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American Beech

Fagus grandifolia Ehrh.

Comments

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A. J. Rehder (1907) argued for recognition of a southern variety ( Fagus grandifolia var. caroliniana ) of this somewhat variable species. The northern populations in general are characterized by cupules with denser, longer prickles, somewhat narrower leaves with a greater proportion of cuneate leaf bases, and larger fruits that exceed the cupules slightly. Others (e.g., W. H. Camp 1951) have suggested the existence of three races within United States F . grandifolia , often referred to as gray beech, red beech, and white beech. I follow J. W. Hardin and G. P. Johnson (1985) and others in not recognizing subspecific or varietal variation within eastern United States F . grandifolia . Examination of material over the geographic range of the species suggests that this variation is broadly clinal and can only be demonstrated statistically, with much variation indicative of the other races within most populations. It seems a matter of taste as to whether such variation be recognized with infraspecific names. In addition, forms with more densely pubescent leaves occur locally in both the north and south; they have been formally recognized by some authors. Clearly, additional taxonomic work on variation in F . grandifolia is desirable; it is possible that other characters that could adequately delimit subgeneric/varietal taxa might yet be identified.

Fagus sylvatica Linnaeus, the European Beech, is cultivated in temperate eastern North America and to a lesser extent in western United States and Canada. Escapes are to be expected. Various cultivars are known, particularly purple-leaf, tricolor-leaf, and cut-leaf forms. When encountered, F . s ylvatica is easily distinguished from F . grandifolia by the crenate leaf margin (without distinct teeth) and the softer, less stout, less reflexed spines of the cupule of F . sylvatica .

Native Americans used various preparations from plants of Fagus sylvatica medicinally for worms, consumption, chancre, and heart trouble, to purify the blood, as a poultice for burns and scalds, and as a wash for poison ivy (D. E. Moerman 1986).

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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
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Flora of North America Vol. 3 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Description

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Trees , to 20(-30) m. Bark gray, smooth. Twigs glabrous at maturity, or with scattered, straight, silky, simple hairs, prominent ringlike bud scale scars at beginning of each year's growth. Buds narrowly fusiform, to 15-20 mm, apex acute, scales few, silky light brown or glabrous. Leaves: petiole 4-12 mm. Leaf blade ovate or narrowly ovate, rarely obovate, 60-120 × 25-75 mm, base cuneate or subacute, apex acuminate; surfaces abaxially with scattered straight silky hairs, these often concentrated on midrib, occasionally glabrous or much more villous. Fruits: cupule brown to reddish brown, 15-20(-25) mm, opening at maturity to reveal nuts; nut 15-20 × 10-18 mm wide, glabrous or puberulent, often hollow even when full-sized.
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copyright
Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 3 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
source
Flora of North America @ eFloras.org
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Flora of North America Editorial Committee
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Distribution

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N.B., N.S., Ont., P.E.I., Que.; Ark., Ala., Conn., Del., Fla., Ga., Ill., Ind., Ky., La., Maine, Md., Mass., Mich., Miss., Mo., N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., R.I., S.C., Tenn., Tex., Vt., Va., W.Va., Wis.
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cc-by-nc-sa-3.0
copyright
Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 3 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Flora of North America @ eFloras.org
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Flora of North America Editorial Committee
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Flowering/Fruiting

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Flowering spring (Apr-Jun).
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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 3 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Habitat

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Rich woods, deciduous forest and mixed broadleaf-conifer forest; 0-1000m.
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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 3 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Synonym

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Fagus grandifolia var. caroliniana (Loudon) Fernald & Rehder
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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 3 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Flora of North America @ eFloras.org
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Broad-scale Impacts of Fire

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Fire wounds may serve as entrance courts for a host of decaying fungi
[41].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Common Names

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American beech
beech
Carolina beech
gray beech
red beech
ridge beech
white beech
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Cover Value

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More info for the term: cover

American beech provides cover for the Carolina chickadee (Parus
carolinensis) and the black-capped chickadee (Parus atricapillus) [10].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Description

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More info for the terms: fruit, tree

Beech is a large, native, deciduous tree. It normally grows 65 to 80
feet (20-25 m) tall but can can grow up to 130 feet (40 m) and can live
to over 300 years old. The bark is blue gray. The leaves are yellow
green during the growing season. The branches are stout and horizontal,
or ascending, with interlocking leaves forming a dense crown. The root
system is shallow and spreading. The fruit is a bur, usually containing
two nuts [4,41].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Distribution

provided by Fire Effects Information System Plants
American beech is distributed from Cape Brenton Island, Nova Scotia west
to Maine, southern Quebec, southern Ontario, northern Michigan, and
eastern Wisconsin; south to southern Illinois, southeastern Missouri,
northwestern Arkansas, southeastern Oklahoma, and eastern Texas; east to
northern Florida; and northeast to southeastern South Carolina. An
isolated variety (var. mexicana) occurs in the mountains of northeastern
Mexico [41,22].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Fire Ecology

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More info for the terms: fire regime, forest, frequency

Thin bark renders American beech highly vulnerable to injury by fire. Postfire
colonization is through root suckering [39].

When fire is absent or of low frequency, beech frequently becomes a
dominant species in mixed deciduous forests [5]. The transition from an
open fire-dominated forest to a closed canopy deciduous forest favors
the beech-magnolia type in the southern portion of beech's range [11].

