Index of Species Information
SPECIES: Picea rubens
SPECIES: Picea rubens
AUTHORSHIP AND CITATION :
Sullivan, Janet. 1993. Picea rubens. In: Fire Effects Information System, [Online].
U.S. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences Laboratory (Producer).
Available: http://www.fs.fed.us/database/feis/ .
Picea rubra (Du Roi) Link
Picea australis Small
Picea nigra var. rubra Engelmann.
SCS PLANT CODE :
COMMON NAMES :
West Virginia spruce
The accepted scientific name for red spruce is Picea rubens Sarg. There
are no subspecies, varieties, or forms [48,64].
Natural hybrids with black spruce (P. mariana) have been reported
LIFE FORM :
FEDERAL LEGAL STATUS :
No special status
DISTRIBUTION AND OCCURRENCE
SPECIES: Picea rubens
GENERAL DISTRIBUTION :
Red spruce occurs from Cape Breton Island, Nova Scotia, and New
Brunswick, west to Maine, southern Quebec, and southeastern Ontario, and
south to central New York, northeastern Pennsylvania, northern New
Jersey, and northeastern Massachusetts. Its range extends south in the
Appalachian Mountains of extreme western Maryland, eastern West
Virginia, northern and western Virginia, western North Carolina, and
eastern Tennessee .
FRES10 White - red - jack pine
FRES11 Spruce - fir
FRES18 Maple - beech - birch
FRES19 Aspen - birch
CT ME MD MA NH NJ NY NC PA TN
VT VA WV NB NS PE PQ
BLM PHYSIOGRAPHIC REGIONS :
KUCHLER PLANT ASSOCIATIONS :
K096 Northeastern spruce - fir forest
K097 Southeastern spruce - fir forest
K108 Northern hardwoods - spruce forest
SAF COVER TYPES :
5 Balsam fir
12 Black spruce
17 Pin cherry
18 Paper birch
21 Eastern white pine
22 White pine - hemlock
23 Eastern hemlock
24 Hemlock - yellow birch
25 Sugar maple - beech - yellow birch
27 Sugar maple
30 Red spruce - yellow birch
31 Red spruce - sugar maple - beech
32 Red spruce
33 Red spruce - balsam fir
34 Red spruce - Fraser fir
35 Paper birch - red spruce - balsam fir
37 Northern white-cedar
60 Beech - sugar maple
107 White spruce
108 Red maple
SRM (RANGELAND) COVER TYPES :
HABITAT TYPES AND PLANT COMMUNITIES :
Red spruce is a common dominant or codominant in the red spruce and the
spruce-fir forests of the northeastern United States and adjacent
Shrub associates of red spruce in the Adirondack Mountains of New York
include red raspberry (Rubus idaeus), dwarfed blackberry (R. pubescens),
hobblebush (Viburnum alnifolium), Canada yew (Taxus canadensis), and
American fly honeysuckle (Lonicera canadensis). Ground layer herbs
include wild sarsaparilla (Aralia nudicaulis), Aster acuminatus, yellow
beadlily (Clintonia borealis), and common wood-sorrel (Oxalis montana).
Common bryophytes found in old-growth red spruce forests in the
Adirondacks include Brotherella recurvans, Schreber's moss (Pleurozium
schreberi), Polytrichum ohioense, mountain fern moss (Hylocomium
splendens), Bazzania trilobata, ptilium (Ptilium crista-castrensis),
Drepanocladus uncinatus, Dicranum scoparium, and D. montanum .
In the southern Appalachian Mountains, arboreal associates include
Fraser fir (Abies fraseri), yellow buckeye (Aesculus octandra), sweet
birch (Betula lenta), and black cherry (Prunus serotina) in addition to
those found in the northern part of its range [59,79,87]. Understory
associates in openings include rhododendrons (Rhododendron spp.),
American mountain-ash (Sorbus americana), and wild raisin (Viburnum
cassinoides). Other understory associates include highbush cranberry
(Viburnum edule), mountain holly (Ilex montana), mountain laurel (Kalmia
latifolia), speckled alder (Alnus rugosa), pin cherry (Prunus
pensylvanica), serviceberry (Amelanchier spp.), raspberries (Rubus
spp.), and blueberries and huckleberries (Vaccinium spp.). In closed
red spruce stands, mosses, lichens, and clubmosses predominate in the
understory along with wood sorrel (Oxalis spp.), trillium (Trillium
spp.), and checkerberry wintergreen (Gaultheria procumbens) .
Publications describing habitat or cover types in which red spruce is
dominant or codominant include:
(1) Proceedings of the Region 9 Land Systems conference on the White
Mountain National Forest 
(2) The Hubbard Brook ecosystem study: composition and dynamics of the
tree stratum 
(3) Ground vegetation patterns of the spruce-fir area of the Great
Smoky Mountains National Park 
(4) Spruce-fir forests of the coast of Maine 
(5) Forest type studies in the Adirondack region 
(6) The classification and evaluation of site for forestry 
(7) The identification and description of forest sites 
(8) Old-growth forests of Adirondack Park, New York 
(9) Vegetation-environment relationships in virgin, middle elevation
forests in the Adirondack Mountains, New York 
(10) Natural ecological communities of New York State 
SPECIES: Picea rubens
WOOD PRODUCTS VALUE :
Red spruce is one of the more important timber species in the
northeastern United States. The wood is light in weight, straight
grained, and resilient. It is used for paper, construction lumber, and
is highly preferred for musical instruments [9,29].
