Effect of Light Intensity on Arundinaria gigantea Growth and

Physiology
Author(s): Margaret C. Cirtain, Scott B. Franklin, and S. Reza Pezeshki
Source: Castanea, 74(3):236-246. 2009.
Published By: Southern Appalachian Botanical Society
DOI: 10.2179/08-060R3.1
URL: http://www.bioone.org/doi/full/10.2179/08-060R3.1

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CASTANEA 74(3): 236246. SEPTEMBER 2009

Effect of Light Intensity on Arundinaria gigantea

Growth and Physiology
Margaret C. Cirtain,1* Scott B. Franklin,2 and S. Reza Pezeshki3
Department of Biological Sciences, University of South Carolina, Columbia,
South Carolina 29208
2
School of Biological Sciences, University of Northern Colorado, Greeley, Colorado 80639
3
Department of Biology, University of Memphis, Memphis, Tennessee 38152
1

ABSTRACT The once dominant Arundinaria gigantea canebrake ecosystems have been
reduced to fragmented populations less than 2% of their former extent resulting in a critically
endangered ecosystem. Restoration of canebrakes is thus necessary for maintaining and
enhancing biodiversity in the southeastern United States. Contemporary fragments of
canebrakes are trapped between anthropogenic development and closed canopy forests. The
goal of this study was to assess the impact of light intensity on rivercane growth and
physiology in both laboratory and field studies and thereby enhance restoration success. A
laboratory experiment was conducted to test light (partial shading and no shading) and
nitrogen (0, 0.5, 5.0, 25, and 100 g/L) effects on seedling growth. Potential interaction occurred
between light and nitrogen. Levels of nitrogen up to ten times the commercially recommended
amount (0.5 g/L) significantly increased seedling growth when plants were grown under nonshaded conditions. A thinning site was established in an existing rivercane population and
forest canopy was reduced 60% by girdling overstory trees. Number of new shoots and new
shoot diameter were both increased by thinning. In addition, light response curves
demonstrated that rivercane had a typical C3 light response pattern and field readings showed
that rivercane maintained net photosynthetic activity throughout the dormant season. Results
indicate that rivercane growth is enhanced with increased light levels. Reduction of overstory
canopy is a potential management tool for enhancing survival and growth of existing
populations.
INTRODUCTION The Arundinaria gigantea
(Walt.) Muhl. (giant cane or rivercane) canebrake ecosystem, once a dominant landscape
component of the southeastern United States,
is now reduced to less than two percent of its
former status (Noss et al. 1995). Rivercane is
restricted to fragmented populations; most
commonly found along stream banks, fence
rows, and edges of agricultural fields and
forests. Fragmented populations have potentially been limited in growth due to two major
landscape modifications, increasing agricultural and urban development, and encroaching closed canopy forest where rivercane does
not receive adequate light (Gagnon and Platt
2008). Due to fragmentation and reduced
extent, an understanding of the physiological
capabilities of rivercane cannot be fully
developed from field studies. Studies of river*email address: mcirtain@biol.sc.edu

canes light and nutrient requirements are

lacking and research on these critical environmental factors is essential to restoration
strategies and success (Brantley and Platt
2001).
As a critically endangered ecosystem, the
declining rivercane population has impacted
faunal species diversity and ecosystem services (Platt and Brantley 2001). Swainsons
Warbler (Limnothlypis swainsonii), now placed
on the Audubon watch list, and the extinct
Bachmans Warbler (Vermivora bachmanii) are
avian species often associated with canebrake
decline (Brawn et al. 2001). Other species
impacted include the Creole Pearly Eye (Enodia creola), which uses rivercane as its host
plant, and mammalian species known to
utilize canebrake habitat (Platt and Brantley
2001), such as swamp rabbit (Sylvilagus aquaticus Bachman), cougar (Puma concolor), and
black bear (Ursus americanus). Additionally,

