dcsimg
Common Names
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American holly
dune holly
hummock holly
scrub holly
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ILEOPA_COMMON_NAMES
Cover Value
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Cavities in American holly provide nesting habitat for the endangered
red-cockaded woodpecker [28].
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Coladonato, Milo. 1991. 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/
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Description
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More info for the terms: dioecious, drupe, fruit, tree

American holly is a native evergreen tree that grows to 50 feet (16 m)
[12,17]. Its evergreen leaves are leathery, with sharp pointed tips and
spiny toothed margins. The branches are short and crooked, and the
crown is rounded or pyramidal. The greenish white flowers are
unisexual, dioecious, and borne on short-stalked, axillary cymose
clusters. The fruit is a round, bright red, orange, or occasionally
yellow, four-seeded drupe or pyrene. The bark is thin, gray, and often
warty [21,40]
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Coladonato, Milo. 1991. 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/
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ILEOPA_GENERAL_BOTANICAL_CHARACTERISTICS
Distribution
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From the maritime forests of Massachusetts, American holly is scattered
along the coast to Delaware. It grows inland to several Pennsylvania
counties and to extreme southeastern Ohio. It occurs abundantly
southward throughout the Coastal Plain, Piedmont, and Appalachians. Its
range extends south to mid-peninsular Florida and west to eastern Texas
and southern Missouri [23,46]. It is cultivated in Hawaii [49]
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Coladonato, Milo. 1991. 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/
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ILEOPA_GENERAL_DISTRIBUTION
Fire Ecology
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More info for the terms: fire regime, root crown

American holly is susceptible to aboveground fire damage. It may persist by
sprouting from the root crown. [1,7,14].

FIRE REGIMES :
Find fire regime information for the plant communities in which this
species may occur by entering the species name in the FEIS home page under
"Find FIRE REGIMES".
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Coladonato, Milo. 1991. 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/
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ILEOPA_FIRE_ECOLOGY_OR_ADAPTATIONS
Fire Management Considerations
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Fire is a very effective agent for controlling American holly.
Seedlings and sprouts can usually be eliminated as a result of normal
underburning regimes in most commercial pine stands [48].
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Coladonato, Milo. 1991. 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/
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ID
ILEOPA_FIRE_MANAGEMENT_CONSIDERATIONS
Growth Form (according to Raunkiær Life-form classification)
provided by Fire Effects Information System Plants
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Coladonato, Milo. 1991. 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/
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ILEOPA_RAUNKIAER_LIFE_FORM
Habitat characteristics
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American holly is primarily a plant of the humid Southeast. It occupies
a wide variety of soils, from nearly sterile Inceptisols of the Atlantic
sandy beaches to fertile but thin mountain Ultisols to an elevation of
approximately 3,000 feet (915 m) [12,23]. The largest trees are found
in the rich bottomlands and swamps of the Coastal Plain. Growth is best
in moist, slightly acidic, well-drained sites such as upland pine sites
and hammocks [8,11]. Trees will not survive flooding or saturated soils
for more than 17 percent of the growing season. In the northeastern
portion of its range, holly is found on sandy soils on the Coast and on
dry gravely soils farther inland [2,12].

American holly is a common understory component in the longleaf pine
(Pinus palustris)-slash pine (P. elliotti) forests of the Coastal Plain.
Other common associates include sweetgum (Liquidambar styraciflua),
flowering dogwood (Cornus florida), American beech (Fagus grandifolia),
red maple (Acer rubrum), white oak (Quercus alba), water oak (Q.
nigra), hickory (Carya spp.), white ash (Fraxinus americana), yellow
poplar (Liriodendron tulipifera), black tupelo (Nysaa sylvatica),
southern red oak (Quercus falcata), and post oak (Q. stellata) [23].
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Coladonato, Milo. 1991. 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/
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ILEOPA_SITE_CHARACTERISTICS
Habitat: Cover Types
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More info on this topic.

This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

More info for the terms: hardwood, swamp

21 Eastern white pine
40 Post oak - blackjack oak
43 Bear oak
44 Chestnut oak
45 Pitch pine
46 Eastern redcedar
51 White pine - chestnut oak
52 White oak - black oak - northern red oak
53 White oak
57 Yellow poplar
58 Yellow poplar - eastern hemlock
59 Yellow poplar - white oak - northern red oak
61 River birch - sycamore
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 oak
89 Live oak
91 Swamp chestnut oak - cherrybark oak
92 Sweetgum - willow oak
96 Overcup oak - water hickory
97 Atlantic white cedar
98 Pond pine
100 Pondcypress
101 Baldcypress
103 Water tupelo - swamp tupelo
104 Sweetbay - swamp tupelo - red bay
110 Black oak
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Coladonato, Milo. 1991. 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/
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ID
ILEOPA_SAF_COVER_TYPES
Habitat: Ecosystem
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This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

FRES10 White - red - jack pine
FRES12 Longleaf - slash pine
FRES13 Loblolly - shortleaf pine
FRES14 Oak - pine
FRES15 Oak - hickory
FRES16 Oak - gum - cypress
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Coladonato, Milo. 1991. 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/
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ID
ILEOPA_ECOSYSTEMS
Habitat: Plant Associations
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More info on this topic.

