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STUDIES OF
THE VEGETATION OF MISSOURI— I

NATURAL PLANT ASSOCIATIONS AND
SUCCESSION IN THE OZARKS OF MISSOURI

BY

JULIAN A. STEYERMARK

ASSISTANT CURATOR OF THE HERBARIUM, DEPARTMENT OF BOTANY

THI "«***<*

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

VOLUME IX, NUMBER 5

DECEMBER 31, 1940

PUBLICATION 485

PRINTED IN THE UNITED STATES OF AMERICA
BY FIELD MUSEUM PRESS

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CONTENTS
LIST OF ILLUSTRATIONS 351

INTRODUCTION 353

NATURAL PLANT ASSOCIATIONS AND SUCCESSION IN THE OZARKS OF MISSOURI . 353

I. Sugar maple-bitternut hickory association 358

II. Sugar maple-white oak association 372

III. Oak-hickory association 387

IV. Oak-pine association 393

V. White oak-red maple association 397

SUBCLIMAX STAGES IN THE OZARKS 401

PLANT INDICATORS OF ACIDITY AND ALKALINITY 404

ECOLOGICAL VARIATIONS INDUCED BY BURNING, LOGGING, CLEARING, AND

OTHER UNNATURAL CAUSES 408

EXAMPLES TAKEN FROM SPECIFIC AREAS SHOWING PARTICULAR STAGES

WITHIN AN ASSOCIATION . .421

349

LIST OF ILLUSTRATIONS

TEXT FIGURES PAGE

9. Salix longipes-Hamamelis vernalis associes on gravel bar at mouth of

Indian Creek and White River, near Baxter, Stone County 359

10. Salix longipes associes on gravel bar, four miles northeast of Waynesville,

Pulaski County 360

11. Dianthera americana forming dominant herbaceous growth along gravel

bar on Bourbeuse Creek at Glaser Ford, three and one-half miles south-
west of Bern, Gasconade County 361

12. Salix nigra, instead of Salix longipes var. Wardii, pioneering on gravel.

Along Moreau River two miles southeast of Jefferson City, Cole County 362

13. Deposit of muddy alluvium above sand on bar on Little North Fork of

White River, one and one-half miles south of Theodosia, Ozark County . 363

14. Acer saccharinum-Populus deltoides associes on right side behind gravel

bar, between Reed Springs and Viola, Stone County 364

15. Ulmus americana-Fraxinus pensylvanica var. lanceolata associes on

alluvium of valley along Jacks Fork of Current River, north of Monteer,
Shannon County 365

16. Along Huzzah Creek, two miles southeast of Davisville, Crawford County . 371

17. Limestone glade along Niangua River, two miles south of Corkery, Dallas

County 372

18. Limestone glade with dominants of Silphium laciniatum, Psoralea tenui-

flora, and Bonteloua curtipendula on bluffs along Osage River, two
miles northwest of Wisdom, Benton County 373

19. Red cedar glade or bald on dolomitic limestone, near Taney County line,

in Stone County 374

20. Bald knob west of Table Rock, west of Branson, Taney County .... 375

21. Diospyros virginiana initiating woody growth on limestone glade along

Niangua River, two miles south of Corkery, Dallas County 376

22. Quercus Muhlenbergii invading an associes with Diospyros virginiana and

Bumelia lanuginosa along the bluffs of Osage River (Lake-of-the-
Ozarks), at Riverview, Morgan County 377

23. Quercus Muhlenbergii-Fraxinus americana associes dominating grassy

slopes along headwaters of Bryant Creek, two miles southeast of
Cedar Gap, Wright County 378

24. Bald knob, east of McClurg, Ozark County 379

25. Similar to Fig. 24, three miles northwest of Vichy, Maries County, except

that forest of Acer saccharum-Quercus alba associes has more nearly
closed the originally open glade 380

26. Acer saccharum-Quercus alba climax associes well developed over a lime-

stone substratum, Taney County 381

27. Bluffs along White River just west of Shreiner's Ferry, Barry County . 382

28. Bee Bluff, along Current River, northeast of Eminence, Shannon County . 383

29. Bee Bluff, along Current River, northeast of Eminence, Shannon County . 385

30. Limestone bluffs along Eleven Points River, near mouth of White Creek,

Oregon County 386

31. A typical Ozark landscape, south of Harley, Barry County 388

32. Porphyritic trachyte glade of Red Rock Mountain, near Irondale,

Washington County 389

351

352 LIST OF ILLUSTRATIONS

PAGE

33. Rhus Copallina invading prairie habitat, one mile from Riverview bluffs,

Morgan County 390

34. Upland woods with dense, grassy cover near head of Mud Creek, twelve

miles north of Poplar Bluff, Butler County 391

35. An oak-hickory associes on a flat upland near Dillard, Crawford County . 392

36. Pinus echinata developed in almost pure stand on Pine Bluff, overlying

Roubidoux sandstone along Meramec River, four miles south of
St. Clair, Franklin County 394

37. Pure pine stand on ridge top north of Bunker, Reynolds County . . . 395

38. South-facing limestone bluffs along White River opposite Table Rock,

Taney County 399

39. Right half of picture in background in white is covered by Juniperus

virginiana overlying a limestone substratum; left half of picture mixed
oak and pine overlying chert substratum. Along highway No. 19, two
miles north of Eminence, Shannon County 400

40. Right half of picture in background in white is covered by Juniperus

virginiana overlying a limestone substratum; left half of picture mixed
oak and pine overlying chert substratum. Along highway No. 19, two
miles north of Eminence, Shannon County 402

41. Effect of a spring burn, showing a very light density of forest floor cover-

age. Note the open patches of soil between the herbs. Twelve miles
north of Poplar Bluff, Butler County 409

42. Damage done by summer burn, along highway No. 49, just south of

Black, Reynolds County 410

43. Damage done by summer burn, along highway No. 49, just south of

Black, Reynolds County 411

44. Flat ridge top in Quercus alba-Quercus stellata woodland, four miles north-

west of Bunker, Reynolds County 413

45. Quercus alba-Quercus velutina-Quercus stellata woodland on flat ridge top,

four miles northwest of Bunker, Reynolds County . . 414

STUDIES OF THE VEGETATION OF MISSOURI— I

JULIAN A. STEYERMARK

INTRODUCTION

The present paper is the result of observations and records made
by the author while collecting and making a systematic study of the
flora of Missouri during the past fifteen years. He does not profess
to be a trained ecologist in the modern usage of that term. As a
taxonomist who, in making a systematic study of the flora, has
collected in every county of the state in order to make the thorough
phytogeographical survey that was necessary to the understanding
of the distribution and limitations of species and varieties, he has
observed, collected, and recorded the flora found in every conceivable
type of association (in the usual sense employed for this expression)
in the state. Specimens of all the plants mentioned in the present
paper may be found in the Herbarium of the Missouri Botanical
Garden and in the Herbarium of Field Museum. Such specimens
form a concrete and definite basis for the observations made in this
paper. Moreover, they serve as the only tangible evidence whereby
the statements made by the author may be verified or refuted.

The terms "association," "consociation," "associes," "society,"
"climax," and "dominant" used in the present paper are to be
applied in the ordinary and usual sense in which they are employed
generally by ecologists in the United States, and agree with the con-
cept of these terms as originally given by Warming,1 Weaver and
Clements,2 and many others. The 164 examples given at the end
of the paper represent actual areas of ground surveyed in detail by
the author in making observations within the Ozarks. They may
be considered as large-scale, natural plots selected because they
represent typical cross sections which occur in diverse areas within
the Ozark region.

NATURAL PLANT ASSOCIATIONS AND SUCCESSION
IN THE OZARKS OF MISSOURI

Some of the stages of succession noted here will be found in parts
of the state other than the Ozarks, for example, in the glaciated

1 E. Warming, Oecology of plants. 422 pp. Clarendon Press. Oxford. England.
1909.

1 J. E. Weaver and F. E. Clements, Plant ecology. 520 pp. McGraw-Hill
Book Company, New York. 1929.

353

354 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

prairie region north of the Missouri River and outside the Ozarks,
or in the southeastern lowland area, or in the unglaciated prairie
region of west-central and southwestern Missouri.

Attention should be called to the fact that the past and intricate
geological history of the Ozark region has been one of the chief factors
in causing the migration and present distributional status of many
of the species. For example, the present distribution of the bald
cypress and other southeastern lowland species in Missouri is de-
limited by the incursion of the Mississippi embayment in past geo-
logical periods. Again, the present northern and northwestern limits
of the southern yellow pine and species with similar distribution in
the Ozarks of Missouri do not result primarily from any factors of
the soil or climate of the present, since similar conditions are
encountered in northern Missouri, but rather from factors of past
geological history, such as the extermination by glaciers of the Ozark
flora north of the Ozark region, or a climatic disturbance causing
a block or change in migration, or elimination of habitats previ-
ously suitable to the species but now exterminated north of the
Ozark region.

The Ozark flora is an extremely varied one, and its composition
has been made much more intricate and complex by the uplifts and
peneplanations of the land, varied rock formations, and long history
as a land area since the end of the Paleozoic era. This accounts for
the great number and variety of stages of development within a
given plant association.

Reference is here made to Plant ecology by Weaver and Clements.
It will be found that a number of conclusions reached in this report
differ in some ways from those made in that publication. The reasons
for this are several. In the first place, no critical study of the natural
plant successions in the Ozark flora has previously been attempted.
Thus, while some general statements in Weaver and Clements' book
hold true, a number of large generalizations made in that work to
cover the region in which the Ozarks are located, concerning climax
formations and associations, do not hold. Further than this, these
authors, by stressing the physical factors of the soil, have not suf-
ficiently considered variations in the chemical nature of the soil or
the past geological history of the region as factors which might cause
different stages in development of an association or account for the
appearance of certain consocies and dominants. Thus, for example,
the maple-beech forest is regarded by them as a typical association
of the deciduous forest climax formation; but, in Missouri, this

STUDIES OF THE VEGETATION OF MISSOURI — I 355

formation is found only in the southeastern portion, and mainly on
Crowleys Ridge, the beech being limited to this part of the state on
account of the past geological history and the meanderings of the
Mississippi River and the relation of this to the Crowleys Ridge area.
The beech could, on the basis of soil and climatic conditions and its
shade tolerance, grow farther west where the maple is found, but
past geological history has restricted it to its present area. In the
Ozarks the sugar maple (Acer saccharum) forms a climax with some
of the oaks (Quercus alba and Q. borealis var. maxima) along rich,
wooded slopes and bluffs, rather than with beech. Furthermore, the
sugar maple climax is not reached unless the soil ranges from neutral
to alkaline, for where acid soils exist, other climax species, such as
Quercus coccinea and Pinus echinata, are found. In the Ozarks the
acidity of the soils plays an important part in causing different stages
and units of succession, and many of the associes owe their appearance
in the development of the vegetation to the particular stage of sub-
stratum decomposition or formation. Thus, in the Ozarks, the basis
for the appearance and difference in the oak-hickory or pine-oak
association on the one hand and the sugar maple association on the
other, lies in the acidity and chemical nature of the soil, rather than,
as Weaver and Clements maintain, in the physical or moisture con-
dition of the soil. The limiting factor, in other words, appears to be
the presence or absence of limestone, rather than soil moisture and
shade. This will be discussed further in this paper.

In Missouri different types of soils within a limited region appear
to result in entirely different associations and climaxes. For so
limited an area this conclusion is at variance with the single climax
type admitted for the region included within the Ozarks by Weaver
and Clements. However, some of the Old World ecologists appear
to confirm the conclusions reached by the present author. Du Rietz,1
for example, points out that "contrary to the monoclimax theory
developed by Clements and Braun-Blanquet, I find it necessary, how-
ever, to maintain the polyclimax theory developed by Domin, Gams,
Gleason, Nichols, Nordhagen, Scharfetter, Tansley, myself, etc., i.e.,
to admit that several climax-phytocoenoses — in many cases with very
little relationship to each other — may occur side by side in the same
vegetation region, but in edaphically different habitats. . . . My own
field experience certainly does not support the theory of the power
of a uniform climate to transform all these edaphically different
habitats into one comparatively uniform 'climax-habitat' ('climax-

1 G. Einar du Rietz, Classification and nomenclature of vegetation. Svensk.
Bot. Tidskr. 24, H. 4, p. 497. 1930.

356 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

soil/ etc.), as postulated by the monoclimatic theory." The author
continues his discussion by showing that in various places in Europe
many climaxes are developed in the same region. He states1 that "in
the Quercus region of south Sweden, for instance, certainly both
Quercus forests and Picea-Pinus forests, among others, must be
admitted as climax-phytocoenoses, the former on better and the
latter on poorer soils. In the case of the alpine vegetation regions of
the Alps and of Scandinavia, there is a complete agreement among
Scandinavian ecologists that the dominating dwarf-shrub-heath and
grass-heath of the acid and moderately snow-covered ground in the
low-alpine and middle-alpine belts respectively, are by no means the
only climax-phytocoenoses of these regions as assumed by Braun-
Blanquet. Parallel with them there also occur not only subneutro-
philous (pH 5.5-7) and neutrophobous climax phytocoenoses both
chionophobous (on wind-exposed ridges) and chionophilous (on snow-
loving, i.e. 'Schnee-boden' communities), consisting of grass, herb,
dwarf-Safe, moss, and lichen communities very different from the
dominant subchionophobous heath."

Tansley2 states that "the so-called 'polyclimax theory' takes
what appear to be permanent types of vegetation under given con-
ditions and calls them climatic conditions of the region and on the
most favorable soils the climatic climax is reached by the succession;
but that on less favorable soils of special character different kinds of
stable vegetation are developed and remain in possession of the
ground, to all appearances as permanently as the climatic climax.
These are called edaphic climaxes, because the differentiating factor
is a special soil type." Other climaxes are described, caused by
physiography, grazing, fire, and mowing. Of these he says,3 "In
each case the vegetation appears to be in equilibrium with all the
effective factors present, including of course the climatic factors, and
the climax is named from the special factor differentiating the vegeta-
tion from the climatic climax. . . . The word climax is used in its
simple and natural signification of a culmination of development —
a permanent or apparently permanent condition reached when the
vegetation is in equilibrium with all the incident factors." Speaking
of the relation of soil on succession, Tansley states3 that while it is
simple to believe that "the soil profile runs pari passu with the de-
velopment of the vegetation it bears, and that consequently the

1 Ibid., p. 499.

2 A. G. Tansley, Vegetational concepts and terms. Ecology 16: 292. 1935.

3 Ibid.

STUDIES OF THE VEGETATION OF MISSOURI — I 357

mature climatic soil type corresponds and co-exists with the climatic
climax community," yet he goes on to state that this is not true,
because "even when profile development under the influence of
climate is perfectly normal and regular, the climatic climax com-
munity may establish itself long before the soil is mature, and may
not be substantially altered by the later stages of profile maturation.
Again a climatic climax may establish itself on a soil which is kept
immature by geological and geographical causes, as on a steep slope.
And finally it is now generally agreed by pedologists that some rocks,
owing to the simplicity of their composition, produce soils which can
never form the normal climatic mature profile, and these may or may
not bear the typical climatic climax vegetation." (The italics are the
present author's.)

The field experience of the author in his observations on the
Missouri flora is in full agreement with the statements quoted above
by du Rietz and Tansley. The Ozarks offer a wonderful example in
soil diversities and give ample opportunities of corroborating the
opinions of these ecologists concerning the influence of soils on the
various types of vegetation within a small region. Thus, we find
that on the acidic substrata of chert, sandstone, or granitic rock an
associes culminating in oak, hickory, southern yellow pine, Acer
rubrum, Nyssa sylvatica var. caroliniana, and Cornus florida is as char-
acteristic a climax association as is that of Acer saccharum, Quercus
Muhlenbergii, and other species developing on a limestone soil. The
fact that several climaxes may exist in the Ozarks is proven by these
permanent, well-marked associations which have originated from
different substrata, yet may exist, as they often do, contiguous to
each other. The fact remains that they are not climatic climaxes,
as argued by Weaver and Clements, because the climate has not
differentiated these diverse, stable associations, one on limestone,
the other on chert, sandstone, or porphyritic trachyte or granite.
Rather, these climaxes are edaphic and owe their diversity to their
separate selections of soil conditions.

Friesner and Potzger1 concluded, after a study of certain areas
in Indiana in which soil reactions between the Fagus-Acer and Quer-
cus-Carya communities were compared, that "the range of reaction
in surface soils for beech-maple does not show much difference from
that of oak-hickory, except that the reaction spread is a little nearer
alkalinity in beech-maple, extending from pH 4.5 to 7.53, and a little

1 Ray C. Friesner and J. E. Potzger, Contrasts in certain physical factors in
Fagus-Acer and Quercus-Carya communities in Brown and Bartholomew counties,
Indiana. Butler Univ. Bot. Stud. 4: 8-9. 1937.

358 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

more acid in oak-hickory, extending from pH 4.1 to 7.24. . . . The
majority of the soil samples gave a decidedly more acid reaction for
oak-hickory than was found in any of the beech-maple areas except
one. It will also be seen that while the oak-hickory range exhibits as
wide a spread as the beech-maple, the majority of the samples fall
within a much narrower range which is more acid than the majority
of the readings for the beech-maple soils." As will be shown in the
following portion of this paper on plant associations in the Ozarks,
the oak-hickory association develops upon an acid substratum, while
Acer saccharum develops from a limestone substratum and Acer
rubrum from an acidic cherty, sandstone, or granitic or porphyritic
trachyte substratum.

In Weaver and Clements' Plant ecology, three associations, i.e.,
the oak-hickory association, maple-beech association, and oak-
chestnut association comprise the three associations in the deciduous
forest climax formation, while in this report five associations are
described.

The climax formation in the Ozarks is a mesophytic forest. The
climate is of the type which effects such a mesophytic development.
This mesophytic climax forest is made up of various associations.

Units for discussion in the Ozarks have arisen from the following
sources: (1) stages in flood-plain formation; (2) ridge tops and plateau
areas; (3) ravines and upper drainage heads of the tributaries in acid
soil areas; and (4) stages in cover over an eroded limestone bluff or
open glade. Some of these associations, therefore, depend on the type
of rock substratum from which they start. Thus, the development
or succession of the vegetation is closely correlated with the type of
rock in the particular area under discussion.

The five associations included within this climax formation are:
(I) sugar maple-bitternut hickory association; (II) sugar maple-
white oak association; (III) oak-hickory association; (IV) oak-pine
association; and (V) red maple-white oak association.

I. SUGAR MAPLE-BITTERNUT HICKORY ASSOCIATION

(Acer saccharum-Carya cordiformis)

(1) The first association, the sugar maple-bitternut hickory
(Acer saccharum-Carya cordiformis) association, is the climax stage
reached from a development of the flood-plain flora. The develop-
mental stages or associes of this association are definitely correlated
with the development and maturity of the stream. Most of the
smaller Ozark streams are still in some stages of youth and conse-

STUDIES OF THE VEGETATION OF MISSOURI — I 359

quently are dashing and swift-flowing types, carrying loads of chert
which have been eroded and borne from the hills. At the stage when
the stream is still small and youthful, with a rocky bed loaded with
gravel deposits, occurs the first developmental stage, the Ward
willow-witch hazel (Salix longipes var. Wardii- Hamamelis vernalis)
associes. This shrubby willow pioneers with the witch hazel along
all the smaller, rock-laden streams or along the gravel bars or rocky
deposits of streams which are in a more mature stage of develop-
ment (fig. 9). In the northern and western border of the Missouri
Ozarks, in much of the area of the Niangua, Osage, lower Gasconade,

FIG. 9. Salix longipes-Hamamelis vernalis associes on gravel bar at mouth
of Indian Creek and White River, near Baxter, Stone County.

and Pomme de Terre rivers, the witch hazel is absent, leaving Ward's
willow as the sole dominant of the first stage of the flood-plain
association (fig. 10). Witch hazel is found all over the eastern and
southern Ozarks.

Sycamore (Platanus occidentalism is always a conspicuous mem-
ber (consocies) of the willow-witch hazel associes. It is particularly
dominant on gravelly and sandy deposits. Along the gravel bars
and sandy deposits of the larger Ozark streams such as the Current
River, Piney River, Gasconade River, Black River, White River,
and others, it and Ward's willow are the pioneers, while alder and
usually witch hazel are absent. Alder (Alnus rugosa) is often
associated with the willow, witch hazel, and sycamore, but is com-
monest along the smaller tributaries and headwaters. These four
species (Ward's willow, witch hazel, sycamore, and alder) make up
the dominants or consocies. Sometimes the sand grape (Vitis rupes-

360 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

tris), ninebark (Physocarpus opulifolius var. intermedius) , and
Cornus obliqua are associated with these. Water willow (Dianthera
americana) is a conspicuous herbaceous dominant in the willow-
witch hazel associes and is found in dense colonies, rooting along the
gravelly, shallower margins of most of the streams, especially the
smaller ones (fig. 11). Chairmakers' rush (Scirpus americanus) is
often associated with water willow on gravelly areas.

FIG. 10. Salix longipes associes on gravel bar, four miles northeast of Waynes-
ville, Pulaski County.

Many of the extensive gravel bars may be covered with:

Ambrosia elatior var. artemisiaefolia

Ambrosia trifida

Amsonia illustris

Commelina erecta

Cyperus esculentus

Cyperus inflexus

Desmodium sessilifolium

Digitaria sanguinalis (common)

Diodia teres

Eragrostis hypnoides

Eragrostis pilosa

Euphorbia corollata

Euphorbia maculata

Euphorbia supina

Hemicarpha micrantha
Ipomoea hederacea
Lespedeza capltalta
Lespedeza virginica
Leucospora multifida
Lindernia dubia var. major
Mollugo verticillata
Oxalis stricta

Panicum virgatum (common)
Polygonum lapathifolium
Polygonum pensylvanicum
Rorippa islandica var. microcarpa
Xanthium pensylvanicum

The above gravel bar species may be expected to occur on most
gravel bars along the streams, the amount of sand on the one extreme
and muddy alluvium on the other changing materially the aspect of
the composition of the flora.

(2) Farther away from the margin of the rocky stream a richer
soil is gradually built up by successive alluvial deposits (fig. 13).

STUDIES OF THE VEGETATION OF MISSOURI — I 361

A new associes of silver maple-cottonwood (Acer saccharinum-
Populus deltoides) develops here. Other dominants (consocies)
usually found with silver maple and cottonwood are river birch
(Betula nigra), black willow (Salix nigra), and buttonbush (Cepha-
lanthus occidentalis) (fig. 14). As the silver maple-cottonwood
associes gradually gains possession of the better soil, witch hazel
disappears, but sycamore and Ward's willow of the preceding associes
persist into this new succession. Instead of cottonwood with silver
maple, as in the southeastern Missouri Ozarks, one finds green ash
(Fraxinus pensylvanica var. lanceolata) in the western Ozarks along

FIG. 11. Dianthera americana forming dominant herbaceous growth along
gravel bar on Bourbeuse Creek at Glaser Ford, three and one-half miles south-
west of Bern, Gasconade County. Note Salix lonyipes var. Wardii immediately
behind Dianthera colony.

the Gasconade River, where cottonwood is quite rare. In the south-
eastern Ozarks local species such asLiquidambar Styraciflua diversify
the alluvial formation, but along the Gasconade River there are no
unusual species diversifying the area. Along the Gasconade River,
Hamamelis vernalis is rare and local. On Piney River the very infre-
quent cottonwood is associated with silver maple and green ash.

The silver maple-cottonwood associes may completely succeed
the willow-witch hazel associes if the rocky stream habitat of the
latter becomes covered over with a rich alluvium favorable for the
development of the former associes, or the two associes may exist
as contiguous ones in relation to the alluvial formation of the stream.

The birch and the buttonbush sometimes come into the succes-
sion before the black willow, cottonwood, and silver maple have made

362 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

their appearance, and this may be explained by the greater ability
of these two species to seed themselves in more gravelly and sandy
stream bars. Along the Osage fork of the Gasconade River in Web-
ster County, for example, river birch is absent as one of the associes
of a later flood-plain succession. In this region Ward's willow and
sycamore are dominant, being followed by black willow, silver maple,
and green ash. On the Current River silver maple and poplar
pioneer together as associate species in much the same way as they
do in parts of areas of northeastern Missouri, as, for example,
on Salt River.

FIG. 12. Salix nigra, instead of Salix longipes var. Wardii, pioneering on
gravel. Along Moreau River, two miles southeast of Jefferson City, Cole County.

These first two associes of a flood-plain association are subject
to sudden stream overflows and variation of water level. The her-
baceous ground flora is, therefore, not varied, is late in developing,
and is often composed of annuals or water-resistant types. There
is practically no herbaceous or moss associes on the rocky habitat
of the willow-witch hazel associes, but where a relatively large
area of alluvial ground exists, with the silver maple-cottonwood
associes are found a few dominant species on the forest floor, such as
Pilea pumila var. Deamii, Aster lateriflorus var. pendulus, Rudbeckia
laciniata, and white grass (Leersia virginica). Such mosses as
Amblystegium riparium and Leskea gracilescens may be present.

After this stage of the flood-plain has been reached, the succession,
once the deposit of alluvium has been formed, proceeds at a faster
rate and more quickly reaches the climax types.

STUDIES OF THE VEGETATION OF MISSOURI — I

363

(3) The next stage, associated with the formation of richer soil
over the original alluvium, is the American elm-green ash ( Ulmus
americana-Fraxinus pensylvanica var. lanceolata) associes (fig. 15).
Sometimes, in places along the border counties of the Ozarks, Fraxi-
nus pensylvanica var. lanceolata may pioneer on gravel bars with
Salix Wardii, sycamore, and Cephalanthus—as on the Osage River
in St. Clair and Benton counties. Another dominant in this phase
of the succession is Quercus Shumardii (locally called water oak,
otherwise named Shumard's red oak). Along Current and Black
rivers, Quercus Shumardii is common in the Ulmus americana-Acer
saccharinum-Fraxinus pensylvanica var. lanceolata formation, but
on Gasconade and Piney rivers it is quite absent most of the
time and found only occasionally. Bur oak (Quercus macrocarpa),

FIG. 13. Deposit of muddy alluvium above sand on bar on Little North Fork
of White River, one and one-half miles south of Theodosia, Ozark County. Shows
how a sand bar is gradually built up with a consequent new associes developing
upon it.

box elder (Acer Negundo), and a few others also make their appear-
ance in this stage, and in parts of the southeastern Ozarks, as along
the Black and St. Francis rivers, sweet gum (Liquidambar Styraciflua)
is found sometimes in this associes. The herbaceous associes on the
forest floor is somewhat more varied than in the last associes, but
stream overflows continue to occur in this associes likewise. Such
herbs as honewort (Cryptotaenia canadensis), Virginia knotweed
(Polygonum virginianum), beefsteak plant (Perillafrutescens), Iresine
rhizomatosa, and Calycocarpum Lyoni are common, while the grass
flora is dominated by spike grass ( Uniola latifolia), fowl meadow

364 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

grass (Glyceria striata), giant cane (Arundinaria gigantea), and satin
grass (Muhlenbergia mexicana). Along alluvial limestone banks at
the base of bluffs, as along the Osage River in St. Clair County
and elsewhere, Uniola latifolia and Panicum virgatum are dominant,
associated with Apocynum sibiricum and Leptochloa filiformis. This
is equivalent to pioneers on a mud stage, discussed under previous
alluvial forms.

FIG. 14. Acer saccharinum-Populus deltoides associes on right side behind
gravel bar, between Reed Springs and Viola, Stone County.

No Arundinaria gigantea occurs along the Gasconade, Osage,
Piney, Meramec, Big, and Niangua rivers or their tributaries in the
elm-ash stage or in other stages. The Arundinaria begins to appear
on Black, Current, and other rivers where shaded alluvial banks and
thickets occur. Smilax Bona-nox together with Arundinaria is com-
mon along the banks of White River.

After the third developmental stage (i.e. the American elm-
green ash associes) has been attained, Ward's willow and button-
bush, more or less shade-intolerant types, disappear, but sycamore,
cottonwood, and black willow persist. In many alluvial associations
in the western Ozarks, however, no cottonwood is present. Some-
times sugar maple also appears in the elm-ash associes, but does not
become dominant until the last stage of the associes. This elm-ash
associes may last indefinitely, and in some regions where the soil is
flooded frequently and worn away or where the surrounding soil
types are too acid, sugar maple may be prevented from entering this
associes, and the elm-ash associes may become a temporary climax
or subclimax.

STUDIES OF THE VEGETATION OF MISSOURI— I 365

Sometimes, in the Gasconade River region, silver maple and
green ash together with sycamore are followed by birch-elm-ash-
sycamore with maple, the two intergrading and slightly different
from the situation which occurs in the southeastern Ozarks, where
silver maple, cottonwood, and birch are often followed by elm and
green ash, with birch, sycamore, and maple lingering over into the
newer associes.

FIG. 15. Ulmus americana-Fraxinus pensylvanica var. lanceolata associes on
alluvium of valley along Jacks Fork of Current River, north of Monteer, Shannon
County. On top of the bluffs on the right are pine trees occurring on cherty and
Roubidoux sandstone substrata.

In the southeastern Ozarks the usual succession is:

(1) Ward's willow, witch hazel, sycamore, alder; succeeded by

(2) silver maple, poplar, birch, black willow; succeeded by

(3) elm, green ash.

In the White River country it is:

(1) Ward's willow, witch hazel; succeeded by

(2) black willow; succeeded by

(3) silver maple, birch, elm, green ash.

In the Gasconade River area it is:
(l)Ward's willow; succeeded by

(2) sycamore, black willow; succeeded by

(3) silver maple, green ash, birch; succeeded by

(4) elm, silver maple.

But the end result is in all cases usually the same, i.e. with elm,
silver maple, and green ash comprising the dominants in the associes.
The elm comes in last in each case, but may have ash with it first or
last, the ash being the varying pioneer.

366 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Variations in Stage No. 3:

(a) On Piney Creek and other areas in Table Rock National
Forest Purchase Unit in Barry and Stone counties, southwestern
Missouri, instead of alder occurring with witch hazel in stage No. 1,
Ilex decidua is found with witch hazel. The latter two plants are
the dominants, together with such species as Ostrya virginiana,
Cornus obliqua, Cornus asperifolia, and Physocarpus opulifolius var.
intermedius (example No. 5).

(6) On Table Rock National Forest Purchase Unit in the White
River country of southwestern Missouri, swamp privet (Forestiera
acuminata) in stage No. 3 is scattered and sometimes is frequent along
the banks with a characteristic alluvial stage of ash, elm, birch, and
silver maple; in later associations it is often found with Carpimis
caroliniana along steep river slopes (example No. 31). This same
condition occurs on the St. Francis River near Chaonia on the Wap-
papella National Forest Purchase Unit in southeastern Missouri.

(c) At the base of steep slopes in the Table Rock National For-
est Purchase Unit of southwestern Missouri, Hamamelis vernalis is
common, but is not the dominant on gravel bars as it usually is else-
where, for example, in the eastern and southern Ozarks. Instead,
Salix Wardii and Platanus dominate the gravel bars. This is
of interest, because Hamamelis is absent also to the east, on the
larger streams, but is present on the smaller ones.

(d) On the Niangua River, on the Little Pomme de Terre River,
and on the Osage Fork of Gasconade River near Rader in Webster
County, there is no birch, alder, or poplar in the alluvial formation.
Instead, one finds that Ward's willow or Ward's willow and syca-
more are followed by black willow and green ash, and then elm.

(e) On the Gasconade River, in stage 2, instead of sycamore
and black willow as co-dominants, there are sycamore, black willow,
sand-bar willow (Salix interior), birch, and Ampelopsis cor data.
Birch may be entirely absent from the associes as, for example, along
the Osage Fork of the Gasconade River in Webster County.

(/) In the southeastern Ozarks sycamore appears at the very
beginning of the succession with Salix Wardii and witch hazel
(examples Nos. 1, 2, 9, and 10), but on the Gasconade River syca-
more follows Salix Wardii or S. interior and is usually mixed with
black willow.

(gf) Along the northern border of the Ozark region the black
willow pioneers on the gravel at the very beginning of the succession
instead of Salix longipes var. Wardii. The latter is here near its

STUDIES OF THE VEGETATION OF MISSOURI— I 367

northern limit of dispersal in the Ozarks and its place is taken by
Salix nigra (fig. 12).

(li) Silver maple, elm, and green ash are usually associated at
the end of the flood-plain succession in the Gasconade and Piney
river areas.

Rosa setigera var. tomentosa is frequent along the river banks in
all the areas. Vitis cordifolia and V. cinerea, especially the latter,
are the common vines in this alluvial formation, but Ampelopsis
cordata and Parthenocissus quinquefolia also occur frequently. Aris-
tolochia tomentosa is common all along the Gasconade, Piney, Black,
Current, and White rivers in the alluvial formation. Variations on
the Gasconade River are found in the pioneering frequencies of either
Salix Wardii or S. interior, the former pioneering on direct gravel,
the latter where more sandy mud has accumulated instead of gravel.
Both species may grow on the same river bar.

