The Silvics of Castanea dentata

(Marsh.) Borkh., American Chestnut, 

Fagaceae (Beech Family) 

United States

Department of

Agriculture

Forest Service

Southern 

Research Station

General Technical

Report SRS-173

G. Geoff Wang, Benjamin O. Knapp, Stacy L. Clark, 

and Bryan T. Mudder
G. Geoff Wang, Professor, and Benjamin O. Knapp, Research Specialist, School 

of Agricultural, Forest and Environmental Sciences, Clemson University, Clemson, 

SC 29634-0317; Stacy L. Clark, Research Forester, U.S. Department of Agriculture, 

Forest Service, Southern Research Station, The University of Tennessee, Knoxville, TN 

37996-4563; and Bryan T. Mudder, Biological Science Technician, U.S. Department of 

Agriculture, Forest Service, Southern Research Station, Clemson University, Clemson, 

SC 29634.

The Authors:

March 2013

Southern Research Station

200 W.T. Weaver Blvd. 

Asheville, NC 28804

www.srs.fs.usda.gov

Cover Photograph:

Clockwise from top left: American chestnut; the spiny burs of the American chestnut; 

catkins in full bloom; an open bur, showing the chestnut inside. Photos reprinted with 

permission from the Southern Research Station Archives. 

The Woody Plant Seed Manual, originally published as USDA Miscellaneous publication 

654 in 1948, was twice revised, most recently in 2008 as Agriculture Handbook 727. 

These publications were the first attempt to summarize life histories, growth habits, and 

cultural practices to regenerate tree species in North America. The Woody Plant Seed 

Manuals included the genus Castanea, but provided only brief descriptions of growth 

habits, blight-resistance breeding, and regeneration practices. “Silvics of Forest Trees 

of the United States” (USDA Forest Service, Agriculture Handbook 271) was the first 

document to comprehensively summarize the silvics of tree species, and contained 

information on 127 species of trees. The handbook did not contain information for the 

genus Castanea. The second publication “Silvics of North America” (USDA Forest Service, 

Agriculture Handbook 654) also did not contain Castanea species. There has been great 

progress in development of blight-resistant American chestnut progeny, work that was 

initiated by the U.S. Department of Agriculture and continued by other organizations. 

Artificial regeneration technology, which will be a requirement for restoration of American 

chestnut, has also greatly improved, in part through the work of the Institute for Tree 

Root Biology of the Forest Service, U.S. Department of Agriculture. The Forest Service 

signed a memorandum of understanding with the American Chestnut Foundation that 

provides a framework for the Forest Service to assist the Foundation in research and 

provide lands for plantings of experimental material. Putative blight-resistant progeny 

was first planted on the national forests in 2009, and planting of chestnut material for 

reforestation was proposed in 2012 in an Environmental Assessment on the Nantahala 

National Forest. Guidelines for silvical requirements, regeneration strategies, and 

management techniques for restoration of this species are needed prior to large-scale 

planting on public and private lands. This publication is modeled after the “Silvics of 

North America” in its content with additional sections unique to American chestnut, such 

as the history of blight-resistance work and current restoration efforts. 

Preface:
G. Geoff Wang, Benjamin O. Knapp, Stacy L. Clark, and Bryan T. Mudder

The Silvics of Castanea dentata (Marsh.) Borkh., 

American Chestnut, Fagaceae (Beech Family)

ABSTRACT

This report describes how the American chestnut (Castanea 

dentata) was ecologically extirpated due to an exotic 

pathogen, the chestnut blight (Cryphonectria parasitica), 

and describes current restoration efforts. The habitat, 

life history, special uses, and genetics of the American 

chestnut are detailed. The American chestnut was an 

important and versatile tree species until its demise from 

the chestnut blight. Backcross breeding techniques, the 

use of hypovirulent blight strains, and genetic engineering 

programs are currently being developed and tested to 

produce trees resistant to the blight fungus. The first 

putative blight-resistant trees have been planted and 

silvicultural techniques to improve competitive ability 

of chestnut are being tested. American chestnut is a fastgrowing species with the ability to persist in shaded 

conditions and it responds favorably to forest management 

techniques that limit competition and increase available 

sunlight. Restoration will require advanced artificial 

regeneration techniques. The effects of damaging agents 

other than blight, including root rot caused by Phytophthora 

cinnamomi, Asiatic oak weevil (Cyrtepistomus castaneus), 

the gypsy moth (Lymantria dispar), and Asian ambrosia 

beetles (Coleoptera: Scolytidae) are not well understood but 

may represent real barriers to restoration.

Keywords: American chestnut, backcross breeding, 

chestnut blight, exotic species, restoration, silvics, 

silviculture. 

INTRODUCTION

American chestnut [Castanea dentata (Marsh.) Borkh.], also 

called sweet chestnut, was once a widespread and dominant 

species throughout the deciduous forests of eastern North 

America (Braun 1950, Russell 1987). American chestnut 

had the fastest growth and highest abundance in Braun’s 

(1950) oak-chestnut forest region, but the species inhabited 

nearly all States east of the Mississippi River (fig. 1). The 

introduction of exotic pathogens, primarily the chestnut 

blight fungus [Cryphonectria parasitica (Murr.) Barr], 

led to the extirpation of the species as a forest canopy 

dominant throughout its native range. Root rot caused by 

Phytophthora cinnamomi (Rands) also contributed to the 

demise of Castanea species growing in lowland or riparian 

areas (Russell 1987). The effects of these exotic pests 

resulted in the current listing of American chestnut as a 

species of special concern in Maine and Tennessee and as 

an endangered species in Kentucky and Michigan (USDA 

Plants Database 2012). The Committee on the Status of 

Endangered Wildlife in Canada listed American chestnut as 

threatened in 1987 and changed the status to endangered in 

2004 (Tindall and others 2004). 

American chestnut was one of the most valued and beloved 

eastern hardwood species due to its use as a timber tree, its 

abundant nut production, and its secondary wood products 

(Emerson 1846, Zeigler 1920). American chestnut grew 

straight and tall, reaching up to 5 feet in diameter and 

120 feet in height, and the species could live for several 

hundred years (Buttrick 1925). It was among the most 

versatile trees on the continent, historically used for 

construction lumber, shingles, fence posts and rails, 

poles, paneling, trim, furniture, and firewood (Ashe 1911, 

Detwiler 1915, Emerson 1846, Zeigler 1920). The wood 

was important in the charcoal iron furnace industry in 

the northern hardwoods region (Emerson 1846) and is 

highly resistant to decay because of its high tannin content. 

Extracted tannins from chestnut once drove the leather 

tanning industry in the Northeastern United States (Youngs 

2000). American chestnut produced copious amounts of 

flavorful nuts that were consumed by humans and wildlife 

(Frothingham 1912, Hawley and Hawes 1925, Hepting 

1974, Smith 2000, Zeigler 1920). 

At its greatest distribution, American chestnut constituted 

up to one-fourth to one-half of the canopy trees in its native 

range, and because of its desirable characteristics, it was 

widely planted in orchards and along streets, roadsides, 

and fencerows for shade, windbreaks, nut production, and 

aesthetics (Burnham 1988, Emerson 1846, Emerson and 

Weed 1908, Russell 1987). In the early 1900s, American 

chestnut made up more than 25 percent of all timber cut in 

the Southern Appalachians, was the most valuable tree in 

southern New England, and constituted almost half of the 

timber cut in Connecticut (Frothingham 1912, Hawley and 

Hawes 1925, Hepting 1974). The species was described as 

“one of the most promising trees for forest management” in 

Tennessee (Ashe 1911) and “one of the most promising trees 

for forestry planting” at the onset of professional forestry in 

the United States (Emerson and Weed 1908).
The demise of American chestnut followed the introduction 

of two exotic fungi and is regarded as the most tragic 

ecological event in the post-glacial history of eastern 

North American forests (Jacobs 2007). Root rot caused by 

Phytophthora cinnamomi was found to affect American 

chestnut as early as 1824, with the greatest impact on trees 

in riparian areas, heavy clay soils, and low-lying valleys 

(Anagnostakis 2002, Crandall and others 1945). The 

disease was initially most damaging in the Piedmont and 

Coastal Plain regions but was also noted in mountainous 

areas, moving from low to progressively higher elevations 

(Anagnostakis 2002, Crandall and others 1945, Russell 

1987). Unlike the chestnut blight, trees infected with 

Phytophthora root rot cannot maintain above-ground stems 

for long before death.

