Documente Academic
Documente Profesional
Documente Cultură
R. H. Whittaker
Ecological Monographs, Vol. 26, No. 1. (Jan., 1956), pp. 1-80.
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T A B L E OF C O N T E S T S
PAGE
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Literature on Area . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Acknowledgments . . . . . . . . . . . .
.......... 2
Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Growth-Forms ...............................
10
Coverages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Diversities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Self-Maintenance of Stands . . . . . . . . . . . . . . . . . . . . 11
Subxeric Sites . . . . . . . . . . . . . . . . . . . . . . . . .
Xeric Sites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Stratal Distributions . . . . . . . . . . . . . . . . . . . . . . . . . .
Trends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
I 1 . D I S C U S S I O NA:N INTERPREPATION
OF
\ T F X 3 ~PATTERNING
~ ~ ~ ~ .~. .~
. . . . . . . . . . . . . . . . . .
Distributions o f Species and the Study o f Genecology
T h e Association-Unit Theory and Individualistic
Hypothesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
T h e Distributional Basis o f Community.Types . . . . . .
Gradation and the Grouping o f Species . . . . . . . . .
I. GRADIENT ANALYSIS
INTRODUCTION
NATURE O F T H E STUDY
''
Zonation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Ecotones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
8 . W h i t e Oak-Chestnut Forest . . . . . . . . . . . . . . . . . .
12 . Grassy Bald . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
T h e Southern Appalachian Subalpine
Forest Center . . . . . . . . . . . . . . . . . . . . . . . . . . . .
13 . Red Spruce Forest . . . . . . . . . . . . . . . . . . . . . . . . .
Distributional Relations . . . . . . . . . . . . . . . . . . . . . . .
T h e Xosaic Chart . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Distributions of Types . . . . . . . . . . . . . . . . . . . . .
of Other V o u n t a i n Ranges . . . . . . . . . . . . . . . . .
Suncnca~.
Y ...................................
LITERATURECITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
A P P E N D I X E S. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
A . Population Charts for Major Tree Species . . . . . .
Kote on Supplementary Publication o f Appendixes
B and C . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
f o r a s t u d y o f t h i s vegetation . T h e w o r k w a s origin a l l y intended t o provide i n f o r m a t i o n o n t h e veget a t i o n f o r t h e sake o f i t s o w n interest and as a basis
f o r studies i n animal ecology ( W h i t t a k e r 1952) . A
m a j o r purpose o f b o t h t h i s and t h e preceding s t u d y .
hou-ever. u-as use o f t h e c o m p l e x p a t t e r n o f natural
communities in the Great Smoky Mountains for recomlnunity units o r assosearch into t h e theory
. F~~ t h i s purpose. the approachto vegetatation w a s based o n sampling w i t h o u t regard t o a p parent associations and analysis o f t h e samples i n
ITTAKER
Ecological Monographs
Vol. 26, No. 1
LITERATURE O N AREA
GEOLOGY A N D CLIAXATE
January. 1956
VEGETATION
OF THE GREATSMOKY
MOUNTAINS
FIG.1. Mature, forest-covered topography of the Great Smoky Mountains, a view of the southeast slope
from Frye Mountain near Bryson City, North Carolina . Reproduced by permission of W. M. Cline Co., Chattanooga, Tenn.
VEGETATION
OF THE GRE
January, 1956
/ K* / I I I1 I 111 I IV / I VI / VII
V
Tsuga canadensis.. . . . . . . . . .
Halesia monlieola . . . . .
Aesculus octandra. . . . . . . . . . .
dcer saccharum . . . . . . . . . . . .
Tilio heferophylla. . . . . . . . .
Fagun grandifolio. . . . . . . .
Betula allegheniensis. . . . . . . .
Liriodendron tulipifma. . . . . .
Magnolta acuminata . . . . . . . .
Ilez opaco . . . . . . . . . . . . .
Carye cordiformis . . . . . . . . .
Frazrnus amencana . . . . . . . .
Cladrastts lulea
.......
Magnolio f.asert . . . . . . . .
Acer rubrum. . . . . . . . .
Oslrya virginlona . . . . . . .
Acer pensylwnvum . . . . . . .
Belula lenta . . . . . . . . . . . . .
Hamamelis mrginiona . . . . . . .
Clelh~aacumimta. . . . . . . . . .
Amelanchier arborea. . . . . . . .
Robrnio pseudoam&. . . . . . .
Castanea denhta (dead).. . . . .
Quercus prinus.. . . . . . . . . . . .
Ozydendrum arboreum.. . . . . .
Nwxa xylwlico. . . . . . . . . . . .
Sassafras altndum. . . . . . . . .
Quercus coccineo.. . . . . . . . . .
Pinus pungena. . . . . . . . . . . . .
Pinus rigido.. . . . . . . . . .
Total stems. . . . . . . . . .
*Kalanu Flats, a cove forest 6 mi. east of transect area, elevation 2800 ft.
Ecological Monographs
Vol. 26, No. 1
tion from a factor gradicnt, may be termed a complex-gradier~t (Whittaker 1954b). The "elevation
gradient" is likewise a complex-gradient, involving
many factors of physical environment, soils, and
natural communities other than temperatures and
growing seasons.
The complex-gradient f r o m valley bottoms to dry
slopes will be called the "moisture gradient," but
with no assumption that moisture factors directly
control the distribution of any plant population along
it. Jfeasurernents of all' factors of environment and
determination of which may be most significant f o r
populations of different plant species are f a r beyond
the scope of the present work. F o r the present study
it may suffice that a complex-gradient exists, in relation to which the distributions of plant populations
may be studied.
Such study is dependent on the definition of relative positions along the gradient. Since these could
not be determined, f o r hundreds of site-samples,
by direct environmental measurement, approaches
through the vegetation itself were sought. Along the
gradient the four moisture classes of trees rise a n d
fall in sequence, forming a set of curves flowing
continuously into one another (Figs. 2, 3, 4 ) . I f , as
is s h o ~ v by
~ l the transects, there is progressive shift in
proportions of trees of different tolerances along the
gradient, then it is not unreasonable to turn from
this fact to its converse and regard the same proportions as expressions of position along the gradient.
I n the following discussion, stands and sites will be
termed mesic, sz*bmesic, subxeric, and xeric according
to which of the moisture classes predominate in stem
numbers.
SITE-SAMPLES AND COhlPOSITE TRANSECTS
The main reliance i n solving the vegetation pattern was on the site-samples and their manipulation.
A site-sample was a vegetation sample from a restricted site of uniform physical habitat-the floor of
a valley, a single hillside slope of the same direction
and inclination, o r the crest of a ridge. I n order to
obtain a n approximately random coverage of the
whole vegetation pattern, samples were taken from
the many trails a t all elevations in the mountains.
The method was to move along a trail recording a
sample from each new slope exposure, inside o r out
of a valley, of sufficient extent to give a homogeneous sample. The site-samples were in no case selected to represent either apparent vegetation types
o r the transitions between them. The bulk of the sitesamples were obtained f r o m the mountains surrounding Greenbrier, Sugarland, and Cades Coves in the
Kational P a r k on the Tennessee o r northwest side
of the range.
A t each site the same data were recorded as i n the
transect stations. Sample size varied with the numher of trees thought necessary to indicate stand composition: fifty were sufficient i n some stands with
one or two dominants, but most counts included about
100, while 200 or 300 were tallied f o r some mixed
types. The dense small stems of Rhododendron
thickets were not counted in forests where they occurred. The 25,000 stems recorded in 300 site-samples
were the total sample analyzed f o r the vegetation
pattern.
Exactlng phytosociological analysis of the undergrawth mas not a n objective. Information on shrubs
i s largely limited to presence and stratal dominance,
that on herbs to visible presence a t the time of sampling. Stratal coverages are estimates, intended only
to permit cornparisons between different stands in
the Smokies. A t 1 5 sample stations f o r another study
(Whittaker 1952) location and coverage of individual
plants were mapped in quadrats 10m square.
The site-samples were manipulated in several ways.
By comparing series of them from north and south
slopes o r other exposures, the alterne effect on vegetation could be determined, a method found particularly effective a t high elevations where alterne effects
are more conspicuous in undergrowth than canopy.
Groups of samples from sites w ~ t hsimilar molsture
conditions within limited ranges of elevations were
compiled into composite stand counts. These counts,
usually f o r about 1000 stems, c o m p e n s a t ~ d f o r the
slnall size of the site-sample counts and were used
f o r the characterization of vegetation types ( P a r t
111). Data from the slte-samples were arranged in
mosaic form on a chart with elevation and topographic sites as axes, to show distribution? of species
2nd vegetation types in relation to e l e v ~ t i o n and
topography ( P a r t 111).
The site-samples were, finally, arranged in composite transects in terms of elevation, o r of topographic site, o r of moisture conditions as indicated
hy the vegetation itself. F o r the elevation transects
come means of comparing stands of equivalent moisture conditions a t different elevations was needed.
The site-samples were consequently classified into
f o u r groups, according to which of the moisture
clasies of trees was predonlinant in a given sample.
TT1thin each of the f o u r classes of stands, the sitesamples were grouped by 200- and 300-ft intervals.
F o u r composite transects were thus arranged to cover
thc whole of the vegetation pattern, showing the
change in levels of plant populations from low elevations to hi& in each of the f o u r classes of stands
and sites recognized.
A more sensitive indication of relative positions
along the moisture gradient is possible through the
use of w e i g h t e d averages as indicator values (cf.
Ellenberg 1948, 1950, 1952, Curtis & McIntosh 1951,
w h i t t a k e r 1954b). I n a given stand the number
of stems in each moisture class is multiplied by a
weight ( 0 f o r mesics, 1 f o r submesics, 2 f o r subxerics, 3 f o r xerics), and the total of weig!~ted stem
numbers is divided by the total number of stems.
Within elevation belts (1500-2500, 2500-3500, and
3500-4500 f t ) t,he site-samples were arranged in sequence f r o m most mesic to most xeric by these
weighted averages, and were then grouped f o r tabulation int,o 1 2 or 1 3 steps along the gradient. This
method of ,arranging the transects involves a n evi-
January, 1956
VEGETATION
OF TIIE GREATSMOKY
~~OUNTAINS
signed to form a grid covering the whole of the yegetation pattern of the Great Smoky Mountains. The
following sections will discuss distributions of p l a n t
populations a n d trends in community composition
shown by these transects. The whole body of tables
cannot be published here. Two tables have already
been published ( W h i t t a k e r 1 9 5 1 ) , a n d t h e other
tables f o r tree populations a r e presented here (with
extension of elevation intervals f r o m 200 to 400 f t
i n tables 5 a n d 6 ) . The full set of tables f o r tree
populations and undergrowth species a r e available
to those desiring them (see Note on Supplenientary
Publication)
TABLE2. Composite transect of moisture gradient between 2500 f t and 3500 f t , distribution of trees along gradient. Transect along the moisture gradient from mesic valley sites (Sta. 1 ) to xeric southwest slope sites (Sta. 13),
based on 67 site counts including 6122 stems from elevations between 2500 and 3500 ft. All figures are percentages
of total stenis in station from 1-in. diameter class up.
STATION
N UMBER
Tree species
1
5 6 1
Acer spicatum. . . . . . . . . . . . .
4
Fraxinus americana. . . . . . . . . 2
Tilia heterophylla . . . . . . . . . . . 17
Aesculus octandra. . . . . . . . . . .
7
Fagus grandifolia. . . . . . . . . . .
Acer saccharum. . . . . . . . . . . . .
6
Magnolia acuminata.. . . . . . . .
x
Zlex opaca.. . . . . . . . . . . . . . . . . . .
Prunus serotina.. . . . . . . . . . . . . .
Tsuga canademis. . . . . . . . . . . 25
Betula allegheniensis. . . . . . . . . 26
Liriodendron tulipifera. . . . . . .
2
Halesia monticola. . . . . . . . . . .
5
Magnolia fraseri. . . . . . . . . . .
2
Acer pensylvanicum.. . . . . . . .
1
Betula lenta. . . . . . . . . . . . . .
2
Acer rubrum.. . . . . . . . . . . .
x
Ilex montana. . . . . . . . . . . . . . . .
Cornus flflorida. . . . . . . . . . . . . . .
Hamamelis virginiana. . . . . . . . . .
Ostrya virginiana. . . . . . . . . . . . . .
Carya glahra.. . . . . . . . . . . . .
Clrthra acvminata . . . . . . . . . . . .
Aralia spinosa.. . . . . . . . . . . . . . .
Carya tomentosa.. . . . . . . . . . . . . .
Pyrularia pubera.. . . . . . . . . . . . .
Amelanchier arborea. . . . . . . . . . .
Robinia pseudoacacia. . . . . . . . . .
Oxydendrum arboreum. . . . . . . . .
Quercus prinus.. . . . . . . . . . . . . . .
Sassafras albidum. . . . . . . . . . . . .
Nyssa sylvatica. . . . . . . . . . . . . . .
Quercus velutina. . . . . . . . . . . . . . .
Quercus alba.. . . . . . . . . . . . . . . . .
Quercus coccinea. . . . . . . . . . . . . .
Pinus rigida.. . . . . . . . . . . . . . . . .
Pinus pungem.. . . . . . . . . . . . . . .
Percents b y classes
RIesic. . . . . . . . . . . . . . . . . . . . . 97
Submesic . . . . . . . . . . . . . . . . . .
3
Subxeric . . . . . . . . . . . . . . . . . . . .
Xeric. . . . . . . . . . . . . . . . . . . . .
Trees in s t a t i o n s . . .
.
. 33;
Site-samples used. . . . . . . . . . .
/ / 1 /
:4
59;
671
41;
518
-------9 ~ l O I l l I l 2 I 1 3
I / 1 1 /
62;
35;
3T
42:
43;
41;
554
5
x, Present below . 5 7 .
*Dead chestnut trees were counted in all stands. Since the smaller stems had ceased t o be identifiable as such in 1947, the number of chestnuts in the tables is smaller
than the number of living stems would have been (see size distributions in Appendix C ) .
R. H. WHITTAKER
DISTRIBUTIONS
OF SPECIES
A LOKG THE
MOISTUREGRADIENT
Distributions of tree populations along the moisture gradient are shown in the three tables for different elevation belts (1500-2500 f t , Whittaker 1951,
table 1 ; 2500-3500 and 3500-4500 ft, present work,
tables 2 and 3 ) . Almost all species show a rounded
or bell-shaped curve of population distribution along
the gradient (see Figs. 21 31 4 ) . P o ~ u l ~ t i ocurves
n
for different species, including many of those in dif-
Ecological Monograph6
Vol. 26, No. 1
mazima( ~ ~ ~ ~ h carya
. 1
glabra, ace,.rubrum,
Carya tomentosa, Castalzea dentata, & u e r c m prinus,
ferent
TABLE3. Composite transect of moisture gradient between 3500 and 4500 f t , distribution of trees along gradient.
Transect along the moisture gradient from mesic valley sites (Sta. 1) to xeric southwest slope sites (Sta. 12), based
on 46 site counts including 4906 stems from elevations between 3500 f t and 4500 ft. All figures a r e percentages of
total stems in station from 1-in. diameter class up.
Tree species
--1
Fagus grandijolia. . . . . . . . . . . . . . . . . . 10
Ilex opaca. . . . . . . . . . . . . . . . . . . . . . . . . .
Picea Tubens. . . . . . . . . . . . . . . . . . . . . . . .