Thus far, we have discussed the succession only among the trees
and shrubs of the various associes of the flood-plain formation
(Acer saccharum-Carya cordiformis association). The herbaceous
succession in this formation is:

(1) Water willow (Dianthera americana)', succeeded by

(2) Stir pus americanus or Sagittaria latifolia in back of water willow; succeeded

by

(3) Leersia oryzoides and Echinochloa pungens.

Associated with the above dominant species in stage No. 3 are:

Bidens polylepis Polygonum pensylvanicum

Panicum capillare Polygonum punctatum

Panicum dichotomiflorum Xanthium chinense

Panicum Gattingeri Xanthium pensylvanicum

On more sandy soil, Scirpus americanus, Lobelia cardinalis, and
Eupatorium coelestinum occur directly behind water willow, while
Sagittaria latifolia grows on more muddy soil behind water willow.
Rotala ramosior var. interior is found usually on muddy alluvium.
Sometimes along the border counties of the Ozarks, as along the
Osage River in St. Clair County, occur on the muddy bar Acnida
tamariscina, Ampelamus albidus, Xanthium pensylvanicum and X,
chinense, Echinochloa pungens, Polygonum coccineum, P. lapathi-
folium, and others, Lippia lanceolata, Lindernia dubia, Ammannia
coccinea, and Leptochloa filiformis. Just within the Xanthium for-
mation or behind it is Hibiscus lasiocarpus. In shaded alluvium,
back of Xanthium-Leersia association occur:

Acalypha rhomboidea Perilla frutescens

Ambrosia trifida Pilea pumila var. Deamii

Eupatorium coelestinum Polygonum punctatum

Impatiens biflora Verbena urlicaefolia

368 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Along the Osage River in Miller County water willow is followed
by an association of the following herbaceous plants: Setaria lutes-
cens, Panicum virgatum, P. capillare, Polygonum pensylvanicum,
Euphorbia supina (E. maculata of authors), andE". serpens. Back of
the gravel accumulates an alluvial deposit of muddy soil dominated
by the following plants: Xanthium chinense, Polygonum pensylvani-
cum, Spermacoce glabra, Euphorbia maculata (E. nutans of authors),
Ipomoea lacunosa, and Panicum dichotomiflorum.

The herbaceous element changes as one goes from an alluvial
type of woods to one with a development of more humus on the
alluvial bottoms and slopes. Thus, in stage No. 1 on the moist
alluvial banks along the Gasconade River the dominant species are:

Stage No. 1

Agrostis perennans Leersia virginica

Cinna arundinacea Muhlenbergia sp.

Elymus virginicus

Associated with the species in stage No. 1 are:

Campsis radicans Spermacoce glabra

Panicum clandestinum Symphoricarpos orbiculatus

Phlox divaricata var. Laphamii Ulmus americana

Pika pumila var. Deamii Uniola latifolia (on Black River)

Ruellia strepens Viola papilionacea

Solidago serotina Xanthium pensylvanicum

Elymus virginicus is a dominant species along the banks of
streams in the White River country.

Stage No. 2, on higher but still flat, alluvial ground, consists of
the following species, developed in a woodland consisting principally
of silver maple and green ash:

Stage No. 2

Acalypha rhomboidea Perilla frutescens
Acer Negundo (rare)

(young plants) Polygonum punctatum
Acer saccharinum (occasional)

(young plants) Polygonum virginianum

Aster lateriflorus var. pendulus Quercus macrocarpa
Benzoin aestivak (occasional; young plants)

(occasional; young plants) Rudbeckia laciniata

Bidens frondosa Ruellia strepens

Boehmeria cylindrica Sambucus canadensis
Cinna arundinacea (occasional; young plants)

Commelina virginica Scukllaria lateriflora

Dicliptera brachiata Smilax hispida

(occasional) Solanum nigrum

Echinocystis lobata Solidago serotina

Fraxinus pensylvanica var. lanceolata Toxicodendron radicans

(young plants) (young plants)

Impatiens biflora Vernonia altissima

Iresine rhizomatosa Viola papilionacea

Mimulus alatus Vitis cordifolia

STUDIES OF THE VEGETATION OF MISSOURI — I

369

The species listed above are in the type of woods described for
the silver maple-cottonwood associes of southeastern Missouri.

Beginning with stage No. 3, where the base of the slope begins
with a richer, better-drained soil, the herbaceous flora changes to an
abundant and luxuriant growth of the following species:

Stage No. 3

Amphicarpa bracteata
Asarum canadense
Asimina triloba

(young plants)
Bkphilia hirsuta
Carya cordiformis

(young plants)

Cystopteris fragilis var. protrusa
Eupatorium purpureum
Eupatorium urticaefolium
Euonymus atropurpureus

(young plants)
Galium triflorum

Geum canadense var. camporum
Impatiens pallida
Laportea canadensis
Menispermum canadense
Parthenocissus quinquefolia
Phytolacca americana
Sanguinaria canadensis
Tilia glabra

(young plants)
Verbesina virginica
Viola eriocarpa var. leiocarpa
Viola striata

As the richer slopes are ascended into drier, rockier ground, there
is a thinning out of alluvial species such as Leersia virginica, Rud-
beckia laciniata, Eupatorium coelestinum, Aster lateriflorus var.
pendulus, and Pilea pumila var. Deamii. Their places are taken by
the following species:

Carex blanda
Carpinus caroliniana

(young plants)
Carya cordiformis

(young plants)
Celtis occidentalis var. canina

(young plants)
Circaea latifolia

Cystopteris fragilis var. protrusa
Dioscprea villosa
Fraxinus americana

(young plants)
Phlox divaricata var. Laphamii

Quercus alba f. latiloba

(young plants)
Quercus borealis var. maxima

(young plants)
Sassafras albidum var. molle

(young plants)
Scutellaria ovala
Symphoricarpos orbiculatus

(young plants)
Ulmus fulva

(young plants)
Verbena urticaefolia

On some slopes covered with talus at this stage the herbaceous
cover is dominated by Impatiens pallida, Hydrophyllum appendicu-
latum, Viola eriocarpa var. leiocarpa, Podophyllum peltatum, Circaea
latifolia, Menispermum canadense, and Asarum canadense. Farther
up the talus, where rocks are nearer the surface and more plentiful,
Polymnia canadensis is dominant and may cover the ground solidly
for more than half the way up. Staphylea trifolia is the dominant
shrub on the talus one-fourth to one-half the way up, while on the
lower, richer slopes, as the final climax stage, Carya cordiformis is
dominant. Tradescantia Ernestiana is one of the herbaceous domi-
nants of the rich slopes in the region of White River and its

370 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

tributaries in southwestern Missouri, but is not found elsewhere
in the state.

As the soil becomes richer and richer and gradually builds up a
deeper humus layer mixed with rich, sandy alluvium, a greater
number of trees, but particularly shade-tolerant shrubs, appear.

(4) This final developmental stage is dominated by sugar maple
and bitternut hickory (Gary a cordiformis), and the associes may be
called the sugar maple-bitternut hickory associes. One or two
understories of shrubs are usually found in this associes. The paw-
paw (Asimina triloba) and American hornbeam or ironwood (Car-
pinus caroliniana) are dominant small trees forming an upper portion
of the understory, while buck brush or coralberry (Symphoricarpos
orbiculatus) is a dominant lower understory shrub. Flowering dog-
wood (Cornus florida) is often very abundant, as is also spicebush
(Benzoin aestivale) and greenbrier (Smilax hispida). Along the
tributaries of Black River, especially in the southeastern half of the
Clark National Forest Purchase Unit, leatherwood (Dirca palustris)
becomes a conspicuous shrub of the lower understory (examples Nos.
43 and 46). It is conspicuous also along Noblett and Indian creeks
in Howell and Douglas counties. These creeks drain into the North
Fork of White River in southern Missouri. Other trees and shrubs
found commonly in the final stage of this flood-plain association are,
in order of abundance:

Black walnut (Juglans nigra) Hackberry (Celtis occidentalis)

Chinquapin oak (Quercus Muhlen- Bur oak (Quercus macrocarpa)

bergii) Winged elm (Ulmus data)

White oak (Quercus alba) Red-flowered buckeye (Aesculus dis-
Sour gum (Nyssa sylvatica) color var. mollis)

Hazelnut (Corylus americana) Bladdernut (Staphylea trifolia)

Butternut (Juglans cinerea) Hop hornbeam (Ostrya virginiana)

Red mulberry (Morus rubra) Elderberry (Sambucus canadensis)

Shingle oak (Quercus imbricaria) Black haw (Viburnum rufidulum)

Redbud (Cercis canadensis) Chittim-wood (Bumelia lanuginosa)

Several species of hawthorn (Crataegus) Moonseed (Menispermum canadense)

Linden (Tilia glabra) Fragrant sumac (Rhus aromatica)

Northern red oak (Quercus borealis var. Black cherry (Prunus serotina)

maxima) Ohio buckeye (Aesculus glabra var.
Honey locust (Gleditsia triacanthos) pallida)

Red-flowered buckeye, found in rich, alluvial woods, is common
only in the southeastern section of the state.

Some of the species mentioned above, namely, linden, white oak,
chinquapin oak, northern red oak, black walnut, butternut, and
sour gum, seem to have invaded the final stages of this associes by
an incursion from the adjacent rich lower regions at the base of
limestone bluffs or slopes where they were originally part of the

STUDIES OF THE VEGETATION OF MISSOURI— I

371

climax flora of these areas and having descended the ravines and
slopes of the limestone areas gradually merged with the climax
vegetation of the alluvial association (fig. 16). They may be con-
sidered a post-climax of the flood-plain association.

Sycamore is a very common species throughout the flood-plain
development and persists into the very last associes. Together with
elm and ash, sycamore and sugar maple are the dominant species in
the final associes of the flood-plain flora. Birch likewise persists,
but in a rich flood-plain is commonest adjacent to the river. Black

FIG. 16.
County.

Along Huzzah Creek, two miles southeast of Davisville, Crawford

willow and buttonbush do not persist into this final stage, but are
represented here and there by isolated plants, usually near the
water's edge.

Some of the herbaceous species common on the forest floor of
the final flood-plain stage are:

Actinomeris alternifolia

Amphicarpa bracteata

Asarum canadense var. acuminatum

Blephilia hirsuta

Cystopteris fragilis var. protrusa

Dioscorea villosa

Eupatorium purpureum

Eupatorium urticaefolium

Galium triflorum

Geum canadense var. camporum

Impaliens pallida and I. biflora

Laportea canadensis

Lappula virginiana

Parthenocissus quinquefolia

(young plants)
Passiflora lutea
Perilla frutescens
Phlox divaricata var. Laphamii
Phytolacca americana
Pilea pumila var. Deamii
Ranunculus abortivus
Sanguinaria canadensis
Sicyos angulatus
Toxicodendron radicans
Verbesina virginica
Viola eriocarpa
Viola striata

372 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Sometimes the herbaceous flora of the rich wooded alluvium of
the final stage will be found in the stage preceding. The two inter-
grade into their floral components, with a greater variety finally
appearing in the final stage. The herbaceous ground flora associes
of this final stage is very poorly represented, the spring flora being
best developed.

II. SUGAR MAPLE-WHITE OAK ASSOCIATION

(Acer saccharum-Quercus alba)

The next association of the deciduous forest climax formation to
be discussed is the sugar maple-white oak association. In its final
developmental stages the flora of this association approaches closely

FIG. 17. Limestone glade along Niangua River, two miles south of Corkery,
Dallas County.

the composition of the flood-plain association of sugar maple-bitter-
nut hickory. The two arrive at nearly the same climax, namely,
the mesophytic, deciduous forest, because the climatic climax is a
mesophytic one, but the flood-plain association (sugar maple-bitter-
nut hickory) begins as a hydrosere (its initial stage beginning in or
near water), while the other (sugar maple- white oak) association
begins as a xerosere (its initial stages beginning where there exists
an extreme deficiency of water, as on bare limestone glades or exposed,
rocky bluffs).

(1) The first stage takes place on bare, rocky, limestone glades
or openings where the rock is exposed and the residual fragments are
everywhere conspicuous. An herbaceous flora with largely a com-
ponent of prairie species covers the glade (fig. 17). This first stage is

STUDIES OF THE VEGETATION OF MISSOURI— I

373

usually represented by the side-oats grama-perennial black-eyed
Susan (Bouteloua curtipendula-Rudbeckia missouriensis) associes.
In other portions of the Ozarks, as in some of the Ozark border coun-
ties, i.e. Benton County along the Osage River, Silphium laciniatum
and Psoralea tenuiflora are dominant instead of Rudbeckia missouri-
ensis, and occur with the co-dominant Bouteloua curtipendula (fig. 18).
In other Ozark glades, as in the border counties of the Ozarks or
in places where a rich prairie or grass flora covers open slopes, one
finds Andropogon provincialis very common and co-dominant with
Bouteloua curtipendula. Many other herbaceous plants are commonly
found. The changing aspects of the glade are the result of the spring,
summer, and autumn societies of plants. In the spring there are

FIG. 18. Limestone glade with dominants of Silphium laciniatum, Psoralea
lenuiflora, and Bouteloua curtipendula on bluffs along Osage River, two miles
northwest of Wisdom, Benton County.

societies of Missouri primrose (Oenothera missourensis), blue wild
indigo (Baptisia minor), sandwort (Arenaria patula), wild hya-
cinth (Camassia scillioides), Leavenworthia uniflora, purple cone-
flower (Echinacea pallida), yellow cone-flower (Echinacea paradoxa),
Galium virgatum, and many others. In summer the conspicuous
dominants are Houstonia nigricans, false aloe (Agave virginica),
E volvulus Nuttallianus (mostly in western Missouri), Petalostemum
purpureum, Centaurium' texense (in western Missouri), Heliotropium
tenellum, Tragia urticaefolia, Croton capitatus, and calamint (Satureja
glabella var. angustifolia). In the autumn the prevailing species are
Aster oblongifolius, A. sericeus, wild onion (Allium stellatum), and
perennial black-eyed Susan (Rudbeckia missouriensis). Panicum

374 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

mrgatum, melic grass (Melica niten-s), and big and little blue-stem
(Andropogon provincialis and A. scoparius), together with side-
oats grama (Bouteloua curtipendula), predominate among perennial
grasses, while Sporobolus vaginiflorus, S. neglectus, and Panicum
capillare predominate among annual grasses. Sporobolus vaginiflorus
is a much more common and dominant grass over the balds of
southwestern Missouri than it is on the eastern Ozark glades.

Herbaceous species found on the glades of the western Ozarks
which are common in addition to the ones given below are Pala-
foxia callosa, Linum sulcatum, Polytaenia Nuttallii, and Parthenium
hispidum. Berchemia scandens is often found. The Tragia, Pala-

FIG. 19. Red cedar glade or bald on dolomitic limestone, near Taney County
line, in Stone County.

foxia, Berchemia, and Baptisia are much more characteristic of and
some are altogether confined to the western and southwestern
Missouri glades and bald knobs.

This first stage was the original condition which existed all over
the Ozarks in limestone areas following uplifts. Then the gradually
elevated, rocky terrain first became carpeted with an herbaceous
flora such as is now found on eroded glades. As will be noted from
what follows, the limestone glades have been and are gradually losing
ground, i.e. are becoming covered by the forest (figs. 19, 20, 21,
24, and 25). In other words, the natural succession, under the present
mesophytic climatic climax, is towards a forest which sooner or later
obliterates the glade.

This pioneer herbaceous stage may last indefinitely. Those
glades and eroded limestone slopes facing west or south remain

STUDIES OF THE VEGETATION OF MISSOURI — I 375

exposed longer than those facing east or north, because the dry sun
and windswept conditions existing on such slopes are much more
favorable for the development of an herbaceous flora than for a
forest coverage (figs. 19, 20, and 21). The glade covering is too
sparse and open to permit good windbreaks to hold leaves for decay,
and those leaves which do remain are quickly scorched and dried in
the sun so that leaf decay, rock disintegration, and soil formation
proceed at a very slow rate. On east- and north-facing areas, more
moisture and protection against prevailing winds exist, and soil
building goes on at a relatively faster pace. Thus the succession
taking place on north- and east-facing slopes culminates in the climax
sugar maple-white oak associes at a relatively more rapid rate when
compared with that taking place on a south- and west-facing area.
(2) The next stage to follow the pioneer herbaceous flora is
usually initiated by the glade variety of fragrant sumac (Rhus tri-
lobata var. serotina) and is soon joined by its associates, red cedar

FIG. 20. Bald knob west of Table Rock, west of Branson, Taney County.

(Juniperus virginiana) and persimmon. Any one of these may be
dominant, but red cedar in most cases becomes the most conspicuous
member and often forms a consociation by itself (fig. 19). "Cedar
glades" are a common, characteristic feature of eroded limestone
slopes and knobs (figs. 19 and 20). The ground cover is sparse under
the red cedars and in a red cedar forest. Melic grass (Melica nitens),
Panicum virgatum, and big and little blue-stem are grasses usually
found in open areas in this associes. Some mosses, such as Orthotri-
chum Porteri and species oiGrimmia, occur on the bare, exposed rocks.

376 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Red cedar is dominant in the eastern Ozarks and in the country
bordering the Piney and Gasconade River region, but on the bald
knobs and bare limestone areas in the White River region of south-
western Missouri persimmon is often the pioneer and dominant
species. On these balds, persimmon (fig. 21) or cedar is first, or the
two occur simultaneously, and around shade in thickets at the same
time occur Rhus trilobata var. serotina, Symphoricarpos orbiculatus,
and Smilax Bona-nox. Sassafras sometimes grows as a pioneer with
persimmon, especially if there is more chert in the rock. Then follows
Bumelia lanuginosa (fig. 22) succeeded by the Cotinus americanus-

FIG. 21. Diospyros virginiana initiating woody growth on limestone glade
along Niangua River, two miles south of Corkery, Dallas County.

Quercus Muhlenbergii stage, which is associated with QuercusShumardii
var. Schneckii, Rhus trilobata var. serotina, Cercis canadensis, and
Chionanthus virginica (example No. 49). On some of the dolomitic
limestone glades and knobs here, persimmon and Rhus trilobata var.
serotina appear to be the pioneers, with cedar absent or suppressed.
If this happens, Cotinus and Quercus Muhlenbergii follow the cedar-
Rhus trilobata var. serotina stage, without the occurrence of cedar.
With persimmon are also Ulmus alata and Bumelia lanuginosa, and
sometimes Viburnum rufidulum, but Rhamnus caroliniana and Chio-
nanthus virginica are later in the succession thanQuercus Muhlenbergii
or Cotinus, or infrequently come at the same time. Aster oblongi-
folius is common on the glades, while Aster sericeus is rare. In some
places persimmon and cedar alone occur, while in others persimmon,
cedar, and winged elm dominate. Sassafras is often dominant with
persimmon on some of the bald knobs. Along draws in the limestone

STUDIES OF THE VEGETATION OF MISSOURI — I

377

glade the most common pioneering plants are Physocarpus, An-
drachne, Ilex decidua, Rhamnus caroliniana, Cornus obliqua, and C.
asperifolia. These draws may occur in slight depressions or valleys
between the balds or may occur on a slope. Chionanthus appears
around draws and drainage crevices and occurs as a substory with
Cotinus, Cornus florida, Acer saccharum, Ilex decidua, Cornus asperi-
folia, Quercus Shumardii var. Schneckii, Rhamnus caroliniana, and
Physocarpus.

The normal glade, thus far discussed, in most of the Ozarks,
including the last two stages discussed, is still open, but gradually
becomes forested (figs. 23, 24, and 25) ; with the incursion and devel-
opment of red cedar and persimmon, the rocks become covered by
leaves. This next stage of development results when the trees gain
a foothold and thereby serve as windbreaks with consequent leaf
preservation on the ground. With the gradual rock disintegration

FIG. 22. Quercus Muhknbergii invading an associes with Diospyros virginiana
and Bumelia lanuginosa along the bluffs of Osage River (Lake-of-the-Ozarks), at
Riverview, Morgan County.

and increase of soil, an incursion of various shrubs appears, and the
third stage of the succession follows.

(3) This -is the chittim-wood-black haw (Bumelia lanuginosa-
Viburnum rufidulum) associes. While these two species are domi-
nant, others also may be abundant and these are usually greenbrier
(Smilax Bona-nox), supplejack (Berchemia scandens), and species of
hawthorn (Crataegus) or hackberry (Celtis pumila var. georgiana
or C. laevigata var. texana).

378 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Once this stage has been reached, other shrubs and trees rapidly
begin to take possession, and red cedar, once so conspicuous on the
glade, gradually loses ground.

(4) An associes that usually follows is that of the winged elm-
Indian cherry ( Ulmus alata-Rhamnus caroliniana) . Associated
with these species usually is the southern hackberry (Celtis laevigata
var. texana), stunted specimens of white ash (Fraxinus americana),
and possum haw (Ilex decidua).

(5) This is quickly followed by the next associes, chinquapin
oak- white ash (Quercus Muhlenbergii-Fraxinus americana), which

FIG. 23. Quercus Muhlenbergii-Fraxinus americana associes dominating
grassy slopes along headwaters of Bryant Creek, two miles southeast of Cedar
Gap, Wright County. Shows incursion of forest over prairie-glade slopes with
gradual elimination of the latter.

dominates west- and south-facing slopes. In the eastern and south-
eastern Ozarks the chinquapin oak-white ash stage appears as one
associes, but on the White River bald knobs this stage is equiva-
lent to the chinquapin oak-Cotinus associes discussed previously.
The chinquapin oak-white ash associes has a rich variety of other
trees and shrubs, among which may be found hop hornbeam (Ostrya
virginiana), redbud, slippery elm ( Ulmus fulva), Quercus Shumardii
var. Schneckii (a variety of red oak sometimes confused with Texas
red oak), and blue ash (Fraxinus quadrangulata). Greenbrier, as
well as species from the preceding associes, such as black haw (Vi-
burnum rufidulum), winged elm ( Ulmus alata), and hop hornbeam
usually occur in this associes (fig. 28). Sometimes blue ash is much
more conspicuous in portions of the Ozarks along limestone bluffs

STUDIES OF THE VEGETATION OF MISSOURI — I 379

and becomes a dominant species, forming an associes with Bumelia
lanuginosa, immediately following an occupation by Viburnum rufi-
dulum, Rhamnus caroliniana, and Rhus trilobata var. serotina, or else
occurring simultaneously with them. On west- and south-facing
slopes and bluffs this associes may last indefinitely and form more or
less of a climax or subclimax stage, the arrested development charac-
terizing such a subclimax being due to exposed conditions which favor
the persistence of such a flora. Chinquapin oak is particularly
abundant in such exposures. On the more rocky slopes in this
associes, a temporary substory of winged elm, hop hornbeam, black
haw, and Bumelia lanuginosa may appear dominant but is finally super-
seded in the following climax forest associes, where these species
become inconspicuous. On rocky slopes in this and the following stage
with sugar maple, Brickellia grandiflora may frequently be dominant.
At this stage the herbaceous flora is dominated on the rocky lime-

FIG. 24. Bald knob, just east of McClurg, Ozark County. Showing the
Acer saccharum-Quercus alba climax reached in the development of a limestone
substratum, and the covering of the glade by forests of this climax stage.

stone slope by Pilea pumila var. Deamii, Impatiens pallida and /.
.biflora, Hydrophyllum appendiculatum, Campanula americana, and
Polymnia canadensis. This may, however, be a derived condition
Drought on by burning. At this stage a sedge (Carex eburnea) usually
occurs in the bluff crevices where a slight accumulation of soil exists,
and grows with mosses such as Rhodobryum roseum and Thuidium
minutulum. A great variety of grasses is present at this stage among
the ledges and rock exposures where soil has accumulated. They
include species of Muhlenbergia, Brachyelytrum erectum, Uniola

380 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

latifolia, wild rye (Elymus virginicus), chess (Bromus purgans), fescue
(Festuca obtusa), bottlebrush (Hystrix patula), and Sphenopholis
intermedia.

On the edges of bluffs (fig. 22) in associes Nos. 2, 3, 4, or 5 of
Quercus Muhlenbergii-Fraxinus americana or Bumelia-Diospyros
is an herbaceous flora characterized by:

Allium mutabile

Asckpias stenophylla

Euphorbia (zygophylloides) missurica

Hekotropium ienellum

Helianthus Maximiliani

Houstonia nigricans

Isanthus brachiatus

Liatris cylindracea

Malvastrum angustum
Mentzelia oligosperma
Mirabilis albidus
Psoralea tenuiflora
Scutellaria parvula
Solidago Drummondii
Solidago radula

FIG. 25. Three miles northwest of Vichy, Maries County. Similar to Fig. 24,
except that forest of Acer saccharum-Quercus alba associes has more nearly closed
the originally open glade. Cf. Figs. 24 and 25 with Figs. 19, 20, and 21.

These open bluff slopes may vary in their herbaceous composition
according to the amount of chert or limestone present, the amount of
lime which has been eroded, and other factors. If there is much
more chert than lime present, acidity prevails and Polygonum tenue,
Acalypha gracilens var. monococca, Linum medium var. texanum,
Psoralea psoralioides var. eglandulosa, Stylosanthes biflora var. his-^
pidissima, Lechea tenuifolia, Crotonopsis elliptica, Scleria pauciflora,
and Crotalaria sagittalis predominate; while, if limestone predomi-*
nates, limestone-loving species occur, such as Malvastrum angustum,
Isanthus brachiatus, Heliotropium tenellum, Houstonia nigricans,
Linum sukatum, Helianthus Maximiliani, Allium mutabik, Eu-
phorbia (zygophylloides) missurica (in the western Ozarks), Verbena
simplex, Asclepias stenophylla, and Mentzelia oligosperma.

STUDIES OF THE VEGETATION OF MISSOURI — I 381

Species commonly occurring along the edges of the bluffs are
Solidago radula, S. Drummondii, Liatris squarrosa, L. cylindracea,
and sometimes on very high, exposed situations of west- or south-
facing bluffs are Arenaria stricta var. texana and Muhlenbergia cuspi-
data (fig. 16).

Some of this flora is, of course, persistent from the original type
which occurred on the limestone glade in the beginning while yet
a part of the original cover. This is true of the types not on the very
edge of the bluff. The very edge of the bluff has distinctive groups
of species, such as Solidago Drummondii, S. radula, and Liatris cylin-
dracea, which are usually not on the open or more level parts of the
glade proper.

(6) At the beginning of this last associes, sugar maple makes its
appearance. Not until sufficient soil has accumulated on the lime-
stone exposures does it usually penetrate the area, although very

FIG. 26. Acer saccharum-Quercus alba climax associes, well developed over a
limestone substratum, Taney County.

little soil is needed to induce it to appear frequently. It is much
more abundant on north- and east-facing slopes at the same stage
when chinquapin oak is dominant on south- and west-facing ones.
Often on these south- and west-facing slopes it may be almost as
common as chinquapin oak. As more soil is formed over the lime-
stone substratum which originally was an open glade (figs. 24 and
25), sugar maple becomes commoner, and finally attains a dominance,
forming, with white oak, the last stage of development, the sugar
maple-white oak associes. On north- and east-facing slopes sugar

382 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

maple becomes the dominant species. The common species of trees
in this climax associes are white oak (Quercus alba), northern red oak
(Quercus borealis var. maxima), linden (Tilia glabra), and butternut
(Juglans cinerea). These north-facing limestone slopes produce the
climax mesophytic forest with white oak, northern red oak, and
sugar maple dominating. Usually on such north-facing slopes the
lower portions are covered by linden, northern red oak, and white
oak, while on the upper slopes northern red oak and sugar maple
are usually dominant (fig. 26).

The dominant understory in this climax associes of sugar maple-
white oak consists of flowering dogwood and American hornbeam or

FIG. 27. Bluffs along White River just west of Shreiner's Ferry, Barry
County, showing at left all stages of succession from a gravel bar in the initial
development of a flood-plain association to the final stage (Acer saccharum-Quercus
alba) of the development over an eroded limestone bluff or open glade.

iron wood (Carpinus caroliniand), while a lower story may be com-
posed of spicebush (Benzoin aestivale) and bladdernut (Staphylea tri-
folia) . As the soil becomes richer, deeper, and consequently of greater
water-holding capacity, these shrubs in the climax associes succeed in
becoming dominant over those of the preceding associes, i.e. hop
hornbeam, winged elm, black haw (Viburnum rufidulum), and Bu-
melia lanuginosa, the understory once dominant on drier exposures.
In some portions of the southeastern Ozarks the running strawberry-
bush (Euonymus obovatus) and the eastern witch hazel (Hamamelis
virginiana) (examples Nos. 78, 79, 80, and 101) may be locally
dominant in the understory flora of north- and east-facing lime-
stone slopes.

STUDIES OF THE VEGETATION OF MISSOURI — I

383

Variations in Stage No. 6:

(a) In some portions of the Ozarks, Cornus florida and Staphylea
are dominant as a substory, while in other parts they are not. In
certain parts of the western Ozarks, as in the Swan Creek territory
of Christian County, Viburnum molle is a local co-dominant in a
sugar maple-chinquapin oak-northern red oak woods as a substory

FIG. 28. Bee Bluff, along Current River, northeast of Eminence, Shannon
County, showing the development of vegetation over a limestone bluff.

co-dominant with pawpaw, Carpinus, Benzoin, and Staphylea (exam-
ple No. 70), while black maple (Acer nigrum) often becomes a local
dominant (example No. 71) instead of sugar maple on rich, north-
facing slopes along Bull Creek, Christian County, along Swan Creek
in Christian and Taney counties (examples Nos. 69 and 70), Mon-
tauk Spring Branch, in Dent County, some portions of the Gas-
conade River in Pulaski County (example No. 89), and Bryant
Creek in Wright and Douglas counties. In this case the climax is

384 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

made up of black and sugar maple, linden (Tilia glabra), and north-
ern red oak.

(6) Similarly, Cornus alternifoUa becomes a local substory co-
dominant with Staphylea on north-facing slopes in certain parts of
the eastern Ozarks, as at Montauk State Park, Dent County (exam-
ple No. 94), Bee Fork of Black River, Reynolds County, certain
parts of the Gasconade River in Pulaski County, certain places
along the North Fork of White River in Douglas County, Bryant
Creek in Wright County, along portions of Jacks Fork of Current
River and tributaries in Shannon and Texas counties, and along
Noblett and Spring creeks in parts of northwestern Howell County.

(c) Sugar maple is often dominant with chinquapin oak on west-
and south-facing areas rather than with red oak in the White River
region of southwestern Missouri. Morus rubra, Quercus Muhlen-
bergii, and Celtis occidentalis are dominant on south- and west-facing
areas. On areas in the chinquapin oak-sugar maple association in the
White River region, Cladrastis lutea is associated with Ulmus ameri-
cana, Cercis, Celtis occidentalis, Morus rubra, pawpaw, benzoin, Sym-
phoricarpos, Viburnum rufidulum, and Juglans nigra (example No.
66). In this association Yucca arkansana occurs in association with
Smilax Bona-nox, Elymus virginicus, and Berchemia. In the White
River area Robinia Pseudo-Acacia is native and part of the substory
with pawpaw, benzoin, and redbud in the sugar maple-northern red
oak-chinquapin oak area, and occurs at the base of bluffs on rich
slopes. This Robinia with its associates may be placed also in the
final stage of the alluvial associations.

(d) White oak is not part of the dominant last associes in this
final association in the White River area. The east-facing slopes
are dominated by sugar maple, northern red oak, linden, Juglans
cinerea (all found in the eastern Ozarks), as well as Celtis occidentalis,
with the lower story or substory dominated by Staphylea, paw-
paw, and Carpinus, as in the eastern Ozarks. Dogwood, which is a
dominant in substory in the eastern Ozarks, practically does not
occur here.

(e) All these local dominants are found only in certain areas,
and their distribution is limited. They do not enter the picture in
all the plant successions within the Ozarks. For example, along
Pomme de Terre River in Hickory County, Carpinus becomes less
abundant and Staphylea and pawpaw predominate.

(/) On north- and east-facing limestone slopes where the climax
type is reached, the herbaceous flora may be dominated by liverleaf

STUDIES OF THE VEGETATION OF MISSOURI — I

385

(Hepatica acutiloba), early meadow rue (Thalictrum dioicum), and
thin grass (Agrostis perennans). Various mosses, such as Mnium
cuspidatum and other species of Mnium, Platygyrium repens and
Orthotrichum pusillum, may be common in this associes.

(</) The herbaceous flora may have also Bromus purgans, Festuca
obtusa, Hystrix patula, Osmorhiza Claytoni, Athyrium pycnocarpon,

FIG. 29. Bee Bluff, along Current River, northeast of Eminence, Shannon
County, in the southeastern Ozarks; showing the development of vegetation on
a limestone bluff.

