeds evapotranspiration [61]. Average annual precipitation on sites supporting inkberry ranges from 40.5 to 64 inches (1,030-1,630 mm) [22,56,61,112,118,184,202]. Average annual temperatures in the Southeast range from 39 to 92 degrees Fahrenheit (15-33 oC) [22,56,61,112,184]. The growing season in the Southeast ranges from 240 to 365 days [184,196]. Northern forms of inkberry are cold hardy to -15 or -20 degrees Fahrenheit (-26 or -29 oC).

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) [9,112,121,131,152]. Inkberry grows on sites with flat to slightly undulating topography [1,50,67,98,118,131,144,194,202].

Site types: Inkberry is found in wetlands [101,104,157,174], bogs [35,47,64,65,104,157,208], seeps [47,64,65], pocosins [8,153,156,157,176,206], swamps [74,134,135,148,170,171,174], lake and pond borders [4,74,164], and creek bottoms [47,57,64,65]. It also occurs in upland areas [57,104] including pine flatwoods [35,47,50,64,65,74,153,207,208], pine barrens [153], prairies [64,65,196], terraces [184], and savannahs [40,47,50,64,65,74,153]. Inkberry is also found on coastal sand dunes [40,196] and swales [35,208].

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 [4,23,40,56,67,70,71,72,74,79,80,121,125,131,150,196], loam [13,70,72,131], clay [56,72,125], marl [40,56,196], and peat [43,71,74,126,148,176]. These soils may have an underlying, impervious clay hardpan [4,114]. Soils often have low organic matter [4,56,74] and nutrient content [4,40,121,126,131,184,185], and are often acidic [1,4,47,64,65,71,121,126,131,134,152,176,184,186]. Soils may be calcareous [45,196], though Halls [71] 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) [72], inkberry growth may be enhanced by pine needle fall due to increased phosphorus concentrations [23].

Inkberry is common on both well-drained [1,40,185,188] and poorly drained sites [1,22,40,58,59,67,70,74,79,98,176] with moist or wet soils [4,13,19,34,58,79,103,133,144,151] and a shallow or fluctuating water table [19,59,74,148,176,206]. The soils are often seasonally wet [4,74,152] or periodically flooded [1,134,184,196], saturated in winter and droughty during the growing season [56,74,148]. 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:
Inkberry occurs in disturbed [8,115], undisturbed [115] and mature [167] vegetation. It is often part of residual vegetation following disturbance [143]. Inkberry was present in an old-growth longleaf pine stand in Alabama that was undisturbed for over 45 years. It remained on the site following 3 understory prescribed burns at 1-year intervals [193]. Inkberry is moderately shade tolerant [47], occurring under both open [1,104] and closed [1,3] canopies. When overstory trees are removed by disturbance, inkberry may dominate successional vegetation in the holly (Ilex spp.)-titi (Cyrilla spp.)-honeycup (Zenobia spp.) association [148].

SEASONAL DEVELOPMENT:
Inkberry flowers from February to late June [15,47,50,71], varying by latitude [50]. In North and South Carolina, inkberry flowers in May and June [153]; in the New England states, it flowers from mid-June into July [174].

Inkberry produces fruit as early as late July [71]; ripening generally occurs between September and November [15,47,98,153]. Inkberry fruits persist on plants through the winter and begin falling when new growth starts [47,71,98]. The fruits are often still present when new flowers emerge [47].

FIRE ECOLOGY

SPECIES: Ilex glabra
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: Inkberry survives fire by sprouting from the root crown and rhizomes [1,29,68,81,82,82,113,166].

Fuels: Though green year-round, inkberry is highly flammable [7,106,168] due to the volatile and easily ignited substances in the plant tissues [11,28,107,163]. 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 [28]. The heat value of inkberry leaves is greater than 5000 calories per gram [88]. 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 [7].