IMPORTANCE TO LIVESTOCK AND WILDLIFE :
Spruce grouse browse the leaves and twigs of red spruce . Mice and
voles consume and store significant amounts of spruce seeds, preferring
red and white spruce to balsam fir . Birds (particularly crossbills
or grosbeaks) will clip the terminal buds of young spruce, as will
porcupines, bears, snowshoe hares, and, rarely, deer [7,55,78]. Red
squirrels clip twigs and terminal buds and also eat reproductive and
vegetative buds [7,72].
In the southern part of its range, red spruce forests are used by only a
few wildlife species. Many of these species are usually only found
farther north, such as snowshoe hare, wood warblers and other songbirds,
rodents, and salamanders .
Red spruce is unpalatable to white-tailed deer .
NUTRITIONAL VALUE :
COVER VALUE :
Red spruce provides thermal and loafing cover for spruce grouse in
VALUE FOR REHABILITATION OF DISTURBED SITES :
Red spruce is occasionally used for revegetation of coal mine sites in
West Virginia, primarily at high elevations, but it is of limited value
for this purpose .
OTHER USES AND VALUES :
Red spruce gum was formerly collected and processed for chewing gum .
OTHER MANAGEMENT CONSIDERATIONS :
Silviculture: Various silvicultural systems may be used to manage red
spruce. Single tree selection, group selection, shelterwood, and strip
clearcut are all practical harvesting methods. Red spruce is subject to
windthrow; partial cuttings are recommended not to exceed half of the
basal area, and a lighter harvest is usually better. Seed tree cuts are
not recommended [6,9]. Frank and Blum  recommend a selection
silviculture where net growth is maximized by a 10-year, intensive
selection system. Clearcuts are contraindicated for many soil types and
fertility levels .
Postharvest red spruce regeneration is entirely dependent on advance
reproduction. If seedlings are not present at the time of logging, any
new spruce seedlings will be quickly overtopped and suppressed by faster
growing hardwoods . Leaf litter may aid in red spruce
regeneration. Harvesting during the dormant season or allowing
harvested trees to dry on site has been recommended to increase litter
. Loucks  noted that in the Maritime Provinces of Canada, red
spruce regeneration is usually good following partial cuts but may be
lacking in clearcuts.
The extent of red spruce forests has decreased following extensive
logging practices and subsequent fire . In the mountains of central
West Virginia, it is estimated that approximately 500,000 acres (200,000
ha) of red spruce present in the late 19th century had been reduced to
less than 60,000 acres (24,000 ha) by 1975, and as little as 17,500
acres (7,000 ha) in 1978 [10,73].
Management for wildlife: Harvest practices have an effect on the
resulting stand structure, and therefore on the numbers and species of
birds that use red spruce habitats. Crawford and Titterington 
identified five seral stages and the corresponding bird species, and
made associated recommendations for management of spruce-fir stands.
They also determined that spruce budworm infestation increases both the
number and diversity of birds. Dense, young stands of red spruce
support a higher population of birds but with less diversity than in
Insects and disease: Red spruce is relatively free from insects and
diseases until it is mature. Mature trees are susceptible to the
following insects: spruce budworm (Choristoneura fumiferana), eastern
spruce beetle (Dendroctonus rufipennis), European spruce sawfly (Diprion
hercyniae), yellowheaded spruce sawfly (Pikonema alaskensis), and
eastern spruce gall adelgid (Adelges abietis) [9,22,23,30]. Diseases of
red spruce have been detailed [9,22,23,30,47].
Red spruce decline: Throughout its range, growth rates of red spruce
have declined and mortality has increased . This decline is
apparently more severe at higher elevations, in older stands, and on
more exposed sites. This decline is not limited to red spruce; balsam
fir and associated white and black spruce appear to be affected also
. A number of studies on the causes of red spruce decline have
failed to make a definitive case for any single cause. There may be no
single cause or the complexity of the situation may not lend itself to a
clear cause-effect relationship [36,42,47]. The combination of climatic
stress and atmospheric pollution is probably the major cause of this
decline, according to a number of researchers [19,36,41,42]. Numerous
other causes have been proposed as well, including a natural cycle of
dieback and recovery [3, 36,]. A survey of the extent and identifiable
causes of mortality and decline was published in 1985 .
BOTANICAL AND ECOLOGICAL CHARACTERISTICS
SPECIES: Picea rubens
GENERAL BOTANICAL CHARACTERISTICS :
Red spruce is a native, evergreen conifer. It is a medium-sized tree,
attaining a maximum height of 115 feet (35 m); the average mature height
is 60 to 75 feet (18-23 m). The ovulate cones are 1.3 to 1.5 inches
(3-4 cm) long, with rigid rounded scales that are often slightly toothed
on the edges. Red spruce is very shallow rooted; most of the feeding
roots occur in the duff and top few centimeters of soil. In Maine, the
average depth of roots was 13 inches (33 cm), with a maximum depth of 22
inches (56 cm) . Red spruce is long-lived, often achieving ages
greater than 350 years .
RAUNKIAER LIFE FORM :
REGENERATION PROCESSES :
Red spruce reproduces exclusively by seed. The first cone crop is
usually produced when the crown first reaches direct light [27,39].