Sanicula canadensis, Actaea pachypoda, Eupatorium urticaefolium,
and Veratum Woodii. Thalictrum dioicum is locally dominant on
rich, north-facing slopes along Swan Creek, Christian County, Bee
Fork, Reynolds County, some parts of the Gasconade River in
Pulaski County, some parts of White and James rivers in Stone,
Taney, and Barry counties, some parts of the Osage River in St.
Clair County, and elsewhere in the Ozarks.

386 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

The climax development of a limestone slope will, therefore, be
seen to simulate mostly the climax associes of a flood-plain develop-
ment, the most common element in both being sugar maple. In
both associations ironwood (Carpinus caroliniana) and flowering dog-
wood are conspicuous members of the understory. The one associa-
tion begins as a hydrosere while the other starts as a xerosere, yet
each ends in the climatic mesophytic climax. Since sugar maple
favors alkaline to neutral soils, it naturally is found in both associa-
tions, in the flood-plain the development appearing on the richest

FIG. 30. Limestone bluffs along Eleven Points River, near mouth of White
Creek, Oregon County, showing somewhat more advanced stage of cover than in
Figs. 28 and 29.

alluvial climax soils, while on the limestone slope development it
responds to the calcareous soils.

Likewise there is much interchange and overlapping of floras of
the two preceding climax associations, particularly where the flora
of the rich lower portions of the limestone slopes merges into that of
the flood-plain flora. This is conspicuous in the case of such plants
as pawpaw, bitternut hickory (Carya cordiformis) , leatherwood, but-
ternut, spicebush (Benzoin aestivale), buck brush (Symphoricarpos
orbiculatus), and linden (Tilia glabra). These species occupy the
lower slopes of the limestone flora, and occur also in the climax
associes of the flood plain.

It should be borne in mind that one limestone hillside may show
all stages of the succession. Very often most of the hillside will
have reached a stage where most of the slope is covered with fairly
deep soil and is well forested except for a few exposed rock outcrops

STUDIES OF THE VEGETATION OF MISSOURI— I 387

(fig. 27). The species of the deeper soil adjacent to the rock out-
crops are constantly pioneering on any new portions of the outcrop
or glade where some soil is available so that they can gain a foot-
hold. This means that they are gradually crowding in on these
rocky habitats marginal to the open glade or rock formation.

In the case of a limestone bluff, the succession starts similarly to
that of the limestone glade from a xerosere stage and eventually
reaches the climax associes of sugar maple-white oak. However,
the succession in the former case is much more rapid, and differs
somewhat from that of the limestone glade on account of the differ-
ence in gradient of the slope, which affects the drainage and moisture
relationships markedly. In the beginning stages of a limestone bluff
formation only a few herbs take hold in the crevices of the bluff.
Common pioneer types are hydrangea (Hydrangea arborescens),
columbine (Aquilegia canadensis), poison ivy (Toxicodendron radi-
cans), bulblet bladder fern (Cystopteris bulbifera), cliff brake fern
(Pellaea glabella and P. atropurpurea) , Virginia creeper (Partheno-
cissus quinquefolia), and alum root (Heuchera puberula and H.
nmericana var. hirsuticaulis) .

As more soil accumulates and rocks disintegrate, a few hardy
trees and shrubs, such as red cedar, Bumelia lanuginosa, Rhus trilo-
bata var. serotina, Physocarpus opulifolius var. intermedius, and others,
gain a foothold and help break up the rock and aid in soil formation
(figs. 28, 29, and 30). More rapid succession takes place as a talus
slope is built up with consequent occupation by many species, such
as leaf cup (Polymnia canadensis) and bladdernut (Staphylea), which
carry the succession rapidly forward eventually toward the climax
associes.

Thus far the associations which have been discussed are those
whose final stages at least have developed largely from neutral to
calcareous soils. We come now to a discussion of associations whose
development proceeds from an acid type of soil.

III. OAK-HICKORY ASSOCIATION

(Quercus-Carya)

The first of these associations to be described is the oak-hickory.
This is the common and prevailing association over most of the
Ozark ridges, uplands, and upper slopes of hills. It develops on
the drier and more acid of the ridge tops and upper slopes (fig. 31)
and is probably the most complex association found in the Ozark
region.

388 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Just as a sugar maple-white oak association originates from an
open limestone glade, so the oak-hickory association takes its origin
from a prairie association beginning on an acid substratum. Many
parts of the plateau now covered with a dense stand of oak-hickory
were, according to the early settlers and records of travelers and
writers, originally a prairie. This again illustrates the natural
tendency under a climatic mesophytic climax for an open barren or
prairie habitat to end in a forested area. The upland prairie flora
of much of the Ozarks overlies areas of acid soils produced by Roubi-
doux sandstone or residual chert. The prairie flora which covers an

FIG. 31. A typical Ozark landscape, showing the abundant coverage by the
oak-hickory association on the ridge tops and plateaus. South of Harley, Barry
County.

upland may also originate on a sandstone, chert, granitic, or por-
phyritic trachyte barren (fig. 32), of which there are many types
scattered throughout the Ozarks.

(1) The first woody plants to invade this prairie flora are sassafras
and winged sumac (fig. 33). Each forms local colonies and together
they comprise the first developmental stage of the association, the
sassafras-winged sumac (Sassafras albidum var. molle-Rhus Copal-
Una) associes. Fragrant sumac (Rhus aromatica) also at times forms
societies in this associes. Persimmon alone frequently initiates
instead of sassafras and winged sumac. Grasses and sedges common
at this stage are wild oats grass (Danthonia spicata), slender fescue
(Festuca octoflora), Cyperus filiculmis, and Panicum linearifolium.

(2) With the next stage the development of the forest flora
becomes conspicuous, for it is at this time that black jack oak and

STUDIES OF THE VEGETATION OF MISSOURI— I 389

post oak penetrate the area. Usually these two trees are already
present in a forest area contiguous to the prairie and merely move
into the open territory afforded them. Black jack oak usually
penetrates the area first, followed by post oak, the two constituting
the black jack-post oak (Quercus marilandica-Quercus stellata)
associes. This associes may form a temporary climax or subclimax
over certain areas, and is particularly well developed on broad, level
expanses of plateau of the Ozark Dome, as is especially well dis-
played on the level uplands in parts of Dent, Crawford, Shannon,
Texas, Phelps, and Howell counties. Incursion of these oaks into

^ -

te

-.'v!

FIG. 32. Porphyritic trachyte glade of Red Rock Mountain, near Irondale,
Washington County, showing a type of habitat which may be invaded by a prairie
flora eventually culminating in an oak-hickory climax.

prairie habitats may frequently be observed along level prairie areas
such as railroad right-of-ways and natural grassy openings bordering
woods (fig. 34). This pioneering tendency of these scrub oaks to
penetrate the prairie habitats is shown by the black jack-post oak
"flats," which, according to earlier records and reports made by
travelers and settlers in the region, were originally level, open, tree-
less expanses. In this associes the grasses of the preceding associes
continue to occur, while broom sedge (Andropogon virginicus) and
several species of Panicum also are found.

(3) The next successional development occurs with the entrance
of black oak (Quercus velutina) into the black jack-post oak associes.
Along with black oak usually appears Ozark hickory1 (Carya Buckleyi

1 The name Ozark hickory is applied here to distinguish this characteristic
small-fruited Ozark species from Carya glabra and C. cordiformis, both known as
pignuts. Since it would be very misleading to speak of a third type of hickory in
Missouri as "pignut," the name "Ozark hickory" is preferred.

390 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

var. arkansana) and the two form a black oak-Ozark hickory associes
(fig. 35). Since black jack and post oaks are light-demanding types,
they begin to lose ground with the incursion into their associes of
black oak. Black oak grows so much faster and eventually taller
than the black jack and post oak that in time the latter become
shaded, and this marks the beginning of an end for them; in other
words, the beginning of another stage of the succession.

This black jack-post oak associes is the first of a later oak-
hickory climax to invade the bald knobs and limestone glades of

FIG. 33. Rhus Copallina invading prairie habitat, one mile from Riverview
bluffs, Morgan County. This is the first woody stage in the development of the
oak-hickory association.

southwestern Missouri. Post oak and black jack oak begin this suc-
cession associated with the Ozark hickory; then black oak and Ozark
hickory enter. On the knobs in later stages of succession are white
oak and black oak, with post oak and Ozark hickory. The herbaceous
flora found in this stage consists principally of Tephrosia virginiana,
Schrankia Nuttallii, Antennaria plantaginifolia, Lespedeza procum-
bens, Aster patens, Ascyrum hypericoides var. multicaule, Desmodium
rotundifolium, Panicum dichotomum, P. sphaerocarpon, P. lanugino-
sum var. fasciculatum, P. flexile, and Acacia angustissima var. hirta.

Usually black oak is the first to enter the black jack-post oak
associes, but the oak-hickory association is a most complex one, and
several other dominants or consociations may enter the black jack-
post oak associes.

(a) There may be, for example, an incursion of white oak, which
becomes dominant and with the black oak forms a white oak-black

STUDIES OF THE VEGETATION OF MISSOURI — I 391

oak associes. Or scarlet oak in the eastern Ozarks, as far west as
Indian Creek and tributaries in Douglas County, may be the domi-
nant oak type to penetrate the black jack-post oak associes.

(6) There may be an associes of scarlet oak- black oak with the
scarlet oak predominant, or an associes of scarlet oak-Ozark hickory
in which black oak is present, or there may be various mixtures of
dominants, such as scarlet oak-Ozark hickory with black oak-Ozark
hickory, or scarlet oak-black oak and white oak mixed.

(c) Sometimes in some of the porphyritic or granitic or sandstone
territory in the southeastern Ozarks, in parts of St. Francois, Madi-
son, Iron, Reynolds, and Wayne counties, winged elm, succeeding
the black jack-post oak stage, may dominate to form an associes of
winged elm-Ozark hickory ( Ulmus alata-Carya Buckleyi var.
arkansana).

FIG. 34. Upland woods with dense, grassy cover near head of Mud Creek,
twelve miles north of Poplar Bluff, Butler County. Illustrates encroachment of
oaks into prairie habitat, and shows the dominance of the blue-stem grasses in
protected and unburned areas of pine or pine-oak forests. The area illustrated
was unburned for many years.

(d) Also, in some parts of the Ozarks, as for example in the
Potosi Ranger District of the Clark National Forest Purchase Unit
in Washington County, northern red oak may follow a black jack-
post oak associes and become dominant, together with black oak
forming a northern red oak-black oak associes.

(e) In parts of southern Missouri southern red oak (Quercus
falcata) may become dominant where it occurs, and with scarlet and
black oak form an associes following a black jack-post oak associes.

392 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

The dominant understory shrubs of these various mixed oak-
hickory associes consist of low-bush huckleberry (Vaccinium vacillans
var. crinitum} (fig. 35), and high-bush huckleberry (Vaccinium
stamineum and its var. neglectum) . Low-bush huckleberry continues to
dominate and exist throughout the Ozark region, but the high-bush

sir . - - T .,

k*- * • *> -*^fc. _j

'*- *^*t • '• ff^^r •*«•
"& -^S^-- *»•- i'/tb f '

*>•

•

FIG. 35. An oak-hickory associes on a flat upland near Dillard, Crawford
County. Vaccinium vacillans var. crinitum is the dominant understory in such
associations.

huckleberry begins to disappear as the northern and western borders
of the Missouri Ozarks are reached. Under a scarlet oak-black oak
associes or scarlet oak-Ozark hickory associes deerberry or farkle-
berry (Vaccinium arboreum var. glaucescens) is dominant. It may
be common also in other associes. While Vaccinium vacillans var.
crinitum is usually the climax understory in an oak-hickory or
pine-oak associes, it may not always be. Sometimes, as in north-
eastern Wright County near Beaver Creek, there is instead of
Vaccinium vacillans var. crinitum a great variety of herbs of an acid

STUDIES OF THE VEGETATION OF MISSOURI — I 393

soil flora, such as species of Desmodium and Lespedeza, Psoralea
psoralioides var. eglandulosa, Aureolaria grandiflora var. cinerea,
Hieracium Gronovii, species of Panicum, Galactia volubilis var.
mississippiensis, Solidago radula, and others. Other common domi-
nants forming the ground cover are summer grape (Vitis aestivalis),
New Jersey tea (Ceanothus americanus), and Lespedeza, procumbens.
Fragrant sumac (Rhus aromatica) and sassafras are commonly found
in the oak-hickory association from the black jack-post oak stage on.
Winged sumac (Rhus Copallina) also is common in the association.

Sometimes black jack oak is absent from an area, leaving black
oak, white oak, post oak, Ozark hickory, and shagbark hickory as
associates. Ozark hickory is commoner usually on ridges or plateau
tops and on south- and west-facing slopes than it is on north- or
east-facing ones.

On the so-called "post oak flats," post oak, white oak, and Ozark
hickory may be the common species, with white oak dominating.
In some associes scarlet oak may be the abundant tree while in others
it may be black oak.

These associes of the upland and ridge tops in the Ozark all
develop on acid substrata and the herbaceous ground cover is toler-
ant of acid soil. In order of abundance throughout all the associes,
in addition to those already listed, they are: dittany (Cunila origanoi-
des), oats grass (Danthonia spicata), pussy toes (Antennaria plan-
taginifolia), goat's rue (Tephrosia virginiana), Lespedeza repens,
Panicum lanuginosum var. fasciculatum, Lespedeza hirta, Panicum
dichotomum, Carex hirsutella, cinquefoil (Potentilla simplex var.
genuina), tick trefoil (Desmodium marilandicum and D. rotundi-
folium), and Carex artitecta. Other common herbaceous plants are
Aster patens, A. turbinellus, A. anomalus, A. linariifolius, Solidago
ulmifolia, and S. petiolaris var. Wardii.

Leucobryum, Catharinea, Polytrichum, and Dicranum are usually
the most abundant mosses in this oak-hickory association; while of
grasses, oats grass (Danthonia spicata), species of Panicum, and
occasionally blue-stem (Andropogon scoparius and A. provincialis)
are most conspicuous.

IV. OAK-PINE ASSOCIATION

(Quercus-Pinus echinata)

The next association, the oak-pine, will require some prefacing.
Weaver and Clements (Plant ecology] consider pine only as a fire
subclimax of the deciduous forest. This will be more fully discussed
later. Suffice it to state at this time that these studies reveal that

394 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

southern yellow pine (Pinus echinata) in the Ozark region has not
developed as a fire subclimax, but has originated as a natural associa-
tion which is usually mixed with various climax species of oak. This
association consists of two associes: (1) a pure pine type with species
of Vaccinium forming the understory, which may be called the pine-

FIG. 36. Pinus echinata developed in almost pure stand on Pine Bluff, over-
lying Roubidoux sandstone along Meramec River, four miles south of St. Clair,
Franklin County.

huckleberry (Pinus echinata-V 'actinium) associes, and (2) the mixed
oak-pine (Quercus- Pinus) associes.

Pine occurs on acid soils associated with such rocks as sandstone,
chert or flint, and granite or porphyritic trachyte, which yield acid
soils. On such soils it is usually found on the upper slopes of hills,
or on narrow ridges underlain by sandstone or chert. It is much
commoner on the narrower ridges than on the broader, flat, undis-
sected plateau portions; in other words, it is most dominant where
the upland is most dissected and the drainage pattern best developed.

STUDIES OF THE VEGETATION OF MISSOURI— I

395

(1) Pure stands of pine are rare today and occur locally along
the tops and upper slopes of the ridges and on bare sandstone or
chert glades or outcrops (figs. 36 and 37). A pure pine associes may
start wherever exposed acid rock or the like occurs. On such bare
or exposed areas pine, free from any other woody species, pioneers by

FIG. 37. Pure pine stand on ridge top north of Bunker, Reynolds County.
Note the sparse herbaceous undergrowth characteristic of the coverage under a
pure pine grove.

seeding itself. This incursion by pine on open ledges and outcrops
free from competitors results in a local pure stand. Species of blue-
berry, locally called huckleberry, low-bush huckleberry (Vaccinium
vacillans var. crinitum), Vaccinium stamineum and its var. neglectum,
and farkleberry (Vaccinium arboreum var. glaucescens) comprise the
climax understory in the pure pine associes. The herbaceous ground
cover under pure pine is very sparse, consisting mostly of Lespedeza
procumbens andL. repens, horsemint (MonardaBradburiana), catgut
(Tephrosia virginiana), dittany (Cunila origanoides), tick trefoil

396 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

(Desmodium marilandicum and D. rotundifolium), and pussy toes
(Antennaria plantaginifolia) . Species of Panicum (P. linearifolium,
P. dichotomum, P. commutatum var. Ashei, and P. lanuginosum var.
fasciculatum), slender fescue grass (Festuca octoflora var. tenella), oats
grass (Danthonia spicata), species of Car ex (C. hirsutella and C.
artitecta), broom sedge (Andropogon virginicus), and blue-stem (An-
dropogon scoparius and A. provincialis), comprise the dominant
grasses and sedges. Such pure stands of pine today occur around
Pickle Creek and River Aux- Vases in Sainte Genevieve County, and
in portions of the Piney and St. Francis River country. Extensive
lumbering has diminished much of the pine stands, and pine
forests were formerly much more numerous in many places than
they are today.

(2) Much more common than the pure pine associes is the mixed
oak-pine associes, which may have several variations. The associes
may consist of (a) black jack and post oak with pine, or (6) black oak,
mockernut hickory, and Ozark hickory with pine, or of (c) white,
black, and scarlet oak, Ozark hickory and shagbark or mockernut
hickory with pine. The last type is the commonest and climax stage,
in which pine is associated with the dominant or climax species of
the oak-hickory association. This last stage develops from the
second, which in turn proceeds from the first stage of succession with
the pine.

Variations in Stage No. 2:

(a) The first type, with black jack, post oak, and pine usually
develops in an area where a pine stand is contiguous to a black jack-
post oak woods. At the zone of contact between the two some of the
black jack and post oak seed themselves in natural openings in the
pine forest, and in time form a mixed forest with the pine. In other
words, the oak has penetrated into the pine forest rather than vice
versa. The black jack and post oak also may enter a pine stand when
the latter is developed on bare sandstone or chert outcrops. Such
exposures of acid rock or even shallow soil areas with acid rock near
the surface may give a foothold here and there for pioneer black jack
and post oak. Once these oaks take hold, they begin to increase in
other openings and again the mixed type occurs.

(6) Pine grows faster and consequently taller than black jack
and post oak and is never actually dominated or towered over by
them. In time, however, black oak, together with the hickories
mentioned in the second type, invades the black jack-post oak-pine
associes, and enters into competition with the pine, finally suppress-

STUDIES OF THE VEGETATION OF MISSOURI — I 397

ing, by over-shading, the black jack and post oak. The oaks or
hickories, while they do not succeed in overtopping the previously
existing growth of pine with its more rapid rate of growth, may elimi-
nate it by over-shading and crowding out the seedling pines.

(c) The third and climax stage of mixed oak and pine occurs after
white and scarlet oak penetrate the last associes. This climax type
is best developed on the narrow ridges and upper slopes of hills or
heads of ravines and draws (fig. 15). A similar understory of species
of V actinium dominates this associes; likewise, the herbaceous ground
flora and grass associes is the same, with the exception that it is
usually more luxuriant and of greater density than in a pure pine
associes. The commonest mosses in this associes are Leucobryum
albidum, Catharinea, Dicranum scoparium, Polytrichum ohiense, Cir-
riphyllum Boscii, and Aulacomnium heterostichum. Other shrubs
besides Vaccinium in this associes are greenbrier (Smilax glauca
var. leurophylla), smooth sumac (Rhus glabra), and winged sumac
(Rhus Copallina). There is usually an abundance of broom sedge
(Andropogon virginicus) and little blue-stem (Andropogon scoparius)
in this stage.

The existence of pure pine stands of greater extent than those
today will be discussed in a later portion of this paper.

V. WHITE OAK-RED MAPLE ASSOCIATION

(Quercus alba-Acer rubrum)

The last association to be discussed is that of white oak-red
maple (Quercus alba- Acer rubrum). This would probably be
united by Weaver and Clements with the oak-hickory association,
but it always takes its origin from the same type of topographical
expression and is so distinct that it seems a most natural one. This
association commonly develops from drainages in acid soil areas,
such as slight draws in ravines, heads of tributaries of streams, and
upper slopes of hills and ravines. It may be initiated in the following
manner: From the run-off following rains along the angle of dip of a
slope, an otherwise level, broad plateau may develop a slight draw
(or wash), which eventually serves as a path (or course) along and
down which rainwater will trickle.

(1) This draw is occupied by the first stage, the flowering dog-
wood-sour gum (Cornus florida-Nyssa sylvaticd) associes. These
two trees pioneer with sassafras.

Sometimes on gravel washes which begin high or far up the
ravines at the headwaters of a tributary, Ozark witch hazel ( Hama-

398 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

melis vernalis) may be found, and in such places Ward's willow may
also be present. The Ozark witch hazel occurs on draws which lead
eventually into a stream or tributary of a stream, but is not found
where an upland slope does not lead directly into a tributary.

On the gravel wash or cherty draw of the first stage of succession
shrubby St. John's-wort (Hypericum prolificum*) and hazel (Corylus
americana) are often found.

(2) As the drainage head becomes more deeply dissected and a
larger and deeper ravine develops, other species enter, and by this
time red maple, which had previously made its appearance in the
first associes, now becomes the dominant member, with mockernut
hickory, to form the second stage of development, the red maple-
mockernut hickory (Acer rubrum-Carya tomentosa) associes. In
this associes smooth sumac (Rhus glabra), winged elm ( Ulmus alata),
and serviceberry (Amelanchier canadensis) are found together with
the other woody species of the preceding associes. Pine may enter
at this stage where footholds on open, rocky places occur. The pine
in such places is scattered because of the discontinuous nature of
the openings and the consequent more or less crowded conditions for
entering the associes. As one approaches the northwestern limit of
the Ozark region, red maple and sour gum, here at their northwestern
distributional limits within Missouri, gradually become less con-
spicuous and drop out of the associes as important species, their
places being taken more conspicuously by serviceberry, Ozark hick-
ory, and flowering dogwood.

As the ravine is deepened and more soil accumulates, the forest
becomes well mixed with other species and a number of oaks of sour
soil types enter, such as white, scarlet, and black oaks. Northern
red oak may occur; in this associes bitternut hickory often appears.

(3) Although the white oak had made its appearance at
an earlier stage of succession, it now becomes the dominant species
with red maple and forms the final stage, the white oak-red maple
(Quercus alba-Acer rubrum) associes. Low-bush huckleberry (Vac-
cinium vacillans var. crinitum), farkleberry (V actinium arbor eum
var. glaucescens), and fragrant sumac (Rhus aromatica) comprise
the main understory shrub layer, and in the southeastern Ozarks
greenbrier (Smilax glauca var. leurophylla) should be included as a
common member of the understory. The herbaceous ground cover
consists of the usual thin, rocky, acid soil dominants of catgut,
dittany, and some others, such as bracken fern (Pteridium latius-
culum), Lespedeza hirta, and tick trefoil (Desmodium marilandicum

STUDIES OF THE VEGETATION OF MISSOURI — I

399

and D. rotundifolium), Aureolaria grandi flora var. cinerea, A. pecti-
nata, A. calycosa (in the southeastern Ozarks), Aster linariifolius,
Solidago hispida, and Houstonia longifolia. Other common species
here are Euphorbia corollata, Rosa Carolina, and R. subserrulafa,

This ravine formation develops in acid soil areas and as the
headwaters enlarge the valley and ravine, the floras of the lower
slopes and valley penetrate the richer slopes and ravine bottom of

FIG. 38. South-facing limestone bluffs along White River opposite Table
Rock, Taney County, showing how a vertical cross section of this locality would
include all stages in the development of a flood-plain association, through the
stages leading to the coverage of a limestone slope, and eventually, over a cherty
substratum, to the stages leading to the oak-hickory climax.

the new drainage with such species as pawpaw, American horn-
beam (Carpinus caroliniana), spicebush, redbud, chinquapin oak,
butternut, shagbark hickory, and mulberry, while the flora of the
ridge tops, plateaus, and upper slopes descends somewhat and overlaps
that of the ravine and drainage types. Thus, the flora may become
very diversified as the head of a ravine is followed down its course.
Red maple, sour gum, white oak, and mockernut hickory comprise
the tallest dominants, with flowering dogwood, sassafras, and service-
berry forming the conspicuous understory. One sees this association
developed in all the ravines having acid soils and on all acid soil
slopes along the dissected courses of streams.

These associations have been traced according to their origin
and place of development. While they have been discussed in detail
separately and disconnectedly, it should be emphasized that they

400 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

are all contiguous and overlapping. A good example of the close
proximity of the associations is in connection with a limestone bluff
along a river course. A vertical cross section of this bluff might
reveal several stages in the development of the flood-plain flora fol-
lowing the river (fig. 38), leading to a limestone bluff with its various
associes from a limestone glade or eroded escarpment to a climax or
semiclimax stand of sugar maple-white oak; while directly overlying
the limestone bluff formation might occur an eroded chert or sand-
stone ridge covered by red maple-white oak or pine-oak association,

FIG. 39. Right half of picture in background in white is covered by Juniperus
virginiana overlying a limestone substratum; left half of picture mixed oak and
pine overlying chert substratum. Shows how two totally unlike associations may
develop side by side if growing on certain substrata. Along highway No. 19, two
miles north of Eminence, Shannon County.

and on the broader plateau portion away from the hill slope might
occur a climax oak-hickory association.

Also, as a result of variations in substratum even on one hill-
side and in the degree of erosion of the substrata, part of an associa-
tion may be in some stage of development located amidst another
association. This is a common occurrence on some limestone slopes
where chert has been exposed along a draw or has been eroded from
the surrounding limestone formation. This natural admixture of
associations often results in a mixture of the understory. The effect
of the substratum upon the plant association which develops on it
is well shown where one part of a hillside which has a limestone sub-
stratum develops the calciphilous red cedar associes, while adjacent
to the latter on a cherty soil is developed the oxylophilous southern
yellow pine associes (figs. 39 and 40).

STUDIES OP THE VEGETATION OF MISSOURI — I 401

SUBCLIMAX STAGES IN THE OZARKS

Throughout the study of natural succession in the Ozarks, vari-
ous subclimax stages were observed in the different associations.

(a) There are two subclimaxes possible in the successional stages
over an eroded limestone substratum covered eventually by a sugar
maple-white oak association: (1) a red cedar subclimax, and (2) a
chinquapin oak subclimax. The former (Juniperus virginiana) type
may persist as long as eroded limestone bluffs or glades remain.
Red cedar is a pioneer and monopolizer of such limestone-barren
areas. Ozark uplifts followed by erosion have been the natural
means of preserving such areas, which have been constantly occupied
and reoccupied by cedar. As long as a bare rocky limestone sub-
stratum occurs, cedar will be its dominant occupant. Dynamiting
of limestone areas to open up new exposures of rock, or artificial
prevention of soil cover over the limestone barren are methods by
which the area could be enlarged, if desired. Also, cutting so that
only cedar is left will again bring on a cedar associes, since, being
a light-demanding species, it will pioneer on open places.

(6) Another subclimax, that of chinquapin oak (Quercus Muhlen-
bergii) is reached on south- and west-facing limestone slopes. As
already explained, the covering up of a red cedar glade eventually
leads to dominance of the chinquapin oak stage. This may eventu-
ally culminate in the climax sugar maple, but a subclimax of the
chinquapin oak may be reached on south- and west-facing slopes.
At least, on such slopes chinquapin oak can last indefinitely because
the greater exposure and drouth, and the windswept nature of such
slopes, tend to keep chinquapin oak dominant and at the same time
prevent sugar maple from encroaching upon such dry exposures.

(c) Another subclimax is reached in the development of the
flood-plain flora. As will be recalled from discussions above, the
flora of the flood-plain areas culminates in a sugar maple-bitternut
hickory climax associes. It is possible before this climax stage is
reached for a subclimax of elm in the elm-ash associes to be developed.
A subclimax of elm may continue as long as conditions are unfavor-
able for the entrance of sugar maple into the associes (example No.
161). This may occur in several ways. If, for example, a flood-plain
flora is developing within a region surrounded by acid soils of sand-
stone or chert, and the acid soils prevent entrance of sugar maple
upon the slopes underlain by such rock, the elm in the valley would
reach its dominance and not become eventually dominated by sugar
maple, as would be the case if it were in a region surrounded immedi-

402 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

ately by limestone substrata. Also, a subclimax of elm would be
reached in a region outside the distributional limits of sugar maple.
Besides, in a flood-plain region developed where there are no rock
exposures, such as in a prairie area, or in a flood plain in a late stage
of valley development where the flood plain is several miles wide
bordered by low hills with few or no outcrops, the subclimax of elm
may be attained.

(d) Just as a subclimax of red cedar may be reached on an eroded
limestone slope, so may a subclimax of southern yellow pine be

FIG. 40. Right half of picture in background in white is covered by Juniper us
virginiana overlying a limestone substratum; left half of picture mixed oak and
pine overlying chert substratum. Shows how two totally unlike associations may
develop side by side if growing on certain substrata. Along highway No. 19,
two miles north of Eminence, Shannon County.

attained on an eroded sandstone or chert slope, ledge, or bare rock
exposure. Pine is a pioneer on such rocky ledges and once it has
gained a foothold on these areas, it monopolizes and takes possession
of them. As long as pine has the open rock free from other woody
species and undisturbed so that it can continue to invade and reseed
on such areas, it will remain dominant and uncontaminated and so
form a pine subclimax. In other words, as long as pine has penetrated
the acid rocky area first, and the thin, acid soils continue as such,
pine will form a dominance of a subclimax stage. This natural
pioneering of pine on undisturbed, light, open territory is the basis
for an understanding of the past history and the future of pine in the
Ozark forests. In the past there were undoubtedly more stands of
pine of larger and in many cases purer extent than there are today.
This is borne out by logging records, by reports by early geologists

STUDIES OF THE VEGETATION OF MISSOURI — I 403

and scientific observers throughout the Ozarks, such as those of
Shumard and Broadhead, and by reports of older settlers and "old-
timers" within the region. Some of the records of pure pine stands
from the latter sources were, no doubt, exaggerated, since early
lumbermen and settlers frequently judged as pure a stand composed
of 80-90 per cent pine, or, in other words, were "pine-minded" where
the pine occurred in thicker and purer stands. According to some
settlers, the entire country was covered with pine. When such re-
ports are more critically traced to their sources, it is usually dis-
covered that pine was found on the upper slopes and narrower ridges
while the ravine bottoms and broader plateau expanses were occu-
pied by oaks and hickories.

The natural pine succession in itself helps to explain the previous
greater dominance and abundance of pine. As has already been
explained, as soon as black jack and other oaks penetrate a pine
stand, that purity of stand is gone forever, since the various hard-
woods continue to compete and the pine is unable to suppress their
continued development, with the result that a mixed oak-pine forest
is developed. The demand by pine for light plus no competition
explains the reason why pine in the Ozarks today will not reseed
itself to a pure pine stand in an existent oak-pine forest. But pure
pine stands may be seen today where pine is spreading into bare,
sterile, or fallow fields overlying acid soil, or onto bare, acid rocks,
or over thin, acid soils recently exposed by burns, road cuts, or dyna-
miting. Since it is well known, by records of scientific travelers
throughout the Ozarks, and by reports of settlers and surveyors,
that many portions of the upland Ozark plateau now forested were
open prairie a hundred years or more ago, and since many such up-
land areas included open sandstone or chert ledges or outcrops, it is
reasonable to believe that pine in pure and large stands pioneered
and occupied such open areas. Present areas of similar nature sup-
port a dense pine growth and it seems likely that past areas would
have had the same ecologic expression.

While pines existed in denser and purer stands many years ago,
there were nevertheless upland forests of oak and mixed oak-pine.
The records of earlier surveyors and scientific travelers not only
record the "pineries" but also call attention to upland forests of oak
and oak-hickory, alternating with pine. The oak and oak-hickory
association always has been common in the Ozark region, but as a
result of lumbering the pine forests, oaks and hickories have pene-
trated more into the areas occupied by the latter and have become
even commoner than previously. This also is borne out by earlier

404 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

settlers' observations. In other words, oak becomes a real rival to
the pine, and the latter can merely hold its own, and is unable to
seed itself to advantage or gain dominance once it becomes mixed
with oak and hickory. If most of the oaks and hickories are cut,
pine has a clearing again of open and light, more or less undisturbed
territory, and will seed itself in such situations. This condition
may readily be seen today in driving over highways in the Ozarks
where pine seedlings may be observed pioneering on the bare, open,
acid soils along the road.

The facts that today on bare or open, acid, sandstone or chert
ledges or acid soil prairies, the blue-stem grasses (Andropogon sco-
parius and A. provincialis) dominate, and that these same grasses
appear commonly in protected or unburned areas in pine and pine-
oak forests, seem to offer proof that pine enters such prairie grass
areas and that under a condition of an open forest these same grasses
continue to grow (fig. 34). Thus, the forest has occupied and is
occupying the prairie, and the blue-stem and other prairie grasses
now found in pine and pine-oak and other forest types merely remind
us of the former existence of that area as a prairie, rather than as a
forest type. In other words, the forest has gradually been encroach-
ing upon the prairie rather than vice versa.

