Beech (Fagus grandifolia) in the Plant Communities of Long Island, New York, and Adjacent Locations, and Some Comparisons Across Eastern North America

Abstract

A review was undertaken of the distribution of Fagus grandifolia (American beech) in ligneous communities on Long Island (Oak-Chestnut Association of the Eastern Deciduous Forest). There are seven forest types described in which beech is a dominant or co-dominant; and three microclimatic variants of those types. On Long Island, beech occupies moist well-drained to overly well-drained sites, either in pure stands, in combination with a few other tree species of moist soils, or in combination with tree species of well-drained uplands. Dwarfed beech forest is represented on Long Island (“Grandifolia Sandhills”), as it is in coastal New England and in the “Beech Gaps” of the southern Appalachian Mountains. Finally, the phytosociological role of Fagus grandifolia in all the forest regions of eastern North America and Mexico is examined.

1 Introduction

American Beech (Fagus grandifolia) is widely distributed in eastern North America from approximately 29 to 47° N latitude (at which northern latitude it has been stable for 8000 years (Maycock 1994); and from the Atlantic Coast at 60° W longitude to 95° W longitude (Little 1971). According to Maycock (1994), it achieves an upper elevational limit of 1875 m in the Appalachian Mountains and reaches 1000 m in the Adirondack Mountains. It occurs in 16 USDA “Forest Cover Types.” It is associated with 18 dominant trees in those communities. The only forest region of eastern North America in which it does not occur is the Maple-Basswood Association. Carpenter (1974) summarized the distribution, description, growth characteristics and properties, as well as commercial aspects of the beech. Maycock (1994) reported that beech reaches 125 ft (38.1 m) tall and averages 60–80 ft (18.3–24.4 m) tall in southern Ontario, near its northern limits. It reaches 400 years of age in Pennsylvania. Camp (1951) argued for the recognition of three basic types of Beech in eastern North America, ‘White’, ‘Red’ and ‘Gray’. He outlined their distributions: White in the southeastern coastal plain, Gray at higher elevations and latitudes, and Red widely in the deciduous forest region. He said the beech of the Beech-Maple climax forest of this region is made up of genetic elements of the three basic types. Maycock (1994) summarized the putative distributions of each of the three races. Subsequent students of beech genetics have sometimes recognized the White Beech as a separate subspecies, F. grandifolia var. caroliniana (viz., A.J. Rehder, cf. Flora of North America 2012). More generally, Camp’s segregation of the three races has not been supported by recent investigations (Hardin and Johnson 1985). In Mexico, Fagus has a restricted distribution in montane forests that contain a number of Holarctic genera that are similarly deciduous (Miranda and Sharp 1950). It has been treated as a variety of Fagus grandifolia (F. g. var. mexicana) by Little (1965) and by Williams-Linera et al. (2003).

From its extensive range, it is clear that beech can tolerate a wide range of average winter temperatures, down to January Normals of −4 °C (Huntley et al. 1989). Maycock (1994) reported that beech tolerates extreme low temperatures of between −30 and −40 °F (−34.4 to −40 °C). He further stated that it reproduces by seeds and root sprouts. It is shade tolerant and co-occurs with Acer saccharum, which is also shade tolerant. It is fire tolerant. It can occur on a wide range of soils, and on two major soil types, Gray-Brown Podzolic in the north, and ‘Lateritic” in the south. Soils range in pH from 4.1 to 6.0, seldom exceeding pH 7.0.

Ecologically, beech occupy soils that are well-drained, mesic to hydro-mesic and not depleted of nutrients (Carpenter 1974). Thus, it often dominates stands on coarse soils at the bases of hills. Yim (1983) states that North American beech-dominated forests occur on mesic sites, on moderately acidic soils of moderate to high fertility, in a variety of topographic sites.

2 Classification of Forest Zones Where Beech Is Present in Eastern North America

Greller (1988) recognized three Forest Zones or Ecotones in eastern North America, the Conifer-Hardwoods Zonoecotone (CD, or Hemlock-White Pine-Northern Hardwoods of Braun 1950), the Deciduous Forest Zonobiome (DF), and the Deciduous Dicot-Evergreen Dicot-Conifer Zonoecotone (DEC). The latter includes most of the Southern Mixed Hardwood Forest of Quarterman and Keever (1962), excluding only those regions where evergreen dicots do not reach the canopy because of cold climate. Figure 10.1 shows all the forest regions of eastern North America, with CD located at the north and DEC at the southern edge and the associations of the Deciduous Forest between them. Figure 10.2 is a H.P. Bailey nomograph that shows the three zones can be distinguished climatically: Conifer-Hardwood (CD) from Deciduous Forest (DF) by warmth, specifically the radian of W = 12.5 °C. Evergreen Broadleaved-Deciduous Broadleaved-Evergreen Conifer Forest (DEC or Southern Mixed Hardwoods, in part) can be distinguished approximately from Deciduous Forest by an annual freeze frequency of 3% hour/year.

Fig. 10.1

Map of Eastern North America to show the forest associations according to Greller (1988), following Braun (1950) and Quarterman and Keever (1962). Code: CD = Conifers-Deciduous Dicots (northern hardwoods); [Decidous Forest Associations:] B-M = Beech-Maple [-northern hardwoods], MB = Maple-Basswood [-northern hardwoods], MM = Mixed Mesophytic, OH = Oak-Hickory, OPH = Oak-Pine [-Hickory-southern hardwoods]; DEC = [Southern] Deciduous Dicots–Evergreen Dicots-Conifers [in part, the Southern Mixed Hardwood Forest of Quarterman and Keever (1962)]

Fig. 10.2

H.P. Bailey Nomograph of the three forest zones of Eastern North America in which deciduous trees dominate the canopy (from Greller 1988). Code for Nomograph: A = Annual range of mean monthly temperatures; T = Mean annual temperature, W = Warmth in °C, % = Percentage hours of the year with temperatures ≤0 °C, td = number of days with temperatures above W, N = Water need [not analyzed]. Code for forest zones: 1–9 = DEC (Greller 1988); 10–23 = Deciduous forest (sensu Greller 1988); 24–35 = CD (Hemlock-White Pine Northern Hardwoods of Braun 1950). Refer to Greller (1988) for names of numbered stations

2.1 Conifer-Hardwoods Zonoecotone (CD)

This is the [Needleleaved] Coniferous/Deciduous [Broadleaved] Dicotyledonous Forest [Greller’s (1989) CD]. The following is a summary of the bioclimatology of the Conifer-Hardwoods Zonoecotone. It is abstracted from Greller (1989, 1990).