The chestnut blight is an aggressive diffuse canker disease 

that was introduced along with imported Castanea spp. 

seedlings from Asia (Anagnostakis 1987). The disease was 

first detected in 1904 at the Bronx Zoological Park in New 

York City (Merkel, 1905, Roane and others 1986); by 1950, 

the disease had spread throughout the range of American 

chestnut, and by 1960 it had killed an estimated 4 billion 

trees and essentially extirpated the species as a canopy 

tree (Anagnostakis 1987, Hepting 1974, McCormick and 

Platt 1980). However, American chestnut is still a common 

component of eastern deciduous forests, persisting as 

understory sprouts that originated from blight-killed trees 

and saplings (Russell 1987, Stephenson and others 1991). 

The cycle of sprouting, infection, and dieback continues, 

with sprouts generally not exceeding small tree size and 

rarely growing to reproductive maturity (Paillet 1984, 2002).

Recent advancements in the production of putatively blightresistant American chestnut trees indicate that restoration 

may be imminent (Clark and others 2011, 2012a). Because 

of the unique history of this species, relatively little is 

known about its ecology and life history. The objective 

of this technical report is to synthesize the current 

understanding of the silvics of American chestnut in order to 

facilitate current and future restoration efforts.

HABITAT 

Native Range 

American chestnut reached its maximum distribution in 

the United States about 2000 years before present, when 

the species established in New England forests for the 

first time (Davis 1983, Paillet 2002). By the late 19th 

century, the American chestnut range extended from central 

Mississippi and Alabama through the Appalachian region 

and into New Hampshire, Vermont, and Maine; the range 

extended west into Ohio and Indiana and into portions of 

southern Ontario, covering an area greater than 

309,000 square miles (Braun 1950, Hawley and Hawes 

1925, Russell 1987, Woods 1953) (fig. 1). Other reports 

suggest the possibility of a more extensive range, for 

example, Woods (1953) found chestnut far south in the Gulf 

Coastal Plain, occurring with longleaf pine (Pinus palustris

Mill.) in southern Alabama, Florida, and Louisiana.

According to Braun (1950), the most concentrated 

abundance of American chestnut in its native range falls 

within the Oak-Chestnut Forest and the Mixed Mesophytic 

Forest regions. In the Oak-Chestnut Forest region, American 

chestnut was often the dominant tree species, comprising 

an estimated 25 percent of hardwood forests (Burnham 

1988). In portions of the Appalachian Mountains, chestnut 

was thought to have made up 40 to 50 percent of the forest 

canopy (Braun 1950), and in the Southern Appalachian 

Mountains, it could occasionally form pure stands (Buttrick 

1915). Saucier (1973) reported that the species made up 15 

billion board feet or 25 percent of the total timber volume 

in the Southern Appalachian Mountains. Chestnut trees 

comprised 50 percent of the timber on the mountainous 

ridges of western Maryland, dominated 26 to 36 percent of 

the slope forests of Maryland, and were estimated to hold 

3.75 billion board feet of timber in the Blue Ridge 

Mountains of North Carolina (Buttrick 1915). Chestnutdominated stands formed 50 percent of second growth 

forests in New Jersey, Pennsylvania, Connecticut, and 

southern New England (Frothingham 1912, Hough 1878, 

Russell 1987). Most of these species composition estimates 

were made after the disturbances associated with European 

settlement, such as grazing, land-clearing for agriculture, 

uncontrolled wildfires, and indiscriminant logging (Abrams 

1992, 1996). Chestnut regeneration probably increased 

following repeated disturbance due to its ability to 

prolifically sprout and occupy abandoned agriculture fields 

(Matoon 1909, Paillet 2002); therefore, historical species 

composition estimates may overestimate chestnut abundance 

prior to European settlement. Nevertheless, chestnut was 

perhaps the most widespread and abundant species in the 

Eastern United States since the last glaciation, with the 

highest concentration of American chestnut in the Southern 

Appalachian Mountains.

Climate

In the native range of American chestnut, mean annual 

precipitation varies from about 32 inches in western 

New York and southern Ontario to about 80 inches in 

the Southern Appalachian Mountains; however, annual 

precipitation for the majority of the native range is between 

40 inches and 48 inches. Annual snowfall ranges from only 

trace amounts in the Southern United States to 100 inches 

2
or more in the Northern United States and in Canada. Mean 

annual temperature is about 40 °F in the northern part of the 

range and 60 °F in the extreme southern part. The frost-free 

period averages 100 days in the Northern United States and 

240 days in the Southern United States.

Although American chestnut is believed to be the most 

cold-hardy species of the Castanea genus (Huang 1998), 

it is prone to frost damage (Gurney and others 2011, 

Paillet 1988, Parker and others 1993, Zon 1904), which 

may have affected its northern range and upper altitudinal 

limits (Russell 1987). Zon (1904) noted a lack of American 

chestnut in the bottoms of ravines and valleys because 

frosts commonly occur on wet, protected soils. Braun 

(1950) reported American chestnut occurring at elevations 

of 1,300 to 4,500 feet in the Southern Appalachian 

Mountains, with the possibility of occurrence at higher 

elevations of 4,000 to 5,500 feet on southern exposures. In 

North Carolina, Buttrick (1925) found American chestnut 

growing between elevations of 500 feet and 5,500 feet. 

American chestnut grew at lower elevations in northern 

latitudes, occurring at elevations < 3,000 feet in the Catskill 

Mountains (Russell 1987) and at elevations < 400 feet in 

New Hampshire (Hough 1878). 

Only one study has examined climate-tree growth 

relationships of American chestnut (McEwan and others 

2006). The relationships between American chestnut growth 

and climate were similar to those of other hardwoods, with 

increasing growth associated with increasing precipitation 

and decreasing Palmer Drought Severity Index values 

during the growing season. However, American chestnut 

growth had strong positive relationships with August 

precipitation of the current year and strong negative 

relationships with August temperature of the previous year, 

suggesting it may be slightly more sensitive to late growing 

season climate than other hardwoods.

Soils and Topography

American chestnut naturally occurs on a wide range of 

soil and topographic conditions but is most commonly 

associated with well-drained, subxeric to mesic soils that 

often occur on sand plains and dry ridges (Abrams and 

Ruffner 1995, McEwan and others 2005, Paillet 2002, 

Russell 1987, Stephenson and others 1991). American 

chestnut is believed to grow poorly in very wet or very 

dry soils, and rarely grows in areas with high pH or on 

limestone-derived soils (Braun 1950; Frothingham 1912; 

3

Figure 1—Original native range of American chestnut in eastern North America, as adapted from Little (1977) by Jacobs (2007) 
Paillet 1988, 2002; Russell 1987). Within a soil pH range 

of 3.3 to 5.5, chestnuts were more abundant at the higher 

end of the range (pH of 4-5) in a study conducted in 

southwestern Virginia (Burke 2011). A recent survey of 

surviving trees in Canada found that American chestnut 

was most likely to occur in deciduous forest habitats with 

high canopy cover (> 50 percent), gentle slopes (0 to 10 

percent) and acidic (pH 4–6), sandy (> 75 percent) soils 

(Tindall and others 2004). In the southern portion of the 

range, the occurrence of American chestnut on northern 

side slopes increased with topographic relief, and the 

species was negatively associated with the more calcium 

rich soils and lower relief of the Coastal Plain region (Black 

and others 2002). The presence of American chestnut often 

indicates acidic soils derived from gneiss, metamorphosed 

sandstone, and sandstone that is deficient in lime and potash 

(Frothingham and others 1926). A recent study also reported 

that American chestnut height growth was negatively 

correlated with pH (Tindall and others 2004). 

Although the Society of American Foresters included 

American chestnut as a characteristic tree in the cove 

hardwood forest type (Frothingham and others 1926), it 

is not clear to what extent American chestnut occurred in 

cove or ravine sites. Frothingham (1912) and Braun (1950) 

indicate that chestnut was not commonly found on moist 

cove sites, but Ashe (1911) and Buttrick (1925) suggest the 

opposite. Ashe (1911) reports that American chestnut grew 

best in rich, deep coves in Tennessee, and Buttrick (1925) 

observed that American chestnut in cove forests produced 

the best timber. More recent evidence supports that 

American chestnut contributed substantially to the canopy 

of ravine and cove sites in the Cumberland Plateau and 

Southern Appalachian Mountains, e.g., more survivors were 

found in ravine sites than on ridge-top sites (Hinkle 1989), 

and chestnut potentially represented 25 to 40 percent of the 

basal area in pre-blight riparian stands (Vandermast and Van 

Lear 2002). Additionally, McCament and McCarthy (2005) 

observed that first-year seedling growth was negatively 

correlated with the proportion of sand in the soil, suggesting 

the superior suitability of finer textured soils in cove sites 

for chestnut growth. 

The dominance of American chestnut may have affected 

nutrient cycling and soil chemistry (Ellison and others 

2005), although a replicated study examining chestnut 

leaf litter effects on soil biogeochemical properties has not 

been conducted. One descriptive study found that most soil 

nutrients on sandy loam soils were higher under chestnut 

canopies compared to areas without chestnut (Rhoades 

2007). As soil texture became finer, this effect disappeared, 

suggesting that effects of American chestnut leaf litter on 

soil chemistry are dependent on soil texture. 

