Vegetation of Limestone and Dolomite Glades in the Ozarks and Midwest Regions of the United States

9 Pages·2000·8.4 MB·

by  Jerry M Baskin

A
VEGETATION
OF LIMESTONE
Jerry M. Baskin' and Carol Baskin'-'
C.
AND
DOLOMITE GLADES
IN
THE OZARKS AND MIDWEST
REGIONS OF THE
UNITED
STATES^
Abstf^act
Lileralure on the vegetation of limestone and dolomite (cedar) glades in the Ozarks of Missouri ami Arkansas and
in the niidwestern United States (Illinois, Indiana, Ohio, Wisconsin) is reviewed. Dominant plants in these glades are
C,^ [)eretniial prairie grasses, of which little bluestem {Schizachyrium scoparium (Micli\.) Nash) the most important.
is
Without removal of invading woody plants by fire or other means, succession in these rocky, calcareous openings
is lo
Q
forest. Tliey differ from cedar glades in the southeastern United States, which are dominated by annual grasses
(primarily Sporobolus vaginijloriis (Torr. ex Gray) ^Xood) and do not require management or natural disturbances
to
We
maintain them. suggest that the anthro[)ogenic, prairie-grass-dominaled openings in the Ozarks and Midwest be
called xeric limestone (or dolomite) f)rairies and that the term cedar glades be used for an edaphic climax dominated
by Cj sunnncr annual grasses in natural openings on limestone or dolomite bedrock.
Key
words: annual Sporobolus species, Midwestern (U.S.A.) limestone glades, Ozark glades, Schizachyrium scopar-
woody
ium, phuit invasion, xeric limestone
prairies
Although the climatic climax vegetation in the terms
eastern United States is forest (Shreve, 1917; Braun, souri). Midwest (Illinois, Indiana, Ohio, Wisconsin),
1964) persisting plant com- and Southeast (see 12.2 (map), 208 Baskin
fig. p. in
&
^ous angiosperms and/ Baskin, 1999) has been mean
interpreted
to that
or cryptogams occur in where bedrock is ex- cedar glade vegetation is the same in the three re-
posed and/or soil depth in most places is too shallow gions (cf. Curtis, 1959; Kuchler, 1964). However,
support Examples
to trees or shrubs. of these edaph- cedar glades of the southeastern United States are
ic climax rock outcrop communities include the mid- dominated by C4 summer annual
grasses, primarily
K
Appalachian
shale barrens 1951;
(Piatt,
uginifl
1983; Braunschweig et al., 1999), granite outcrops Midwest are dominated by C^ perennial grasses,
&
of the southeastern piedmont (Costing Anderson, primarily Schizachyrium scoparium (Baskin
et
al.,
1943 Keever
et al., 1951; Bur- 1994, 1995). In a recent review of the literature on
1964; Palmer, 1970; Shure, 1999), cedar glades of the southeastern United Bas-
States,
and
the cec (limestone) glades of the Central kin and Baskin concluded
1 (1999) that: Sporo-
(1)
(Nashville) Basin in Tennessee (()uarterman 1950; bolus vaginiflorus is the most important species in
&
Baskin Baskin, 1985; Somers et al., 1986; Drew, this limestone/dolomite rock outcrop vegetation
and
1991) of the southeastern UUnniitteedd SSttaatteess iinn geeenn-- type; and neither Schizachyrium scoparium nor
(2)
1999) any other perennial grass species an important
is
The
terms "cedar glades" and "limestone component No
(or do- of the vegetation. such review of
lomite) glades" are also used to describe grass/forb- quantitative information has been published on
the
dominated openings over shallow limestone or do- vegetation of calcareous glades in the Ozarks and/
lomite soil on ridgetops and side slopes in the or Midwest.
Ozarks
(Steyermark, 1940; Erickson et al., 1942; Thus, the purpose of this paper is to review the
& &
Kucera Ladd
Martin, 1957; Nelson, 1982; literature on the vegetation of limestone and dolo-
&
Nelson Ladd,
1983; Nelson, 1985; Fig. 1) and mite glades in the Ozarks and Midwest. In partic-
Midwest
Curtis, 1959; Kurz, 1981; Aldrich
(e.g., et ular, quantitative and/or qualitative evidence
&
Heikens
al., 1982; Robertson, 1995). Thus, the presented that, indeed, Schizachyrium scoparium
is
We
thank Kayri Havens and several anonymous reviewers comments
for their very helpful on various
' drafts of the
manuscript.
^School
of Biologieal Sciences, University of Kentucky, Lexington, Kentucky 40506-0225,
U.S.A.
Department Agronomy,
of University of Kentucky, Lexington, Kentucky 40546-0091,
U.S.