This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the term: forest

K089 Black Belt
K090 Live oak - sea oats
K095 Great Lakes pine forest
K100 Oak - hickory forest
K104 Appalachian oak forest
K110 Northeastern oak - pine forest
K111 Oak - hickory - pine forest
K112 Southern mixed forest
K113 Southern floodplain forest
K114 Pocosin
K115 Sand pine - scrub
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Coladonato, Milo. 1991. 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/
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ILEOPA_KUCHLER_PLANT_ASSOCIATIONS
Immediate Effect of Fire
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Fire easily top-kills American holly [37]. Its thin bark is easily
injured by fire [24,26]. The cambium layer is destroyed and the leaves
and crown defoliated. Even large trees may be killed by light fires in
the understory [27,48].
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Coladonato, Milo. 1991. 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/
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ILEOPA_IMMEDIATE_FIRE_EFFECT_ON_PLANT
Importance to Livestock and Wildlife
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More info for the term: fruit

Birds are the principal consumers of American holly fruit, although
deer, squirrels, and other small animals eat them. At least 18 species
of birds, including songbirds, mourning doves, wild turkeys, and
northern bobwhite, are known to eat the fruit [42,46]. Cattle and deer
sometimes browse the foliage [9,18].
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Coladonato, Milo. 1991. 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/
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ID
ILEOPA_IMPORTANCE_TO_LIVESTOCK_AND_WILDLIFE
Life Form
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More info for the terms: shrub, tree

Tree, Shrub
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Coladonato, Milo. 1991. 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/
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ILEOPA_LIFE_FORM
Management considerations
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More info for the term: shrub

American holly is considered an undesirable shrub that competes with
pines and desirable hardwoods for light, moisture, and nutrients.
Streamline basal application of the herbicide Garlon 4 is an effective
means of controlling American holly [38].

The greatest damage to American holly trees is the indiscriminate
harvesting of foliage with berries for Christmas decorating. Before
laws were passed in Maryland and Delaware to protect the holly, there
was a "roadside" market for holly collected from trees that did not
belong to the harvesters. Trees were left mutilated and many died [23].
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Coladonato, Milo. 1991. 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/
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ILEOPA_OTHER_MANAGEMENT_CONSIDERATIONS
Nutritional Value
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More info for the term: forest

Lay [32] listed nutrient percentage values for American holly browse
collected in winter and summer on a pine-hardwood forest in Newton
County, Texas:

N-Free Phosphoric
Protein Fat Fiber extract Ash acid Calcium
Su 5.50 --- --- --- --- 0.14 ---
W 6.73 3.16 26.10 46.17 2.84 0.14 0.70

These levels are low for protein and deficient for phosphoric acid, but
high for calcium [32].
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Coladonato, Milo. 1991. 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/
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ILEOPA_NUTRITIONAL_VALUE
Occurrence in North America
provided by Fire Effects Information System Plants
AL AR CT DE FL GA HI KY LA MA
MD MS MO NC OH OK PA RI SC TN
TX VA WV
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ILEOPA_STATES
Other uses and values
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More info for the terms: forest, tree

The attractiveness of American holly's foliage is its principal value,
whether as a forest tree, planted ornamental, or Christmas decoration.
It can be used for yard, street, park planting, or for hedges [6,47].
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Coladonato, Milo. 1991. 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/
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Palatability
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The palatability of American holly to white-tailed deer and cattle is
considered poor. Deer and cattle generally consume American holly only
when more preferred browse is unavailable [33].
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Coladonato, Milo. 1991. 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/
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ILEOPA_PALATABILITY
Phenology
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More info on this topic.