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]:

Ether extractives Benzene-ethanol extractives Nonextractives
% of total C (%) H (%) % of total C (%) H (%) % of total C (%) H (%)
11.8 76.5 11.4 32.8 50.8 6.4 55.4 49.4 6.7

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 [71,84,147,192]. 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 [24]. 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.

Open stands*, age of rough Dense stands**, age of rough
1 year 2 years 3-5 years 10-15 years 1 year 2 years 3-5 years 10-15 years
4.4 5.8 5.1 7.5 8.0 8.8 8.2 13.0
* open stands had no pines over 5 feet tall within 20 feet of the sampling area
** dense stands were those with at least 4 pines with DBH of 4 inches or greater within 10 feet of the sampling area

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.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
mangrove Avicennia nitida-Rhizophora mangle 35-200 [141]
sugarberry-America elm-green ash Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica < 35 to 200
Atlantic white-cedar Chamaecyparis thyoides 35 to > 200 [195]
northern cordgrass prairie Distichlis spicata-Spartina spp. 1-3 [145]
black ash Fraxinus nigra < 35 to 200 
yellow-poplar Liriodendron tulipifera < 35 [195]
Everglades Mariscus jamaicensis < 10 
melaleuca Melaleuca quinquenervia < 35 to 200 [141]
wheatgrass plains grasslands Pascopyrum smithii < 35 [145]
shortleaf pine Pinus echinata 2-15 
shortleaf pine-oak Pinus echinata-Quercus spp. < 10 
slash pine Pinus elliottii 3-8 
slash pine-hardwood Pinus elliottii-variable < 35 
sand pine Pinus elliottii var. elliottii 25-45 [195]
South Florida slash pine Pinus elliottii var. densa 1-5 
longleaf-slash pine Pinus palustris-P. elliottii 1-4 [141,195]
longleaf pine-scrub oak Pinus palustris-Quercus spp. 6-10 [195]
pitch pine Pinus rigida 6-25 [25,77]
pocosin Pinus serotina 3-8 
pond pine 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 pine Pinus taeda 3-8 
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 [195]
aspen-birch Populus tremuloides-Betula papyrifera 35-200 [48,195]
black cherry-sugar maple Prunus serotina-Acer saccharum > 1000 
oak-hickory Quercus-Carya spp. < 35
northeastern oak-pine Quercus-Pinus spp. 10 to < 35 [195]
oak-gum-cypress Quercus-Nyssa-spp.-Taxodium distichum 35 to > 200 [141]
southeastern oak-pine Quercus-Pinus spp. < 10
white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra < 35 
bear oak Quercus ilicifolia < 35 >
chestnut oak Q. 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 
live oak Quercus virginiana 10 to< 100 [195]
cabbage palmetto-slash pine Sabal palmetto-Pinus elliottii < 10 [141,195]
blackland prairie Schizachyrium scoparium-Nassella leucotricha < 10
Fayette prairie Schizachyrium scoparium-Buchloe dactyloides < 10
tule marshes Scirpus and/or Typha spp. < 35 
southern cordgrass prairie Spartina alterniflora 1-3 [145]
baldcypress Taxodium distichum var. distichum 100 to > 300 
pondcypress Taxodium distichum var. nutans < 35 [141]
eastern hemlock-yellow birch Tsuga canadensis-Betula alleghaniensis > 200 [195]
elm-ash-cottonwood Ulmus-Fraxinus-Populus spp. < 35 to 200 [48,195]

POSTFIRE REGENERATION STRATEGY [183]:
Tall shrub, adventitious bud/root crown
Rhizomatous shrub, rhizome in soil

FIRE EFFECTS

SPECIES: Ilex glabra
IMMEDIATE FIRE EFFECT ON PLANT:
Fire generally top-kills inkberry [29,71,95,113,161], reducing height for 2 to 3 years [29]. A single fire frequently consumes or kills much of the foliage of inkberry, but seldom kills the entire plant [85].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
No entry