As one goes northward and northwestward along the border of the
Ozarks, pine becomes less and less conspicuous, while oaks and
hickories become commoner. The chances of the occurrence of pure
pine stands diminish as one goes toward the boundary of the Ozark
region, while the chances of the occurrence of mixed pine-oak and
oak-hickory stands increase. Thus, around the Dillard camp area
in the Clark National Forest Purchase Unit and in other places in
that region pine is absent, being too near its northern limit or as far
as its past geological migration has permitted it to go. However,
post and black jack oak and Ozark hickory in that region are sup-
ported by an upland acid soil, and pine could easily be planted on
such a soil and could become common, provided the necessary light,
open, undisturbed conditions could be given. This would apply
elsewhere on existent acid soil areas.

PLANT INDICATORS OF ACIDITY AND ALKALINITY

On account of the unglaciated condition of the Ozarks, with
residual soils and rock outcrops everywhere, the relationship be-
tween the occurrence of a species and its underlying substratum is
very obvious. It is true that other factors, such as the exposure

STUDIES OF THE VEGETATION OF MISSOURI — I 405

of the slope, i.e. whether north or south, or the proximity to water, or
exposure to sunlight, play an important r61e in affecting distribution
of the various species, but the factor of the soil acidity in relation
to the rock from which the soil has been derived appears to be the
most important one in determining distributions of many species.
This is most apparent wherever the soil is thin and the rock sub-
stratum is near the surface or exposed as bluffs, ledges, and the like.
So frequently is the same species found on particular soils of an acid
or alkaline reaction, that many species are actually indicators of
certain types of soils.

The acid soils of the Ozarks are derived from sandstone, cherts,
and flints, or granitic rocks, while the alkaline types originate from
limestones and dolomites. The ridge tops and upper slopes of many
of the hills in the Ozarks are mostly acid in soil reaction because they
are underlain by either sandstones or cherts. This is one reason why
pine occurs largely on ridge tops and upper slopes. The limestones,
on the other hand, are exposed mainly as bluffs and in outcrops
along the streams and from the bases of hills to two-thirds or three-
fourths the way up the slope, and these give rise to alkaline soils.
Draws in the heads of ravines which usually originate in the upper
cherty portions of the upland are generally of an acid nature. Plant
species of alkaline soils are often found on neutral soils as well. The
alluvial soils and those found in the development of the flood-plain
flora are usually types about neutral in reaction and there no plant
indicators are found on them. White oak is found on all types of
soils, both acid and alkaline, dry and rocky as well as alluvial, and
does not indicate any particular acidity type.

(a) The following species in the Ozarks are plant indicators of
acid soils, and where limited to certain associes the name of that
associes is given immediately after the scientific name:

Agrostis (Agrostis Elliottiana)

Aster (Aster linariifolius)

Bastard toadflax (Comandra Richardsiana)

Bedstraw (Galium arkansanum)

Bedstraw (Galium pilosum)

Black jack oak (Quercus marilandica)

Black or sour gum ( Nyssa sylvatica)

Blue curls (Trichostema dichotomum)

Blue-stem (Andropogon scoparius var. genuinus)

Bluet (Houstonia longifolia)

Bracken (Pteridium latiusculum)

Broom sedge (Andropogon Elliottii)

Broom sedge (Andropogon virginicus var. genuinus)

Bulbostylis (Bulbostylis capillaris)

Bush clover (Lespedeza hirta)

406 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Bush clover (Lespedeza procumbens)

Bush clover (Lespedeza repens)

Bush clover (Lespedeza virginica)

Catgut, hoary pea (Tephrosia virginiana)

Cinquefoil (Potentilla canadensis var. villosissima)

Cinquefoil (Potentilla simplex)

Crotonopsis (Crotonopsis elliptica)

Dropseed (Sporobolus vaginiflorus)

Early bunch grass (Sphenopholis nilida)

Fall witch grass (Leptoloma cognatum)

False dandelion (Krigia Dandelion)

Farkleberry (Vaccinium arboreum and var. glaucescens)

Goldenrod (Solidago hispida)

Greenbrier (Smilax glauca var. leurophylla)

Harvey's buttercup (Ranunculus Harveyi)

Hedgehog club-rush (Cyperus filiculmis)

Hedgehog club-rush (Cyperus ovularis)

High-bush huckleberry (Vaccinium stamineum and var. neglectum)

Indian grass (Sorghastrum nutans)

Knotweed (Polygonum tenue)

Ladies' tobacco (Antennaria plantaginifolia)

Liverleaf (Hepatica americana)

Low-bush huckleberry (Vaccinium vacillans and var. crinitum)

Mockernut hickory (Carya tomentosa)

Ozark hickory (Carya Buckleyi var. arkansana)

Panic grass (Panicum depauperatum)

Panic grass (Panicum dichotomum)

Panic grass (Panicum lanuginosum var. fasciculatum)

Panic grass (Panicum linearifolium)

Panic grass (Panicum perlongum)

Panic grass (Panicum sphaerocarpon)

Paspalum (Paspalum ciliatifolium var. Muhlenbergii)

Pencil flower (Stylosanthes biflora)

Pine- weed (Hypericum gentianoides)

Pinweed (Lechea tenuifolia)

Pinweed (Lechea villosa)

Post oak (Quercus stellata)

Poverty grass (Aristida dichotoma)

Red maple (Acer rubrum)

Rough buttonweed (Diodia teres)

Rush (Juncus marginatus)

St. Andrew's cross (Ascyrum hypericoides var. multicaule)

Scarlet oak (Quercus coccinea)

Sedge (Carex artitecta)

Sedge (Carex hirsutella)

Sedge (Carex umbellata)

Sensitive pea (Cassia nictitans)

Serviceberry (Amelanchier canadensis)

Slender fescue grass (Festuca octoflora var. tenella)

Southern red oak (Quercus falcata)

Southern yellow pine (Pinus echinata)

Spiderwort (Tradescantia longipes)

Star violet ( Houstonia pusilla)

Sundrops (Oenothera linifolia)

Tick trefoil (Desmodium laevigatum)

Tick trefoil (Desmodium marilandicum)

STUDIES OF THE VEGETATION OF MISSOURI — I 407

Tick trefoil (Desmodium nudiflorum)

Tick trefoil (Desmodium obtusum)

Tick trefoil (Desmodium rotundifolium)

Triple-awned grass (Aristida oligantha)

Turkey pea (Clitoria mariana)

Violet (Viola triloba and var. dilatata)

Wild honeysuckle, pink azalea (Rhododendron nudiflorum var. roseum)

Wild oats grass (Danthonia spicata)

Woodrush (Luzula campestris var. bulbosa)

Yellow star grass ( Hypoxis hirsuta)

(6) The following species in the Ozarks are plant indicators of
alkaline soils, and where limited to certain associes, the name of that
associes is given immediately after the scientific name:

Asclepiodora (Asclepiodora viridis) ; Bouteloua-Rudbeckia associes

Bedstraw (Galium virgatum) ; cedar associes

Blue ash (Fraxinus quadrangulata) ; chinquapin-ash associes

Buckthorn (Rhamnus lanceolaia)

Calamint (Satureja glabella var. angustifolia) ; Bouteloua-Rudbeckia associes

Chinquapin oak (Quercus Muhlenbergii) ; chinquapin-ash associes

Chittim-wood (Bumelia lanuginosa)

Cowbane (Oxypolis rigidior)

Crosswort (Lysimachia longifolia)

Early meadow rue (Thalictrum dioicum)

Evolvulus (Evolvulus Nuttallianus) ; Bouteloua-Rudbeckia associes

False aloe (Agave virginica) ; Bouteloua-Rudbeckia associes

Wood's hellebore (Veratrum Woodii)

False pennyroyal (Isanthus brachiatus)

Feverfew (Parthenium hispidum) ; Bouteloua-Rudbeckia associes

Goldenrod (Solidago Drummondii)

Grass-of-parnassus (Parnassia grandifolia)

Heliotropium ( Heliotropium tenellum) ; cedar associes

Houstonia ( Houstonia nigricans) ; cedar associes

Indian cherpy (Rhamnus caroliniana var. mollis) ; cedar associes

Kentucky blue grass (Poa pratensis)

Leavenworthia uniflora; Bouteloua-Rudbeckia associes

Liverleaf (Hepatica acutiloba)

Melic grass (Melica nitens) ; cedar associes

Missouri primrose (Oenothera missourensis) ; cedar associes

Mountain mint (Pycnanthemum virginianum)

Prairie clover (Petalostemum purpureum); Bouteloua-Rudbeckia associes

Red cedar (Juniperus virginiana)

Rudbeckia (Rudbeckia palustris)

Running strawberry-bush (Euonymus obovatus)

Rush (Juncus diffusissimus)

Sandwort (Arenaria patula) ; Bouteloua-Rudbeckia associes

Satin grass (Muhlenbergia sobolifera)

Schneck's red oak (Quercus Shumardii var. Schneckii) ; chinquapin-ash associes

Scurfy pea (Psoralea tenuiflora); cedar associes

Sedge (Carex eburnea)

Sedge (Carex leptalea)

Side-oats grama grass (Bouteloua curtipendula)

Slippery elm ( Ulmus fulva)

Sugar maple (Acer saccharum var. glaucum)

Tansy mustard (Descurainia pinnata var. brachycarpa)

408 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Umbrella grass (Fuirena simplex)

Vervain (Verbena simplex) ; cedar associes

Whitlow Grass (Draba cuneifolia) ; Bouteloua-Rudbeckia associes

Widow's cross (Sedum pulchellum)

Wild columbine (Aquilegia canadensis); chinquapin-ash associes

Wild onion (Allium stellatum); cedar associes

Most of the acid soil indicators occur in woods of either red maple-
white oak, oak-pine, or oak-hickory associes. Those most indicative
of calcareous soils are found, especially in the first few stages, in the
covering of the limestone glade, first the limestone glade, then the
"cedar" stage, and finally on to the chinquapin oak and eventually
sugar maple stage.

ECOLOGICAL VARIATIONS INDUCED BY BURNING,
LOGGING, CLEARING, AND OTHER UNNATURAL CAUSES

The various units and their succession as above discussed repre-
sent the natural and typical manner in which the vegetation has
developed. There have, however, been various factors at work, such
as burning, overgrazing, logging, and erosion, which have often
helped to change the natural aspect of the succession. The present
picture, therefore, in places, represents a hodgepodge, because while
the association is still recognizable as a unit, the component species
of its various stages have been suppressed at one time or in one section
while accelerated at another period or in some other portion. This
has led to an unbalanced and often unnatural appearance of the
association. The various factors which cause this variation also
work unevenly. Burning alone may cause certain changes in the
normal course and rate of development, while over-grazing or logging
alone may effect still other changes. When, however, instead of one
of these acting alone there is a combination of burning, logging, and
over-grazing, the effect of an unequal rate of development of the
various species may be visualized.

Burning. — Burning has very definite results on succession.
In taking stock of burns both the forest cover and the herbaceous
ground flora must be considered; too frequently only the effect on
the former has been stressed. Spring burns do considerably less
damage than summer burns, so far as the effect on the succession is
concerned, as they only retard it or keep it at a standstill. The
herbaceous ground flora does not suffer extensively from a spring
burn; a few of the more tender species may perish here and there as a
result of diminished humus cover, but in general the ground cover
remains at a standstill from year to year (fig. 41). The effect of
spring burns on the forest and understory is even less than that

STUDIES OF THE VEGETATION OF MISSOURI — I

409

on the herbaceous ground cover. The trees and shrubs after spring
burns throw out new foliage; some become scorched here and there,
but the killing effect is negligible. The quality of the timber does not
suffer nearly as greatly from spring burns as from the scarred effects
left on the wood by summer burns. Repeated annual spring burning
for a few years merely destroys the yearly leaf litter, and thereby
keeps the soil light and more or less open, preventing the accumula-
tion of a dense leaf litter. This naturally allows those species which

FIG. 41. Effect of a spring burn, showing a very light density of forest floor
coverage. Note the open patches of soil between the herbs. Twelve miles north
of Poplar Bluff, T. 26 N., R. 7 E., sec. 20 and 21, Butler County.

favor light, open ground, relatively free from leaves, to become
commoner and more or less dominant.1

Gradually, however, annual spring burnings wear down the soil
cover, and if continued over many years — perhaps 20 or 30 — soil
may be worn down to its underlying rock substratum, which in the

1 At this point it may be stated that most of the burns observed occurred on
acid soil substrata, either upland slopes, ridges, and broad plateaus, or ravines and
draws in acid soil areas; therefore, in the oak-hickory, red maple- white oak, or
oak-pine associations. No burns were observed within the limestone soil areas of
the sugar maple-white oak association. This may be due to the fact that most of
the limestone areas are in broken, rough country in the form of bluffs or glades,
usually next to streams, a type of country seldom fired by the Ozark people, while
the commonly fired hilltops and adjoining uplands and ravines have an underlying
acid substratum of eroded chert, flint, or sandstone. Thus, there are perpetuated
upon such places species found on light, open, acid soils, such as catgut (Tephrosia
rirginiana), dittany (Cunila origanmdes), pussy toes (Antennaria plantaginifolia),
horsemint (Monarda Bradburiana), bracken fern (Pteridium latiusculum), and
various species oiLespedeza and tick trefoil (Desmodium rotundifolium, D. Dillenii,
D. •marilandicum, D. obtusum, and others), sensitive pea (Cassia niditans), cin-
quefoil (Potentilla simplex), and oats grass (Danttwnia spicata).

410 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

case of much of the Ozarks is Roubidoux sandstone or chert. Thus,
in many parts of the Ozark region many flinty hills are so exposed
that they resemble a barren rock pile. In such cases the soil is worth-
less and will support only a sparse cover, much more sparse than
before, and this sparse cover is very often dominated by the worth-

FIG. 42. Damage done by summer burn, along highway No. 49, just south
of Black, Reynolds County. The burn occurred on June 25 and the picture was
taken August 14 of the same year. Note how the area was burned down to the
chert substratum, and how the sumac (Rhus Copallina) has already sprouted at
the bases of the stems.

less oats grass (Danthonia spicata). If practiced for many years,
spring burning leads eventually to destruction of a soil cover and
causes the herbaceous ground cover to revert to an earlier stage of
succession.

The effect of a spring burn on the succession is rather slow com-
pared with that of a summer burn. Summer burns cause serious
scars on the bark which leave impressions in the wood, so that its

STUDIES OF THE VEGETATION OF MISSOURI — I

411

quality is injured. Summer burns do not kill the older trees but
only the lower branches of some of the younger ones, but very often
one-year-old to twenty-year-old saplings are killed. Saplings are
killed from the tops to the base of the stem, and if they come back
thereafter, it is from the sprouting of underground shoots (fig. 42).

FIG. 43.
vegetation.

Same burn shown in Fig. 42. Note damage done to the herbaceous

The understory succession may be set back from one to twenty years
as a result of a summer burn. Usually a summer burn throws back
the stage of succession only a few years.

The effect of a summer burn on the herbaceous ground cover is
much more abrupt and deteriorating (fig. 43). The more tender
herbaceous growth is killed outright and many species usually do not
recover enough to come up again from sprouts. Thus many species
are lost in that area for a while and may not return for many years,
and in the case of some selective types, may never return. As a rule,

412 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

however, members of the Leguminosae, Compositae, Euphorbiaceae,
and Gramineae survive summer burns sufficiently to come back from
underground shoots. Species of Lespedeza and Desmodium, catgut
(Tephrosia}, Rosa, Antennaria, Panicum, Danthonia, Car ex, winged
and smooth sumacs (Rhus Copallina and R. glabra), flowering
spurge (Euphorbia corollata), and a few others come up again after
summer burns. The succession of the herbaceous ground cover is,
however, set back several years, because summer burns wear the soil
away to the rocky substratum stage so much sooner (fig. 42), and
the more this condition is approached, the more rapidly the earlier
stages of ground vegetation are reached.

These spring and summer burns do not at the time of their oc-
currence destroy all the herbaceous ground cover of an area. Most
of it is taken, but patches here and there are left untouched, and
species from these unburned patches have the advantage of repro-
ducing their kind over the barren area without much interference,
so that certain species sooner or later may dominate the burned
areas. Summer burns may be said to wear down the soil cover more
rapidly than spring burns and throw back the succession more
quickly to an earlier stage of development.

Burning in general has a deteriorating effect by diminishing most
of the palatable forage grasses, and this is an important issue in the
entire question of burns. The grass that used to be the commonest
and best range grass in the Ozarks was blue-stem (Andropogon
scoparius mostly, with some A . provincialis) . This covered large tracts
in open oak-hickory and oak-pine woodlands and occurred in great
density. Blue-stem and broom sedge (Andropogon virginicus), species
of Panicum, Sorghastrum nutans, and species of Carex (C. artitecta, C.
hirsutella, and others), were common. Wild oats grass (Danthonia
spicata) was much less common then than today, because the other
grasses prevented its development. As fires increased and annual
burning became commoner with increased settlement, the density of
the important grasses became less and less. Finally, in many places
the stage was reached where bare, open places existed, and became
commoner as grass and other herbage grew less dense. This type of
light, open, rocky soil habitat became the territory which wild oats
grass (Danthonia spicata) most favored. It increased in such open,
rocky areas and in time became dominant, until now it is the com-
monest grass of the upland acid slopes and ridges. It is not a good
forage grass, and its establishment on light, open, rocky soil areas
caused by burning indicates the harm caused by repeated and un-

STUDIES OF THE VEGETATION OF MISSOURI — I

413

controlled burning. If the people in the Ozarks burn their forests to
get a cover of more grass for forage, they are being deceived into
believing that oats grass is good, while as a matter of fact it is
inferior.

The rate of deterioration or reversion to an earlier and earlier stage
of the ground cover is directly influenced by the slope gradient. In

aiS^-- >v*i; •ta'ir^^^t."-^

~**fe^

FIG. 44. Flat ridge top in Quercus alba-Quercus stellata woodland, four miles
northwest of Bunker, Reynolds County. Note thick leaf covering with resultant
scarcity of undergrowth.

the case of a steep slope, the soil is worn down much more rapidly to
flint or sandstone than in that of the flat upland ridge or plateau.
With the latter a very much longer period of time is required to wear
the ground down to a rock substratum.

Shading and the density of leaf litter on the ground have definite
effects on the herbaceous ground flora. For the most part, shading
and a dense ground cover produce a sparse ground flora, and a sparse

414 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

grass flora in particular. The following conditions serve to show the
various types found within the Ozarks:

(1) If the soil has been worn down as a result of continued fires
to the flinty or sandstone substratum, and at the same time the woods
are more or less open, the herbaceous cover is poor and the inferior
oats grass (Danthonia spicata) is dominant among the grasses present.

(2) If the soil as a result of scattered burns has been worn down
so that it is still intact but also rocky and at the same time the woods

FIG. 45. Quercus alba-Quercus velutina-Quercus stellata woodland on flat
ridge top, four miles northwest of Bunker, Reynolds County, near ridge top shown
in Fig. 44. Note the dense sprout undergrowth, but poor herbaceous ground flora.

are more or less open, the result is usually a rich and dense herba-
ceous vegetation with grasses and legumes dominating, and usually
the better forage grasses, such as blue-stem (Andropogon scoparius
and A. provincialis) and panic grasses, predominate, while oats grass
is inconspicuous. This is a desirable condition.

(3) If the soil on a flat upland ridge or plateau is built up and
covered by a dense forest of younger trees, the effect of shade and at
the same time a dense leaf litter, is such as to produce a very sparse
ground flora, the grasses being almost negligible (fig. 44).

(4) If a forest such as is found in the preceding type, with built-up
soil, is burned over and a dense stump sprout growth springs up, the
sprouts act as wind-breaks and allow dense coverings of leaves to
accumulate in their vicinity. Wherever these dense leaf piles appear,
the grasses are absent, especially oats grass. Some legumes, such
as Lespedeza procumbens, may be present, but the flora as a whole is

STUDIES OF THE VEGETATION OF MISSOURI — I 415

sparse. Only in light and rocky openings, between leaf piles or where
rock is exposed, or around the bases of trees, does the ground flora
become denser.

(5) In areas protected from burns and at the same time with an
open woodland, a variety of grasses return and in time approach their
original dense growth.

(6) In areas protected from burns but with a dense, crowded
woodland, the shade and leaf litter are too great for a rich ground
flora, and consequently the herbaceous flora is very poor, especially
in grasses (fig. 45). The ground cover may then be dominated by
the shade-enduring low-bush huckleberry (fig. 35).

It appears that the densest growth of grasses of the better forage
type is found in open woods protected from fire with a light, semi-
rocky soil not extensively covered with leaf litter. If an area is
repeatedly burned down, eventually to a rock substratum, oats grass
becomes the most conspicuous grass.

A light, open, rocky soil also harbors a rich variety of legumes,
many of which favor such a rocky, thin, open substratum. Thus,
after continued burning with reduction of soil, such species increase
in numbers because of an increase in their habitats. It is therefore
incorrect to speak of such species as indicators of burning; rather
they are species which appear in areas opened up by burning. The
various herbs which are characteristic or commonest in areas opened
by burns, or which tolerate burns, are the following:

Bracken fern (Pteridium latiusculwrri)

Bush clover (various species) (Lespedeza procumbens, L. repens, L. hirta, and

L. virginicd)

Catgut (Tephrosia virginiana)
Cinquefoil (Potentilla simplex var. typica)
Dittany (Cunila origanoides)
Fleabane (Erigeron ramosus)
Flowering spurge (Euphorbia corollata)
Forked chickweed (Paronychia fastigiata)
Greenbrier (Smilax glauca var. leurophylla)
Horsemint (Monarda Bradburiana)
Ironwood (Vernonia Baldwini)
Japanese clover (Lespedeza slriata)
Pencil flower (Stylosanthes biflora)
Pennyroyal (Hedeoma pulegioides)
Pinweed (Lechea tenuifolia)
Pussy toes (Antennaria plantaginifolia)
Rose (Rosa subserrulata)
Sensitive pea (Cassia nictitans)
Sweet everlasting (Gnaphalium obtttsifolium)
Tick trefoil (various species) (Desmodium rotundifolium, D. obtusum, and

D. Dillenii) .
Wild indigo (Baptisia leucantha and B. leucophaea)

416 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

It will be seen that a great majority of these species belong to
the pea and composite families. Catgut, dittany, greenbrier, Les-
pedeza procumbens, and Desmodium rotundifolium are especially
common.

The grasses which are commonest in such burned areas are the
following:

Oats grass

Species of Panicum (P. dichotomum and P. lanuginosum var. fasciculatum)

Hair grass (Agrostis scabra)

Sedge (Carex hirsutella)

In woods which have been burned over and opened up, winged
sumac (Rhus Copallina) and sassafras are the commonest woody
plants to enter and consequently are indicators of burning in that
sense.

If, as these studies indicate, a great variety and density of legu-
minous species appear on light, thin soils of the rocky, open type —
the type most directly produced by burning, especially summer
burns — it might be advantageous on turkey ranges or other similar
areas where leguminous foods are of high value, to burn, under con-
trolled management, every five to ten years in order to revert the
herbaceous ground cover to an earlier stage of succession where the
leguminous flora will be at its maximum development. If this is
not done, the natural succession of the ground flora, where fires
are eliminated and where leaves consequently would accumulate
and shade increase, would be towards a suppression of grasses and
legumes, resulting in less abundance of turkey food.

Logging. — The effects of burning are thus found to be varying
in degree and must be considered from different angles. The effects of
logging on the natural succession of a forest are equally interesting.
When tall trees are removed from a forest, the result is an open, sunny
place where previously more or less shaded conditions existed. This
creates a habitat within the forest equivalent to a prairie opening,
and allows species which favor open and sun-lit conditions to enter,
penetrate, or spread. Very often some species which come in on
repeatedly cleared areas are the same as appear in burned-over areas,
because similar habitats are created by burning and repeated clear-
ing. In the case of a minor amount of cutting in a forest, done only
rarely, the migration into that area of light-demanding species is not
great. However, where repeated clearing continues, a number of
species are commonly found which are more or less characteristic of
open areas. The herbaceous species which may be called indicators

STUDIES OF THE VEGETATION OF MISSOURI — I 417

of repeated clearing, although they are merely species coming in on
light, open soils, are the following:

Bitterweed (Helenium tenuifolium)

Black-eyed Susan (Rudbeckia hirta)

Bracken fern (Pteridium latiusculum)

Broom sedge (Andropogon virginicus)

Bush clover (Lespedeza virginica)

Buttonweed (Diodia teres)

Dewberry (Rubus flagellaris)

Everlasting (Gnaphalium purpureum)

Field sorrel (Rumex Acetosella)

Fleabane (Erigeron ramosus)

Flowering spurge (Euphorbia corollata)

Forked chickweed (Paronychia fastigiata)

Greenbrier (Smilax glauca var. leurophylla)

Ironweed (Vernonia Baldwini)

Japanese clover (Lespedeza striata)

Mullein (Verbascum Thapsus)

Pennyroyal (Hedeoma pulegioides and H. hispida)

Pinweed (Lechea tenuifolia)

Pussy toes (Antennaria plantaginifolia)

Thistle (Cirsium lanceolatum)

Wild indigo (Baptisia leucantha and B. leucophaea)

Wild petunia (Ruellia caroliniensis)

Yarrow (Achillea Millefolium)

Yellow hop clover (Trifolium procumbens)

The commonest species which appear on such areas are bitter-
weed, yarrow, greenbrier, and flowering spurge. Oats grass is by far
the commonest of the grasses. Among the commonest shrubs are
smooth sumac (Rhus glabra) and winged sumac (Rhus Copallind).
Sassafras, persimmon, post and black jack oak, and winged elm are
the only trees which are commonly found in or come into repeatedly
cleared areas. The first woody species to spread into an opening in a
forest induced by repeated clearing or logging are the smooth and
winged sumac and sassafras. The natural home for these species is
open prairie, borders of woods, or openings in woods. Thus, when
conditions are created within a forest by clearing which establish an
open habitat, these shrubs and trees are first to pioneer in response to
their light-demanding requirements.

Repeated Clearing and Logging. — The effects of repeated clearing
and logging are not as harmful as repeated burning. Logging and
clearing merely open up the soil cover and understory, but do not
materially disturb the understory or reduce the soil cover to the
bare substratum as does repeated burning. If grasses of the better
forage type are already present in sufficient numbers when the clearing
occurs, they are thereby benefited, because new open places and an
increase of favorable habitats are created. Thus, under such con-

418 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

ditions, grasses will increase in density, as will also legumes and
composites and other light-demanding herbs. If, however, the soil
has been worn down badly as a result of repeated burns, and the area
is simultaneously or subsequently cut over, the worthless oats grass
(Danthonia spicata) becomes more plentiful. In many instances it
was observed that grasses were most plentiful and in greatest density
in areas which had been opened up by logging but protected from
fires. Thus, for best grazing ranges, it might be advantageous to cut
over in part and at different times an area already covered with good
forage grasses, so as to create more openings and sun-lit areas to be
occupied by an increasing number of grass species.

Repeated clearing and logging result in such a mixture and con-
fusion in the natural forest succession that it is at times difficult to
interpret the stage at which it has arrived in a given forest. There-
fore certain species are thinned out, and instead of the succession
carrying on toward the climax stage, the earlier species of the succes-
sion may become once more dominant, and the previous dominant
species may become temporarily suppressed. Thus, a temporary
standstill or reversion may occur. In time, however, the species
originally dominant usually persist and, if left to grow again from
saplings or sprouts, finally enter the climax stages of the association.

Over-Grazing. — Many portions of the Ozark region are over-
grazed. These areas usually occur on the upland. Usually a greater
number of animals is turned loose over an area than can be supported
on that area. The average density of the herbaceous ground cover
varies according to different influences. The density of an herbaceous
cover under a dense, unburned post oak-black jack oak stand is
always very poor; that under an unburned, open, oak-hickory or
pine-oak association is usually better; that of a logged but unburned
area is usually good; that under an annually burned-over, open woods
is usually quite poor, as is also one under a very much cut-over and
burned-over woods. Since the normal density is not great, and does
not average very much in an ordinary open oak-hickory or pine-oak
forest, it is easy to allow too many animals to graze in such an area.
When, in addition, open range hogs are turned loose to grub roots
and eat herbage and nuts of various kinds indiscriminately, their
effect on making a low density even lower should be emphasized.
There is too much grazing on a limited amount of land in the Ozarks
already. When it is considered that excessive over-burning in cut-over
or non-cut-over areas lowers the ordinary density of herbaceous
growth within a forest, and when to this is added the burden of over-

STUDIES OF THE VEGETATION OF MISSOURI — I 419

grazing, it is evident that the already sore conditions may be further
aggravated. The Ozarks, if properly protected from fire and too
much grazing, may afford valuable pasturage, a high density yield
being possible in an open oak-hickory or oak-pine association. If
too frequent burning and over-grazing are permitted, the low density
at present encountered may be expected to continue. And in many
areas in the Ozarks grazing in the uplands and slopes is not now
practicable on a large scale.

When an upland area that has been cleared or cut over is then
grazed, the constant cropping of herbage by too many cattle during
summer leads to deterioration of the herbaceous cover, which in
time becomes thinned out, exposing numerous bare patches of ground.
When this condition has been reached, various species, mostly weedy
and colonizing types, take possession. These are indicators of over-
grazing, and may be the same as those on a sterile or fallow field.

The herbs which indicate over-grazing, because they are the ones
which remain in a grazed-over area after the more delicate species
have been eaten or have perished, are:

Bitterweed ( Helenium tenuifolium)

Blue vervain (Verbena stricta)

Bracted plantain (Plantago aristata)

Dwarf fleabane (Erigeron dlvaricatus)

English plantain (Plantago lanceolata)

Everlasting (Gnaphalium purpureum)

Field sorrel (Rumex Acetosella)

Hairy pennyroyal ( Hedeoma hispida)

Hogwort (Croton monanthogynus)

Horse nettle (Solanum carolinense)

Japanese clover (Lespedeza striata)

Lesser ragweed (Ambrosia bidentata)

Mullein (Verbascum Thapsus)

Pinweed (Lechea tenuifolia)

Plantain (Plantago elongata)

Rough buttonwood (Diodia teres)

Rush (Juncus macer)

Sundrops (Oenothera linifolia)

Sweet everlasting (Gnaphalium obtusifolium)

Thistle (Cirsium lanceolatum)

White boneset (Eupatorium serotinum)

Yarrow (Achillea Millefolium)

Yellow hop clover (Trifolium procumbens)

The following grasses are characteristic indicators of over-grazing :

Crab grass (Digitaria sanguinalis)

Elliott's hair grass (Agrostis Elliottiana)

Fescue grass (Festuca octoflora and var. tenella)

Hair grass (Agrostis scabra)

Poverty grass (Aristida dichotoma)

Sporobolus vaginiflorus

Triple awn grass (Aristida oligantha)

420 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Winged sumac (Rhus Copallina), sassafras, and winged elm are
woody species that indicate over-grazing. These species all enter
areas which have been over-grazed.

Interestingly enough, most of these species occur also on open,
rocky, sandstone or chert or granite glades. In other words, the
same conditions of thin, bare, worn-down, or acid soils prevail on
both barrens of acid rock and on over-grazed fields. An over-grazed
soil usually becomes worn down to the extent that bare patches of
soil exist with stunted herbage; this type of condition approaches a
glade in its sterile, open, dry, wind-swept soil, and thus, species
favoring such areas pioneer quickly on such types.

Other Unnatural Conditions.— During dry summers in the Ozarks,
cattle were observed to have an injurious effect on the vegeta-
tion of natural sink-hole ponds. Within certain parts of the eastern
Ozarks, particularly in Reynolds, Shannon, Oregon, Texas, and
Howell counties, are many such ponds. During ordinary seasons they
usually remain filled with water. Most of those that dried up during
the summer might have retained water had it not been for the cattle,
which were allowed to wade in and out at any time; every time they
emerged they took away water. Ordinarily evaporation during
a summer merely reduces the water level, but does not dry up
the pond. However, cattle shuffling in and out of the ponds caused
many of the ponds to dry up completely. As the ponds become dried,
the plants existing on the tussocks of sedges or in the lower crotches
of shrubs are killed, both by lack of water and by being eaten by the
cattle. Many of these plants at ordinary times furnish excellent
forage, but when over-grazed and eaten to the roots by cattle, they
are badly injured or killed. Since in these ponds many rare species
of plants exist which are found nowhere else in the state, some way
should be devised to prevent cattle from ruining the plants that
serve as food for turkey, quail, duck, and other upland game life
of the region.