1
Cornus alternijolia. . . . . . . . . . . . . . . . .
Aesculusoctand~a. . . . . . . . . . . . . . . . . . 8
Tilia hete~ophylla.. . . . . . . . . . . . . . . . . 29
Acer spicatum. . . . . . . . . . . . . . . . . . . . .
Acersaccharum . . . . . . . . . . . . . . . . . . . . 17
2
Prunus se~otina.. . . . . . . . . . . . . . . . . .
1
Fraxinus americana.. . . . . . . . . . . . . . .
5
Betula allegheniemis. . . . . . . . . . . . . . .
Magnolia acuminala . . . . . . . . . . . . . . . .
Magnolia jraseri. . . . . . . . . . . . . . . . . . .
Tsuga canadensis.. ............
20
5
Halesia monticola.. . . . . . . . . . . . . . . . .
1
Ilex montana.. . . . . . . . . . . . . . . . . . . .
1
Ace~pensylvanicum. . . . . . . . . . . . . . . .
Amelanchie~laeciis. . . . . . . . . . . . . . . . . . .
Quercus borealis. . . . . . . . . . . . . . . . . . . . .
Acer rubrum. . . . . . . . . . . . . . . . . . . . . . . .
Prunus pensylvanica. . . . . . . . . . . . . . . . .
Betula lenta.. . . . . . . . . . . . . . . . . . . . . . .
Clethra acuminata.. . . . . . . . . . . . . . . . . . .
Hamamelis vi~giniana. . . . . . . . . . . . . . . .
Cornus jiorida. . . . . . . . . . . . . . . . . . . . . . .
Li~iodendrontulipijera. . . . . . . . . . . . . . .
Rhododendron calendulaceum . . . . . . . . . .
C a ~ y aglabra . . . . . . . . . . . . . . . . . . . . . . . .
C a ~ y atomentosa.. . . . . . . . . . . . . . . . . . . .
Carya ovalis.. . . . . . . . . . . . . . . . . . . . . . . .
Nyssa sylvatica. . . . . . . . . . . . . . . . . . . . . .
Oxydend~umarboreum.. . . . . . . . . . . . . . .
Cmtanea dentata (dead). . . . . . . . . . . . . .
Sassaj~asalbidum.. . . . . . . . . . . . . . . . . . .
Quercus alba. . . . . . . . . . . . . . . . . . . . . . .
Rolrinia pseudoacacia. . . . . . . . . . . . . . . . .
Quercus prinus. . . . . . . . . . . . . . . . . . . . . .
Quercus velutina. . . . . . . . . . . . . . . . . . . . .
Quercus coccinea.. . . . . . . . . . . . . . . . . . . .
Pinus rigida. . . . . . . . . . . . . . . . . . . . . . . .
Pinus pungem. . . . . . . . . . . . . . . . . . . . . .
Percents by classes
hfesic . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
2
Submesic . . . . . . . . . . . . . . . . . . . . . . . . .
Subxeric . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Xeric. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Treesinstations . . . . . . . . . . . . . . . . . . 377
1
Site-samples used. . . . . . . . . . . . . . . . . .
x, Present below .5%.
NUMBER
STATION
P
-
5
1
x
1
9
11
16
7
1
1
17
s
..
22
8
x
x
x
1
1
4
1
..
1
x
..
..
..
4
9
11
1
10
2
1
..
1
1
1
15
..
..
20
34
4
4
62
1
1
3
x
..
..
..
..
1
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
2
1
..
98
2
..
95
4
1
90
..
..
597
7
520
4
..
..
..
..
4
1
..
..
..
..
1
..
..
..
..
..
..
..
..
..
1
3
1
1
2
x
1
..
..
..
2
..
5
..
..
1
4
..
..
..
1
5
1
2
2
..
..
.
3
..
..
..
..
78
19
2
22
62
16
..
..
..
232
3
449
4
594
9
1
x
1
1
x
3
2
x
10
37
1
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
41
..
..
..
..
..
..
..
..
1
..
..
..
..
..
..
..
..
..
.
..
..
..
..
..
..
..
..
..
..
..
..
..
. .
..
..
..
. .
..
..
..
..
4
21
..
..
..
..
..
..
..
..
..
..
..
..
15
13
11
10
2
8
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
1
1
..
1
..
..
..
2
..
1
..
..
..
..
..
..
..
..
..
..
..
..
..
..
40
6
..
..
..
..
..
x
13
1
3
x
2
1
2
..
1
x
..
..
..
..
..
..
..
..
..
..
..
..
..
x
x
6
14
17
5
x
1
4
x
1
18
9
1
8
x
2
1
..
..
..
..
..
..
..
.
..
..
..
. .
..
..
..
..
..
2
7
1
..
4
..
1
x
..
..
1
4
..
..
..
..
x
2
6
5
2
..
..
..
..
x
...
..
4
'7
2
1
2
1
8 1 4 1 6
3
1
12
7
9
10
1
1
1
4
x
.
x
24
1
8
10
4
5
1
3
8
3
11
4
4
16
15
x
x
l
l
.
.
1
..
..
..
..
11
1
1
..
7
1
54
4
..
..
17
x
..
..
..
..
..
..
. .
..
..
..
..
..
46
49
5
70
23
1
3
44
46
7
1
39
58
2
..
. .
26
69
5
12
23
65
96
472
266
4
369
4
378
4
297
3
355
Ecological Monographs
Vol. 26, No. 1
HITTAKER
ft
Diversity of the tree stratum can best be a p proached through the alplta values of Fisher (Fisher
r t al. 1943 ; Williams 1947, 19.50 ; Whittaker 1952).
January, 1 9 5 6
VEGETATION
OF THE GRE
FIG..5. Pattern of tree species diversities (alpha diversity values, for all tree stems i n the composite stand
counts of Appendix C).
Alpha values cannot be computed f o r the undergrowth data available. Analysis of the transects
through average numbers of species listed per sample
provides a more limited indication of diversity trends
in the herb and shrub strata. F o r shrubs the average
numbers of species recorded in mesic, submesic, subxeric, and xeric stands a r e : 5.2, 7.6, 6.2, 6.6. A submesic maximum corresponding to that f o r trees is
thus suggested; but the shrub stratum in xeric sites
may be more diverse than that in subxeric ones, often
strongly dominated by Kalmia latifolia. Corresponding average numbers of herb species a r e : 19.1, 10.6,
7.1, 8.6. The herb stratum is thus richest in species,
as well as of highest coverage, in mesic sites and
Stature and stem diameter of canopy trees in general decrease along the moisture gradient. I n mesic
sites canopy trees are more than 100 f t high and 3-4
f t or more in diameter, in xeric sites they are mostly
50-75 f t high and 1.0 to l..5 f t in diameter. The number of tree stems per unit area In general increases
along the gradient (cf. Ilvessalo 1921, Lutz 1932),
in inverse relation to tree stature. The cove forests
have mostly between 7.50 and 1000 stems per hectare
from the 1-in. class u p (except in stands of higher
elevations where there are many small stems of Acer
spicatum) , the more xeric stands have mostly 2000
to 2500 stems per hectare. I n p a r t the increase in
stem numbers toward xeric sites reflects the smaller
stature and denser growth of canopy trees; but the
numerous small stems in more xeric sites are predominantly made u p of small-tree species. These
small-tree species (Carpinus carol~nlana,Magnolia
trzpetnla, Ostrya virg~nlana, Ilez opaca; Cornzis
jlortdn, Betzcla lenta, Acer r ~ l b r ~ l mHamamelis
,
vlrginiamn, Cletltra aczcminata, Acer pensylvaniczcm;
Robinia pseudoacacia, Oxydendr~lmarboreum, Sassaf r a s albiclzcm; Q~lerczcsmarilandica) are relatively unimportant in most mesic sites ( a s low as 1-2% of
stems in some cove forests) and most xeric sites
(10-15%). I n submesic and subxeric stands of lower
and middle elevations, however, the small-tree species
comprise around 50% of stem numbers.
Trends in stand composition have been much a f fected by death of the chestnuts (Castanea dentata).
I n many submesic and subxeric stands chestnut
formed 30-60% of the canopy stems, and death of the
chestnuts both removed many of the larqest stems
from the stand and permitted heavy reproduction of
other species. Effects of death of the chestnuts are
most evident in chestnut oak-chestnut forests, in
which maxirriunl numbers of tree stenis per unit area
now occur, and in which 70% of the s t e r ~ ~ins sollie
stands are now of the sniall-tree species.
Trends in tree sizes are illustrated in a family of
curves (Fig. 6 ) , in which steepness of slope reflects
normal survival of small trees into larger size classes.
The more xeric the site, the steeper the curve and
the smaller the proportion of growth and survival
into larger size classes. The oak-chestnut curve is
altered by death of the chestnuts and increased reproduction of other species; the dotted curve is a n
interpolation of what might be expected otherwise.
The hemlock forests are exceptional, for large sizes
are even more heavily represented than in cove hardwoods forests. Fig. 7 indicates the effect of the same
gradient on growth and survival in the populations of
red maples (Acer rubrum).
Curves such as those illustrated in Figs. 6 and 7
ore expressions of the dynamics of stands, the manner in which the tree population is maintaining it-
HITTAKER
Ecological Monographs
Vol. 26, No. 1
7.
b r m in different sites.
" .
...
FIG.
VEGETATION
OF TFIE GREATSMOKY
MOUNTAINS
January, 1956
Acer spicatum. . . . . . . . . . . . . 14
Aesculus octandra. . . . . . . . . . . 11
Betula allegheniensis.. . . . . . . . 10
Acer pemylvanicum. . . . . . . . . . 1
Acer saccharum. . . . . . . . . . . . . 1
Tilia heterophylla. . . . . . . . . . . x
Sorbus americana. . . . . . . . . . . x
Cornus alternifolia. . . . . . . . . . . x
Fraxinus americana. . . . . . . . . x
Amelanchier laevis . . . . . . . . . . . 4
Fagus gandifolia. . . . . . . . . . . 50
Zlez montana. . . . . . . . . . . . . . . . .
Prunus serotina.. . . . . . .
.. . . . .
Halesia monticola. . . . . . . . . . . 2
Quercus borealis.. . . . . . . . . . . . 2
Tsuga canadensis.. . . . . . . . . . . 1
Acer rubrum.. . . . . . . . . . . . . . . 2
Hamamelis wirginiana. . . . . . . . . .
Betula lenta. . . . . . . . . . . . . . . . .
Vaccinium constablaei. . . . . . . . . .
Rhododendron calendulaceum. . . .
Magnolia fraseri. . . . . . . . . . . . . .
Magnolia acuminata.. . . . . . . . . .
Ozydendrum arboreum. , , . , . , . .
Castanea dentata (dead).:....
1
Sassafras albidum. . . . . . . . . . . . .
Quercus alba.. . . . . . . . . . . . . . . . .
Robinia pseudoacacia. . . . . . . . . .
Nyssa sylvatica. . . . . . . . . . . . . . .
Quercu.7 velutina. . . . . . . . . . . . . . .
Prunus pemylvanicn.. . . . . . . . . .
Pinus pungem. . . . . . . . . . . . . . . .
Pinus rigida. . . . . . . . . . . . . . . . . .
Pinus strobus.. . . . . . . . . . . . . . . .
Liriodendron tulipifera., . , . . . . .
Total stems. . . . . . . . . . . . . . . .
Site-samples used. . . . . . . . . . .
-, seedling^ recorded.
13
Ecological Monographs
10
16 20
24
28
32 36
44
2 3 4 6 40
7 8 . 48 52 56
stands, increases through red oak-chestnut to white
Tree species
.
x
oak-chestnut stands and the forest-edge of grassy Fagus grandtfolia.. . . . . 6
1 2 ' 6 1 6
14 23 10
5
8
8
balds, and is low in the grassy balds. Herb coverage Tsuga mnadensis.. . . . 12
Halesia mont~cola . . . . . . 3
12 13 18 30
1
4
increases from the north-slope to the south-slope Prazznus amerimna.. . . x
2
2
1
6
1
1
3 15 15 20 22
4
Lzriodendron tulipifera. . 1
2 4 1 x x . .
lower elevations, herb and shrub coverages are in- Aesculus octandra . . . . . 4
Betula ollegheniensis. . . . 1
5 16 11
8
8 10
4
2 1 2
4
5 1 2
erease from mesic sites into submesic and subxeric Magnolza fraseri. ...... 2
3 5 2 x . .
1 . . . . . . . .
ones, where oaks predominate; evergreen tree species Magnolia tripetola.. ... 1
Magnolia acuminata. . . . . .
1 ..
1 ..
x
1
are almost absent from these forests. I n the shrub Carptnus carolzniana.. . . 10
. . . . . . . . . .
x
3
. . . . . . .
. . . . 1 x x .
Cladrastis luteo. . . . . . . . .
. 4 . . . . .
.
~ 1 4 1 7 3
0
in the oak-chestnut forests.
Prunus se~ottna
. . .
. . . . . x
1
1
1
Amelanchter laenis. . . . .
. . . . . .
DISTRIBUTIONSSPECIES
OF
I N RELATION
TO ELEVATION
MESIC SITES
Cmnus allernifolb.. . . .
Cmnusflmido... . . . . . . . 14
mazima . . . . . . . . 3
Amelanchter arborea. . . . x
BeluIa lenlc . . . . . . . . . 8
Acei pensgdnanicum. . . . 1
Acer rubrum . . . . . . 12
Ilez montona. . . . . . . . . . x
Carua glabra.. . . . . . . . . . 2
Carua lomentosa.. . . . . . . . .
Carua ooalts. . . . . . . . .
x
Quer,usprinw. . . . . . . 2
Nussa sulwrlim . . . . . . . 1
Quercus a l h . . . . . . . . . . 1
Oz~aendrumarbmeum.. . 2
Pinus sfrobus... . . . . . . x
Sassafras a b d u m . . . . . . x
Robinia pseudoacucia . . . . .
Sorbus americana. . . . . . . . . .
. .I
1 1 1 1
4 I
1 1
. . . . . . . . .
.
2 .
x
..
13
3
2
1
1
x
1
2
3
5
6
2
6
1 0 3 2
.. x x
5
1
1
X . . X
. . . : . .
2
2
x
1
1 2
..
. . . .
. . . . . .
. . . . . . . . . .
3
. . . . . . . .
2 . .
. . . . . . . . . . . .
1
x . .
. . . . . . .
. . . . . . . .
x
x . . . . .
. . . . . . . . .
. I
. . . . .
Total stems.. . . . . . . . . ROO 841 518 639 793 429 358 468 646 360 406
I 5 2
S i t e . ~ m ~ l ~ u ~ e d . .2
5
I
Betula allegheniensis, Halesia monticola, Acer saccharztm, Tilia heterophylla, Aesculus octandra, and
Faglis grandifolia ("red" and ((gray" populations).
The decline t o w a d higher elevations of Tsuga canadensis and Magnolia fraseri does not reflect their
true distributions (cf. Appendix A ) , for toward
higher elevations these species are increasingly segregated into hemlock stands which were not included in
the transect. The most significant change in composition of stands occurs a t 4500 f t ; a t this elevation
1500 2d00 2&0
30b0 35b0 4d00 45b0 5000
January. 1956
VEGETATION
OF
THE
GREATSMOKY
MOUNTAINS
Distributions of some submesic tree species ( W h i t taker 1951; table 2 ) are indicated in F i g . 9. Carya
tomentosa is largely restricted to elevations below
2500 f t , Cornus florida and the low-elevation population of Carya glabra (see Appendix A ) to elevations below 3000 f t . The most significant change in
the tree stratum occurs in the elevations between
3500 and 4000 f t , a s Querczis borealis Michx. f . increases to become the major submesic tree a t higher
elevations. Some of the major shrubs of suhmesic
sites (Rhododendron m a x i m u m , Clethra acuminata,
Gaylussacia ursina, Pyrularia pubera, V i b u r n u m
acerifoli~tm, Calycanthus fertilis) have their u p p e r
Limits of distribution a t elevations of 3500-4500 ft.