PLANT RESPONSE TO FIRE:
Inkberry resprouts from rhizomes and the root crown following fire [1,29,68,81,82,82,113,166]. Sprouting occurs within months, and often results in a greater number of stems than were present before the fire [29,71,82,113,116]. When inkberry is top-killed, 2 to 4 new shoots grow from the base of the old stem or from nearby along the rhizome. Stems may grow 2 feet (0.6 m) or more in the 1st growing season following top-kill; without fire, normal growth is 4 to 6 inches (10-15 cm) in a season. Where stems are defoliated but not killed, twigs may be produced earlier in the season than on unburned plants, and new sprouts may also develop [116]. A number of sprouts die the 2nd year after fire, but foliage cover per stem increases gradually in the absence of fire. After 2 or 3 years, the number of stems and the amount of foliage cover return to approximate prefire levels [82]. Some stems eventually surpass and suppress others, and as the stand ages the number of stems is again reduced. The percent cover of inkberry on southeastern coastal plain sites was 5.6% 1 year postfire, increasing to 8.3% after 8 years, and decreasing to 5.5% after the 9th postfire growing season [116]. Inkberry may actually thrive on occasional fires, especially on moist sites [66].

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 [84].

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 [22]. Prescribed burns at 3- to 4- year intervals restrict the presence and cover of inkberry, though it resprouts quickly [68,192]. 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 [83] recorded number of inkberry stems per acre:

  Sites
shrub height Charlotte, FL Baker, FL Berrien, GA Beaufort, SC
<1 foot tall 3,267 3,025 605 363
1-3 feet tall 6,292 15,246 19,239 15,004
3+ feet tall 0 0 14520 726

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]:

  Prefire 6 postfire months  12 postfire months  18 postfire months  24 postfire months  36 postfire months 
height <1.6 feet 0.4 6.0 8.4 7.9 4.9 3.2
height >1.6 feet  13.7 0.0 1.9 3.0 8.7 14.3

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 [71,199,209], 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 [71]. On plots in Georgia last burned in 1941, inkberry percent cover increased from 7% in 1942 to 14% in 1951 and 39% in 1963 [82].

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 [71,82].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
No entry

FIRE MANAGEMENT CONSIDERATIONS:
Inkberry grows vigorously on sites where the overstory has been removed by severe fire [46]. It shows high fruit production a few years after fire [34,111]; burning at 3-year intervals may optimize fruit production in open slash pine forests [111].

Contribution to fuels: Without frequent fire, the herbaceous understory of southeastern pine flatwoods is replaced by highly flammable species, including inkberry [36,169,198], 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" [78]. On areas protected from fire in longleaf pine forests, inkberry attains much greater size than on frequently burned areas [81]. On sites in Georgia, prescribed fire reduced vegetative inkberry fuel 0 to 70% [85].

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].

Time since fire (years)
1 2 3 4 12 (control)
Florida inkberry 224 429 -- 747 708
South Carolina inkberry 152 244 681 775 1,504

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 [31] 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 [61]. However, Clewell [36] 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 [54]. Inkberry thrives with frequent winter burning [71,95]; though fire top-kills the plants, within a few months as many or more new sprouts grow [95]. Hilman and Hughes [82] 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 [82,95]. 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 [95].

Month burned Effect of 3 annual, consecutive prescribed burns
  Height Growth Foliage Cover
January Little or no effect on annual height growth of sprouts  
April Little or no effect on annual height growth of sprouts  
June Little or no effect on annual height growth of sprouts Decreased substantially following 3rd annual burn, though 1 or 2 annual burns had little effect
August Reduced annual height growth of sprouts Decreased substantially following 3rd annual burn, though 1 or 2 annual burns had little effect
October Reduced annual height growth of sprouts Decreased substantially following 3rd annual burn, though 1 or 2 annual burns had little effect

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 [95,161].

Burning in ungrazed and lightly grazed longleaf pine forests may offer effective control of inkberry; however, burning is ineffective on heavily grazed sites [69]. A 1965 study found that prescribed fire following chemical control of inkberry may enhance the recovery of other native forage species in areas formerly occupied by inkberry, though with fire exclusion, forage production may decline sharply [82].