236

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CIRTAIN ET AL.: LIGHT INTENSITY ON ARUNDINARIA

canebrakes may have contributed significantly to ecosystem services such as erosion

control and water quality. Rivercane, with
its extensive rhizomal system, has been
considered for stream bank and roadway
embankment stabilization (Zaczek et al.
2004, Dattilo and Rhoades 2005). Agricultural sediment and nutrients can be reduced by
using rivercane buffer strips to improve water
quality in riparian restoration projects
(Schoonover et al. 2006). This loss of faunal
species and ecosystem services indicates that
reestablishment of canebrakes is necessary for
maintaining and enhancing faunal biodiversity and improving environmental quality in
the southeastern United States.
In a previous study, and based on the
location of remnant rivercane in the landscape, Cirtain et al. (2003) suggested that
light may be a limiting factor for rivercane
growth, while nitrogen had little effect. Rivercane remnant populations are often found
in forest understory where canopy limits
irradiance, potentially restricting population
recovery. This reduction in radiant energy
limits photosynthetic processes on which the
fitness success of a plant depends (Fitter and
Hay 2002). Canopy gaps, often disturbance
related, allow light to reach understory
plants, thereby significantly affecting plant
growth (Battaglia et al. 2000, Saitoh et al.
2002; Wang et al. 2006). Plant response to
this light fluctuation varies due to differences
in regeneration patterns in canopy gaps, as
has been shown with rhododendron in Appalachian oak forests (Clinton 2003) and bottomland hardwood forests (Sharitz et al.
1992). In addition, evergreen species may
potentially possess a phenological gap
advantage (Lei and Koike 1998), continuing
photosynthetic activity during the temperate
forest dormant season during diminished leaf
area. Rivercane populations may be responding to canopy gap and disturbance related
openings in the canopy (Gagnon et al. 2007,
Gagnon and Platt 2008) and, as an evergreen,
continue to photosynthesize during winter
months.
An additional critical environmental factor
for canebrake reintroduction is the impact
nutrients have on rivercane. Currently, populations of rivercane are often found along
stream banks, low-lying wetlands, and hillsides which are areas undesirable for agriculture. These current stands, as well as sites

237

being considered for reintroduction of rivercane, are often subjected to increased nutrient
input during periods of periodic inundation
and flooding. As such, these populations are
ideal for nutrient buffer strips and erosion
control. However, little is known of the
physiology of rivercane and its use and
tolerance of agricultural byproducts, typically
nitrogen and phosphate. For any restoration
project utilizing rivercane, understanding the
impact and tolerance levels that both established stands and introduced plants have for
these nutrients is necessary.
While the need for restoring canebrakes is
clear, methods for restoration and limitations
on rivercanes potential for restoration are
unclear. The goal of this study was to gain a
better understanding of the impact of light on
rivercane growth and physiology by examining light effects in both field and laboratory
studies. This research thus focuses on: 1)
examining the effect of light levels and
nitrogen on rivercanes growth and biomass
partitioning; 2) effects of light intensity on
rivercane in a site where light levels were
manipulated by thinning the overstory; and
3) determining leaf photosynthetic efficiency
by constructing photosynthetic light-response
curves and preliminary testing for photosynthesis during the dormant season. We hypothesized that: 1) both light and nitrogen
would increase rivercane seedling growth, 2)
thinning would enhance rivercane growth,
and 3) rivercane would continue photosynthetic activity during the temperate dormant
season.

MATERIALS AND METHODS

Study Species
Arundinaria gigantea, commonly known as
giant cane or rivercane, is one of only three
native bamboo species [includes two sister
species, Arundinaria tecta (Walt.) Muhl.
(switch cane) and Arundinaria appalachiana
Triplett, Weakley and L. G. Clark (hill cane)]
found in the United States (McClure 1973,
Calderon and Soderstrom 1980, Campbell
1985, Platt and Brantley 1997, Triplett et al.
2006). Its distribution is primarily limited to
the southeastern United States, from Florida
north to New York and west to Missouri,
Arkansas, and Texas. This species is monocarpic and gregarious, flowering approximately on a twenty to thirty year cycle
(Hughes 1951). Individual plants are lepto-

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CASTANEA

morphic, with an extensive underground

rhizome system. The culm or stem ranges
from 1 to 4 meters in height and from 0.5 to
2 centimeters in diameter, with lancelate
leaves that are persistent (Judziewicz et al.
1999, Triplett et al. 2006).