Rhodorlendron calendulaceum and Vaccinium constablaei extend to elevations above 5000 ft. Polystichum acrostichoides, Aureolaria laevigata, and
other major submesic herb species are confined to
lower and nriddle elevations, hut some (Smilacina
I5
40-
40-.
50-
ELEVATION IN
FEET
One population of Qzurcus alba is largely restricted to elevations below 2500 f t , the other t o
elevations above 3500 f t (Table 6 , Fig. 9 ) . Quercus
prinzls is centered a t lower elevations and has its
normal upper limit below 4500 f t . Castanea dentata
extends throughout the elevation range of the transect. At elevatlons below 3500-4000 f t Quercus plinus and Castanea dentata are dominant subxeric
trees; above these elevations Castartea dentata shares
dominance with the high-elevation populations of
Quercus alba and Q . borealis.
Kalmia latifolia is the dominant shrub In the oakchestnut heaths from lowest elevations to about 3500
f t ; Rhododewdrofz m a x i m u m and Gayltusacia ursina
also occur. A t elevations above 2500 f t Rhoclodendron calendulaceum and Vaccinium constablaei a r e
important shrubs; a t elevatlons above 3500 f t Kalmia
lntifolia shares dominance with these until Kalmia
becomes a minor species in forests above 4000 f t .
Galax aphylla is the most important subxeric herb a t
all elevations. Chimaphila maculata, Campanula
divaricata, the submesic Aureolaria laevigata, and
xeric Epigaea repens are major herb species in subx e r ~ csltes; and all extend through most of the elevations represented in the transect (1500-5000 f t ) .
XERIC SITES
Ecological M o n o g r a p h s
Vol. 26. No. 1
R. H. WHITTAKER
16
ELEVATION IN
FEET
TRENDSI N RELATION
TO ELEVATION
GROWTH-FORMS
No trends i n growth-form composition of communities as striking as those along the moisture gradient a p p e a r along the elevation gradient. Viewing
the vegetation pattern a s a whole, three growthforms-abietine
trees, ericaceous shrubs, a n d ground
heaths-are
of increasing importance toward higher
elevations. Among the abietine trees, Tsuga canadensis is of increasing importance from low elevations to about 4500 f t ; and Picea rubens and Abies
fraseri dominate most forest stands above that elevation in the northeast half of the range. The great
development of ericaceous shrub communities is one
of the most characteristic features of the vegetation
I.
Station .... l*
Tree species
Tsuoa canadensts . . . .
1
1.triodendron tulipl jna . . . . . 1
Halesia monticola. . . . . . .
Mapnolia fraseri . . . . . . . . . .
Acer sacchrum. . . . . . . . . . . .
Mapnolia aeuminata.. . . . ... .
Iler opaca.. . . . . . . . . . . . .
Betula allegheniensis . . . . . . .
Amelanchier laevis. . . . . . . . . . .
Acer rubrum.. . . . . . . . . . . . 20
A c ~ rpensyluanicum . . . . . . . .
Cornus florida. . . . . . . . . . . 5
dmelanchter arborea.. . . . . . . x
Retula lenla. . . . . . . . . . . . x
Carya glabra. . . . . . . . . . 4
Carya tomentosa. . . . . . . . . 3
Carsa ouahs.. . . . . . . . . . . .
Clethra acuminata.. . . . . .
ller moninna. . . . . . . . .
dralta sptnosa.. . . . . . . . . . .
Nyssa sylmtica. . . . . . . . . . . . 1
Orydendrum arborpum. . . . 15
Quercus prinus. . . . . . . . . . . . 24
Quercua nelutina. . . . . . . . . . . 4
Quercua alba.. . . . . . . . . . . . . . . 5
Quercus stellato. . . . . . . . . . .
Sassafras a h d u m . . . . . . . . . . . 1
M n i a peudnacneia.. . . . . . . 1
Castanea dentata ( d e a d ) .. . . . 5
Pinus strobus. . . . . . . . . . . . . .
Q w c u s mccinea. . . . . . . . . . . 4
Pinw airpiniana . . . . . . . . . . 2
Pinus ripido . . . . . . . . . . . .
Pinus pungens . . . . . . . . . . . .
Total stems.. . . . . . . . . . . . . .
Site-samples used. . . . . . . . . .
January, 1956
VEGETATION
OF THE GREATSMOKY
MOUNTAINS
17
TABLE7. Composite elevation transect in xeric sites, distribution of trees. All figures are percentages of total
stems in station from 1-in. diameter class up.
Station . . . .
Elevation, hundred feet. . . .
l*
14
2
17
3
20
--- --- --
Tree species
Tsuga canadensis. . . . . . . . . . . . . . . . . .
2
Liriodendron tulipifera. . . . . . . . . . .
Liquidambar styraciflua. . . . . . . . . . .
x
Amelamhier laeztis . . . . . . . . . . . . . . . .
Cornus florida.. . . . . . . . . . . . . . . . . .
x
Quercus borealis. . . . . . . . . . . . . . . . . . .
Carya glabra.. . . . . . . . . . . . . . .
x
Carya tomentosa.. . . . . . . . . . . . . . . . . . 1
Carya ovalis.. . . . . . . . . . . . . . . . . . .
Hamamelis virginiana . . . . . . . . . . . . .
Acerrubrum . . . . . . . . . . . . . . .
6
Quercus falcata. . . . . . . . . . . . .
1
Stewartia ovata.. . . . . . . . . . . .
x
Pinus strobus. . . . . . . . . . . . . . . . . . . .
Robinia pseudoacacia. . . . . . . . . . . . .
Sassafras albidum.. . . . . . . . . .
1
Nyssasyluatica . . . . . . . . . . . . . . . . . . 4
Ozydendrum arboreum.. . . . . . . . . . . . . 8
Castanea dentata (dead). . . . . . . . . . . . 5
Quercus velutina.
. . . . . . . . . . . . . . . .
Quercus a l b a . . . . . . . . . . . . . . . . . . . . .
1
Quercusprinus . . . . . . . . . . . . . . . . . . x
Quercus marilandica. . . . . . . . . . . . . . . . 7
Quercuscoccinea . . . . . . . . . . . . . . . . . 10
Pinus rrirginiana.. . . . . . . . . . . . . . . . . 40
Pinus rigida. . . . . . . . . . . . . . . . . . . . 13
Pinus pungens. . . . . . . . . . . . . . . . . . . .
..
..
..
..
..
. .
..
.
4
23
5
26
6
29
7
32
8
35
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
14
..
..
..
..
..
..
10
..
..
..
..
..
..
..
..
..
..
..
12
56
12
"
21:
..
..
..
2
x
..
4
6
~
2
..
..
2
8
2
48
10
2
2
8
38
30
..
..
11
44
12
47
..
..
..
1 :. .
..
..
..
. .
..
. .
..
..
..
..
1
..
..
14
..
26
1
..
..
..
..
..
..
..
..
..
10
..
..
..
..
..
3
2
1
..
..
..
14
1
..
..
..
..
1
3
4
3
..
..
2
9
..
..
.
..
5
1
2
3
x
10
41
..
x
9
38
..
..
x
2
2
1
2
..
..
..
. .
1
9
2
3
..
..
..
1
x
3
2
1
..
..
. .
..
14
. . -
..
..
..
..
8:
57;
..
..
..
4
2
3
2
..
..
. .
..
..
..
..
11
11
..
..
..
9
..
..
. .
..
..
..
'7
/ 1 1 1 1
/ 1
36:
..
..
..
..
..
..
16
62
10
55
"
21;
..
. ..
. .
..
..
..
37;
41:
24;
36;
12:
..
87
90
s overage^:
R. H. WHITTAKER
18
FAVORABI,EIE,?ESS
EcologicalVol.
Monographs
2 6 . No. 1
borealis.
January, 1956
VEGETATIOX
OF
THE
GREATSMOKYMOUXTAINS
19
TABLE8. Transect data for subalpine forests. Percentages of coverage or stems for stratal groupings in different
sites along the moisture gradient i n five steps: 1, most mesic valley and sheltered-slope sites; 2, open north-facing
slopes; 3, east and west slopes; 4, south-facing slopes; and 5, ridges. Figures are based on averages from several sitesamples of stem numbers and estimated coverages.
I
Elevation belt. . . .
REDSPRUCE
FORESTS SPRUCE-FIR
FORESTS FRASER
FIR FORESTJ
5600-6200 ft.
454B5500 ft.
above 6200 it.
---
site . . . . . . . . . . . . .
--
1 / 2 / 3 / 4 / 5
'Not observed.
I n the tree stratum composition of subalpine forests changes more conspicuously with elevation than
along the moisture gradient. At 5000 f t on south-facing slopes 100% of the conifers may be spruce (Picea
rubens); above 6300 f t 100% may be fir (Abies
fraseri), while intermediate sltes have mixed stands.
Spruce and fir cross in numbers of stems a t around
5600 f t if exposure differences are averaged out.
The change in canopy dominance occurs higher, near
6000 f t , because of the greater size of spruce. Elevation alone, however, does not determine proportions in the mixture of these two dominants; a t any
elevation fir increases relative to spruce toward more
mesic sites (Table 8 ) .
Sorbus americana is confined to high elevations
and is centered in the more mesic forest stands above
6200 f t . Fagus grandzfola ("gray" population)
dominates stands in the concave slopes of "gapi" in
the area of spruce-fir forests, and it occurs also to a
limited extent in stands dominated by spruce and fir.
All other deciduous tree species which occur in the
spruce-fir forests are a t least equally important in
deciduous stands below 4500 f t . The high-elevation
population of yellow birch (Betula lutea) is centered
in the transition from cove forests to spruce forests
a t 4500 ft, but is of wide extent a t higher elevations
as the major deciduous tree species of spruce-fir
forests. Yellow birch and other "ecotonal-mesic"
species (Acer spicatum, Amelanchier laecis, Cornus
alternifolia) decrease in importance with departure
from their centers in mesic stands near 4500 f t to-
ward higher elevations and less mesic sites. The coveforest tree species which occur in subalpine forests
(Aesculus octaadra, Tilia heterophylla, Halesia rnonticola, Acer saccharurn, Fraxinus arnerzcana, and the
submesic Acer pensylranicurn) are similarly related
to the gradients; their numbers decrease toward
higher elevations and more xeric sites.
Three shrub unions with different distributional relations have been distinguished in the spruce-fir forests. The distributional relation of the deciduous
high shrubs of the ecotonal-mesic union is the same
as that of Betula lutea and Acer spicaturn; their importance declines with departure from mesic stands
near 4500 f t . The shrub species of the group differ
in extent and importance in the spruce-fir forest pattern; Viburnum alnifolurn is the most wide-spread
and much the most important of them. The low-shrub
union dominated. by Vacciniurn erythrocarpurn is
centered in middle elevations of the spruce-fir forests.
rather than lower ones, and in mesic sites (northfacing slopes and flats) other than the valley sites
where the ecotonal-mesic species are most important.
The high heath union is most important on ridges,
hut the heath species also occur with lower coverage
in south-slope stands and on some intermediate
slopes. Of these shrub species Vacciniurn constablaei
occurs more widely in spruce-fir forests than the
heath dominants Rhododendron catazclbiense and R.
carolinianum.
I n valley sites the mesic herbs strongly dominate
the herb stratum. Most of the herb species listed
are of wide extent in spruce-fir forests (other than
those of most xeric sites), but their coverage is lol~.
outside valleys. The moss stratum, the low-herb
stratum, and the fern stratum are, in contrast, centered in stands of north slopes and flats. These three
unions also are of wide extent through the spruce-
Ecological Monographs
Vol. 26, No. 1
R. H. WHITTAKER
20
in contrast with the pattern of forests a t lower elevations. Deciduous trees decrease in importance from
valleys toward xeric sites; ericaceous shrubs increase
in importance from intermediate sites to ridges.
Herb coverage and species diversity decrease along
the moisture gradient. There is thus a trend of decrezsing importance of non-evergreen elements in all
three strata toward more xeric sites. Valley sprucefir forests resemble high-elevation cove forests in
most respects, even though the canopy is dominated
by conifers. At the xeric extreme, however, evergreen elenlents are strongly dominant both in canopy
and in the dense ericaceous shrub stratum, and ground
heaths predominate in the sparse herb stratum.
Along the elevation gradient within the spruce-fir
forests, moss coverage increases strongly from lower
elevations toward higher ones (Table 8 ) . Coverage
of shrubs other than the heath is in general lower at
highest elevations, but undergrowth coverage trends
with elevation are not otherwise conspicuous. Representation of deciduous trees other than Sorbzis americana decreases toward higher elevations, and (since
Sorbus is a small-tree species) the fraction of deciduous trees in the canopy decreases toward higher elevations. Species diversity of the tree stratum decreases toward higher elevations, and the same is
probably true of other strata of vascular plants.
The most conspicuous change in canopy of subalpine forests is the increase of fir and decrease of
ipruce toward higher elevations, but the most con-
SPWCE
- FIR
BOREAL
FORESTS
5-STORY
UNDERGROWTH
UNDERGROWTH
SPARSE
UMRCRMll-M
EASTERN FOREST
FOREST
HEMLOCK
SYSTEM
3000 FEET
OAK-CtESTNT
0L)cCWISTNT
FOREST
HEATH
S.W.,UPPER
S L O P E S
KEY:
9
4
?
"LO
(fl-X
W
ClS7WT,DLAO
WE-*
uubc 8-*r
LYLIVILLN
4/.
VYCIMWY
TRUER n R
CUTER.
TMO
T * m
P I E WIATH
mATW
ClOUD K I M
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YY FERNS
ulvuv
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oxut)
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January, 1956
VEGETATION
OF THE GR
22
R. H. WHITTAKER
((natural" in the sense that they are clearly determined by the data in his possession. I f species are
quite variously distributed, forming no well-defined
group. distinctly separate f r o m other groups, the
groupings recognized are free or "arbitrary." They
are not clearly determined by the data, but represent one anlong a series of groupings which might
be rccognized without violating the more evident distributional relations of species in the data. Relative
arbitrariness and naturalness of species groupings
may thus depend on the degree to which the ecologist is clea1.1y guided and limited in his choice of
groupings by actual distributional relations of species in the field and in adequately representative
saiirple data.
The basis of groupings in the present study is the
loration of centers of maximum population level f o r
species along gradients, and the problem of naturalness 1.r. arbitrariness of groupings may be further
definrd on this basis. Various manners in which species population centers might be related to one another may he conceived in terms of degrees of clustering along a hypothetical gradient from 0 to 1 0 : (1)
Species populations are absolutely clustered, so that
species of one group are centered a t 2.0 and those of
another a t 8.0, and none are centered het~veenthese
points. ( 2 ) Species populations are relatively clustered along the gradient, so that in the sequence of
their centers along the gradient (1.3, 1.6, 2.0, 2.3, 3.5,
6.0, 7.2, 7.5, 8.1, 9.0) groupings niay be recognized,
even though the centers are somewhat scattered and
so~l!cspecies arp intermediate. ( 3 ) Species are distributed irregularly along the gradient (0.9, 1.4. 2.1, 3.5,
3.2, 5.5, 6.0, 7.3, 8.1, 9 . 0 ) , and thc spacing of their
centers approaches randon~ness. (3) Distribution of
centers along the gradient is less irregular than randonlness would require, and the centers are approxiiiiately equidistant froin one another (1.0, 2.0, 3.0,
4.0. . . .).