More info for the terms: fruit, tree

American holly begins flowering in April in the southern parts of its
range and in June at its northern limits. The fruit ripen from
September through December and remain on the tree through most of the
winter [46,47].
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Coladonato, Milo. 1991. 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/
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ILEOPA_SEASONAL_DEVELOPMENT
Plant Response to Fire
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More info for the term: forest

American holly sprouts from surviving basal buds following fire [31].
Initial growth after fire is slow, averaging about 6 feet (1.8 m) in 16
years under medium shade [23]. Three annual fires in a southern pine
forest reduced the number of fruit-producing holly trees by 95 percent
[27].
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Coladonato, Milo. 1991. 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/
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ILEOPA_PLANT_RESPONSE_TO_FIRE
Post-fire Regeneration
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More info for the terms: caudex, root crown, seed

survivor species; on-site surviving root crown or caudex
off-site colonizer; seed carried by animals or water; postfire yr 1&2
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Coladonato, Milo. 1991. 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/
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ILEOPA_POSTFIRE_REGENERATION_STRATEGY
Regeneration Processes
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More info for the terms: fruit, seed, selection

Sexual: American holly is pollinated by insects, including bees, ants,
wasps, and night-flying moths [23]. Seed germination is slow, requiring
16 months to 3 years in nature [46]. Seeds are mainly dispersed by
birds and small mammals [10,13]. Seed production may be low in years of
heavy spring rain, as rain can diminish the wide dissemination of
pollen. A frost can kill the spring flowers, eliminating the fruit
crop. Frequent prescribed burning will also reduce fruit production
[20].

Vegetative: American holly sprouts from basal dormant buds [23,44].

Propagation: Transplanting of young holly trees should be done during
the dormant season, usually November through March. Small plants may be
dug bare-rooted if roots are kept moist, but larger plants should be
balled and burlapped. When wild hollies are transplanted from the
woods, tops should be severely pruned and most of the remaining leaves
removed. Small trees should be allowed to flower before transplanting
to ensure the selection of fruit-bearing individuals. Root pruning to a
depth of 2 to 3 feet (0.6-0.9 m) a year before lifting improves
transplanting success. Holly can be produced from cuttings taken in
August or September and December. Cuttings should be taken from the
current season's ripened wood, with a small section of 2-year-old wood
including several leaves. Cuttings should be set slanting in about 6
inches (15 cm) of moist peatmoss, with the leaves lying flat on the
surface [23].
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Coladonato, Milo. 1991. 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/
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ILEOPA_REGENERATION_PROCESSES
Successional Status
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More info on this topic.

More info for the terms: competition, forest, natural

American holly is seldom dominant because of its slow growth and
relatively short stature [3,16]. It is very shade tolerant and can
survive in the understory of most forest canopies [20]. American
holly's slow growth allows faster growing species to overtop it. Shade
and root competition in natural stands reduces the average height of
hollies compared with those growing in full sunlight. Crown area is
reduced by more than one-third under medium shade and by more than
one-half under heavy shade [23,30].
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Coladonato, Milo. 1991. 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/
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ILEOPA_SUCCESSIONAL_STATUS
Taxonomy
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The currently accepted scientific name of American holly is Ilex opaca
Ait. The Ilex genus consists of 13 species belonging to the holly
family (Aquifoliaceae). There are no forms or subspecies of American
holly. Recognized varieties include [21,34]:

I. opaca var. opaca American holly
I. opaca var. arenicola (Ashe) Ashe scrub or hummock holly

I. opaca hybridizes with I. cassine to produce I. x attenata Ashe [47].
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Coladonato, Milo. 1991. 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/
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ILEOPA_TAXONOMY
Value for rehabilitation of disturbed sites
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More info for the term: forest

American holly is useful for rehabilitating areas that have been damaged
by salt spray. It is more resistant to damage from salt spray than any
associated woody species in the maritime forest of New England [23,25].
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Coladonato, Milo. 1991. 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/
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ILEOPA_VALUE_FOR_REHABILITATION_OF_DISTURBED_SITES
Wood Products Value
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The wood of American holly is heavy, tough, and close-grained. It
shrinks considerably, checks or warps badly unless properly seasoned,
and is not durable under exposure [47]. The wood is used for veneer and
to a limited extent as pulpwood and lumber. The greates use of the wood
is for cabinets, interior finish, novelties, handles, fixtures, and
scientific instruments. When dyed black to resemble ebony, it is used
for piano keys, violin pegs, and fingerboards [23].
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Coladonato, Milo. 1991. 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/
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ILEOPA_WOOD_PRODUCTS_VALUE
Associated Forest Cover
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In longleaf pine-slash pine (Pinus palustris-P. elliottii) forests of the coastal plain, frequent prescribed fires are more limiting to the presence of American holly than site. Within that timber type, therefore, association is mainly with trees of bottom lands, swamps, or other sites not subject to burning. In the loblolly pine-shortleaf pine and upland hardwood types where fire is not so common, holly, as well as numerous other hardwoods, is found beneath the pines on a wide range of sites. Because of its slow growth and relatively short stature, holly is seldom dominant. It is an understory component of a number of forest cover types (10). Common associates of holly in various parts of its geographic range are shown in table 1.



