Journal of the Torrey Botanical Society 135(3), 2008, pp. 309–316
Reproductive and seedling ecology of a semelparous native
bamboo (Arundinaria gigantea, Poaceae)1
Paul R. Gagnon2,3 and William J. Platt
Louisiana State University, Department of Biological Sciences, Baton Rouge, LA
GAGNON, P. R. AND W. J. PLATT (Louisiana State University, Department of Biological Sciences, 202 Life
Sciences Building, Baton Rouge LA 70803). Reproductive and seedling ecology of a semelparous native
bamboo (Arundinaria gigantea, Poaceae). J. Torrey Bot. Soc. 135: 309–316. 2008.—Canebrakes were
monodominant stands of bamboo once common in bottomlands throughout the southeastern U.S. They
were habitat for many wildlife species, and have declined drastically since the 18th century. Knowledge of the
reproduction of canebrake bamboos is sparse and often contradictory. We studied reproduction and the role
of disturbance in one canebrake bamboo, Arundinaria gigantea (Walt.) Muhl., at two field sites in Louisiana.
We noted flowering with little or no seed-set during each of four years at our primary study site, and a massflowering event at a second site from which we collected 2,000 seeds. Of these, 82–95% proved viable in
germination trials. We used these seeds in a 2 3 2 factorial design at the Buckhorn Wildlife Management
Area in NE Louisiana to test the effects of prior windstorm and fire on seed germination and seedling
survival. Results indicate that A. gigantea is capable of reproducing in the leaf litter and partial shade typical
of open forest habitat but may have problems reproducing in burned-over areas with bare mineral soil. We
identified three potential bottlenecks in the species’ regeneration and proposed that successful outcrossing
may drive synchronized flowering events in these and other bamboos. We presented implications for
canebrake restoration efforts, plus several questions that might be examined by future studies.
Key words: Arundinaria gigantea, bamboo reproduction, bottomland hardwood forest, canebrakes, fire
ecology, giant cane seeds, mass-flowering, multiple disturbances, tornados, windstorms.
Canebrakes are monodominant stands of
bamboo once common in bottomlands
throughout the southeastern U.S. The dense
and expansive canebrakes described by early
explorers have declined by an estimated 98%
since the 18th century (Noss et al. 1995).
Causes of this decline include overgrazing,
land conversion to agriculture, and alteration
of the disturbance regime in bottomland
hardwood forests (Platt and Brantley 1997,
Gagnon and Platt 2008). Canebrakes are
habitat for numerous wildlife species, including several threatened and endangered species
(Roosevelt 1908, Platt et al. 2001). Two of the
three recognized native bamboos in the genus
Arundinaria are known to form canebrakes:
1 This research was funded by U.S. Environmental Protection Agency S.T.A.R. Fellowship #
U916181, a Louisiana Board of Regents Fellowship,
and grants from the American Bamboo Society and
the J. Bennett Johnston Science Foundation.
2 For their support and ideas we thank Tommy
Tuma, Kenny Ribbeck, Randy Ewing, Heather
Passmore, and Jim Cronin. We thank the Louisiana
Department of Wildlife and Fisheries for logistical
support. We thank three anonymous reviewers for
their suggestions with this manuscript.
3 Author for correspondence. Current address:
Department of Wildlife Ecology and Conservation,
110 Newins-Ziegler Hall, PO Box 110430, Gainesville, FL 32611-0430. E-mail: [email protected]
Received for publication December 4, 2007, and
in revised form May 5, 2008.
Arundinaria gigantea (Walt.) Muhl. (Poaceae;
giant cane) and A. tecta (switch cane; Triplett
et al. 2006). These bamboos still occur in
small, sparse stands, often along forest edges
and in gaps. Due in part to their value as
wildlife habitat, there is considerable interest
in restoring these native bamboo stands to
monodominant canebrake-like structure.
Information about reproduction in canebrake bamboos is sparse and often contradictory. This is in part because, like other woody
bamboos, species of Arundinaria are thought
to be semelparous; they grow for decades
before reproducing one time and dying
(McClure 1966, Judziewicz et al. 1999).