The problen~ of relative "naturalnesh" of species
groupings map thus be approached through degrees
of clustering suggestive of the degrees of "contapion" o r clustc~ringof individuals in a stand. Condition ( 1 ) would imply that associations of species
are "natural" groupings (though community-types
might still be continuous with one another). Condition ( 2 ) would define varying degrees of relative
('naturalness" 1.8. relative '(arbitrariness" of groupings which would not necessarily represent either
obligate associations of species or discrete community-types. Groupings of species distributed according to conditions ( 3 ) and ( 4 ) would be primarily
arbitrary. The transect data and nomograms ( A p pendix A ) apparently exclude conditions ( 1 ) and
( 3 ) f o r the distributions of species in the Smokies.
The data show no evidence of high degree of clusteririg according to condition (2). S o sensitive statistical tests are available a t present to distinguish con(!ition ( 3 ) from low degrees of clustering in condition ( 2 ) and slight departure from randomness toward the regularity of condition (4). The impres-
Ecological Monographs
Vol. 26, No. 1
J a n u a r y , 1956
VEGETATION
OF THE GR
D O U I S B N C E I N RELATION TO C O M M U N I T Y COMPOSITION
HITTAKER
Ecological Monographs
Vol. 26, No. 1
~ t e dand homogeneous commun~ty-unitj. These state7. Picea rubens, dominating suhalpine forests bements may imply (5) that, in general, com~nunity- tween 4500 and 6000 f t .
8. A b i e s fraseri and S o r b u s americana, centered in
units are more "arbitrary" products of classification
than "natural" units clearly defined in the field. suhalpine forests of highest elevations.
Thls possibility must be f u r t h e r considered in relaI I . S h r u b strata
tion to other problems of ecological theory and loglc
1.
Low-elevation
mesic shruh union : E n o n y m tts
in following sections ( P a r t 11).
americanus and L i n d e r a benzoin.
SVMMARY
O F DISTRIBUTIOXAL
GROUPIXGS
2. Mesic heath union : R h o d o d e n d r o n ma.cim ton
Dlstrihutional grouplngs of plant specles will he a n d Leucotho? editorum.
listed to summarize some results of the gradient anal3. H y d r a n g e a arborescens, widespread i n mesic
yst\ and reduce r e p e t i t ~ o nof specles l ~ s t sIn type de- and suhmesic sites.
s c r ~ p t i o n s ( P a r t 111). The concept~onon w h ~ c hthe
4. Suhmesic shrub union, identical with suhmesic
grouplngs a r e based is that of the "commod~um" (see tree unlon listed ahove but also including shrubs:
P a r t 11) Species a r e grouped together whose modes Pyrularict pubera, R h o d o d e n d r o n calendulaceztm, G a y o r distribut~onalcenters are relatirely close together lussacia ursina, V i b t ~ r n u m acerifolium, Calycanthzts
In relation to environmental gradients. Specles thus ferttlis and f . nanzcs, C e a n o t h u s americanus, Aralia
grouped tend to occur together In many stands; hut spinosa, and S m i l a x r o t l ~ n d i f o l i a .
5. Suhxeric heath union: K a l m i a latifolia with
they I\-111 also occur separately in other standi, slnce
IIO t n o have the same d i s t r ~ h u t ~ o n sF o r reasons alI y o n i a llgztstrina, P a c c i n i u m constablaet, and S m i l a x
ready indicated, the grouplngs a r e conridered pn- glauca
marlly arhitrary In the sense that the number iecog6. Xeric shrub union, with low Vaccinioideae
nlzed, their l ~ m ~ and
t s r e l a t ~ o n sto the gradients, a l e dominant : Vac.c.intum twcillans (including P . pallideterrnlned by suhjectlve cholce and convenience The drcm), T7. Itirslttttm, P . s t a m i n e u m (including V .
grouplngs will be termed ztnions and ~ 1 1 1In most crtndzcan\ ( C . l f o h r ) Sleumer and T'. melanocarcases he named f o r enviromnental r e l a t ~ o n s rather p u m ) , Gaylussacia baccata, Pieris floribunda, I1r.r
thali major species Herh-layer grouplngs nlaj he m o n t a n a v. beadlei.
considered also groups of unlons, slnce specles are
7. Ecotonal-mesic shruh union, i d e n t ~ c a lmith econot separated according to 11fe-forn~s.
tonal-meslc unlon l ~ s t e d ahove hut also lncludlng
ihruhs : V t b u r n t t m alnzfoltum, Sambuczts pubens,
I T r e e strata
R t b e s c!/nosbatt
1. J I e s ~ ctree class o r unlon, centered In cove for8 Lon-shrub unlon of subalplne forests T'acegts helon 4500 f t Ttlza h ~ t e r o p h v l l a ( ~ n c l u d l n gTar. cznitcm c r y t h r o c a r p u m , lllenziesia ptlosa, Dterrilla
mzchauxzz and T t r u n c a t a Spach ) , Aesculus oc- 9esszltfolta.
t a n d r a , Halecta monttcola, B e t z ~ l a alleghenzensis,
9 Loner-elevat~onheath hald shrub unlon. Ii'alT s u g a canadensts, A c e r saccharum, L i r t o d e n d r o n mzo ltttifolta, Pibztrnum cassinoides, A r o n i a m r l a n o tzcltpzfera, F a g u s grandtfolta ("white" and "red"
carpa
populations), F r a x z n u s amerzcana and v. bzltmore10. H ~ g h - e l e ~ a t ~heath
on
hald shrub unlon: Riioa n a , Magnolza aczcmtnata, M fraserz, M . trzpetala, r7or7enrlron catclrtbtrnse, R. caroltntanum, L e t o p h ~ l l Cladrastzs lntea, P r u n u s serottna, C a r y a cordzformis, lltm lr/alrt S x eet.
I l e x opaca, C a r p t n u s caroltnzana, O s t r y a I3trgtntana.
I I I . H e r b strata
2. Submesic tree class o r unlon, centered In oak1. Mesic herb union: D r y o p t e r i s spinulosa v. i n hlckory and oak-chestnut forerts . Quercus borealzs
and r. m a r z m a , Carva glabra, C . tomentosa, C. 01-ctlts, terinerlia, 9 t h t j r t u m thel!/pterioides, Eupatoritcm
A c e r r u b r u m , B e t u l a lenta, B e e r p e n s y l ~ a n z c u m , rttgosztm, Ctnztctfuga racemosa, I m p a t t e n s pallida,
C o ~ n z t s flortdn, Hamamelzs ~ % i r g t n z a n aA, m e l a n c h t e r Cuulophylltcm thalictrotdes, L a p o r t e a canadensis, Trzll t u m erectum f . albtflorum, A s t e r d t ~ a r t c a t u s ,A d t a n arborra, I l e x m o n t a n a , Clethra acumznata.
3 S u h x e r ~ ctree class or unlon, centered In oak- t u m p e d a t u m , A c t a e a pachypoda, T h a l i c t r u m dzoicum,
chestnut heath and drier oak-chestnut forests. Cas- Hepatzca acutiloba, V i o l a rotundifolia and s p p ,
tanea dentata, Qurrcus przntts ( Q m o n t n n a W ~ l l d), Stellaria pubera, Ttarella cordzfolza, Mztella d t p h y l l a ,
Q. alba, Q ~ e l u t t n a ,N y s s a s t / l ~ a t z c a , O x y d e n d r u m Etconymus oboratus, Galium trtflorum, Mitchella renrboreztm, Pztzus strobus, Robznza pseudoacacta, S a s - pens, Osmorhiza longistylis.
2. lfesic high-elevation herh union. Rudbecliza
safrac albzdum.
laciniata, Circaea a l p i n a , D r y o p t e r i s spinulosa r.
4. X e r ~ ctree class o r unlon, centered In plne forests
and plne heaths. Pztzus t'zrgznzana, P rtgtda, P . amerzcana, Allzum trzcoccum, S o l i d a g o glomerata,
Chelone lyont, Clintonia borealis, Dzphylleia cymosa,
pungc2ns, Quercus coccznea, Q martlandzca.
5. F a g n s grandtfolta ("gray" population) In meslc Ltlizcm s u p e r b u m , Ocalis m o n t a n a , Senecio rugelia,
lllontrrda d t d y m a , A s t e r acumznatus, S t r e p t o p u s rodeciduous stands ahove 4500 ft.
6. Ecotonal-mes~cunlon, centered In meslc s ~ t e sa t sezts, Prtlltzcm erectum.
3. Submesic herh union: P o l y s t i c h u m acrostzelevat~onsaround 4500 f t : B e t u l a lutea, A c e r spzchozdes, Smtlacina racemosa, Ptola hastata, A u r e o c a t u m , Amelanchzer l a e ~ z s ,Corntts alternifolta
January, 1956
VEGETATION
OF T I ~ EGREATSMOKY
~~OUNTAINS
11. LIISCTSSION: A N I X T E R P R E T A T I O T O F
VEGETATIOS PATTERXIKG
DISTRIRUTIOXS
O F SPECIES4 N D T H E STVDYO F
G~xec.or.oclI n P a r t I a number of curves f o r distributions of
plant populations along environmental gradients were
~llustrated Such cur\es are doubtless of fundamental
significance In ecology, c l a r ~ f y l n gas they do the relatlons of specles populations to one another, to h a b ~ tats, and to community-types. Real understandmy of
phenomena of p o p u l a t ~ o n d y n a m ~ c s w h ~ c h underlle
these curves must a l v a ~ t future research. Present
knowledge of the genetics of species populations
mag, however, permit a t least partial interpretation
of some distributions ohserved in the Great Smoky
Mountains. F o r much of the genetic information used
in this discussion the author is indebted to IT. H.
Camp, who has given the author the benefit of his
experience with plant populations in the Smokies
through personal communication a s well as published
work ( C a m p 1945, 1951; Camp 8: Gilly 1943).
An ideal situation may first be imagined: a n unbroken complex-gradient of environment along which
a population consisting of a single hiotype is distrihuted and is maintaining itself. A maxinium population level should occur a t t h a t point of the gradient
which represent,^ optimum conditions f o r the biotype,
and the number of individuals should decrease with
increasing departure from the optimum in either direction. The curve of distrihution f o r this population
would be essentially a curve of probability of sur-
25
HITTAKER
Ecological Monographs
Vol. 26, No. 1
January, 1956
VEGETATION
OF
THE
GREATSMOKY
~IOUNTAIXS
27
28
R. H. WHITTAKER
(Huxley 1938, 1940, 1943) through series of p a r tially or wholly discontinuous populations. Cline
is customarily applied to the populations through
which a genetic trend exists as well as to the trend
itself. Along a segmented cline ecotypes may be
recognized as relatively discontinuous populationtypes, marked by a distinct population mode, a t least.
The term ecotypic populatzon is conveniently a p p l ~ e d
to a population-type which 1s recognized as distinct ~ v ein its adaptation to habitat, when i t is not known
whether this is a clznal populatzor, continuous with
other populations along a cline, or a n ecotype relatively discontinuous with other ecotypes.
Because of the separation of populations into local
habitats not arranged along gradients, and because of
the possibility of genetic d r i f t within local populations, these will not always form clines. I n some species there may be much ecotypic variation which cannot be reduced to clines and related to environmental
gradients. I t may further be observed that ecotypes
occur within ecotypes (e.g., edaphic within climatic
ecotypes, Zherebina 1931, Kruckeberg 1951, 19541,
that ecotypes occur along clines and clines within
ecotypes, that clines nlay cross one another in various
ways, and that rninor clines may occur within seg111ents of 11~ajorones. Ecotypes and clines, a s defined
here, occur in all possible combinations. Clines and
ecotypes a r e thus alternative and complementary
n e a n s of abstrar4ing fro111 the ecotypic variation of
species, emphasizing relative continuity and relative
discontinuity, respectively, applicable according to
the particular cornbination of continuity and discont ~ n u i t e~x i s t ~ n gIn a given species population (cf.
S t e b b ~ n s1950, Whittaker 1954b).
Both c l ~ n e sand ecotypes may be thought funda~nentallgphysiological ( i n a broad sense) in meaning
though sorliet~~nesrecognizable also in n3orphology.
The ecotype may be thought to represent a relatively
favorable genetic combination within the species, a n
adaptive peak in relation to environment which has
actually heen attained. I t seems likely that such
favorable genetic cornbinations are usually expressed
in local maxirna of population density, in population
peaks such a s a r e illustrated in Appendix A. The
occurrence of clines and the ridge-like shape of some
of the population figures in Appendix A (granting
that the relative lengths of the ordinate and abscissa
are wholly arbitrary) suggest that adaptive "peaks"
may take the form of "ridges" i n relation to combinations of gradients. The genetic pattern and the
more abstract adaptive landscape of a complex species, or of a genus o r compariuln, may be visualized
a s a complex topography of hills, peaks, and ridges
of different heights and extents, variously related to
one another and variously separated by cols, ravines,
and level valley bottoms of different depths and
widths.
The role of introgressive hybridization i n modifyi n g the genetic pattern of species has been referred
to. An introgressive cline suggestive of that i n the
northern red oaks appears arnong the blueberries,
Ecological Monographs
Vol. 26, No. 1
January, 1956
VEGETATION
OF
THE
GREATSMOKY~IOUNTAINS
29
Ecological Monographs
Vol. 26, No. 1
One of the most nearly universal ideas among different schools of ecology is that of vegetation units,
which have most commonly been termed associations.
Some representative statements on associations from
different schools are the following: Flahault &
Schroter (1910 :152, Pavillard 1935), "An association is a plant community of definite floristic composition, presenting a uniform physiognomy, and groming in uniform habitat conditions. The association
is the fundamental unit of synecology." BraunBlanquet (1921), "The association is a plant community, characterized by certain floristic and sociological marks, mhich reveals a certain independence
through the presence of character-species." Nichols
(1923), "Viewed in the abstract, an association may
be described as a vegetation-unit characterized by its
essentially constant physiognomy and ecological
structure and by its essentially constant floristic composition, a t least with regard to dominant species."
Conard (1939), "The association of individuals and
species is much more than a chance meeting. I t is
part of the order of nature. . . . Associations of
plants on land are definable entities, susceptible of
naming and classifying." Du Rietz (1929), ". . . the
association is a concrete part of actual vegetation,
though mostly split u p into a large number of isolated parts. I am quite unable to regard each of
these isolated parts as a fundamental unit of vegetation analogous to the individuals, the real fundamental unit being the sum of all the parts showing
essentially the same composition." Tansley (1920),
"But if me admit, as everyone who has worked at
the subject does admit, that vegetation forms natural
units which have an individuality of their own, and
that these units owe their existence to the interaction
of individual plants of different species with their
environment, then it becomes clear that a mere study
of the distribution of species as specirs cannot form
the basis of the science of vegetation. We have instead to focus our attention on the vegetational units
themselves."