The natural succession here is a series of stages eventually leading
to a filling up of the pond and the establishment of a mesophytic
forest thereon, and since such destructive over-grazing and drying
up of these ponds by mismanagement of cattle grazing on them
hastens their extinction, it seems highly important that steps be
taken to protect ponds. Such ponds are valuable water reservoirs
for game in the national forests, and every effort should be taken to
keep cattle from destructive "stamping" in and out of the water from
the pond.

STUDIES OF THE VEGETATION OF MISSOURI — I 421

Each of the following localities visited by the writer illustrates
some special stage of development of a plant association discussed
in the present paper. Usually the most conspicuous members of
the plant association are listed under dominant trees or shrubs, and
dominant herbaceous plants, as well as sub-dominants wherever it
has seemed pertinent. The species listed as dominant are the most
numerous in a given area.

Herbarium specimens of all the examples of plant associational
cross sections in the following pages are to be found in the Herba-
rium of the Missouri Botanical Garden and the Herbarium of Field
Museum of Natural History.

I. ACER SACCHARUM-CARYA CORDIFORMIS
ASSOCIATION

(1) Salix longipes var. Wardii-Hamamelis vernalis Associes

1. Gravel bar at Hall's Bluff, south of Davidson's Blue Spring
along Black River, sec. 4, T. 27 N., R. 4 E., four miles southeast of
Mill Spring, Wayne County.

Dominant Trees

Ward's willow (Salix longipes var. Sycamore (Platanus occidentalis)
Wardii)

2. Porphyritic stream bed in "shut-in" along Telleck Branch,
sec. 23, T. 35 N., R. 1 E., three and one-half miles northwest of
Shepherd, Iron County.

Dominant Tree

Sycamore (Platanus occidentalis)

Dominant Shrubs

Ward's willow (Salix longipes var. Ozark witch hazel (Hamamelis vernalis)
Wardii) Smooth alder (Alnus rugosa)

Associate

Sand grape (Vitis rupestris)

3. Gravel bar along Big Piney River between Dial's Camp, east
of Big Piney and Ormsby's Place at Spring Creek crossing, T. 34-
35 N., R. 10-11 W., Pulaski County.

Dominant Woody Plant

Ward's willow (Salix longipes var. Wardii)

Dominant Herbaceous Non-Grass Plants

Water willow (Dianthera americana) Chairmaker's rush (Scirpus americana)

4. Rocky stream bed along Noblett Creek, sec. 19, T. 26 N.,
R. 10 W., eight miles west of Burnham, Howell County.

422 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Shrubs

Ward's willow (Salix longipes var. Swamp dogwood (Cornus obliqua)

Wardii) Ozark witch hazel ( Hamamelis vernalis)

Smooth alder (Alnus rugosa)

5. Gravelly part of Piney Creek, sec. 19, T. 23 N., R. 24 W.,
five miles southwest of Cape Fair, Stone County.

Dominant Shrubs
Ozark witch hazel (Hamamelis vernalis) Possum haw (Ilex decidua)

Other Shrub Associates

Hop hornbeam (Ostrya virginiana) Swamp dogwood (Cornus obliqua)

Rough-leaved dogwood (Cornus asperi- Ninebark (Physocarpus opulifolius var.
folia) intermedius)

6. Gravel bar along Pomme de Terre River, three miles north-
east of Elkland, Hickory County.

Dominant Shrub

Ward's willow (Salix longipes var. Wardii)

Dominant Herbaceous Non-Grass Plant

Water willow (Dianthera americana)

7. Gravel bar along Osage River, sec. 2, T. 40 N., R. 13 W.,
west of Capps, Miller County.

Foxtail grass (Setaria lutescens) Lady's thumb (Polygonum pensylvani-

Panic grass (Panicum virgatum) cum)

Witch grass (Panicum capillare) Spurge (Euphorbia supina)

Spurge (Euphorbia serpens)

8. Dry gravel bar along headwaters of Meramec River, sec. 14,
T. 33 N., R. 4 W., two miles southeast of Max, Dent County.

Dominant Shrubs

Swamp dogwood (Cornus obliqua) Shrubby St. John's-wort (Hypericum

Ninebark (Physocarpus opulifolius var. prolificum)

intermedius) Sassafras (Sassafras albidum)
Sand grape (Vitis rupestris)

Dominant Herbaceous Non-Grass Plants

Flowering spurge (Euphorbia corollata) Bush clover (Lespedeza virginica)
Tick trefoil (Desmodium sessilifolium) Bush clover (Lespedeza capitata)
Star tickseed (Coreopsis pubescens)

Dominant Grass

Panic grass (Panicum virgatum)

9. Gravel bar along Salt River, at Asher Ford, south of Spalding,
Rails County.

Dominant Woody Plants

Ward's willow (Salix longipes var. Sycamore (Platanus occidentalis)
Wardii)

STUDIES OF THE VEGETATION OF MISSOURI — I 423

Dominant Herbaceous Non-Grass Plant

Water willow (Dianthera americana)

Other Herbaceous Non-Grass Plant Associates

Small white morning glory (Ipomoea Wild bean (Strophostyles helvola)

lacunosa) Water hemp (Acnida tamariscina)

Lady's thumb (Polygonum lapathi- Mule tail (Erigeron canadensis)

folium) Carpet weed (Mollugo verticillata)

Lady's thumb (Polygonum pensyl- Nodding spurge (Euphorbia maculata)

vanicum) Sida (Sida spinosa)

Smartweed (Polygonum punctatum) Small ragweed (Ambrosia artemisiae-

Smartweed (Polygonum Hydropiper var. folia var. elatior)

projectum)

Dominant Grasses

Panic grass (Panicum virgatum) Big blue-stem (Andropogon provincialis

Slough grass (Spartina pectinata)

Other Grass Associates

Panic grass (Panicum dichotomiflorum) Crab grass (Digilaria sanguinalis)
Love grass (Eragrostis cilianensis)

10. Gravel bed of Little Fourche a Renault Creek, sec. 2, T. 37 N.,
R. 1 E., two miles southeast of Floyd, Washington County.

Dominant Trees

Sycamore (Platanus occidentalis) Persimmon (Diospyros virginiana)

Dominant Understory

Swamp dogwood (Cornus obliqua) Buck brush (Symphoricarpos orbiculatus)

Ward's willow (Salix longipes var. Shrubby St. John's-wort (Hypericum

Wardii) prolificum)

Sand grape (Vitis rupestris) Rough-leaved dogwood (Cornus asperi-

Ninebark (Physocarpus opulifolius var. folia)

intermedius) Black haw (Viburnum rufidulum)

(2) Acer saccharinum-Populus deltoides Associes

11. Alluvial banks along Black River between Mill Spring and
Williamsville, sec. 30, T. 27 N., R. 5 E., Wayne County.

Dominant Trees

Cottonwood (Populus deltoides) Sycamore (Platanus occidentalis)

Dominant Understory

Ward's willow (Salix longipes var. Black willow (Salix nigra)
Wardii)

12. Gravel bar at base of limestone slopes along Brushy Creek,
sec. 8, T. 27 N., R. 3 E., six miles north of Ellsinore, Carter County.

Dominant Trees

River birch (Betula nigra) Sycamore (Platanus occidentalis)

Dominant Shrubs

Ozark witch hazel (Hamamelis vernalis) Button-bush (Cephalanthus occidentalis)

424 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

13. Sandy mud along Big Piney River, between Dial's Camp,
east of Big Piney and Ormsby's Place at Spring Creek crossing,
T. 34-35 N., R. 10-11 W., Pulaski County.

Dominant Woody Plants

Sandbar willow (Salix interior) Black willow (Salix nigra)

Dominant Herbaceous Non-Grass Plants
Blue mist flower (Eupatorium coeles- Cocklebur (Xanthium pensylvanicum)

tinum) Lady's thumb (Polygonum pensyl-

Cardinal flower (Lobelia cardinalis) vanicum)

Beggar-ticks (Bidens polylepis)

Other Herbaceous Non-Grass Plant Associates

Broad-leaved arrow-head (Sagittaria Smartweed (Polygonum punctatum)
latifolia) Hibiscus (Hibiscus lasiocarpos)

Dominant Grasses

Barngrass (Echinochloa pungens) Rice cut grass (Leersia oryzoides)

Other Grass Associates

Panic grass (Panicum dichotomiflorum) Panic grass (Panicum Gattingeri)
Witch grass (Panicum capillare)

14. Alluvial woods along lower part of spring branch of Onandago
Spring, Crawford County.

Dominant Trees

Silver maple (Acer saccharinum) Black willow (Salix nigra)

Other Tree Associates

Sycamore (Platanus occidentalis) Box elder (Acer Negundo)^

River birch (Betula nigra) Honey locust (Gleditsia triacanthos)

Bur oak (Quercus macrocarpa) American elm (Ulmus americana)

Dominant Understory

Elder (Sambucus canadensis)

Dominant Herbaceous Non-Grass Plants

Wild goldenglow (Rudbeckia laciniata) Aster (Aster lateriflorus var. pendulus)

15. Alluvial bottoms at Hall's Bluff, south of Davidson's Blue
Spring, along Black River, sec. 4, T. 27 N., R. 4 E., four miles south-
east of Mill Spring, Wayne County.

Dominant Trees

Silver maple (Acer saccharinum) Green ash (Fraxinus pensylvanica var.

River birch (Betula nigra) lanceolata)

Cottonwood (Populus deltoides) Black willow (Salix nigra)

Dominant Understory

Button-bush (Cephalanthus occidentalis)

16. Alluvial woods along lower part of spring branch of Schlicht
Spring, sec. 30, T. 36 N., R. 12 W., two miles southeast of Swede-
borg, Pulaski County.

STUDIES OF THE VEGETATION OF MISSOURI — I 425

Dominant Trees

Sycamore (Platanus occidentalis) Silver maple (Acer saccharinum)

Black willow (Salix nigra) American elm (Ulmus americana)

17. Alluvial banks along Big Piney River, between Dial's Camp,
east of Big Piney and Ormsby's Place at Spring Creek crossing,
T. 34-35 N., R. 10-11 W., Pulaski County.

Dominant Trees

Green ash (Fraxinus pensyhanica var. Silver maple (Acer saccharinum)
lanceolata) American elm (Ulmus americana)

Other Tree Associates

Cottonwood (Populus deltoides) Shumard's red oak (Quercus Shumardii)

18. Alluvial mud along Big Piney River, between Dial's Camp,
east of Big Piney and Ormsby's Place at Spring Creek crossing,
T. 34-35 N., R. 10-11 W., Pulaski County.

Dominant Trees

Black willow (Salix nigra) Sycamore (Platanus occidentalis')

Dominant Herbaceous Non-Grass Plants

Three-seeded mercury (Acalypha rhom- Clearweed (Pilea pumila var. Deamii)

boidea) Spotted jewel-weed (Impatiens biflora)

Blue mist flower (Eupatorium coeles- White vervain (Verbena urticaefolia)

tinum) Beefsteak plant (Perilla frutescens)

Horseweed (Ambrosia trifida)

19. Mud flat along Osage River, sec. 2, T. 40 N., R. 13 W., west
of Capps, Miller County.

Lady's thumb (Polygonum pensylvani- Cocklebur (Xanthium chinense)

cum) Button-weed (Spermacoce glabra)

Small white morning glory (Ipomoea Panic grass (Panicum dichotomiflorum)

lacunosa) Spurge (Euphorbia maculata)

(3) Ulmus americana-Fraxinus pensylvanica var. lanceolata

Associes

20. Alluvial woods along Gasconade River, south of Hazel Green,
Pulaski County.

Dominant Trees

Black willow (Salix nigra) Green ash (Fraxinus pensylvanica var.

Silver maple (Acer saccharinum) lanceolata)

Sycamore (Platanus occidentalis)

Other Tree Associates

American elm ( Ulmus americana) Shumard's red oak (Quercus Shumardii)

River birch (Betula nigra) Box elder (Acer Negundo)

426 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Dominant Understory

Virginia creeper (Parthenocissus quin- Winter grape (Vitis cinerea)
quefolia) Frost grape (Vitis cordifolia)

Other Understory Associates

Dutchman's pipe (Aristolochia tomen- Hispid greenbrier (Smilax hispida)
tosa)

Dominant Herbaceous Non-Grass Plants

Blue mist flower (Eupatorium coeles- Goldenrod (Solidago serotina)
tinum) Sweet William (Phlox paniculata)

21. Alluvial woods along Sniabar Creek, four and one-half miles
south of Napoleon, Lafayette County.

Dominant Trees

Black willow (Salix nigra) Cottonwood (Populus deltoides)

Green ash (Fraxinus pensylvanica var. Silver maple (Acer saccharinum)
lanceolate) Sycamore (Platanus occidentalis)

Other Tree Associates

Box elder (Acer Negundo) Bur oak (Quercus macrocarpa)

American elm (Ulmus americana) Hackberry (Celtis occidentalis)

22. Alluvial woods along Pomme de Terre River, three miles
northeast of Elkland, Hickory County.

Dominant Trees

Green ash (Fraxinus pensylvanica var. Sycamore (Platanus occidentalis)
lanceolata)

23. Low, alluvial woods along Noblett Creek, sec. 19, T. 26 N.,
R. 10 W., eight miles west of Burnham, Howell County.

Dominant Trees

Green ash (Fraxinus pensylvanica var. Sycamore (Platanus occidentalis)
lanceolata) American elm ( Ulmus americana)

Dominant Understory

Leatherwood (Dirca palustris)

24. Alluvial ground along King's River, south of Prentiss
Ford, sec. 24, T. 21 N., R. 25 W., five miles south of Viola, in Barry
County.

Dominant Trees

Green ash (Fraxinus pensylvanica var. American elm (Ulmus americana)

lanceolata) Sugar maple (Acer saccharum var.

Sycamore (Platanus occidentalis) glaucum)

River birch (Betula nigra)

25. Low, alluvial woods along Gasconade River, east of Mossy
Spring, sec. 2, T. 36 N., R. 11 W., seven miles northeast of Waynes-
ville, Pulaski County.

STUDIES OF THE VEGETATION OF MISSOURI — I 427

Dominant Trees

Green ash (Fraxinus pensylvanica var. American elm (Ulmus americana)
lanceolata) Silver maple (Acer saccharinum)

Other Tree Associates

Bur oak (Quercus macrocarpa)

Dominant Understory

Buck brush (Symphoricarposorbiculatus) Poison ivy (Toxicodendron radicans)

Other Understory Associates

Trumpet creeper (Campsis radicans) Elderberry (Sambucus canadensis)

(Young) Silver maple (Acer sacchari- Hispid greenbrier (Smilax hispida)

num) Frost grape (Vitfe cordifolia)

(Young) Box elder (Acer Negundo) Spicebush (Benzoin aestivale)

Dominant Herbaceous Non-Grass Plants

Virginia knotweed (Polygonum vir- Clear weed (Pilea pumila var. Deamii)

ginianum) Wild goldenglow (Rudbeckia laciniata)
Common blue violet (Viola papilionacea) Ironweed (Vernonia altissima)

Goldenrod (Solidago serotina) Aster (Aster lateriflorus var. pendulus)

Blue phlox (Phlox divaricata var. La- Juba bush (Iresine rhizomatosa)

phamii) Three-seeded mercury (Acalypha rhom-

Ruellia (Ruellia strepens) boidea)

Other Herbaceous Non- Grass Plants

Blue mist flower (Eupatorium coelesti- Horseweed (Ambrosia trifida)

num) Water pepper (Polygonum punctatum)

Beefsteak plant (Perilla frutescens) False nettle (Boehmeria cylindrica)

Spotted touch-me-not (Impatiens bi- Black nightshade (Solanum nigrum)
flora)

Dominant Grasses

White grass (Leersia virginica) Thin grass (Agrostis perennans var.

Wild rye grass (Elymus virginicus) aestivalis)

Wood reed grass (Cinna arundinacea)

26. Alluvial woods along Current River, sec. 34, T. 25 N., R. 1 E.,
six miles northeast of Bennett, Ripley County.

Dominant Trees

Shumard's red oak (Quercus Shumardii) Sycamore (Platanus occidentalis)

American elm (Ulmus americana) Black willow (Salix nigra)

White ash (Fraxinus americana) Silver maple (Acer saccharinum)
Cottonwood (Populus deltoides)

27. Alluvial banks along Black River between Mill Spring and
Williamsville, sec. 15, T. 27 N., R. 4 E., Wayne County.

Dominant Trees

Cottonwood (Populus deltoides) American elm (Ulmus americana)

River birch (Betula nigra) Sycamore (Platanus occidentalis)

Silver maple (Acer saccharinum)

Other Tree Associates

Green ash (Fraxinus pensylvanica var. Shumard's red oak (Quercus Shumardii)
lanceolata)

428 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Dominant Understory

Winter grape (Vitis cinerea) Dutchman's pipe (Aristolochia tomen-

Raccoon grape (Ampelopsis cordata) tosa)

Dominant Grasses

Spike grass (Uniola latifolia) Switch cane (Arundinaria gigantea)

28. Alluvial woods along Salt River at Asher Ford, south of
Spalding, Rails County.

Dominant Trees

Hackberry (Celtis occidentalis var. American elm (Ulmus americana)
canina) Bur oak (Quercus macrocarpa)

Other Tree Associates

Red mulberry (Moms rubra) Honey locust (Gleditsia triacanthos)

Red haw (Crataegus mollis) Sycamore (Platanus occidentalis)

Dominant Understory

Elderberry (Sambucus canadensis) Bristly greenbrier (Smilax hispida)

Dominant Herbaceous Non- Grass Plants

Aster (Aster lateriflorus var. pendulus) Three-seeded mercury (Acalypha rhom-
Climbing buckwheat (Polygonum scan- boidea)

dens) Goldenrod (Solidago serotina)

One-seeded bur cucumber (Sicyos angu- Clearweed (Pilea pumila var. Deamii)

latus) Horseweed (Ambrosia trifida)

Pale-flowered touch-me-not (Impatiens Actinomeris (Actinomeris alternifolia)

pallida) Slender nettle (Urtica procera)

Dominant Grasses

Wild rye (Elymus canadensis) White grass (Leersia virginica)

29. Alluvial woods along Hazel Creek, sec. 14, T. 36 N., R. 1 W.,
one mile southwest of Palmer, Washington County.

Dominant Trees

Sycamore (Platanus occidentalis) Black cherry (Prunus serotina)

American elm (Ulmus americana) White ash (Fraxinus americana)

Honey locust (Gleditsia triacanthos Chinquapin oak (Quercus Muhlenbergii)

var. inermis) Shingle oak (Quercus imbricaria)
Bur oak (Quercus macrocarpa)

Dominant Understory

Smooth alder (Alnus rugosa) Redbud (Cercis canadensis)

Ward's willow (Salix longipes var. Button-bush (Cephalanthus occidentalis)
Wardii) Ironwood (Carpinus caroliniana)

30. Alluvial woods along spring branch of Montauk Spring in
Montauk State Park, near Montauk, Dent County.

Dominant Trees

River birch (Betula nigra) American elm (Ulmus americana)

Black walnut (Juglans nigra) Black willow (Salix nigra)

Shingle oak (Quercus imbricaria)

STUDIES OF THE VEGETATION OF MISSOURI — I 429

Dominant Understory

Chittim-wood (Bumelia lanuginosa) Black haw (Viburnum rufidulum)

Smooth alder (Alnus rugosa)

31. Alluvial, lime-bouldered banks along White River between
Forsyth and Moore's Ferry, Taney County.

Dominant Trees

Sugar maple (Acer saccharum var. Green ash (Fraxinus pensylvanica var.

glaucum) lanceolate.)

American elm ( Ulmus americana)

Other Tree Associates

Hackberry (Celtis occidentalis) Bur oak (Quercus macrocarpa)

River birch (Betula nigra) Shumard's red oak (Quercus Shumardii)

Catalpa (Catalpa speciosa)

Dominant Understory

Ironwood (Carpinus caroliniana) Swamp privet (Forestiera acuminata)

32. Alluvial woods at Hall's Bluff, south of Davidson's Blue
Spring along Black River, sec. 4, T. 27 N., R. 4 E., four miles south-
east of Mill Spring, Wayne County.

Dominant Trees

Box elder (Acer Negundo) Hackberry (Celtis occidentalis)

American elm (Ulmus americana) Sweet gum (Liquidambar Styraciflua)

(4) Acer saccharum-Carya cordiformis Associes

33. Alluvial woods along Eleven Points River, sec. 18, T. 24 N.,
R. 2 W., three and one-half miles south of Wilderness, Oregon
County.

Dominant Trees
American elm (Ulmus americana) Sycamore (Platanus occidentalis)

Other Tree Associates

Shumard's red oak (Quercus Shumardii) Chinquapin oak (Quercus Muhkn-

Bur oak (Quercus macrocarpa) bergii)

Black willow (Salix nigra) Linden (Tilia glabra)

River birch (Betula nigra) Bitternut hickory (Carya cordiformis)

Northern red oak (Quercus borealis var. Pignut hickory (Carya ovalis var. obo-

maxima) valis
Persimmon (Diospyros virginiana)

Dominant Understory

Pawpaw (Asimina triloba)

Other Understory Associates

Button-bush (Cephalanthus occidentalis) Ironwood (Carpinus caroliniana)
Hop tree (Ptelea trifoliata) Chittim-wood (Bumelia lanuginosa)

430 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

34. Alluvial woods at base of west-facing limestone slopes along
Otter Creek, sec. 18, T. 27 N., R. 6 E., near Taskee, Wayne County.

Dominant Trees

Sugar maple (Acer saccharum var. Winged elm (Ulmus alata)

glaucum) American elm (Ulmus americana)

Sycamore (Platanus occidentalis) Box elder (Acer Negundo)

Bitternut hickory (Carya cordiformis) River birch (Betula nigra)

Dominant Understory

Button-bush (Cephalanthus occidentalis) Red-flowered buckeye (Aesculus dis-
Hop hornbeam (Ostrya virginiana) color var. mollis)

Alder (Alnus rugosa) Spicebush (Benzoin aestivale)

Mulberry (Morus rubra) Persimmon (Diospyros virginiana)

Black haw (Viburnum rufidulum) Hazelnut (Corylus americana)

This is not an alluvial flora as developed on large streams or
flood plains, but is partly alluvial and partly some of the flora of the
adjacent lower slopes.

35. Alluvial woods along Huzzah Creek, two miles southeast of
Davisville, Crawford County.

Dominant Tree
Sugar maple (Acer saccharum var. glaucum)

Other Tree Associates

American elm (Ulmus americana) Butternut (Juglans cinerea)

Green ash (Fraxinus pensylvanica var. Blue ash (Fraxinus quadrangulata)

lanceolata) Honey locust (Gleditsia triacanthos)

Sycamore (Platanus occidentalis) Big shell-bark hickory (Carya laciniosa)
Bur oak (Quercus macrocarpa)

Dominant Understory

Ironwood (Carpinus caroliniana) Pawpaw (Asimina triloba)

Other Understory Associates

Ohio buckeye (Aesculus glabra f . pallida) Raccoon grape (Ampelopsis cordata)

Dominant Herbaceous Non-Grass Plants

Beggar's lice (Lappula virginiana) Three-seeded mercury (Acalypha rhom-

Beefsteak plant (Perilla frutescens) boidea)

36. Rich woods built up on alluvial formation at the base of
Hall's Bluff, south of Davidson's Blue Spring along Black River,
sec. 4, T. 27 N., R. 4 E., four miles southeast of Mill Spring, Wayne
County.

Dominant Trees

Sugar maple (Acer saccharum var. Mulberry (Morus rubra)

glaucum) Chinquapin oak (Quercus Muhlenbergii)

Kentucky coffee tree (Gymnocladus Northern red oak (Quercus borealis var.
dioica) maxima)

Walnut (Juglans nigra) Persimmon (Diospyros virginiana)

Butternut (Juglans cinerea) Hackberry (Celtis laevigata)

STUDIES OF THE VEGETATION OF MISSOURI — I 431

Dominant Understory

Spicebush (Benzoin aestivale) Pawpaw (Asimina triloba)

American hornbeam (Carpinus caro- Flowering dogwood (Cornus florida)

liniana) Hazelnut (Corylus americana)

American bladdernut (Staphylea tri- Greenbrier (Smilax hispida)

folia)

Dominant Herbaceous Non- Grass Plants

Honewort (Cryptotaenia canadensis) Virginia tickseed (Lappula virginiana)

Blue phlox (Phlox divaricata var. Blopdroot (Sanguinaria canadensis)

Laphamii) Maidenhair fern (Adiantum pedatum)

Virginia knotweed (Polygonum virgini- Jack-in-the-pulpit (Arisaema triphyl-

anum) lum)

Dominant Grass

Fescue grass (Festuca obtusa)

37. Rich, alluvial bottom woods along Cane Creek, sec. 17,
T. 26 N., R. 4 E., six miles southwest of Williamsville, Wayne
County.

Dominant Trees

Bitternut hickory (Carya cordiformis) American elm (Ulmus americana)

White oak (Quercus alba) Sweet gum (Liquidambar Styraciflua)

Northern red oak (Quercus borealis var. River birch (Betula nigra)

maxima) Honey locust (Gleditsia triacanthos)

Shingle oak (Quercus imbricaria) Silver maple (Acer saccharinum)

American ash (Fraxinus americana) Cottonwood (Populus deltoides)

Walnut (Juglans nigra) Sycamore (Platanus occidentalis)
Mulberry (Morus rubra)

Dominant Understory

American hornbeam (Carpinus caro- Buck brush (Symphoricarpos orbicu-

liniana) latus)

Flowering dogwood (Cornus florida) Hazelnut (Corylus americana)

Redbud (Cercis canadensis) Pawpaw (Asimina triloba)

Black haw (Viburnum rufidulum) Alder (Alnus rugosa)

Red-flowering buckeye (Aesculus dis- Elderberry (Sambucus canadensis)

color var. mollis)

Dominant Grasses

Spike grass (Uniola latifolia) Fowl meadow grass (Glyceria striata)

Wood reed grass (Cinna arundinacea)

38. Along Ten Mile Creek, loaded with chert from adjacent
slopes, sec. 9, T. 25 N., R. 4 E., 13 miles northwest of Poplar Bluff,
Butler County.

Dominant Trees

Northern red oak (Quercus borealis var. Red maple (Acer rubrum)

maxima) Silver maple (Acer saccharinum)

White oak (Quercus alba) White ash (Fraxinus americana)

Shingle oak (Quercus imbricaria) Sweet gum (Liquidambar Styraciflua)

River birch (Betula nigra) Winged elm (Ulmus alata)
Mockernut hickory (Carya tomentosa)

432 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Understory

Ozark witch hazel (Hamamelis vernalis) Smooth alder (Alnus rugosa)
Ward's willow (Salix longipes var. Button-bush (Cephalanthus occidentalis)
Wardii)

Other Understory Associates

Persimmon (Diospyros virginiana) Chittim-wood (Bumelia lanuginosa)

Black haw (Viburnum prunifolium) Red-flowered buckeye (Aesculus dis-

Black haw (Viburnum rufidulum) color var. mollis)

Buck brush (Symphoricarpos orbiculatus) Ironwood (Carpinus caroliniana)

Dogwood (Cornus florida) Shadbush, serviceberry (Amelanchier

Dwarf sumac (Rhus Copallina) canadensis)

Smooth sumac (Rhus glabra) Ninebark (Physocarpus opulifolius var.

Deerberry (Vaccinium stamineum var. intermedius)

neglectum) Hop tree (Ptelea trifoliate)

Dominant Herbaceous Non-Grass Plants

Rough button-weed (Diodia teres var. Water willow (Dianthera americana)
setigera)

39. Rich, low woods along Big Barren Creek, near "Blue Spring,"
sec. 4, T. 25 N., R. 1 W., two and one-half miles southwest of East-
wood, 10 miles northwest of Bennett, Carter County.

Dominant Trees

Linden (Tilia glabra) Northern red oak (Quercus borealis var.

Bitternut hickory (Carya cordiformis) maxima)

White ash (Fraxinus americana) Sycamore (Platanus occidentalis)

American elm (Ulmus americana) Winged elm (Ulmus alata)

Butternut (Juglans cinerea) Sour gum (Nyssa sylvatica var. caro-

White oak (Quercus alba) niana)

Chinquapin oak (Quercus Muhlenbergii)

Dominant Understory

Ironwood (Carpinus caroliniana) Fragrant sumac (Rhus aromatica)

Spicebush (Benzoin aestivale) Buck brush (Symphoricarpos orbicu-

Hazelnut (Corylus americana) latus)

Black haw (Viburnum rufidulum) Redbud (Cercis canadensis)

40. Alluvial woods along Eleven Points River opposite west-
facing slopes, sec. 11, T. 24 N., R. 3 W., one mile southeast of
Turner's Mill, four miles southwest of Wilderness, Oregon County.

Dominant Trees

American elm (Ulmus americana) Sycamore (Platanus occidentalis)

Other Tree Associates

Shumard's red oak (Quercus Shumardii) Persimmon (Diospyros virginiana)

Bur oak (Quercus macrocarpa) Linden (Tilia glabra)

Black willow (Salix nigra) Butternut (Juglans cinerea)

River birch (Betula nigra) Bitternut hickory (Carya cordiformis)

Northern red oak (Quercus borealis var. Pignut hickory (Carya ovalis var.

maxima) obovalis)

Chinquapin oak (Quercus Muhlenbergii)

Dominant Understory

Pawpaw (Asimina triloba)

STUDIES OF THE VEGETATION OF MISSOURI — I 433

Other Understory Associates

Button-bush (Cephalanthus occidentalis) Ironwood (Carpinus caroliniana}
Hop tree (Ptelea trifoliata) Dogwood (Cornus florida)

Chittim-wood (Bumelia lanuginosa)

41. Low, alluvial woods around Iron Mountain Lake, five miles
south of Bismarck, St. Francois County.

Dominant Trees

Black willow (Salix nigra) Bitternut hickory (Carya cordiformis)

Swamp white oak (Quercus bicolor) Shagbark hickory (Carya ovata)

Shingle oak (Quercus imbricaria) White ash (Fraxinus americana)

White oak (Quercus alba) Sycamore (Platanus occidentalis)
Northern red oak (Quercus borealis var. Black walnut (Juglans nigra)

maxima) Sugar maple (Acer saccharum var.

Red maple (Acer rubrum) glaucum)
American elm (Ulmus americana)

Dominant Shrubs

Hazelnut (Corylus americana) Smooth alder (Alnus rugosa)

Bristly greenbrier (Smilax hispida)

42. Wooded valley floor along Brazil Creek, sec. 28, T. 38 N.,
R. 1 W., six miles northeast of Berryman, Washington County.

Dominant Trees

White oak (Quercus alba) Sycamore (Platanus occidentalis)

Sugar maple (Acer saccharum var. Bitternut hickory (Carya cordiformis)
glaucum)

Dominant Understory

Flowering dogwood (Cornus florida) Hop hornbeam (Ostrya virginiana)

43. Alluvial woods along Bee Fork of Black River, sec. 23, T. 32
N., R. 1 W., four miles northwest of Centerville, Reynolds County.

Dominant Trees

Sugar maple (Acer saccharum var. Black walnut (Juglans nigra)

glaucum) Sycamore (Platanus occidentalis)

Butternut (Juglans cinerea) American elm (Ulmus americana)
Chinquapin oak (Quercus Muhlenbergii)

Dominant Understory

Ironwood (Carpinus caroliniana) Buck brush (Symphoricarpos orbicu-
Pawpaw (Asimina triloba) latus)

Leatherwood (Dirca palustris) Flowering dogwood (Cornus florida)

Elderberry (Sambucus canadensis) Frost grape (Vitis cordifolia)

44. Alluvial woods along Brushy Creek, sec. 24, T. 32 N., R. 1 W.,
one mile north of Moses Store, Reynolds County.

Dominant Trees

Shumard's red oak (Quercus Shumardii) Bitternut hickory (Carya cordiformis)

American elm (Ulmus americana) Sugar maple (Acer saccharum var.

Sycamore (Platanus occidentalis) glaucum)
Butternut (Juglans cinerea)

434 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Dominant Understory

Pawpaw (Asimina iriloba) Buck brush (Symphoricarpos orbicu-

Spicebush (Benzoin aestivale) latus)

Moonseed (Menispermum canadense)

45. Alluvial woods along Big Piney River, nine miles west of
Licking, Texas County.

Dominant Trees

Sugar maple (Acer saccharum var. Hackberry (Celtis occidentalis var.

glaucum) canina)

Sycamore (Platanus occidentalis) Black walnut (Juglans nigra)

River birch (Betula nigra) White oak (Quercus alba)

American elm (Ulmus americana) Bur oak (Quercus macrocarpa)

White ash (Fraxinus americana) Shumard's red oak (Quercus Shumardii)

Black gum (Nyssa sylvatica var. caro- Chinquapin oak (Quercus Muhlen-

liniana) bergii)

Sassafras (Sassafras albidum var. molle)

Dominant Understory

Ohio buckeye (Aesculus glabra f. pallida)

46. Alluvial woods along Big Creek, sec. 22, T. 31 N., R. 3 W.,
one and one-half miles west of Rat, Shannon County.

Dominant Trees

Sugar maple (Acer saccharum var. Shumard's red oak (Quercus Shumardii)
glaucum) Sycamore (Platanus occidentalis)

Other Tree Associates

Bitternut hickory (Carya cordiformis) Chinquapin oak (Quercus Muhlenbergii)

Dominant Understory

Leatherwood (Dirca palustris) Pawpaw (Asimina iriloba)

Ironwood (Carpinus caroliniana)

Other Understory Associates

Buck brush (Symphoricarpos or biculatus) Flowering dogwood (Cornus florida)

Dominant Herbaceous Non-Grass Plant

Beefsteak plant (Perilla frutescens)

II. ACER SACCHARUM-QUERCUS ALBA ASSOCIATION

(2) Juniperus virginiana-Rhus trilobata var. serotina Associes

47. Upland limestone glade of bald knob in Hercules Game
Refuge, sec. 10-11, T. 24 N., R. 18 W., two miles southwest of
Hercules, Taney County.