Accounts of flowering in Arundinaria vary
widely, ranging from reports of occasional
culms flowering yearly, to synchronized massflowering events that produce millions of seeds
and massive die-offs (Hughes 1951, Marsh
1977, Platt and Brantley 1997 and references
therein, Judziewicz et al. 1999). Arundinaria
seed and seedling ecology is similarly enigmatic (but see Hughes 1951, Marsh 1977, Cirtain
et al. 2004, and Platt et al. 2004). These
bamboos are disturbance-dependent (Hughes
1957, Gagnon et al. 2007, Gagnon and Platt
2008), but little is known about how postdisturbance conditions influence seed germination and seedling survival.
We studied the role of reproduction in the
life history of Arundinaria gigantea. We used
309
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[VOL. 135
both descriptive and experimental approaches
to: 1) determine the effects of post-disturbance
conditions on seed germination and seedling
survival and 2) identify and describe critical
stages in cane sexual reproduction, including
flowering and seed/seedling ecology. Based on
our prior study of the responses of full-size
plants to wind and fire disturbances (Gagnon
et al. 2007, Gagnon and Platt 2008), we
expected that A. gigantea seeds and seedlings
would respond positively to the high light
conditions of previously disturbed sites. Specifically, we hypothesized that rates of both
germination and seedling survival would be
higher in post-windstorm (i.e., large forest
gaps), post-fire, and combined post-windstorm and post-fire conditions than under
undisturbed forest canopy. Results of our
study offer insights into the ecology of
Arundinaria and other bamboos, and inform
canebrake restoration efforts.
FIG. 1. The Buckhorn Wildlife Management
Area, where we planted seeds of Arundinaria
gigantea, including photos of: A) the bottomland
hardwood forest, B) large tornado-generated gap, C)
a stand of A. gigantea growing in a small forest gap,
and D) prescribed burning of A. gigantea prior to
Materials and Methods. SITE DESCRIPTIONS.
Our primary site was the Buckhorn Wildlife
Management Area (WMA) in northeast
Louisiana. Located in Tensas Parish within
the lower Mississippi alluvial valley (32u 029
410 N, 91u 219 530 W), the holding was
administered by the Louisiana Department
of Wildlife and Fisheries. Approximately
three-quarters of its 5,000 ha contained bottomland hardwood forest, and the balance
was reclaimed agriculture fields. Soils were
somewhat poorly drained alluvial clays on
gently undulating ridge/swale terrain ranging
in elevation from 10 to 25 m, and 0–3u in
slope. The forested portion was managed
using single-tree and group selection silviculture. In November 2000, a large F2 tornado
crossed the Buckhorn WMA, leaving a very
large blowdown gap. The central zone of this
gap was approximately 1 km wide running the
length of the holding. Within this area
virtually all trees were snapped off or blown
over (Fig. 1B; for a more detailed description
of this tornado-generated gap see Gagnon et
al. 2007). Our study species (Arundinaria
gigantea) occurred throughout the holding
along forest edges, under forest canopy and
in the large tornado blowdown.
r
planting. Photo credits: Virginia Velez Thaxton (A)
and Paul R. Gagnon (B, C, D).
2008]
GAGNON AND PLATT: ECOLOGY OF GIANT CANE
A secondary study site was Tunica Hills
Wildlife Management Area. The site was
located approx. 150 km south of the Buckhorn WMA, immediately east of the Mississippi River in the ‘‘Loess Hills’’ (30u 569 190
N, 91u 309 120 W), at elevations ranging from
50–75 m. Terrain included bluffs, rugged hills,
and ravines. Vegetation was mixed mesophytic
hardwood forests on well-drained fertile silt
loam soils (Delcourt and Delcourt 1974). The
focal bamboo was common in the understory
and along forest edges.
OBSERVATIONS OF FLOWERING, SEEDING,
GERMINATION, AND ESTABLISHMENT. We documented Arundinaria gigantea reproduction at
the Buckhorn WMA every winter and spring
for four years. We recorded the presence of
flowering individuals within patches, and
made note of the size of patches. Whenever
flowering culms were observed, we looked for
seeds in their synflorescences (aggregations of
spikelets terminal to the culm or lateral branch
that compose bamboo flowering structures;
Judziewicz et al. 1999), and we checked the
ground for seedlings. We examined every seed
we encountered for evidence of seed predation.