Soil nutrient requirements—American chestnut grows 

and remains competitive over a wide range of soil 

nutrient regimes. It was frequently found on relatively 

nutrient deficient to moderately fertile soils (Russell 

1987, Zon 1904). Although American chestnut tolerates 

relatively low soil nutrient levels, it responds well to 

increasing soil nutrients. Latham (1992) demonstrated 

improved seedling growth with increasing soil nutrient 

availability in a greenhouse study. Furthermore, positive 

growth and/or biomass response reported for American 

chestnut seedlings has been associated with increasing 

availability of magnesium (leaf mass and area, root mass), 

potassium (diameter, specific leaf area) (McCament and 

McCarthy 2005), and nitrogen (stem, root, leaf parameters) 

(McCament and McCarthy 2005, Rieske and others 2003). 

Soil moisture regime—American chestnut is a relatively 

drought tolerant species, as suggested by its historical 

dominance on upland sites with well-drained, sandy soils. 

During an early-season drought in a hardwood forest in 

Pennsylvania, American chestnut saplings maintained higher 

leaf water potential than several associated oak (Quercus) 

species that are known for their drought tolerance (Abrams 

and others 1990). Recently, Bauerle and others (2006) 

reported high water use efficiency of American chestnut 

seedlings exposed to drought compared to published reports 

of co-occurring hardwood species. However, American 

chestnut is reported to require more moisture than associated 

oaks, including black oak (Quercus velutina Lam.), scarlet 

oak (Quercus coccinea Münchh.), and chestnut oak 

(Quercus prinus L.) (Frothingham 1912). Alternatively, 

American chestnut growth was less related to drought 

during the growing season of the previous year compared 

to other hardwoods in the same stand (McEwan and others 

2006). The reported dominance of American chestnut 

in cove forest sites, as well as on well-drained upland 

sites, suggests that the species can compete across a wide 

range of soil moisture regimes (Ashe 1911, Buttrick 1925, 

Vandermast and Van Lear 2002). 

Associated Forest Cover

American chestnut is capable of forming pure stands, but 

historical reports suggest that pure stands were rare (Buttrick 

1915, Frothingham 1912, Zon 1904). In Braun’s (1950) 

classification of forest communities, American chestnut was 

the dominant tree species in the Oak-Chestnut Forest region 

(Frothingham and others 1926, Paillet 2002, Russell 1987). 

The Oak-Chestnut Forest region included five sections: 

(1) the Southern Appalachian section; (2) the Northern Blue 

Ridge section; (3) the Ridge and Valley section; (4) the 

Piedmont section; and (5) the Glaciated section.

4
Common associates of American chestnut in the Southern 

Appalachian Mountains included buckeyes (Aesculus spp.), 

basswood (Tilia americana L.), yellow poplar (Liriodendron 

tulipifera L.), eastern hemlock [Tsuga canadensis (L.) 

Carrière], white pine (Pinus strobus L.), pitch pine (Pinus 

rigida Mill.), table mountain pine (Pinus pungens Lamb.), 

northern red oak (Quercus rubra L.), white oak (Q. alba

L.), black oak, scarlet oak, and chestnut oak. The canopy 

trees were often associated with an ericaceous shrub layer 

consisting of Rhododendron spp., Azalea spp., Kalmia spp., 

Vaccinium spp., Leucothoe spp., Menziesia spp., and others 

depending on geographic region and elevation (Braun 1950, 

Russell 1987, Vandermast and Van Lear 2002). 

In the Glaciated section, American chestnut was commonly 

associated with red oak, scarlet oak, white oak, chestnut 

oak, pignut hickory (Carya glabra Mill. Sweet), American 

beech (Fagus grandifolia Ehrh.), and yellow poplar with 

an ericaceous understory (Braun 1950). In the Catskill 

Mountains of New York, American chestnut was observed 

in mixed deciduous forests along with American beech, 

northern red oak, butternut (Juglans cinerea L.), sugar 

maple (Acer saccharum Marsh.), and an understory of 

Rhododendron spp., Kalmia spp., and other ericaceous 

shrubs (Russell 1987). 

In the Northern Blue Ridge section, American chestnut was 

commonly associated with yellow poplar, northern red oak, 

white oak, chestnut oak, sugar maple, eastern hemlock, and 

basswood, with Rhododendron species, mountain laurel 

(Kalmia latifolia L.), and blueberry (Vaccinium spp.) as 

common understory species (Braun 1950). 

American chestnut associates in the Ridge and Valley 

section included northern red oak, scarlet oak, white oak, 

chestnut oak, white pine, eastern hemlock, and yellow 

poplar, with an ericaceous understory similar to that already 

mentioned (Braun 1950). 

In the Piedmont section, chestnut is being replaced by white 

oak, black oak, northern red oak, post oak (Quercus stellata

Wangenh.), blackjack oak (Quercus marilandica Münchh.), 

chestnut oak, yellow poplar, and American beech, with an 

understory of dogwoods (Cornus spp.) and hickories (Carya

spp.) mixed with the heath layer (Braun 1950). 

American chestnut was also one of the dominant trees in 

the Mixed Mesophytic Forest region that encompasses 

the Cumberland Mountains, Cumberland and Allegheny 

Plateau, and Allegheny Mountain sections. It was commonly 

associated with American beech, yellow poplar, basswood, 

sugar maple, yellow buckeye (Aesculus flava Aiton), 

northern red oak, white oak, and eastern hemlock (Braun 

1950). American chestnut was the most dominant species 

in the Allegheny Mountain section, where it was commonly 

associated with northern red oak, white oak, black oak, 

scarlet oak, yellow poplar, hickories, black gum (Nyssa 

sylvatica Marsh.), black birch (Betula lenta L.), basswood, 

sugar maple, and beech.

Associated forest cover of American chestnut is highly 

dependent on disturbance history. The composition of many 

forests once containing a major component of chestnut are 

succeeding from an oak-hickory dominated association 

(Keever 1953, McCormick and Platt 1980) to one dominated 

by maple or shade-intolerant species (Abrams 1998, Lorimer 

1984). In a recent survey of surviving American chestnut 

sprouts and trees in the Coastal Plain, Highland Rim, and 

Cumberland Plateau regions of Kentucky and Tennessee, 66 

percent of the sampled American chestnut trees 

(n > 2,000) occurred within 49 feet of red maple trees 

(Schibig and others 2005). Oak species were recorded in 

< 30 percent of plots that surrounded American chestnut. 

Red maple (Acer rubrum, L.) was historically not a major 

component of these forest ecosystems (Braun 1950), but 

it has become more common with changes in disturbance 

dynamics (Abrams 1998, Nowacki and Abrams 2008). 

LIFE HISTORY

Reproduction and Early Growth

Flowering and fruiting—American chestnuts are 

monoecious, with male and female reproductive parts 

on the same tree (Brown and Kirkman 1990, Zon 1904). 

Flowers are staminate catkins that are borne on a central 

axis 5 to 10 inches long and are produced in clusters 

from the axils of leaves. Flowers appear from late May to 

June, after frosts have passed (Brown and Kirkman 1990; 

Zon 1904). Weak lower lateral shoots consist of all male 

flowers, while stronger upper vertical shoots have male 

flowers on upper shoots and female flowers on lower shoots. 

American chestnut pollen is not as abundantly dispersed as 

that of many other wind-pollinated tree species and often 

requires another tree within 328 feet to reproduce (Paillet 

2002, Russell 1987). Pollen is not dispersed until after full 

leaf development has occurred, and only pollen escaping 

above the forest canopy can be dispersed for long distances 

(Russell 1987). The fruit of American chestnut, maturing 

between September and October, is a brown edible nut with 

a sweet taste (Brown and Kirkman 1990), and the species 

was sometimes referred to as sweet chestnut due to its 

flavorful nut (Van Fleet 1914). The nut develops within a 

round prickly green bur that produces two to three nuts in 

each bur (Saucier 1973, Zon 1904). 

5
Seed production and dissemination—American chestnut 

is a prolific seed producer and one of the most fruitful 

among all American nut-producing species. It is able to 

produce fruit as early as age 4 when open-grown, or at about 

8 to 20 years old when competing with other trees in the 

forest (Paillet and Rutter 1989). Reproduction starts earlier 

and yields more nuts in trees reproduced by coppice than in 

trees regenerated from seed. A mature tree can produce 

1.5 to 3 bushels or up to 6,000 nuts per year (Paillet and 

Rutter 1989), and unlike other nut producers that display 

variable masting behavior, such as white oak, American 

chestnut is a dependable seed producer every year. The 

dependable seed production is probably due to its late 

flowering that avoids frost damage. The fruit typically drops 

with the first frosts of autumn, which opens the bur, and the 

seeds germinate the following spring (Zon 1904). 