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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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Fire Management Considerations

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More info for the terms: litter, tree

Fire could create favorable conditions for beech production. Fire could
reduce the litter and humus layer, expose roots, or injure the parent
tree, creating conditions for the production of sucker shoots [39].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Growth Form (according to Raunkiær Life-form classification)

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More info on this topic.

More info for the term: phanerophyte

Phanerophyte
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Habitat characteristics

provided by Fire Effects Information System Plants
More info for the term: mesic

Beech is found at low elevations in the North and relatively high
elevations in the South. Local soil and climatic factors probably
determine whether beech grows at the higher elevations. In the
Adirondack Mountains, low temperatures and wind keep beech below 3,200
feet (975 m) in contrast to the Appalachian Mountains where on the
warmer slopes it grows at elevations up to 6,000 (1,830 m) feet. At
altitudes in the middle of its range, beech is more abundant on the
cooler, moister, northern slopes than on the southern slopes [41].

Beech is usually found within two principal soil groups: the gray
podzolic (Hapludalf) and the laterite (Acrothox) and is prevalent on
podzols. It is seldom found on limestone soils except in the western
edge of its range. Beech populations are higher on coarse textured, dry
to mesic soils in the northern part of its range [2,41].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

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):

20 White pine - northern red oak - maple
21 Eastern white pine
22 White pine - hemlock
23 Eastern hemlock
24 Hemlock - yellow birch
25 Sugar maple - beech - yellow birch
26 Sugar maple - basswood
27 Sugar maple
28 Black cherry - maple
30 Red spruce - yellow birch
31 Red spruce - sugar maple - beech
32 Red spruce
33 Red spruce - balsam fir
34 Red spruce - Fraser fir
35 Paper birch - red spruce - balsam fir
52 White oak - black oak - northern red oak
55 Northern red oak
57 Yellow poplar
58 Yellow poplar - eastern hemlock
59 Yellow poplar - white oak - northern red oak
60 Beech - sugar maple
108 Red maple
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

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
FRES11 Spruce - fir
FRES12 Longleaf - slash pine
FRES13 Loblolly - shortleaf pine
FRES14 Oak - pine
FRES15 Oak - hickory
FRES16 Oak - gum - cypress
FRES17 Elm - ash - cottonwood
FRES18 Maple - beech - birch
FRES19 Aspen - birch
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Coladonato, Milo. 1991. Fagus grandifolia. 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/

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 terms: bog, forest

K081 Oak savanna
K083 Cedar glades
K084 Cross timbers
K089 Black Belt
K090 Live oaks - sea oats
K093 Great Lakes - fir forest
K094 Conifer bog
K095 Great Lakes pine forest
K096 Northeastern spruce - fir forest
K097 Southeastern spruce - fir forest
K098 Northern floodplain forest
K099 Maple - basswood forest
K100 Oak - hickory forest
K101 Elm - ash forest
K102 Beech - maple forest
K103 Mixed mesophytic forest
K104 Appalacian oak forest
K106 Northern hardwoods
K107 Northern hardwoods - fir forest
K108 Northern hardwoods - spruce forest
K109 Transition between K104 and K106
K110 Northeastern oak - pine forest
K111 Oak - hickory - pine forest
K112 Southern mixed forest
K113 Southern floodplain forest
K115 Pocosin
K116 Subtropical pine forest.
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Immediate Effect of Fire

provided by Fire Effects Information System Plants
Fire usually top-kills American beech. Mortality of young trees is related
to fire severity: Cool fires kill 40 to 50 percent of the seedlings and
saplings [16,36].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Importance to Livestock and Wildlife

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More info for the term: mast

Beech mast is eaten by a variety of birds and mammals, including mice,
squirrels, chipmunks, black bear, deer, foxes, ruffed grouse, ducks, and
bluejays [41].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Key Plant Community Associations

provided by Fire Effects Information System Plants
More info for the terms: codominant, forest

American beech is either a dominant or codominant species in the
northern hardwoods of the Northeast, Lake States, and the Appalachian
Mountains. Common associates include sugar maple (Acer saccharum), red
maple (A. rubrum), yellow birch (Betula alleghaniensis), American
basswood (Tilia americana), black cherry (Prunus serotina), southern
magnolia (Magnolia grandiflora), eastern white pine (Pinus strobus), red
spruce (Picea rubens), hickories (Carya spp.), and oaks (Quercus spp.)
[3,41].

Published classification schemes listing American beech as dominant or
codominant in habitat types (hts) are listed below:

Area Classification Authority

n MI, ne WI forest hts Coffman, Alyanak &
Rosovosky 1980
n WI forest hts Kotar & others 1989
n WI, n MI forest hts Kotar 1986
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Life Form

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More info for the term: tree

Tree
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Management considerations

provided by Fire Effects Information System Plants
More info for the terms: competition, forest

Even-aged silviculture adversely affects beech production and favors
production of associated hardwoods. Beech seedlings may be overtopped
in clearcuts by less shade-tolerant species such as birches (Betula
spp.) and oaks, which respond vigorously to increased light. Repeated
clearcutting at short intervals may eliminate beech. Shelterwood cuts
allow beech to develop with little competition from more intolerant
species [40,12,30].