Table 1- Trees commonly associated with American holly on various sites throughout its range Tree species Range¹ Sweetgum (Liquidamber styraciflua) A C D F G I Flowering dogwood (Cornus florida) A B D G I American beech (Fagus grandifolia) A B E G I Red maple (Acer rubrum)
A F G H White oak (Quercus alba) B C F G Water oak (Quercus nigra) A C G I Hickory (Carya spp.) C D G White ash (Fraxinus americana) C D F Yellow-poplar (Liriodendrom tulipifera) D F G Black tupelo (Nyssa sylvatica) A D G Soutern red oak (Quercus falcata) A F G Post oak (Quercus stellata) A C H Key to range symbols:
A. East Texas (15,25).
B. Southeast Louisiana (7).
C. Mississippi River Delta (24).
D. Georgia Piedmont (24).
E. Kentucky; western hills (24).
F. Southeast Pennsylvania (24).
G. Virginia Coastal Plain (35).
H. New Jersey and New York; maritime forests (28)
I. North-central Florida (29).

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Silvics of North America
ID
Ilex_opaca_silvics_Associated_Forest_Cover
Climate
provided by Silvics of North America
Like the southern pines, American holly is primarily a plant of the humid Southeast. Annual precipitation throughout its range is 1020 to 1650 mm (40 to 65 in) with over 2030 mm (80 in) in the southern Appalachians. Growing season varies from about 150 days in the Appalachian Mountains, the northeastern limit of the range, to almost yearlong in the central Florida peninsula. Average minimum temperature in the coastal plain portion of its range is above -18° C (0° F) but in the mountains of West Virginia, holly grows where the average low temperature is below -23° C (-10° F) (20). Holly cultivars in a northern Ohio arboretum, north of its natural range, have survived -29° C (-20° F) (12). American holly is the hardiest known broadleaf evergreen tree (23).

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Damaging Agents
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The greatest damage to holly trees is indiscriminate harvesting of foliage with berries for Christmas decorating. Before laws were passed in Maryland and Delaware to protect the holly, there was a "roadside" market for holly vandalized from trees that did not belong to harvesters. Trees were left mutilated and many died (17).

Fire is another deadly enemy of American holly. Most commercial pine timberland is burned often enough to eliminate holly seedlings or sprouts, especially where livestock graze. Burning where hollies are in the midstory can seriously damage the bark and kill trees. Three annual fires in a southern pine forest reduced the number of fruit-producing holly trees by 95 percent (19).

The thick evergreen leaves, which remain on the trees until the spring of their third year (18), are year-long hosts to many foliage diseases and insects. Few threaten the health of the trees, but many may reduce the esthetic and commercial value of the foliage. Diseases include 14 species of leaf spot fungi, six species of black mildews, two powdery mildews, and one rust. The most common and widespread of the leafspots are caused by the fungi Cercospora pulvinula, Phacidium curtissii, Phyllosticta opaca, and Physalospora ilicis. The rust Chrysomyxa ilicina is known only from the southern Appalachian area. Hollies of the northeast are subject to a serious leaf and twig blight caused by Corynebacterium ilicis (16).

Although nearly 30 species of insects are known to attack holly, only a few are serious pests. The southern red mite (Oligonychus ilicis) causes a reduction in leaf and twig growth and undesirable foliage color. The native holly leafminer (Phytomyza ilicicola) can damage foliage severely, causing leaves to drop prematurely. The holly midge (Asphondylia ilicicola) feeds on the berries causing them to remain green in color. Several species of scale insects feed on holly, including the holly scale (Asterolecanium puteanum) (1,24).

Strong winds cause spines of mature leaves to puncture other leaves, rendering the foliage less valuable for decoration (24). In northern portions of its range, twigs and branches can be killed during extreme cold periods (12), although holly is quite hardy (23).

Holly is more resistant to damage from salt spray than associated woody species in the maritime forests of New England, enabling it to dominate coastal stands (29). Hollies are intolerant of flooding and may die if their roots are inundated for a period of several weeks (31).

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Silvics of North America
ID
Ilex_opaca_silvics_Damaging_Agents
Flowering and Fruiting
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Hollies are dioecious; male (staminate) and female (pistillate) flowers, similar in appearance, with four to six small white petals, are produced on separate plants on the current season's growth. The male-to-female ratio for 1,930 seedlings from 10 randomly chosen, open-pollinated pistillate trees was 1.03 to 1.00 after 9 years had elapsed and all seedlings had produced flowers. Flowering began as early as age 3 and the latest bloomed at age 9, staminate plants flowering somewhat earlier on the average than pistillate. For this reason, the male-to-female ratio at age 5 was about 5 to 1 (5).