Chestnuts were historically an important food for wildlife, to 

the point that seed predation combined with insect damage 

limited the success of reproduction from seed (Detwiller 

1915, Hawley and Hawes 1925). Chestnut provided a more 

stable and more abundant source of mast for wildlife than 

oaks, hickory, and beech species (Diamond and others 

2000). Additionally, the nuts were extensively harvested by 

humans as a source of food and income (Hawley and Hawes 

1925, Hepting 1974, Saucier 1973). As a result of the high 

demand for chestnuts by wildlife and humans, Paillet and 

Rutter (1989) report that only 1 to 5 viable seeds germinated 

into seedlings that survived for more than 1 year. When 

seeds were not consumed by animals or damaged by insects, 

chestnuts generally had high germinate success, with 60 to 

70 percent germination observed in the field (McCament 

and McCarthy 2005) and > 90 percent germination in the 

greenhouse (Wang and others 2006) and in commercial tree 

nurseries (Clark and others 2012b.).

Like many other large-seeded trees, American chestnut 

seeds were dispersed by wildlife species that included birds 

and squirrels (Sciurus spp.). The American crow (Corvus 

brachyrhynchos), blue jay (Cyanocitta cristata), wild turkey 

(Meleagus gallaparo silvestris), ruffed grouse (Bonasa 

umbellus) and passenger pigeon (Ectopistes migratorius) 

were major dispersers of large fruit and probably important 

for chestnut dispersal (Russell 1987, Webb 1986). Large 

mammals such as white-tailed deer (Odocoileus virginianus) 

or black bear (Ursus americanus) may have also played 

an important role in the dispersal of American chestnut, 

as the bur is thought to be an evolutionary adaptation for 

hitchhiking on mammalian fur.

Seedling development—Natural regeneration from seeds 

was probably rare in pre-blight years. The nuts were 

damaged by insects (Detlefson and Ruth 1922, Russell 

1987) and highly utilized as a wildlife food source (Detwiler 

1915). Small seedlings were easily killed by fire or frosts 

(Ashe 1911). 

Artificial regeneration of nursery-grown seedlings will be 

a crucial component of restoration. Once sufficient lines 

of blight-resistant seedlings are developed and tested, 

they can be planted, cross-pollinate with one another, and 

sexually reproduce. However, restoration of this species 

will probably always involve planting, due to the loss of 

seedlings from nut predation and high seedling mortality.

Seedlings grow rapidly in the greenhouse or in the field 

as nursery-grown bare-root seedlings or direct-seeded 

regeneration. Artificially regenerated chestnuts grow best 

under moderate- to high-light conditions but are capable of 

surviving under low-light conditions. Seedlings averaged 

3 to 4 feet in height and 0.5 to 0.6 inches in diameter at 

the root collar after 1 year in a commercial tree nursery 

after receiving steady applications of fertilizer throughout 

the growing season (Clark and others 2009, 2011, 2012b). 

Large, nursery-grown, seedlings generally outperform 

smaller seedlings due to the ability of the larger seedlings to 

reach heights above browse and natural woody competitors 

(Clark and others 2011). After 1 year in the field, nurserygrown seedlings that were not browsed by deer averaged 7 

inches in height growth and had 80-93 percent survival if 

not affected by root rot caused by Phytophthora cinnamomi

(Clark and others 2012a). Larger nursery seedlings appear to 

allocate more growth to the root system after being fieldplanted than do smaller nursery seedlings and, consequently, 

are better suited to resist transplant shock (Clark and others 

2009, 2011). Three growing seasons after planting nurserygrown seedlings in the field, seedlings averaged 0.91 inches 

and 0.94 inches in root collar diameter and 63 inches 

and 91 inches in height, respectively, when grown in 100 

percent and 34 percent full sunlight (Anagnostakis 2007). 

Early results indicate that trees bred for blight-resistance 

are similar to American chestnut but differ from Chinese 

chestnut with respect to height growth rates (Clark and 

others 2011, 2012a).

Direct-seeding American chestnuts may provide an 

alternative to out-planting nursery-grown seedlings on 

certain sites but will require intensive management to avoid 

predation by mammals and to maintain the competitive 

status of the seedlings. Two-year survival from direct 

seeding ranged from 40 to 50 percent in a field study in 

Ohio (McCament and McCarthy 2005). After 2 years of 

growth under partial forest shade (~ 25 percent open sky), 

direct-seeded seedlings averaged ~ 0.23 inches in root 

collar diameter and ~ 14 inches in height (McCament and 

McCarthy 2005). Under full sunlight, seedlings grown 

from seed in the greenhouse averaged ~ 0.20 inches in root 

collar diameter and ~ 12 inches in height after one growing 

6
season; seedlings allocated < 30 percent of the total carbon 

gain to below-ground biomass, compared to > 65 percent 

allocation to below-ground biomass for white oak (Wang 

and others 2006). Latham (1992) reported that first-year 

American chestnut seedlings grew faster in height, leaf area, 

and total dry mass under different levels of nutrients when 

compared to co-occurring species, including mockernut 

hickory [Carya alba (L.) Nutt], northern red oak, American 

beech, blackgum (Nyssa sylvatica) and tulip poplar. 

However, root-to-shoot ratios were among the lowest for 

American chestnut (Latham 1992), further indicating that 

early growth is allocated to above-ground biomass rather 

than root systems. 

Vegetative reproduction—American chestnut is a 

tenacious sprouter, and its ability to repeatedly produce 

large numbers of fast-growing sprouts following dieback 

is largely responsible for its persistence in forests today 

(Hawley and Hawes 1925). Natural sprouts will probably 

exist in the forest for years to come, but American chestnut 

will never again be a significant component of the forest 

canopy without the introduction of blight-resistant seedlings. 

Sprouts that live long enough to bear fruit are valuable 

because they can be used to maintain genetic diversity in 

breeding programs (Pierson and others 2007) and for in situ 

tests of blight resistance and hypovirulence (Anagnostakis 

2001, Griffin and others 1983). 

Frothingham (1912) attributes the “prodigious sprouting 

capacity” and rapid growth of American chestnut as 

reasons for the species’ great abundance in second-growth 

Connecticut forests prior to the blight. Prolific sprouting 

is considered an adaptation for long-term survival in the 

forest understory (Paillet 1984, 2002). Sprouts of American 

chestnut arise from dormant buds that may exist for several 

years before developing into a shoot or from adventitious 

buds that develop following localized stimulation (Mattoon 

1909, Paillet 1984). Chestnut rarely develops stool sprouts 

because the narrow region of sprouting is generally in 

contact with the ground or just above it. However, lowcut stumps tend to have sprouts of deeper origin and show 

a tendency to develop new lateral roots (Mattoon 1909). 

In a study by Zon (1904), chestnut was found sprouting 

principally from the root collar, and the best sprouts were 

found on low stumps. 

Although American chestnut is reportedly able to produce 

sprouts from trees up to 170 years old, the optimum age 

for sprouting is probably < 75 years (Mattoon 1909, Zon 

1904). Mattoon (1909) observed that old stumps produce a 

full thicket of short, spindling sprouts, while young stumps 

generate a smaller number of tall, stout sprouts. Paillet 

(1984) characterized the growth form of sprouting American 

chestnuts in the northeast and found it to be highly variable. 

Sprouts developed a shrub-like form when grown in the 

shade but expressed strong apical dominance and rapid 

growth when released, suggesting a growth strategy well 

suited to periodic disturbance and release from the canopy. 

The most vigorous sprouts are produced at periods of 

maximum growth in height, which occurs within the first 

decade in coppice trees (Zon 1904) or 20-30 years in stands 

regenerated from seedlings (Mattoon 1909). 

Coppice trees grow faster than trees regenerated from 

seed during the first 20 years. Because of their fast initial 

growth, sprouts reach their maximum average height growth 

rate during the first decade of growth. Mattoon (1909) 

observed rapid height growth (4 to 7 feet per year) in the 

first year but then a sharp decrease (2 to 3 feet per year) 

in the second and third year. Diameter growth continues 

well after height growth has decreased (Zon 1904). An 

increase of nearly 1 inch per year in diameter during the 

first 8-15 years has been observed under the most favorable 

conditions (Mattoon 1909). 

Dormant season cutting favors the production and growth 

of sprouts. Mattoon (1909) reportes that midwinter 

cutting produced sprouts of superior first-year growth 

when compared to May cutting: 6.2 feet versus 3.5 feet 

in height and 0.42 inches versus 0.23 inches in diameter, 

respectively. Early recommendations for cutting include: 

(1) late fall to early spring cutting to avoid production of 

weak sprouts too tender to stand frost; (2) cutting stumps 

low and at a slight angle to allow water runoff; and (3) 

avoiding damage to bark which may in turn damage buds 

(Frothingham 1912, Zon 1904). 