Ann. Missouri Bot. Card. 286-294.
87: 2000.
Volume Number &
2 Baskin Baskin 287
87,
2000 Limestone Glade
Vegetation
Figure Top, Valley View Glade, Jefferson County, Missouri. Bottom, a glade in the Hercules Glades Wilderness
1.
Area, Taney County, Missouri. Photographs taken by Carol C. Baskin on 24 July 1996 (top) and 26 July 1993 (bottom).
288
Annals
the
of
Garden
Missouri Botanical
the dominant plant species of these calcareous DC. Berchemia scandens K. Koch a
tallii (Hill) is
common
glades and that annual grasses are relatively un- woody vine in this pioneer stage in the
important. In addition, this review contrasts the cal- southwestern Ozarks.
A
careous glades of the Ozarks and Midwest with second serai stage begins when Rhus aroma-
those of the southeastern United States. tica, Diospyros virginiana, Juniperus virginiana,
and woody
other plants invade these limestone
(or
The Ozarks dolomite) glades. Juniperus virginiana typically be-
comes
the most conspicuous plant, especially on
Steyermark (1940; see also Steyermark, 1959) eroded slopes and "bald knobs," rming what
fo
recognized five climax vegetation associations (sen- Steyermark referred to as "cedar glades," "red ce-
su the polyclimax theory) of the Missouri Ozarks dar glades," or "red cedar balds." These terms in-
and qualitatively described stages in primary sue- dicate scattered redcedar trees in a matrix of her-
He
cession leading to each of them. also recognized baceous vegetation dominated by prairie grasses, in
several subclimaxes in these seres. However, these particular A, gerardii, virgatum^ and scopar-
P. 5.
presumed successional sequences probably repre- ium. In the White River region of southwestern
sent vegetation sequences along environmental gra- Missouri, virginiana and/or virginiana may
/. /).
dients that are unrelated to per se invade limestone glades and become the dominant
The open limestone and dolomite glades of the species. Sassafras alhidum Nees may co-oc-
(Nutt.)
Missouri Ozarks are the first two stages of what cur with D. virginiana as a pioneer woody species.
Steyermark (1940) presented as a six-stage sere be- Other investigators also have found that prairie
ginning on bare, rocky slopes and ending with a grasses are the dominants of cedar glades in the
sugar maple-white oak climatic climax: Boute- Ozarks of Mi Kucera and Martin (1957) re-
(1)
i.
loua curtipendula (Michx.) Torr-Rudbeckia mis- ported Schizachyrium scoparium as the dominant
51%
souriensis Engelm.; Rhus aromatica Ait -Dios- species (85% frequency, of herbaceous cover)
(2)
pyros virginiana L-Juniperus virginiana L. (with a in cedar glades in the Ozarks of southwestern Mis-
redaeceeadar subclimax persisting on eroded limestone souri. Locally dominant grasses included A. gerar-
slopes and knobs); Bumelia lanuginosa (Michx.) B. curtipendula, virgatum, Sorghastrum nu-
(3) dii, P.
-Viburnum Ulmus
Vi^^xs rufidulum Raf.; alata tans, Sporoholus heterolepis (A. Gray) A. Gray,
(4)
Michx -Rhamnus caroliniana Walt.; (5) Quercus Tridens flavus (L.) A. Hitchc, and the C^ summer
muhlenbergii Engelm. -Fraxinus americana L. (with annual Sporoholus neglectus. Hedyotis nigricans was
Q, muhlenbergii forming a subclimax on southern the most important herbaceous dicot, Rhus aro-
and western exposures); and Acer saccharum matica the most common shrub, and Juniperus
(6) vir-
Marsh-Quercus alba L. Climax vegetation in the giniana the most common tree. Other typical woody
White may
River region of southwestern Missouri species included Bumelia lanuginosa, Cotinus
be Q. muhlenbergii-Cotinus obovatus Raf. Thus, the obovatus, Diospyros virginiana, Ilex decidua Walt.,
pioneer stage in this sere is a limestone (or dolo- and Rhamnus caroliniana Walt. Hicks's (1981) re-
mite) glade (sensu Steyermark, 1940: 392) ". suits of a vegetational analysis of seven stands of
. .
<
30%
with largely a component of prairie species. ." open cedar glades with woody plant canopy
. .
The dominants are Bouteloua curtipendula and cover and 2 Steyermark scheme
(stages 1 in s of
Rudbeckia
missouriensLs {B. curtipendula, Psorali- xerarch succession to a sugar maple-white oak cli-
dium
tenuiflorum (Pursh) Rydb., and Silphium la- max) in the Hercules Glades Wilderness Area in
ciniatum Andropogon and
L,, or gerardii Yii. B. cur- southwestern Missouri were quite similar to those
some
tipendula in parts of the Ozarks). In addition of Kucera and Martin Schizachyrium
(1957). sco-
to A. gerardii and B. curtipendula, other important parium was the dominant species, and H. nigricans,
Panicum
C4 grasses on these glades are: virgatum Rhus aromatica, and virginiana were the most
/.