The central idea of these conceptions of vegetation
may be termed the association-unit theory, the belief
that vegetation consists of distinct, natural units
called associations. With the association, so f a r as
basic philosophy is concerned, may be grouped such
other units as the formation or biome, sociation,
union and socion, site-type, life-zone, etc. We may
regard the association-unit theory as a form of a
broader community-unit theory, which assumes that
natural communities are made up of units and that
these units are themselves the proper objects of
study. Like the cell theory and the molecular theory,
the community-unit theory is a conception of the
structure and organization of the phenomena with
which the scientist deals. Whatever the present
status of the community-unit theory, for its role in
the development of ecology and for its influence
January, 1 9 5 6
VEGETATION
O F T l I E GREATSMOKY
~IOUNTAINS
31
32
R. H. WHITTAKER
EcologicalVol.
Monographs
26, No. 1
January. 1956
VEGETATION
OF THE G
Ecological Monographs
Tol. 26, S o . 1
FIG. 13. Distributioris of plant populatioris in relation to "zones." Above-the transition from core forests below 4500 f t to mesic spruce-fir forests above,
)lased on coniposite transect data. Zoned distribution of
evergreen and deciduous growth-forms relative to one
another: a, all deciduous, broad-leaved trees; b, all evergreen, needle-leaved trees ( P i c e a rubelis, A b i e s fraseri,
a ~ i d ,below 4i00 feet, T s u g a canadensis) ; c, Picea r u b a n s : d, A b i e s frascri. Distrib~itionof other species in
relation to the transition and spruce-fir forests: e and e',
Retula allegheniensis and B. l u t e a ; f , Acer spicaturn:
g, C o r ~ i ~ lalternifolia;
s
h, Amelancltier laevis; i, V i bitrnunt alntfolium (estimated coverage percents) ; j,
Sorbus americana. Belon-the
transition from core
forests below 4500 f t to gray beeeli forests above, based
or1 a field t r n ~ ~ s e eint Trillium Gap. Zoned distribution
of gray beeeli relatire to otlier tree species: a , major
mesic tree sl~ecies ( H a l e s i a monticola, Retrrla allegheni-
elisis, Aesculus octandra, Tsrrga canadensis, Acer saccharunt, T i l i a heterophylla, Frasinils americana, Liriodendron t u l i p i f e r a ) ; b, red, arid c, gray populatio~lsof
beech ( F a g u s g r a n d i f o l i a ) . Distribution of otlier spe: Betilla alleghcnicies in relation to the t r a ~ ~ s i t i o nd,
ensin; e, Cornus alternifolia; f , A~ncrrlrts o c t a n d r a ; g,
A C P T s p i c a t u ~ n ;h, P r u n u s nr>rofina.
tioh ( h c ~ ~ halds),
th
and coniferous forests (spruce-fir
zone). The g r a y beech is a deciduous tree, but differs
froin many of the cove forest trees in its sniall stature
a n d nlanner of reproduction. A t a contact between
tlift'erent doininant growth-forms, such as t h a t of the
grassy halds and forests, e n v i r o n ~ i ~ e n t aconditions
l
relatetl to don~inanceof one growth-fonn iilay largely
prevent individuals of t h r other fro111 establishing
thc~~:sc,lves.The two dorninant growth-forms inay
thus, in some cases, replace one another ahruptly
stands along a wide s p a n
rather than f o r l ~ ~ i nniixed
g
of t h r gradient. Because stands of n ~ i x e ddoininanctx
a r c liinited, the two g r o w t h - f o r ~ i ~asp p e a r , in hulllan
interpretation, relatively '(inco~npatihle."
Tlics relation of the rclatively discontinuous typtbs
to the gradients i.: of interest. All art. types of high
elt.rationi; and the t\vo I\-hich a r c il~ost clearly tlis-
January, 1956
VEGETATIOK
O F T I l E GREAT
SMOKY
;\IOUNTAIXS
35
36
R. H. WHITTAKER
Ecological Monographs
Vol. 2 6 , No. 1
January, 1 9 5 6
VEGETATIOK
OF THE GREATSMOKY
MOUXTAINS
cornrnodiu~n of species centered in or near the ecotone is illustrated for the transition from cove forest
to rnesic spruce forests (Fig. 13) ; and a related cornmodiurn appears between cove forest and gray beech
forest. Several woody species appear in the forestedge between deciduous forests rind grassy balds. Ko
such grouping appears between subalpine forest and
heath bald, although S o r b u s americana may have an
ecotonal mode there. These ecotones are characterized
by smaller woody species; and another grouping of
shrubs and small trees appears in the Smokies, the
suhrnesic comrnodium between covr forests and oakchestnut forests. Species of the three transitional
cornmodia which are best developed are:
Ecotonal-mesic union
B e t u l a lutea
A c e r spicatztm
Amelanchier laevis
V t b u r n z t m alnifoliztm
Cornus alternifolin
Rl~oclodendron calelzcl~ilnceum
V a c c i n i u m corzstablaei
L yonza li,qustrirm
Crataegzts m a c r o s p e r m a r.
roanensis
R u b u s canadensis
S a l i x spp.
37
Submesic union
B e t u l a lenta
A c e r pensylcanicztm
A c e r rubrztm
Amelanchier urborea
V i b z t r n u m acerifolzum
C o r n u s jlorida
R l ~ o d o d e n d r o ncalendulac e u m
Vaccirzium conctablari
P y r u l a r i a pubera
I l e x montancr
C a l y c a n t h u s fertilis
Cea?lothzrs americtrnus
R. H. WHITTAKER
Sn:okies might he regarded a s postclimax a n d preclimax, respectively. Oak-chestnut forests, occupying
many of the lower slopes of the mountains, might
then he interpreted a s clinratic cliniax. Alternatively,
by the polyclirnax theory (Sichols 1917, 1923; Tansley 1935), the five types distinguished along the
nioisture gradient a t low elevations a r e so many polyclimaxes. Of these oak-chestnut forest would preiuniably be designated climatic cliniax a n d the remainder physiographic or topographic climaxes.
I n the present study these five types a r e regarded
not a s discrete associations but a s p a r t s of a single
vegetation continuum. I f stands a t all points along
the gradient are accepted a s self-maintaining, and
therefore climax in this sense, the continuum itself
should be regarded a s the cliniax. R a t h e r than a
single clinratic c l i n ~ a xo r set of polyclimaxes, climax
vegetation i n the Smokies con~prises a whole vegetation pattern ranging f r o m cove forest to pine forest
along the ntoisture gradient and f r o m these low-elevation types to spruce and fir forests i n one p a r t
of t h ~range and t o other deciduous types in the
other p a r t of the range a t high elevations.
I n such conditions a s those of the Smokies, the
pattern of vegetation along the ~tioisture gradient
provides a reasonable description of climax vegetation In a glven elevation belt. Through the moisturegradient patterns ( a n d the transect data representing
them), composition of vegetation a t different elevations may be compared. Population levels of species
can be c o n ~ p a r e dfrom one elevation to another; thus
the changing role and importance of hemlock a t
different elevations may be revealed ( F i g . 1 5 ) . Within
the c11111axpattern f o r a given elevation, relative importance and relative position along the moisture
gradient of species may he compared. W h e n different cliniax patterns a r e cornpared, s t a t e n ~ e n t s may
he made about the r e l a t ~ v eimportance _of a given
species population or conlmunity fraction i n different
patterns, relative irrrportance of growth-forms or
life-forms i n different patterns, relative variety of
species or of dominants in different patterns, relative
extent of change in coniposition f r o m most rnesic to
inost xeric sites in different patterns, etc. Some of
these possib~litiesmay he ~ l l u s t r a t e din the follo\v~ng
statements on cliniax coniposition in the Great Smoky
Ilountains :
1 . The climate of low elevations (1500-2500 f t ) is
a deciduous forest climate, a s expressed in cornposition of cliriiax stands. Deciduous trees form 73%,
henllock 14%, and pine species 1 3 % of total canopy
stenis i n the low-elevation moisture-gradient transect
(Whittaker 1951, table 3 ) .
2. The clin~ate of high elevations (4500-5500 f t )
in the northeastern half of the range is a coniferous
(taiga) climate. Two ahietine dominants, Picea ruhens and A b i e s fraseri, make u p 87% of canopy
stenis i n the sample, deciduous trees only 13%. Toward still higher elevations the deciduous trees decline to 7 p e r cent (5600-6100 f t ) a n d 1 % (6200-6600
f t ) of canopy sterns. I n the southwestern half of the
Ecological Nonographs
Vol. 26, No. 1
range, however, a pattern i n which deciduous broadleaved trees form 100% of canopy stems occurs a t
c,levations between 4500 and 5500 f t .
3. Alajor cliriiax tree species a t low elevations are,
in order of over-all importance with percentages of
canopy stems f o r the whole transect (Whittaker 1951,
t,ahle 3 ) : Quercus prinus 14, T s u g a canadensis 14,
Castanea dentata 13, Quercus borealis v. m a x i m a 6,
Pallus cirginiana 6, A c e r spccharz-lm 4, A, r u h r u m 4,
T i l i a heteropll!llla 4, P i n u s rigida 4, Liriodendron
t u l i p i f e r a 4, P i n u s strobus 3, Aescullis octandra 3,
Quercus alba 3, C a r y a glabra 3, Halesia monticola
2, B e t u l a allegheniensis 2, Quercus coccinea 2, C a r y a
tomerztosn 2. S o other species is represented by more
than 1 p e r cent of canopy sterns.
4. I n contrast to these diverse stands of low elevations, composition of canopy trees a t highest elevations, above 6100 f t , is simple: A b i e s fraseri 90,
Picea rubens 9, a n d B e t u l a lutea 1%of canopy sterns.
rndergro~vths of these stands a r e varied, hut in the
canopy there is profound simplification i n composition from lowest elevations to highest.
5. I f a criterion is chosen f o r the oak-forest groupi n g (all stands i n which oaks plus other subn~esic
a n d suhxeric species make u p a t least 50% of stems),
then 57% of the climax stands sampled a t low elevations a r e oak forests, a s defined. Among the same
samples only 28% a r e oak-chestnut forests, if these
a r e defined a s all stands i n which Castarzea dentata
a n d Qzterc~is prirzus form a t least 50% of canopy
stems.
6. The relative irtiportance of oak stands in the
climax pattern decreases somelvhat with elevation.
F o r the elevation belts 1500-2500, 2500-3500, 35004500, and 4500-5500 f t outside the range of spruce,
percentages of stands classified as oak forests were
-57, 46, 44, and 41. The relative importance of
rnesophytic stands correspondingly increased-11,
28,
41, 59-while that of pine stands decreased-32,
25,
15, 1 .
7. Stand composition a t the niid point of the riioisture gradient may be deterniined-by the middle station of the transect, by a "median" stand so chosen
t h a t half the samples have inore highly n~esic and
half more highly xeric moisture-class formulas, or by
percentage-similarity comparison with stands representative of the extremes of the gradient. X o r e cornplicated means of finding a n "elective mean" in a
multi-dimensional pattern a r e described by Rainensky
(1930). The three techniques mentioned give equivalent results i n the Smokies. Using the first a s the
simplest, intermediate stands a t different elevations
a r e : stands dominated by Quercus prinus a t 15002500 feet, stands i n which Q. prinus and C a s t a ~ l e a
dentata share don~inancea t 2500-3500 f t , and stands
i n which C . dentata a n d Quercus borealis share dominance a t 3500-4500 f t . X o r e limited data f o r high
elevations give stands in which C. dentata a n d Q.
borealis share dominance ( 4 5 a n d 40% of canopy
sterns), with A c e r r u b r u m , Halesia monticola, a n d
F a g u s grandifolia a s m a j o r subordinate species.
J a n u a r y , 1956
~ E G E T A T ~ OONF T H E
GRE
R. H. WHITTAKER
40
3. Percentage sirililarity comparisons with the extrelnes suggest a stand of the chestnut-sugar mapletuliptree segregate (Braun 1950, table 5 ) or correkponding drier mixed niesophytic (Braun 1950, table
1 B ) as intermediate climax stand in the Cumberlands, in contrast to the oak-chestnut intermediate
climax of the Srnokies.
4. I n composition and diversity the all-deciduous
mixed ~nesophyticforest (Braun 1940a :237) is closely
similar to the most highly direrse stands in the
Sn~okies, those described as cove forest transition.
D i v e r s l t ~conditions of stands in the two ranges appear to be similar, although the sanlples are not
strictly comparable. The sum of a set of tables which
approximate a moisture-gradient pattern (Braun
1950, tables 2, I A , 5, 7, 8) gives a diversity value
(5.5) scarcely different fro111 the 5.8 obtained for
canopy stems a t low elevations in the Sn~okies(Whittaker 1951, table 3 ) .
5 . Indicated half-changes along the nloisture gradient in this same set of tables are slightly more than
2 in terms of species and approximately 2 in terms
of moisture classes, if the pine stands of special sites
are excluded.
Coniparing the regetation pattern of the Cuniberlands with that of the Smokies. it m a r be said that
the former is more clearly mesophytic in character
and of comparable species diversity, but less diverse
in terms of stand types along the moisture gradient.
The Cumberlands pattern largely corresponds to the
niesic half of the Smokies patteni (cove forest, cove
forest transition, and oak-chestnut forest), while the
more xeric types of the Smokies (oak-chestnut heath
and pine forest) are reduced in the Cumberlands to
stands of special edaphic situations. The two climax
patterns are thus related through a shift of relative
clnphasis of the ~iiesophytic grouping and the oakforest grouping.
Such shift in relative importance of parts of climax patterns is further illustrated in Braun's treatment of vegetation of the Southern Appalachians, the
Ozarks, and the area between. Three regions are
recognized fro111 the Cumberlands west: a region of
mixed mesophytic climax in and around the Cumberlands, a region of oak-hickory climax in the Ozarks
and west and north from them, and between these
the Western Mesophytic Region, regarded as ecotonal, and forriling "a mosaic of like and unlike climaxes" (Braun 1947). Of this last region Braun
(1950:123) writes, "Tn its eastern part, mixed nlesophytic forests are of frequent occurrence; westward
they become more and more lirnited in extent and
more closely dependent on very favorable habitat
conditions.. . Because of the many mixed forest communities, and of the gradual change from east to
west in extent of mixed mesophytic communities
and in cornposition of forests, and of the increasing
frequency of communities in which oaks are dominant, this is considered a transition region."
The description seerns most expressive of gradation on a grand scale, along a major climatic gradi-
Ecological Monographs
Vol. 26, No. 1
is the distributional relations of species which cause communities are interpreted. I n sampling, an ecoloecologists to recognize coir~~nunity-types.The a p - gist must usually choose certain stands as "typical"
proach to species distributions through co~n~nunity- or "representative" while other stands are passed by
types thus involves a methodological inversion, pro- as atypical, transitional or mixed, or disturbed. When
ceeding from a higher to a lower level and fro111 sec- a number of stand samples are presented in a comondary phenon~ena to the distributional relations piled table f o r a comir~unity-type,the table and the
which underlie them. I f , on the other hand, prob- properties of the coir~munity-type indicated by it
lems of the validity and meaning of associations are represent not merely "the community," but an ecoloapproached by way of the associations themselves, gist's conception of the community as this influences
other difficulties of method appear. Sarilples are his choice of samples. A constancy value of 100%
taken from the vegetation, and it is difficult to take for species A may sornetirnes mean little more than
such samples without preference for those which con- that an ecologist regards as typical only stands conform to subjective ideas of associations and some ex- taining species A. Consistent difference and apparent
clusion of those which do not. I f , now, the samples discontinuity between compiled tables for two com.are used to study validity of associations, a clear cir- inunity-types may imply only that intermediate stands
cularity is introduced into the procedure; associations were not sampled. Statenlent that a community-type
are being studied in terms of samples taken in terms is homogeneous or heterogeneous may reflect the narof associations. The student is thus caught in a circle rowness or broadness of the ecologist's conception, as
of studying secondary phenomena and his own ab- this directs his sampling. The author would emphastractions from them in terms of themselres, in place size both the value of such cornpiled tables and the
of the more reasonable procedure from lower levels fact that sample choice, and therefore statements
to higher. The contrary viewpoints of Du Rietz about the comn~unity-typeand constancy and fidelity
of species, can seldoin be wholly independent of a
(1921 2 1 5 ) and Tansley (1920) may be noted.