Walter’s (1979) zone: ZE VI–VIII; W = 11.5–12.5 °C (Cool); M = 32–51 (Intemperate-Subtemperate); f ≤ 0 °C > 10% hour/year; Growing season (td) = 104.1–137.9 days; S = 4.4–15.1 inches (Semiarid-Humid)

In the present study this type is considered to be separate from the Deciduous Forest, a practice that Braun (1950) acknowledges as common. According to Braun this region ranges from “Minnesota to Atlantic Coast, occupying a position between the Deciduous Forest and the Boreal (Conifer) Forest.” The CD, which Braun calls Hemlock-White Pine-Northern Hardwood Forest (HWPNH), is characterized by the pronounced alternation of deciduous dicot, coniferous, and mixed communities. The name Conifer/Deciduous Dicot Forest (CD), gives more emphasis to the physiognomy, and allows one to include Küchler’s (1964) Spruce-Hardwood forest type as a subdivision of equal rank to the HWPNH. “[Acer saccharum, Fagus grandifolia, Tilia americana, Betula lutea, Tsuga canadensis, and Acer saccharum, Fagus grandifolia, Tilia americana, Betula lutea, Tsuga canadensis, and Pinus strobus] are the climax dominants of the region as a whole…. [Pinus resinosa] and, in the east, [Picea rubens], are characteristic species of the region, the former in dominantly coniferous communities, the latter in dominantly deciduous communities” (Braun 1950). Referring to Little (1971), it is clear that the arborescent dominants of the CD are broadleaved deciduous taxa and evergreen conifers of northern distribution in eastern and central North America. In Wisconsin, Curtis (1959: 535) listed Dirca palustris and Sambucus pubens as characteristic shrubs of CD mesic forests; other common shrubs are Corylus cornuta and Lonicera oblongifolia. The diversity of shrub and herb layers is lower in CD than in DF mesic forests, and decreases with an increase of coniferous trees (Braun 1950; Rogers 1982). Curtis (1959) lists 21 common angiospermous herbs.

2.2 Deciduous Forest Zonobiome (DF)

This forest zone (Zonobiome) has, as arborescent dominants, taxa that are cold-tolerant and deciduous. Deciduous taxa also dominate a well-developed subcanopy tree layer, a discontinuous shrub layer, and a dense, rich, seasonally varied herb layer. Evergreen dicotyledons, when present, are confined to the subcanopy tree, shrub, and herb layers; they comprise little of the total basal area and belong to only a few taxa. Dominant deciduous trees have wide distributions in the eastern United States, for example: Quercus, Fagus, Acer, Tilia, Carya, Fraxinus, Ulmus, Betula, Liriodendron, and (formerly) Castanea. Tsuga canadensis, an evergreen conifer, is a dominant on some mesic sites. The following is a summary of the bioclimatology of the Deciduous Forest Zonobiome (DF) of eastern North America. It is abstracted from Greller (1989, 1990). Greller (2013) treated the Oak-Pine[-Hickory] Association of DF as a separate ZE for purposes of climatic analysis.

Deciduous [Broadleaved] Forest [Greller’s (1989) DF].

Walter’s (1979) zone: ZB VI: W = 12.5–14.9 °C (Cool/Mild); M = 36–53 (Subtemperate-Temperate); f ≤ 0 °C = 3 to 10% hours/year; S = 6.8–13.3 inches (Subhumid-Humid)

Growing season (td) = 137.9–209.0 days.

2.3 Deciduous Dicot-Evergreen Dicot-Conifer Zonoecotone (DEC)

This is the zonal type of the Gulf and Atlantic Coastal Plains. Here the canopy comprises three physiognomic types: deciduous angiosperms, evergreen broadleaved angiosperms, and evergreen conifers. Among the dominant deciduous angiosperms are Fagus grandifolia, Liquidambar styraciflua, Carya glabra, Quercus spp., and often, Acer spp. Among the dominant evergreen angiosperms are: Magnolia grandiflora, Quercus hemisphaerica, Ilex opaca and, locally, Q. virginiana. Dominant conifers are: Pinus glabra, P. taeda and P. echinata. The following is a summary of the bioclimatology of the Deciduous Dicot-Evergreen Dicot-Conifer Zonoecotone (DEC) of southeastern North America. It is abstracted from Greller (1989, 1990).

Deciduous Dicot—Evergreen Dicot—Coniferous Forest (Greller’s [1989] DEC). It is equivalent to Quarterman and Keever’s (1962) Southern Mixed Hardwoods forest, excluding the stands where evergreen trees do not reach the canopy.

Walter’s zone = VI–V; W = 14.7–15.7 °C (Mild/to Warm); M = 49–53 (Temperate); f ≤ 0 °C = 0.5 to 3.0% hours/year; Growing season (td) = 203.1–233.7 day; S = 8.8–13.0 inches (Humid).

2.4 Mexican Holarctic Dicotyledonous Forest (MHDF, cf., Greller 1990)

Another region of North America possesses small and scattered populations of Fagus (F. mexicana or F. grandifolia var. mexicana). That is the montane zone of the Sierra Madre Oriental, from Tamulipas to Veracruz States. When analyzed using the H.P. Bailey nomograph, Mexican Holarctic Dicotyledonous Forest (MHDF, cf., Greller 1990) weather stations in that forest type are congruent in Warmth with the DEC of Greller 1990 (in part, Southern Mixed Hardwoods Forest of Quarterman and Keever 1962) and with the northern coastal stations of the Temperate Broadleaved Evergreen Forest (TBEF, sensu Greller 1980, 1990).