L., Sorghastrum nutans (L.) Nash, Schizachyrium important herbaceous dicot, shrub, and tree, re-
and
scoparium, Sporoholus neglectus Nash. Impor- spectively (Table Schizachyrium scoparium had
1).
Agave
tant forbs include virginica (L.) Rose, Allium the highest percent importance value (%I.V.) in
all
Calam-
stellatum Roth., Aster oblongifolius Nutt., seven stands sampled by Hicks.
+
intha arkansana (Nutt.) Shinn., Dalea purpurea Hall (1955) determined frequency of herbs
m
Vent,, Echinacea pallida (Nutt.) Nutt., E. paradoxa seedlings and of sprouts of woody plants 0.27 or
(Norton) Britt. \ ax. paradoxa (an Ozark cedar glade less in height and the number of Juniperus virgi-
endemi(!), Hedyotis nigricans (Lam.) Fosb., Heli- niana seedlings/saplings an open dolomite glade
in
otropium tenellum Oenothera macro- Shaw
(Nutt.) Torr., in the Missouri Botanical Garden's Arbore-
carpa Nutt. subsp. macrocarpa, and Polytaenia nut- tum, near the northeastern border of the Ozark Pla-
Volume Number 2 Baskin & Baskin 289
87,
2000 Limestone Glade Vegetation
Table Average percent importance values (%I.V.) Table Continued.
1. 1.
for herbs, shrubs/vines, and trees in seven stands of open
%
cedar glades in the Ozarks of southwestern Missouri. Only
^
Species constancy %1.\
plants with a %1.V. of 1.0% are listed. Percent con-
stancy and %I.V. were calculated from data Hicks
in L.
J. Rhamnus 86
caroliniana Walt. 4.77
(1981).
100 4.43
Celtis tenuifolia Nutt.
Fraxinus americana L. 71 3.67
%
Cornas drummondii C. Meyer 57 1.96
%I.V
Species constancy
c
43
canadensis 1.54
ercis de. L.
Herbs Quercus prinoides Willd. 29 1.30
6 3.09
other species
Schizachyrium scoparium (Michx.)
100.09
Total
Nash 100 29.21
Hedyotis nigricans (I^m.) Fosb. 100 10.47
Nash 100 9.47
Sporobolus
neglectiis
Carex 100 7.37
sp.
Understory species with the highest frequen-
Rudbeckia missouriensis Engelm, 100 7.30 teau.
Panicum virgaium 100 6.40 cies were Sporobolus neglectus (100%), Carex
L.
Croton capitatus Michx./C monan- crawei Dewey (90%), Schizachyrium scoparium
thogynus Michx. 100 5.10 Euphorbia
(80%), Hedyotis nigricans (75%), corol-
Sorghastrum nutans Nash 100 3.81
(L.) lata Engelm. (60%), and Rudbeckia missouriensis
Andropogon 71 2.57
gerardii
Vil.
The number
(55%). total of juniper seedlings in the
ramo-
Tragia betonicifoUa Nult./7^
99.7%
glade was 6694, and of these were in the
100
sa Torr. 1.73
number
0.46-m or less height class. Average of ju-
Silphium terebinthinaceum Jacq. 71 1.51
niper seedlings per m^ was 0.84. Hall (1955: 177)
Sporobolus heterolepis (A. Gray) A.
stated, "The Glade a 'prairie' association with
Gray 86 is
1.44
[=5.
& Andropogon scoparium] and Rud-
scoparius
Palafoxia callosa (Nutt.) Torr.
A. Gray 71 1.41 beckia missouriensis contributing most to its aspect
&
Evolvulus nuttallianus Roemer and Andropogon scoparius and Sporobolus neglectus
57
Schultes 1.31 contributing most to cover." Hall concluded that
86
Heliotropium tenelluin (Nutl.) Torr. 1.31 dolomite glade was an edaphic subclimax, as
this
Oenothera macrocarpa Nutt. subsp
did Erickson et al. (1942) for the dolomite glades
86
macrocarpa 1.17
of the northeastern Ozarks in general.