Laboratory Experiment
Light and nitrogen effects were tested with a
two-factor, experimental design with two
levels of light intensity typical of thinned
and unthinned forest understory of the study
area, control (200400 mmol/m2/s PAR) and
shaded (25% of control), and five levels of
nitrogen (0, 0.5, 5.0, 25.0, and 100.0 g/L).
Pots were constructed from 5.08 cm diameter
PVC pipe approximately 40 cm in length that
were capped on one end with three holes
drilled on two sides in a line about 6 mm from
the base for drainage. Pots were filled with a
soil composition of 60% play sand: 40%
commercial topsoil (Organic Valley Top Soil,
Home Depot). Two seedlings were planted in
each pot after recording initial measurements
of shoot length, root length, and number of
leaves. Seedlings were fertilized with Peters
20:20:20 followed by a two-week period for
acclimation during which seedlings were
watered every other day. Laboratory temperature averaged 72uC with supplemental light
maintained at 200 mmol/m2/s PAR (light
cycle 10 hour day, 14 hour night). Experimental setup consisted of 50 pots; 25 shaded
(25% of control: 50100 mmol/m2/s PAR) with
fabric, and 25 non-shaded control (200
400 mmol/m2/s PAR). Nitrogen was supplied
in applications of 50 ml per pot every other
day using ammonium nitrate. A concentration of 50 mg/L NH4NO3 was used in a prior
experiment and resulted in little effect (Cirtain et al. 2004); therefore, increased concentrations were used for this experiment. Levels
of ammonium nitrate applied were 0.5 g/L,
5.0 g/L, 25 g/L, and 100 g/L. The standard
commercial field application for ammonium
nitrate is the lowest concentration of 0.5 g/L,
and the highest concentration of 100 g/L was
designed to mimic agricultural and livestock
impoundment runoff concentrations (Carpenter et al. 1998, Schoonover and Willard
2003).
Eight weeks from treatment initiation,
seedlings were removed from pots and roots
were carefully rinsed to remove residual
potting soil. Measurements were taken for

VOL. 74

shoot and root length, number of shoots, and

number of leaves present. Shoot, root and
rhizome were separated, and subsequently
weighed both before and after drying for
48 hours at 60uC. Measurement values for
two seedlings from the same pot were averaged for statistical analyses. Biomass data
(shoot, root, and rhizome) and length (shoot
and root) and number (leaf and shoot) data
were analyzed using multivariate analysis of
variance (SPSS) and Tukey post hoc analysis
for main effects found significant in the
ANOVA. Shaded and unshaded groups were
analyzed separately and are only compared
through descriptive analyses.

Field Study
The field study site was located in Dahomey
National Wildlife Refuge (DNWR), approximately 3640 ha of bottomland hardwood
wetlands in the northwest portion of Mississippi, southwest of Cleveland, Mississippi
(N33u41.45 W90u55.24 Elevation 39 m).The
refuge provides habitat for migratory birds,
including waterfowl and songbirds, and public hunting for white-tailed deer and eastern
wild turkey. Historically, DNWR was dominated by extensive stands of rivercane which
were periodically flooded by the Mississippi
River (Survey conducted by Mr. Whitford on 4
December 1834). Remnant populations remain in forest understory and at forest-field
edges. Precipitation averaged 154.5 cm and
temperature averaged 17.7uC during the
experimental period (20012006). The frostfree period was March through November,
and vegetation was classified as bottomland
hardwood wetland. The study site was clear
cut in 1969, and the dominant tree species
was currently sweet gum (Liquidambar styraciflua L.) with some Southern red oak (Quercus
falcata Michx.) (B. Rosamond, United States
Fish and Wildlife Service, pers. comm.). Soils
are predominately Chromic Epiaquert (United
States Geological Service, Soil Survey 1958).
Paired sites, with treatments separated by
100 meters, were established in 2003 in
existing rivercane stands. The site was selected in the forest interior to eliminate edge
effects (approximately 75 meters from the
edge) and site boundaries were determined
by extent of rivercane population. Treatment
occurred in 20032004 dormant season by
girdling selected trees and treating the wound
with herbicide to reduce the canopy cover by