We documented a mass-flowering event of
Arundinaria gigantea at Tunica Hills WMA in
spring 2004. Unlike the flowering events we
witnessed at the Buckhorn WMA, A. gigantea
at Tunica Hills flowered synchronously and
extensively. The event produced a very large
quantity of well-developed seeds, and the
mature plants died immediately thereafter.
On April 22, 2004 we collected approximately
2,000 of these seeds by shaking or lightly
stripping them from culms. We monitored
seedling growth during several subsequent
visits to Tunica Hills and censused the area
at the beginning of the next growing season
(44 weeks after collecting the seeds).
SEED AND SEEDLING EXPERIMENTS. To estimate the viability of seeds, we performed three
germination trials. The day after collecting, we
placed 21 well-developed seeds on a moist
paper towel and kept them in partial light at
room temperature. Well-developed seeds were
easily distinguished by their greater size and
mass (the large majority of the Tunica Hills
seeds appeared to be well-developed). Six days
after harvesting the seeds we randomly chose
90 to plant into a sterile, unfertilized 2:1:1 mix
of peat moss, perlite, and vermiculite. We then
311
stored remaining unplanted seeds in an airtight container at 4 uC in a dark refrigerator.
Four weeks after collecting seeds, we chose
100 well-developed seeds to plant into a sterile
2:1:1 mix of peat moss, vermiculite, and
fertilized perlite. We kept all planted seeds at
room temperature in partial shade/sun, and
tallied number of germinated seeds three
weeks after each planting.
We used a 2 3 2 factorial design at the
Buckhorn WMA to test the effects of prior
windstorm and fire on seed germination and
seedling survival. We set up eight study plots
in the large tornado blowdown (windstorm
treatment) and eight under surrounding forest
canopy (windstorm control; Fig. 1). Half of
the plots in each habitat had been burned
three weeks prior to planting seeds (fire
treatment; Fig. 1). Within each plot, we
located one pair of 1 m2 subplots 2 to 5 m
apart; each was devoid of competing vegetation, although adjacent vegetation was typically present. We chose only well-developed
seeds for the experiment. We planted 32 seeds
into each subplot by pressing seeds lightly
into the soil or leaf litter. We marked the
location of every seed using flags, and recensused the plots three, 11, and 44 weeks
after planting. We collected data on germination, establishment, survival, and growth of
each seed/seedling. We also observed the size
and health of seeds and seedlings, and noted
any signs of seed predation. We used ‘‘seedling
survival’’ to refer specifically to the likelihood
of seedlings in one census surviving to the next
census.
We quantified light in forest and blowdown
plots. We took hemispherical canopy photos
in each plot at 1.5 m in height and from these
estimated percent total transmitted light using
Gap Light Analyzer 2.0 (Frazer et al. 1999).
Mean total transmitted light in forest plots
was 16.9% 6 1.68 SE. Mean total transmitted
light in blowdown plots was 89.9% 6 1.97 SE.
This difference was significant (F1,14 5 798, P
, 0.001).
STATISTICAL ANALYSES. We used generalized
linear mixed models to test effects of fire and
windstorm on germination and seedling survival. We coded germination as a binary
response variable weighted by the number of
seeds originally planted into each subplot (32).
We performed repeated measures analyses on
survival, which we calculated as the number of
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JOURNAL OF THE TORREY BOTANICAL SOCIETY
seedlings alive during a given census weighted
by the number alive during the previous
census. Our three censuses yielded data for
two transition periods as the repeated measures (time) variable. We performed these
analyses using Proc Glimmix in SAS with a
binomial distribution and logit link (SAS
Institute 2005). In addition to the linear model
analyses, we explored among-plot differences
by ranking plots, and by determining individual plot contributions to overall means.
Results. OBSERVATIONS OF GIANT CANE
FLOWERING, SEEDING, GERMINATION, AND ESTABLISHMENT. In each of four years, Arundinaria gigantea flowered in a limited way at our
primary research site (Buckhorn WMA).