86
Dalea purpurea Vent. 1.11
The and community ecology
flora qualitative of
104 other species 9.18
and
limestone dolomite glades (as well as that of
101.87
Total
sandstone, chert, and igneous glades) of the Mis-
SHRUBS/VllNES
Ozarks have been studied extensively by Nel-
souri
Rhus aromatica 100 58.77
Ait.
son and Ladd (1982, 1983) and Nelson (1985). Nel-
Symphoricarpos
orbiculatiis
and Ladd
son (1982: stated, "Characteristic
5)
Moench 86
9.71
dominant vascular plants of these [dolomite] glades
Koch 43
Berchemia scandens K. 7.24
(Hill)
Andropogon Bouteloua
include scoparius, curtipen-
Rosa Michx. 57 5.40
setigera
and Sorghastrum and limestone
dula, nutans^^; for
Andrachne phyllanthoides
(Nutl.)
J.
glades, "Dominant vascular plants on these glades
Coulter 14 5.34
Mimosa 29 4.99 are Andropogon scoparius and Bouteloua curtipen-
quadrivalvis L.
&
Vitis aestivalis Michx. 57 4.84 dula'' (Nelson Ladd, 1982: 5). Nelson (1985) list-
Smilax bona-nox L. 57 2.10 ed A. scoparius and B. curtipendula as the dominant
Parthenocissiis quinquefolia (L.) plants on limestone glades and A. scoparius, B. cur-
29
Planch. 1.37
and dominants
tipendula, Sporobolus heterolepis as
Kun-
Toxicodendron radicans
(L.)
on dolomite glades.
14 0.37
tze
Ver Hoef et (1993) quantitatively sampled the
al.
100.13
Total
32 on Eminence and Gasconade
vegetation of glades
Trees
dolomites in southeastern Missouri. Schizachyrium
100 41.17
Juniperus virginiana
L.
scoparium clearly was the most important species in
Cotinus obovatus Raf. 100 19.17
these rocky forest openings. This C4 perennial grass
86 12.15
Diospyros virginiana L.
100%
had a constancy of in glades on each of the
Bumella lanuginosa (Michx.) Pers 100 6.84
two geologic formations, and was in the highest
it
%
=
3L6) 32
cover class average cover in all
(4,
290
Annals
the
of
Garden
Missouri Botanical
10-100%
glades, cover class, geometric mean Table 2. The most important species in 20 limestone
i.e.,
=
31.6. Other important species on the dolomite glaJes studied by Logan (1992) in northwest Arkansas,
50%
glades were Andropogon gerardiU Calamintha ^"^y species with a presence of at least and/or an
ar-
abundance
UEch- ^^^'^^^ of at least 1.00 are included in the
kansarm, Carex meadii, Coreopsis lanceolata
P^^^^'^S^ F^^"^^ ^^^^^g^ abundance values
^'''- ^"^^
inaeea pallida, Fimbristylis pubemla (Michx.) M.
,T v^ere calculated from data in Loean (1992).
T7 T'
Van, l, var. pub,erul1a, tied1yoti's ni'gri'cans, Liatn's aspera
~
Panicum
Michx., cylindracea Michx.
Liatris virga-
j Average
%
Rudbeckia Silphium
turn, missouriensis, terebinthin-
abundance
aceum
Jacq., Smilax bona-nox L., Sorghastrum nu~ Species value^ presence
tarn, Sporobolus clandestine (Biehler) A. Hitchc,
Schizachyrium scoparium
and The most woody
5. vaginiflorus. important spe-
(Michx.) Nash 4.60 95
cies on these glades were Juniperus virginiana, Bu-
Junlperus virginiana 2.60 70
L.
melia lanuginosa, Diospyros virginiana, Pinus echin- Helianihus hirsuius Raf. 2.05 70
Andropogon
ata Miller, Quercus prinoides Willd., Q. stellata gerardii Vit. 2.00 55
Wangenh
Wangenh., and Ulmus Quercus stellata 1.95 60
alata.
UlmiLS alata Michx. 1.65 70
Keeland sampled
(1978) quantitatively the veg-
'?"^'^'" prinoides Willd. 1.40 60
etation of 13 calcareous glades in the Ozarks of
60
1.35
^
He
northwestern Arkansas. recognized four general ^".
J'^
^"f^"^". ^^"^""t^
r^'
OpuntiQ numijusa 60
(Kal.) Raf. 1.25
community types, wh, i. ch1 i. n an ap^p^arent successi. on- r
^^
tupaton• um t
^ , ali tissimum 1.25 45
L.
sequence and
al are: grass cedar; cedar;
(1) (2) (3)
30
Coreopsi, tinctoria Nutt. 1.15
cedar-hardwood; and hardwood. Keeland
(1978:
(4) DUhanthellum acuminatum
(Sw.)
a
12) described the grassland-cedar type as &
Gould Clark 1.15 40
t
grassy slope with a few scattered woody species." Fraxinus amerlcana 50
L. 1.15
Further, ". the typical Ozark cedar glade as Rueltia humilis Null. 1.15 60
is
it
. .
