Table Of ContentAND
COMPOSITION POTENTIAL COMPETITORS
FLORISTIC IN
^
MEUSSIFOLIA (LAURACE COLONIES
LINDERA AE) IN MISSISSIPPI
WITH REFERENCE TO HYDROLOGIC REGIME
Hawkins 2 Daniel Skojac 3 Nathan M. and Theodor Leininger
1 A. Schiff, D.
Tracy - Jr.
S.
, ,
USDA
Forest Service
Center Bottomland Hardwoods Research
for
U.SA
3
Stoneville, Mississippi 776,
1
ABSTRACT
fill
RESUMEN
INTRODUCTION
Species
member
United This
^melissifolia (Walter) Blume a dioecious shrub endemic to the southeastern States.
is
*l»aceae, commonly was South Carolina and described by Walter (1788)
called pondberry, collected in
®
Uurus few herbarium specimens
wtissaefolia. By the mid-20th century, Steyermark (1949) found that
“
made
dbeen having been
collected subsequent with only three collections
description,
to Walter’s (1788)
^
18 United
throughout the
first half of the twentieth Herbarium studies by Steyermark (1949)
century.
^confirmed Missouri,
Louisiana,
Georgia,
Alabama, Florida,
the historical presence of in
L. melissifolia
Tennessee,
T^Grolina, made in
and South Carolina. Although Gattinger (1901) reference to L. melissifolia
er® rk(1949) pubescent Lindera Benzoin, since
la
misidentified
^ believed this report probably referred “...to
*
Wtnentic University of Tennessee...
herbarium of the
material of has been found in the
L. melissaefolia
7^’ with
only one historic report
(1924) excluded from “Shrubs of Indiana” citing
L. melissifolia
joucher concluded that
Steyermark (1949)
specimen.
Based on the number of preserved L. melissifolia,
ma
Cies
be
^ one United
y of the rarest shrubs in the States.
J^atly, Arkansas, Georgia, Mississippi,
Alabama,
extant present in
populations of are
L. melissifolia
and In the
Louisiana Florida.
North be extirpated in
Carolina, and South believed to
Zj*"’ Carolina.
It is
2**
margin of seasonally
grows along the
Coastal
^ Plain (South Carolina and Georgia), L. melissifolia
&
(Alenc
Brongn.
de and Taxodium ascendens
J*" pressional Walter
wetlands dominated by Nyssa
biflora
depres-
grows along the edge of a forested
2005). The Alabama
known
only population to occur in
,
382 Journal of the Botanical Research
Institute of
Texas
4(1)
Lower (LMAV),
2005), In the Mississippi Alluvial Valley L. melissifolia populations in Mississippi,
Arkansas,
and Missouri grow in periodically flooded, bottomland hardwood forests underlain by hydric
soils. Forest
LMAV
canopy composition among composed
the sites in the is similar, primarily of trees designated
as
facultative wetland or obligate wetland species (Hawkins et 2009a). Liquidambar
al. styraciflua, Acer rubrum,
Q. lyrata Q. nuttallii, and phellos are important canopy components; however, the relative importance
Q.
of
these and other canopy species varies between sites (Hawkins et 2009a).
al.
Lindera melissifolia populations consist of spatially segregated, unisexual colonies (Hawkins
etal. 2009b).
Colony sizes range from approximately 20 stems to >1,000 stems (Morris 1987; Devall et 2001; Hawkins
al.
LMAV,
pers. obs.). In the populations are strongly male-biased, with male female colony
to ratios ranging
from (Wright 1994) (Hawkins
7:1 to 19:1 et 2009b).
al.
In both male (pollen-bearing) and female (seed/fruit-bearing) L. melissifolia, anthesis occurs in bte
spring, often during flooded conditions, and precedes leafing out. Small, yellow flowers are produced on
axillary inflorescences (Fig. 1A), followed by production of green drupes on female plants. Approximately
90
days following anthesis, drupes contain a single, fully developed seed (Connor et 2007) and upon
al.
maturation drupes
are bright red (Fig. IB, 1C). Drupes are dispersed from to early winter (Smith etal.
fall
and may
2004) fruit pedicels remain on the plant until the following spring. Female L. melissifolia appear
to invest heavily in sexual reproduction (Connor et 2007); however, seedlings are rarely observed in
al.
naturally occurring populations (Wright 1990). Vegetative propagation of ramets from rhizomes appears to
be the predominant form
of reproduction (Wright
1990, 1994).