Chestnut sprouts remain an important component of 

understory vegetation throughout the species’ former 

range (Paillet 1984). However, Paillet (1984, 1988, 

2002) concluded that a majority of these sprouts were not 

associated with canopy trees killed by the blight but instead 

probably originated from former seedlings that had been 

through several cycles of fungal infection and resprouting or 

were from sprouts developed from canopy trees cut before 

the blight. Griffin and others (1991) observed that sprouts 

generally did not exceed 2 inches in diameter at breast 

height (d.b.h.) 10 years after sprouting in a clearcut before 

dying back from blight. Paillet (1984) alludes to sprouting 

as a reproductive strategy for chestnuts awaiting a crown 

opening because sprouts are capable of rapid transformation 

from small, suppressed stems to straight, vigorous saplings 

in a short time. Suppressed chestnut stems that have escaped 

blight for several decades develop a form with a single 

stem or a single stem with at least one weak secondary stem 

(Paillet 2002). 

7
Sapling and Pole Stages to Maturity

Growth and yield—Early studies indicate that American 

chestnut is highly competitive and fast-growing during 

early growth (Ashe 1911, Graves 1905, Zon 1904). These 

historical observations have been confirmed in recent 

studies. Jacobs and Severeid (2004) reported that juvenile 

plantation growth of American chestnut on blight-free sites 

in Wisconsin greatly exceeded that of interplanted black 

walnut (Juglans nigra L.) or northern red oak. In a recent 

study (Jacobs and others 2009), American chestnut averaged 

3.5 inches in d.b.h. and 28.5 feet in height 8 years after 

planting and 10 inches d.b.h. and 45 feet in height after 

19 years. Paillet and Rutter (1989) described the ability of 

introduced American chestnut to rapidly out-compete and 

eventually replace native tree species (e.g., Quercus spp., 

Carya spp.) as the dominant canopy tree in a Wisconsin 

forest. Studying American chestnut development in this 

same forest, McEwan and others (2006) documented that 

the growth of American chestnut trees released from the 

forest canopy following a logging event exceeded that of 

associated hardwoods by nearly a factor of two. 

The timber yield of American chestnut varied greatly 

depending on the dominance of American chestnut in 

the stand. Buttrick (1925) estimated that pure stands of 

American chestnut could yield as high as 20,000 board feet 

per acre and yielded approximately 20 to 30 billion board 

feet in the United States (Buttrick 1915). In mixed stands, 

Buttrick (1925) reported yield at an average of 4,000 board 

feet per acre with an estimated maximum of 10,000 board 

feet per acre for cove forests. The yield on slopes was 

estimated at 2,000 to 3,000 board feet per acre and that on 

ridges was reported at 1,500 board feet per acre. A study in 

Tennessee reported average annual yield of approximately 

500 board feet per acre per year over a 60-year rotation 

(Holmes 1925). In a recent study, Jacobs and others (2009) 

reported that biomass accumulation reached almost 80 tons 

per acre in a 19-year-old plantation.

Rooting habit—Little is known about the rooting habit 

of American chestnut, and inconsistent descriptions were 

found in the literature. Some describe American chestnut 

as having a shallow root system (Buttrick and Holmes 1913, 

Paillet 2002), yet others report that American chestnut is 

a deep-rooted species with a tap root similar to oak and 

lower lateral roots that spread up to 3.3 feet deep in the soil 

(Smith 2000, Zon 1904). Chestnut tap roots may divide into 

many vertical roots that extend 3 to 6 feet, each of which 

may develop many lateral roots (Smith 2000). Studies of 

American chestnut growing in the nursery indicate that 

chestnut seedlings have a main tap root with many lateral 

roots, similar to that of oak species (Clark and others 2009), 

and the number of lateral roots is positively correlated to 

both height and root collar diameter (Clark and others 2009, 

2012b).

Reaction to competition—Although American chestnut has 

been shown to outgrow competing hardwood species, the 

presence of abundant competition reduces chestnut growth 

(Griffin and others 1991). Insight into the competitive nature 

of American chestnut can be traced back to historical pollen 

records. Range expansion of American chestnut during the 

Holocene from glacial refugia was the most recent of windpollinated trees (Paillet 1982, 2002, Russell 1987), and its 

rapid expansion to canopy dominance implicates American 

chestnut as an exceptionally competitive species. 

The shade tolerance of American chestnut is still under 

debate (Joesting and others 2009, Wang and others 2006). 

Early observations suggest that American chestnut is 

relatively intolerant (Frothingham 1912, Hawley and Hawes 

1925) to moderately tolerant (Zon 1904) of shade. Recent 

studies have classified American chestnut as either shade 

tolerant (Wang and others 2006) or intermediately shade 

tolerant (Joesting and others 2009). In the first study to use 

contemporary instrumentation on American chestnut, Wang 

and others (2006) suggested that American chestnut is shade 

tolerant based on physiological characteristics that include 

low light compensation and saturation points measured in a 

greenhouse study, and these results were later confirmed in 

field studies (Joesting and others 2007, 2009). 

The shade tolerance of American chestnut is supported by 

other studies on morphology, survival, and growth under 

canopy shade. Paillet (1982) found that chestnut can survive 

in deep shade under the canopy for up to three decades and 

is more shade tolerant than co-occurring sub-canopy species. 

When growing in a light-limited environment, American 

chestnut increases its specific leaf area (Joesting and others 

2009, King 2003, McCament and McCarthy 2005, Wang 

and others 2006) and develops canopy architecture that is 

optimal for harvesting light (Paillet 1982). Anagnostakis 

(2007) found that seedlings growing in the field under 

63 percent shade cloth had more above-ground mass than 

those growing in full sunlight after three growing seasons. 

However, the shade tolerance of artificially regenerated 

seedlings may be different than those growing as sprouts. 

Seedlings planted as bare-root nursery stock may not be able 

to withstand shade due to the relatively small carbohydrate 

reserves stored in the newly developed root system, 

compared to a native sprout with an older, well-established 

root system.

Joesting and others (2009) reported that natural chestnut 

seedlings and saplings can subsist in the understory for 

8
many years, have a low dark respiration rate, high quantum 

efficiency, low N percent, low leaf mass per area, and 

high light-induced morphological plasticity, all of which 

are characteristic of a shade tolerant species. However, 

Joesting and others (2009) classified the shade tolerance of 

American chestnut as intermediate based on their finding 

that American chestnut has a maximum photosynthesis 

rate comparable to shade intolerant species. To justify 

their classification, Joesting and others (2009) also cited 

previous observations that American chestnut can rapidly 

assume a canopy position following the creation of canopy 

openings (Jacobs and Severeid 2004, McEwan and others 

2006) and that seedlings respond well to increasing light 

both in greenhouse (Latham 1992, Wang and others 2006) 

and field conditions (Boring and others 1981, Griffin 1989, 

McCament and McCarthy 2005, Tindall and others 2004). 

However, we argue that the ability to fill gaps in the canopy 

does not confer shade intolerance to a tree species. In fact, 

this strategy is commonly used by many shade tolerant 

species, e.g., red maple. 

Considering all the published evidence, we maintain that 

American chestnut should be classified as shade-tolerant. 

Indeed, the continuous survival of chestnut sprouts from 

blight-killed advanced regeneration in former American 

chestnut forests (Paillet 2002) is the best testament 

of its shade tolerance. Its strong ability to survive for 

prolonged periods in deep canopy shade (Paillet 1982), 

coupled with its ability to grow well under partial shade 

(Anagnostakis 2007), suggests that chestnut maintains its 

presence in the understory in anticipation of disturbance 

events (Jacobs 2007), with a strategy similar to that of 

other Fagaceae species, e.g., oaks. The rapid height growth 

of American chestnut following release or after planting 

in full sun (Billo 1998; Clark and others 2009, 2012a; 

McEwan and others 2006; Paillet 1982, 2002; Paillet and 

Rutter 1989) distinguishes American chestnut from cooccurring intermediately shade tolerant species such as oaks 

(Paillet 2002). 

Competition can also affect blight development on American 

chestnut seedlings. Field tests indicate that blight may be 

more common on sprouts growing in open or clear-cut sites 

compared to sprouts growing in the understory of mature 

forests (Griffin 1989, Griffin and Elkins 1986, Reynolds 

and Burke 2011). However, after blight has infected the 

site, surviving trees appear to better withstand the blight 

if released from competition. The causal factor may be 

related to hypovirulence, which may be more prevalent if 

competition is controlled and chestnut stems are allowed to 

take a dominant canopy position (Griffin 1989). Therefore, 

mesic sites may offer the best opportunity for biocontrol of 

the fungus through hypovirulence (Griffin 1992), but such 

sites will also have the most need for competition control. 