3 Classification of Forests Containing Beech on Long Island

Long Island beech trees usually have a single trunk, and mature trees reach a height of 70–100 ft. Sexual reproduction resulting in the production of beech nuts is relatively uncommon; asexual reproduction via clone production is common.

Greller (1977, 2001, 2002, cf. Reschke 1990) summarized a classification of Long Island forests. Those types that include Fagus grandifolia in the canopy are given in an extracted and slightly modified form below.

(MACROCLIMATIC TYPES)

• I. Upland Forests, Woodlands, and Shrublands (Well-Drained Sites)

  • i. Non-Oak Hardwoods Dominant

    • A. Tuliptree Series

      • 1. Mixed Mesophytic Association (tuliptree, beech, red oak, red maple, black birch, sweetgum, white ash. ex., Nassau Co.: Grace Forest, North Hills (largely extirpated), Glen Cove, Welwyn Preserve; Queens Co.: Tuliptree Trail, Alley Park.

    • B. Beech Series

      • 2. Beech-Mixed Hardwoods (incl. red oak, red maple) Association; ex., Nassau Co.: Meadowbrook Park (north), William Cullen Bryant Preserve, Roslyn.

      • 3. Beech Consociation; ex., Nassau Co.: southern edge of Hempstead Plains (Harper 1918), Fox Hollow Preserve, Syosset; Suffolk Co.: Cold Spring Harbor vic., Coastal Sand Dunes on the Bluffs Above Long Island Sound, Riverhead.

  • ii. Oaks Dominant

    • D. Black oak Series

      • 7. Black oak-beech-black birch Association; ex., Queens Co.: Oakland Lake; Nassau Co.: Whitney Est., Manhasset; Glen Cove; Suffolk Co.: Jayne’s Hill, West Hills Co. Pk.; Lloyd Neck, Huntington; Riverhead Bluffs, Riverhead.

• II. Swamp Forests and Related Shrubland (Edaphic Forests of Flooded Soil)

  • i. Hardwood Swamps

    • B. Red Maple Series (red maple dominant or co-dominant)

      • 8. Red Maple-American Beech Association; ex., Suffolk Co.: Point Woods, Camp Hero SP, Montauk (now rare; Taylor 1923).

    • F. Beech Series (Beech dominant or co-dominant)

      • 19. Beech-sweetgum Association; ex., Nassau Co.: Wantagh Creek, North Bellmore

      • 20. Beech-Sourgum-Bitternut-Red Maple Association; ex., Suffolk Co.: Point Woods, Montauk Point SP (common).

(MICROCLIMATIC TYPES)

• III. Coastal Variants of Forests on Well Drained Sites

  • A. Black Oak Series

    • 1. Black Oak-Beech [-Black Birch]/American Holly Association ex., Suffolk Co.: Montauk Pt. St. Pk.

  • B. Tuliptree Series

    • 2. Mixed Mesophytic/Ilex opaca var. opaca/Polystichum acrostichoides; Nassau Co.: Cove Neck (Edinger et al. 2008)

• IV. Maritime Forests and Shrublands

  • A. Beech Series

    • 1. Dwarfed Beech Low Woodland Association ex., Suffolk Co. : Friar’s Head (on “Grandifolia Sandhills”), Riverhead

4 Results and Discussion

4.1 Beech in the Deciduous Forest Zonobiome (DF) (All Photos by A.M. Greller Unless Otherwise Indicated)

4.1.1 Beech in the Oak-Chestnut Forest Association of the Eastern Deciduous Forest (DF)

4.1.1.1 Long Island

4.1.1.1.1 Mixed Mesophytic Community (Fig. 10.3)

I. Upland Forests, Woodlands, and Shrublands (Well-Drained Sites); Non-Oak Hardwoods Dominant; (A). Tuliptree Series; (1). Mixed Mesophytic Association (tuliptree, beech, red oak, red maple, black birch, sweetgum, white ash); ex., Nassau Co.: Grace Forest, North Hills (largely extirpated), Glen Cove, Welwyn Preserve; Matinecock, Coffin Woods; Queens Co.: Tuliptree Trail, Alley Park

Fig. 10.3

Fagus grandifolia with Liriodendron tulipifera, Coffin Woods, Matinecock, Nassau County, NY

Greller et al. (1978) described in some detail a Mixed Mesophytic stand at North Hills, Nassau County, N.Y. The dominant trees were: Liriodendron tulipifera (51.3% rel. dom.), Quercus [rubra] 11.9%), Fagus grandifolia (7.5%), Acer rubrum (4.5%), Quercus velutina (4.1%), and Quercus coccinea (3.5%). Other sub-dominants include: Acer saccharum, Betula lenta, Carya glabra, Cornus florida, Fraxinus americana, Liquidambar styraciflua, and Quercus palustris, among others. At 35–54 cm below the surface, the soil was mottled. Thus, the water table was high and it fluctuated. This explains the relative paucity of Quercus and Carya species on the site.

4.1.1.1.2 Beech-Black Birch-Black Oak Forest (Fig. 10.4)

Upland Forests, Woodlands, and Shrublands (Well-Drained Sites); Oaks Dominant; (D) Black oak Series; (7). Black oak-beech-black birch Association; ex., Queens: Oakland Lake; Nassau Co.: Greentree (fmr. Whitney Est.), Manhasset; Glen Cove; Suffolk Co.: Jayne’s Hill, West Hills Co. Pk.; Lloyd Neck, Huntington; Riverhead Bluffs, Riverhead.

Fig. 10.4

Beech-Black Birch-Black Oak forest at “Greentree,” Manhasset, Nassau County, NY

Good and Good (1970) presented quantitative data on a forest that included oaks, beeches, birches and other hardwood trees on the bluffs above Long Island Sound, north of Riverhead (Friar’s Head). This type of forest is perhaps the most common one in which beech occurs on Long Island. Fagus, Quercus velutina and Betula lenta appear to be regularly scattered throughout the forest. Acer saccharum occurs here with more frequency than in other forest types of Long Island, as does Ostrya virginiana.