Flowering began in late winter, at which time
leaves started to brown and dry. In March,
straw-colored flowering patches grew increasingly obvious as the surrounding (non-flowering) patches began to flush new leaves. On
close inspection, synflorescences resembled
large heads of barley or rice (Fig. 2A). On
multiple occasions we noted a single culm
flowering alone. Every year small patches
(presumably individual genets) flowered, but
did not produce seed. On two occasions we
found patches that flowered a second year but
produced no seeds in either year. Twice during
the four years we found somewhat larger
flowering patches that did produce a limited
number of seeds. These few seeds did not
detach quickly, and we observed various
insects consuming the seeds, including weevils
(Curculionidae), chinch bugs (Blissus sp.,
Lygaeidae), and a small Lepidopteran larva
that burrowed into and consumed seeds. Too
few seeds were produced (fewer than two per
culm) for germination experiments. Flowering
culms always died, sometimes after a one-year
lag.
The mass-flowering event at our secondary
research site (Tunica Hills WMA) was highly
productive. When we first visited the South
Tract of this site in early April 2004, virtually
every culm we examined over a large area (.
500 ha) was laden with scores or even
hundreds of well-developed seeds (Fig. 2B).
When we returned on April 22, the greater
part of the seed crop had already fallen,
apparently during a windy frontal system that
passed just prior. The remaining seeds detached readily, and synflorescences fractured
in even a light breeze. Within the flowering
[VOL. 135
area, all adult culms were already dead or
dying. Seedlings of Arundinaria gigantea recruited abundantly following the mass-flowering event. On hilltops 44 weeks after seed
drop, we observed approximately 12 new
seedlings within a 1 m radius of each dead
adult culm. Seedling density was strongly
affected by local topography, and density
was much higher down-slope, where seeds
were apparently washed by rain. Seedlings had
green leaves in late February, and many of
these were new, suggesting they were actively
photosynthesizing well before the forest overstory flushed spring leaves.
SEED AND SEEDLING EXPERIMENTS. A large
majority of A. gigantea seeds from Tunica
Hills proved viable in our germination trials.
Many seeds germinated within a week of
planting, and the large majority germinated
within three weeks of planting. When placed
on a moist paper towel at room temperature
immediately following seed harvest, 20 of 21
fully developed seeds (95.2%) germinated.
When planted into sterile potting mix 10 days
after seed harvest, 74 of 90 seeds unsorted by
size (82.2%) germinated. When planted into
sterile potting mix four weeks after seed
harvest, 88 of 100 fully developed seeds
(88.0%) germinated. These results and the
large number of seeds produced per synflorescence indicate high fertilization success
during the Tunica Hills flowering event.
In our field experiments, seed germination
was highly variable among plots. To count as
‘‘germinated’’ in our field trials at the Buckhorn WMA, seeds had to both be viable and
survive long enough in-place to germinate. In
three of the four treatment combinations, field
germination was distinctly bimodal. In half of
burned forest, unburned forest and unburned
blowdown plots, field germination rates exceeded 50% (sometimes substantially), while
germination in the other half was lower than
30% (in some instances much lower; Fig. 3). In
contrast, germination rates in all burned
blowdown plots were lower than 20% (often
much lower; Fig. 3). This apparent difference
between burned blowdown plots and the other
three types was not statistically significant
because the bimodality of germination rates in
the other three treatment combinations made
for high variances. For the same reason,
overall tests of windstorm effects and fire
effects were not significant. During our first
2008]
GAGNON AND PLATT: ECOLOGY OF GIANT CANE
313
FIG. 2. Cane reproduction and regeneration, including photos of: A) flowers of Arundinaria gigantea
with extruded anthers, B) seeds of A. gigantea, C) seedlings of A. gigantea 3 weeks after planting of seeds and
D) seedlings of A. gigantea 11 weeks after planting. Photo credits: Paul R. Gagnon
two censuses, we found evidence (i.e., the
stripped off bracts of missing seeds) that many
seeds had been pulled up and consumed,
probably by rodents and/or birds. Seeds might
also have been at risk of desiccation, especially
those planted into the large blowdown in full
sun. Of our 16 seedling plots, six accounted for
76% of total germination in the field, and the
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JOURNAL OF THE TORREY BOTANICAL SOCIETY
[VOL. 135
accounted for more than three-quarters (76%)
of remaining seedlings, and the one-quarter of
plots in the burned blowdown accounted for
only 6.5%.