40
described by Hall (1955) and Kucera and Martin Lespedeza capitata Michx. 1.10
Sporobolus ex
and 99 vaginifloriis (Torr.
(1957) in Missouri Hite (1959) in Arkansas
Wood
Gray) 1.10 30
(Keeland 1978: For the representative stand of
'9)U, Jur^iJT-
...
.1-.
Dalea purpurea Vent. 1.10 55
1
descnbed
this type by Keeland, Juniperus vireini-
r. i,. .*,
^ Echi inacea , 40
„ pallida (NuU.) Nutt. 1.05
J
ana and Uuercus stellata were t1he most i• mportant
r
^ ^ i-
tupnj orma corolfl? at. a th-ngeli m. 1.00 55
overstory trees, Rhus glabra L. the most important
,^f^^ aromatica 1.00 55
Ait.
shrub, and Schizachyrium scoparium the most im-
Desmanthus
illinoemis (Michx.)
portant herb. Overstory tree basal area on this ce- MacMillan ex Robins. & Fern 1.00 35
dar glade was low, ca. 4.8m^/ha. Schizachyrium sco- Bumella lanuginosa (Michx.)
parium also was the most important species Pers. 0.90 50
in
50
limestone cedar glades quantitatively sampled by Celtis tenuifolia Nutt. 0.85
Logan =
(1992) in the Buffalo National River area in Abundance
scale from one
-* 1 to 5: 1 rare (only or
= =
the Ozarks of northwestern Arkansas (Table two individuals present); 2 occasional; 3 frequent; 4
2).
= =
connnon; and 5 abundant (widespread with high cov-
Skinner (1979) placed O.Ol-m^ and 0.1-m' quad-
a dominant or near-dominant).
er,
around Ozark
rats individual plants of the cedar
glade endemic Penstemon cobaea
Nutt. var. pur-
pureus Pennell, Centaurium texense (Griseb.) Fern., and 59 vs. 16 for Penstemon, Centaurium, and
and
Stenosiphon Heynh. The
linifalius (Nutt.) to deter- Stenosiphon, respectively. increase in percent
mine
species associates (as well as other site char- occurrence of S. neglectus with increase in size of
acteristics) of these three rare plant species in lime- quadrats was less dramatic: 46 58
vs. 32, vs. 33,
stone glades of the southwestern Missouri Ozarks. and 11 10 Penstemon, Centaurium, and
vs. for
Sporobolus neglectus had the highest percent oc- Stenosiphon, respectively (Skinner, 1979).
currence in quadrats of both sizes placed around In a dolomite glade in the Ozarks of southeastern
Penstemon and Centaurium, and Schizachyrium Missouri, two contiguous 7 X 7-m sample
plots
scoparium had
the highest percent occurrence in within a whole-glade sample unit of 100 contiguous
X
those of both sizes placed around Stenosiphon, 7 7-m plots dominated by Schizachyrium sco-
±
which grows deeper cm, mean parium and
in soil (15.7 6.1 classified as "glades" open glades)
(i.e.,
±
SD) than Penstemon (10.9 5.6 cm) or Cen- were divided into 200 70 X 70-cm plots (Ver Hoef
±
taurium (5.3 2.8 cm). Percent occurrence of S. et al., 1993). At this small scale of sampling, the
scoparium was considerably higher In the 0.1-m^ glade was divided into rocky glade, shallow
soil
<
than in the 0.01 -m^ quadrats: 22 vs. 5, 37 vs. 5, glade, glade, and deep soil glade zones. Sporobolus
Volume Number &
2 Baskin Baskin 291
87,
2000 Limestone Glade Vegetation
vaginiflorus was the dominant species in the rocky nutans^ Thus, annual Sporobolus can be the dom-
glade and shallow soil glade zones, and 5. scopar- inant grass on overgrazed Ozark glades.
ium in the glade and deep soil glade zones (Ver
Hoef et 1993). However, in two other dolomite The Midwest
al.,
glades southeastern Missouri sampled by Ver
in
Hoef In the Midwest, cedar or limestone glades have
et al. (1993), as described above, 5. vagini-
was not an important species. been reported in Wisconsin, Illinois, and Indiana.
florus
Schizachyrium scoparium was most prominent
Results of studies by Skinner (1979) and Ver the
Hoef made ground-layer species in what Curtis (1959) called
et al. (1993) support a statement by
cedar glades in Wisconsin, with a presence of
Kucera and
Martin (1957: 290) in their study of
100%
and an average frequency of 38%. Other
glades in the Ozarks of southwestern Missouri:
species in the ground layer with relatively high val-
[=
''Andropogon scoparius Schizachyrium scopar-
"^^ presence and average frequency were the
^^^
ium] was the principal dominant; Sporobolus neg-
C3 herbaceous perennial forbs ^^ui/e^ia canarfem^
lectus was generally frequent and locally abun-
Anenwne
cylindHca A. Gray, Dalea purpurea.