4.1.1.1.3 Beech-Mixed Hardwoods (Fig. 10.5)

Upland Forests, Woodlands, and Shrublands (Well-Drained Sites); Non-Oak Hardwoods Dominant; (B). Beech Series; (2). Beech-Mixed Hardwoods (incl. red oak, red maple) Association; ex., Nassau: Meadowbrook Park (north); William Cullen Bryant preserve, Roslyn.

Fig. 10.5

Beech-Mixed Hardwoods, William Cullen Bryant Preserve, Roslyn, New York

In the north-facing, forested ravine in William Cullen Bryant Preserve, Roslyn, Nassau County, beech occurs from the upper rim of the ravine down nearly to the coast of Long Island Sound. At the rim, Fagus occurs scattered with Quercus spp., Carya spp., Liriodendron, Acer spp., Ostrya virginiana, and Betula lenta. On steeper slopes in the ravine, beech occurs with Quercus species such as Q. rubra, Q. velutina and Q. montana.

4.1.1.1.4 Beech Consociation (Fig. 10.6)

Upland Forests, Woodlands, and Shrublands (Well-Drained Sites); (i). Non-Oak Hardwoods Dominant; (B). Beech Series; (3). Beech Consociation: Nassau Co.: Southern edge of Hempstead Plains (Harper 1918); Suffolk Co.: Coastal sand dunes on the bluffs above Long Island Sound; scattered throughout moist sites in Point Woods, Montauk.

Fig. 10.6

Stand of pure beech (consociation) in Point Woods, Montauk, Suffolk Co., NY

Lamont (1998) gave a detailed description of the sand dunes that cap the remnant of the terminal moraine north of Riverhead, Long Island. These dunes stretch about 1.25 miles and have a maximum width of about 0.5 mile; they have been known to science for over 120 years. “Myron L. Fuller (1914), in his geological survey of Long Island, extensively discussed the sand dunes, devot[ing] a full page plate to photographs of sand dunes.” Elias Lewis, Jr. recorded the dunes as rising 150 ft above tide, but standing on the bank (moraine) which is half that height, so that 75 ft of that elevation is drifting sand. At least five distinct but interrelated plant communities occur in the sandhills: (1) Coastal Beech Forest, (2) Maritime Dwarf Beech Forest, (3) Maritime Dunes, (4) Pitch Pine-Oak Duneland, and (5) Maritime Shrubland. The first four communities are rare occurrences in New York.

One site for Beech Consociation occurs at Wildwood State Park, approximately 3 miles west of the Riverhead sandhills; another site is located approximately 2.5 miles east of the sandhills. Frederick C. Schlauch included approximately 250 acres of beech-dominated vegetation on coastal dunes in that area. According to Lamont (1998), coastal beech forest occurs throughout the protected depressions of the undulating duneland south of the bluffs. Nowhere else in New York does such a mature climax forest occur on sand dunes. Sugar maple (Acer saccharum) occurs scattered throughout the coastal beech forest; and a there is a grove of Canada hemlock (Tsuga canadensis). Both species occur much more widely further north, but are uncommon south, in forests of the Oak-Chestnut Association. Good and Good (1970) published a detailed phytosociological description of the coastal beech forest of Friars Head, north of Riverhead. Beech can be seen in pure stands on old stabilized dunes in the Point Woods, Montauk.

4.1.1.1.5 Beech-Mixed Hardwoods Swamp (Figs. 10.7 and 10.8)

II. Swamp Forests and Related Shrubland (Edaphic Forests of Flooded Soil); (i). Hardwood Swamps; (B). Red Maple Series (Red Maple dominant or co-dominant); (8). Red Maple-American Beech Association; ex., Suffolk: Point Woods, Montauk Point SP (Taylor 1923, p. 55); F. Beech Series (Beech dominant or co-dominant); (20). Beech-Sourgum-Bitternut Hickory-Red Maple Association; ex., Suffolk Co.: Big Reed Pond Trail, Montauk Point.

Fig. 10.7

Beech with Carya cordiformis near Big Reed Pond, Montauk

Fig. 10.8

Beech and red maple swamp. Camp Hero, Point Woods, Montauk

At Big Reed Pond, Montauk, beech occurs in a continuum of soils from well drained to very moist. It is seen on wetter soils where Symplocarpus foetidus is present in springs or seeps, intermixed with Nyssa sylvatica or Carya cordiformis, or above Acer rubrum on slightly better drained sites (pers observ., Mar 27–30, 2012). Taylor (1923) described extensive Beech-Red Maple stands at Point Woods, Montauk. Today, Beech-Red Maple is a fragmentary association and is rarely encountered in the many swamps of the Montauk Point Woods.

4.1.1.1.6 Oak (Quercus)-Beech (Fagus)/Ilex opaca/Kalmia latifolia (Fig. 10.9)

III. Coastal and Highland Variants of Forests on Well Drained Sites; (A). Black Oak Series; (1). Black Oak-Beech-Black Birch/American Holly Association ex. Suffolk Co.: Montauk Pt. St. Pk.

Fig. 10.9

Fagus grandifolia with Quercus spp., Ilex opaca and Kalmia latifolia on morainal till in Point Woods, Montauk

In the Point Woods, Montauk, Fagus can be seen on well-drained morainal uplands in combination with Quercus spp., and an occasional Betula lenta, with an understory of Ilex opaca and a shrub layer of tall Kalmia latifolia.

4.1.1.1.7 Maritime Dwarfed Beech Forest (Fig. 10.10)

IV. Maritime Forests and Shrublands; (A). Beech Series; (1). Dwarfed Beech Low Woodland Association ex., Suffolk: Friar’s Head [on “Grandifolia Sandhills”], Riverhead).