Dominant Tree

Smoke-tree (Cotinus americanus)

STUDIES OF THE VEGETATION OF MISSOURI — I 435

Other Tree Associates

Persimmon (Diospyros virginiana) Red cedar (Juniperus virginiana)

Dominant Understory

Fragrant sumac (Rhus trilobata var. Greenbrier (Smilax Bona-nox)
serolina)

Understory Associate

Buck brush (Symphoricarpos orbiculatus)

Dominant Herbaceous Non- Grass Plants

Prairie clover (Petalostemum purpureum) Heliotropium tenellum

Stinging nettle (Tragia urticaefolia) Sandwort (Arenaria patula)

Hogwort (Croton capitatus) Palafoxia callosa
Linum sulcatum

Dominant Grasses

Side-oats grama (Bouteloua curti- Big blue-stem (Andropogon provincialis)
pendula)

48. Limestone bald knobs around Swan U. S. Lookout Tower,
sec. 20, T. 25 N., R. 18 W., four miles east of Garrison, Christian
County.

Dominant Trees

Persimmon (Diospyros virginiana) Winged elm (Ulmus alata)

Red cedar (Juniperus virginiana)

Dominant Understory

Supple-jack (Berchemia scandens)

Dominant Herbaceous Non- Grass Plants

Calamint (Satureja glabella var. angustifolia)

Other Herbaceous Non-Grass Plant Associates

Blue false indigo (Baptisia minor) Black-eyed Susan (Rudbeckia missouri-

Feverfew (Parthenium hispidum) ensis)

Dominant Grass

Side-oats grama (Bouteloua curtipendula)

49. Limestone slopes in small ravine between upper exposed
parts of bald knobs in Hercules Game Refuge, sec. 10-11, T. 24 N.,
R. 18 W., two miles southwest of Hercules, Taney County.

Dominant Trees

Schneck's red oak (Quercus Shumardii Smoke tree (Cotinus americanus)
var. Schneckii) Chinquapin oak (Quercus Muhlenbergii)

Other Tree Associates

Redbud (Cercis canadensis) Fringe tree (Chionanthus virginica)

Dominant Understory

Fragrant sumac, pole-cat bush (Rhus trilobata var. serotina)

436 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

(4) Ulmus alata-Rhamnus caroliniana Associes

50. Low woods along branch in Stilwell Hollow, Low Gap Game
Refuge, southeast of Black U. S. Lookout, sec. 12, T. 32 N., R. 1 E.,
four miles southwest of Black, Reynolds County.

Dominant Trees

Winged elm (Ulmus alata) White ash (Fraxinus americana)

Bitternut hickory (Carya cordiformis) Chinquapin oak (Quercus Muhlenbergii)

Dominant Understory

Pawpaw (Asimina triloba) Redbud (Cercis canadensis)

Buck brush (Symphoricarpos orbiculatus)

Dominant Herbaceous Non-Grass Plant

Beefsteak plant (Perilla frutescens)

(5) Quercus Muhlenbergii-Fraxinus americana Associes

51. Limestone glades and ledges on bluffs along East Fork of
Bull Creek, sec. 26, T. 26 N., R. 20 W., three miles southwest
of Chadwick, Christian County.

Dominant Trees

Schneck's red oak (Quercus Schumardii Chinquapin oak (Quercus Muhlenbergii)
var. Schneckii)

Dominant Shrubs

Chittim-wood (Bumelia lanuginosa) Winged elm (Ulmus alata)

Black haw (Viburnum rufidulum)

Other Shrub Associates

Redbud (Cercis canadensis) Persimmon (Diospyros virginiana)
Indian cherry (Rhamnus caroliniana) Rough-leaved dogwood (Cornus aspen-
Fragrant sumac (Rhus trilobata var. folia)
serotina)

Dominant Herbaceous Non- Grass Plants

Black-eyed Susan (Rudbeckia missouri- Houstonia nigricans
ensis)

Other Herbaceous Non-Grass Plant Associates

Wine-cups (Callirhoe digitata) Gum-plant (Grindelia lanceolata)

Dominant Grasses

Blue-stem (Andropogon scoparius var. Side-oats grama (Boutelouacurtipendula)
genuinus) Big blue-stem (Andropogon provincialis)

52. Limestone bluffs along Hazel Creek, sec. 14, T. 36 N., R. 1 W.,
one mile southwest of Palmer, Washington County.

Dominant Trees

Slippery elm (Ulmus fulva) White ash (Fraxinus americana)

Chinquapin oak (Quercus Muhlenbergii)

STUDIES OF THE VEGETATION OF MISSOURI — I 437

Dominant Understory

Rough-leaved dogwood (Cornus asperi- Swamp dogwood (Cornus obliqua)
folia) Black haw (Viburnum rufidulum)

53. Northwest-facing slopes along Marble Creek, sec. 20, T. 32
N., R. 5 E., two miles southwest of French Mills, Madison County.

Dominant Trees

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.

glaucum) maxima)

Chinquapin oak (Quercus Muhlenbergii)

Other Tree Associates

White ash (Fraxinus americana) Linden (Tilia glabra)

Butternut (Juglans cinerea) American elm (Ulmus americana)

Kentucky coffee tree (Gleditsia triacan- Blue ash (Fraxinus quadrangulata)

thos) Slippery elm (Ulmus fulva)

Dominant Understory

Indian cherry (Rhamnus caroliniana) Black haw (Viburnum rufidulum)
Ninebark (Physocarpus opulifolius var. Hop hornbeam (Ostrya virginiana)
intermedius) Red cedar (Juniperus virginiana)

54. Lower, west-facing, limestone slopes along Eleven Points
River, sec. 11, T. 24 N., R. 3 W., one mile southeast of Turner's
Mill, four miles southwest of Wilderness, Oregon County.

Dominant Trees

Chinquapin oak (Quercus Muhlenbergii) Hackberry (Celtis occidentalis)

White ash (Fraxinus americana) American elm (Ulmus americana)

Redbud (Cercis canadensis) Blue ash (Fraxinus quadrangulata)

Butternut (Juglans cinerea) Shumard's red oak (Quercus Shumardii)

Linden (Tilia glabra) Box elder (Acer Negundo)
Sugar maple (Acer saccharum)

Dominant Understory

Buck brush (Symphoricarpos orbicu- American bladdernut (Staphylea tri-

latus) folia)

Pawpaw (Asimina triloba) Black haw (Viburnum rufidulum)

Ironwood (Carpinus caroliniana) Rough-leaved dogwood (Cornus asperi-

Spicebush (Benzoin aestivale) folia
Red mulberry (Morus rubra)

55. West-facing, wooded, limestone slopes along Castor River,
at Hahn's Mill, sec. 16, T. 33 N., R. 8 E., four miles northeast of
Cornwall, Madison County.

Dominant Trees

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.

glaucum) maxima)

White oak (Quercus alba)

Other Tree Associates

Red maple (Acer rubrum) Linden (Tilia glabra)

White ash (Fraxinus americana) Chinquapin oak (Quercus Muhlenbergii)

Shagbark hickory (Carya ovata)

438 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Understory
Ironwood (Carpinus caroliniana)

Other Understory Associates

Black haw (Viburnum rufidulum) Hop hornbeam (Ostrya virginiana)

Flowering dogwood (Cornus florida) Bladdernut (Staphylea trifolia)

Smooth alder (Alnus rugosd)

56. North-facing, wooded, limestone slopes along Brazil Creek,
sec. 28, T. 38 N., R. 1 W., six miles northeast of Berryman, Wash-
ington County.

UPPER PORTION OF SLOPES

Dominant Trees

Sugar maple (Acer saccharum var. Schneck's red oak (Quercus Shumardii

glaucum) var. Schneckii)

Northern red oak (Quercus borealis var. Chinquapin oak (Quercus Muhlenbergii)

maxima)

LOWER PORTION OF SLOPES

Dominant Trees

Linden (Tilia glabra) Chinquapin oak (Quercus Muhlenbergii)

Sugar maple (Acer saccharum var. American elm ( Ulmus americana)
glaucum)

Dominant Understory

Chittim-wood (Bumelia lanuginosa) Redbud (Cercis canadensis)

Black haw (Viburnum rufidulum) Ironwood (Carpinus caroliniana)

Indian cherry (Rhamnus caroliniana Bladdernut (Staphylea trifolia)

var. mollis) Hackberry (Celtis pumila var. georgiana)

57. One-half to two-thirds the way up south-facing limestone
slopes along Bee Fork of Black River, sec. 29, T. 32 N., R. 1 W.,
seven miles west of Centerville, Reynolds County.

Dominant Tree

Chinquapin oak (Quercus Muhlenbergii)

Tree Associate

Shagbark hickory (Carya ovata)

Dominant Understory

Flowering dogwood (Cornus florida) Black haw (Viburnum rufidulum)

Other Understory Associates

Hop hornbeam (Ostrya virginiana) Chittim-wood (Bumelia lanuginosa)

Greenbrier (Smilax Bona-nox)

58. Two-thirds to three-fourths the way up south-facing lime-
stone slopes along Bee Fork of Black River, sec. 29, T. 32 N., R. 1 W.,
seven miles west of Centerville, Reynolds County.

STUDIES OF THE VEGETATION OF MISSOURI — I 439

Dominant Trees

Schneck's red oak (Quercus Shumardii White oak (Quercus alba)
var. Schneckii)

Other Tree Associates

Post oak (Quercus stellata) Slippery elm (Ulmusfulva)

Shingle oak (Quercus imbricaria) Persimmon (Diospyros virginiana)

Dominant Understory

Dwarf hackberry (Celtis pumila var. Wild plum (Prunus americana)
georgiana)

59. Base of limestone slopes along Eleven Points River, sec. 11,
T. 24 N., R. 3 W., one mile southeast of Turner's Mill, four miles
southwest of Wilderness, Oregon County.

Dominant Trees

Northern red oak (Quercus borealis var. American elm (Ulmus americana)

maxima) White ash (Fraxinus americana)

Chinquapin oak (Quercus Muhlenbergii) Black walnut (Juglans nigra)
Linden (Tilia glabra)

Tree Associate

Persimmon (Diospyros virginiana)

Dominant Understory

American bladdernut (Staphylea tri- Pawpaw (Asimina triloba)
folia) Redbud (Cercis canadensis)

Other Understory Associates

Ironwood (Carpinus caroliniana) Hazelnut (Corylus americana)

Spicebush (Benzoin aestivale) Black haw (Viburnum rufidulum)

60. West-facing limestone slopes along Current River, sec. 34,
T. 25 N., R. 1 E., five miles northeast of Bennett, Ripley County.

Dominant Trees

Sugar maple (Acer saccharum) Persimmon (Diospyros virginiana)

Sweet gum (Liquidambar Styraciflua) White ash (Fraxinus americana)

Linden (Tilia glabra) Red cedar (Juniperus virginiana)
Chinquapin oak (Quercus Muhlenbergii)

Dominant Understory

Black haw (Viburnum rufidulum) Ninebark (Physocarpus opulifolius var.
Redbud (Cercis canadensis) intermedius)

Possum haw (Ilex decidua) Ironwood (Carpinus caroliniana)

Hop hornbeam (Ostrya virginiana) Indian cherry (Rhamnus caroliniana)

Rough-leaved dogwood (Cornus asperi- Sassafras (Sassafras albidum var. molle)
folia)

Dominant Herbaceous Non-Grass Plants

Dutchman's pipe (Aristolochia tomen- Rattle weed (Astragalus canadensis)
tosa)

Dominant Grass

Switch cane (Arundinaria gigantea)

440 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

61. Wooded limestone bluffs along Beaver Creek, sec. 21 and 22,
T. 24 N., R. 18 W., three miles southwest of Bradleyville, Taney
County.

Dominant Trees

Chinquapin oak (Quercus Muhlenbergii) Smoke tree (Cotinus americanus)

Other Tree Associates

Sugar maple (Acer saccharum var. Butternut (Juglans cinerea)

glaucum) American elm (Ulmus americana)

Hackberry (Celtis occidentalis var. ca- Linden (Tilia glabra)
nina)

Dominant Understory
Spicebush (Benzoin aestivale)

Understory Associate

Persimmon (Diospyros virginiana)

62. Limestone glade on bluffs along Eleven Points River, sec. 11,
T. 24 N., R. 3 W., one mile southeast of Turner's Mill, four miles
southwest of Wilderness, Oregon County.

Dominant Trees

Chinquapin oak (Quercus Muhlenbergii) Hackberry (Celtis laevigata var. texana)

Dominant Understory

Winged elm (Ulmus alata) Chittim-wood (Bumelia lanuginosa)

Hop hornbeam (Ostrya virginiana) Fragrant sumac (Rhus aromatica)

Black haw (Viburnum rufidulum) Shadbush (Amelanchier canadensis)

Buck brush (Symphoricarpos orbiculatus) Greenbrier (Smilax Bona-nox)

63. West-facing limestone slopes along Middle Fork of Black
River, sec. 20, T. 34 N., R. 1 E., one mile north of Redmondville,
Iron County.

Dominant Trees

Sugar maple (Acer saccharum var. Chinquapin oak (Quercus Muhlenbergii)
glaucum) Slippery elm (Ulmus fulva)

Other Tree Associates

Schneck's red oak (Quercus Shumardii Red cedar (Juniperus virginiana)
var. Schneckii) White ash (Fraxinus americana)

Dominant Understory

Chittim-wood (Bumelia lanuginosa) Black haw (Viburnum rufidulum)

Hop hornbeam (Ostrya virginiana)

Dominant Grasses

Wild chess (Bromus purgans) Sphenopholis (Sphenopholis intermedia)

(6) Acer sacchamm-Quercus alba Associes

64. At base of south-facing limestone slopes along Bee Fork of
Black River, sec. 29, T. 32 N., R. 1 W., seven miles west of Center-
ville, Reynolds County.

STUDIES OF THE VEGETATION OF MISSOURI — I 441

Dominant Trees

Sugar maple (Acer saccharum var. Slippery elm (Ulmus fulva)
glaucum) Winged elm (Ulmus alata)

White oak (Quercus alba) Sassafras (Sassafras albidum var. molle)

Northern red oak (Quercus borealis var. Red mulberry (Morus rubra)
maxima) Bitternut hickory (Carya cordiformis)

Tree Associate

Persimmon (Diospyros virginiana)

Dominant Understory

Black haw (Viburnum rufidulum) Redbud (Cercis canadensis)

Other Understory Associates

Fragrant sumac (Rhus aromatica) Ironwood (Carpinus caroliniana)

Greenbrier (Smilax Bona-nox) Smooth sumac (Rhus glabra)

Frost grape (Vitis cordifolia)

65. Northeast-facing, limestone, wooded slopes along Black
River, north of Markham Spring, sec. 23, T. 27 N., R. 4 E., three
miles west of Williamsville, Wayne County.

Dominant Trees

Sugar maple (Acer saccharum var. American elm (Ulmus americana)

glaucum) White ash (Fraxinus americana)

Chinquapin oak (Quercus Muhlenbergii) Sweet gum (Liquidambar Styraciflua)

Northern red oak (Quercus borealis var. Sycamore (Platanus occidentalis)

maxima) Bitternut hickory (Carya cordiformis)

Butternut (Juglans cinerea) Linden (Tilia glabra)

Dominant Understory

Ironwood (Carpinus caroliniana) Frost grape (Vitis cordifolia)

Rough-leaved dogwood (Cornus asperi- Button-bush (Cephalanthus occidentalis)

folia) Black haw (Viburnum rufidulum)

Spicebush (Benzoin aestivale) Burning bush (Euonymus atropurpu-

American bladdernut (Staphylea trifolia) reus)

Hydrangea (Hydrangea arborescens) Hop hornbeam (Ostrya virginiana)

Flowering dogwood (Cornus florida) Poison ivy (Toxicodendron radicans)

Bristly greenbrier (Smilax hispida) Smooth alder (Alnus rugosa)

Dominant Herbaceous Non-Grass Plant

Liverleaf (Hepatica acutiloba)

Dominant Grasses

Fescue grass (Festuca obtusa) Switch cane (Arundinaria gigantea)

66. Along south-facing limestone bluffs of King's River, one
mile southwest of Prentiss Ford, sec. 19-24, T. 21 N., R. 25 W., five
miles south of Viola, Barry County.

Dominant Trees

Sugar maple (Acer saccharum var. Chinquapin oak (Quercus Muhlenbergii)
glaucum)

Other Tree Associates

Yellow-wood (Cladrastis lutea) Red mulberry (Morus rubra)

American elm (Ulmus americana) Black walnut (Juglans nigra)

442 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Understory

Redbud (Cerds canadensis) Hackberry (Celtis occidentalis)

Other Understory Associates

Indian cherry (Rhamnus caroliniana Spicebush (Benzoin aestivale)

var. mollis) Black haw (Viburnum rufidulum)

Red cedar (Juniperus virginiana) Buck brush (Symphoricarpos orbicu-

Pawpaw (Asimina triloba) latus)

Dominant Herbaceous Non-Grass Plants

White snakeroot (Eupatorium urticae- Thimble weed (Anemone virginiana)

folium) Beggar's ticks (Desmodium acuminatum)

Black snakeroot (Sanicula canadensis)

Other Herbaceous Non- Grass Plants

Frost grape (Vitis cordifolia) Solomon's seal (Polygonatum canalicu-

Yellow violet (Viola eriocarpa) latum)

Lopseed (Phryma Leptostachya) Virgin's bower (Clematis virginiana var.
American bellflower (Campanula ameri- missouriensis)

cana) Leather flower (Clematis versicolor)

Wild licorice (Galium circaezans var. Skull-cap (Scutellaria ovata)

hypomalacum) Skull-cap (Scutellaria ovalifolia)

Virginia knotweed (Polygonum vir- Bear grass (Yucca arkansana)

ginianum)

Dominant Grasses

Spike grass (Uniola latifolia) Wild chess (Bromus purgans)

67. East-facing limestone bluffs along Huzzah Creek, two miles
southeast of Davisville, Crawford County.

Dominant Trees

Northern red oak (Quercus borealis var. Chinquapin oak (Quercus Muhlenbergii)
maxima)

Other Tree Associates

Sugar maple (Acer saccharum var. American elm (Ulmus americana)

glaucum) Blue ash (Fraxinus quadrangulata)

Linden (Tilia glabra) Red mulberry (Morus rubra)

White ash (Fraxinus americana) Sour gum (Nyssa sylvatica var. caro-
Black walnut (Juglans nigra) liniana)

Bitternut hickory (Carya cordiformis) Shagbark hickory (Carya ovata)

Dominant Understory
Bladdernut (Staphylea trifolia)

Other Understory Associates

Spicebush (Benzoin aestivale) Black haw (Viburnum rufidulum)

Sassafras (Sassafras albidum var. molle) Pawpaw (Asimina triloba)

Burning bush (Euonymus atropur- Redbud (Cercis canadensis)

pureus) Rough-leaved dogwood (Cornus asperi-
Flowering dogwood (Cornus florida) folia)

Persimmon (Diospyros virginiana)

Dominant Herbaceous Non-Grass Plants

Bristly greenbrier (Smilax hispida) Wild yam (Dioscorea quaternata)

Maidenhair fern (Adiantum pedatum) Leafcup (Polymnia canadensis)

STUDIES OF THE VEGETATION OF MISSOURI — I 443

68. Wooded talus at base of east-facing limestone bluff, "Log-
slide Bluff," along White River, sec. 22-27, T. 22 N., R. 23 W., two
miles west of Marmaros, Stone County.

Dominant Trees

Linden (Tilia glabra) Bitternut hickory (Carya cordiformis)

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.
glaucum) maxima)

Other Tree Associates

Butternut (Juglans cinerea) Chinquapin oak (Quercus Muhknbergii)

Hackberry (Celtis occidentalis var. Yellow-wood (Cladrastis lutea)
canina)

Dominant Understory

Bladdernut (Staphylea trifolia) Pawpaw (Asimina triloba)

Ironwood (Carpinus caroliniana)

Understory Associate

Spicebush (Benzoin aestivale)

Dominant Herbaceous Non- Grass Plant

Leafcup (Polymnia canadensis)

Other Herbaceous Non- Grass Plant Associates

Pale touch-me-not (Impatiens pallida) Enchanter's nightshade (Circaea lati-

Waterleaf (Hydrophyllum appendicu- folia)

latum) Wild ginger (Asarum canadense var.

Yellow violet (Viola eriocarpa var. acuminatum)

leiocarpa) Celandine poppy (Stylophorum diphyl-

May-apple (Podophyllum pellatum) lum)
Moonseed (Menispermum canadense)

69. Rich, wooded, north-facing, limestone slopes along spring
branch tributary to Swan Creek, sec. 34, T. 26 N., R. 19 W., four
miles southeast of Chadwick, Christian County.

Dominant Trees

Black maple (Acer nigrum) Sugar maple (Acer saccharum var.

Chinquapin oak (Quercus Muhlenbergii) glaucum)

Other Tree Associates

Northern red oak (Quercus borealis var. Bur oak (Quercus macrocarpa)

maxima) American elm (Ulmus americana)

Blue ash (Fraxinus quadrangulata)

Dominant Understory

Pawpaw (Asimina triloba) Spicebush (Benzoin aestivale)

Bladdernut (Staphylea trifolia)

Understory Associate

Arrow-wood (Viburnum molle)

Dominant Herbaceous Non-Grass Plants

Waterleaf (Hydrophyllum appendicu- Liverleaf (Hepatica americana)

latum) Moonseed (Menispermum canadense)

Leafcup (Polymnia canadensis)

444 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Other Herbaceous Non-Grass Plant Associates

Cup-seed (Calycocarpum Lyoni) False spikenard (Smilacina racemosa)

Dominant Grasses

Wild chess (Bromus purgans) Bottle brush (Hystrix patula)

Fescue grass (Festuca obtusa) Diarina festucoides

70. Northwest-facing wooded slopes along Swan Creek, sec. 34,
T. 26 N., R. 19 W., four miles southeast of Chadwick, Christian
County.

Dominant Trees

Black maple (Acer nigrum) Linden (Tilia glabra)

Sugar maple (Acer saccharum var. Bitternut hickory (Carya cordiformis)
glaucum)

Other Tree Associates

White ash (Fraxinus americana) Slippery elm (Ulmus fulva)

Dominant Understory

Arrow-wood (Viburnum molle) Bladdernut (Staphylea trifolia)

Pawpaw (Asimina triloba) Spicebush (Benzoin aestivale)

Ironwood (Carpinus caroliniana)

Understory Associate

Leatherwood (Dirca palustris)

Dominant Herbaceous Non- Grass Plants

Liverleaf (Hepatica acutiloba) Black snakeroot (Cimicifuga racemosa)

Early meadow rue (Thalictrum dioicum) Leaf cup (Polymnia canadensis)

Other Herbaceous Non-Grass Plant Associates

Liverleaf (Hepatica americana) Indian turnip (Arisaema triphyllum)

Joe-Pye weed (Eupatorium purpureum) Wood's hellebore (Veratrum Woodii)

71. North-facing limestone slopes along valley bordering Stin-
son Creek, near Ballinger Springs, sec. 1, T. 46 N., R. 9 W., two
miles northeast of Hams Prairie, Callaway County.

Dominant Trees

Sugar maple (Acer saccharum var. Linden (Tilia glabra)
glaucum) Black maple (Acer nigrum)

Other Tree Associates

Butternut (Juglans cinerea) Kentucky coffee tree (Gymnocladus dio-

Bitternut hickory (Carya cordiformis) ica)

Dominant Understory

Pawpaw (Asimina triloba) Ironwood (Carpinus caroliniana)

Alternate-leaved dogwood (Cornus alter- Bladdernut (Staphylea trifolia)
nifolia)

Understory Associate
Arrow-wood (Viburnum molle)

Dominant Herbaceous Non-Grass Plant

Woollen breeches ( Hydrophyllum appendiculatum)

STUDIES OF THE VEGETATION OF MISSOURI — I 445

Other Herbaceous Non-Grass Plants

Celandine poppy (Stylophorum diphyl- White avens (Geum canadense var.

lum) camporum)

Goldenrod (Solidago latifolia) Wild geranium (Geranium maculatum)

Clearweed (Pilea pumila var. Deamii) Bloodroot (Sanguinaria canadensis)

Narrow-leaved spleenwort (Athyrium Spiderwort (Tradescantia subaspera)

pycnocarpori) Maidenhair fern (Adiantum pedatum)
Leafcup (Polymnia canadensis)

72. North-facing limestone bluffs with wooded talus slopes
below, along Osage River, sec. 2, T. 40 N., R. 13 W., west of Capps,
Miller County.

Dominant Trees

Sugar maple (Acer saccharum var. Chinquapin oak (Quercus Muhlenbergii)
glaucum)

Other Tree Associates

Linden (Tilia glabra) Northern red oak (Quercus borealis var.

Butternut (Juglans cinerea) maxima)

White oak (Quercus alba)

Dominant Understory

American hornbeam (Carpinus caro- Bladdernut (Staphylea trifolia)
liniana) Flowering dogwood (Cornus florida)

Other Understory Associates

Benzoin (Benzoin aestivale) Pawpaw (Asimina triloba)

Hydrangea ( Hydrangea arborescens)

Dominant Herbaceous Non-Grass Plant

Goldenrod (Solidago latifolia)

73. North-facing limestone bluffs along Little Niangua River,
sec. 5, T. 38 N., R. 18 W., two and one-half miles south of Barnum-
town, Camden County.

Dominant Trees

Sugar maple (Acer saccharum var. Bitternut hickory (Carya cordiformis) —

glaucum) at base

Black maple (Acer nigrum)

Other Tree Associates

Linden (Tilia glabra) Northern red oak (Quercus borealis var.

Butternut (Juglans cinerea) maxima)

Dominant Understory

Bladdernut (Staphylea trifolia) American hornbeam (Carpinus caro-

Benzoin (Benzoin aestivale) liniana}

Pawpaw (Asimina triloba)

Dominant Herbaceous Non-Grass Plants

Pale touch-me-not (Impatiens pallida) Leafcup (Polymnia canadensis)

Other Herbaceous Non-Grass Plants

Woollen breeches (Hydrophyllum appen- Narrow-leaved spleenwort (Athyrium
diculatum) pycnocarpon)

446 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

74. West-facing, wooded, limestone slope along Brushy Creek,
sec. 8, T. 27 N., R. 3 E., six miles north of Ellsinore, Carter County.

Dominant Tree

Sugar maple (Acer saccharum var. glaucum)

Other Tree Associates

Northern red oak (Quercus borealis var. White ash (Fraxinus americana)

maxima) Linden (Tilia glabra)

White oak (Quercus alba) Mulberry (Moras rubra)

Chinquapin oak (Quercus Muhlenbergii) Slippery elm (Ulmusfulva)

Bitternut hickory (Gary a cordiformis) Winged elm (Ulmus alata)
Shagbark hickory (Carya ovata)

Chief Understory

Bladdernut (Staphylea trifolia) Flowering dogwood (Cornns florida)

Other Understory Associates

Possum haw (Ilex decidua) Hop hornbeam (Ostrya virginiana)
Chittim-wood (Bumelia lanuginosa) Sassafras (Sassafras albidum var. molle)
Hackberry (Celtis laevigata var. texana) Eastern witch hazel (Hamamelis virgin-
Wild plum (Prunus americana) iana)
American hornbeam (Carpinus caro- Black haw (Viburnum rufidulum)

liniana) Redbud (Cercis canadensis)
Hydrangea ( Hydrangea arborescens)

Dominant Grass-like Plant

Carex eburnea

Other Grass or Grass-like Associates

Wild chess (Bromus purgans) Fescue grass (Fesluca paradoxa)

Indian grass (Sorghastrum nutans) Blue-stem grass (Andropogon scoparius)

75. Lower portion of northeast-facing limestone slopes along St.
Francis River, just west of Greenville, Wayne County.

Dominant Trees

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.

glaucum) maxima)

Chinquapin oak (Quercus Muhlenbergii)

Dominant Understory

Black haw (Viburnum rufidulum) Hazelnut (Corylus americana)

White ash (Fraxinus americana) Redbud (Cercis canadensis)

Pawpaw (Asimina triloba) Ironwood (Carpinus caroliniana)

Indian cherry (Rhamnus caroliniana) Hop tree (Ptelea trifoliata)

Dominant Herbaceous Non-Grass Plants

Liverleaf (Hepatica acutiloba) Early meadow rue (Thalictrum dioicum)

Dominant Grasses

Fescue grass (Fesluca obtusa) Sphenopholis (Sphenopholis intermedia)

76. North-facing, wooded, limestone slopes along Brushy Creek,
sec. 8, T. 27 N., R. 3 E., six miles north of Ellsinore, Carter County.

STUDIES OF THE VEGETATION OF MISSOURI — I 447

Dominant Trees

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.
glaucum) maxima)

Other Tree Associates

White oak (Quercus alba) Sassafras (Sassafras albidum var. molle)

White ash (Fraxinus americana) Red maple (Acer rubrum)

Bitternut hickory (Carya cordiformis) Sycamore (Platanus occidentalis)

Butternut (Juglans cinerea) Sour gum (Nyssa sylvatica var. caro-
River birch (Betula nigra) liniana)

Dominant Understory

Dogwood (Cornus florida) Ironwood (Carpinus caroliniana)

Other Understory Associates

Smooth alder (Alnus rugosa) Redbud (Cercis canadensis)

Buck brush (Symphoricarpos orbicu- Button-bush (Cephalanthus occidentalis)

latus) Spicebush (Benzoin aestivale)

Black haw (Viburnum rufidulum) Hazelnut (Corylus americana)

Shadbush, serviceberry (Amelanchier Hop hornbeam (Ostrya virginiana)

canadensis) Ozark witch hazel (Hamamelis vernalis)

American bladdernut (Staphyka tri- Ward's willow (Salix longipes var.

folia) Wardii)

Fragrant sumac (Rhus aromatica) Hydrangea (Hydrangea arborescens)

Dominant Herbaceous Non-Grass Plant

Liverleaf (Hepatica acutiloba)

Dominant Grasses

Thin grass (Agrostis perennans) Fowl meadow grass (Glyceria striata)

77. Rich, low woods and adjacent lower limestone slopes along
spring branch tributary to Current River, sec. 34, T. 25 N., R. 1 E.,
six miles northeast of Bennett, Ripley County.

Dominant Trees

Sugar maple (Acer saccharum var. Red mulberry (Morns rubra)

glaucum) Black walnut (Juglans nigra)

Red maple (Acer rubrum) Linden (Tilia glabra)

Chinquapin oak (Quercus Muhlenbergii) White ash (Fraxinus americana)

Bitternut hickory (Carya cordiformis) Northern red oak (Quercus borealis var.
Sour gum (Nyssa sylvatica var. ca.ro- maxima)

liniana) American elm (Ulmus americana)

Dominant Understory

Spicebush (Benzoin aestivale) Ironwood (Carpinus caroliniana)

Hazelnut (Corylus americana) Pawpaw (Asimina triloba)

Redbud (Cercis canadensis) Bladdernut (Staphyka tnfolia)

78. North-facing, wooded, limestone slopes along Big Creek,
sec. 18, T. 31 N., R. 2 W., three miles northeast of Rat, Shannon
County.

Dominant Trees

Sugar maple (Acer saccharum var. American linden (Tilia glabra)
glaucum) Butternut (Juglans cinerea)

448 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Dominant Understory
Eastern witch hazel (Hamamelis virgini- Flowering dogwood (Cornus florida)

ana) Running strawberry-bush (Euonymus

Spicebush (Benzoin aestivale) obovatus)

Ohio buckeye (Aesculus glabra f . pallida) Bristly greenbrier (Smilax hispida)
Bladdernut (Staphylea trifolid)

Dominant Herbaceous Non-Grass Plant

Wood's hellebore (Veratrum Woodii)

Dominant Grasses and Grass-like Plants

Brachyelytrum erectum Sedge (Carex eburnea)

79. North-facing, wooded, limestone slopes along Bee Fork of
Black River, sec. 23, T. 32 N., R. 1 W., four miles northwest of
Centerville, Reynolds County.