I t has been indicated that the approach to species giren ecologist's conception of the con~munity-tvpe.
distributions along the moisture gradient (but not I n practice con~piledtablei are often presented and
along the elevation gradient) used in this paper also inferences drawn from them as if the table repreinvolves a circularity. I t is further clear that the sented "the community" in nature, wholly independapproach to the moisture gradient proceeds from the ently of the ecologist's interpretation of it.
Two very general difficulties appear here, in the
higher l e d of the community gradient to the lower
one of the assurned gradicnt of physical environlnrnt failure to allow for the ecologist's interpretation of
-the approach is inferential. The point is not vegetation as guiding choice of data on which further
that such procedures are excluded from ecological interpretation is based, and the confusion of levels
methods; it is that they should be understood, allowed i~lvolved in the word "comnlunity." At least three
for, and made explicit. The author has sought to levels of phenomena may be recognized among the
state such limitations in his own methods and feels applications of this term (cf. Dice 19% :426) : ( 1 )
that certain liri~itationsof other methods which may communitzes or stands which are essentially homob e a d ~ a n c e das alternati~esshould also be obser~ed. geneous over some area of the earth's surface and
The relation of abstract types to the stands from comprise all organisms above and below the soil surwhich they are abstracted involves further difficulties. face in this area, ( 2 ) community-fractions and microI n preliminary field work a type or assuciation is commumrties, comprising some organisms of the comrecognized, and a number of samples conforming to munity or stand which are for some reason grouped
it are taken. The samples are relatively uniform and together, as in a stratal community of plants or
of anirnals of a particular
as a group distinct from those taken to represent animals, a co~~imunity
other associations; the further sampling thus rein- taxonomic category, o r the community of a special
forces the conctlption of the association. This, now, micro-environment, and ( 3 ) commumity-complexes,
is likely to be seen as a "good," "real," or "valid" unit, comprising a number of distinguishable communities
verified by the sampling. The abstract conception is or stands arranged in a rnosaic or pattern in space,
likely to be projected back into nature and regarded
such as landscapes, biotic provinces, the complexes
as a real unit of structure of that which has been of bogs and frost-determined landforms, ;topographic
sampled; the unit in the ecologist's mind is reified patterns, etc. On each level abstract t y p e s of conor hypostatized (cf. Tansley 1929). The initial choice munities may be recognized, such a s : ( 1 ) formations,
of a "climatic climax" niay silriilarly be reinforced associations, sociations, dominance-types, site-types,
by observations of successions and used as a basis for
( 2 ) unions, socions, niicro-associations, ( 3 ) !andscapefurther interpretation, until the "climatic climax" types, bog types, climax-complexes, the generalized
seen~san established and necessary fact. I t is all too topographic patterns of the present work. The ameasy to moTe from the view that such abstractions biguity of the term conlmunity results not from the
have real usefulness to the view that they are part of, accepted convention that it may apply on any level of
con~~nunity-size
fro111 the inhabitants of a termite'.,
or must correspond to, reality.
The reification of the comn~unity-typeis often a p - gut to those of an ocean basin, but from its applicaparent in the manner in which quantitative data on tion on a t least two levels of abstraction: the ron-
Ecological Monographs
Vol. 26, No. 1
..
J a n u a r y , 1056
43
Ecological Monographs
Vol. 26, No. 1
more than one physiognomic t y p e ; they are consequently not formations. The word LLsystem"has been
used f o r them here, with no intention of establishing
this as a new vegetation unit. These two major p a r t s
of the vegetation pattern a r e named, f o r their geographic relations, the "Eastern Forest System" and
the "Boreal Forest Svstem."
Physiognomic types within each system are a logical basis of further division. Vegetation of the
Smokies includes types dominated by f o u r major
growth-forms-needle-leaved
evergreen or coniferous
trees, deciduous broad-leaved trees, evergreen-sclerophyllous shrubs, and grasses. I t is useful f o r the
present study to divide the tree growth-forms
further. The ~ i n e sof thr Smokies, forming open,
xeric, lower-elevation stands are conveniently distinguished as a growth-form from the abietine trees
(Picea, Abies, Tsuga) forming denser and less xeric
stands, predolninantly of higher elevations. Oaks
affect lower strata through late leafing and type of
leaf-litter differently from many other rleciduous trees
( B r a u n 193,5b, Kucera 19.52) and may be (listinguished as a growth-form. Along the moisture grarlient in the "eastern" forest pattern, non-quercine trees
predominate in mesic sites, oaks i n intermediate ones,
and pin-s in xeric ones (see P a r t I ) . Ericaceous
shrubs are a major growth-form in the Smokies,
dominating heath balds and sharing dominance with
a n open tree ctratum in forest-heaths. The heath
strata are predominantly evergreen but contain, and
are in some types dominated by, deciduous Vaccinioideae. Physiognomic types recognized in the
Smokies are t h u s : abietine forest, non-quercine deciduous forest, oak forest, pine forest, forest-heaths
(pine heaths and oak-chestnut heath), heath bald,
and grassy balrl.
Vegetation types are further distinguished accorrling to their (laminant species. All but two of the
vegetation types described below a r e defined by dominance of one species or a pair of species. The two
exceptions (cove fprest and heath bald) are dominated by several important species, the proportions
of which vary from one stand to another. These types
are consequently defined by rlistributional groupings
or commodia ( P a r t 11) of species.
Since the vegetation units are not defined primarily
by character-species, according to the system of
Braun-Blanquet (19.51), they are not necessarily associations in the sense of that term mhich most nearly
has international acceptance. They consequently will
be designated only as "types" and will not be given
formal, latinized names. I n deference to phytosociological practice, however, character-species have been
designated f o r these types as f a r as possible. To indicate these without repetition, names of characterspecies are marked with asterisks ( * ) in the rlescription of the type to which they apply. I n view of the
distributional data already discusserl, it need hardly
be emphasizeil that these character-species a r e of low
degree (mostly preferential or holde). Very few species of the Smokies can be considered exclusive o r
January, 1956
VEGETATION
OF THE Gii
Ecological Monographs
Vol. 26, No. 1
The shrub stratum is poorly developed or absent gests, a naturalist wandering, unknown to him, into
m stands of valley flats at middle elevations. E u o n y - some Tertiary forests might see no very conspicuous
rnzu. americanus* and L i n d e r a benzoin* occur iri low- difference except the presence of Ginkgo and Seelevation stands, C o r n u s alternifolia, V i b z ~ r n z ~alnim
quoia (or Metasequoia) .
folium, and R i b e s cynosbati in high-elevation ones.
Various cornbinations of dominants appear locally
H y d r a n g e a arborescens occurs locally at all eleva- in the cove forests: A c e r saccharum-Halesia montitions; R h o d o d e n d r o n m a x i m u m occurs along streams cola, A. saccharum-Talia heterophylla, T s z ~ g a canaand (often with LeucothoB e d i t o r u m ) in some other densis-T. Iteterophylla, Aescrclz~s octandra-T. heterosites, particularly under hemlocks. There is, holvever, phylla, A. o c t a n d r a - B e t z ~ l a allegheniensis, P. canano shrub grouping characteristic of the cove forests densis-Liriodendroa t u l i p i f e r a , P. canadensis-Fagus
as a whole.
grandifolia, P. canadensis-B. allegheniensis, A. sacThe herb stratum is the richest in the mountains. c h a r u m - F . grandifolia. None of these (cf. Cain 1943)
Cain (1943) lists over a hundred species for the represents s well-defined and extensive type. The
spring and summer aspects, and Braun gives com- most significant change in the cove forests, other
parable lists in her papers (1935b) 1940a, 1942). than the transition to oak types, is the gradual
Summer herb coverage is as high as 80 percent in rise of A e s c u l z ~ s octandra, T i l i a Iteteropltylla, and
some sites, with a luxuriant growth of mesic ferns B e t z ~ l a alleghenzensis toward their different modes
and herbs of spreading or umbrella-shaped foliage at higher elevations, where they form the so-called
form. Most species of the mesic herb union are cen- northern hardwoods dominated by these three
tered in the cove forests and may be regarded as southern tree species (cf. Braun 1950 :207). The
character-species for them. The most abundant spe- upper cove forests may be regarded as a sub-type of
cies of the union in summer are E z ~ p a t o r i u m rzego- the cove forests with the same dominants in different
sum*, C i m i c i f z ~ g a racemosa*, C a u l o p h y l l u m thalic- proportions, but with Liriodendron and Tsuga often
troides*, I m p a t i e n s pallida*, L a p o r t e a c a n a d ~ n ~ i s * , absent, and with A c r r s p i c a t u m , Amelancltier laevis,
T r i l l i u m e r e c t u m v. albiflorum*, A s t e r divaricatus*, Vibzirnzim a l n i f o l i t ~ m ,and C o r n u s alternafolia domiand the ferns D r y o p t e r i s spinulosa v. intermedia*, nating the stratum of shrubs and low trees. M o and A t h y r i z l m t h e l y p t e r i o i d e s f . Beneath them grow n a r d a dadyrna, R u d b e c l i a lacaniata, Oxalis m o n t a n a ,
smaller species-Viola
spp., Stellaria pubera, T i - and other species of the high-elevation mesic union
arella cordifolia", G a l i z ~ mtriflorum, and E z ~ o n y m u s appear in the herb stratum.
obovatz~s. The beautifully developed herb community
An additional subtype may be recognized in the
is stratified as the forest is, with a canopy of tall cove forest transition between the mesic forest and
spreading herbs and ferns and an undergrowth of oak forest groupings. As less mesic sites are aplower and prostate plants. Somr species of the sub- proached from the cove forests, increasing percentmesic herb union occur, and toward higher elevations ages of submesic species occur in the stands. Subspecies of the mesic high-elevation union form a con- xeric species ( C a s t a n e a dentata, O x y d e n d r u m arspicuous part of the stratum.
boreum, Quercus alba and Q. p r i n z ~ s ,N y s s a s y l v a t i c a )
The relation of cove forest and mixed mesophytic also appear in significant numbers. I n the cove forest
to the Arctotertiary forests has been indicated by transition 25 or more tree species, representing all
Braun (1935a, 1947, 1950). Cain (1943) studied the' classes but the xeric, may be encountered among 200
Tertiary character of the cove forests in the Smokies, or 300 stems. I n the shrub stratum R h o d o d e n d r o n
using fossil records of genera and distributions, par- m a x i m u m is most important, and many of the shrub
ticularly the frequent eastern Asia-eastern North species of oak-chestnut forests appear. With their
America disjuncts. Cain considered that all genera heavier growth of small trees and greater shrub covof ferns and shrubs and most genera of trees and erage (20-50%), these stands are less open and spaherbs were of Tertiary history. His figures for trees cious in appearance than the cove forests themselves.
and flowering herbs were 86 and 75% of genera with Herb coverages in cave forest transition vary widely
known or indicated Tertiary history, including the
(5-,50%), the extremes of this range corresponding
minor species in his lists. On the basis of his con- to the low coverages in oak-chestnut forests and the
stancy-class V or the most abundant species of the high ones in cove forests.
low-elevation mesic herb union listed here, all prin,%fixed M e s o p h y t i c i n t h e S m o k i e s a n d C u m b e r l a n d s
cipal herb species are of Tertiary history. The union
The cove forests of the Smokies are closely related
is related, and linked by its predominant growthto the mixed mesophytic forests of the Cumberland
form, to Lippmaa's (1939) Galeobdolon-Asperula- Mountains (Braun 193513, 1940a, 1942. 1950), but
A s a r u m Union of herbs surviving from Eurasian the mesophytic communities of the two ranges differ
Arctotertiary forests. Of the trees listed by Cain, in some respects (see also Part 11). With the greater
if Tulipastrum is grouped with Magnolia, and area occupied by mixed mesophytic in the Cumberif Robinia pseudoacacia and O x y d e n d r t ~ marboreurn, lands, there is a greater differentiation of the assowhich are extraneous to the cove forests, are set aside, ciation into segregates; some of the combinations
all are of Tertiary history. The Tertiary history of mentioned by Braun were not observed in the
the genera certainly does not imply that extensive Smokies. Sotable among these are the beech ( F a g u s
changes have not occurred. However, as Cain sug- g r a n d i f o l i a ) ravine coiilnlunities, Quercus alba-F.
January, 1956
VEGETATION
OF
THE
GREATSMOKE'
D~OUNTAINS
47
these mesophytic forests in the Smokies have discouraged the author from following this course:
1. T i l i a heterophglla and Aesculus octandra are indicated by Braun (1950 :43) to be characteristic species
of Nixed Mesophytic. I n the Smokies, a t least, other
character-species may be added to these, notably H a lesia monticola, B e t u l a allegheniensis, Cladrastis lutea, and the local population of A c e r saccharurn. The
populations of these character-species are centered
not in the cove forest transition but in cove forests
proper.
2. The optimum development of mesophytic forests,
with the richest mesophytic herb stratum as well a s
strongest dominance by five of the character-species
above, is in the cove forests of deep valleys, not in
the cove forest transition.
3. Castanea dentata and other tree species charactcristic of the oak forests are almost absent from the
cove forests proper and extend into cove forest transition only with the tails of their distributions. It is
difficult to interpret these species as belonging to the
cove forests and segregating into oak forests. Rather
than this, their distributions appear simply to overlap in p a r t with those of cove forest species.
4. Applied to the Smokies, the Mixed hlesophytic
in Braun's sense seems less a definable vegetation
type than a range of stand conditions from cove forests to more mesophytic oak forests. Mixed hlesophytic seems too broad and heterogeneous a grouping, bringing together into one association species
whose relations to the moisture gradient are too
widely different.
I n the .Cumberlands, stands of mesic species mixed
with oak-forest species form the prevailing climax
type. It is consequently reasonable to emphasize
these highly mixed stands and to regard the mesic
and xeric extremes as secondary to them. The diverse
stand types of the Cumberlands may then be given
unity in terms of the monoclimax theory, very broad
clefinition of associations, and the conception of association-segregates. I n the Smokies it seems preferable to identify the mesophytic association in question with the cove forests and to regard stands highly
mixed with oaks and Castanea dentata as transitional.
Applied to the Cumberlands, the author's interpretation would suggest that the Mixed Mesophptic comprises: ( 1 ) truly mesophytic forests (sugar maplebasswood-buckeye), ( 2 ) a wide range of highly mixed
transitional stands between these and oak forest
types, and ( 3 ) various segregate types dominated by
species which are represented also in stands of the
preceding.