Fig. 10.10

Dwarfed beech emerging from dune sand near a large Quercus velutina, McQuaid Property, Riverhead, NY

Lamont (1998) summarized information on the dwarfed beech forests of the Riverhead, Long Island area as follows: they occur on the bluff face, on the bluff top, and on dune ridges. He stated that this unique plant community is considered to be globally rare. Good and Good (1970) published a detailed phytosociological description of the maritime dwarfed beech-forest of Friars Head, north of Riverhead. In addition to those on the 140 acres of sandhills at Friars Head, Riverhead (“Grandifolia Sandhills”), several small, disjunct occurrences of the dwarf beech forest have been located and described by Mary Laura Lamont. The dominant species is a dwarf form of American beech; mature individuals are stunted [usually less than 12 ft (3.66 m) tall], often multi-stemmed from the base, and extremely gnarled and contorted. Century old trees often grow horizontally before sending twisted limbs skyward, only to be pruned back by constant exposure to salt spray, sand blowup, cold wind, and winter ice. Asexual reproduction by root crown sprouts may play a significant role in the success of the species at the “Grandifolia Sandhills,” the major stand of L.I. Maritime Dwarfed Beech Forest. Colonies of dwarf beech trees on the seaside bluffs probably constitute a single persistently, organically connected clone (Lamont 1998). Other species associated with the dwarf beech include red maple (Acer rubrum), hickory (Carya glabra and C. tomentosa), black oak (Quercus velutina), and shadbush (Amelanchier canadensis). Lamont used an increment borer to determine the age of one of the old dwarf beech trees as 138 years, in 1998.

4.1.1.2 Other Areas Near Long Island

Westchester and Putnam Cos

North of New York City, at Ice Ponds Conservation Area, Putnam County, New York, Beech-Maple mesic forest occurs where the water table is high (but below the surface) and the soil is slightly acidic (pH 5–6). Native tree, shrub and herb diversity is highest here. The dominant trees are Liriodendron, Fagus, Betula lenta and B. allegheniensis, Quercus rubra and Q. velutina, Acer rubrum, Prunus serotina, Ulmus rubra, and Carya spp. Lindera benzoin, Viburnum acerifolium and Hamamelis virginiana are the common shrubs. The herb layer contains Trillium, Asarum canadense, Polystichum, Caulophyllum, Maianthemum canadense and Sanguinaria. Lianas include much Vitis aestivalis and some Celastrus scandens.

New Jersey

West of New York City, Collins and Anderson (1994) report that Fagus grandifolia is present in both physiographic regions—the coastal plain of “South Jersey” and the piedmont and highlands of “North Jersey.”

In Southern New Jersey, only a small part of the Inner and Outer Coastal Plains can be classified as mesic uplands. Two major forest types occur here, Mixed Oak and Beech-Oak.

• A. Mixed Oak. In the western and southwestern part of the coastal plains, upland forests comprise the following trees: Quercus alba, Q. velutina, Fagus grandifolia, Carya glabra and C. alba, Juglans nigra, Liriodendron, and Acer rubrum; less common are: Quercus falcata, Diospyros virginiana, Liquidambar, Pinus virginiana and Ilex opaca. Understory trees are: Cornus florida, Carpinus caroliniana, Prunus serotina and Sassafras. Vines are Lonicera japonica (exotic and invasive), Parthenocissus quinquefolia, Vitis spp., Toxicodendron radicans. Fagus is a relatively less important constituent of this forest, after Quercus spp., Ilex and Sassafras.

• B. Beech-Oak. Fagus is a more important constituent of the Beech-Oak Forest. Near Camden, N.J., pure stands of Fagus were reported by W. Stone in 1910 [Fagus Consociation]. In general, beech intergrades into the Mixed Oaks Forest. In the Mount Holly area, Chesterfield and Burlington Cos., Fagus occurs in large numbers in all size classes, with Liriodendron, Quercus velutina, Carya ovata, C. glabra and Acer rubrum; Cornus florida and Carpinus caroliniana are scattered throughout. At Mill Creek Park in Willingboro, Burlington Co., Fagus composes more than 50% of the forest, associated with Quercus alba, Q. montana, Q. velutina, Liriodendron, Ilex opaca; other trees are Prunus serotina, Quercus falcata, Sassafras, Liquidambar, Acer rubrum, Pinus virginiana, Diospyros virginiana, Carya [alba] and C. glabra. In Monmouth Co., at the junction of the coastal plain and piedmont regions, beech occurs with black birch (Betula lenta) and tuliptree (Liriodendron). Other constituents of this mixed mesophytic community are Quercus rubra and Q. velutina, Liquidambar and Sassafras; understory comprises Cornus florida, Viburnum acerifolium, Viburnum dentatum, Lindera benzoin, Prunus serotina and Smilax glauca.

In Northern New Jersey, on Kittatiny Limestone, in High Point State Park, Collins and Anderson (1994) recognize a Sugar Maple-Mixed Hardwood Forest. Fagus is a minor component of this forest. Dominant species are: Acer saccharum, with Quercus alba, Q. velutina, Q. rubra; also Fraxinus americana, Liriodendron tulipifera, Betula lenta, B. allegheniensis, Acer rubrum, Tilia americana, Fagus grandifolia, Carya spp., scattered Tsuga canadensis, Pinus strobus, Ulmus americana, Juglans nigra.

4.1.1.2.1 Coastal New England

Busby and Matzkin (2009) described a Dwarf Beech Forest in coastal New England “on morainal knobs and ridges characterized by excessively well-drained, sandy soils and well-developed E horizons; soil organic horizon …absent or minimal” due to high winds that remove it. They state that tall beech appears in adjacent depressions on fine-texture soils, with greater soil fertility and protection from the wind. In summary they said, “dwarf growth is a response to harsh edaphic conditions including chronic nutrient depletion, drought stress and wind exposure.”