Dominant Trees
Sugar maple (Acer saccharum var. Butternut (Juglans cinerea)

glaucum) Red mulberry (Morus rubra)

Northern red oak (Quercus borealis var. American elm (Ulmus americana)

maxima) Linden (Tilia glabra)

Sour gum (Nyssa sylvatica var. caro- Bitternut hickory (Carya cordiformis)

liniana)

Dominant Understory of Upper Portion
Eastern witch hazel (Hamamelis virgini- Flowering dogwood (Cornus florida)

ana) Juneberry, serviceberry (Amelanchier

Hop hornbeam (Ostrya virginiana) canadensis)

Redbud (Cercis canadensis)

Dominant Understory of Lower Portion

Leatherwood (Dirca palustris) Spicebush (Benzoin aestivale)

Flowering dogwood (Cornus florida)

Dominant Herbaceous Non- Grass Plant

Liverleaf (Hepatica acutiloba)

80. East-facing limestone slopes along Brushy Creek, sec. 24,
T. 33 N., R. 1 W., one mile north of Moses Store, Reynolds County.

Dominant Trees

Northern red oak (Quercus borealis var. Sugar maple (Acer saccharum var.

maxima) glaucum)

Linden (Tilia glabra)

Tree Associate
Black cherry (Prunus serotina)

Dominant Understory

Eastern witch hazel (Hamamelis virgini- Flowering dogwood (Cornus florida)
ana)

Other Understory Associates

Ironwood (Carpinus caroliniana) Juneberry, serviceberry (Amelanchier

Spicebush (Benzoin aesiivale) canadensis)

Leatherwood (Dirca palustris) Sassafras (Sassafras albidum var. molle)

Dominant Herbaceous Non-Grass Plants

Early meadow rue (Thalictrum dioicum) Wild ginger (Asarum canadense var.
Liverleaf (Hepatica acutiloba) reflexum)

Black cohosh (Cimicifuga racemosa)

STUDIES OF THE VEGETATION OF MISSOURI— I 449

81. West-facing, cherty limestone slopes along Otter Creek,
sec. 18, T. 27 N., R. 6 E., near Taskee, Wayne County.

Dominant Trees

Sugar maple (Acer saccharum var. Black walnut (Juglans nigra)

glaucum) White ash (Fraxinus americana)

Winged elm (Ulmus alata) Sour gum (Nyssa sylvatica var. caro-

White oak (Quercus alba) liniana)

Northern red oak (Quercus borealis var. Mockernut hickory (Carya tomentosa)
maxima)

Dominant Understory

Dogwood (Cornus florida) Hop hornbeam (Ostrya virginiana)

Fragrant sumac (Rhus aromatica) Redbud (Cercis canadensis)

Sassafras (Sassafras albidum var. molle) Shadbush (Amelanchier canadensis)

American bladdernut (Staphylea tri- Indian cherry (Rhamnus caroliniana)

folia) Iron wood (Carpinus caroliniana)

82. Base of rich, low, wooded slopes just above alluvial woods
along Gasconade River, east of Mossy Spring, sec. 2, T. 36 N.,
R. 11 W., seven miles northeast of Waynesville, Pulaski County.

Dominant Trees

Northern red oak (Quercus borealis var. Bitternut hickory (Carya cordiformis)

maxima) Linden (Tilia glabra)

White oak (Quercus alba f. latiloba)

Other Tree Associates

Hackberry (Celtis occidentalis var. Slippery elm (Ulmus fulva)
canina)

Dominant Understory

Ironwood (Carpinus caroliniana) Pawpaw (Asimina triloba)

ironwood (Larpmus carolimana)
Burning bush (Euonymus atropurpureus)

Other Understory Associates

Moonseed (Menispermum canadense) Sassafras (Sassafras albidum)
Buck brush (Symphoricarpos orbicula- Virginia creeper (Parthenocissus quin-
tus) quefolia)

Dominant Herbaceous Non-Grass Plants

Pale touch-me-not (Impatiens pallida) White violet (Viola striata)

White crown-beard (Verbesina virginica) Wild ginger (Asarum canadense var.

Clearweed (Pilea pumila var. Deamii) acuminatum)

Pokeweed (Phytolacca americana)

Other Herbaceous Non-Grass Plant Associates

Blephilia (Blephilia hirsuta) Skullcap (Scutellaria ovata)

Wood nettle (Laportea canadensis) Joe-Pye weed (Eupatorium purpureum)

Yellow violet (Viola eriocarpa var. Wild yam (Dioscorea villosa)

leiocarpa) Enchanter's nightshade (Circaea lati-

Fragile fern (Cystopteris fragilis var. folia)

protrusa) Wild blue phlox (Phlox divaricata var.

Hog-peanut (Amphicarpa bracteata) Laphamii)

Fragrant bedstraw (Galium triflorum) Vervain (Verbena urticaefolia)
Bloodroot (Sanguinaria canadensis)

450 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

83. East-facing, cherty, limestone slopes along Ottery Creek,
sec. 34, T. 34 N., R. 1 E., two miles northeast of Edgehill, Iron
County.

Dominant Trees

Sugar maple (Acer saccharum var. Bitternut hickory (Carya cordiformis)

glaucum) White ash (Fraxinus americana)

Shagbark hickory (Carya ovata) Linden (Tilia glabra)

Northern red oak (Quercus borealis var. White oak (Quercus alba)
maxima) Butternut (Juglans cinerea)

Dominant Understory
Flowering dogwood (Cornus florida)

84. West-facing, porphyritic, trachyte slopes along Ottery Creek,
sec. 26, T. 34 N., R. 1 E., three miles northeast of Edgehill, Iron
County.

Dominant Tree

Winged elm ( Ulmus alata)

Other Tree Associates

Sugar maple (Acer saccharum var. White oak (Quercus alba)
glaucum) Shagbark hickory (Carya ovata)

Dominant Understory

Hop hornbeam (Ostrya virginiana) Hackberry (Celtis laevigata var. texana)

85. Limestone bluffs at mouth of spring branch of Pulltight
Spring, Shannon County.

Dominant Trees

Sugar maple (Acer saccharum var. Chinquapin oak (Quercus Muhlenbergii)
glaucum) Schneck's red oak (Quercus Shumardii

Red cedar (Juniperus virginiana) var. Schneckii)

Chittim-wood (Bumelia lanuginosa) . Persimmon (Diospyros virginiana)

86. Rich, east-facing slopes at base of limestone bluffs along
King's River, south of Prentiss Ford, sec. 24, T. 21 N., R. 25 W., five
miles south of Viola, Barry County.

Dominant Trees

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.

glaucum) maxima)

Chinquapin oak (Quercus Muhknbergii)

Dominant Understory

Black locust (Robinia Pseudo-acacia) Spicebush (Benzoin aestivale)
Pawpaw (Asimina triloba) Redbud (Cercis canadensis)

87. Limestone draw between upper part of bald knob in Hercules
Game Refuge, sec. 10-11, T. 24 N., R. 18 W., two miles southwest of
Hercules, Taney County.

STUDIES OF THE VEGETATION OF MISSOURI — I 451

Dominant Trees
Sugar maple (Acer saccharum var. Schneck's red oak (Quercus Shumardii

glaucum) var. Schneckii)

Smoke tree (Cotinus americanus)

Dominant Understory

Flowering dogwood (Cornus florida) Possum haw (Ilex decidua)

Swamp dogwood (Cornus obliqua) Ninebark (Physocarpus opulifolius var.

Rough-leaved dogwood (Cornus asperi- intermedius)

folia) Andrachne phyllanthoides

Other Understory Associates

Indian cherry (Rhamnus caroliniana) Fringe tree (Chionanthus virginica)

88. North-facing, wooded, limestone slopes along Pomme de
Terre River, three miles northeast of Elkland, Hickory County.

Dominant Trees

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.
glaucum) maxima)

Dominant Understory

Bladdernut (Staphyka trifolia) Pawpaw (Asimina triloba)

Other Understory Associates

Prickly ash (Zanthoxylum americanum) Flowering dogwood (Cornus florida)
Bittersweet (Celastrus scandens) ~Burningb'ash(Euonymusatropurpureus)

Dominant Herbaceous Non-Grass Plants

Tick trefoil (Desmodium acuminatum) Tall bellflower (Campanula americana)
Joe-Pye weed (Eupatorium purpureum) Lopseed (Phryma Leptostachya)
Agrimony (Agrimonia rostellata)

Dominant Grass

Brachyelytrum erectum

89. Northeast-facing, wooded, limestone slopes at base of bluffs
along Gasconade River, sec. 8, T. 36 N., R. 12 W., five miles north-
west of Waynesville, Pulaski County.

Dominant Trees

Black maple (Acer nigrum) Linden (Tilia glabra)

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.
glaucum) maxima)

Dominant Understory

Alternate-leaved dogwood (Cornus alter- Bladdernut (Staphyka trifolia)
nifolia)

Other Understory Associates

Arrow-wood (Viburnum molle) Dogwood (Cornus florida)

Ironwood (Carpinus caroliniana)

90. North-facing, shaley, limestone slopes along Deepwater
Creek, sec. 20, T. 40 N., R. 29 W., one and one-half miles southwest
of Spruce, Bates County.

452 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Trees

Northern red oak (Quercus borealis var. Black oak (Quercus velutina)
maxima) White oak (Quercus alba)

Dominant Understory

Rough-leaved dogwood (Cornus asperi- Bladdernut (Staphylea trifolia)
folia) Pawpaw (Asimina triloba)

91. Vertical, northeast-facing, limestone bluffs of Hall's Bluff,
south of Davidson's Blue Spring, along Black River, sec. 4, T. 27
N., R. 4 E., four miles southeast of Mill Spring, Wayne County.

Dominant Shrubs

Poison ivy (Toxicodendron radicans) Hydrangea (Hydrangea arborescens)

Virginia creeper (Parthenocissus quin- Ninebark (Physocarpus opulifolius var.
quefolia) intermedius)

Dominant Herbaceous Non-Grass Plants

Alum root (Heuchera parviflora var. Rock cress (Arabis laevigata)
Rugelii) Cliffbrake (Pellaea atropurpurea)

Bulblet bladder fern (Cystopteris bulbi- Gray polypody (Polypodium polypodi-
fera) oides var. Michauxianum)

Liverleaf (Hepatica acutiloba)

Dominant Grasses

Wild chess (Bromus purgans) Melic grass (Melica mutica)

Bottlebrush ( Hystrix patula)

92. North-facing, limestone slopes along Current River, sec. 3,
T. 24 N., R. 1 E., five miles northeast of Bennett, Ripley County.

Dominant Trees

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.

glaucum) maxima)

Butternut (Juglans cinerea)

Dominant Understory

American bladdernut (Staphylea tri- Ironwood (Carpinus caroliniana)
folia) Dogwood (Cornus florida)

Dominant Grasses and Grass-like Plants

Wild chess (Bromus purgans) Sedge (Carex eburnea)

93. West-facing, limestone slopes along Coldwater Creek, sec. 15,
T. 35 N., R. 8 E., five miles northeast of Womack, Sainte Genevieve
County.

Dominant Trees

Sugar maple (Acer saccharum var. Chinquapin oak (Quercus Muhlenbergii)
glaucum)

Dominant Understory
Redbud (Cercis canadensis)

94. Lower portion of north-facing, wooded, limestone slopes
along spring branch of Montauk Spring, Montauk State Park, near
Montauk, Dent County.

STUDIES OF THE VEGETATION OF MISSOURI — I 453

Dominant Tree

White oak (Quercus alba)

Other Tree Associates

Butternut (Juglans cinerea) Chinquapin oak (Quercus Muhlenbergii)

Linden (Tilia glabra)

Dominant Understory

Pawpaw (Asimina triloba) Spicebush (Benzoin aestivale)

Alternate-leaved dogwood (Cornus alter- Running strawberry-bush (Euonymus

nifolia) obovatus)

Bladdernut (Staphylea trifolia)

95. Upper portion of north-facing, wooded, limestone slopes
along spring branch of Montauk Spring, Montauk State Park, near
Montauk, Dent County.

Dominant Trees

Northern red oak (Quercus borealis var. Sugar maple (Acer saccharum var.
maxima) glaucum)

Other Tree Associates

White ash (Fraxinus americana) Slippery elm (Ulmusfulva)

Dominant Understory

Ironwood (Carpinus caroliniana) Flowering dogwood (Cornus florida)

Understory Associate

Hop hornbeam (Ostrya virginiana)

96. East-facing upper portion of wooded limestone slopes along
Hazel Creek, sec. 14, T. 36 N., R. 1 W., one mile southwest of Palmer,
Washington County.

Dominant Trees

Sugar maple (Acer saccharum var. Linden (Tilia glabra)

glaucum) Butternut (Juglans cinerea)

Chinquapin oak (Quercus Muhlenbergii)

Dominant Understory

Bladdernut (Staphylea trifolia)

Other Understory Associates

Rough-leaved dogwood (Cornus asperi- Flowering dogwood (Cornus florida)
folia)

97. Lower portion of east-facing limestone slopes along Hazel
Creek, sec. 14, T. 36 N., R. 1 W., one mile southwest of Palmer,
Washington County.

Dominant Tree

American elm ( Ulmus americana)

Dominant Understory

Pawpaw (Asimina triloba) Spicebush (Benzoin aestivale)

454 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

98. East-facing limestone slopes along Fourche a Renault Creek,
sec. 15, T. 37 N., R. 1 E., one and one-half miles north of Shirley,
Washington County.

Dominant Trees

Sugar maple (Acer saccharum var. Linden (Tilia glabra)
glaucum) Chinquapin oak (Quercus Muhlenbergii)

Other Tree Associates

Slippery elm (Ulmus fulva) Red mulberry (Moms rubra)

Butternut (Juglans cinerea) Winged elm (Ulmus alata)

Red cedar (Juniperus virginiana)

Dominant Understory

Hop hornbeam (Ostrya virginiana) Ironwood (Carpinus caroliniana)

Chittim-wood (Bumelia lanuginosa) Redbud (Cercis canadensis)

99. Intermediate zone of east-facing limestone slopes between
upper, drier, cedar-clad slopes and lower, richer soil slopes along
Fourche a Renault Creek, sec. 15, T. 37 N., R. 1 E., one and one-half
miles north of Shirley, Washington County.

Dominant Trees

Sugar maple (Acer saccharum var. Linden (Tilia glabra)

glaucum) Chinquapin oak (Quercus Muhlenbergii)

Schneck's red oak (Quercus Shumardii White ash (Fraxinus americana)

var. Schneckii) Winged elm (Ulmus alata)

Dominant Understory

Rough-leaved dogwood (Cornus asperi- Buck brush (Symphoricarpos orbicu-

folia) latus)

Hop hornbeam (Ostrya virginiana) Ninebark (Physocarpus opulifolius var.

Redbud (Cercis canadensis) intermedius)
Smooth sumac (Rhus glabra)

100. Upland slopes following along gravel bar of Little Fourche
a Renault Creek, sec. 2, T. 37 N., R. 1 E., two miles southeast of
Floyd, Washington County.

Dominant Trees

White oak (Quercus alba) Mockernut hickory (Carya tomenlosa)

Northern red oak (Quercus borealis var. Bitternut hickory (Carya cordiformis)
maxima) Black walnut (Juglans nigra)

Dominant Understory

Redbud (Cercis canadensis)

101. North-facing, wooded, limestone slopes along Big Creek,
sec. 18, T. 31 N., R. 2 W., three miles northeast of Rat, Shannon
County.

Dominant Trees

Sugar maple (Acer saccharum var. Linden (Tilia glabra)

glaucum) Butternut (Juglans cinerea)

STUDIES OF THE VEGETATION OF MISSOURI — I 455

Dominant Understory

Eastern witch hazel (Hamamelis virgini- Running strawberry-bush (Euonymus

ana) obovatus)

Spicebush (Benzoin aestivale) Flowering dogwood (Cornus florida)

Ohio buckeye (Aesculus glabra f . pallida) Bladdernut (Staphylea trifolia)

102. Acid, wooded valley floor along headwaters of branch south
of Enough, Iron County.

Dominant Trees

White oak (Quercus alba) Northern red oak (Quercus borealis var.

Black oak (Quercus velutina) maxima)

Other Tree Associates

Red maple (Acer rubrum) Honey locust (Gleditsia triacanthos)

Sycamore (Platanus occidentalis) Winged elm (Ulmus alata)

Butternut (Juglans cinerea) Sassafras (Sassafras albidum var. mplle)

Mockernut hickory (Carya tomentosa) Bitternut hickory (Carya cordiformis)

Dominant Understory

Smooth alder (Alnus rugosa) Hazelnut (Corylus americana)

Ironwood (Carpinus caroliniana) Black haw (Viburnum rufidulum)

Ozark witch hazel (Hamamelis vernalis) Hop hornbeam (Oslrya virginiana)

Flowering dogwood (Cornus florida) Bristly greenbrier (Smilax hispida)

Juneberry, serviceberry (Amelanchier Ward's willow (Salix longipes var.

canadensis) Wardii)

Redbud (Cercis canadensis)

103. Valley floor of shallow ravine along headwaters of Sinking
Creek, sec. 10, T. 32 N., R. 3 W., three miles northwest of Bunker,
in Dent County.

Dominant Trees

Sycamore (Platanus occidentalis) Bitternut hickory (Carya cordiformis)

Other Tree Associates

Slippery elm (Ulmus fulva) Black gum (Nyssa sylvatica var. caro-

Northern red oak (Quercus borealis var. liniana)

maxima) Shumard's red oak (Quercus Shumardii)
White oak (Quercus alba)

Dominant Understory

Buck brush (Symphoricarpos orbicu- Hazelnut (Corylus americanus)
latus)

Other Understory Associates

Redbud (Cercis canadensis) Smooth alder (Alnus rugosa)

Ironwood (Carpinus caroliniana) Swamp dogwood (Cornus obliqua)

Flowering dogwood (Cornus florida) Ward's willow (Salix longipes var.

Pawpaw (Asimina triloba) Wardii)
Black haw (Viburnum rufidulum)

104. Sandy, alluvial woods along Marble Creek opposite lime-
stone bluffs, sec. 21, T. 32 N., R. 5 E., two and one-half miles south-
west of French Mills, Madison County.

456 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Trees

Red cedar (Juniperus virginiana) Red maple (Acer rubrum)

Southern yellow pine (Pinus echinatd) Sour gum (Nyssa sylvatica var. caro-

Sugar maple (Acer saccharum var. liniana)

glaucum) Chinquapin oak (Quercus Muhlenbergii)
White oak (Quercus alba)

Dominant Understory

Leatherwood (Dirca palustris) Indian cherry (Rhamnus caroliniana

Chittim-wood (Bumelia lanuginosa) var. mollis)

Ironwood (Carpinus caroliniana)

III. QUERCUS-CARYA ASSOCIATION
(2) Quercus marilandica-Quercus stellata Associes

105. Upland, level, wooded ridge top between highway No. 32
and Dillard, Crawford County.

Dominant Trees

Post oak (Quercus stellata) Black jack oak (Quercus marilandica)

Other Tree Associates

White oak (Quercus alba) Black oak (Quercus velutina)

106. West-facing, upland, granite glade at Granite Bend, sec. 10,
T. 27 N., R. 4 E., five miles northwest of Williamsville, Wayne
County.

Dominant Trees

Post oak (Quercus stellata) Ozark pignut hickory (Carya Buckleyi

Black jack oak (Quercus marilandica) var. arkansana)

Winged elm (Ulmus alata)

Dominant Herbaceous Non-Grass Plants

Crotonopsis elliptica Rock pink (Talinum calycinum)

False aloe (Agave virginica)

Dominant Grasses and Grass-like Plants

Wild oats grass (Danthonia spicata) Slender fescue grass (Festuca octoflora)

Panic grass (Panicum linearifolium) Cyperus filiculmis var. macilentus

(3) Quercus velutina-Carya Buckleyi var. arkansana Associes

107. Upper valley wooded portion of Stilwell Hollow, in Low Gap
Game Refuge, sec. 12, T. 32 N., R. 1 E., southeast of Black U. S.
Lookout, four miles southwest of Black, Reynolds County.

Dominant Trees

Mockernut hickory (Carya tomentosa) Black oak (Quercus velutina)
White oak (Quercus alba) Shagbark hickory (Carya ovata)

Dominant Understory

Fragrant sumac (Rhus aromatica)

STUDIES OF THE VEGETATION OF MISSOURI — I 457

108. Upland oak-hickory woods in grazing plot southwest of
Bunker CCC camp F-7, sec. 3, T. 33 N., R. 3 W., five miles north-
west of Bunker, in Dent County.

Dominant Trees

White oak (Quercus alba) Black gum (Nyssa sylvatica var. caro-
Post oak (Quercus stellata) liniana)

Black oak (Quercus velutina) Mockernut hickory (Carya tomentosa)

Black jack oak (Quercus marilandica) Bitternut hickory (Carya cordiformis)

Ozark pignut hickory (Carya Buckleyi Shagbark hickory (Carya ovata)
var. arkansana)

Dominant Understory

Low-bush huckleberry (Vaccinium vacil- Smooth sumac (Rhus glabra)
lans var. crinitum)

Dominant Grasses and Grass-like Plants

Wild oats grass (Danthonia spicata) Panic grass (Panicum dichotomum)
Panic grass (Panicum lanuginosum var. Panic grass (Panicum lanuginosum var.

fasciculatum) Lindheimeri)

Panic grass (Panicum linearifolium) Blue-stem (Andropogon scoparius var.

Panic grass (Panicum Boscii) genuinus)
Sedge (Carex cephalophora)

109. Level, upland, oak woods two miles northwest of Bunker
CCC camp F-7, sec. 32, T. 33 N., R. 3 W., seven miles northwest
of Bunker, in Dent County.

Dominant Trees

White oak (Quercus alba) Black oak (Quercus velutina)

Post oak (Quercus stellata)

Other Tree Associates

Ozark pignut hickory (Carya Buckleyi Sassafras (Sassafras albidum var. molle)

var. arkansana) Shagbark hickory (Carya ovata)

Mockernut hickory (Carya tomentosa)

Dominant Understory

Low-bush huckleberry (Vaccinium vacil- Hazelnut (Corylus americana)
lans var. crinitum) Flowering dogwood (Cornus florida)

Dominant Herbaceous Non-Grass Plant

Bush clover (Lespedeza procumbens)

Dominant Grasses and Grass-like Plants

Wild oats grass (Danthonia spicata) Sedge (Carex artitecta)

Sedge (Carex retroflexa)

110. Upland cherty woods, sec. 7, T. 25 N., R. 4 E., 10 miles
southeast of Ellsinore, Carter County.

Dominant Trees

Scarlet oak (Quercus coccinea) Post oak (Quercus stellata)

Other Tree Associates

Black oak (Quercus velutina) Ozark pignut hickory (Carya Buckleyi

Black jack oak (Quercus marilandica) var. arkansana)

White oak (Quercus alba)

458 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Understory

Low-bush huckleberry (Vaccinium vacil- Dwarf sumac (Rhus Copallina)
lans var. crinitum)

Understory Associate

Sassafras (Sassafras albidum var. molle)

Dominant Herbaceous Non-Grass Plants

Cinquefoil (Potentilla simplex) Bush clover (Lespedeza procumbens)

Goat's rue (Tephrosia virginiana) Pussy toes (Antennaria plantaginifolia)

Dittany (Cunila origanoides)

Herbaceous Non- Grass Associate

Forked chickweed (Paronychia fastigiata var. typica)

Dominant Grasses and Grass-like Plants

Sedge (Carex hirsutella) Wild oats grass (Danthonia spicata)

Grass Associate

Hair grass (Agrostis hyemalis)

111. Upland oak-hickory woods, sec. 20 and 21, T. 26 N., R. 7 E.,
three miles north of Rombauer, Butler County.

Dominant Trees

Scarlet oak (Quercus coccinea) Black jack oak (Quercus marilandica)

Black oak (Quercus velutina) Shagbark hickory (Carya ovata)

Post oak (Quercus stellata) Mockernut hickory (Carya Buckleyi var.
Southern red oak (Quercus falcata) arkansana)

Winged elm ( Ulmus alata)

Dominant Understory

Fragrant sumac (Rhus aromatica) Sassafras (Sassafras albidum var. molle)

Dwarf sumac (Rhus Copallina)

Dominant Herbaceous Non-Grass Plants

Pussy toes (Antennaria plantaginifolia) Dittany (Cunila origanoides)

Cinquefoil (Potentilla simplex) Pinweed (Lechea villosa)

Goat's rue (Tephrosia virginiana) Bush clover (Lespedeza repens)

Pencil flower (Stylosanthes biflora var. Bergamot (Monarda Bradburiana)

hispidissima) Bastard toadflax (Comandra Richard-
Sensitive pea (Cassia nictitans) siana)
Field goldenrod (Solidago nemoralis) Wild aster (Aster patens)
Bush clover (Lespedeza procumbens) Brown-eyed Susan (Rudbeckia hirta)
Tick trefoil (Desmodium obtusum)

Dominant Grasses and Grass-like Plants

Wild oats grass (Danthonia spicata) Sphenopholis (Sphenopholis nitida)

Beard grass (Andropogon virginicus var. Paspalum (Paspalum ciliatifolium var.
genuinus) Muhlenbergii)

112. Upper, cherty slopes above limestone bluffs along Hazel
Creek, sec. 14, T. 36 N., R. 1 W., one mile southwest of Palmer,
Washington County.

Dominant Trees

Black oak (Quercus velutina) Northern red oak (Quercus borealis var.

White oak (Quercus alba) maxima)

STUDIES OF THE VEGETATION OF MISSOURI — I 459

Other Tree Associates

Black gum (Nyssa sylvatica var. caro- Red maple (Acer rubrum)
liniana) Sassafras (Sassafras albidum var. molle)

113. West-facing, porphyritic trachyte slopes of Johnson Moun-
tain, sec. 24, T. 35 N., R. 1 E., two miles south of Peoria, Iron County.

Dominant Trees

Ozark pignut hickory (Carya Buckleyi Shagbark hickory (Carya ovata)

var. arkansana) Black jack oak (Quercus marilandica)

Post oak (Quercus stellata)

Dominant Understory

Fragrant sumac (Rhus aromatica)

Other Understory Associates

Flowering dogwood (Cornus florida) High-bush huckleberry (Vaccinium

Low-bush huckleberry (Vaccinium vacil- stamineum)
lans var. crinitum)

114. Porphyritic trachyte outcrops on top of Johnson Mountain,
sec. 24, T. 35 N., R. 1 E., two miles south of Peoria, Iron County.

Dominant Trees

Northern red oak (Quercus borealis var. Ozark pignut hickory (Carya Buckleyi

maxima) var. arkansana)

Post oak (Quercus stellata)

Dominant Understory

Winged sumac (Rhus Copallina) Fragrant sumac (Rhus aromatica)

Smooth sumac (Rhus glabra)

Dominant Grasses and Grass-like Plants

Big blue-stem (Andropogon provincialis) Panic grass (Panicum lanuginosum var.
Fescue grass (Festuca octoflora var. fasciculatum)
tenella) Sedge (Carex artitecta)

115. Upland, level, oak woods near Dillard, Crawford County.

Dominant Trees

Scarlet oak (Quercus coccinea) Black jack oak (Quercus marilandica)

Other Tree Associates

White oak (Quercus alba) Black oak (Quercus velutina)

Dominant Understory

Low-bush huckleberry (Vaccinium vacillans var. crinitum)

116. Cherty upper slope south of Black, sec. 33, T. 33 N., R. 1 E.,
Reynolds County.

Dominant Trees

Black oak (Quercus velutina) Mockernut hickory (Carya tomentosa)

White oak (Quercus alba) Shagbark hickory (Carya ovata)

Scarlet oak (Quercus coccinea) Red mulberry (Morus rubra)

460 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Understory

Low-bush huckleberry (V actinium Smooth sumac (Rhus glabra)

vacillans var. crinitum) Sassafras (Sassafras albidum var. molle)

Winged sumac (Rhus Copallina) Bristly greenbrier (Smilax his-pida)

Fragrant sumac (Rhus aromatica) Dewberry (Rubus flagellaris)

117. Upper south-facing portion near top of Little Pilot Knob,
sec. 36, T. 38 N., R. 1 W., three miles northwest of Floyd, Washington
County.

Dominant Trees

Ozark pignut hickory (Carya Buckleyi Shingle oak (Quercus imbricaria)

var. arkansana) Black oak (Quercus velutina)

Northern red oak (Quercus borealis var. White oak (Quercus alba)

maxima) Post oak (Quercus stellata)

Dominant Understory

High-bush huckleberry (Vaccinium Low-bush huckleberry (Vaccinium
stamineum var. neglectum) vacillans var. crinitum)

118. Porphyritic trachyte soil on top of Little Pilot Knob, sec. 36,
T. 38 N., R. 1 W., three miles northwest of Floyd, Washington
County.

Dominant Trees

White oak (Quercus alba) Post oak (Quercus stellata)

Other Tree Associates

Ozark pignut hickory (Carya Buckleyi Shagbark hickory (Carya ovata)
var. arkansana)

Dominant Understory

Smooth sumac (Rhus glabra)

119. Upper east-facing portion near top of Little Pilot Knob,
sec. 36, T. 38 N., R. 1 W., three miles northwest of Floyd, Washington
County.

Dominant Trees

Northern red oak (Quercus borealis var. White oak (Quercus alba)

maxima) Black oak (Quercus velutina)

Shagbark hickory (Carya ovata)

Dominant Understory

Juneberry, serviceberry (Amelanchier New Jersey tea (Ceanothus americanus)
canadensis) Flowering dogwood (Cornus florida)

120. Upland oak-hickory woods, sec. 20 and 21, T. 26 N., R. 7 E.,
three miles northwest of Rombauer, Butler County.

Dominant Trees

Black oak (Quercus velutina) Shingle oak (Quercus imbricaria)

White oak (Quercus alba) Ozark hickory (Carya Buckleyi var.
Black jack oak (Quercus marilandica) arkansana)

Scarlet oak (Quercus coccinea) Shagbark hickory (Carya ovata)

STUDIES OF THE VEGETATION OF MISSOURI — I 461

Dominant Understory
Farkleberry (Vaccinium arboreum var. glaucescens)

121. South-facing wooded slopes by headwaters of Meramec
River along dry bed of river, sec. 14, T. 33 N., R. 4 W., two miles
southeast of Max, Dent County.

Dominant Trees

Post oak (Quercus stellata) Black jack oak (Quercus marilandica)

Ozark pignut hickory (Carya Buckleyi Shagbark hickory (Carya ovata)
var. arkansana) Mockernut hickory (Carya tomentosa)

122. North-facing wooded slopes by headwaters of Meramec
River along dry bed of river, sec. 14, T. 33 N., R. 4 W., two miles
southeast of Max, Dent County.

Dominant Trees

Northern red oak (Quercus borealis var. White oak (Quercus alba)
maxima) Black oak (Quercus velutina)

IV. QUERCUS-PINUS ECHINATA ASSOCIATION

(2b) Quercus marilandica-Quercus stellata-Pinus echinata

Associes

123. Woods off North Fork Hollow, near Cane Creek, sec. 8,
T. 26 N., R. 4 E., six miles east of Ellsinore, south of Upalika, near
Wayne County line, in Butler County.

Dominant Tree

Southern yellow pine (Pinus echinata)

Subdominant Trees
Post oak (Quercus stellata) Black jack oak (Quercus marilandica)

Chief Understory

White oak (Quercus alba f. latiloba) Ozark hickory (Carya Buckleyi var.

Fragrant sumac (Rhus aromatica) arkansana)

Winged sumac (Rhus Copallina) Mockernut hickory (Carya tomentosa)

Winged elm (Ulmus alala) Shingle oak (Quercus imbricaria)

Low-bush huckleberry (Vaccinium Shagbark hickory (Carya ovata)
vacillans var. crinitum)

Dominant Herbaceous Non-Grass Plants

Pussy toes (Antennaria plantaginifolia) Bush clover (Lespedeza repens)
Bush clover (Lespedeza procumbens)

Dominant Grass

Wild oats grass (Danthonia spicata)

123a. Pine stand on southeast-facing portion of porphyritic
trachyte slopes of Little Pilot Knob, sec. 36, T. 38 N., R. 1 W., three
miles northwest of Floyd, Washington County.

462 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Tree
Southern yellow pine (Pinus echinata')

Tree Associate

Ozark pignut hickory (Carya Buckleyi var. arkansana)

Dominant Understory

Low-bush huckleberry (Vacdnium vacillans var. crinitum)

Dominant Herbaceous Non-Grass Plants

Dittany (Cunila origanoides) Pussy toes (Antennaria plantaginifolia)

Aster (Aster patens)

Dominant Grasses and Grass-like Plants

Panic grass (Panicum lanuginosum var. Sedge (Carex artitecta)

fasciculatum) Wild oats grass (Danthonia spicata)

Sedge (Carex hirsutella)

(2c) Mixed Oak-Hickory-Pine Associes

124. Upper wooded ravine slopes of Stilwell Hollow, in Low
Gap Game Refuge, sec. 12, T. 32 N., R. 1 E., southeast of Black U. S.
Lookout, four miles southwest of Black, Reynolds County.