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CIRTAIN ET AL.: LIGHT INTENSITY ON ARUNDINARIA

239

Table 1. ANOVA treatment effect F values and r.F for biomass (shoot, root and rhizome) and length
(shoot and root)/number (leaf and shoot) from an Arundinaria gigantea seedling greenhouse experiment.
Treatments included two levels of light (control and shaded) and five levels of NH4NO3 (0, 0.5, 5.0, 25, and
100 g/L).
Treatment

Nitrogen and full light

Variable

F value

r.F

F value

r.F

42.59
17.49
11.95

,0.0001
,0.0001
,0.0001

18.55
5.21
104.27

,0.0001
0.005
,0.0001

4.48
16.78
38.37
9.65

0.010
,0.0001
,0.0001
,0.0001

6.91
4.81
39.56
10.71

0.001
0.007
,0.0001
,0.0001

Biomass
Shoot
Root
Rhizome
Length/Number
Shoot length
Root length
Shoot number
Leaf number

60% (Pickett and White 1985, Johnson 1993).

Post-treatment photosynthetic active radiation (PAR) readings conducted mid-day 12
May 2006 averaged 32 mmol/m2/s in the
control site and 781 mmol/m2/s in the thinned
site. Readings were taken at points from (1)
the site boundary nearest the edge of the
forested area, (2) mid-site, and (3) the site
boundary most distant from forest edge.
To monitor rivercane response, five 2 m2
plots were set up along each of four transects
(6 m apart) at two meter intervals for a total
of 20 plots in each site. Measurements were
taken on stem age, condition, diameter and
height on all individuals within study plots
prior to treatment and for three years (2004
2006) following treatment. Data from plots
were averaged for comparison of treatment
and control sites for each year of the study.
Three additional paired sites had been established for this study but were eliminated due
to interfering conditions, preventing statistical analysis of the data (n51).

Light Response of Rivercane

Photosynthetic light curves were conducted on
rivercane new shoots grown in the greenhouse
to gain a better understanding of its light
requirements and light utilization efficiency.
Plants were watered regularly and fertilized
two weeks prior to conducting measurements
to ensure no limiting effect of moisture and
nutrients. There were no additional treatments on these plants. The temperature in
the greenhouse averaged 20 to 25uC with
natural lighting. Measurements were conducted on March 5, 2007 using a CIRAS II
Portable Photosynthesis System (PP Systems,
Amesbury, Massachusetts). Photosynthetic

Nitrogen and shade

active radiation (PAR) levels were changed

in a series of intensities as follows: 0, 100, 200,
400, 600, 800, 1000, 1200, 1400, 1600, 1800
and 2000 mmol/m2/s. The measurements
were repeated with new shoots of three
healthy plants.
Photosynthetic measurements were also
examined during the temperate dormant
season (14001500 hour, 17 Nov 2006 and
26 Jan 2007) in the field on mature culms and
new shoots (,1 year old) at the Edward J.
Meeman Biological Field Station, western
Tennessee. Soil classification at this site was
predominately Memphis silt loam; fine-silty,
mixed, active, thermic Typic Hapludalfs (Soil
Survey Team 2008). A total of sixteen (17 Nov
06) and eight (26 Jan 07) individuals were
measured on the third leaf of a top branch, a
mid branch, and lower branch of each
individual plant. Air temperature and leaf
temperature averaged 13uC in November and
15uC in January. First frost in the area was 3
November 2006. Data collected included
stomatal conductance, photosynthetic rate,
and the calculated internal CO2 concentration (Ci), and are presented here using
descriptive statistics.