4.1.1.2.2 Great Smoky Mountains

The Oak-Chestnut Association is centered on the southern Appalachian Mountains. Above about 1100 m, beech occurs in forests of all ranges of “mesic” (Yim 1983). The reader is referred to Whitaker (1956) for a thorough discussion of forest composition in the Great Smoky Mountains. The only significant associates of Fagus grandifolia in the Oak-Chestnut region are: Acer saccharum, Acer rubrum, Quercus rubra, Betula lenta, Liriodendron tulipifera, and Quercus alba. The following dominance types, sampled by Braun (1950), are listed by Maycock (1994): Fagus grandifolia-Acer saccharum, Fagus grandifolia, Fagus grandifolia-Quercus alba, Fagus-Castanea dentata. The first listed type was more commonly sampled than all the other types combined.

4.1.2 Beech in the Oak-Pine[-Hickory] Forest Association of the Eastern Deciduous Forest (DF)

In this association (for detailed tree composition see Greller 2013) on the Atlantic and Gulf Coastal Plains, beech is confined to the narrow zone of mesic soils of higher nutrient content. It is absent from most of the upland sites, which are sandy, dry and nutrient-poor (Yim 1983). The mixed mesophytic community in this forest association differs from that in the Oak-Chestnut Association in having a greater, often leading dominance by Fagus grandifolia, greater dominance by Quercus, especially Q. alba, and an understory that includes Ilex opaca. Nesom and Treiber (1977) listed the following taxa and their percentage dominance in three stands: Fagus grandifolia (49, 47, 82% dom.), Quercus alba (9, 10, 8), Liquidambar styraciflua (20, 5, 1), Liriodendron tulipifera (12, 8, 1), Quercus rubra (6, 5, 4), Carya [alba] (1, 10, 1), Nyssa sylvatica (0, 5, 2), Quercus falcata (0, 5, 1), as well as others. Ware (1970), working on the coastal plain in Williamsburg, Virginia, described a mesic forest dominated by Querus alba (21.9% dominance), Pinus taeda (16.5), Fagus grandifolia (13.1), Quercus falcata (13.1), Liriodendron (10.8), Pinus virginiana (5.1), Quercus rubra (4.6), as well as lesser percentages of Acer rubrum var. trilobum, Carya spp., Liquidambar, Cornus florida, and Quercus nigra. Yim (1983) summarized previous work on an Oak-Beech Forest near Washington, D.C. Here, Quercus alba accounted for 41.0% basal area and Fagus 24.2%; (MAT = 14.1 °C; MAP = 1029. Habitat was a slope of 10–25%).

4.1.3 Beech in the Beech-Maple Forest Association of the Eastern Deciduous Forest (DF)

According to Maycock (1994) extreme southern Ontario between three Great Lakes, below 43° N latitude, has a summergreen deciduous forest of 85 species of trees. This region is the northern limit of the Beech-Maple Association of the Eastern Deciduous Forest. It is a limestone region of low elevation 125–400 m a.s.l; Clay Plains and Sand Plains account for 90% of site types, but Till Plains and morainal hills are present. Acer saccharum, Fagus grandifolia, Quercus rubra and Ulmus americana are the leading dominants. Other important taxa are Acer rubrum, Acer saccharinum, Quercus velutina and Q. alba; other “significant” species are: Acer negundo, Betula papyrifera, Carya cordiformis, C. ovata, Celtis occidentalis, Populus deltoides, P. tremuloides, Quercus macrocarpa, Q. palustris, Salix amygdaloides, S. nigra, Tsuga canadensis and Ulmus rubra. Maycock (1994) gave detailed composition and structure data for some of these stands.

Ohio comprises three of the Associations of the Eastern Deciduous Forest, Beech-Maple, Mixed Mesophytic, and Western Mesophytic in that order of areal extent. Gordon (cf. Yim 1983) recognized five beech-dominated forest types in Ohio: Beech-Sugar Maple, Wet Beech, Hemlock-Beech, Beech-Maple-Tuliptree, and Beech-White Oak. “Wet beech” forests occur on mounds above floodplains, but may endure occasional inundation. Beech forests do not occur on well-drained or “droughty” soils in Ohio. In Cincinnati, Ohio, a till plain mesophytic forest has 50% basal area for Fagus; the MAT= 9.9 °C and MAP = 1208 mm (cf. Yim 1983). In the Ontario lowlands of western New York State, where the substrate is lowland glacial drift of dolomitic limestone, Miller in 1973 (cf. Yim 1983) describes a Maple-Beech Forest where Acer saccharum accounts for 61% of basal area; here the MAT = 8.4 °C. In the “Beech Border” of Wisconsin, Ward in 1961 (cf. Yim 1983) describes a Beech-Maple Forest where Fagus accounted for 9.4% of basal area. Soil pH ranges from 5.5 in the north to 6.5 in the south.

For Warren Woods, Michigan, Yim (1983) summarizes previous work on a Beech-Maple Forest in a till plain, in which Fagus accounts for 50% of basal area, and Acer saccharum 12%; where MAT = 9.9 °C and MAP = 1020 mm.

4.1.4 Beech in the Mixed Mesophytic Forest Association of the Eastern Deciduous Forest (DF)

Braun (1950) compiled extensive data on forest composition in this association. Maycock (1994) summarized the stand data as follows: Fagus is the sole dominant in 9 of Braun’s stands, Fagus-Liriodendron co-dominated in 4 stands, Fagus-Tsuga canadensis co-dominated in 3 stands, Fagus-Acer saccharum co-dominated in 2 stands, Fagus-Tilia heterophylla co-dominated in 2 stands. The following co-dominated stands occurred once in Braun’s data tables: Liriodendron-Fagus, Liriodendron-Tsuga canadensis-Fagus, Castanea dentata-Fagus (sic). In addition, Maycock reported the following Fagus co-dominated types in the Mixed Mesophytic Association region: Fagus-Quercus alba, Quercus alba-Fagus, Fagus-Magnolia acuminata, Fagus—Picea rubens, and Tsuga canadensis-Fagus.