Therefore, red spruce can bear cones as early as 15 to 20 years of age;
cone production peaks about 15 years later. In dense, even-aged stands,
full cone crops are rare until the trees are 40 to 50 years old .
Good seed crops are produced every 3 to 8 years, with light crops in
intervening years. Cones are dropped shortly after they are mature .
The seeds are wind or rain disseminated. The maximum distance for
dispersal by wind is approximately 201 feet (61 m) . Seeds do not
exhibit dormancy. Most germinate the spring following dispersal;
occasionally germination will occur in the fall soon after seeds drop
from the tree. Seeds are usually not viable after 1 year. Germination
is largely controlled by moisture availability. Seeds will germinate in
almost any medium except sod. Seeds that germinate in thick duff are
subject to overheating and/or drought mortality. Drought and
frost-heave are the major causes of seedling mortality the first year
Successful reproduction appears to depend more on seedling survival than
on germination requirements . Seedling establishment is usually best
on shallow, less fertile soils that discourage competitive hardwoods
. The primary roots of red spruce seedlings do not penetrate litter
and forest duff to any depth . Red spruce seedlings have a root
system of finely branched rootlets and no strong laterals; they depend
entirely on the humus for nutrients and water .
SITE CHARACTERISTICS :
Red spruce grows in climates with cool, moist summers and cold winters
. In the northeastern United States, the mean annual precipitation
ranges from 36 to 52 inches (910-1,320 mm) and is often higher in the
mountainous terrain where red spruce occurs, due to fog drip. Snow
cover averages 80 to 160 inches (203-406 cm), with 100 to 140 days of
snow cover per year .
Most of the soils on which red spruce occurs are developed from glacial
deposits. The most productive soils are derived from parent materials
of unsorted glacial drift and till deposited on the midslopes of hills
and mountains. Soils on red spruce sites are usually acid Spodosols,
Inceptisols, and sometimes Histosols with thick mor humus and a
well-defined A2 horizon. Soil pH ranges from 4.0 to 5.5. Red spruce is
often found on sites that are unfavorable for other species, such as
organic soils overlying rocks in mountainous locales, on steep rocky
slopes with thin soils, and in wet bottomlands .
In the northern part of its range, red spruce occurs at elevations
ranging from sea level to 4,500 feet (0-1,370 m), above which it is
usually replaced by balsam fir (Abies balsamea). The elevational
zonation of species is defined as follows :
up to 1,485 feet (450 m) northern hardwoods (hemlock phase)
1,486 to 2,508 feet (451- 760 m) northern hardwoods (spruce phase)
2,508 to 4,026 feet (761-1,220 m) subalpine (spruce-fir phase)
4,027 to 4,785 feet (1,221-1,450 m) subalpine (fir phase)
In the southern Appalachian Mountains, red spruce occurs at elevations
from about 3,200 feet to 6,200 feet (980-1,890 m); above 6,200 feet
(1,890 m), red spruce tends is usually replaced by Fraser fir (Abies
SUCCESSIONAL STATUS :
On shallow, acidic, glacial till soils, red spruce is considered climax.
It is usually subclimax on fertile, well-drained slopes and on abandoned
fields and pastures where is is replaced by shade-tolerant hardwoods
such as sugar maple and beech. Other types, such as red spruce-balsam
fir and red spruce-yellow birch are usually climax .
Red spruce is tolerant of shade. Seedlings of red spruce can establish
in as little as 10 percent of full sunlight, but for optimum growth, at
least 50 percent of full sunlight is needed [9,75,81]. Growth tends to
be suppressed in shade, but such suppression can persist for many years
without killing the tree. For example, suppressed understory
individuals may be 4 to 5 feet (1.2-1.5 m) tall, and be more than 50
years old. In comparison, open-grown red spruce can reach sawtimber
size at 50 years [9,29].
Red spruce responds to canopy removal even after many years of
suppression. The taller and older a seedling or sapling is, the greater
is its response to release, up to about 55 years of age after which
response to release starts to decline. However, the amount of response
does not revert to seedling levels until the tree is around 100 years of
age. Umbrella-shaped saplings 40 to 80 years old that have been
suppressed will respond to release after a delay of several years, and
in fact have an advantage because they are taller than smaller,
healthier saplings which respond more quickly to canopy opening. More
than half of mature red spruce second growth arises from larger but
suppressed advance growth, as opposed to having arisen from small
advance growth or new seedlings . Upon release, 60-year-old red
spruce growth exceeds that of same-age balsam fir and therefore tends to
dominate the canopy .
Leak  defined red spruce in New Hampshire as a dominating climax
species on shallow, dry, wet, or poorly aerated soils; it is a minor
component in young stands but increases markedly over time until it is a
canopy dominant. He estimated that, if undisturbed, red spruce can
reach densities of 70 to 80 percent in a minimum of 250 years. Red
spruce is a long-lived species and, once established, persists as a
dominant for a long time.
Davis  observed young spruce-fir stands in coastal Maine originating
in open sites and as the understory to early seral hardwoods such as
paper birch. The young, open-grown stands may be dominated by white
spruce, red spruce, or balsam fir in any proportions. A spruce-fir
stand originating as understory tends to be dominated by red spruce
and/or balsam fir, though white spruce is often present. Moore 
found red spruce forests to be even-aged in groups, indicating that
establishment and/or canopy achievement tends to occur in openings.