Allelopathy—Leachate from American chestnut 

litter may have allelopathic properties that limit the 

development of some common competitors in the southern 

Appalachians, including eastern hemlock and rhododendron 

(Rhododendron maximum L.). Good (1968) found that 

chestnut leaf extracts (1.8 ounce leaf per 1 fluid ounce 

distilled water) inhibited the germination of eastern 

hemlock and significantly reduced shoot length of black 

birch, yellow birch (Betula allegheniensis Britton), tomato 

(Lycopersicon esculentum Mill.), and eastern hemlock in 

a laboratory bioassay; he also found that adding chopped 

chestnut leaf material (5 g dry weight) to germination pots 

significantly reduced root length of tomato and root, stem, 

and leaf biomass of black birch. Vandermast and others 

(2002) found that chestnut leaf extracts (0.04 ounces per 

0.7 fluid ounces of distilled water) significantly reduced seed 

germination of lettuce (Lactuca sativa L.), rhododendron, 

and eastern hemlock and also reduced the length of radicles 

of germinating lettuce and rhododendron. Although these 

studies clearly demonstrate the allelopathic potential of 

American chestnut, the nature of these studies prevents 

extrapolation of the results to field conditions.

Damaging agents—American chestnut might be the most 

susceptible tree in the Eastern United States to damaging 

agents. The most widespread damaging agent to American 

chestnut is the chestnut blight. The fungus was originally 

believed to be a new species and was named Diaportha 

parasitica, but it was discovered that it likely came into 

the United States on imported Asian Castanea species. 

The fungus was renamed Endothia parasitica, and then 

Cryphonectria parasitica, and it is most commonly 

referred to as the chestnut blight fungus (Anagnostakis 

1987, 1992). 

Common symptoms of chestnut blight include the presence 

of mycelium arranged in buff-colored fans on the inner 

bark, sunken or raised cankers on the infected tree, the 

appearance of stromata (orange fruiting bodies) on the 

smooth bark surrounding cankers, the development of 

sprouts below cankers, and small, dying leaves and nuts 

on affected stems (Anagnostakis 1987, Gravatt 1925, 

Hawley and Hawes 1925). Disease symptoms also include 

yellowing or wilting leaves that tend to persist on the tree 

past leaf abscission in autumn (Griffin and Elkins 1986). 

Cankers are usually found at the base of the tree and 

usually have an elliptical shape, extending up and down 

the trunk. Old trees susceptible to blight typically produce 

sunken cankers, while more vigorous or blight resistant 

trees produce slightly swollen cankers (Gravatt 1925, 

Griffin 2000, Griffin and Elkins 1986). The bark over the 

injury is typically darker red in color than the surrounding 

bark, and exposed wood is often at the canker center 

(Griffin and Elkins 1986). 

9
As an ascomycete, the fungus produces small reddishbrown fruiting bodies on the surface of cankers where two 

different types of spores are formed (Gravatt 1925). Shortlived conidia are summer spores exuded in sticky masses. 

After summer rains these spores stick to the feet of birds, 

insects, and mammals and are carried long distances to 

infect other trees (Burnham 1988, Hepting 1974). Ascopores 

are winter spores that are carried long distances by wind 

to infect other trees by entering at a wound or split in 

the bark, where they germinate and enter the inner bark, 

killing vital cells as they invade (Burnham 1988, Hepting 

1974). American chestnut trees commonly have wounds 

and punctures caused by insects, woodpeckers, and natural 

bark cracks (Ashe 1911). The fungus enters through these 

bark wounds, where it grows and has a girdling effect on 

the cambium of the tree. Although the blight has effective 

dispersal mechanisms, humans helped the spread by 

shipping diseased nursery stock, carrying the fungus on 

chestnut wood, and transporting spores on clothing, shoes, 

and tools (Hepting 1974).

Although the chestnut blight has certainly had the greatest 

impact on American chestnut populations across the native 

range, the species is also susceptible to other damaging 

agents (Jacobs 2007). Records of dying chestnuts caused by 

the root rot fungus Phytophthora cinnamomi can be traced 

back to 1824 (Crandall and others 1945). Woods (1953) 

reported that an epidemic of P. cinnamomi in the southern 

portion of the United States nearly eliminated chestnut from 

that part of its range from 1825–75. Crandall and others 

(1945) attributed P. cinnamomi to the disappearance of 

chestnut from the Gulf and Atlantic States as well as the 

foothills and mountains of Mississippi, Alabama, Georgia, 

Tennessee, Maryland, Virginia, North Carolina, and South 

Carolina. Ashe (1911) reported declining populations of 

chestnut throughout lower elevations of the Appalachians 

in Tennessee, and although not recognized at the time, the 

probable cause was P. cinnamomi. 

This pathogen spreads primarily through free-moving 

soil water as zoospores that colonize fine-feeder roots; 

the fungus then can move into healthy cells of the main 

tap root and into the above-ground stem, forming lesions 

above the root collar (Crandall and others 1945, Robin and 

others 1992). Common symptoms of P. cinnamomi include 

sudden yellowing or wilt of the leaves during the growing 

season, followed by defoliation and death. The pathogen 

can also cause a gradual reduction in the size of the leaves 

over several years (Crandall and others 1945). Lesions of 

varying size with an ink-like exudate appear on the roots 

and root collar cankers, leading to the name “ink disease.” 

The disease also affects red oaks, although it usually does 

not cause death (Robin and others 1992); however, it has 

been determined to cause the death of white oak and has 

been found in pine stands as well (Balci and others 2007, 

Campbell and Hendrix 1967). 

Phytophthora cinnamomi is found in many forest types 

throughout the Southeastern United States (Campbell and 

Hendrix 1967, Hendrix and Campbell 1970, McLaughlin 

and others 2009) but is not tolerant of cold environments 

with annual minimum temperatures less than -20 °C (Balci 

and others 2007, Benson 2002). It is generally not found 

north of 40° latitude (Balci and others 2007) or in higher 

elevations at lower latitudes. In an experimental study with 

American chestnut seedlings, Rhoades and others (2003) 

found that the occurrence of P. cinnamomi infection was 

greater in wet soil than in dry soil, but around 25 percent of 

seedlings became infected regardless of soil moisture. Many 

observers noted the local decline and death of large numbers 

of American chestnut trees adjacent to one another, and such 

epidemics often began at poorly drained sites with heavy 

soils and then spread to higher, drier sites (Crandall and 

others 1945). However, susceptible trees on well-drained 

soils have also succumbed to the disease if the soil becomes 

infected (Balci and others 2007, Clark and others 2009, 

2012a). Clark and others (2009, 2012a)1 attributed failure in 

the first-year survival of pure American chestnut plantings 

in the Southeastern United States to P. cinnamomi that was 

probably brought in from nursery stock.

Phytophthora cinnamomi represents a large threat to 

restoration of American chestnut in the areas with climate 

and soil conditions favorable to the disease, including the 

Southern Piedmont, the Blue Ridge, and some areas within 

Tennessee and Kentucky. There currently exists no effective 

control of the disease in the nursery, in the field, or through 

breeding programs. Screening by the American Chestnut 

Foundation and the Connecticut Agriculture Experiment 

Station for resistance to the disease is in the beginning 

stages and shows promise (Anagnostakis 2002, James 

2011, Jeffers and others 2009). The genes for resistance 

appear to be incompletely dominant but are different from 

genes controlling resistance in blight (Anagnostakis 2001). 

Therefore, it could be many years before trees resistant to 

both fungal pathogens are produced. 

The Asian chestnut gall wasp (Dryocosmus kuriphilus) 

is a recently introduced pest that also poses a threat to 

American chestnut (Anagnostakis and others 2011, Jacobs 

10

1 Personal communication. 2011. Griffin, G.J. Professor Emeritus, Virginia Polytechnic and State University; and President, American Chestnut 

Cooperators Foundation, 2667 Forest Service Road 708, Newport, VA 24128.
2007). Introduced in Georgia in 1975, the species has 

quickly spread through the range of American chestnut 

and is now found throughout Tennessee and as far north 

as Cleveland, OH. Both American chestnut and Chinese 

chestnut are susceptible to Asian chestnut gall wasp, making 

it unclear how this species may affect chestnut hybrids 

developed for restoration (Anagnostakis and others 2011). 

The insect has been found on blight resistant hybrids 

planted in the field, but effects on seedling growth have not 

been yet determined.2

Other potentially harmful exotic pests include the Asiatic 

oak weevil (Cyrtepistomus castaneus Roelofs Coleoptera: 

Curculionidae), the gypsy moth (Lymantria dispar L. 