RESULTS
Laboratory Experiment
Arundinaria gigantea seedlings treated with
25 and 100 g/L nitrogen did not survive
through the eight-week experimental period. ANOVA results show treatment effects
vary among variables (Table 1). Seedling
growth was greater in all measurements taken for full light at 0.5, and 5 g/L
nitrogen levels (Figure 1a, b, c, d, and e). Leaf
and shoot number increased with nitrogen

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VOL. 74

Figure 1. Comparison of mean 6 standard error shoot height (a), root length (b), number of leaves (c), number
of shoots (d), shoot biomass (e), root biomass (f), and rhizome biomass (g) of Arundinaria gigantea seedlings grown
in the greenhouse under control and shaded conditions with nitrogen level applications of 0, 0.5, 5.0, 25, and
100 g/L. Tukey post hoc analyses indicated significant differences among nitrogen levels for (c) number of leaves
(r50.004) and (d) number of shoots (r50.002) under non-shaded conditions only, as letters indicate.

2009

CIRTAIN ET AL.: LIGHT INTENSITY ON ARUNDINARIA

241

application and was the greatest at 5 g/L

nitrogen in full light. Shoot biomass increased
with 0, 0.5, and 5 g/L nitrogen in full light.
Root and rhizome biomass increased with 0
and 0.5 g/L nitrogen in full light with a
decline in root and rhizome biomass at
5 g/L nitrogen.

Field Study
Results from the DNWR field site indicated a
greater number of new shoots in thinned plots
compared to control plots for both Fall 2004
and Fall 2005 (Figure 2). Individuals in the
thinned plot began to flower in 2006 and
could no longer be compared to individuals in
control plots. However, data on new shoot
height and diameter indicated reduced aboveground growth prior to flowering (Figure 3a
and b). There was little difference in the new
shoot height between control and thinned
sites (Figure 3a), and new shoot diameter
appeared to increase following thinning in
both control and thinned sites for one year.
During the following two years, new shoot
diameter decreased in both control and
thinned sites. However, in the control site,
there was a greater reduction in new shoot
diameter from Fall 2004 to Fall 2005 with a
slight increase from Fall 2005 to Fall 2006, but
well below new shoot diameter of the thinned
site (Figure 3b).

Light Response of Rivercane

Rivercane individual plants responded to
changes in light levels as typical C3 plants
with a photosynthetic light compensation
point between PAR of 30 and 50 mmol/m2/s,
and reaching a light saturation point
between 800 and 1000 mmol/m2/s PAR
(Figure 4). To date, there has been no
published photosynthetic response curve
on this species.
Photosynthetic rate of sixteen individuals
measured 17 Nov 2006 ranged from 2.7 to
11.6 mmol/m2/s (averaged 6.6 mmol/m2/s)
while stomatal conductance ranged from 59
to237 mmol/m2/s (averaged 131 mmol/m2/
s). Calculated internal CO2 concentration
(Ci) values ranged from 219 to 351 mmol/
mol (averaged 295 mmol/mol). Photosynthetically active radiation (PAR) values for 17
November 2006 averaged 755 mmol/m2/s and
ranged from 252 to 1273 mmol/m2/s. The
temperature averaged 13uC during data collection.

Figure 2. Mean 6 standard error (20 plots/site) of

Arundinaria gigantea new shoots/m2 from a thinned site
and a control site before and after thinning treatment.
Thinning mimicked a first shelterwood cut, reducing
the canopy by approximately 60%. Measurements
were taken each fall from 2003 through 2006.