Red spruce and red spruce-fir cover types are self-maintaining. Stand
composition may vary with stand age. Both red spruce and its two fir
associates (balsam and Fraser) are shade tolerant, and both spruce and
fir reproduction are found under spruce-fir canopies [6,16]. In the
Catskill Mountains of New York, balsam fir reproduction predominated
under both spruce and balsam fir stands. Both red spruce and balsam fir
reproduction occurred at low densities under hardwood stands (mostly
yellow birch) . McIntosh and Hurley  do not believe that red
spruce forests form a self-perpetuating climax in this area. Their
conclusion may be biased, however, since balsam fir outcompetes red
spruce in early stages, but is usually overtopped or outcompeted by red
spruce in more mature forests . Flieger  described 350-year-old
stands of red spruce which were characterized by irregular stocking and
variable crown heights and widths, with at least two age classes
apparent. Most virgin red spruce forests are uneven-aged, indicating
that the forests did no originate following stand-destroying
disturbances, and that red spruce is able to reproduce under its own
SEASONAL DEVELOPMENT :
Red spruce vegetative buds begin growth from May 26 to June 3 .
Needles are shed early in summer . Reproductive cones open in late
April to early May [29,72]. Red spruce cones mature the first autumn
from mid-September to mid-October [29,39]. Dissemination of seeds
begins soon after cones are ripe and continues until March .
SPECIES: Picea rubens
FIRE ECOLOGY OR ADAPTATIONS :
Red spruce forests persist without fire. Red spruce is easily killed by
fire due to its thin bark, shallow roots, flammable needles, and lack of
self-pruning [9,23,39]. Its slow early growth rate delays the formation
of a corky layer, which increases the fire susceptibility of young trees
. In a study based on a survey of foresters, Starker  rated the
fire resistance of 22 New England tree species based on fire mortality
and fire avoidance (occurrence in habitat that does not burn very
often). Red spruce was not resistant in terms o fire mortality but
moderately or very resistant in terms of fire avoidance, and was ranked
Red spruce habitat is subject to few fires; fires that occurred in
presettlement times were usually of low severity . Saunders 
noted that old-timers claimed that forest fires would stop when they
reached the spruce-fir forest boundary. Electrical storms are common in
this area but are usually accompanied by sufficient rain, and fuels are
usually moist . Severe surface fires probably occurred
infrequently, during periods of prolonged drought, and usually affected
forests that were breaking up due to wind, ice storm damage, or similar
events that generate surface fuels [25,32,60,61,87].
The estimated natural fire return intervals for the northeastern United
States and adjacent Canada range from 330 to 3,300 or more years
[25,32,51,52,84]. Estimates of natural fire frequency have been
complicated by human activities. Logging in these forests has resulted
in an increase in fire frequency and intensity, particularly in logging
slash [18,32,52]. The catastrophic fires of the 19th and 20th centuries
can be attributed to human activities [21,32,52]. However, even with
the increase in fires due to human activity, most fires are small and
quickly suppressed. There should be sufficient time between fires for
red spruce to regain dominance on most sites unless deliberately and/or
It has been suggested that, in presettlement forests, the increase of
dead fuels following spruce budworm outbreaks increased the likelihood
of fire [21,25,32]. Such outbreaks are more common in
balsam-fir-dominated forests than in red-spruce-dominated forests, but
the two species usually occur together, in varying proportions.
Before settlement by Europeans, forests in northern New England, the
Adirondack Mountains, and the hillier sections of southern New England
and Pennsylvania were not deliberately burned by Native Americans as
were other areas in the northeastern United States .
POSTFIRE REGENERATION STRATEGY :
Tree without adventitious-bud root crown
Secondary colonizer - off-site seed
SPECIES: Picea rubens
IMMEDIATE FIRE EFFECT ON PLANT :
Red spruce is easily killed by fire . Surface or ground fires that
consume the litter and organic layers covering the superficial roots of
red spruce are almost certain to severely injure the roots . Fire
kills mature trees by exposing roots, subjecting the tree to water
stress and/or windthrow, which may result in the eventual death of the
DISCUSSION AND QUALIFICATION OF FIRE EFFECT :
PLANT RESPONSE TO FIRE :
Red spruce does not sprout. Seed germination is greater on burned areas
with exposed mineral soil than in duff; mortality, however, is also
greater due to increased surface temperature and drought .
Burned red spruce or spruce-fir stands are initially restocked by aspen
(Populus spp.) or birch (Betula spp.) via wind-disseminated seed; paper
birch (Betula papyrifera)-aspen stands are particularly diagnostic of
fire in upland red spruce forests . Red spruce seedlings appear a
few years after fire, developing as an understory in the aspen-birch
complex, and eventually penetrate the overstory after 50 or 60 years.
Birch and aspen become decadent after 75 to 80 years and red spruce or
red spruce and balsam fir regain dominance if left undisturbed
[49,52,65]. On better sites, northern hardwoods, chiefly sugar maple
and American beech, may replace red spruce, and in some areas, balsam
fir will dominate the late postfire succession. Postharvest/postfire
restocking by red spruce is extremely slow where the organic layers are
destroyed by severe fire (particularly where harvest has been heavy)
In Nova Scotia, mature spruce forests have few herbs and shrubs in the
understory. After a fire, herbs increase in the first 6 years and
dominate for 40 or more years while conifers slowly establish .