Dominant Trees

Black oak (Quercus velutina) Scarlet oak (Quercus coccinea)

Other Tree Associates

Ozark pignut hickory (Carya Buckleyi Southern yellow pine (Pinus echinata)
var. arkansana) Slippery elm (Ulmus fulva)

Dominant Grass

Wild chess (Bromus purgans)

125. Upland cherty ridge near Cornwall, Madison County.

Dominant Trees

Scarlet oak (Quercus coccinea) White oak (Quercus alba)

Black oak (Quercus velutina) Southern yellow pine (Pinus echinata)

Other Tree Associates

Post oak (Quercus stellata) Ozark pignut hickory (Carya Buckleyi

Mockernut hickory (Carya tomentosa) var. arkansana)

126. Upper, cherty, west-facing slopes along Eleven Points
River, sec. 11, T. 24 N., R. 3 W., one mile southeast of Turner's Mill,
four miles southwest of Wilderness, Oregon County.

Dominant Trees

Southern yellow pine (Pinus echinata) Mockernut hickory (Carya tomentosa)
White oak (Quercus alba) Shagbark hickory (Carya ovata)

Black oak (Quercus velutina) Black jack oak (Quercus marilandica)

Red maple (Acer rubrum)

Dominant Understory

Dogwood (Cornus florida) Fragrant sumac (Rhus aromatica)

STUDIES OF THE VEGETATION OF MISSOURI — I 463

Dominant Herbaceous Non-Grass Plants

Goat's rue (Tephrosia virginiana) Dittany (Cunila origanoides)

Dominant Grass

Panic grass (Panicum dichotomum)

127. Upper, west-facing, cherty slopes, sec. 7, T. 25 N., R. 4 E.,
10 miles southeast of Ellsinore, Carter County.

Dominant Tree

Southern yellow pine (Pinus echinata)

Other Tree Associates

White oak (Quercus alba) Ozark pignut hickory (Carya Buckkyi

Scarlet oak (Quercus coccinea) var. arkansana)

Post oak (Quercus stellata)

Dominant Understory

Low-bush huckleberry (Vaccinium vacillans var. crinitum)

Dominant Herbaceous Non-Grass Plants

Dittany (Cunila origanoides) Goat's rue (Tephrosia virginiana)

Bush clover (Lespedeza hirta) •

Dominant Grasses and Grass-like Plants

Sedge (Carex hirsutella) Panic grass (Panicum dichotomum)

128. Upland, cherty, sandstone area, sec. 22, T. 26 N., R. 3 E.,
three miles south of Ellsinore, Carter County.

Dominant Trees

Southern yellow pine (Pinus echinata) Black jack oak (Quercus marilandica)

Scarlet oak (Quercus coccinea) Ozark hickory (Carya Buckleyi var.

Spanish oak (Quercus falcata) arkansana)
Black oak (Quercus velutina)

Dominant Understory

Winged sumac (Rhus Copallina) Sassafras (Sassafras albidum var. molk)

Dominant Herbaceous Non-Grass Plants

Dittany (Cunila origanoides) Catgut, goat's rue (Tephrosia virginiana)

Pussy toes (Antennaria plantaginifolia) Bush clover (Lespedeza repens)
Bush clover (Lespedeza procumbens)

129. Cherty, steep exposures above bluffs along Huzzah Creek,
two miles southeast of Davisville, Crawford County.

Dominant Trees

Post oak (Quercus stellata) Northern red oak (Quercus borealis var.

Black oak (Quercus velutina) maxima)

Winged elm ( Ulmus alata)

Tree Associate

Southern yellow pine (Pinus echinata)

464 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Understory

Low-bush huckleberry (Vaccinium vacillans var. crinitum)

Understory Associate

Hop hornbeam (Ostrya virginiana)

130. Cherty slopes along Stilwell Hollow, Low Gap Game Refuge,
southeast of Black U. S. Lookout, sec. 12, T. 32 N., R. 1 E., four
miles southwest of Black, Reynolds County.

Dominant Trees
Scarlet oak (Quercus coccinea) Black oak (Quercus velutina)

Other Tree Associates

Southern yellow pine (Pinus echinata) Shagbark hickory (Carya ovata)
Mockernut hickory (Carya tomentosa) Post oak (Quercus stellata)

131. Cherty ridge top above limestone slopes along Fourche a
Renault Creek, sec. 15, T. 37 N., R. 1 E., one and one-half miles
north of Shirley, Washington County.

Dominant Tjees

Black jack oak (Quercus marilandica) Black walnut (Juglans nigra)

Post oak (Quercus stellata) White oak (Quercus alba)

Shagbark hickory (Carya ovata) Southern yellow pine (Pinus echinata)

Scarlet oak (Quercus coccinea) Bitternut hickory (Carya cordiformis)

Dominant Understory

High-bush huckleberry (Vaccinium Flowering dogwood (Cornus florida)
stamineum var. negkctum) Winged sumac (Rhus Copallina)

132. East-facing cherty slopes above limestone bluffs along
Brazil Creek, sec. 28, T. 38 N., R. 1 W., six miles northeast of Berry-
man, Washington County.

Dominant Trees

Southern yellow pine (Pinus echinata) Shagbark hickory (Carya ovata)
Post oak (Quercus stellata) Black oak (Quercus velutina)

Black jack oak (Quercus marilandica) Black walnut (Juglans nigra)

Dominant Understory
High-bush huckleberry (Vaccinium stamineum var. neglectum)

133. Cherty slopes along Coldwater Creek, sec. 33, T. 35 N.,
R. 8 E., three miles northeast of Womack, St. Francois County.

Dominant Trees

White oak (Quercus alba) Scarlet oak (Quercus coccinea)

Black oak (Quercus velutina) Southern yellow pine (Pinus echinata)

Other Tree Associates

Post oak (Quercus stellata) Northern red oak (Quercus borealis var.

Black jack oak (Quercus marilandica) maxima)

Shagbark hickory (Carya ovata)

STUDIES OF THE VEGETATION OF MISSOURI — I 465

Dominant Understory

Red maple (Acer rubrum) Farkleberry (Vaccinium arboreum var.

Flowering dogwood (Cornus florida) glaucescens)

Azalea (Rhododendron nudiflorum var. Low-bush huckleberry (Vaccinium

roseum) vacillans var. crinitum)

Sassafras (Sassafras albidum var. molle)

134. Upland, level, wooded tract, just east of Bardley CCC camp,
sec. 23, T. 24 N., R. 2 W., one and one-half miles west of Pine, in
Ripley County.

Dominant Trees
Black jack oak (Quercus marilandica) Shagbark hickory (Carya ovata)

Other Tree Associates

Southern yellow pine (Pinus echinata) Southern red oak (Quercus falcata)
Scarlet oak (Quercus coccinea) Ozark pignut hickory (Carya Buckleyi

Post oak (Quercus stellata) var. arkansana)

White oak (Quercus alba)

Dominant Understory

Dwarf sumac (Rhus Copallina) New Jersey tea (Ceanothus americanus)

Other Understory Associates

Low-bush huckleberry (Vaccinium Smooth sumac (Rhus glabra)
vacillans var. crinitum) Dogwood (Cornus florida)

High-bush huckleberry (Vaccinium Sour gum (Nyssa sylvatica var. caro-
stamineum var. neglectum) liniana)

Summer grape (Vitis aestivalis)

Dominant Herbaceous Non-Grass Plant

Bush clover (Lespedeza procumbens)

Dominant Grasses

Panic grass (Panicum dichotomum) Panic grass (Panicum Boscii)

Panic grass (Panicum sphaerocarpon)

135. Ridge top in sec. 30, T. 26 N., R. 4 E., six miles southeast
of Ellsinore, in Carter County.

Dominant Trees

Scarlet oak (Quercus coccinea) Southern yellow pine (Pinus echinata)

Other Tree Associates

Black oak (Quercus velutina) Pignut hickory (Carya ovalis var.

Spanish oak (Quercus falcata) obovalis)

White oak (Quercus alba) Sour gum (Nyssa sylvatica var. caro-

Post oak (Quercus stellata) liniana)

Black jack oak (Quercus marilandica)

Chief Understory

Winged sumac (Rhus Copallina) Catbrier (Smilax glauca var. leuro-

Farkleberry (Vaccinium arboreum var. phylla)
glaucescens)

Other Understory Associates

Staghorn sumac (Rhus glabra) Flowering dogwood (Cornus florida)

Fragrant sumac (Rhus aromatica) Persimmon (Diospyros virginiana)

466 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Herbaceous Non-Grass Plants

Pinweed (Lechea tenuifolia) Horsemint (Monarda Bradburiana)

Cinquefoil (Potentilla simplex var. Pennyroyal (Hedeoma pulegioides)
typica) Dittany (Cunila origanoides)

Other Herbaceous Associates

Bush clover (Lespedeza procumbens) Bush clover (Lespedeza repens)
Bush clover (Lespedeza virginica) Forked chickweed (Paronychia fasti-
Sweet everlasting (Gnaphalium dbtusi- giata var. typica)
folium) Fleabane (Erigeron ramosus)

Dominant Grasses and Grass-like Plants

Panic grass (Panicum lanuginosum var. Panic grass (Panicum dichotomum)
fasciculatum) Rush (Juncus macer)

Other Grasses and Grass-like Associates

Poverty grass (Aristida dichotoma) Sedge (Carex hirsutella)

136. West-facing, cherty, limestone slopes along Fourche a
Renault Creek, sec. 15, T. 37 N., R. 1 E., one and one-half miles north
of Shirley, Washington County.

Dominant Trees

Southern yellow pine (Pinus echinata) Scarlet oak (Quercus coccinea)
White oak (Quercus alba)

Other Tree Associates

Black walnut (Juglans nigra) Black gum (Nyssa sylvatica var. caro-
Sassafras (Sassafras albidum var. molle) liniana)
Shingle oak (Quercus imbricaria)

Dominant Understory

Low-bush huckleberry (Vaccinium vacillans var. crinitum)

Other Understory Associates

Farkleberry (Vaccinium arboreum var. Winged sumac (Rhus Copallina)
glaucescens)

Dominant Herbaceous Non- Grass Plants

Dittany (Cunila origanoides) Pussy toes (Antennaria plantaginifolia)

Dominant Grasses and Grass-like Plants

Blue-stem (Andropogon scoparius var. Wild oats grass (Danthonia spicata)

genuinus) Broom sedge (Andropogon virginicus

Sedge (Carex artitecta) var. genuinus)

137. South and southeast-facing, porphyritic trachyte slopes of
Johnson Mountain, sec. 24, T. 35 N., R. 1 E., two miles south
of Peoria, Iron County.

Dominant Trees

White oak (Quercus alba) Ozark pignut hickory (Carya Buckleyi

Scarlet oak (Quercus coccinea) var. arkansana)

Black oak (Quercus velutina) Northern red oak (Quercus borealis var.

Southern yellow pine (Pinus echinata) maxima)

Black gum (Nyssa sylvatica var. caro- Sassafras (Sassafras albidum var. molle)

liniana) Shagbark hickory (Carya ovata)

Post oak (Quercus stellata) Red maple (Acer rubrum)

STUDIES OF THE VEGETATION OF MISSOURI — I 467

Dominant Understory

Flowering dogwood (Cornus florida) Low-bush huckleberry (Vaccinium

Farkleberry (Vaccinium arboreum var. vacillans var. crinitum)
glaucescens) Winged sumac (Rhus Copallina)

V. QUERCUS ALBA-ACER RUBRUM ASSOCIATION

(1) Cornus florida-Nyssa sylvatica Associes

138. Gravelly draw in upland, level woods around Little Scotia
Pond, sec. 25-26, T. 33 N., R. 4 W., four miles east of Turtle, Dent
County.

Dominant Trees

Red maple (Acer rubrum) Flowering dogwood (Cornus florida)

Dominant Understory

Farkleberry (Vaccinium arboreum var. Shrubby St. John's-wort (Hypericum
glaucescens) prolificum)

(2) Acer rubrum-Carya tomentosa Associes

139. Cherty upper slopes overlying cherty acid soil between
Saunders and Pine Ridge, Christian County.

Dominant Trees

Sour gum (Nyssa sylvatica var. caro- Red maple (Acer rubrum)
liniana)

Tree Associate
Southern yellow pine (Pinus echinata)

Dominant Understory

Low-bush huckleberry (Vaccinium vacillans var. crinitum)

Other Understory Associates

Wild rose (Rosa Carolina) Wild rose (Rosa subserrulata)

Dominant Herbaceous Non-Grass Plant

Flowering spurge (Euphorbia corollata)

Dominant Grass

Broom sedge (Andropogon virginicus var. genuinus)

140. North-facing, porphyritic trachyte slopes of Johnson Moun-
tain, sec. 24, T. 35 N., R. 1 E., two miles south of Peoria, Iron County.

Dominant Trees

White oak (Quercus alba) Black gum (Nyssa sylvatica var. caro-
Northern red oak (Quercus borealis var. liniana)

maxima) Shagbark hickory (Carya ovata)

Red maple (Acer rubrum) Mockernut hickory (Carya tomentosa)

Dominant Understory

Flowering dogwood (Cornus florida)

468 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

141. Lower cherty slopes of Stilwell Hollow, Low Gap Game
Refuge, southeast of Black U. S. Lookout, sec. 12, T. 32 N., R. 1 E.,
four miles southwest of Black, Reynolds County.

Dominant Trees

Southern yellow pine (Pinus echinata) Red maple (Acer rubrum)

Dominant Understory

Hop hornbeam (Ostrya virginiana) Smooth sumac (Rhus glabra)

Serviceberry, Juneberry (Amelanchier Winged sumac (Rhus Copallina)

canadensis) High-bush huckleberry (Vaccinium

Low-bush huckleberry (Vaccinium stamineum var. neglectum)

vacillans var. crinitum) Farkleberry (Vaccinium arboreum var.

Wild honeysuckle (Lonicera flava) glaucescens)

142. Slight draw on upper south-facing portion of porphyritic
trachyte slopes of Little Pilot Knob, sec. 36, T. 38 N., R. 1 W., three
miles northwest of Floyd, Washington County.

Dominant Trees

Red maple (Acer rubrum) Sassafras (Sassafras albidum var. molk)

Sour gum (Nyssa sylvatica var. caro- Southern yellow pine (Pinus echinata)

liniana) White oak (Quercus alba)
Black oak (Quercus velutina)

(3) Quercus alba— Acer rubrum Associes

143. Draw in ravine in pine-oak woods south of Bunker CCC
camp F-7, sec. 3, T. 32 N., R. 3 W., five miles northwest of Bunker,
near Reynolds County line, in Dent County.

Dominant Trees

Red maple (Acer rubrum) Ozark pignut hickory (Carya Buckleyi

Sour gum (Nyssa sylvatica var. caro- var. arkansana)

liniana) White oak (Quercus alba)

Dominant Understory

Hazelnut (Corylus americana) Sassafras (Sassafras albidum var. molk)

Dwarf sumac (Rhus Copallina)

144. Cherty, sandstone, shallow slopes along headwaters of Big
Creek, sec. 10, T. 31 N., R. 3 W., south of Bunker U. S. Lookout,
five miles southwest of Bunker, in Shannon County.

Dominant Trees

Mockernut hickory (Carya tomentosa) Northern red oak (Quercus borealis var.
White oak (Quercus alba) maxima)

Other Tree Associates

Scarlet oak (Quercus coccinea) Red maple (Acer rubrum)

Southern yellow pine (Pinus echinata) Sour gum (Nyssa sylvatica var. caro-

Sassafras (Sassafras albidum var. molle) liniana)

Winged elm (Ulmus alata)

STUDIES OF THE VEGETATION OF MISSOURI — I 469

Dominant Understory

Ozark witch hazel (Hamamelis vernalis) Buck brush (Symphoricarpos orbiculatus)

Other Understory Associates

Flowering dogwood (Cornus florida) Juneberry, serviceberry (Amelanchier

Low-bush huckleberry (V actinium canadensis)
vacillans var. crinitum) Black haw (Viburnum rufidulum)

145. Cherty, upper, south-facing slopes along Big Barren Creek,
sec. 4, T. 25 N., R. 1 W., three miles southwest of Eastwood, 10
miles northwest of Bennett, Carter County.

Dominant Trees

White oak (Quercus alba) Sour gum (Nyssa sylvatica var. caro-

Southern yellow pine (Pinus echinata) liniana)
Red maple (Acer rubrum)

Other Tree Associates

Post oak (Quercus stellata) Scarlet oak (Quercus coccinea)

Dominant Understory

Shadbush (Amelanchier canadensis)

Other Understory Associates

Fragrant sumac (Rhus aromatica) High-bush huckleberry (Vaccinium

Dwarf sumac (Rhus Copallina) stamineum var. negkctum)

Hazelnut (Corylus americana) Farkleberry (Vaccinium arboreum var.

Low-bush huckleberry (Vaccinium glaucescens)
vacillans var. crinitum)

Dominant Herbaceous Non-Grass Plants

Dittany (Cunila origanoides) Tick trefoil (Desmodium rotundifolium)

Bracken fern (Pteridium latiusculum var. Goat's rue (Tephrosia virginiana)

pseudocaudatum) Christmas fern (Polystichum acrosti-

Bush clover (Lespedeza hirta) choides)

Tick trefoil (Desmodium marilandicum)

Dominant Grass

Fescue grass (Festuca paradoxa)

146. Cherty slopes along headwaters of Big Creek, sec. 5, T. 31
N., R. 3 W., two miles south of Melton, Shannon County.

Dominant Trees

White oak (Quercus alba) Scarlet oak (Quercus coccinea)

Other Tree Associates

Southern yellow pine (Pinus echinata) Red maple (Acer rubrum)
Sour gum (Nyssa sylvatica var. caro- Black oak (Quercus velutina)
liniana)

147. Wooded, north-facing, cherty slopes along Eleven Points
River between mouth of White's Creek and Bliss Spring, sec. 7,
T. 24 N., R. 2 W., two and one-half miles southwest of Wilderness,
Oregon County.

470 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Trees

Northern red oak (Quercus borealis var. White oak (Quercus alba)

maxima) Scarlet oak (Quercus coccinea)

Bitternut hickory (Carya cordiformis)

Other Tree Associates

Sassafras (Sassafras albidum var. molle) Persimmon (Diospyros virginiana)

Red maple (Acer rubrum) Sour gum (Nyssa sylvatica var. caro-

Mockernut hickory (Carya tomentosa) liniana)

Dogwood (Cornus florida) Slippery elm (Ulmus fulva)

Red mulberry (Moms rubra)

Dominant Understory

Pink azalea (Rhododendron nudiflorum Hop hornbeam (Ostrya virginiana)

var. roseum) Smooth sumac (Rhus glabra)

Hazelnut (Corylus americana) Rough-leaved dogwood (Cornus asperi-

Redbud (Cercis canadensis) folia)

Shadbush (Amelanchier canadensis) Low-bush huckleberry (Vaccinium

Pawpaw (Asimina triloba) vacillans var. crinitum)

148. Cherty slopes south of Rockhouse Spring, sec. 33, T. 23 N.,
R. 26 W., two miles west of Lohmer, Barry County.

Dominant Trees

Black oak (Quercus velutina) Sour gum (Nyssa sylvatica var. ca.ro-

White oak (Quercus alba) liniana)

Red maple (Acer rubrum) Chinquapin (Castanea ozarkensis)

Other Tree Associates

Northern red oak (Quercus borealis var. Mockernut hickory (Carya tomentosa)
maxima)

Dominant Understory

Flowering dogwood (Cornus florida) Low-bush huckleberry (Vaccinium

Juneberry, serviceberry (Amelanchier vacillans var. crinitum)
canadensis)

Other Understory Associates

High-bush huckleberry (Vaccinium Hazelnut (Corylus americana)
stamineum)

Dominant Herbaceous Non-Grass Plants

Bracken (Pteridium latiusculum var. Pussy toes (Antennaria plantaginifolia)

pseudocaudatum) Horsemint (Monarda Bradburiana)

Dittany (Cunila origanoides)

Other Herbaceous Non-Grass Plant Associates

Dwarf dandelion (Krigia biflora) Bush clover (Lespedeza procumbens)

Bedstraw (Galium arkansanum) Lead plant (Amorpha canescens f. gla-

Rue anemone (Anemonella thalictroides) brata)

Tick trefoil (Desmodium rotundifolium) Bracted wild indigo (Baptisia leuco-

Tick trefoil (Desmodium Dillenii) phaea)

Bush clover (Lespedeza hirta) Flowering spurge (Euphorbia corollata)

Black cohosh (Cimicifuga racemosa)

Dominant Grass-like Plant

Sedge (Carex cephalophora)

STUDIES OF THE VEGETATION OF MISSOURI — I 471

149. Northwest-facing, cherty, sandstone, wooded slopes along
Little Sinking Creek, sec. 26, T. 32 N., R. 3 W., two miles southwest
of Bunker, Dent County.

Dominant Trees

Northern red oak (Quercus borealis var. Red maple (Acer rubrum)
maxima)

Other Tree Associates

Black gum (Nyssa sylvalica var. caro- White oak (Quercus alba)
liniana)

Dominant Understory

Low-bush huckleberry (Vaccinium Hop hornbeam (Ostrya virginiana)
vacillans var. crinitum) Ironwood (Carpinus caroliniana)

150. Cherty ravine floor and adjacent lower slopes in Stilwell
Hollow, Low Gap Game Refuge, southeast of Black U. S. Lookout,
sec. 12, T. 32 N., R. 1 E., four miles southwest of Black, Reynolds
County.

Dominant Trees

White oak (Quercus alba) Northern red oak (Quercus borealis var.

Black gum (Nyssa sylvatica var. caro- maxima)
liniana)

Other Tree Associates

Shagbark hickory (Carya ovata) Sassafras (Sassafras albidum var. molle)

Black oak (Quercus velutina) Butternut (Juglans cinerea)

Scarlet oak (Quercus coccinea) Sycamore (Platanus occidentalis)

Southern yellow pine (Pinus echinata) Slippery elm (Ulmus fulva)

Red maple (Acer rubrum) Red mulberry (Morus rubra)

Dominant Understory

Hazelnut (Corylus americana) Low-bush huckleberry (Vaccinium

Serviceberry, Juneberry (Amelanchier vacillans var. crinitum)

canadensis) Fragrant sumac (Rhus aromatica)

Flowering dogwood (Cornus florida) Spicebush (Benzoin aestivale)

Ironwood (Carpinus caroliniana) Elderberry (Sambucus canadensis)

Black haw (Viburnum rufidulum) Alternate-leaved dogwood (Cornus alter-

Ozark witch hazel (Hamamelis vernalis) nifolia)

Smooth alder (Alnus rugosa) Climbing rose (Rosa setigera var. tomen-

tosa)

Dominant Grasses and Grass-like Plants

Panic grass (Panicum Boscii) Panic grass (Panicum lanuginosum var.

Panic grass (Panicum xalapense) Lindheimeri)

Panic grass (Panicum dichotomum) Sedge (Carex cephalophora)

151. Woods along headwaters of Big Creek, sec. 5, T. 31 N.,
R. 3 W., two miles south of Melton, Shannon County.

Dominant Trees

Northern red oak (Quercus borealis var. White oak (Quercus alba)
maxima)

472 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. IX

Other Tree Associates

Mockernut hickory (Carya tomentosa) Sassafras (Sassafras albidum var. molle)

Red maple (Acer rubrum) Winged elm (Ulmus alata)

Scarlet oak (Quercus coccinea) Southern yellow pine (Pinus echinata)

Dominant Understory
Ozark witch hazel ( Hamamelis vernalis; along stream bed)

Other Understory Associates

Juneberry, serviceberry (Amelanchier Flowering dogwood (Cornus florida)

canadensis) Black haw (Viburnum rufidulum)

Buck brush (Symphoricarpos orbiculatus) Winter grape (Vitis cinerea)

152. Cherty quartzitic slopes of ravine tributary to Huzzah
Creek, two miles southeast of Davisville, Crawford County.

Dominant Trees

White oak (Quercus alba) Sour gum (Nyssa sylvatica var. caro-

Northern red oak (Quercus borealis var. liniana)
maxima}

Other Tree Associates
Winged elm (Ulmus alata) Red mulberry (Morus rubra)

Dominant Understory
Flowering dogwood (Cornus florida)

Other Understory Associates

Sassafras (Sassafras albidum var. molle) Hop hornbeam (Ostrya virginiana)

Dominant Herbaceous Non-Grass Plants

Dittany (Cunila origanoides) Bedstraw (Galium arkansanum)

Honeysuckle (Lonicera flava)

153. Gravelly wash in draw in upland off North Fork Hollow,
near Cane Creek, sec. 8, T. 26 N., R. 4 E., six miles east of Ellsinore,
south of Upalika, near Wayne County line, in Butler County.

Dominant Trees

Sour gum (Nyssa sylvatica var. caro- Red maple (Acer rubrum)
liniana) White oak (Quercus alba)

Other Tree Associates

Mulberry (Morus rubra) American elm (Ulmus americana)

Shingle oak (Quercus imbricaria)

Chief Understory

Hazelnut (Corylus americana) Prairie willow (Salix humilis)

Flowering dogwood (Cornus florida) Shrubby St. John's-wort (Hypericum

Black haw (Viburnum rufidulum) prolificum)
Possum haw (Ilex decidua)

154. Small gravel wash along slight gradient of ravine slope,
near Cane Creek, sec. 8, T. 26 N., R. 4 E., six miles east of Ellsinore,
south of Upalika, near Wayne County line, in Butler County.

STUDIES OF THE VEGETATION OF MISSOURI — I 473

Dominant Trees

White oak (Quercus alba) Sour gum (Nyssa sylvatica var. coro-

Red maple (Acer rubrum) liniana)

Other Tree Associates

Shingle oak (Quercus imbricaria) American elm (Ulmus americana)

Dominant Understory

Possum haw, deciduous holly (Ilex Hazelnut (Corylus americana)

decidua) Black haw (Viburnum rufidulum)

Dogwood (Cornus florida) Red mulberry (Moms rubra)

Prairie willow (Salix humilis)

155. Lamotte sandstone bluffs along Pickle Creek, Pickle Springs,
sec. 20, T. 36 N., R. 7 E., six miles northwest of Coffman, Sainte
Genevieve County.

Dominant Trees

Scarlet oak (Quercus coccinea) Black oak (Quercus velutina)

White oak (Quercus alba) Red maple (Acer rubrum)

Southern yellow pine (Pinus echinata)

Dominant Understory

Farkleberry (Vaccinium arboreum var. Greenbrier (Smilax glauca var. leuro-
glaucescens) phylla)

156. Lamotte sandstone ravine along Pickle Creek, Pickle
Springs, sec. 29, T. 36 N., R. 7 E., six miles northwest of Coffman,
Sainte Genevieve County.

Dominant Trees

Red maple (Acer rubrum) White oak (Quercus alba)

Sour gum (Nyssa sylvatica var. caro- Black oak (Quercus velutina)

liniana) Sassafras (Sassafras albidum var. molle)

Northern red oak (Quercus borealis var. Bitternut hickory (Carya cordiformis)

maxima) Mockernut hickory (Carya tomentosa)

Dominant Shrubs

Flowering dogwood (Cornus florida) Greenbrier (Smilax glauca var. leuro-

Juneberry, serviceberry (Amelanchier phylla)

canadensis) Smooth alder (Alnus rugosa)

Hop hornbeam (Ostrya virginiana) Low-bush huckleberry (Vaccinium

Ironwood (Carpinus caroliniana) vacillans var. crinitum)

157. Ravine bottom woods along Coldwater Creek, sec. 33,
T. 35 N., R. 8 E., three miles northeast of Womack, St. Francois
County.

Dominant Trees

White oak (Quercus alba) Sycamore (Platanus occidentalis)

Other Tree Associates

Red maple (Acer rubrum) American elm (Ulmus americana)

Sugar maple (Acer saccharum) Sour gum (Nyssa sylvatica)

Shagbark hickory (Carya ovata)

474 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. IX

Dominant Understory

Smooth alder (Alnus rugosa) Hazelnut (Corylus americana)

Other Understory Associates

Spicebush (Benzoin aestivale) Ward's willow (Salix longipes var.

Dogwood (Cornus florida) Wardii)

158. Wooded floor of ravine tributary to headwaters of Brazil
Creek, sec. 28, T. 38 N., R. 1 W., six miles northeast of Berryman,
Washington County.

Dominant Trees

White oak (Quercus alba) Bitternut hickory (Carya cordiformis)

Scarlet oak (Quercus coccinea) Sassafras (Sassafras albidum var. molle)

Mockernut hickory (Carya tomentosa) Red maple (Acer rubrum)

Dominant Understory

Flowering dogwood (Cornus florida) Hazelnut (Corylus americana)

159. Slight draw on upper east-facing portion of porphyritic
trachyte slopes of Little Pilot Knob, sec. 36, T. 38 N., R. 1 W., three
miles northwest of Floyd, Washington County.

Dominant Trees

Southern yellow pine (Pinus echinata) Northern red oak (Quercus borealis var.
Sassafras (Sassafras albidum var. molle) maxima)
Sour gum (Nyssa sylvatica var. caro- Black oak (Quercus velutina)
liniana) Persimmon (Diospyros virginiana)

Dominant Understory

Flowering dogwood (Cornus florida)

160. Cherty slopes along Little Fourche a Renault Creek, sec. 2,
T. 37 N., R. 1 E., two miles southeast of Floyd, Washington County.

Dominant Trees

Southern yellow pine (Pinus echinata) White oak (Quercus alba)
Red maple (Acer rubrum)

Dominant Understory

Flowering dogwood (Cornus florida)

SUBCLIMAX STAGE C

Ulmus americana Subclimax

161. Upper, cherty portion of northeast-facing limestone slopes
along St. Francis River, just west of Greenville, Wayne County.

Dominant Trees

American elm (Ulmus americana) Slippery elm (Ulmus fulva)

Dominant Understory

Shadbush, serviceberry (Amelanchier Winged elm (Ulmus alata)
canadensis) Hop hornbeam (Ostrya virginiana)

STUDIES OF THE VEGETATION OF MISSOURI — I 475

MIXED ASSOCIATIONS WHERE ONE HAS PENETRATED
AND INTERGRADED INTO ANOTHER

162. Porphyritic trachyte "shut-in" along Marble Creek, sec. 21,
T. 32N.,R.5E., two and one-half miles southwest of French Mills,
Madison County.

Dominant Trees

Winged elm (Ulmus alata) White oak (Quercus alba)

Scarlet oak (Quercus coccinea)

Other Tree Associates

Southern yellow pine (Pinus echinata) River birch (Betula nigra)

Red cedar (Juniperus virginiana) Sour gum (Nyssa sylvatica var. co.ro-

Northern red oak (Quercus borealis var. liniana)

maxima) American elm (Ulmus americana)

Mockernut hickory (Carya tomentosa) Red maple (Acer rubrum)

Dominant Understory

Ozark witch hazel (Hamamelis vernalis) Sycamore (Platanus occidentalis)

Smooth alder (Alnus rugosa) Greenbrier (Smilax glauca var. leuro-

Swamp dogwood (Cornus obliqua) phylla)
Greenbrier (Smilax Bona-nox)

163. Porphyritic trachyte slopes along St. Francis River, sec. 10,
T. 32 N., R. 5 E., two miles north of French Mills, Madison County.

Dominant Trees

Sugar maple (Acer saccharum var. Northern red oak (Quercus borealis var.

glaucum) maxima)

Red maple (Acer rubrum) Linden (Tilia glabra)

Sour gum (Nyssa sylvatica var. caro- Red mulberry (Morus rubra)

liniana) White oak (Quercus alba)

Sweet gum (Liquidambar Styraciflua) Winged elm (Ulmus alata)

Dominant Understory

Flowering dogwood (Cornus florida) Hop hornbeam (Ostrya virginiana)

Pawpaw (Asimina triloba) Bristly greenbrier (Smilax kispida)

Ironwood (Carpinus caroliniana) Greenbrier (Smilax Bona-nox)

164. Porphyritic trachyte slopes along "shut-in" of Telleck
Branch, sec. 23, T. 35 N., R. 1 E., three and one-half miles northwest
of Shepherd, Iron County.

Dominant Trees

Southern yellow pine (Pinus echinata) Northern red oak (Quercus borealis var.

Red cedar (Juniperus virginiana) maxima)

Sugar maple (Acer saccharum var. Black gum (Nyssa sylvatica var. caro-

glaucum) liniana)

White oak (Quercus alba) Red maple (Acer rubrum)

Winged elm ( Ulmus alata) Persimmon (Diospyros virginiana)

Dominant Understory
Indian cherry (Rhamnus caroliniana) THE LIBRARY OF THF

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