4.1.5 Beech in the Western Mesophytic Association of the Eastern Deciduous Forest (DF)

Maycock (1994) reported that in this association, beech domination occurs with trees of southern distribution such as Tilia neglecta, Quercus schumardii, Carya glabra, Tilia heterophylla, Aesculus octandra, Carya [alba] and Magnolia acuminata. The following tree species had Constancy greater than 70%: Acer saccharum, Fraxinus americana, Carya ovata, Quercus alba and Liriodendron tulipifera. Fagus grandifolia accounts for 42% of all trees in the stands in which it dominates, accompanied by Acer saccharum (11%), Liriodendron (8), Quercus alba (5), Fraxinus americana (4), Carya ovata (3), Tilia heterophylla (3), and Aesculus octandra (3). Maycock listed the following forest types for this association: Fagus, Fagus-Acer saccharum, Fagus-Liriodendron, Fagus-Quercus alba, Fagus-Aesculus octandra, Fagus-Tilia heterophylla. The most commonly sampled type was Fagus-Acer, followed by Fagus and Fagus-Liriodendron. Greller (2013) assigned the southern portion of this Association, in the Mississippi Valley, to the Oak-Pine Forest Association.

4.1.6 Beech in the Oak-Hickory Forest Association of the Eastern Deciduous Forest (DF)

This is the western Association of the Deciduous Forest. Here the natural uplands are often vegetated by savanna and prairie. Beech-dominated forests occur on middle or lower slopes. In northwestern Arkansas and the Boston Mountains south of the Ozark Mountains, in a Beech-Oak forest, Fagus grandifolia occurs with Nyssa sylvatica and Quercus rubra, Q. alba, Ulmus americana, Tilia floridana, Acer saccharum, Carya ovata and C. cordiformis (Maycock 1994).

4.1.7 Beech in the Beech-Birch-Maple-Basswood Forest Association of the DF in the Southern Appalachian Mountains of the Eastern United States

Greller (1988) recognized a combination of non-oak deciduous trees at elevations of approximately 1000–1400 m in the Southern Appalachian Mountains, as constituting a new forest Association in eastern North America, (BBMB), based on the Mesophytic Forest of Rheinhardt and Ware (1984). Those authors described a few types of forest in this zone: Beech-Maple, Mesophytic and Dwarf Beech. They gave composition data for each type. It is their Mesophytic type that distinguishes the Association.

Mesophytic Forest (Fig. 10.11) is the most variable type of forest in the Balsam Mountains of southwestern Virginia. Rheinhardt and Ware (1984) distinguish it from the Mixed Mesophyic forest and cove forests of the Oak-Chestnut Association because BBMB-Mesophytic type lacks Liriodendron, Tsuga, Quercus alba, Nyssa sylvatica, Juglans nigra and Carya spp. It does however include species that are lacking or uncommon in those lower elevation Mixed Mesophytic communities, such as Tilia heterophylla, Acer saccharum, Fagus grandifolia, Fraxinus americana, Betula lutea, Aesculus octandra, Quercus rubra and Acer rubrum. Mesophytic forest at high elevations is dominated by Acer saccharum (28%), Tilia heterophylla (22%), Fagus grandifolia (14%), Fraxinus americana (12%) and Betula lutea (6%). On south-facing slopes the leading dominants are Quercus rubra, Acer spp. and Fagus grandifolia. On north-facing slopes Tilia heterophylla and Fraxinus americana co-dominate, with lesser percentages of Acer saccharum and Fagus grandifolia.

Fig. 10.11

Mesophytic type of BBMB Association, with prominent Aesculus octandra trunks and abundant leaves of Fagus grandifolia from smaller trees; on Blue Ridge Parkway, North Carolina (2005)

The Beech-Maple type occurs on sheltered slopes and flat land at higher elevations. Here the beech is said to be the ‘Red’ race, and is usually the leading dominant, although Acer saccharum can be the leading dominant. Rheinhardt and Ware (1984) gave the following composition data: Fagus grandifolia (38% dominance), Acer saccharum (35%) and Betula lutea (10%).

The Dwarf Beech (Fig. 10.12) type has the following composition: (Fagus grandifolia (69% dominance), Aesculus octandra (13%) and Betula lutea (7%). This type occurs on and immediately below exposed summits above 1400 m a.s.l. Rheinhardt and Ware (1984) considered the Fagus grandifolia to be of the ‘Gray’ race. Accompanying the beech are small numbers of Aesculus octandra, Betula lutea and Acer saccharum. Fagus and Aesculus are stunted, with few stems greater than 6 m tall and 40 cm in circumference.

Fig. 10.12

Dwarf beech forest on the Blue Ridge Parkway in North Carolina (2005)

“Beech Gaps” (Fig. 10.13) are areas where beeches dominate in low shrub-like communities between peaks in the southern Appalachian Mountains. Whitaker (1956) gave some environmental and compositional data for that type in the Great Smoky Mountains: they occur at 1586–1708 m; MAT = 7–9 °C; MAP = 1250 mm; beech composes 50–80% of stems ≥2.5 cm, on mesic, rich soils.

Fig. 10.13

Beech Gap near the Blue Ridge Parkway, North Carolina

4.2 Beech in the Conifer-Hardwoods Zonoecotone (CD)

For central Ontario, Maycock (1994) gave some data on forest composition for a Dry-Mesic Forest and for a Mesic Forest. In the Dry-Mesic Forest the leading trees were: Fagus grandifolia (100 Constancy/129 Importance Value), Acer saccharum (100/70), Fraxinus americana (100/6), and Prunus serotina (100/1); other trees of high Constancy and IV are: Quercus rubra, Ostrya virginiana, Pinus strobus, Acer rubrum and Tsuga canadensis. For the Mesic Forest, the following taxa reached 100% Constancy: Fagus grandifolia, Acer saccharum, Tilia americana, Acer rubrum, Prunus serotina Tsuga canadensis and Betula lutea. Similar compositions occur in dry-mesic Beech-Maple forests in central Quebec, where Betula lutea, Acer pensylvanica and Abies balsamea show 100% Constancy values. In southern Quebec, in the Monteregian Hills (including Mt. Royal, Mt St. Bruno, etc.) composition of Dry-Mesic and Mesic stands of Beech-Maple Forest is similar to central Ontario, with the addition of Betula papyrifera, Acer pensylvanicum, and Prunus pennsylvanica, which also achieve high Constancy ratings.