Flowering begins in April in the southern part of the range of American holly, and in June at the northern limits. Pollination is accomplished by insects, including bees, wasps, ants, yellowjackets, and night-flying moths (3). Staminate trees should be planted close to fruit-producing trees (34). Although some female hollies are apparently isolated by distance from pollen-bearing trees, good fruit crops are produced regularly. The fruit, commonly called "berries," are technically four-seeded drupes or pyrenes. They ripen from September through December and remain on the tree through most of the winter unless consumed by birds or other wildlife. The fruit is round to ellipsoid, about 6 to 12 mm. (0.25 to 0.50 in) in diameter, and bright-but not shiny-red, orange, or occasionally yellow in color.

The four seeds in each fruit are bony with a coarsely reticulated, ridged surface (34). Seed germination is very slow, requiring from 16 months to 3 years in nature. Germination tests over a 2.5-year period indicated only 33 to 56 percent germination capacity. Overwinter storage or cold, moist stratification improves germination. Sowing immediately after collection has been recommended although complete germination does not occur until the second or third spring (4).

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Genetics
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Population Differences American holly has 36 chromosomes, differing from the basic number for the genus Rex of 9 or 10 (24). Although leaf spininess and fruit colors vary, the coastal-dune hollies are usually smaller than those on the rich bottom lands of the Mississippi River Delta. Only one botanical variety other than the typical variety is recognized (21). Dune holly (Ilex opaca var. arenicola) grows on deep sandy soils in north and central Florida. It has lanceolate or oblanceolate leaves with forward-pointing teeth and oval, shallowly grooved nutlets. Yellow-fruited holly, once named a variety, is now considered only the expression of the recessive yellow color gene present in nearly all red-fruited hollies. Spineless leaves were once the basis for segregating another variety but the trait is highly variable; spiny and spineless leaves often grow on the same plant (34).

More than 1,000 cultivars of American holly have been named, although not all have been registered with the International Registration Authority. These do not necessarily represent different forms of Ilex opaca; many were selected because of unusual growth habit, fruit color, size or shape, or degree of leaf spininess (9).

Hybrids Topel holly (I. x attenuata Ashe) is a hybrid of I. opaca and dahoon holly (I. cassine), with long spiny-pointed leaves, that grows in South Carolina and northwestern Florida (21,34). Several cultivars registered under flex opaca, such as Foster, Hume, Savannah, and East Palatka, are actually I. x attenuata (9). Crosses have occurred between American holly and myrtle dahoon (I. myrtifolia), which, like dahoon holly, is an evergreen, red-fruited shrub or small tree found on wet sites of the coastal plain (24).

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Growth and Yield
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Because of its intolerance of fire, holly is found as scattered trees, even on good holly sites. Its slow growth and limby habit detract from its timber value. American holly under intensive culture is capable of 0.9 to 1.2 m (3 to 4 ft) of height growth per year (3).

Holly dominates some of the maritime forests of the Atlantic coast near the northern limit of its range, associated with salt-intolerant species such as black cherry (Prunus serotina), eastern redcedar (Juniperus virginiana), and hackberry (Celtis occidentalis). One of the best developed coastal stands reported was at Sandy Hook in New Jersey, where 97 percent of the tree basal area of a 30-ha (74-acre) forest was American holly. The oldest holly was 144 years old and 62 cm (24 in) in d.b.h. Height of holly trees in these sandy coastal forests ranges from 4.6 to 9.1 m (15 to 30 ft). Older trees or those on better sites may reach 15.2 m (50 ft) (28).

The "national champion" American holly, in the Congaree Swamp of South Carolina, is 30.2 m (99 ft) tall, with a circumference of 248 cm (98 in), a trunk diameter of 79 cm (31 in), and a crown diameter of 12.2 m (40 ft) (2). Hollies 30 to 90 cm (24 to 36 in) in diameter measured near the ground are common in the Mississippi River Delta (24). Trees 30.5 m (100 ft) tall and 122.0 cm (48 in) in d.b.h. have been recorded (18), but such trees were over 100 years old.

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Reaction to Competition
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Holly is classed as very shade tolerant and can survive in the understory of most forest canopies, but growth may be slowed and flowering and fruit set reduced under shade (22). Leaf area increased and leaves were greener under shade (30). In a mesic pine-hardwood forest of east Texas, dominated by loblolly pine (Pinus taeda), southern magnolia (Magnolia grandiflora), American beech (Fagus grandifolia), white oak (Quercus alba), and water oak (Q. nigra), holly was the principal understory species (15). Its slow growth allows faster growing species of the same age to overtop it. Shade and root competition in natural stands reduced average height of hollies at age 16 by about 0.3 m (1 ft) in medium shade and 0.61 m (2 ft) in heavy shade, compared with those growing in full sunlight. Crown area was reduced by one-third under medium shade and by more than one-half under heavy shade (32).