L.,
dant." Hall (1955), Hicks (1981), and Logan (1992)
Tradescantia ohioensis Raf., Euphorbia corollata,
also found that either Sporobolus neglectus or va-
5.
^^^^«"«
^^"^^^ Michx., Solidago nemoralis Dryan-
was
giniflorus a relatively important species in the
der, Antennaria neglecta E. Greene, Viola pedata
Ozark glades they studied. Nelson (1985) listed S.
and Scutellaria leonardu Epling; the C, peren-
L-,
neglectus as a characteristic plant of dolomite
^^^^^ Andropogon gerardu, Bouteloua hirsuta
"'^^
glades, but not of limestone glades, in the Ozarks
Lagasca, and Bouteloua curtipendula; and the C3
of Missouri. Thus, both quantitative and qualitative
1.1.
R
,1. ic perennial erass Koeleria pyramidata (Lam.)
r.1 • u
studies show that annual Sporobolus species can be
L^
i-irx
/^^
. i<
Beauv. (Curtis, 1959: table XVI-10, appendix,
p.
locally dominant within the larger prairie-like com-
573). The most important tree species was Junipe-
munity on limestone and dolomite glades in the
rus virginiana (Curtis, 1959: table XVI-11, appen-
As
Ozarks. such, cedar glades of the type described
dix, 574).
p.
&
for the southeastern United States (Baskin Bas-
80-
Schizachyrium scoparium had a frequency
of
kin, 1999) occur in microhabitats within xeric lime-
100%
in 30 of 32 limestone glades sampled by Kurz
stone prairies of the Ozarks.
(1981) in Illinois. Other important grasses were the
Schizachyrium scoparium, also the dominant
is
C4 perennials Bouteloua curtipendula, Sorghastrum
plant in Ozark glades in terms of dry mass produc-
nutans, and Sporobolus aspera (Michx.) Kunth. The
In the Ozarks of southwestern Missouri,
tion. total
CAM
important forbs were the leaf succulent Agave
annual herbage production on glades ungrazed
for
virginica\ the C3 perennials Aster oblongif
alius.
4 and 21 years was 312.2 and 241.5 g/m^, respec-
Echinacea
Brickellia eupatorioides Shinn., pal-
(L.)
v^
^' ° ^ ^ I ;• lida^ Euphorbia corollata, Hedyotis nigricans, and
duction by scoparium on these two exclosures
S.
and sum-
p;,^,^,^,^,-^ virginiana Benth.; the C3
(L.)
mean
term used by Buttery areas fenced
(the to off ^^^^
^^^ monanxhogynus
^^^^^^ Michx. Schiza-
was and
to prevent cattle grazing) 175.2 112.3 ^^^^^
g/
scoparium also was the most important spe-
m^
respectively, and by Sporobolus neglectus 24.7
^-^^ -^
limestone glades of southern
^j^^ Illinois
and g/m^
7.9 respectively. In the 4-year exclosure,
^y Heikens and Robertson
^i^^^-^^ (1995).
production by Andropogon gerardii and Sorghas-
Limestone glades have been reported from
five
trum nutans was 52.2 and 34.8 g/m^, respectively,
counties in southern Indiana (Aldrich et al., 1982;
and in the 21-year exclosure 29.2 and 74.7 g/m^
Bacone Homoya,
al, 1983; 1987; Maxwell,
et
Thus
respectively. the C^ perennial grasses sco-
5. 1987), and they obviously are dominated by peren-
84%
parium, and nutans produced
A, gerardii,
5. For example,
nial prairie grasses. referring to a
89%
and of the total herbage on the 4- and 21- Bacone
limestone glade Perry County,
in et al.
year exclosures, respectively (Buttery, 1960). Re-
(1983: 368) stated, "Indian grass {Sorghastrum nu-
garding herbage production on grazed glades, But-
bluestem {Andropogon and
tans), big gerardii)
little
tery (1960: 235) stated, "In 1956, as now, the bluestem scoparius) are common grasses." Re-
{A.
glades outside the exclosures were producing about Max-
ferring to two cedar glades in Clark County,
400 pounds of oven dry herbage per acre [44.9g/ well (1987: 413) stated, "Little bluestem dominates
^^ ^^ ^^ ^^_ '
m^], mostly the less desirable baldgrass [S. neglec- both glades as a xeric, bunch-grass surrounded by
tus] and black-eyed Susan {Rudbechia hirta L.) with patches of rocky pavement, 99
a scattering of the more desirable bluestem The Ozarks and Midwest limestone and dolomite
little
(Andropogon scoparius Michx.) and Indiangrass glades described above are similar vegetationally
[S. to
292
Annals
the
of
Garden
Missouri Botanical
Table Average percent frequency of native plant component
3. of the vegetation of limestone glades in
X
1-m
taxa in 147 1 quadrats in five stands of xeric prai- Indiana (Aldrich et al, 1982; Bacone et 1983;
al.,
Adams
rie vegetation on the Peebles dolomite in County,
Homoya,
1987; Maxwell,
1987).