After fire in the southern Appalachians, blackberry (Rubus ursinus) and
red raspberry colonize the site. Pin cherry and yellow birch follow.
Blackberry and raspberry are too competitive for red spruce and must be
shaded out by the hardwoods before red spruce can establish .
In West Virginia, postlogging and postfire succession in red spruce
forests follows a similar pattern: ferns and raspberry are followed by
other shrubs, then hardwoods (particularly hawthorn [Crataegus spp.]),
and eventually spruce. In many areas, this successional pattern has
been extremely slow; heaths or barrens form that do not appear as if
they will ever return to forest . Martin  studied
postlogging/postfire succession in Nova Scotia and found that red spruce
was present on most sites after the second postfire year, becoming more
numerous and dominant in the later seres. He concluded that repeated
heavy cuttings and light fires on the poorer soils of the southern
upland of Nova Scotia encourages the invasion of heath plants, which
limits the rate and amount of tree regeneration.
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE :
FIRE MANAGEMENT CONSIDERATIONS :
Some managers believe that prescribed fire may be a useful silvicultural
tool for managing red spruce on some sites. On such sites, the exposed
mineral soil must have plentiful moisture, soil temperatures must be
moderate, and competition must be minimal . In general, however,
fires in red spruce habitat are of little silvicultural value .
Slash burning following logging kills advance reproduction and creates
rank postfire vegetation that delays any new seedling establishment
The fire management plan for Acadia National Park, Maine, dictates the
suppression of natural fires. Prescribed fires may be used on occasion
to reduce fuels . Patterson and others  estimated fuel loadings
for a number of stands in Acadia National Park that contained red
spruce. They concluded that fire exclusion was probably resulting in
increased fuel loads.
Alexander  compiled slash fuel indices for red spruce and compared
actual fire spread, intensity, and slash and organic layer depletions
with those predicted by the Canadian Forest Fire Danger Rating System.
Freeman and others  developed equations to determine average crown
weight per tree as a function of tree height and diameter for use in a
method to predict slash weight after logging red spruce.
SPECIES: Picea rubens
1. A. D. Revill Associates. 1978. Ecological eff. of fire and its mgmt. in
Canada's national parks: a synthesis of the literature. Vols 1&2. Lit.
Rev. & Annot. Bibliography. Ottawa, ON: Parks Canada, National Parks
Branch, Natural Resources Division. 345 p. 
2. Abbott, Herschel G. 1962. Tree seed preferences of mice and voles in the
Northeast. Journal of Forestry. 60: 97-99. 
3. Adams, Harold S.; Stephenson, Steven L. 1989. Old-growth red spruce
communities in the mid-Appalachians. Vegetatio. 85: 45-56. 
4. Alexander, Martin E. 1984. Prescribed fire behavior and impact in an
eastern spruce-fir slash fuel complex. Canadian Forestry Service
Research Notes. 4(1): 3-10. 
5. Fay, Stephen C.; Alvis, Richard. 1993. White Mountain landscapes.
Laconia, NH: U.S. Department of Agriculture, Forest Service, Region 9,
White Mountain National Forest. 76 p. Working draft. 
6. Benzie, John W.; Blum, Barton M. 1989. Silviculture of northeastern
conifers. In: Burns, Russell M., compiler. The scientific basis for
silvicultural and management decisions in the National Forest System.
Gen. Tech. Rep. WO-55. Washington, DC: U.S. Department of Agriculture,
Forest Service: 18-30. 
7. Blum, Barton M. 1977. Animal damage to young spruce and fir in Maine.
Res. Note NE-321. Upper Darby, PA: U.S. Department of Agriculture,
Forest Service, Northeastern Forest Experiment Station. 4 p. 
8. Blum, Barton M. 1988. Variation in the phenology of bud flushing in
white and red spruce. Canadian Journal of Forest Research. 18: 315-319.
9. Blum, Barton M. 1990. Picea rubens Sarg. red spruce. In: Burns, Russell
M.; Honkala, Barbara H., technical coordinators. Silvics of North
America. Volume 1. Conifers. Agric. Handb. 654. Washington, DC: U.S.
Department of Agriculture, Forest Service: 250-259. 
10. Bones, James T. 1978. The forest resources of West Virginia. Upper
Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern
Forest Experiment Station. [Pages unknown]. 
11. Bormann, F. H.; Siccama, T. G.; Likens, G. E.; Whittaker, R. H. 1970.
The Hubbard Brook ecosystem study: composition and dynamics of the tree
stratum. Ecological Monographs. 40(4): 373-388. 
12. Chandler, Robert F.; Jr. 1943. Amount and mineral nutrient content of
freshly fallen needle litter of some northeastern conifers. Proceedings,
Soil Science of America Society. 8: 409-411. 
13. Core, Earl L. 1929. Plant ecology of Spruce Mountain, West Virginia.
Ecology. 10(1): 1-13. 
14. Crandall, Dorothy L. 1958. Ground vegetation patterns of the spruce-fir
area of the Great Smoky Mountains National Park. Ecological Monographs.
28(4): 337-360. 
15. Crawford, Hewlette S.; Titterington, Richard W. 1979. Effects of
silvicultural practices on bird communities in upland spruce- fir
stands. In: DeGraaf, Richard M.; Evans, Keith E., compilers. Management
of north central and northeastern forests for nongame birds: Proceeding
of the workshop; 1979 January 23-25; Minneapolis, MN. Gen. Tech. Rep.