MANAGEMENT CONSIDERATIONS

SPECIES: Ilex glabra
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Domestic livestock eat inkberry foliage sparingly [71]. Though not a preferred food of wildlife, inkberry is so abundant and produces so prolifically and consistently that it is likely an important emergency food source for many species [98].

The fruit, leaves, and twigs of inkberry are an important component of white-tailed deer [71,98,160,177,205] and Sambar deer [177] diets; inkberry is moderately preferred browse for white-tailed deer [39]. It is browsed by white-tailed deer primarily in  the fall and winter [71,177,205], and by Sambar deer in both the fall and spring [177].

Inkberry fruit may be important in the diets of small mammals [71,111], and it is an important component of the raccoon winter diet in Alabama [98]. The fruit and leaves provide food for black bears in the fall [44,71,73,75].

Inkberry fruit and leaves are a minor component of wild turkey [52,71,98,160], bobwhite quail [71,98,140,160], and songbird diets [10,71,98].

PALATABILITY:
Inkberry, with its bitter fruit [98], is generally described as unpalatable [70,98]. Inkberry has been specifically identified as unpalatable for cattle [94,178].

NUTRITIONAL VALUE:
The nutrient content of inkberry on burned and unburned plots on the southeastern coastal plain has been recorded as follows [116]:

  % moisture Chemical composition (% of dry weight)
Air dried Oven dried N (total) Cellulose Lignin Ether extract N-free extract Ca P Other Total
Early leaf stage
   Burned 63.9 64.8 1.62 13.25 19.87 5.16 70.09 0.58 0.16 2.30 3.03
   Unburned 61.6 62.4 1.64 11.58 21.54 6.59 71.02 0.55 0.17 2.12 2.64
Full leaf stage
   Burned 61.49 66.93 1.16 15.90 13.74 5.32 68.51 0.40 0.12 2.09 2.61
   Unburned 60.54 66.15 1.17 10.75 15.75 6.74 71.14 0.32 0.13 2.03 2.48

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 [90]:

  % of dry weight
  Total ash N P K Ca Mg S
Live foliage 2.1 + 0.16 1.0  + 0.03 0.05 + 0.005 0.29 + 0.02 0.61 + 0.03 0.18 + 0.008 0.07 + 0.003
Live stems <0.6 cm 3.0  + 0.30 0.6  + 0.02 0.06 + 0.007 0.20 + 0.02 0.51 + 0.03 0.13 + 0.012 0.05 + 0.004
Live stems 0.6-2.5 cm 1.4  + 0.20 0.3  + 0.02 0.04 + 0.006 0.17 + 0.03 0.26 + 0.04 0.05 + 0.005 0.03 + 0.003
Live stems 2.5-7.6 cm 1.4  + 0.20 0.3   + 0.01 0.04 + 0.005 0.17 + 0.02 0.25 + 0.04 0.05 + 0.004 0.03 + 0.003
Dead stems <0.6 cm 1.8  + 0.24 0.4  + 0.02 0.04 + 0.007 0.14 + 0.02 0.40 + 0.04 0.07 + 0.006 0.05 + 0.003
Dead stems 0.6-2.5 cm n.d. 0.4  + 0.08 0.01 + n.d. 0.15 + 0.04 0.12 + 0.01 0.06 + 0.012 0.04 + 0.012

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].

Season Year plantation fertilized % of dry matter
IVOMD* Ca P N
 Summer 1980 (August)

 

None 27.8 0.511 0.042 1.08
1972 27.6 0.454 0.054 1.08
1976 30.0 0.435 0.053 1.04
 Spring 1981(April)

 

None 39.3 0.223 0.114 1.62
1972 34.9 0.213 0.146 1.86
1976 36.0 0.198 0.141 1.76
*In-vitro organic matter digestibility

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:
No information

VALUE FOR REHABILITATION OF DISTURBED SITES:
Inkberry is useful for wetland mitigation [47].