During
one
the first to three years following seedling and/or ramet emergence, L. melissifolia plants
are morphologically very
similar to L. benzoin Blume. In during Steyermark’s (1949) investigation
(L.) fact,
of the species, he noted numerous misidentified herbarium specimens of resulting from this
L. melissifolia
close similarity. Although leaf pubescence and leaf size sometimes differ between the two species, these
may
characteristics be dependent on season and locality. On the other hand, the angle of the lateral veins
may
in the leaf blade be used to distinguish the congeners. In the lowest two pairs of lateral
L.
melissifolia,
veins of the leaf blade diverge at a 45°-50° angle from the midrib, and successive veins diverge
distal to this,
at approximately 35° to the
(Steyermark, 1949). In other words, the lower two veins are not parallel
lateral
successive upper same
veins. In contrast, all lateral veins of L. benzoin leaves diverge from the midrib at the
(194#*r
angle (35°-45°);
therefore, are parallel (Steyermark Both and Steyermark
1949). Nuttall (1818)
scribed the fruits of L. melissifolia as “larger” than those of L. benzoin. The senior author has found the seeds
L
of the congeners be While seeds of
to consistently
reliable for definitive identification of fruiting plants.
biow»
melissifolia are spherical and brown brown and dark
light to yellowish those of L benzoin are oval
(Fig. ID).
and
In 1986, was Fish
L. melissifolia listed under the Federal Endangered Species Act of 1973 (U.S.
(Hawknwjw^
Wildlife Service
1986). Recent
research has provided some ecophysiology
insight into the
& wi*T
2009b; Kirkman
Aleric
2005) of and associated
this species, types and structure
forest forest
LMAV
speck*
melissifolia in the have been suggested
reported (Hawkins Wright
(1990)
2009a).
et
al.
peri**
Brunnichia, Rubus, and Smilax main and that
as competitors Arkansas,
of northeast
L. melissifolia in
flooding served very
minimize
to competitive Beyond report,
interactions. Wright’s (1990) initial
P^
known
about
is the to
microhabitat or biotic interactions in colonies. In an effort
L. melissifolia
concise information disjo**
regarding native,
the microhabitat and three
general ecology
of L. melissifolia,
populations asse®
in Mississippi were monitored for three years. The objectives of our study were to 1)
vascular plant P*#**
checklist of groundcover identify
species growing within colonies, 2)
L. melissifolia
competitors
of
L. and
melissifolia, monitor
3) hydrologic regime colonies.
for
L. melissifolia
Lindera melissifolia colonies
L,
and seeds
L. melissifolia, (D)
!
384 Journal of the Botanical Research
Institute of Texas
4<1)
METHODS
Study
Sites
Two The North
of the study are in Sharkey County, Mississippi. Delta National Forest (NDNF)
sites site
a
is
25-ha tract of Delta National Forest that has been under management by the U. S. Forest Service since 1938
&
Ramp
(Devall 1992). The South Delta National Forest (SDNF) site is a 10-ha tract of forest approximately
km
NDNF.
9 southeast of The third study site (BC) is in Bolivar County, Mississippi, and is a privately owned
30-ha fragment surrounded by Hawkins 2009a map and
forest agricultural fields (see et al. for detailed
Dowling
descriptions). Soil association for the three sites is (very fine, smectitic, nonacid, thermic Venic
Chromic
Endoaquepts)-AUigator thermic Dystraquerts)-Sharkey
(very-fine, smectitic, (very-fine, smectitic,
thermic Chromic Epiaquerts), an association characterized by poorly drained, fine-textured clayey
surface
&
and formed from Mean
soils subsoils Mississippi River alluvium (Rogers 1958, Scott Carter 1962). annual
temperature for Sharkey and Bolivar counties approximately 18.0°C, and total annual precipitation ranges
is
mm mm &
from 1228
1319
to (Rogers 1958, Scott Carter 1962).