NC-51. St. Paul, MN: U.S. Department of Agriculture, Forest Service,
North Central Forest Experiment Station: 110-119. 
16. Davis, Ronald B. 1966. Spruce-fir forests of the coast of Maine.
Ecological Monographs. 36(2): 79-94. 
17. Davis, William C. 1991. The role of advace growth in regeneration of red
spruce and balsam fir in east central Maine. In: Simpson, C. M, ed.
Proceedings of the conference on natural regeneration management; 1990
March 27-28; Fredericton, NB. Fredericton, NB: Forestry Canada,
Maritimes Region: 157-168. 
18. Day, Gordon M. 1953. The Indian as an ecological factor in the
northeastern forest. Ecology. 34(2): 329-346. 
19. DeHayes, D. H.; Waite, C. E.; Ingle, M. A.; Williams, M. W. 1990. Winter
injury susceptibility and cold tolerance of current and year-old needles
of red spruce trees from several provenances. Forest Science. 36(4):
20. Eyre, F. H., ed. 1980. Forest cover types of the United States and
Canada. Washington, DC: Society of American Foresters. 148 p. 
21. Flieger, B. W. 1971. Forest fire and insects: the relations of fire to
insect outbreak. In: Proceedings, annual Tall Timbers fire ecology
conference; 1970 August 20-21; Fredericton, NB. No. 10. Fredericton, NB:
Tall Timbers Research Station: 107-114. 
22. Frank, Robert M.; Bjorkbom, John C. 1973. A silvicultural guide for
spruce-fir in the northeast. NE-6. Upper Darby, PA: U.S. Department of
Agriculture, Forest Service, Northeastern Forest Experiment Station. 29
23. Frank, Robert M.; Blum, Barton M. 1978. The selection system of
silviculture in spruce-fir stands--procedures, early results, and
comparisons with unmanaged stands. Res. Pap. NE-425. Upper Darby, PA:
U.S. Department of Agriculture, Forest Service, Northeastern Forest
Experiment Station. 15 p. 
24. Freeman, Duane R.; Loomis, Robert M.; Roussopoulos, Peter J. 1982.
Handbook for predicting slash weight in the Northeast. Gen. Tech. Rep.
NC-75. St. Paul, MN: U.S. Department of Agriculture, Forest Service,
North Central Forest Experiment Station. 23 p. 
25. Furyaev, V. V.; Wein, Ross W.; MacLean, David A. 1983. Fire influences
in Abies-dominated forests. In: Wein, Ross W.; MacLean, David A., eds.
The role of fire in northern circumpolar ecosystems. Scope 18.
Chichester; New York: John Wiley & Sons: 221-234. 
26. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others].
1977. Vegetation and environmental features of forest and range
ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of
Agriculture, Forest Service. 68 p. 
27. Govindaraju, Diddahally R. 1988. Life histories, neighbourhood sizes,
and variance structure in some North American conifers. Biological
Journal of the Linnean Society. 35: 69-78. 
28. Halliday, W. E. D. 1937. A forest classification for Canada. Forest
Service Bulletin No. 89. Ottawa: Canadian Department of Forestry,
Resource Development. 50 p. 
29. Hart, Arthur C. 1959. Silvical characteristics of red spruce. Paper No.
124. Durham, NH: U.S. Department of Agriculture, Forest Service,
Northeastern Forest Experiment Station. 18 p. 
30. Hawksworth, F. G.; Shigo, A. L. 1980. Dwarf mistletoe on red spruce in
the White Mountains of New Hampshire. Plant Disease. 64(9): 880-882.
31. Heimburger, Carl C. 1934. Forest-type studies in the Adirondack Region.
Memoir 165. Ithaca, NY: Cornell University, Agricultural Experiment
Station. 122 p. 
32. Heinselman, Miron L. 1981. Fire intensity and frequency as factors in
the distribution and structure of northern ecosystems. In: Mooney, H.
A.; Bonnicksen, T. M.; Christensen, N. L.; [and others], technical
coordinators. Fire regimes and ecosystem properties: Proceedings of the
conference; 1978 December 11-15; Honolulu, HI. Gen. Tech. Rep. WO-26.
Washington, DC: U.S. Department of Agriculture, Forest Service: 7-57.
33. Hills, G. A. 1952. The classification and evaluation of site for
forestry. Res. Rep. No. 24. Toronto, ON: Ontario Department of Lands and
Forests, Division of Research. 41 p. 
34. Hills, G. A. 1954. Field methods for investigating site: A. The detailed
site description form. (Part IV of Bulletin "The forest sites of
Ontario). Site Research Manual No. 4. Toronto, ON: Department of Lands
and Forests, Research Division. 104 p. 
35. Hornbeck, J. W.; Smith, C. T.; Martin, Q. W.; [and others]. 1990.
Effects of intensive harvesting on nutrient capitals of three forest
types in New England. Forest Ecology and Management. 30: 55-64. 
36. Johnson, Arthur H.; McLaughlin, Samuel B. 1986. The nature and timing of
the deterioration of red spruce in the northern Appalachian Mountains.
In: Acid deposition: long term trends. [Place of publication unknown]:
National Academy Press: 200-230. 