Beech is seriously affected by beech bark disease. The saddled is its
most serious defoliator, and the forest tent caterpillar, gypsy moth,
fall cankerworm ruce spanworm (Operophtera bruceata) occasionally cause
heavy damage [41].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Occurrence in North America

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AL AR CT DE FL GA IL IN KY LA
MA MD ME MI MN MO MS NC NH NJ
NY OH OK PA SC TN TX VT VA WI
WV NS ON PQ MEXICO
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Coladonato, Milo. 1991. Fagus grandifolia. 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/

Other uses and values

provided by Fire Effects Information System Plants
Beechnuts are roasted and eaten or used a coffee substitute. The leaves
and bark are used to make dyes [24].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Palatability

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More info for the term: frequency

Beech is regarded as a poor deer browse. The frequency of its use in
some areas is due to the low availability of more preferable browse [7].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Phenology

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Flowering occurs from March to May. Fruiting occurs from September to
October. Seeds are released in October or November after frost [33,42].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Plant Response to Fire

provided by Fire Effects Information System Plants
Beech trees that survive a fire regenerate by root suckering or stump
sprouting [29,41].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Post-fire Regeneration

provided by Fire Effects Information System Plants
More info for the terms: caudex, root crown, seed

survivor species; on-site surviving root crown or caudex
survivor species; on-site surviving rhizomes
off-site colonizer; seed carried by animals or water; postfire yr 1&2
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Regeneration Processes

provided by Fire Effects Information System Plants
More info for the terms: adventitious, litter, seed, tree

Sexual reproduction: Beech begin producing seed when 40 years old and
by 60 years old may produce large quantities. Beech produces seed at 2-
to 8-year intervals. Beech seeds average about 1,600 per pound
(3,500/kg) [41]. Most seeds drop to the ground. A few are carried by
rodents but dispersal is limited. Bluejays may transport seeds several
kilometers [9]. Most of the seeds will germinate in the 1st year; after
that, the seeds lose viability [41].

Beech seeds germinate from early spring to early summer. Chilling is
required to break dormancy. Germination is good on mineral soil or
leafy litter, but poor on excessively wet sites. Seedlings grow best
under a moderate canopy or in protected small openings where the soil
does not dry out below the depth of the shallow roots [28].

Vegetative Reproduction: Beech can regenerate by root suckers or by
stump sprouts [6]. Sprouts may develop on the trunk of a tree
immediately below a wound and from the top of stumps. Adventitious buds
develop in callus tissue of the cambial layers of stumps. Sprouts can
also develop from the exposure of the roots to air or elevated
temperatures. Sometimes root sprouts develop when no apparent injury
has occurred [14].

The advance of beech bark disease, with its resultant mortality of
overstory beech stems, is likely to result in an increase in root
suckering [23]. Beech is more likely to develop by sprouting than by
seedling establishment [19].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Successional Status

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More info for the terms: climax, forest

American beech is a climax species that grows slowly underneath an
overstory of conifers or hardwoods. Beech grows faster in canopy
openings and eventually ascends into the overstory [1,8]. In an
old-growth forest in New Hampshire, beech replaced yellow birch and
sugar maple and then was able to maintain itself via root suckering
[36].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Taxonomy

provided by Fire Effects Information System Plants
The currently accepted scientific name of American beech is Fagus
grandifolia (Ehrh.) Little [26]. Some authorities hold that the
southern beeches vary and describe the southern form as F. grandifolia
var. caroliniana (Loud) Fernald & Rehder [32]. The variety F.
grandifolia var. mexicana (Martinez) is found in Mexico [26].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Value for rehabilitation of disturbed sites

provided by Fire Effects Information System Plants
Beech grows relatively slowly and has a low tolerance to fire. Its
value as a colonizer is limited [6,21,39].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Wood Products Value

provided by Fire Effects Information System Plants
More info for the terms: density, fuel

Beech wood is used to make flooring, furniture, veneer plywood, and
railroad ties. It is especially favored as fuel wood because of its
high density and good burning qualities. Coal tar made from beech wood
is used to protect wood from rotting. The creosote made from beech wood
is used to treat various human and animal disorders [31,41].
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bibliographic citation
Coladonato, Milo. 1991. Fagus grandifolia. 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/

Associated Forest Cover

provided by Silvics of North America
Within its wide range in eastern North America, beech is associated with a large number of trees. Some of the principal associates are sugar maple, red maple, yellow birch (Betula alleghaniensis), American basswood (Tilia americana), black cherry (Prunus serotina), southern magnolia (Magnolia grandiflora), eastern white pine (Pinus strobus), red spruce (Picea rubra), and several hickories (Carya spp.) and oaks Quercus spp.). Beech is included in 20 forest cover types and is a major component in the following three (5): Sugar Maple-Beech-Yellow Birch (Society of American Foresters Type 25), Red Spruce-Sugar Maple-Beech (Type 31), and Beech-Sugar Maple (Type 60). Beech is a minor species in 17 other cover types.

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Climate

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Within the range of beech, annual precipitation usually is from 760 mm to 1270 mm (30 to 50 in) (39); however, some beech is found in Michigan where precipitation is about 580 mm (23 in), and in Canada where about 640 mm (25 in) fall annually. Precipitation during the growing season varies from 250 mm to 460 min (10 to 18 in). Beech is a mesophytic species; it uses twice as much water for transpiration and growth processes annually, compared to some drought resistant oaks and even lesser amounts by some pines.

The growing season for beech varies from 100 to 280 days; the species is found in one county in Michigan where the growing season is only 92 days.

Mean annual temperatures range from 4° to 21° C (40° to 70° F). Beech can exist under temperature extremes lower than -42° C (-44° F) and 38° C (100° F). Higher than average summer temperatures may be unfavorable for beech growth.