There are no signposts in nature to tell us whether
we should see mixture occurring in one direction or
segregation in the other, where in a vegetational continuum we should locate our associations and where
our transitions. The interpretations of Braun and
the author are different, possible patterns of abstraction from the vegetation of the Southern Appa-
48
R. H. WHITTAKER
FIG.15. Rise of Tsuga canadensis toward higher elevations, distribution in moisture-gradient transects: a,
1500-2500 f t ; b, 2500-3500 f t ; c , 3500-4500 f t .
EcologicalVol.
Monographs
26, No. 1
Magnolia fraseri*, and sometilnes Liriodendron tulzpifera or Fagzrs grandifolia. A low-tree layer of
Ilex opaca*, Betula lenta, Acer pensylcanicum, and
most other small-tree species of the submesic and
ecotonal-mesic unions, is well developed in some
stands but relatively unimportant in others in which
Rhododendron dominates the undergrowth.
On steep slopes the shrub layer may be strongly
developed as a hlgh heath. At higher elevations heath
coverage may be 60 to 80% with Rhododendron
marimum or R. catazcbiense dominant. The TsugaRhododendron combination resembles the type described by Oosting & Billings (1939), but the combination with Vaccinium (Polycodiuni) was not encountered in the Smokies. Lezicothoe editorurn is
frequent in these forests, Hydrangea arborescens,
Kalmza latifolia, and others are occasional. Some
stands on lower flats, more mixed with hardwoods,
have llttle shrub undergrowth.
The herb stratum may be nonexistent in dense
stands with heavy heath undergrowth. The effect of
hemlock needles and the superficial root system is to
produce a surface soil much more acid than that of
hardwoods stands and to reduce water penetration to
lower levels, drying the soil (Daubenmire 1930, 1931).
Although hemlock stands occur in mesic sites, they
are relatively xeric and acid for undergrowth plants.
A dense Rhododendron heath may develop under
these conditions, and the forest floor may be a sterile
surface of hemlock and Rhododendron leaves. I n
stands with less heath an herb stratum of low cover,
age may appear. Dryopterzs spznulosa v. ivetermedia,
Mitchella repens*, with Goodyera repens v. ophioides"
often growing anlong its stems, and Tiarella cordifolia are the most successful herbs. Polypodium virgznianum is frequent on exposed rocks and roots.
Herb coverage vanes from zero up, related to the
proportion of hardwoods and development of heath.
The hemlock forest type is most distinctive, with
heaviest hemlock dominance, densest heath growth,
and most impoverished herb stratum on steep slopes
a t higher elevations. At lower elevations stands are
more mixed with hardwoods, with less heath and
more herbs. Below 2500 f t the hemlock forest gradually merges with the cove forests; and hemlock becomes only one of t h e dominants, though the most
important one, of lower-elevation cove forests.
3. GRAY BEECH FOREST
Above 4500 f t the high-elevation or "gray" populzition of beech (Fagus grandifolia*) is the mosL important tree of mesic deciduous forests (see also
Russell 1953). Gray beech forms small-tree forests.
The beech itself seldom exceeds 1 5 in. diameter;
and the bulk of canopy stems may be in the 8- to
12-in. classes, though cove-forest hardwoods occurring
with it may reach fair size. I n two quadrat studies
canopy stems (8-10 inches) were only 25 to 40
f t high, with crowns 12 to 20 f t from the ground.
Small stems may be numerous, but frequently the
undergrowth is slight, with few seedlings and few or
J a n u a r y , 1956
VEGETATION
OF
THE
GREATSMOKY
MOUNTAIXS
49
50
R. H. WHITTAKER
Ecological I f o n o g r a p h s
Vol. 26, No. 1
January, 1956
VEGETATION
OF THE GR
becomes strongly dominai~t on soille exposed southwest ridges, with a rather open growth of small trtvs
and a fairly rlch herb l a j e ~x l t h grasses often conspicuous. I n structure and appearance tht3se forests
iuggest the white oak \\nod> of hills f a r n e s t of the
Smokles, though ver) dlfferrnt in thelr florlstics. I n
7 . RED OAK-CHESTSLT FORk3ST
home stands chestnut (Cactanea clentutu) is dominnnt
The red oak-chestnut cornbinatlon 1s submeslc in
(cf. Cam 1 9 3 1 ) . Quercrrs borcrrlzs IS almoit a1wa)s
composition of both tree canopy and l o x e r strata, present; and Carya glabra and Qzterc~ts prl?zu\ ocand 1s the hlgh-altltude equlr alent of chestnut oak- cur In the canup) a t lower r l e ~ a t l o n s Of the srnall
chestnut forest S o r t h e r n red oak (Qllercus bore- trees d t t r t l t b r u m 1s most lrnpoltant, 0 I l/denrl~11171
alzs*) and chestnut ( C a s t a n e a d e n t n t a ) for111 7 0 or a r b o r ~ j t m and Kobznzu psecctloucucra :Ire falrl) nu807{ of the canopy, n l t h the former usually more 1111- inerouy ton ,ard lon el elelations
portant. Although riiost of the chestnuts a r e dead,
The shrub layer is pledominantly erlcnceous, n i t h
nlany large chestnut trees In these and other high- Rhododenrlron calendulacpunz and Vaccznzrcm t o t / had a t least a few branches living In stablarz most impoltant. Iicrlmta lat~folzcr1s frequent
e l e ~ a t ~ oforests
n
1947. S o canopy associate reaches a n y subdantial a t l o u e r ele\arions and I > o f t ~ nthe most impoltant
percentage In r~roststands, of other species d e e r ru- shrub spec1t.s thele. S h r u b coverage is f a d \ h1g21brzlln is most i ~ n p o r t a n t . The lower-elelation stands gt3ncrally 3 0 to GO? belo\! and 20 to 50': nbolc
may h a ~ ea f a l r percentage of n~esics-Tzlza hetero- 1500 f t . The heib s t ~ a t u m Is a m ~ x t u l eof f e i n s
plrrjlla, P~zcnzls serotlrzcr, Retzlla a l l e g h e n ~ e n s ~ sd,e s ( D r ~ / o p t e r l sno1 eborctcerzszs, Atltyrlcitn fill I - f e m z n a \
c u l ~ t soctamdra, F r a ~ z n z c samerzcatza-, but few meslc u ~ p l e n z o ~ d e sn) ~ t hother herbs of the subineslc. -ubtrees except Fagzcs gratzclzfolza enter the canopy of xenc, and high-ele~ation suhniesic unions Graqses,
stand, above 4.500 f t . The submesic s~rialltrees, ~71th Pterzd?~irnaqu~lzrlzcmv, l a t ~ r l s c u l u m ,and other xerlc
Hale.>za ~nontzcolajoinlng them here, form 30 to 5 0 5 species also a r e prominent. H e r b coverage is 10 t o
of the stems, h a ~ i n gtaken advantage of the death of 50% below 2nd 20 to 50r! above 4500 f t .
the chestnuts along with northern red oak. The chestI'itze S t a u d s crrzd Tlrcir Jfaintenatzce
nuts were less nunierous i n submeslc stands of higher
Nost
steep, ope11 south- and southwest-facing
elelatlons, h o w e ~er, and reproduction and appearance
have not been altered a s much by thelr death a s in slopes on the Ten11essc.e side of the mountains a r e
occupied by pint. forest and pine heath. Tsually the
the chestnut oak-chestnut forests.
S h r u b colerage a b o \ e 4500 f t is lo\\, 1~1thR h o d o - population of a single pine stand s h o ~ v sa bimodal
dendron calrrzdulaceum and V a c c ~ r z z u m corzstcrblacz size distribution; often other modcs are represented
the p r ~ n c ~ p aspecles.
l
V a c c ~ n z z l m pallzdzlnz and V . b y a few old trees of earlier generations (Fig. 1 6 ) .
I1~rs2ctzlmoccur in some stands, especially on ridges, I n these d r y sites occasional severe fires destroy the
and Vaccznzzcm erytlrrocarpum and V ~ b u r t z z c malni- forests, and following the fires dense stands of yo!lng
f o l ~ ~ c r na r r occaslonal. I n forests below 4500 f t pines develop. As these trees matllre, age, and die,
Rlrocloclendrotz calerzdzclace~imis most Important ; but the canopy is opened; and a i1ew generation of pines
most other submesic shrubs a p p e a r 111 some stands, establish themselves. W h r n these in t u r n mat~rrc,,a
and H~ldrarzgea arborescens is occas~onal. I i a l m i a third generation of seedlings appears, while sonle
c
above scattered stems of the first a r e still prrseiit. The
latzfolta is alnnost absent froin s u b i i ~ e s ~stand,
3800 f t . S h r u b coverage \arles from 20 to 5 0 4 below effects of fires in permitting the estnblishmt~nt of
initial single-age stands thus introduces into the
4.500 i t to only 0 to 2 0 4 above
maintenance
of the pine stands a rhythrn of gelicvaThe herb layer a b o ~ 4500
e
f t often resembles closely
that of south-slope beech stands, but with lower tions which may persist f o r some tin~t.. The c o ~ i ~ p o s coverage Sedges ( C a r e s s p p . ) and f e r n s (mostly ite stand counts fro111 mature stancis s~lioothout the
d t h y r z z l m fil~x-fernznav asplenioldes) a r e abundant. double-age composition of individual stands, and the
A feu7 meslc species, notably Ezcpatorzrn rzlgosurn, composite curves then take on the characteristic form
Laportea canadensis, and A s t e r d i ~ a r i c a t u s ,a p p e a r of self-maintaining forests ( F i g . 6 ) . The dense
along with the submesic a n d high-elevation submesic small-tree stands on disturbed and recently burned
unlons and P o t e n t ~ l l a canadenszs v. c a r o l ~ n i a n a . Be- sites were not included in these counts; but reprolow 4500 f t the herb flora 1s much the same a s in duction a p p e a r s adequate to replace the stnntls inchestnut oak-chestnut forests, with only the subalpine definitely.
Since chance may determine the species of pine
species T r ~ l l i u mundzllatum, Solidago g l o ~ n e r a t a ,and
Diplryllria cymosn dlst~nguishingthe list f r o m that of seeds available to restock a stand a f t e r a fire, cha~ice,
the lower type. H e r b coverage is 1 0 t o 40% in the a s Cain et 01. ( 1 9 3 7 ) noted, may determine whirh of
the pines available a t a given elevation first domilower stands, 20 to 60% above 4500 f t .
nates the stand. Once dominant. the sr~eciestend.; t o
8. W H I T E OAK-CHESTXUT FOREST
maintain its dominance through several generations
Below 4500 f t white-oak chestnut forests a r e sel- while other pines and oaks may gradually increase to
dom clearly dlst~nguishablefrom red oak-chestnut. approach a balanced stand f o r that ele\ation a n d
Above 4500 f t , however, white oak ( Q ~ i e r c u salba ) site. Most inimature pine stniltls in the Sniokies a r e
heaths Gaultherza procctmbetzs and Epzgaea repens,
P a t z ~ c u msp., C a m p u n u l u dzzarzcata , and drireoluria
laerlgata a r e falrly frequent. H e r b coverage is quite
low, 1 to 5 5 , with herbs other than Galax and the
grourld heaths well below 1 % .
R. H. WHITTAKER
52
DIWFTLR
Ecological Monographs
Vol. 26, No. 1
January, 1956
VEGETATION
OF TIIE GREATSMOKY
~IOUNTAINS
53
Center
51
R. H. WHITTAKER
Ecological N o n o g r a p h s
T o l . 26. No. 1
January, 1956
VEGETATION
OF THE GREAT
SMOKY
?IIOUNTAINS
55
FIG.l'i. Fraser fir forest, near summit of Mt. Le('onte. Reproduced by permissioll of Thompsons, Inc.,
lino\villr, Tenn.
HITTAKER
Ecological Monographs
Vol. 26, No. 1
January, 1956
VEGETATION
OF THE GREATSMOKY
MOUNTAINS
57
58
R. H. WHITTAKER
EcologicalTTol.
Monographs26, No. 1
of the cornplex vegetation; the final charts are presented as Figs. 19 and 20. About 40 of the 300 sitesamples were outside the lines drawn for their types.
A few limitations of the pattern niay be noted.
Gentle slopes often support vegetation a step less
xeric than indicated on the pattern. Although sharp
peaks are more xeric than slopes below them, rounded
o r level sun~rnitsmay support rnore mesic vegetation
than their south slopes. The beech gaps i n certain
concave slopes in the subalpine forests are not indicated in the boreal forest pattern. The two bald
types appear in similar topographic positions f o r
the two systems, but are show11 in contact n i t h only
one other type in each case. The contact indicated
i i the critical one of the most closely related type,
norrnally adjoining the bald on the south slope.
Other types meet the balds on other slopes through
abrupt transitions, and the grassy balds thus contact red oak-chestnut and beech forests. The heath
balds may come in contact nith oak-chestnut and pine
heaths and occasionally with oak-chestnut and hemlock forests. Wherever the environmental gradient
is telescoped, as along a sharp ridge, two types
whether adjacent ones o r not may meet with a telescoped transition between them.
One noteworthy feature of the pattern is the expansion of mesic forests toward higher elevations.
Vertical lines on the chart extend through topographically siniilar sites a t different elevations, but
not through sites of similar nioisture-balance. Change
of moisture conditions in topographically similar
January, 1956
VEGETATION
OF THE GREAT
SMOKY
MOUNTAINS
59
Throughout its range, a s the cove forests expand toward higher elevations, hemlock nioves ahead into
sites which a r e one step less niesic.
3. Gray beech forest. The south-slope subtype with
sedge floor occupies gaps, draws, a n d concave sheltered slopes facing southeast through south to west
above 4500 f t . The north-slope subtype with beech
o r beech-mixed canopy a n d herbaceous floor occupies
c o n ~ p a r a b l e sites of northerly orientation. Outside
the range of spruce forests the north-slope subtype
occupies open northerly slopes.
4. Red oak-pignut hickory forest. Oak-hickory
forests in the Smokies a r e restricted to the lower
slopes of the range, where they occur in draws a n d
ravines, on sheltered northerly slopes, and on lower
sheltered southeait and west s l o ~ e s .
5. Chestnut oak-chestnut forest. Sheltered southDISTRIBUTIONS OF TYPES
erly a n d open northerly slopes a r e occupied by chestTopographic distributions of types on the north- n u t oak-chestnut forests a t lower elevations, but towest slope of the Sniokies may be s u n ~ n ~ a r i z e d :
ward middle elevations the oak-chestnut heath ex1. Cove hardwoods forest. A t low elevations cove p a n d s t o 'eplace them on most open slopes a n d sonie
forests a r e restricted to valleys and lower slopes, sheltered southerly ones. The oak-chestnut forests
with cove foreit transition occupying smaller r a - expand a t the same time t o take the place of oakvines and flats of shallow valleys. Toward higher hickory on sheltered slopes a n d in draws, a n d to adelevations cove forests spread outward on sheltered join cove forest transition. A t 3700 f t chestnut oaka n d northerly slopes, occupying all valley sites ex- chestnut forests occupy a wide range of sites from
cept dry, open draws on south slopes. The u p p e r draws through sheltered slopes to open northerly
limit is near 4500 f t , where either spruce or beech slopes; above t h a t elevation they a r e replaced by red
usually becollie. dolninant. The cove forest transition oak-chestnut forests.
extends from its lower sites upward on the xeric cide
6. Chestnut oak-chestnut heath. At lower cslelaof hemlock, to become above 2500 f t a transition be- tions the t y p e occupies intermediate slopes. Toward
tween hemlock and oak-chestnut forests.
tl~iddleelevations, above 2000 f t , i t expands to occupy
most open slopes except south and south\vest ones.