37. Kartesz, John T.; Kartesz, Rosemarie. 1980. A synonymized checklist of
the vascular flora of the United States, Canada, and Greenland. Volume
II: The biota of North America. Chapel Hill, NC: The University of North
Carolina Press; in confederation with Anne H. Lindsey and C. Richie
Bell, North Carolina Botanical Garden. 500 p. 
38. Klein, Richard M.; Perkins, Timothy D.; Tricou, Jeffery; [and others].
1991. Factors affecting red spruce regeneration in declining areas of
Camels Hump Mountain, Vermont. American Journal of Botany. 78(9):
39. Korstian, Clarence F. 1937. Perpetuation of spruce on cut-over and
burned lands in the higher Southern Appalachian Mountains. Ecological
Monographs. 7(1): 125-167. 
40. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation
of the conterminous United States. Special Publication No. 36. New York:
American Geographical Society. 77 p. 
41. LeBlanc, David C.; Raynal, Dudley J.; White, Edwin H.; Ketchledge, Edwin
H. 1987. Characterization of historical growth patterns in declining red
spruce trees. In: Jacoby, G. C., Jr.; Hornbeck, J. W., eds.
International symposium on ecological aspects of tree-ring analysis;
[Date of conference unknown]; Marymount College, NY. [Place of
publication unknown]. [Publisher unknown]. 360-371. 
42. LeBlanc, David C.; Raynal, Dudley J. 1990. Red spruce decline on
Whiteface Mountain, New York. II. Relationships between apical and
radial growth decline. Canadian Journal of Forest Research. 20(9):
43. Leak, W. B. 1975. Age distribution in virgin red spruce and northern
hardwoods. Ecology. 56: 1451-1454. 
44. Leak, William B. 1991. Secondary forest succession in New Hampshire,
USA. Forest Ecology and Management. 43: 69-86. 
45. Leak, William B.; Graber, Raymond E. 1974. Forest vegetation related to
elevation in the White Mountains of New Hampshire. NE-299. Upper Darby,
PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest
Experiment Station. 7 p. 
46. Leak, William B.; Solomon, Dale S.; Filip, Stanley M. 1969. A
silvicultural guide for northern hardwoods in the northeast. Res. Pap.
NE-143. Upper Darby, PA: U.S. Department of Agriculture, Forest Service,
Northeastern Forest Experiment Station. 34 p. 
47. Leopold, Donald J.; Reschke, Carol; Smith, Daniel S. 1988. Old-growth
forests of Adirondack Park, New York. Natural Areas Journal. 8(3):
48. Little, Elbert L., Jr. 1979. Checklist of United States trees (native
and naturalized). Agric. Handb. 541. Washington, DC: U.S. Department of
Agriculture, Forest Service. 375 p. 
49. Little, Silas. 1974. Effects of fire on temperate forests: northeastern
United States. In: Kozlowski, T. T.; Ahlgren, C. E., eds. Fire and
ecosystems. New York: Academic Press: 225-250. 
50. Long, H. D. 1952. Forest types and sites of the Acadia Forest Experiment
Station. Unpublished paper on file at: Canada Department of Resources
and Development, Forestry Branch, [Place of publication unknown]: [Pages
51. Lorimer, Craig G. 1977. The presettlement forest and natural disturbance
cycle of northeastern Maine. Ecology. 58: 139-148. 
52. Lorimer, Craig G. 1980. The use of land survey records in estimating
presettlement fire frequency. In: Stokes, Marvin A.; Dieterich, John H.,
technical coordinators. Proceedings of the fire history workshop; 1980
October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S.
Department of Agriculture, Forest Service, Rocky Mountain Forest and
Range Experiment Station: 57-62. 
53. Loucks, O. L. 1959. A forest classification for the Maritime Provinces.
Proceedings, Nova Scotian Institute on Science. 25: 86-167. 
54. Martin, J. Lynton. 1956. An ecological survey of burned-over forest land
in southwestern Nova Scotia. Forestry Chronicle. 32: 313-336. 
55. McIntosh, R. P.; Hurley, R. T. 1964. The spruce-fir forest of the
Catskill Mountains. Ecology. 45(2): 314-326. 
56. Meyer, Walter H. 1929. Yields of second-growth spruce and fir in the
Northeast. Tech. Bull. No. 142. Washington, DC: U.S. Department of
Agriculture. 52 p. 
57. Moore, Barrington. 1917. Some factors influencing the reproduction of
red spruce, balsam fir, and white pine. Journal of Forestry. 15(7):
58. Moore, Barrington. 1922. Humus and root systems in certain northeastern
forests in relation to reproduction and competition. Journal of
Forestry. 20: 233-254. 
59. Oosting, H. J.; Billings, W. D. 1951. A comparison of virgin spruce-fir
forest in the northern and southern Appalachian system. Ecology. 32(1):
60. Patterson, William A., III; Saunders, Karen E.; Horton, L. J. 1983. Fire
regimes of the coastal Maine forests of Acadia National Park. OSS 83-3.
Boston, MA: U.S. Department of the Interior, National Park Service,
North Atlantic Region, Office of Scientific Studies. 259 p. In
cooperation with: U.S. Department of Agriculture, Forest Service, State
and Private Forestry, Broomall, PA. 
61. Patterson, William A., III; Saunders, Karen E.; Horton, L. J.; Foley,
Mary K. 1985. Fire management options for coastal New England forest:
Acadia National Park and Cape Cod National Seashore. In: Lotan, James
E.; Kilgore, Bruce M.; Fischer, William C.; Mutch, Robert W., technical
coordinators. Proceedings--symposium and workshop on wilderness
fire;1983 November 15-18; Missoula, MT. Gen. Tech. Rep. INT-182. Ogden,
UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest
and Range Experiment Station: 360-365. 