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Damaging Agents

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In regions with low winter temperatures, long frost cracks often appear in the tree trunks. These cracks are sometimes superficial but sometimes extend deep into the bole. In the Northeast, beech has been damaged or killed by temperatures of -40° to -45° C (-40° to -50° F) preceded by severe droughts (39). Injured trees died the following summer and winter. Beech can be severely damaged by late spring frosts.

In a Kentucky study of effects of flooding, beech was one of the more sensitive species. Beech trees were killed by 2 weeks of submergence of their root crowns in summer. An 18-day period of flooding in winter had no apparent adverse effect, however.

Beech's susceptibility to glaze-storm breakage is no greater than that of its associated hardwoods and may be somewhat less than the average for a mixed stand (39). Except on shallow soils, beech is rather windfirm.

The thin bark of beech renders it highly vulnerable to injury by fire (large shallow roots are especially vulnerable), sunscald, logging, pruning, or disease. When large branches are broken they heal comparatively slowly (38) and serve as entrance courts for a host of decay fungi (12,32).

More than 70 decay fungi (a record for a hardwood species) have been reported for beech (12). The most important include Daedalea unicolor, Ganoderma applanatum, Fomes fomentarius, Phellinus igniarius, Hericium erinaceus, H. coralloides, Steccherinum septentrionale, Inonotus glomeratus, and Ustilina vulgaris. The shoestring fungus, Armillaria sp., the most important root pathogen, attacks and girdles roots of weakened trees. Beech roots are also parasitized by the broomrapes, Conopholis americana and Epifagus virginiana. The latter, beech drops, is specific to beech (8,34).

The thinness of beech bark also makes it vulnerable to an unusually large number of sucking insects, including the beech blight aphid, Fagiphagus imbricator, and the giant bark aphid, Longistigma caryae. Continuous heavy outbreaks of the oystershell scale, Lepidosaphes u1mi, have resulted in severe crown dieback and even in the death of entire stands (1). Xylococculus betulae, another scale, causes roughened spots on stems of young trees and is especially devastating to the sprout thickets that have emerged in the aftermath of beech bark disease , the most serious problem of this species (13,31).

Beech bark disease is initiated when yet another scale insect, the beech scale, Cryptococcus fagisuga, attacks the bark of beech trees and renders it susceptible to bark canker fungi of the genus Nectria (3,33). The insect component of this scale-Nectria "complex" was introduced to Nova Scotia from Europe around 1890 and is now found throughout New England, New York (15) and northern Pennsylvania (37). In 1981, a 70,000-acre infestation was detected in northeastern West Virginia, many miles south of the nearest previously known infestation (28). More recently, the disease has been reported as far west as Toronto, Ontario, and the scale is now present in northeastern Ohio and northwestern Virginia (29). In North America, Nectria coccinea var. faginata is the fungus most commonly associated with the disease in the Maritime Provinces, New England, and northern New York. In western Pennsylvania, West Virginia, and some New York stands however, N. galligena is the predominant associated species. As the disease and forest interact for the first time, mortality may be so severe that a large proportion of the big, mature beech trees are killed. Mortality is now especially high in some southern and western areas of the Adirondack Mountain region. The percent stocking of beech was reported (7) to remain the same after the killing front of the beech bark disease moved through a managed stand; the disease mainly affected the larger trees. Although such mortality is rare in stands emerging in the aftermath of the disease, severe defect may be caused by the now-endemic causal complex together with Xylococculus betulae (13).

Defoliation by insects can occasionally be a serious problem (1). The most damaging is the saddled prominent, Heterocampa guttivitta, although the forest tent caterpillar (Malacosoma disstria), gypsy moth (Lymantria dispar), fall cankerworm. (Alsophila pometaria), and the Bruce spanworm. (Operophtera bruceata) occasionally cause heavy defoliation in local areas. Insect defoliation often renders trees susceptible to attack by the shoestring root fungus.

Beech is seldom severely browsed by white-tailed deer. When other, more desirable tree species are available, beech is usually nipped only sparingly (36).

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Flowering and Fruiting

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In the Northern and Central States, beech flowers appear in late April or early May when the leaves are about one-third grown; the species is monoecious. The flowers are quite vulnerable to spring frosts. Male flowers occur in long-stemmed heads; female flowers in clusters of two to four (40). Beechnuts require one growing season to mature and they ripen between September and November. Two or (rarely) three nuts may be found within a single bur. The first nuts to fall are usually wormy or aborted.

Seed fall begins after the first heavy frosts have caused the burs to open and is completed within a few weeks. Some empty burs remain on the trees throughout the winter.

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Genetics

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Fagus grandifolia Ehrh. is the only type species of American beech now recognized in North America (9,10,23). Some botanical authorities hold that Northern and Southern beeches vary, and have described the southern form as F grandifolia var. caroliniana (Loud.) Fern. & Rehd., Carolina beech (4,6). A previously named species in the mountains of Mexico (39) has been renamed a variety, F grandifolia var. mexicana (Martinez) Little (22).

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Growth and Yield

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Beech's period of radial growth may continue for 80 to 89 days in the Georgia Piedmont and for approximately 60 days in Indiana (39). Annual height growth of beech saplings is complete in about 60 days; 90 percent of this growth occurs between May 10 and June 10, American beech has a lower site index than any associated hardwood in the northern Lake States.

The radial growth period is influenced by available soil moisture. Under normal conditions, it may end in the middle of July, but drought may end it in mid-June. A few individual trees may continue their growth into August and September. In dry years, annual rings may not grow in the basal sections of some beech trees. In general, radial growth of beech begins when the leaves are fully expanded.