Data and
Collection Analysis
autumn BC * NDNF
In 2003, rectangular plots were established for selected L. melissifolia colonies at (Ncoiony 6),
= and SDNF = SDNF was numerous many
( Ncoiony 10), (Ncoiony 1). In 2000, habitat for L. melissifolia colonies,
with >200 stems (GSRC 2000). However, during reconnaissance of this area we found only one remaining
lm
colony and were able to establish only one study plot. The perimeter of each plot was positioned beyond
m
A
cm
the outermost L. melissifolia stems of a colony. 1.2 wooden stake x 5 cm) was established at each
(5
comers
of the four of the rectangular plot. Colony sizes were variable, and thus plot areas varied with colony
lm lm
size. Within each plot, x quadrats were marked with pin along the diagonals of the rectangular
flags
plot; therefore, the percent of plot area sampled was the same among
plots.
In June 2004, May 2005, and June 2007, L. melissifolia stems in each lm x lm quadrat within the {Jots
Common
were and were
counted, groundcover species were and stems were counted. plants
identified
TSH When were
identified in the field by and DAS. was problematic, plants in question
field identification
top-cropped and by Godfrey
taken supplemented
to the lab for identification using Radford
et (1968),
al.
and Wooten
(1979, 1981).
From
10 November 2004 22 May When were not Hooded,
to 2007, plots were visited bi-weekly. plots
mewl
two
samples were cm
soil collected from each plot and placed in individual 141 3 hermetically sealed,
,
an oven at
containers. Soil samples were taken directly to the lab, weighed to the nearest 0.01 dried in
g,
70°C for 48 hr, then weighed again. Percent moisture each sample was calculated by dividing the
for soil
W-
by
difference of and quotient
initial (wet) final (dry) weights by the weight and multiplying the
initial
and
During comer post
flooded cm
conditions, water depth was measured each plot
to the nearest 0.1 at
mean
the SE) used
(± to represent water depth
for the
plot.
NDNF, colonies
At were
there substantial differences in water depth among some of the L. melissifolia
mean analysis
one-way
therefore, water
levels for each sampling date each colony were compared using a
at
of variance (ANOVA; SAS different (p
2001). For colonies where mean water was not significantly
level
0.2421), data were pooled producing one mean ± standard error for each of three groups of colonies.
AND
RESULTS
DISCUSSION
defini-
Hydrologic broad
regime
for forests with Lindera melissifolia populations often described within the
is
tion of
“periodically flooded”. For populations occurred in
in Mississippi, flooding generally
BC
The
-spring.
vever, flooding events and among
flood duration were found vary sites.
to
d and
annual
flooding events of comparable depth and duration in 2005, 2006, 2 ^
regime
t this site is artificially controlled by the lan refore, hydrologic i
from
isistent year
to year.
.
m #0*
Flood
duration and
time NDNF. occurred
initial of flooding varied with year Flooding
at
c
2006, but not in 2007 monitored
i (Fig. 2B). Although water depth differed among some of the ten
NDNF, had no influence on time and duration of individual colony inundation
this (Fig. 2B). Difference
in
among
water depth colonies is the result of surface topography (e.g. sinks or sloughs). Lack of difference
in
time and duration of flooding between colonies results from rapid rising and dropping of water. In
contrast
NDNF
SDNF
to and BC, did not experience flooding in 2005, 2006, and 2007 (Fig.
3).
When
L. melissifolia colonies were not flooded, soil moisture content was comparable among the three
20% 30%
sites, ranging from to throughout the year, with the exception of mid- to late-summer, when
soil
NDNF
moisture contents dropped as low as 15% to 18% for BC and (Figs. 2A and 2B); and as low as 10%
SDNF
(Fig
at
3).
The combined
annotated list of vascular plants (including L. melissifolia for the three study sites includes
)
70 species in 57 genera in 45 families (Appendix and of these, 9 species have the potential to become
1)
weedy or invasive (SWSS 1998). The contribution of L. melissifolia to overall colony composition ranged from
20%-40% NDNF 5%-15% SDNF
approximately at and BC, and at (Fig. Other species growing within the
4).
colonies are typical of bottomland hardwood forests in this area of the LMAV, and tend to reflect hydrologic
regime at each study site. Forty-nine percent of the 69 species identified as growing in association with L.