62. Pielou, E. C. 1988. The world of northern evergreens. Ithaca, NY:
Cornell University Press. 174 p. 
63. Place, I. C. M. 1955. The influence of seed-bed conditions on the
regeneration of spruce and balsam fir. Bulletin 117. Ottawa, Canada:
Department of Northern Affairs and National Resources, Forestry Branch,
Forest Research Division. 87 p. 
64. Radford, Albert E.; Ahles, Harry E.; Bell, C. Ritchie. 1968. Manual of
the vascular flora of the Carolinas. Chapel Hill, NC: The University of
North Carolina Press. 1183 p. 
65. Randall, Arthur G. 1976. Natural regeneration in two spruce-fir
clearcuts in eastern Maine. Research Life Sciences. [University of
Maine]; 23(13): 1-10. 
66. Raunkiaer, C. 1934. The life forms of plants and statistical plant
geography. Oxford: Clarendon Press. 632 p. 
67. Reiners, William A,; Lang, Gerald E. 1979. Vegetational patterns and
processes in the balsam fir zone, White Mountains, New Hampshire.
Ecology. 60(2): 403-417. 
68. Roman, John Ross. 1980. Vegetation-environment relationships in virgin,
middle elevation forests in the Adirondack Mountains, New York.
Syracuse, NY: State University of New York. PhD. Dissertation.
Dissertation Abstracts International. 41(3): 807-B. 
70. Rowe, J. S. 1972. Forest regions of Canada. Publication No. 1300.
Ottawa: Department of the Environment, Canadian Forestry Service. 172 p.
71. Reschke, Carol. 1990. Ecological communities of New York State. Latham,
NY: New York State Department of Environmental Conservation, New York
Natural Heritage Program. 96 p. 
72. Safford, L. O. 1974. Picea A. Dietr. spruce. In: Schopmeyer, C. S., ed.
Seeds of woody plants in the United States. Agric. Handb. 450.
Washington, DC: U.S. Department of Agriculture, Forest Service: 587-597.
73. Saunders, Paul Richard. 1979. The vegetational impact of human
distubance on the spurce-fir forests of the southern Appalachian
Mountains. Durham, NC: Duke University. 188 p. Dissertation. 
74. Saunders, Paul R.; Smathers, Garrett A.; Ramseur, George S. 1983.
Secondary succession of a spruce-fir burn in the Plott Balsam Mountains,
North Carolina. Castanea. 48(1): 41-47. 
75. Shirley, Hardy L. 1943. Is tolerance the capacity to endure shade?.
Journal of Forestry. 41: 339-345. 
76. Starker, T. J. 1934. Fire resistance in the forest. Journal of Forestry.
32: 462-467. 
77. Stickney, Peter F. 1989. Seral origin of species originating in northern
Rocky Mountain forests. Unpublished draft on file at: U.S. Department of
Agriculture, Forest Service, Intermountain Research Station, Fire
Sciences Laboratory, Missoula, MT; RWU 4403 files. 7 p. 
78. Telfer, Edmund S. 1972. Browse selection by deer and hares. Journal of
Wildlife Management. 36(4): 1344-1349. 
79. Trimble, George R., Jr.; Patric, James H.; Gill, John D.; [and others].
1974. Some options for managing forest land in the central Appalachians.
Gen. Tech. Rep. NE-12. Upper Darby, PA: U.S. Department of Agriculture,
Forest Service, Northeastern Forest Experiment Station. 42 p. 
80. U.S. Department of Agriculture, Soil Conservation Service. 1982.
National list of scientific plant names. Vol. 1. List of plant names.
SCS-TP-159. Washington, DC. 416 p. 
81. Vezina, P. E.; Pech, Gy. 1964. Solar radiation beneath conifer canopies
in relation to crown closure. Forest Science. 10(4): 443-451. 
82. Vogel, Willis G. 1981. A guide for revegetating coal minesoils in the
eastern United States. Gen. Tech. Rep. NE-68. Broomall, PA: U.S.
Department of Agriculture, Forest Service, Northeastern Forest
Experiment Station. 190 p. 
83. Wahlenberg, W. G. 1951. Planting in the Appalachian spruce-fir type.
Journal of Forestry. 49(8): 569-571. 
84. Wein, Ross W.; Moore, Janice M. 1979. Fire history and recent fire
rotation periods in the Nova Scotia Acadian Forest. Canadian Journal of
Forest Research. 9: 166-178. 
85. Weiss, Melvyn J.; McCreery, Lew R.; Millers, Imants; [and others]. 1985.
Cooperative survey of red spruce and balsam fir decline and mortality in
New Hampshire, New York, and Vermont--1984. Durham, NH: U.S. Department
of Agriculture, Forest Service, Northeastern Area, Forest Pest
Management. 130 p. 
86. Westveld, Marinus. 1931. Reproduction on pulpwood lands in the
Northeast. Tech. Bull. No. 223. Washington, DC: U.S. Department of
Agriculture. 52 p. 
87. Wright, Henry A.; Bailey, Arthur W. 1982. Fire ecology: United States
and southern Canada. New York: John Wiley & Sons. 501 p. 
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