The annual diameter increment of beech of pole and small saw-log size averages from around 1.8 to 2.3 mm (0.07 to 0.09 in) in undisturbed second-growth stands to 3.8 to 4.8 min (0.15 to 0.19 in) in trees released by partial cuttings (35,39). Annual growth of poles for 5 years after heavy release, leaving from 1.1 to 4.6 m/ha (5 to 20 W/acre) of basal area, ranged from 5.6 mm (0.22 in) to 7.6 min (0.30 in); growth was better in the most heavily stocked stands and on trees with good crown development (26).

Under optimum conditions, beech trees may become 37 in (120 ft) high; however, they generally average 18 to 24 in (60 to 80 ft). Growth data for beech in the Lake States are shown in table 1.

Table 1- Characteristics of American beech growing in the lake states. Age D.b.h. Height Volume (yr) (cm) (m) (m³) 20 2 4 -- 40 6 8.5 -- 60 10 11.9 0.03 80 14 14.6 0.1 100 18 17.4 0.22 150 29 22.9 0.76 200 40 25.6 1.58 250 51 26.8 2.69 (yr) (in) (ft) (ft³) 20 0.7 13  -- 40 2.3 28 -- 60 3.8 39 1 80 5.4 48 3.7 100 7.1 57 7.9 150 11.5 75 27 200 15.7 84 56 250 19.9 88 95 Among 12 broad-leaved species rated according to their longevity, beech was exceeded only by white oak and sugar maple. Beech trees older than 366 years have been found in Pennsylvania. The distribution of numbers of trees by age is "J" shaped, typical of tolerant long-lived species (21). One of the largest beeches on record, growing in Michigan, is 135 cm (53.2 in) in d.b.h., 49 m (161 ft) tall, and has a crown 32 in (105 ft) wide.

Beech trees prune themselves in well-stocked stands. Open-grown trees, however, develop short, thick trunks with large, low, spreading limbs terminating in slender, somewhat drooping branches that form a broad, round-topped head.

Beech trees that have been injured or suddenly exposed by stand cuttings often develop epicormic branches. In one stand where 65 percent of the basal area had been cut, 40 percent of the remaining beech trees had epicormic branches 5 years later, whereas in a similar but uncut stand, only 17 percent of the trees had such branches (39). Epicormic branching of beech trees has also been observed after glaze damage and after low-temperature injury (27). One report on winter injury showed epicormic branches to be restricted largely to trees with d.b.h. of 10 cm (4 in) or less (2).

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Reaction to Competition

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Beech is classed as very tolerant of shade. In some parts of its range, beech is the most tolerant species. Its tolerance is partly due to its very low respiration rate (24) and the quick response of the stomata, which open when light suddenly increases and rapidly close when light intensity diminishes. Beech stomata are more responsive than those of red maple, red oak Quercus rubra), or yellow-poplar (Liriodendron tulipifera), which are less tolerant (43). On very poor soils or in very cold climates, beech may be less tolerant. The tolerances of beech and associated sugar maple are about the same (25), although locally one species or the other may predominate in the forest understory. Factors other than the ability to endure shade appear to govern the relative success of beech and its common tolerant competition, sugar maple, eastern hemlock (Tsuga canadensis), and balsam fir (Abies balsamea). Beech may be more competitive under somewhat adverse site and climate conditions (39).

Beech and sugar maple are recognized as climatic climax species in the northern hardwood types of the Northeast, Lake States, and Appalachian Mountains. In the Southeast, relict areas of beech suggest that an original maple-beech association has been displaced by the once subclimax oak-hickory community.

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Rooting Habit

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Young seedlings have a taproot that gives way to a heart root system as the tree matures (41). The root system is generally shallow but may penetrate to 1.5 m (5 ft) or more in deep soils. The fine roots form a dense mat in our soil types. Beech root systems are more shallow than the associated yellow birch and sugar maple. Few tree species are less tolerant of flooding during the growing season than American beech.

Root exudates of beech contain more organic acids than those of sugar maple or yellow birch.

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Seed Production and Dissemination

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Beech ordinarily begins to produce a substantial amount of seeds when about 40 years old, and by the time it is 60 years old may produce large quantities. Good beech seed crops are produced at 2- to 8-year intervals (40).

Beech seeds, averaging about 3,500/kg (1,600/lb), are relatively heavy. Most of the seeds simply drop to the ground under the parent trees. Rodents may carry some of them short distances and on steep terrain a few may roll down slopes, but dispersal is quite restricted. Bluejays may transport many beech seeds several kilometers (16,17).

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Seedling Development

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Beech seeds germinate from early spring to early summer. Germination is epigeous and chilling is required to break dormancy. On either mineral soil or leaf litter, germination is good, but on excessively wet sites it is poor. Both germination and survival tend to be better on mor humus than on mull humus soil (39,40).

Beech seedlings develop better under a moderate canopy or in protected small openings than they do on larger open areas where the surface soil may dry out below the depth of the shallow roots. Height growth of seedlings is about the same in dense (87 percent) or moderate (55 percent) shade, but total dry weight and root development are greatest under moderate shade. Height growth, dry weight, and root development in the open are less than in shade (25). Seedlings are found in large numbers beneath even the densest stands, but under such conditions their growth is slow. Beech reproduction can start under, and come through, fern and raspberry cover.