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 [47]. 

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 [47]. Good germination is achieved by planting cleaned, collected seeds in moist, fungus-free medium at 60 degrees Fahrenheit (15.5 oC) until seeds sprout [71,93]. 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 [93]. 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) [15].

OTHER MANAGEMENT CONSIDERATIONS:
In field conditions, inkberry is parasitized by hemiparasitic chaffseed (Schwalbea americana), a federally endangered species whose range has been limited to Florida, Georgia, Mississippi, North Carolina, South Carolina, and New Jersey. In laboratory tests, 94.7% of inkberry seedlings planted with chaffseed were parasitized [76].

Inkberry thrives under browsing pressure [66]. 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 [69]. 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 [70,71], reducing the grazing capacity of these areas for domestic livestock [71]. 

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 [27]; 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 [38]. 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].

Studies in 1961 and 1972 demonstrated that control of inkberry might be achieved with a combination of burning and herbicide treatments [29,119]. Up to 80% control of inkberry may be achieved with a variety of herbicide combinations, including imazapyr, picloram, and picloram+triclopyr applications [180]. Good control is achieved with isopropyl ester applications, and fair control is achieved with sodium trichloracetate applications [192]. Information on fire control can be found under "Fire Management Considerations" in the "Fire Effects" section of this summary.

Ilex glabra: References


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2. Abrahamson, Warren G. 1984. Species response to fire on the Florida Lake Wales Ridge. American Journal of Botany. 71(1): 35-43. [9608]

3. Abrahamson, Warren G.; Abrahamson, Christy R. 1996. Effects of fire on long-unburned Florida uplands. Journal of Vegetation Science. 7: 565-574. [28455]

4. Abrahamson, Warren G.; Hartnett, David C. 1990. Pine flatwoods and dry prairies. In: Myers, Ronald L.; Ewel, John J., eds. Ecosystems of Florida. Orlando, FL: University of Central Florida Press: 103-149. [17388]

5. Abrahamson, Warren G.; Johnson, Ann F.; Layne, James N.; Peroni, Paricia A. 1984. Vegetation of the Archbold Biological Station, Florida: an example of the Southern Lake Wales Ridge. Florida Scientist. 47(4): 209-250. [20272]

6. Adler, Gregory H. 1988. The role of habitat structure in organizing small mammal populations and communities. In: Szaro, Robert C.; Severson, Kieth E.; Patton, David R., technical coordinators. Management of amphibians, reptiles, and small mammals in North America: Proceedings of the symposium; 1988 July 19-21; Flagstaff, AZ. Gen. Tech. Rep. RM-166. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 289-299. [7118]

7. Agee, James K. 1973. Prescribed fire effects on physical and hydrologic properties of mixed-conifer forest floor and soil. Contribution Report No. 143. Davis, CA: University of California, Water Resources Center. 57 p. [12337]

8. Ash, A. N.; McDonald, C. B.; Kane, E. S.; Pories, C. A. 1983. Natural and modified pocosins: literature synthesis and management options. FWS/OBS-83/04. Washington, DC: U.S. Fish and Wildlife Service, Division of Biological Sciences. 156 p. [16178]

9. Austin, Daniel F.; Posin, Freda R.; Burch, James N. 1987. Scrub species patterns on the Atlantic Coastal Ridge, Florida. Journal of Coastal Research. 3(4): 491-498. [9340]

10. Beal, F. E. L. 1915. Food of the robins and bluebirds of the United States. Bulletin No. 171. Washington, DC: U.S. Department of Agriculture. 31 p. [24990]

11. Beran, Robert A.; Masters, Robert A.; Gaussoin, Roch E. 1998. Revegetation of leafy spurge-infested grasslands with native grass and legume mixtures. In: Christianson, Kathy, ed. Proceedings, Western Society of Weed Science; 1998 March 10-12; Waikoloa, HI. [Place of publication unknown]: Western Society of Weed Science; Volume 51: 131. Abstract. [26950]

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