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Rooting Habit
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No information available.

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Seed Production and Dissemination
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Seed production may be low in years of heavy spring rain, as rain can diminish the wide dissemination of pollen. A late frost can kill the spring flowers, eliminating a fruit crop. Frequent prescribed burning also drastically reduces fruit production. Such crop failures are rare and localized, however, and an abundance of seed can be found each year (19,24). Clean seeds average approximately 61,730/kg (28,000/lb) and about four units (by weight) of fruit yield one unit of clean seeds (4). Seeds are dispersed mainly by birds and small mammals.

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Seedling Development
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Germination is epigeal. Following germination, holly seedlings rapidly develop a taproot and numerous lateral roots. American holly is very shade tolerant and may become established from bird droppings in the understory of upland pine plantations or bottom land hardwoods. It is very susceptible to fire and may be completely lacking in regularly or even occasionally burned forests (35). Initial growth is slow, averaging about 1.8 in (6 ft) in 16 years under medium shade (32). The bark is easily injured by fire and even large trees may be killed by light fires in the understory. Fastest growth of American holly was probably achieved in a North Carolina holly plantation; after 9 years of cultural practices such as mowing, mulching, and fertilizing, 10-year-old hollies averaged about 6.7 in (22 ft) in height and 3.7 in (12 ft) in crown spread and produced abundant fruit (24).

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Soils and Topography
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Holly survives on a wide variety of soils from near sterile Inceptisols of Atlantic sandy beaches to fertile but thin mountain Ultisols to an elevation of approximately 915 m (3,000 ft) (8). Largest trees are found in the rich bottom lands and swamps of the coastal plain. Growth is best on moist, slightly acid, well-drained sites such as upland pine sites and hammocks. Trees will not survive flooding or saturated soils for more than 17 percent of the growing season (31). In the northeastern portion of its range, holly is found on sandy coastal soils or dry gravelly soils farther inland (16).

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Special Uses
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The attractiveness of its foliage is American holly's principal value, whether as a forest tree, planted ornamental, or Christmas decoration. The development of commercial holly orchards and the education of landowners in the value and harvesting of holly foliage have lessened the exploitation of wild hollies (13).

The wood of American holly is tough and hard but not strong. It is close-grained and moderately heavy, weighing about 640.7 kg/m³ (40 lb/ft³). Specific gravity is 0.61 (oven-dry) and about 0.50 green. It is one of the whitest woods known, with white sapwood and ivory-white heartwood. Growth rings are almost indistinct. The wood is used for veneer and to a limited extent as pulpwood and lumber. Greatest use of the wood is for specialty items such as fancy cabinet inlays, small pieces of furniture, brush backs, handles, novelties, wood engravings, scroll work, woodcuts and carvings, and measuring scales and rules for scientific instruments; when dyed black to resemble ebony, it is used for piano keys, violin pegs, and fingerboards (6,18,33).

Birds are the principal consumers of the fruit, although deer, squirrels, and other small mammals also eat them. Cattle sometimes browse the foliage. At least 18 species of birds including songbirds, mourning doves, wild turkeys, and the bobwhite are known to eat the fruit (14,34). Perhaps the most important in seed dispersal, however, are the large winter-migrating flocks of small birds such as the cedar waxwing and American goldfinch. The complete stripping of all berries from a 10.7 m (35 ft) tall holly in a few seconds by a flock of cedar waxwings has been observed.

Despite the presence of saponins in the leaves and berries, American holly is not considered poisonous to man or animals (36). Although not as well known as gallberry (Ilex glabra) as a honey plant, its nectar makes excellent honey (24).

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Vegetative Reproduction
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Young holly trees usually sprout if the tops are removed by cutting or burning. Because of a taproot and a profilic lateral root system, young hollies can be transplanted without much difficulty (6). Transplanting should be done during the dormant season, usually November through March. Small plants may be dug bare-rooted if roots are kept moist, but larger plants should be balled and burlapped (34). When wild hollies are transplanted from the woods, tops should be severely pruned and most of the remaining leaves removed (16). Small trees should be allowed to flower before transplanting to ensure the selection of fruit-bearing individuals. Root pruning to a depth of 0.6 to 0.9 m (2 to 3 ft), a year before lifting, improves transplanting success (27). In Ohio, outside the natural range of American holly, better outplanting success has been obtained with plants 60 to 120 em (24 to 48 in) tall than those 30 cm (12 in) or less, because of winterkill of the younger plants (11). Holly can be produced from cuttings taken in August or September and December. Cuttings should be taken from the current season's ripened wood, with a small section of 2-year-old wood including several leaves. Cuttings should be set slanting in about 15 cm (6 in) of peatmoss and soil moisture, with the leaves lying flat on the surface. Treating with indolebutyric acid (IBA) and growing under high humidity with bottom heat is also recommended (6,24). "Quick dips" in IBA at high concentrations (up to 20,000 p/m) are recommended for cultivars that are normally hard to propagate (27). Root cuttings are unsatisfactory.