Dormancy of beech seedlings can be broken in spring and growth can be prolonged in fall by supplemental light. Decreasing day length plays the major role in inducing dormancy in the fall, but day length may be secondary to temperature in controlling resumption of growth in the spring. That is, day length probably becomes adequate for growth to resume in the spring before temperatures are high enough for growth to occur. Temperature, therefore, exerts the final control over growth resumption.

Beech continues growing all winter in a greenhouse when daylight is supplemented by continuous artificial light.

The height of beech seedlings growing in the intense competition of a virgin hemlock-hardwood stand in northern Pennsylvania (39) was as follows:



Age Total height (yr) (m) (ft) 6 0.3 1 10 0.6 2 14 0.9 3 17 1.2 4 18 1.4 4.5 20 1.5 5 22 1.8 6 25 2.1 7

When forest stands are heavily cut, beech reproduction tends to grow more slowly than that of most associated hardwood species. This is especially true in clearcuttings. Here the beech reproduction may be overtopped by less tolerant species, such as the birches and white ash (Fraxinus americana), that respond vigorously to increased light. A number of studies have shown that heavy cutting or clearcutting results in fewer beech in the new stand than in the old (39). Repeated clearcutting on short rotations may nearly eliminate beech. Under partial cuttings, especially single-tree selection cuttings, intolerant species offer little competition and the tolerant beech reproduction is able to develop. The beech may be further favored by its virtual immunity to deer browsing.

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Soils and Topography

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Beech is found generally within two principal soil groups: the gray-brown podzolic (Hapludalf) and the laterite (Acrorthox) and is prevalent on podzols; it is seldom found on limestone soils except at the western edge of its range. These soils are contained in the orders Alfisols, Oxisols, and Spodosols. Soils of loamy texture and those with a high humus content are more favorable than lighter soils (39). The largest trees are found in the alluvial bottom lands of the Ohio and the lower Mississippi River valleys, and along the western slopes of the southern Appalachian Mountains.

Beech populations frequently are higher on coarse-textured, dry-mesic soils in the northern part of its range (38). In Indiana, beech is more sensitive to reduced soil moisture than is white oak (Quercus alba), sugar maple (Acer saccharum), American elm (Ulmus americana), and slippery elm (U. rubra). It will grow on poorly drained sites not subjected to prolonged flooding and may grow where the water table is within 15 cm to 25 cm (6 to 10 in) of the surface. It is markedly less tolerant of such conditions than are red maple (Acer rubrum) and sweetgum (Liquidambar styraciflua). Beech trees on poorly drained sites have shallower root systems than those on better drained sites (39). 'Ember stands containing considerable numbers of beech are found on soils ranging from pH 4.1 to 6.0 (39), but seldom where pH exceeds 7.0.

Beech is found at low elevations in the North and relatively high elevations in the southern Appalachians. Local soil and climatic factors probably determine whether beech grows at the higher elevations. In the Adirondacks of New York, low temperatures and wind keep beech below 980 m (3,200 ft), in contrast to the southern mountains where on the warmer slopes it grows at elevations up to 1830 m (6,000 ft). At latitudes in the middle of its range, however, beech is more abundant on the cooler and moister northern slopes than on the southern slopes (39).

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Special Uses

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Beech mast is palatable to a large variety of birds and mammals, including mice, squirrels, chipmunks, black bear, deer, foxes, ruffed grouse, ducks, and bluejays. Beech is the only nut producer in the northern hardwood type. Beech wood is used for flooring, furniture, turned products and novelties, veneer, plywood, railroad ties, baskets, pulp, charcoal, and rough lumber. It is especially favored for fuelwood because of its high density and good burning qualities.

Creosote made from beech wood is used internally and externally as a medicine for various human and animal disorders. (It is important to note that coal tar creosote, the kind used to protect wood from rots, is highly toxic to humans.)

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Vegetative Reproduction

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Beech sprouts well from the stumps of young trees, but this ability diminishes after trees reach 10 cm (4 in) in d.b.h. Sprouts from stumps 25 cm to 38 cm (10 to 15 in) in diameter usually are short lived and do not attain tree stature. Numerous sprouts may develop on the trunk of beech immediately below a wound, and from the tops of stumps; here adventitious buds develop in callus tissue of the cambial region.

Beech trees may develop large numbers of root sprouts or suckers. Studies (30) have shown that reproduction is almost exclusively by suckering in the "beech gaps" and is abundant in the Adirondack Mountains of New York, in Maine (13), and in many other areas, often those near the northern and western limits of its range (11,42) where environments are severe (27). Suckering is stimulated only slightly by removal of the stem (18). Injury to roots appears to be necessary for the initiation of root sprouts in beech (19). Root sprouts arise from adventitious buds that form within callus tissues associated with wounds. Experimental injuries to roots in November resulted in fewer sprouts than did injuries inflicted in spring (20). Sometimes root sprouts develop where no apparent injury has occurred (39). There were relatively more root sprouts on southerly slopes in areas where freeze-thaw action tended to injure shallow or exposed roots and stimulate sprout formation, and where late spring frosts tended to injure or kill young seedlings. In Ohio, seedling regeneration was positively associated with northerly exposures and root sprout regeneration with southerly exposures (11).

In an undisturbed stand of mature beech in the Adirondacks, 1,730 to 2,220 root sprouts per hectare (700 to 900/acre; 7 to 12/tree) were counted (39). Casual observations elsewhere indicate that the number per tree may greatly exceed this figure.