4.3 Beech in the Deciduous Dicot-Evergreen Dicot-Conifer Zonoecotone (DEC)

Beech-dominated forests occur on floodplain terraces of small streams, in addition to lower slopes adjacent to stream floodplains. In those sites, beech is secondary to Magnolia grandiflora, Quercus spp., and others. Rarely, it is dominant on those sites. Those sites are mesic and of relatively high fertility compared to the surrounding sandy or loess-derived soils (Yim 1983). Ware et al. (1993) did a thorough review of the vegetation of this phytogeographic region, which is centered on the Gulf and Atlantic coastal plains. Beech (Fagus grandifolia ‘White’ race or Fagus grandifolia var. caroliniana) occurs in the following types of forest:

• 1. Southern Mixed Hardwoods (SMHF), where it occurs with a wide variety of deciduous trees, evergreen trees and evergreen conifers. Varying combinations of dominants have been described as: Lower Slope Hardwood Pine Forest, Magnolia-Beech-Holly (Fig. 10.14), Beech-Magnolia[-Maple], Beech-Spruce Pine-Magnolia, Oak-Hickory-Pine, Magnolia-Oak-Loblolly Pine. Beech is a canopy dominant in those so indicated.

• 2. Pine-Mixed Hardwoods, a mesic forest on nutritionally poorer sites than the SMHF, where Fagus grandifolia occurs as a sub-dominant in a forest dominated by Pinus palustris, P. taeda, P. elliottii var. elliottii, and Magnolia grandiflora, Liquidambar styraciflua, Quercus hemisphaerica, Quercus alba, Q. nigra, Q. stellata, Liriodendron tulipifera, Nyssa sylvatica, with Cornus florida in the understory.

Fig. 10.14

Fagus grandifolia, with Magnolia grandiflora and Ilex opaca in a lower slope forest, Indian Mounds Wilderness, Sabine Co., Texas, by J.T. Williams

4.4 Beech in Forests (MDHF) Dominated by Holarctic Taxa in the Sierra Madre Oriental of Mexico

Holarctic Dicotyledonous Forest (MHDF) or Bosque Mexicano de Dicotyledoneas Holarcticos, is the name Greller (1990) proposed for the mainly dicotyledonous mountain forests of eastern Mexico. MHDF has a canopy dominated by taxa of Holarctic floristic affinities, both deciduous and evergreen, as detailed by Miranda and Sharp (1950). It is endemic to Mexico and northern Central America, for nowhere else in North America do taxa such as Oreomunnea (Engelhardia; Juglandaceae), Clethra (Clethraceae), Dendropanax (Araliaceae), Podocarpus (Podocarpaceae), and Weinmannia (Cunoniaceae) dominate in the same stands as Quercus, Carpinus, Liquidambar, Juglans, Carya, and Pinus. Podocarpus and Weinmannia belong to the Austral or Southern Hemisphere Temperate Flora, and it is the admixture of Southern Hemisphere taxa into a matrix of Holarctic dominants that makes this ZE unique. Miranda and Sharp (1950) strongly emphasized this conglomeration of disparate floristic groups in arguing for the uniqueness of their predominantly Holarctic forests. MHDF is distributed from approximately 1000 to 2000 m elev., and occurs above the Orizaban Rain Forest (e.g., Gómez-Pompa 1973: Figs. 20, and 21). Miranda and Sharp (1950) listed six types of forest dominated by dicotyledons: “oak-mixed hardwoods; Liquidambar; Platanus; Fagus; Weinmannia; and Engelhardia [Oreomunnea].” The first type is most widely distributed. The canopy averages 20 m tall, but reaches 40 m, and includes: Quercus spp., Clethra quercifolia, Liquidambar [mexicana], Meliosma alba, Carpinus [tropicalis], Ostrya virginiana, Engelhardia [Oreomunnea] mexicana, and Cornus spp. There is a small-tree layer reaching 10 m tall, with Cyathea (a tree fern), Inga (Leguminosae), Oreopanax (Araliaceae), Xylosoma (Flacourtiaceae), Turpinia (Staphyleaceae), Symplocos (Symplocaceae), Podocarpus [matudae], some other tropical taxa (Beilschmiedia and Rapanea) and other Holarctic taxa (Vaccinium, Sambucus, Leucothöe, and Cornus). Epiphytes of Orchidaceae, Bromeliaceae, Cactaceae, and Araliaceae are present. A shrub layer is present, containing a number of Holarctic taxa and some tropical taxa (Drypetes, Randia). Lianas and vines are present, as well as some dicotyledonous herbs.

Williams-Linera et al. (2003) examined ten stands of Mexican Beech (Fagus grandifolia var. mexicana; Figs. 10.15 and 10.16). They stated that

species richness varied between 3 and 27 tree species in the canopy, and from 9 to 29 species in the understorey. Basal area of trees ≥5 cm dbh varied between 27.87 and 70.98 m² ha⁻¹, density from 370 to 1290 individuals ha⁻¹. Beech represented 22–99.6% of total basal area and 6.8–83.3% of total density. Beech dominance varied from monodominance to codominance with Carpinus caroliniana, Quercus spp., Liquidambar styraciflua, Magnolia schiedeana and Podocarpus spp. Beech population ranged from 180 to 8300 trees with a total of fewer than 1300 individuals on four of the sites. Anthropogenic disturbance remains a major threat to these forests. It is uncertain whether Mexican beech will be able to survive without conservation efforts.

Fig. 10.15

Beech forest in the Sierra Madre Oriental (from Adele Rowden, “Conservation genetics of a threatened Mexican tree species, Fagus grandifolia var. mexicana,” Ph.D. Thesis, University of Edinburgh, UK)

Fig. 10.16

Cattle grazing in a Fagus grandifolia var. mexicana forest in the Sierra Madre Oriental (from Adele Rowden, “Conservation genetics of a threatened Mexican tree species, Fagus grandifolia var. mexicana,” Ph.D. Thesis, University of Edinburgh, UK)