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Distribution
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From the maritime forests of Massachusetts, holly is scattered along the coast to Delaware. It grows inland into several Pennsylvania counties and abundantly southward throughout the coastal plain, Piedmont, and Appalachian system. The range extends south to mid-peninsular Florida, west to eastern Texas and southeastern Missouri (20). It corresponds roughly to the combined ranges for loblolly and shortleaf pines.


-The native range of American holly.


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Brief Summary
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Aquifoliaceae -- Holly family

H. E. Grelen

When the Pilgrims landed the week before Christmas in 1620 on the coast of what is now Massachusetts, the evergreen, prickly leaves and red berries of American holly (Ilex opaca) reminded them of the English holly (Ilex aquifolium), a symbol of Christmas for centuries in England and Europe (13,26). Since then American holly, also called white holly or Christmas holly, has been one of the most valuable and popular trees in the Eastern United States for its foliage and berries, used for Christmas decorations, and for ornamental plantings.

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Ilex opaca
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 src= Wikimedia Commons has media related to Ilex opaca.

Ilex opaca, the American holly, is a species of holly, native to the eastern and south-central United States, from coastal Massachusetts south to central Florida, and west to southeastern Missouri and eastern Texas.[2][3]

Description

Ilex opaca is a medium-sized broadleaved evergreen tree growing to 10–20 m (33–66 ft) tall, exceptionally up to 30 m (98 ft) tall, with a trunk diameter typically up to 50 cm (20 in), exceptionally 120 cm (47 in). The bark is light gray, roughened by small warty lumps. The branchlets are stout, green at first and covered with rusty down, later smooth and brown. The winter buds are brown, short, obtuse or acute.

The leaves are alternate, 5–7.5 cm (2.0–3.0 in) long and 2–4 cm (0.79–1.57 in) wide, stiff, yellow green and dull matte to sub-shiny above (distinctly less glossy than the otherwise fairly similar European holly Ilex aquifolium[4]), often pale yellow beneath; the edges are curved into several sharp, spike-like points, and a wedge-shaped base and acute apex; the midrib is prominent and depressed, the primary veins conspicuous; the petiole is short, stout, grooved, thickened at base, with a pair of minute stipules. The leaves remain on the branches for two to three years, finally falling in the spring when pushed off by growing buds.[2][5][6]

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Ripe fruit

The flowers are greenish white, small, borne in late spring in short pedunculate cymes from the axils of young leaves or scattered along the base of young branches. The calyx is small, four-lobed, imbricate in the bud, acute, margins ciliate, persistent. The corolla is white, with four petal-like lobes united at the base, obtuse, spreading, hypogynous, imbricate in bud. The flower stem is hairy with a minute bract at base. Like all hollies, it is dioecious, with separate male and female plants; only female plants produce the characteristic red berries. One male can pollenize several females. Male flowers have four stamens, inserted on the base of the corolla, alternate with its lobes; filaments awl-shaped, exserted in the sterile, much shorter in the sterile flower; anthers attached at the back, oblong, introrse, two-celled, cells opening longitudinally. The pistil on female flowers has a superior ovary, four-celled, rudimentary in staminate flowers; style wanting, stigma sessile, four-lobed; ovules one or two in each cell.

The fruit is a small red drupe 6–12 mm diameter containing four seeds; it is often persistent into winter.[2][5][6]

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Branch full of ripe fruit
Subspecies and varieties[1]
  • Ilex opaca subsp. arenicola (Ashe) A.E. Murray
  • Ilex opaca var. laxiflora (Lam.) Nutt.
  • Ilex opaca subsp. opaca
  • Ilex opaca var. opaca

Ecology

Ilex opaca typically grows as an understory tree in moist forests of the east-central, southeastern, and south-central United States. It is found in sparse numbers in the northern part of its range from Cape Cod, Massachusetts, south to northern New Jersey (including southern Connecticut and southeastern New York). It is abundant further south on the Gulf and Atlantic lowlands. The branches are short and slender. The roots are thick and fleshy. It will grow in both dry and swampy soil, but grows slowly.[2][6] Ilex opaca var. arenicola, or scrub holly, is found as a shrub component in xeric scrub habitats of the Florida peninsula.