Root sprouts can develop into desirable trees. Isozyme genetic studies have shown that some groups of overstory beech trees with similar phenotypic traits are clones (14). Sometimes root sprouts are ephemeral. In one reproduction study, made after a 60-year-old stand of beech was cut, all of the root sprouts died within 4 years. On the other hand, the trees in a 40-year-old beech stand of sprout origin averaged 10 cm (4 in) in d.b.h. and 11.6 in (38 ft) in height.

Beech limbs root in a single year when layered. Interspecific root grafting is common.

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Brief Summary

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Fagaceae -- Beech family

Carl H. Tubbs and David R. Houston

American beech (Fagus grandifolia) is the only species of this genus in North America. Although beech is now confined to the eastern United States (except for the Mexican population) it once extended as far west as California and probably flourished over most of North America before the glacial period (39). This slow-growing, common, deciduous tree reaches its greatest size in the alluvial soils of the Ohio and Mississippi River Valleys and may attain ages of 300 to 400 years. Beech wood is excellent for turning and steam bending. It wears well, is easily treated with preservatives, and is used for flooring, furniture, veneer, and containers. The distinctive triangular nuts are eaten by people and are an important food for wildlife.

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Distribution

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American beech is found within an area from Cape Breton Island, Nova Scotia west to Maine, southern Quebec, southern Ontario, northern Michigan, and eastern Wisconsin; then south to southern Illinois, southeastern Missouri, northwestern Arkansas, southeastern Oklahoma, and eastern Texas; east to northern Florida and northeast to southeastern South Carolina. A variety exists in the mountains of northeastern Mexico.


-The native range of American beech.


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Fagus grandifolia

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Fagus grandifolia, the American beech or North American beech, is a species of beech tree native to the eastern United States and extreme southeast of Canada.

Description

Fagus grandifolia is a large deciduous tree[2] growing to 16–35 metres (52–115 feet) tall,[3] with smooth, silver-gray bark. The leaves are dark green, simple and sparsely-toothed with small teeth that terminate each vein, 6–12 centimetres (2+144+34 inches) long (rarely 15 cm or 6 in), with a short petiole. The winter twigs are distinctive among North American trees, being long and slender (15–20 millimetres or 5834 inch by 2–3 mm or 33218 in) with two rows of overlapping scales on the buds. Beech buds are distinctly thin and long, resembling cigars; this characteristic makes beech trees relatively easy to identify. The tree is monoecious, with flowers of both sexes on the same tree. The fruit is a small, sharply-angled nut, borne in pairs in a soft-spined, four-lobed husk. It has two means of reproduction: one is through the usual dispersal of seedlings, and the other is through root sprouts, which grow into new trees.[4]

Taxonomy

Trees in the southern half of the range are sometimes distinguished as a variety, F. grandifolia var. caroliniana, but this is not considered distinct in the Flora of North America. The Mexican beech (F. grandifolia var. mexicana), native to the mountains of central Mexico, is closely related, and is sometimes treated as a subspecies of American beech, but some botanists classify it as a distinct species. The only Fagus species found in the Western Hemisphere (assuming F. mexicana is treated as a subspecies), F. grandifolia is believed to have spanned the width of the North American continent all the way to the Pacific coast before the last ice age.

Two subspecies are generally recognized:[5]

Etymology

The genus name Fagus is Latin for "beech", and the specific epithet grandifolia comes from grandis "large" and folium "leaf".[6]

Distribution and habitat

The American beech is native to eastern North America, from Nova Scotia west to southern Ontario in southeastern Canada, west to Wisconsin and south to eastern Texas and northern Florida in the United States.[7] Mature specimens are rare in lowland areas as early settlers quickly discovered that the presence of the tree indicated good farmland.

The American beech is a shade-tolerant species,[3] commonly found in forests in the final stage of succession. Few trees in its natural range other than sugar maple match it for shade tolerance. Ecological succession is essentially the process of forests changing their composition through time; it is a pattern of events often observed on disturbed sites.[8] Although sometimes found in pure stands, it is more often associated with sugar maple (forming the beech–maple climax community), yellow birch, and eastern hemlock, typically on moist, well-drained slopes and rich bottomlands. Near its southern limit, it often shares canopy dominance with southern magnolia. Although it has a reputation for slow growth (sometimes only 13 feet in 20 years), rich soil and ample moisture will greatly speed the process up. American beech favors a well-watered, but also well-drained spot and is intolerant of urban pollution, salt, and soil compaction. It also casts heavy shade and is an extremely thirsty tree with high moisture requirements compared to oaks, so it has a dense, shallow root system.[9]

Ecology

The mast (crop of nuts) from American beech provides food for numerous species of animals. Among vertebrates alone, these include various birds[3] including ruffed grouse and wild turkeys, raccoons,[3] foxes,[3] white-tailed deer,[3] rabbits, squirrels,[3] opossums, pheasants, black bears,[3] and porcupines.[3] Some Lepidoptera caterpillars feed on beeches. Deer occasionally browse on beech foliage,[3] but it is not a preferred food. Beech nuts were one of the primary foods of the now-extinct passenger pigeon;[3] the clearing of beech and oak forests is pointed to as one of the major factors that may have contributed to the bird's extinction.[10]

Diseases and pests

Beech bark disease has become a major killer of beech trees in the Northeastern United States. This disease occurs when the European beech scale insect, Cryptococcus fagisuga, attacks the bark, creating a wound that is then infected by Neonectria ditissima or Neonectria faginata, two species of fungi. This causes a canker to develop and the tree is eventually killed.[11]