FIG. 3. Proportion of cane seeds that germinated in each of four treatment combinations. Each
unfilled circle represents one plot composed of two
subplots, each with 32 seeds planted therein. Black
circles represent means of the four plots in each
treatment combination.
one-quarter of plots in the burned blowdown
accounted for only 7% of germination.
Seedling survival varied over time, but the
different post-disturbance environments had
no measurable effect on seedling survival. On
average, only 14% (6 7.1% SE) of seedlings
that germinated prior to our week-3 census
survived until our week-11 census, suggesting
that early establishment was a critical, highrisk stage for cane seedlings. In contrast, 44%
(6 15.2% SE) of seedlings alive at our week-11
census were still alive 33 weeks later for our
week-44 census, indicating a marked decline in
mortality once seedlings became established.
Statistical analyses of seedling survival indicated no significant differences among our
burned and unburned forest and blowdown
plots (Table 1). As with our field germination
trials, among-plot variation was high. However, seedling survival trials demonstrated no
suggestive trends among the treatment combinations. By the study’s end, three of 16 plots
Discussion. Our study demonstrates that
Arundinaria gigantea can reproduce successfully in conditions typical of partially-open
forests or canebrakes, but may face problems
in bare, burned-over areas. The hypothesis
that A. gigantea seeds and seedlings would do
best in post-disturbance environments was
unsupported by our results. Our disturbance
treatments had no measurable effect on seed
germination or seedling survival, and local
environmental factors appeared to dictate
seedling establishment more than the degree
of forest canopy closure. Because of variability
in seed survival, germination rates were quite
high in some plots and nil in others and
followed a bimodal distribution in three of our
four treatment combinations. We observed
that seed survival and germination was reduced on the bare mineral soil in burned
blowdown plots, but this effect was not
statistically significant because of high variability in all treatment combinations. Once
established, seedlings did have a statistically
higher rate of survival than either seeds or new
seedlings.
We observed that plots with the highest
rates of both seed germination and seedling
establishment shared two traits. First, they
received partial sunlight at ground-level. We
observed that seeds rarely survived and
germinated in full, intense sun. In contrast,
seeds in deep shade appeared to germinate
without problem, but we observed that seedlings most often failed to establish. Second,
successful plots had a layer of leaf-litter on the
ground. We suspect leaf litter facilitated
seedling establishment both by 1) hiding seeds
Table 1. Result of mixed model ANOVA with repeated measures for seedling survival in Arundinaria
gigantea. NDF 5 numerator degrees of freedom; DDF 5 denominator degrees of freedom based on
Kenward-Roger approximation.
Source of Variation:
Fixed effects on seedling survival
Stand type (forest or tornado blowdown)
Fire (previously burned or unburned)
Stand type 3 fire
Census
Census 3 stand type
Census 3 fire
Census 3 stand type 3 fire
NDF
DDF
F
P
1
1
1
1
1
1
1
11.1
11.1
11.1
14.1
14.1
14.1
14.1
1.20
0.61
0.19
8.46
2.04
0.39
1.24
0.2959
0.4511
0.6753
0.0114
0.1747
0.5448
0.2833
2008]
GAGNON AND PLATT: ECOLOGY OF GIANT CANE
from predators and 2) by moderating local
moisture regimes. The former (predator protection) agrees with a recent study by Kitzberger et al. (2007) in Chile who found 100%
mortality by seed predators for seeds of a
bamboo (Chusquea culeou E. Desvaux) exposed on the soil surface, but only 20% for
seeds beneath 1–3 cm of leaf litter. The latter
(moisture moderation) is consistent with a
study by Dattilo and Rhoades (2005) who
found that transplanted clumps of Arundinaria
gigantea grew significantly better when
mulched, which they attributed to reduced
desiccation in mulched soils.