Lepidoptera: Lymnantriidae), and the Asian ambrosia 

beetles (Coleoptera: Scolytidae). The Asiatic oak weevil 

is an exotic pest from Japan that prefers to feed on species 

in the Fagaceae family, particularly oaks (Ferguson and 

others 1991, Frederick and Gering 2006). This species 

is especially damaging because larvae feed on roots and 

emerging radicles, while adults feed on leaves (Roling 1979, 

Triplehorn 1955). The Asiatic oak weevil was the most 

abundant insect caught in traps in a Missouri Ozark forest 

(Linit 1986) and is probably contributing to oak decline 

in those forests. The Asiatic oak weevil was found to be 

more prevalent on chestnut trees than oak trees, where it 

was found to completely defoliate young chestnut seedlings 

(Johnson 1956). 

The gypsy moth is an exotic pest that was introduced 

from Europe, and the first gypsy moth outbreaks began 

around the same time as the chestnut blight fungus. The 

species has devastated vast areas of oak-dominated forests 

in the Appalachian Mountains in Virginia and northward 

through eastern deciduous forests. The gypsy moth could 

be a potential problem in defoliation of blight-resistant 

chestnuts, as it has been observed feeding aggressively on 

native chestnuts since 1915 (Mosher 1915). The gypsy moth 

was found to grow larger when fed leaves from second 

generation backcross chestnut hybrids when compared to 

leaves from pure American seedlings (Rieske and others 

2003); similarly, gypsy moth grew better on transgenic 

chestnuts when compared to native American chestnuts 

(Post and Parry 2011). The preference of the moth for the 

BC3F3 generation of chestnut hybrid is currently unknown. 

The Asian ambrosia beetles were first imported into eastern 

North America in the 1930s, and new species have recently 

been discovered (Atkinson and others 1990, Schiefer and 

Bright 2004). The insect bores into small saplings and trees, 

creating tunnels and galleries and potentially introducing 

fungi into the wounds. Xylosandrus crassiusculus

Motschulsky has been found to negatively impact Chinese 

chestnuts in orchards in middle Tennessee (Oliver and 

Mannion 2001), but there is little information on effects 

these beetles will have on American chestnut seedlings, 

hybrid seedlings, or transgenic trees. Mortality caused 

by this insect was observed on the research plots of the 

American Chestnut Cooperators Foundation, typically on 

American chestnut stem < 4 in diameter.3 Another recently 

imported Asian ambrosia beetle (Xylosandrus mutilatus

Blandford) is known to be a pest to Chinese chestnut in 

its native China, but this beetle is currently south of the 

American chestnut species range (Scheifer and Bright 2004, 

Six and others 2009).

Two native insects that affect chestnut include the chestnut 

sawfly [Craesus castaneae Marshall Hymenoptera: 

Tenthrediniae] and the twolined chestnut borer (Agrilus 

bilineatus Weber. Coleoptera: Buprestidae). The chestnut 

sawfly feeds on leaves, and the only paper that has reported 

sawfly on chestnut showed that it was more damaging to 

trees planted in shaded environments than those planted in 

open conditions. In addition, the sawfly preferred hybrid 

(BC2F3) and American chestnut seedlings over Chinese 

chestnut seedlings (Pinchot and others 2011). The chestnut 

sawfly is a rare species and the impact of defoliation on 

chestnut restoration is unclear. 

Historically, the American chestnut was the primary host 

of the twolined chestnut borer, but red oaks have become 

the primary host since the demise of chestnut (Haack and 

Acciavatti 1992). The insect is known to kill trees that 

are already stressed by first attacking the crown and then 

moving down into the trunk until the tree dies after 

2–3 years (Cote and Allen 1980). The chestnut borer can 

invade after gypsy moth infestations, but populations can be 

reduced by removing dying trees and trees with poor vigor 

(Muzika and others 2000). No information exists on the 

impact this species will have on chestnut restoration, but the 

insect will likely only attack mature trees of low vigor.

11

2 Clark, S.L. 2012. The first research plantings of third-generation, third-backcross American chestnut (Castanea dentata) in the Southeastern 

United States. Manuscript in preparation. Proceedings from the 5th International Chestnut Symposium, Sheperdstown, WV, September 5-8, 

2012. Author can be reached at 2431 Joe Johnson Drive, Room 274, Knoxville, TN 37996-4563.

3 Griffin, S.L. 2011. Observations on ambrosia beetle damage. Unpublished data. On file with: American Chestnut Cooperators Foundation, 2667 

Forest Service Road 708, Newport, VA 24128.
The effects of fire on American chestnut are not well 

understood. Some early observations report that chestnut 

is negatively affected by fire because of its thin bark and 

shallow root system, suggesting that anthropogenic fire 

in the Appalachian oak-chestnut forest may have limited 

natural chestnut regeneration (Ashe 1911, Buttrick and 

Holmes 1913, Paillet 2002, Russell 1987). Frothingham 

(1912) found that American chestnut was mostly absent 

from frequently burned forests of pitch pine and scrub 

oak. Fire damage to American chestnut bark may increase 

susceptibility to disease and insect infestation (Hawley 

and Hawes 1925, Russell 1987). However, the vigorous 

sprouting of American chestnut suggests that it may 

be able to persist following infrequent burning. Paillet 

(2002) noted evidence of fire preceding sharp increases in 

the proportion of American chestnut pollen in sediment. 

McEwan and others (2011) related patterns of fire frequency 

with changing forest composition from oak-chestnut forests 

to forests dominated by maple. To our knowledge, there 

exists only one experimental fire study using pure American 

chestnut seedlings to determine effects of fire on seedling 

sprouting and growth response, but this study is still in 

the early stages.4 The complex interactions between site 

conditions, fire intensity, fire frequency, and fire season 

make it difficult to predict how American chestnut will 

respond to fire without additional research. However, the 

contrasting pattern of below- and above-ground biomass 

allocation between chestnut and oaks/hickories (Wang and 

others 2006) suggests that American chestnut may not be 

adapted to fire as well as oaks and hickories. 

Extreme weather events, including excessive heating, low 

winter temperature, frost stress, and drought, may damage 

American chestnut. Excessive heating of the soil can 

injure chestnut when young (Hough 1878), and sprouts are 

sensitive to frost when green, tender, and close to the ground 

(Mattoon 1909, Zon 1904). The health and vigor of sprouts 

is impaired by excessive heat or cold (Zon 1904). Gurney 

and others (2011) reported that American chestnut saplings 

are approximately 5 °C less cold tolerant than red oak and 

sugar maple, suggesting that the northern limit of the range 

may be limited by cold tolerance.

SPECIAL USES 

Perhaps the best known product of American chestnut 

has nothing to do with its wood value. The flavorful nut 

it produces was enjoyed raw, roasted, or boiled by Native 

Americans and settlers, and it was an important food source 

for livestock and wildlife such as turkeys and squirrels 

(Clapper and Gravatt 1943, Hepting 1974, Youngs 2000). 

For those living in the Appalachians, chestnuts were a 

supplemental food and provided a source of income, 

60 cents to $5 per bushel (Zon 1904). During the late 

19th century and early 20th century, chestnuts were a 

popular treat in large cities and could be a profitable small 

business venture.

American chestnut wood has a narrow whitish to light 

brown sapwood and grayish brown heartwood that 

darkens with age. American chestnut is an ideal tree for 

many common forest products because it is rot-resistant, 

straight-grained, and moderately light-weight, with an 

average specific gravity of 0.4. It experiences moderate 

shrinkage and has little tendency to warp or check when 

drying (Youngs 2000). In the past, American chestnut was 

used for heavy lumber, shingles, fuelwood, telegraph and 

telephone poles, rails, posts, pilings, trolley and railroad 

ties, and even musical instruments such as pianos (Hepting 

1974, Smith 2000, Zon 1904). Frothingham (1912) 

reported that chestnut was used for building, bridge and 

car construction, interior finish, furniture, molding, and 

cabinet work, and because of its uniform density, American 

chestnut, was ideal for annealing brass, a process requiring 

good temperature control. 

American chestnut was once the major source of tannins 

for leather production in the United States. With tannin 

content of 6 to 11 percent and numerous large logs 

available, American chestnut was an obvious choice for 

the tanning industry (Youngs 2000). Processed chestnut 

ended up in tanneries all over the Appalachians but also 

in Massachusetts, the leading leather-producing State. 

Dependence of the industry was such that even after most 

chestnuts were killed by the blight, dead trees were being 

harvested for tanneries (Clapper and Gravatt 1943). 

GENETICS

American chestnut is thought to have the lowest genetic 

variability within the Castanea genus (Dane and others 

2003, Huang and others 1998). The proportion of genetic 

diversity found among American chestnut populations 

is much lower than averages reported for species with a 

wide geographic range, species with any seed dispersal 

mechanism, species with similar modes of reproduction, 

or temperate species. Most genetic diversity appears to be 

within populations (Huang 1998). Low genetic diversity 

could be related to the demise of the species due to ink 

disease and chestnut blight. 