FACW
67%-99%
melissifolia (Appendix have a wetland indicator status of and a probability of occurrence
1)
NRCS SDNF
wetland (USDA,
in a area 2008). However, the presence of Callicarpa americana at is atypical
LMAV
for bottomland forests in the and represented a county record reported in 2007 (Skojac et al.
first
SDNF
Lack
2007). of inundation at (Fig. has possibly allowed establishment of C. americana, as well as
3)
other species, such as Asplenium platyneuron and Phytolacca americana, that generally are not found in forests
prone
to flooding.
Of 69 immediate
the species growing in association with few appeared to pose an
L. melissifolia,
above-ground competitive threat and the ratio of stem density for these plants to L. melissifolia stem density
remained
relatively stable throughout our study The predominant growth habit in L. melissifolia
(Fig.
4).
colonies was vines (Fig 4). Wright (1990) considered Brunnichia ovata as a plant with potential to be an ag-
gressive competitor of L. however, we observed early-summer emergence of B. ovata, as well as
melissifolia;
Toxicodendron and radicans
radicans, after L. melissifolia plants had flowered and leafed out. Both B. ovata T.
remained
prostrate throughout the season and did not compromise capture by L. melissifolia leaves,
light
nor twine around or climb On bona-nox, glauca, S.
L. melissifolia stems. the other hand, Smilax spp. (S. S.
become
rotundifolia, S. tamnoides) and Vitis spp. palmata, have potential to
(V. aestivalis, V. V. rotundifolia
)
stems
strong competitors, by remaining upright throughout and some using melissifolia
the year, in cases, L.
above-ground
for
support.
Many have direct
of the associated species in do not appear to
L. colonies in Mississippi
melissifolia
those
competitive impact on and/or
growth habit
L. populations. However, with vining
melissifolia species a
response
with potential become weedy The
to or invasive (Appendix should continue to be monitored.
1),
interactions
of these species to natural or anthropogenic competitive
disturbances has the potential to alter
within
these
L. melissifolia populations.
APPENDIX
1
(Monilophytes
Plants included
in this checklist are compiled in alphabetical order by family within two major groups
names
Angiosperms). Genera and common
species are alphabetical within each nomenclature and
family. Scientific
(USDAN
The
Plants Database (USDA, NRCS Database
2008). Family circumscriptions monilophytes follow The Plants
for
aredeno^
and
2008), for angiosperms, APG 998)
(Stevens become weedy (SWSS,
2008). Plants with potential to or invasive 1
with an
asterisk (*) before the species name. Region (AL AR,
Species wetland indicator status for the Southeast
(67%'W*^
NC,SC,TN) =
noted
obl wetland
is as: obligate (99% =
probability ofoccurrence wetlands); facw facultative
in
ability of occurrence <*
in wetlands, but occasionally found = (equal probability
in non-wetlands); fac facultative
m
wetlands w
or non-wetlands); =
facu facultative upland (occurrence non-wetlands, occasional
usually in
and - -
ni unable to determine wetland indicator status based solely on genus. Locality data are noted as: (1) BotlV M-
-
MS; 2
North County.
( ) Delta National Forest, Sharkey County, MS; and 3 = South Delta National Forest, Sharkey
( )
Journal of the Botanical Research
388 Institute of Texas4(1)
Trees/Shrubs Vines Pondberrv
Forbs/Herbs
KS
|
BC
-NDNF SDNF
1
1
per unit area (stems/m2
).
Melothria pendula L, Guadeloupe
ci
Cyperaceae
Carex crus-corvi, Shuttlw. ex Kunze,
r
coralbead, fac, (2,3)
Carex LH, Carolina
louisianica Bailey, Louisiar *Cocculus carolinus DC.,
(L.)
Carex tribulokles Wahlenb., blunt
brc
Moraceae
swamp obl (2)
privet,
Forestiera acuminata (Michx.) Poir.,
33)
Marsh, green ash, facw, (1
Fraxinus pennsylvanica
pumpkin ash, obl, •
Fraxinus profunda (Bush) Bush,
Gleditsia triacanthos L, honeylocust, (13
fac,
Ludwigia glandulosa Walter,
Fagaceae
Quercus
lyrata Walter, overcup oak, (U)
obl,
Orchidac
Quercus nigra water n
oak,
L., fac,
(3)
tresses,
October lady's
ovalis Lindl,
?s
Quercus
pheltos L, willow oak, facw,
(1,23)
Quercus texana
Buckley, Texas
red oak,
obl,
(1
passionflower, fa
yellow
*Passifiora lutea L,
Uquidambar Phytolaccaceae
sweetgum,
styraciflua L,
fac,
(1,
pokeweed, f*
* American
Phytolacca americ i L„
Carya
aquatica
(Michx. Nutt, water
f) hickor
389
Smilax glauca Walter, cat greenbrier,
fac,
i
ovata (Walter) Shinners, redvine, facw, Smilax rotundifolia roundleaf greenbrk
unnichia (1,2,3) L.