Based on our field observations, three
potential bottlenecks appeared to affect Arundinaria gigantea regeneration. 1) First was
successful outcrossing. At our primary research site, A. gigantea flowered in isolated
patches every year but produced few seeds. In
contrast, the flowering event at our secondary
research site (Tunica Hills WMA) included
virtually all A. gigantea culms and multiple
genets over ca. 500 ha, and produced abundant well-developed seeds with high rates (82–
95%) of germination in our indoor trials.
Given their comparatively low rate of seed set
at our primary site, we suspect the few culms
flowering there may have been pollen-limited.
2) A second potential reproductive bottleneck
was seed survival. Based on our field observations, large A. gigantea seeds were consumed
by various seed predators, including at least
three different insects, and probably rodents
and/or birds. Of the few seeds produced at the
Buckhorn WMA, the majority appeared to fall
prey to insect seed predators before ever
detaching from synflorescences. There also
appeared to be a substantial risk of predation
to seeds planted in our field experiments.
Although we were unable to quantify this
effect, we saw evidence suggesting that seed
predation was one driver of variability in our
field germination trials. 3) A third potential
reproductive bottleneck was seedling establishment. Established seedlings had higher
rates of survival over time than either seeds
or very young seedlings. Mortality of newly
germinated seedlings was markedly higher
during the first 11 weeks after planting than
during the next 33 weeks (through the end of
the seedlings’ first winter).
Mass-flowering events in bamboos have
been a biological curiosity for decades (Janzen
1976). Based on our observations at two
315
Louisiana study sites, we propose that the
underlying driver of synchronized flowering
events in Arundinaria gigantea and perhaps
other bamboos is outcrossing success. Woody
bamboos are wind pollinated and have mechanisms to facilitate outcrossing (stamens are
exserted first, and stigmas appear after pollen
is shed; Judziewicz et al. 1999). This fact and
our observations of localized flowering events
both suggest that genets flowering individually
produce few viable seeds. Self-incompatibility
and constraints on outcrossing could be the
sole synchronizer of mass-flowering, or these
could operate together with predator satiation
(as per Janzen 1976) to select against out-ofphase individuals.
Our findings have several implications for
canebrake restoration efforts. First, for seeds
of Arundinaria gigantea planted directly at
restoration sites, a layer of leaf litter and
controls on seed-predators and herbivores
should improve seed survival and seedling
establishment. Second, seedlings can tolerate
and perhaps even thrive in moderate, though
perhaps not deep shade. Third, use of multiple
planting sites encompassing a range of conditions may be the best way to ensure successful
seedling establishment because among-site
variability in seed and seedling survival rates
could be high. Fourth, growing seedlings in a
greenhouse until one year of age may avoid
the bottlenecks of seed survival and seedling
establishment. Finally, it is possible that for
success in the long term, canebrake restoration
projects might need to include multiple genetic
individuals that flower in-phase.
Our study leaves many remaining questions
about the role of sexual reproduction in the
life history of Arundinaria gigantea that
subsequent studies might seek to address. First
is the degree to which A. gigantea is selfincompatible, which might be answered using
bagging experiments of synflorescences. Second is the proportion of viable seeds that
result from selfing vs. outcrossing, which
might be answered by genotyping. Third is
the predation rate of seeds still attached to
synflorescences, which might also be answered
using bagging experiments. Fourth is rate of
predation after seeds fall to the ground, which
might be answered using different predator
exclosures. Fifth are the factors controlling
seed germination, which might be answered in
part by plots with and without leaf litter and
by greenhouse studies at varying temperature,
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JOURNAL OF THE TORREY BOTANICAL SOCIETY
light, and moisture regimes. Sixth are the
factors controlling seedling establishment,
which might be answered in part by using
plots with/without leaf litter, plots in sun and
shade, and predator exclosures. Seventh are
the effects of different soils and moisture
regimes on seeds/seedlings, which might be
answered by monitoring soil composition and
moisture levels and by greenhouse experiments. We believe that a joint field- and
greenhouse-based approach might best answer
many of these remaining questions.
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