The flowers are pollinated by insects, including bees, wasps, ants, and night-flying moths. The berries are reputedly poisonous to humans, but are important survival food for birds, who will eat the berries after other food sources are exhausted. The tree also forms a thick canopy which offers protection for birds from predators and storms. Songbirds including thrushes, mockingbirds, catbirds, bluebirds and thrashers frequently feed on the berries.[2][6]

Cultivation and uses

The wood is very pale, tough, close-grained, takes a good polish, and is used for whip-handles, engraving blocks and also cabinet work. It can also be dyed and used as a substitute for ebony. It has a density of 0.58 to 0.64.[clarification needed] The sap is watery, and contains a bitter substance used as an herbal tonic.[2][6]

Leaves from the American holly can be used to make a tea-like beverage. American holly tea does not contain caffeine.[7][unreliable source?]

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Mature plants often display a pyramidal shape

Ornamental plant

Ilex opaca is often cultivated by plant nurseries for use as a broadleaf evergreen ornamental plant, planted as a shrub or slower growing ornamental tree. Over 1,000 cultivars have been selected, including plants selected for cold tolerance ('Cobalt', a male cultivar, is able to tolerate temperatures as low as −32 °C), growth form (e.g. dwarf forms such as 'Cardinal Hedge', a female plant growing to 1.2 m tall), and color and abundance of fruit (notable female cultivars including the large-berried 'Yule', and the yellow-berried 'Canary' and 'Morgan Gold').[4]

The holly in winter

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Not only is the holly associated with winter decoration, it serves as a source of food and shelter during inclement weather

Holly is a popular winter Christmas and holiday season decoration. In English poetry and English stories the holly is inseparably connected with the merry-making and greetings which gather around the Christmas time. The custom is followed in North America, and holly and mistletoe are widely used for decoration of homes and churches.

The European holly is smaller than the American holly, but closely resembles the American holly. The leaves of both species are similar in outline and toothed and bristled very much the same way, but the leaves are brighter in the American holly and larger. The American holly, called the evergreen or Christmas holly (Ilex opaca Aiton) was named the state tree of Delaware on 1 May 1939.[8]

Holly fruit (drupes) appear late in the season, and whether due to the need to ripen or being a food of last resort, often last until midwinter. Cedar Waxwings will strip the trees of fruit if they are not already bare during their northward migration.[9]

References

  1. ^ a b The Plant List, Ilex opaca Aiton
  2. ^ a b c d e f Grelen, H. E. (1990). "Ilex opaca". In Burns, Russell M.; Honkala, Barbara H. Hardwoods. Silvics of North America. Washington, D.C.: United States Forest Service (USFS), United States Department of Agriculture (USDA). 2 – via Southern Research Station (www.srs.fs.fed.us)..mw-parser-output cite.citation{font-style:inherit}.mw-parser-output q{quotes:"""""'"'"}.mw-parser-output code.cs1-code{color:inherit;background:inherit;border:inherit;padding:inherit}.mw-parser-output .cs1-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Lock-green.svg/9px-Lock-green.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-limited a,.mw-parser-output .cs1-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Lock-gray-alt-2.svg/9px-Lock-gray-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/Lock-red-alt-2.svg/9px-Lock-red-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration{color:#555}.mw-parser-output .cs1-subscription span,.mw-parser-output .cs1-registration span{border-bottom:1px dotted;cursor:help}.mw-parser-output .cs1-hidden-error{display:none;font-size:100%}.mw-parser-output .cs1-visible-error{font-size:100%}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration,.mw-parser-output .cs1-format{font-size:95%}.mw-parser-output .cs1-kern-left,.mw-parser-output .cs1-kern-wl-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right,.mw-parser-output .cs1-kern-wl-right{padding-right:0.2em}
  3. ^ "Ilex opaca". County-level distribution map from the North American Plant Atlas (NAPA). Biota of North America Program (BONAP). 2014.
  4. ^ a b Huxley, A., ed. (1992). New RHS Dictionary of Gardening. Macmillan ISBN 0-333-47494-5.
  5. ^ a b Oklahoma Biological Survey: Ilex opaca
  6. ^ a b c d e Keeler, H. L. (1900). Our Native Trees and How to Identify Them. New York: Charles Scriber's Sons.
  7. ^ "Hollies: Caffein & Antioxidants". Eat the Weeds.
  8. ^ "Delaware Code Title 29 § 305".
  9. ^ Gil Nelson (2010). The Trees of Florida: A Reference and Field Guide (2nd ed.). Pineapple Press. p. 98.
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Ilex opaca: Brief Summary
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Ilex opaca, the American holly, is a species of holly, native to the eastern and south-central United States, from coastal Massachusetts south to central Florida, and west to southeastern Missouri and eastern Texas.

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