'Sr
Polygonum virginianum L, jumpseed, fa Smilax tamnoides L, bristly greenbrier, fai
(2)
c,
Styracaceae
d leatherflower, facw, Styrax americanus Lam, American snowbell, facw, (2)
(2)
Ulmaceae
supplejack, facw, Planera aquatica Gmel, water elm, obl,
J.F. (2)
i
Ulmus americana American elm,
L, facw,
(1 ,2,3)
Rubus Michx, Boehmeria cylindrica Sw, false nettle, facw, (23)
trivialis (L.)
buttonbush, Callicarpa americana L, American beautyberry, facu,
obl, (3)
(2)
Violaceae
Viola sp. L, violet, ni, (3)
*Ampelopsis arborea (L) Koehne, peppervine, fac, (133)
*Parthenocissus quinquefolia Planch, Virginia creeper.
(L.)
fac, (1,23)
summer
Michx. grape,
Vitis aestivalis fac, (2)
palmata Vahi, catbird grape, facw,
Vitis (1,2)
Michx, muscadine,
rotundifolia fac, (23)
Vitis
ACKNOWLEDGMENTS
Ihe
authors thank Greg Comer, Stephanie Skojac, and Theran Stautz for their assistance in the field, U. S,
Fish and Wildlife Service for permits, the U. S. Army Corps of Engineers for underwriting the cost of this
research, and Drs. Charles Bryson, Emile Gardiner, and Edward W. Chester for review of an earlier draft
°f
this manuscript.
REFERENCES
Alewc
endangered
K-M. and LK. Kirkman. 2005. Growth and photosynthetic responses of the federally shrub,
'
Lindera
Amer. 92:682-689.
melissifolia (Lauraceae), to varied light environments. Bot.
J.
^
Cqnn
and!
K, G5. 2007.
Schaefer, Donahoo, M. Gardiner! Hawkins, D. Wilson, N. Schiff, Hamel, Leininger.
Devall, P.
J. E.
Development, endangered pondberry
fatty acid composition, and storage of drupes and seeds from the
(Undera
melissifolia). Cons. 137:489-496.
Biol.
DtAM.CC.
1924. Shrubs of Indiana. Indiana Department of Conservation, Indianapolis. Pub. No. 44.
Dewu
MS.
and and protection of old growth in the
» P.F. Ramp. 1992. U.S. Forest Service research natural areas
Nat.
Areas.
2:75-85.
J.
1
N
endangered pondberry, Undera
Schiff, and D. Boyette 2001 Ecology and reproductive biology of the
-
.
Melissifolia
(Walt.) Blume. Nat. Areas 21 :250-258.
J.
*®*
A
I1 - 190! The flora of Tennessee and a philosophy of botany, respectfully dedicated to the citizens of
enn
essee.„
Press of Gospel Advocate Publishing Co, Nashville, TN.
R
k and United Monocotyle-
- J.W. Wooten. 979. Aquatic and wetland plants of the southeastern States.
1
5***
Unive
rsity of Georgia Athens.
Press,
^
UK.
and J.W. Wooten. 198! Aquatic and wetland plants of the southeastern United States. Dicotyledons,
diversity
of
Georgia
Athens, GA.
Press,
South
pondberry Prepared
Research Corporation (GSRQ. 2000. survey report: reevaluation of in Mississippi.
Final
US. Army
Corps of Engineers, Vicksburg Gulf South Research Corp, Baton Rouge, LA.
^ District,
DS. Bottomland the
and 2009a. forests in
Skojac, B.R. Lockhart, T.D. Leininger, M.S. Devall, NJVI. Schiff.
UJvver
Mississippi endangered Undera Castanea 74:105-1 13.
with the melissifolia.
Alluvial Valley associated