Ohio. Only plants with a frequency of >:20% are listed
(calculated from data in Braun, 1928).
Woody
Plant Ozark and
Invasion into
% Midwest Glades
Species frequency
Schizachyrium scoparium (Michx.) Nash 79.4
14
Martin (1955: 106) stated that even the
f t
Euphorbia corollata Engelm. 53.6
thin-soiled areas the Ozarks of southwestern
[in
BlephiUa ciliata (L.) Benth. 50.8
Missouri] are being invaded by eastern redcedar
Lithospermum Lehm.
canescens (Michx.) 45.2
{Juniperus virginiana)^ winged elm {Ulmus
alata)^
Agave Rose
virginica 43.2
(L.)
and associated drought-tolerant Using 1938
trees."
Ruellia humilis (Nees) Lindau 41.8
USDA
and 1975 photographs showing
aerial lime-
Silphium
trifoliatum L. 38.2
and
Bouteloua curtipendala (Michx.) 32.4 stone dolomite glades in southwestern Missou-
Torr.
Kimmel
Thaspium sp. 32.0 and Probasco (1980) found a dramatic
ri,
Pycnanthemum 0—15%
flexuosum (Walt.) Britton, decrease in glade area with woody cover
&
Stems Poggenb. 29.8
and
a corresponding increase in glade area with
Serwcio plattensis Nutt. 29.4 50-100%
woody cover. Lowell and Astroth (1989)
Sorghastrum nutans Nash 29,0
(L.) USDA
used aerial photographs study natural
to
Comandra
umbellata 27.6
(L.) Nutt.
succession of glades to forest in the Hercules
Helianthus hirsutus Raf. 27.6
Glades Wilderness Area (southwestern Missouri)
Andropogon
gerardii 26.2
Vit.
from 1938 1986. They found "Even
to that those
Zizia aptera (A. Gray) Fern. 25.4
areas most favorable glades [Gasconade
for
Fragaria soil,
vesca L. 25.2
m
305-365
Prunella vulgaris L. 23.4 (a.m.s.l.), south or southwest slopes]
&
Delphinium exaltatum Ait. 23.2 continue to convert to forest" (Lowell Astroth,
Lobelia spicata Lam. 22.8 1989: 78). The area most favorable for glades and
pannda 2L4
Scutellaria Michx.
occupied by glades decreased from about 285 ha
Solidago nemoralis 20.0
Ait.
to about 200 ha (30%) via conversions of glade land
&
to forest (see 3 in Lowell Astroth, 1989).
fig.
Using 1955, 1966, and 1984 photographs,
aerial
what Braun (1928) called xeric prairies on Silurian Ver Hoef et (1993) also showed that the sizes of
al.
Adams
Dolomite in County in southern Ohio. Here, dolomite glades in the Ozarks of southeastern Mis-
prairie grasses, primarily Schizachyrium scoparium, souri had decreased due to woody plant invasion.
also are the dominant plants (Table Interesting- Kimmel and Probasco (1980) concluded that the
3).
Braun pointed out that although there some most important reason for the increase woody
ly, is in
overlap in species composition between the xeric plant cover on open glades between 1938 and 1975
Adams
prairies of County and the cedar glades of was that the U.S. Forest Service had not used
fire
manage
the Nashville Basin, the vegetation is different. to glade (cattle) range. Lowell and Astroth
Neither annual species oiSporobolus nor of other (1989: 78) thought that their study ". appears
to
. .
seem
grasses to be an important component of ce- support the theory held by some ecologists (USFS,
dar or limestone glades that have been sampled pers. comm.) that the glades are not a naturally
quantitatively in Wisconsin (Curtis, 1959) or lUi- occurring ecotype[?] which can maintain them-
&
nois (Kurz, 1981; Heikens Robertson, 1995). selves independent of and/or human interven-
fire
However, in one of the xeric prairies. Agave Ridge, tion." Likewise, Ver Hoef et (1993) attributed
al.