Beech leaf disease is caused by the nematode, Litylenchus crenatae mccannii. It was discovered in Ohio in 2012 and identified as far south as Virginia in 2022.[12] Beech leaf disease causes severe damage to the American beech and also to the European beech, Fagus sylvatica[13]

The beech leaf-miner weevil, a species native to Europe, has been identified in North America as a cause of defoliation of American beech trees.[14]

Beech blight aphids colonize branches of the tree, but without serious harm to otherwise healthy trees. Below these colonies, deposits of sooty mold develop caused by the fungus Scorias spongiosa growing saprophytically on the honeydew the insects exude. This is also harmless to the trees.[15]

Despite their high moisture needs, beeches succumb to flooding easily and their thin bark invites damage from animals, fire, and human activities. Late spring frosts can cause complete defoliation of the tree, although they typically recover by using reserve pools of sugar.[16] The trunks of mature beeches often rot and develop cavities that are used by wildlife for habitation.[17]

Uses

American beech is an important tree in forestry. The wood is hard and difficult to cut or split, although at 43 pounds per cubic foot (0.69 g/cm3) it is not exceptionally heavy, and it also rots relatively easily. It is used for a wide variety of purposes, most notably bentwood furniture as beech wood easily bends when steamed. It also makes high quality, long-burning firewood.

Like European beech bark, the American beech bark is smooth and uniform, making it an attraction for people to carve names, dates, decorative symbols such as love hearts or gang identifiers, and other material into its surface.[18] One such beech tree in Louisville, Kentucky, in what is now the southern part of Iroquois Park, bore the legend "D. Boone killed a bar. 1775" in the late 18th century. The beech finally fell over in 1916 during a storm; its age was estimated at around 325 years. Its trunk is now on display at the Filson Historical Society.

It is sometimes planted as an ornamental tree, but even within its native area, it is planted much less often than the European beech. Although American beech can handle hotter climates,[19] its European cousin is faster-growing and more pollution-tolerant, in addition to being easier to propagate.

American beech does not produce significant quantities of nuts until the tree is about 40 years old. Large crops are produced by 60 years. The oldest documented tree is 246 years old.[20] The fruit is a triangle-shaped shell containing 2–3 nuts inside, but many of them do not fill in, especially on solitary trees. Beech nuts are sweet and nutritious,[3] can be eaten raw by wildlife and humans,[21] or can be cooked.[22] They can also be roasted and ground into a coffee substitute.[3]

The leaves are edible when cooked.[3] The inner bark can be dried and pulverized into bread flour as an emergency food.[3]

References

  1. ^ Barstow, M. (2017). "Fagus grandifolia". IUCN Red List of Threatened Species. 2017: e.T62004694A62004696. doi:10.2305/IUCN.UK.2017-3.RLTS.T62004694A62004696.en. Retrieved 11 April 2020.
  2. ^ "Trees of the Adirondacks: American Beech | Fagus grandifolia". wildadirondacks.org.
  3. ^ a b c d e f g h i j k l m n o Angier, Bradford (1974). Field Guide to Edible Wild Plants. Harrisburg, PA: Stackpole Books. p. 32. ISBN 0-8117-0616-8. OCLC 799792.
  4. ^ Farahat, Emad; Lechowicz, Martin J. (2013). "Functional Ecology of Growth in Seedlings Versus root Sprouts of Fagus grandifolia Ehrh". Trees. 27 (1): 337–340. doi:10.1007/s00468-012-0781-9. S2CID 17109648.
  5. ^ "Fagus grandifolia Ehrh. - Plants of the World Online". Plants of the World Online. Retrieved 2023-04-24.
  6. ^ Hyam, R. & Pankhurst, R.J. (1995). Plants and their names : a concise dictionary. Oxford: Oxford University Press. pp. 169, 217. ISBN 978-0-19-866189-4.
  7. ^ Tubbs, Carl H.; Houston, David R. "Fagus grandifolia E h rh". www.srs.fs.usda.gov. United States Department of Agriculture. Retrieved June 22, 2021.
  8. ^ Horn, Henry S. (1974). "The Ecology of Secondary Succession". Annual Review of Ecology and Systematics. 5: 25–37. doi:10.1146/annurev.es.05.110174.000325.
  9. ^ North Carolina Cooperative Extension, Extension Gardener Tool Box, Fagus Grandifolia. North Carolina State University, Raleigh, NC, accessed July 9, 2022
  10. ^ Jon M. Conrad, "Open access and extinction of the passenger pigeon in North America", Natural Resource Modeling, Vol. 18, no. 4, pp. 501–519. 2005
  11. ^ Reed, Sharon E.; Volk, Daniel; Martin, Danielle K.H.; Hausman, Constance E.; Macy, Tom; Tomon, Tim; Cousins, Stella (January 2022). "The distribution of beech leaf disease and the causal agents of beech bark disease (Cryptoccocus fagisuga, Neonectria faginata, N. ditissima) in forests surrounding Lake Erie and future implications". Forest Ecology and Management. 503: 119753. doi:10.1016/j.foreco.2021.119753.
  12. ^ Kantor, M.; Handoo, Z.; Carta, L.; Li, S. (1 June 2022). "First Report of Beech Leaf Disease, Caused by Litylenchus crenatae mccannii , on American Beech ( Fagus grandifolia ) in Virginia". Plant Disease. 106 (6): 1764. doi:10.1094/PDIS-08-21-1713-PDN. PMID 34668402.
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Fagus grandifolia: Brief Summary

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Fagus grandifolia, the American beech or North American beech, is a species of beech tree native to the eastern United States and extreme southeast of Canada.

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