Toward higher elevations chestnut oak-chestnut forests occupy sotl!e of the open slopes, a n d above 3700
f t the oak-chestnut hcaths d r o p out xvith expansion
of o a k - c h e ~ t n u t forest and replacenlent of chestnut
oak by red and white oaks.
7. Red oak-chestnut forest. Red oak-chestnut f o r ests occupy sheltered southerly slopes. more xeric
than those occupied by heinlock, and open northerly
sloues above 3700 f t . Toward higher elevations thev
Fro 31. Topographic disposition of vegetatlou types.
occur on open slopes w h ~ l e hen~lock occupies the
V l e ~ r of Idealized mountaln and valley, looklng east,
s
h ~ l t e r e dones, a n d above 4500 f t red oak-chestnut
\\lth 6500 f t peak bearnlg subnlpnle forest on left,
lo\\er 5500-ft peak covered up to summit bald n i t h de- occupies open intermediate a n d southerly slopes a s
ciduous forest on rlght. Vegetation types:
beech forests occupy open northerly slopes. Outqide
BG-Beech
Gap
OH-Oak-Hlckory
Forest
the range of spruce, high cols and ridges, except the
CF-Cove Forest
P-Pine
Forest a n d P i n e
niost exposed ones, bear red oak-chestnut.
F-Fraser
F i r Forest
Heath
8 White oak-chestnut forest. Slopes more xeric
GB-Grassv
Bald
ROC-Red
Oak-Chestnut
than those corered by red oak-chestnut a r e occupied
H-Hemlock
Forest
Forest
HB-Heath
Bald
S-Spruce
Forest
b~ red and a h i t e - or white oak-chestnut foreits. BeOCF-Chestnut
OakSF-Spruce-Fir
Forest
l o ~ v4500 feet white oak-chestnut occurs on some open
Chestnut Forest
WOC-White
Oak-Chestnut
southeast a n d west slopes. Above 4500 f t open south
O('H-Chestnut
OakForest
Chestnut Heath
and southwest slopes and sonie exposed ridge. ancl
summits s u p p o r t white oak-chestnut.
2. Eastern hemlock forest. A t elevations below 3600
9. Virginia pine forest. South- and south\vest-faof t hemlock stands occupy valley flats which a r e ele- ing slopes and summits of low hills bear Virginia
vated above or remote frorn the stream. Toward pine forests, and successional stands a r e frequent
higher elevations, as cove forest occupies these sites, on other sites. Virginia pine extends above 2300 f t
henilock moves u p on sheltered northerly slopes and, but seldoni doiiiinates stands above t h a t elevation.
10. Pitch pine heath. Open south and southwest
still higher, onto sheltered southerly slopes and ridges.
R. H. WHITTAKER
60
One aspect, a most interesting one, of the distribution of types remains. I n the northeast half of the
range sites above 4500 f t , n i t h some exceptions, are
occupied by forests of spruce and fir. South\\-est of
the middle of the range, not f a r beyond Clingnians
Dome, spruce-fir forests are absent from the high
mountains even though considerable areas above 4.500
f t exist there. I n the southwestern Smokies the place
of the subalpine forests is taken by high-elevation
deciduous forests-beech-mixed
and beech in mesic
sites, red and white oak-chestnut in more xeric ones.
The change between the two halves of the range is
not one of gradual tapering away and dilution of the
spruce-fir forests toward the south but of their presence, with full dominance, i n one area and conlplete
absence in the other. Climates in the two halves of
the range cannot differ greatly, for vegetation patterns below 4500 f t are the same. Precipitation above
4500 f t may be less i n the southwestern half of the
range, but there is no reason to doubt that the high
slopes of these mountains could support spruce-fir
forests. The fact that they do not do so is a striking
example of failure of physiognomic convergence of
vegetation in similar cliriiates (cf. Beadle 1951).
EcologicalVol.
Monographs
26. No. 1
FIG.22. Profile of Great Smoky Mountains to illustrate hypothesis on southern limit of Appalachian
spruce-fir forests. Distribution of spruce-fir forests on
higher peaks in northeast half of the range is indicated
by diagonal shading. Grassy balds are indicated by
black caps on lower peaks of the southwest half of the
range, otherwise covered by deciduous forest. Climatic
warming during the xerothermic period sufficient to displace the lower limits of spruce-fir forests upward from
4500 f t to approximately 5700 f t would account for
present distribution of these forests.
January, 1956
VEGETATION
OF THE GREAT
SMOKY
BIOUNTAINS
61
e s t ~are niore narrowly restricted to valleys, and pine woods and hemlock hardwoods in mesic sites (swamp
stands are much more limited. The apparent incon- types al\o occur), red oak and chestnut oak forests
sistency, with both niesic and xeric types reduced, on slopes, ~vhite oak-pignut hickory on northerly
should be dne to effects of the niountains thenuelves faces of hill tops, and pitch pine-scrub oak (Q. ilicion climate. Because of the rain-shadow effect, there folia) on southerly faces. Southern exposures of
is less precipitation to the southeast to support ineiic suiilmits have small areas of natural meadow intervalley types; but the open slopes and ridge\ are spersed with srrub oak, suggestive of grassy balds.
somewhat protected froni the ~vesterlywinds which Because in the Smokies an Appalachian vegetation
contribute to the dryness of pine sites on the north- pattern of maximunl diversity appears, intermediate
west side. The result is an expansion of intermediate, to the mixed nlesophytic and oak-chestnut prevailing
oak-chestnut types a t the expense of the extreines on cliniaxes of the Cumberlands and Blue Ridge, the
the southeast, as coinpared with the northwest side author has regarded the vegetation of this range as
of the range. Farther east of the Sniokies in the "central" to that of other Appalachian Mountains
Blue Ridge, the oak-chestnut grouping is further not too distant.
expanded as a prevailing climax, and mesophptic
The Southern Appalachians have long been recogconrmunities are very restricted.
pized (Adams 1902; Harshberger 1903, 1911:198,
West from the Smokies in the Cuniberland Moun- 209-211) as a geographic center for the forests of the
tains, the mesophytic portion of the pattern expands eastern r n i t e d States. I n the interpretation of Braun
to become prevailing climax, while oak-chestnut and the Appalachian center has been given special proniipine types are much restricted (see also P a r t 1 1 ) . nence, and the hlixed Ilesophytic Region in and
Seen from the Sniokies the vegetation of the Cuiiiber- around the Cuinberland hlountains is regarded as the
lands is a sornenhat reduced, niore highly niesophptic center. I t is Braun's thesis (Braun 1950 :39) that
version of the Sniokies pattern, to which are added "The hliuecl hlesophytic association, which characlo\\--elevation types not recognized in the Snlokies. terizes this region, is the niost conlplex and the oldest
Seen from the Smokies the vegetation of the Blue association of the Deciduous Forest Forniation. I t
Ridge is a coinparable transforination by expansion occupies a central position i n the deciduous forest as
of the oak-chestnut types and contraction of the a whole, and from it or its ancestral progenitor, the
inesophytic coinniunities. Seen from the Cumber- mixed Tertiary forest, all other climaxes of the delands the pattern of the Smokies represents a re- riduous forest have arisen." The choice of which of
duction of the mesophytic part of the pattern and two ranges of the Southern Appalachians is to be
expansion of oak-chestnut, and a transition toward rleiignated as central is not a inatter of great niothe oak-chestnut prevailing climax of the Blue Ridge inent. With due respect for the case for the Cumberand beyond.
lands tleveloped by Braun, certain other consideraChanges In vegetation patterns along clin~atic tions related to the Smokiei and the ITnaka Moungradients and from one lnountain range to another tain\ of which they are part, may be mentioned.
I. "Centers" are products of h u ~ n a ninterpretation
may be interpreted on the basis of changes in relative
- - ~
i
i
62
R. H. WIIITTAKER
Ecological Monographs
Yol. 2 6 , No. 1
and, like such other concepts as "association" and fixed intervals along environmental gI*adients, and
"clinlatic clinlax," should not be taken f o r granted. site-samples taken widely through the vegetation
Designation of a center JllaY be thought llleaningful pattern as an approach toward randomization, were
only as the criterion or criteria (Adanls 1902, Cain used as means of vegetation sampling unprejudiced
by assun~ptionsabout associations.
19-14) one chooses to define a center are stated.
2. There is no reason, a priori, for locating a center
2. Arbitrary classes or ecological groups of tree
in an area possessing a well-defined prevailing or species and weighted averages based on these classes
elinlatic clinlax. I f , on the other hand, diversity is Tvere used for indication of relative positions of
regarded as a primary criterion, then location of stands along the nloisture gradient. The site-sanlples
centers in areas like the Srliokies with yegetation too were arranged into composite transects, showing the
varied to possess a well-defined prevailing climax is relation of plant populations to environmental gradiperhaps to be expected.
ents. Major environnlental gradients, those of eleva3. Designation of one area, the Smokies, as transi- tion and lnoisture conditions of site, are conceived
tional between other areas which are non-transitional as c o m p l e x - g r a d i e n t s of rnany interrelated factors
seenis of little intrinsic meaning. Areas of great affecting plants.
vegetational diversity are 111ost conreniently regarded
3. The sequence of vegetation types from nlesic
as transitional in a fralllework of thought which in- sites to xeric at low elevations is: cove forest (mixed
eludes climatic clinlaxes and clinlax regions as major n~esophytic),oak-hickory forest, oak-chestnut forest,
concepts. These concepts should not necessarily be oak-chestnut heath, and pine forest. Deciduous trees
other than oaks predonlinate in nlesic sites, oaks in
applied to the biogeographic proble111s of "centers."
4. A traditional
criterion of a center, intermediate sites, and pines in xeric sites; but relative proportions of these growth-forms change conthough not the only possible one, is biotic diversitytinuously along the moisture gradient through the
richness in species in general, or in a particular
group being studied. Some secondary criteria are various types. Along the salne gradient, stature of
features usually associated with areas of ~naxii~lum trees and canopy coverage in general decrease; but
biotic diversity-age of the area, \vide range of habi- the number of tree stems is greater in xeric sites than
tats, and occurrence of narrow endemisin anlong its ill mesic. At all points along the gradient1 undisturbed forest stands are self-maintaining; and a
species.
5 , The greater ranges of elevation and nloisture curve relating stem nunlbers and dianleters in these
climax stands is suggested.
or
conditions of the Slllokies imply greater floristic dias greater .r,ariety of vegetation types. richness in species, of the tree stratum is 111axin1ali n
versity, as
low-elevation stands transitional between cove forThis fact, as Well as the secondary criteria
point to the slnokies rather than the cumber- e s t s a n d oak forests; and diversity values decrease
with increasing departure from these stands t o ~ ~ a r d
lands as "central,, for the southern Appalachians.
Both ranges are doubtless to be regarded as parts more mesic o r more xeric conditions and toward
of the broader Southern Appalachian center. Within higher
4. Coverage of the shrub stratulll in general inthis area the different criteria of B~~~~ and the aucreases
along the nloisture gradient toward xeric
thor lead rather naturally to their different eruphasis
of the ~
~
~ and ~the Snlokies,
b
~ A nlajor
~
lcon- ~ sites.~ Deciduous,
d
~ non-ericaceous shrubs predotninate
in
niesic
sites,
evergreen
ericaceous shrubs in intermetern of the present paper is With vegetation
in relation to environmental gradients, and it is diate sites, and deciduous ericads (Vaccinioideae) in
thought that in the Snlokies the eastern forests shall- xeric sites. With increase of shrub coverages
their greatest diversity and differentiation into types the gradient, herb coverages decrease; Coverages f o r
Within a
area. The pattern is doubtless one of the two strata are inversely related. Ferns and delithe lllost conlplex on the continent. Also in the Siski- cate, unlbrella-shaped herbs prevail in mesic forests
you $fountains
Klamath &go.in,
somewhat
hut decrease in coverage along the
gradient
parable forest center of the mest, an ohserver is at as other herb grolvth-forms (rosette plants, grasses,
first overwhelnled by variety of vegetation types; hut ground heaths, etc.) increase. Herb coverages in gengradients and eral increase toward higher elevations, but shrub
this variety is expressive of
variety of soil Illaterials as well as of elevation and coverages are more conlplexly related to elevation.
5 . I n ~ u b a l p i n eforests Picea r ~ b e n sis dominant
local llloisture conditions. The author knows no pat4500 f t u p through the lower part of the forterll of clinlax vegetation, occurring within a limited
area on relatively
soil
lllore varied est" A b i e s f r a s e r i is dominant above 6000 f t . Subalpine forests show lin~iteddifference in canopy comthan that of the Great Srnoky Nountains.
position along the nloisture gradient but striking difSt-JIMART
ference in undergrowths. The satne herbs and shrubs
as those in mesic deciduous forests appear in valleys;
I. Gradient Analysis
a coniplex undergrowth of. five strata (Hylocomium,
1. Vegetation of a range of the Southern Appa- Oxalis, Dryopteris, Vacciniuln, and Viburnum) oclachians, the Great Smoky Mountains of Tennessee, curs on north and east slopes and decreases in coverwas analyzed. Field transects, with saniples taken a t age around to \Test and south slopes; and a dense
patterns
January, 1956
VEGETATION
OF THE GREATSMOKY
R~OUNTAINS
63
64
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Ecological Monographs
Vol. 26, No. 1
J a n u a r y , 1956
VEGETATION
OF THE
GREATSMOKY
RIOUNTAINS
65
66
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Ecological Monographs
Vol. 2 6 , No. 1
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the virgin forest of the north t e m l ~ e r a t e zone. Yew
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VEGETATION
OF THE ORE
Ecological Monographs
Vol. 26, No. 1
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Wangerin, W. 1925. Beitrage zur pflanzensoziologischen
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Ward, R. D. 1925. The climates of the United States.
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Weaver, J. E. & F. E. Clements. 1929. P l a n t ecology.
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-----. 1953. A consideration of climax theory: The
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January, 1966
VEGETATIOX
OF THE GRE
--
R. H. WHITTAKER
70
) Acer pensylvanicum
A b i e s fraseri
stems
stems
8
50
!
1mLATS
MAS
SIJlJ~
FYI?
HWfE
Ecological Monographs
VoL 26, No. 1
January, 1 9 5 6
VEGETATION
OF THE GREAT
SMOKY
~IOCNTAINS
Betula allegheniensis
stems
65
B e t u l a lenta
stems
65
71
72
Ecological Monographs
Vol. 26. No. 1
January. 1956
VEGETATION
OF
THE
GREATSMOKYMOUNTAINS
Cladrastis Iutea
Cornus florida
stems
73
74
R. H. WHITTAKER
Ecological Monographs
Val. 26, No. 1
January, 1956
VEGETATION
OF THE GREATSMOKY
NOUNTAINS
75
R. H. WHITTAKER
76
stems
Ecological Monographs
Tol. 26, No. 1
January. 1956
VE~ETATION
OF THE GREATSMOKY
MOUNTAINS
77
Pinus r i g i d ~
stems
BS00-
6000[ ~ E A L
5MO-
WTSI)
p2i
78
R. H. WHITTAKER
Ecological Monographs
Vol. 26, No. 1
January, 1956
VEGETATIOS
OF THE GREATSMOKY
~\IOUSTAISS
79
80
R. H. WHITTAKER
Ecological Monographs
Vol. 26, No. 1