Photosynthesis of eight individuals measured 26 Jan 2007 ranged from 3.6 to

11.8 mmol/m2/s (averaged 7.1 mmol/m2/s)
while stomatal conductance ranged from 37
to 293 mmol/m2/s (averaged 124 mmol/m2/
s). Calculated internal CO2 concentration (Ci)
ranged from 156 to 325 mmol/mol (averaged
259 mmol/mol). PAR values averaged 491
mmol/m2/s and ranged from 239 to 973
mmol/m2/s; temperature averaged 16uC. Our
results document that gas exchange and
photosynthetic carbon fixation continued
during the temperate dormant season.

DISCUSSION In an effort toward canebrake restoration, the goal of this study was
to assess the impact of light on rivercane
growth in both field and laboratory studies.
Results from all studies indicated that rivercane growth was enhanced with increased
light levels, supporting our hypotheses. These
results show light is a critical factor for both
maintaining and initiating canebrake restoration activities.
We had hypothesized that both light and
nitrogen would increase rivercane seedling
growth. As typical with many C3 grasses, light
and nitrogen are limiting factors, and the
addition of either elicits greater growth (Kozlowski and Pallardy 1997). Our data suggest
increasing nitrogen up to ten times (5.0 g/L)
the commercially recommended amount
(0.5 g/L) may significantly increase seedling
growth with plants grown under non-shaded
conditions. European beech seedlings (Fagus
sylvatica L.) were shown to have a similar

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VOL. 74

Figure 4. Light response curve of three Arundinaria

gigantea individual plants (new shoots) expressed as
mean 6 standard error.

Figure 3. Mean 6 standard error of Arundinaria

gigantea (a) new hoot height (m) and (b) diameter
(mm) from a thinned site (,60% canopy removal) and
control site at Dahomey National Wildlife Refuge.
Measurements were taken each fall from 2003
through 2006.

response (Johnson et al. 1997). Results indicated seedlings had greater biomass when
grown in organic soil when compared to
mineral soil and under increased light conditions. Additionally, there was a significant
shift in biomass allocation from roots to
shoots. Our laboratory study showed a different response. Shoot, root, and rhizome components of rivercane biomass were greater in
the non-shaded control compared to shaded
plants with nitrogen application up to 5 g/L.
These results suggest an increase in carbon
allocation to all plant components (Table 1,
Figure 1e, f, g), as opposed to a targeted
reallocation of carbon. Such a significant
increase in growth, particularly rhizomal,
could prove critical for establishment and
survival of plants at a restoration site.
However, higher N concentrations, similar to
those reported for agricultural runoff (Schoonover et al. 2006), may prove deleterious to

seedlings. Such high nitrogen levels decreased

seedling growth and survivorship in our
study. Thus, rivercane plantings should be
limited to more mature plants in areas with
excessive N levels.
Our second hypothesis was that canopy
thinning would enhance rivercane growth,
suggesting closed canopy forest species were
inhibiting canebrake expansion. Our field
study corroborates experimental and anecdotal evidence that rivercane responds positively to canopy thinning where we documented increased number of new shoots and
their diameter. Our results confirm those of
Widmir (1998) who showed an increase in
number of new shoots, culm height and
diameter in canopy gaps for Chusquea spp.
in Costa Rica. Likewise, Sasa palmate (Bean), a
Japanese dwarf bamboo, was shown to have
significantly greater above and below-ground
biomass and more new shoots and leaves in
higher light environments (Saitoh et al.
2002). Similar positive responses to canopy
gaps have been shown in a number of
understory species in the United States. Jordan
and Hartman (1996) were able to show
increased coastal sweet pepperbush (Clethra
alnifolia L.) growth with higher light resulting
from gap formation in the canopy. Rhododendron (Rhododendron maximum L.) and
mountain laurel (Kalmia latifolia L.) have
formed dense understory growth in Appalachian forest, possibly as a result of large gaps
due to heavy logging in the mid-twentieth
century (Sharitz et al. 1992, Hedman and van
Lear 1995, Clinton 2003). Disturbance, both
fire and wind, have been shown to impact
rivercane with a two-fold increase in culm

2009

CIRTAIN ET AL.: LIGHT INTENSITY ON ARUNDINARIA

number over control sites (Gagnon et al.