12

4 Clark, S.L. 2011. Thinning and burning effects on northern red oak and American chestnut. Manuscript in preparation. Author can be reached at 

2431 Joe Johnson Drive, Room 274, Knoxville, TN 37996-4563.
American chestnut will readily hybridize with Chinese 

chestnut and Japanese (Castanea crenata) chestnut, crosses 

of which are being developed in search of a blight resistant 

tree (Anagnostakis 2002, Burnham and others 1986, 

Hebard 2005). A study by Clapper (1952) crossed American 

chestnut with Chinese, Japanese and Seguin (Castanea 

seguinii) chestnuts, as well as native chinquapin species, 

many of which have been more recently classified as 

varieties of Castanea pumila. 

RESTORATION

Over the past 90 years, much research has been conducted 

to produce blight-resistant chestnuts using breeding 

techniques, biological control, and genetic engineering. 

Chestnuts were bred as early as the 1890s, primarily to 

improve nut production using hybrids of native chestnut and 

chinquapins with Asian and European varieties (Burnham 

and others 1986, Van Fleet 1914). Early breeding for blight 

resistance was first conducted in the 1920s by the Office 

of Forest Pathology, U.S. Department of Agriculture, and 

by Arthur Graves in his work with the Brooklyn Botanical 

Garden. Grave’s work was later transferred in the 1940s to 

the Connecticut Agriculture Experiment Station (Burnham 

and others 1986). Early efforts were primarily focused 

on developing a single cross with the desired traits that 

could be vegetatively propagated (Diller and Clapper 

1969, Graves 1942), but the programs failed by the 1960s 

primarily due to an inadequate understanding of inheritance 

of resistance and desired form (Burnham and others 1986). 

Alternative breeding programs have been attempted to 

develop crosses that promote natural resistance among 

existing American chestnut populations (Griffin 2000). 

Researchers with the American Chestnut Cooperators 

Foundation cross surviving American chestnuts, and have 

yielded a relatively high frequency of progeny with low 

levels of blight resistance (Griffin 2000). Since 1986, the 

American Chestnut Cooperators Foundation has planted 

over 135,000 trees of these progenies across the native range 

of American chestnut.

Hypovirulence of the chestnut blight has been tested 

as a means to control the disease (Anagnostakis 1982, 

Fulbright and others 1983, Jaynes and Elliston 1980). 

Research from Europe and the United States suggests that 

hypovirulent strains of the chestnut blight can weaken the 

pathogen (Anagnostakis 2001, Griffin and others 1983, 

MacDonald and Fulbright 1991). Hypovirulent strains 

of the blight can survive and spread for long periods in 

natural populations (Anagnostakis 2001). While widespread 

efforts to control the blight through hypovirulence alone 

are probably not realistic, biological control appears to 

be an important part of the solution (Anagnostakis 1987, 

Milgroom and Cortesi 2004). 

Currently, the most promising option for widespread 

restoration of American chestnut is the development of 

a blight resistant chestnut variety through the backcross 

breeding programs of the American Chestnut Foundation 

and the Connecticut Agricultural Experiment Station 

(Anagnostakis 2002, Hebard 2005). The technique of using 

multiple backcrosses with the resistant parent did not begin 

until 1981. The breeding program includes the hybridization 

of American chestnut with Chinese chestnut or Japanese 

chestnut species that are naturally resistant to the blight. 

After the initial hybridization, subsequent back crossings 

with American chestnut progressively reduce the proportion 

of genetic material from the Asian parent, ultimately 

resulting in the third backcross hybrids (BC3F1) that are 94 

percent American chestnut but retain the genes for blight 

resistance. After screen testing, BC3F1 hybrids that exhibit 

blight resistance are then intercrossed for two generations to 

create the BC3F3 hybrids, which retain 94 percent American 

chestnut genetics in progeny as well as high levels of 

blight resistance. The American Chestnut Foundation is 

currently in the testing phase of its BC3F3 seedlings through 

inoculations in controlled orchard settings. Early testing 

indicates that morphological traits of the BC3F3 hybrids are 

consistent with American chestnut (Diskin and others 2006), 

but more testing is needed to confirm that blight resistance 

will not break down as the trees age (Hebard 2005). The 

latest tests indicate the BC3F3 families are more resistant 

than the American parent but did not have as high of a blight 

resistance as Chinese chestnut. Selection is not yet complete 

in The American Chestnut Foundations’ BC3F2 orchards.5

The Forest Service, U.S. Department of Agriculture, also 

plants BC3F3 material in forest settings to test blightresistance and morphological growth characteristics (Clark 

and others 2011, 2012a). Early results indicate that the 

BC3F3 generation has height growth similar to the American 

parent but bud break phenology that was slightly more 

developed than American chestnut (Clark and others 2011). 

Genetic engineering of American chestnut is currently under 

way. Progress has been made in somatic embryogenesis 

to rapidly produce seedlings from multiple genotypes that 

have undergone genetic engineering using a wheat gene 

(Merkle and others 2007, Pijut and others 2011). The 

widespread deployment of genetically engineered trees on 

public lands will require addressing a number of ethical 

and social questions, as well as a relatively large funding 

13

5 Personal communication. 2011. F.V. Hebard. Data from Meadowview blight-resistance screening tests. Author can be reached at 

29010 Hawthorne Dr., Meadowview, VA 24361-3349.
base (Merkle and others 2007). Deployment of trees bred 

using traditional breeding efforts is now under way (Clark 

and others 2011, 2012a) but selections of superior families 

are not yet complete. It is now understood that two or three 

incompletely dominant genes control blight resistance 

(Hebard 2005, Kubisiak and others 1997), and use of genetic 

marker-enhanced screening would speed up traditional 

breeding efforts substantially (Wheeler and Sederoff 

2009). Candidate genes for resistance are currently being 

identified (Barakat and others 2009) that can later be used 

in biotechnology efforts to produce a blight-resistant tree or 

to assist in DNA-marker assisted selection. DNA-marker 

assisted selection is a promising technique that could refine 

the selection of traits in the traditional breeding programs, 

resulting in more efficient breeding and reducing the time 

required to develop resistance (Kubisiak and others 1997, 

Wheeler and Sederoff 2009). Therefore, it is possible that a 

large quantity of seeds of blight resistant American chestnut 

and American chestnut varieties could be produced in the 

near future for the purpose of restoration. Another critical 

need in American chestnut restoration is to test the durability 

of blight resistant American chestnut and American chestnut 

varieties developed by the American Chestnut Foundation 

and the American Chestnut Cooperators Foundation, 

especially under various environmental stresses that are 

known to affect the vigor of American chestnut (Griffin and 

others 2006). 

ACKNOWLEDGMENTS

We thank Dr. Sandra Anagnostakis, of the Connecticut 

Agricultural Experiment Station, and Dr. Gary Griffin, of 

the American Chestnut Cooperators Foundation, for their 

thorough review of an early version of this manuscript. 

14

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Wang, G. Geoff; Knapp, Benjamin O.; Clark, Stacy L.; and Mudder, Bryan T. 2013. 

The Silvics of Castanea dentata (Marsh.) Borkh., American chestnut, Fagaceae (Beech Family). 

Gen. Tech. Rep. SRS-173. Asheville, NC: U.S. Department of Agriculture Forest Service, 

Southern Research Station. 18 p. 

This report describes how the American chestnut (Castanea dentata) was ecologically extirpated 

due to an exotic pathogen, the chestnut blight (Cryphonectria parasitica), and describes current 

restoration efforts. The habitat, life history, special uses, and genetics of the American chestnut are 

detailed. The American chestnut was an important and versatile tree species until its demise from the 

chestnut blight. Backcross breeding techniques, the use of hypovirulent blight strains, and genetic 

engineering programs are currently being developed and tested to produce trees resistant to the 

blight fungus. The first putative blight-resistant trees have been planted and silvicultural techniques 

to improve competitive ability of chestnut are being tested. American chestnut is a fast-growing 

species with the ability to persist in shaded conditions, and responds favorably to forest management 

techniques that limit competition and increase available sunlight. Restoration will require advanced 

artificial regeneration techniques. The effects of damaging agents other than blight, including root rot 

caused by Phytophthora cinnamomi, Asiatic oak weevil (Cyrtepistomus castaneus), the gypsy moth 

(Lymantria dispar), and Asian ambrosia beetles (Coleoptera: Scolytidae) are not well understood, but 

may represent real barriers to restoration.

Keywords: American chestnut, backcross breeding, chestnut blight, exotic species, restoration, 

silvics, silviculture.

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