Adams
on Silurian dolomite in County, Ohio, sam- woody plant invasion into calcareous glades in
pled by Braun (1928), 5. vaginiflorus had a fre- southeastern Missouri to fire suppression. Fire fre-
68%
quency of in 25 1-m^ quadrats. The dominants quency was high in the Missouri Ozarks in preset-
&
of this prairie were Schizachyrium scoparium tlement times (Guyette McGinnis, and
1982),
fire
(100% and
frequency) Bouteloua curtipendula probably played an important role in the origin and
(84% frequency). Braun referred Agave Ridge maintenance of limestone and dolomite glades
to in
and one of the four prairies she sampled on Peebles that area.
Dolomite as being more xerophytic than the other According to Curtis (1959), cedar glades in Wis-
However, may
three. the average frequency of 5. vagin- consin: have originated by invasion of red-
(1)
iflorus on the five Peebles dolomite prairies was cedar into a dry prairie on sites protected from
fire;
only 14.8%. Neither is 5. vaginiflorus an important and (2) would in time succeed to oak forest.
Volume Number & 293
2 Baskin Baskin
87,
2000 Limestone Glade Vegetation
Heikens (1991) speculated that, in the absence of ture of the vegetation of the hmestone and dolomite
much
the perennial grass-dominated limestone glades in the Ozarks and Midwest more
fire, is
glades in southern Illinois would succeed to bar- similar to the xeric prairies on Silurian dolomite
Adams
rens ("savannas") and then to forest. described by Braun (1928) in County, Ohio,
By studying aerial photographs of limestone and to the xeric prairies on Mississippian limestone
glade areas in southern Indiana, Aldrich et al. described by Baskin et al. (1994) on and adjacent
(1982) and Bacone et al. (1983) determined that to the Kentucky Karst Plain, than it is to what tra-
these openings in the forest are decreasing in size. ditionally has been called cedar glades in the
Aldrich et (1982: 484) stated, "Aerial photo- southeastern United States. Kuchlers (1964) vege-
al.
graphs document the continuing shrinkage of these tation map of the United States shows the same
glades Harrison County], as they were nearly vegetation type for cedar glades in the Ozarks and
[in
Bacone middle Tennessee-northern Alabama. However,
double their present size in the 1940s." et in
(1983: 372) stated, 'The aerial photographs ample evidence is available in the literature to
al.
show a remarkable decrease in size [of the forest show that the glade vegetation of these two regions
we
openings in Perry County] in the last forty years is quite different. Thus, suggest that the rocky.
91
due encroachment woody anthropogenic, prairie-grass-dominated
of species. cal
to
,
Braun (1928: 425) thought the xeric prairies in openings in the Ozarks and Midwest be called xeric
Adams County, Ohio, ". antedate settlement by limestone prairies, in contrast to cedar glades,
. ,
white man and are undoubtedly primary." However, which are an edaphic climax dominated by C^ sum-
USDA
mer
using aerial photographs taken in 1938, annual grasses.
and Annala and
1950, 1965, 1971, (1983)
et al.
Annala and Kapustka found has Literature Cited
(1983) that prairie
&
succeeded (is succeeding) to forest. For example, Aldrich, R., A. Bacone M. D. Hutchison. 1982.
J. J.
prairie openings in the Lynx Prairie Pre Limestone glades of Harrison County, Indiana. Proc. In-
480-485.
diana Acad. 91:
Sci.
where the are "shallow and poorly developed"
soils
&
Annala, A. E. L. A. Kapustka. 1983. Photographic
47%
(Annala 1983: covered of the Pre-
et al., 22),
encroachment
history of forest in several relict prairies
16%
1938 1971 (Annala
serve in but only in et al, Adams
of the edge of Appalachia preserve system,
&
1983; also see Fore Guttman, 1996). Based on County, Ohio. Ohio Sci. 83: 109-114.
J.
&
mass between D. Dubois L. A. Kaputska. 1983. Prairies
lack of difference in opal (phytolith) , J.
A Adams
33-year history of two sites in
lost to forest:
soils presently under prairies and those presently
County, Ohio. Ohio 83: 22-27.
Sci.
J.
Ad-
under cedar-hardwood forests on dolomite in &
Bacone, A., L. A. Casebere M. D. Hutchison. 1983.
J.
ams County, Boettcher and Kalisz (1991: 127) con- Glades and barrens of Crawford and Perry counties. In-
367-373.
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that
. .
^
^^^^"'^J:
has been generally uniform over areas on do- *5^:„^:„?„^'''^
all
..^.^^^^^^^^
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lomite regardless of present prairie
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. .
meaning
mary and secondary has
prairies erlands.
little
&
1999. Cedar glades of southeastern
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.
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-J*
A
Adams
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r
m
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oi ^ is ^ 00 100
/., ,
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