2007, Gagnon and Platt 2008). Finally, many
land stewards point to historical canopy
opening events for where remnant canebrakes are found today.
Data supporting our hypotheses regarding
light included rivercane showing a typical C3
photosynthesis response curve, and maintaining net photosynthetic activity during
the temperate dormant season. Our results
documented a compensation point (LCP)
between PAR 30 and 50 mmol/m2/s and
saturation point (LSP) between PAR 800 and
1000 mmol/m2/s, typical of C3 plants (Fitter
and Hay 2002) and typical of species found
under forest canopy. For example, the dwarf
bamboo Sasa senanensis (Rupr.) Makino et
Shibata has a LCP between 20 and 30 mmol/
m2/s and a LSP of 500 mmol/m2/s (Lei and
Koike 1998), and the shrub Lindera melissifolia
(Walt) Blume has a LCP of 33 mmol/m2/s and
a LSP of 490 mmol/m2/s (Aleric and Kirkman
2005).
As an evergreen species, rivercane potentially continues to photosynthesize throughout the winter months, storing carbohydrate
reserves in rhizomes. Our data from individual plants on different days during the
temperate dormant season demonstrates rivercane takes advantage of this phenological
gap. In this regard, rivercane has adapted in
a manner similar to conifers in deciduous
forests (Waring and Franklin 1979). Competing species coexist (Reich et al. 2003, Cavemder-Bates et al. 2004) by utilizing seasonal
variation which is an example of niche
partitioning. This phenological adaptation is
common to bamboo species in Asia and
Central America (Taylor et al. 2004, Widmer
1998), as well as forest species in the United
States. Understory woody plants utilize seasonal light variation to persist under canopy
(Augspurger et al. 2005, Katahata et al. 2007,
Xu et al. 2007).
Two predominant characteristics of rivercane that enable populations to persist and to
be restored are clonal growth and evergreen
leaves. One of the most intriguing aspects of
rivercane populations is the ability of this
clonal species to survive in communities
where competition for essential parameters
is reduced by ramets sharing of limited
resources. Stuefer et al. (1994) noted that
clonal plants may benefit from a heterogeneous light environment, due to the sharing

243

of resources among ramets where some are

in a resource rich environment and others in
a poorer environment. Clonal dwarf bamboo growth was greater not only in increased light but also when clonal connection remained intact (Saitoh et al. 2002).
Successful growth of plants in varying
environments is dependent upon efficient
usage of available resources. Rivercane may
have a competitive advantage as a clonal
plant, and examining its clonal integration
is a fruitful area for research. Our data
simply suggest light and nitrogen fertilization increase rhizome biomass in seedlings
and inferred competitive advantage due to
clonal sharing of resources.
The potential for rivercane to take advantage of a phenological gap is another area
deserving further research. Do winter conditions control vegetative production the following growing season? Is winter photosynthetic rate the reason for the northern border
of rivercane? Regardless, our studies show
light and nitrogen to be critical factors for
rivercane growth, necessary to understand for
successful restoration efforts. In conclusion,
these findings indicate canopy reduction and
nutrient supplementation should be considered as viable management tools.

ACKNOWLEDGMENTS Seedlings were germinated from seeds supplied by Paul Gagnon (Louisiana State University). We are
grateful to the staff of Dahomey National
Wildlife Refuge for their cooperation and
assistance. This research was partially supported by the National Fish and Wildlife
Foundation.
We appreciate the valuable statistical advising of Dr. John M. Grego and his student
Stoyan Gamishev of the University of South
Carolinas Department of Statistics.
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Salk. 2005. Light gains and physiological capacity of understorey woody plants
during phenological avoidance of canopy
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