1

INTRODUCTION

In recent years, the generic name Arundinaria has become restricted in usage to the three
native ‘cane’ species of eastern North America: gigantea (= macrosperma), tecta and
appalachiana (Triplett et al. 2006, 2009, 2010). The closest living relatives of these bamboos
are in East Asia, where they are now classified into several distinct genera (Li et al. 2006,
Triplett and Clark 2010). The purpose of this paper is to summarize what is known,
superficially, about the biology of Arundinaria, as applied to problems in horticulture,
restoration and ecology.

Arundinaria has several unusual or unique characters, when compared to other native
plants of eastern North America. These characters are also typical of many bamboos in
temperate regions of East Asia. In flowering behavior, however, species of Arundinaria differ
from most of their long-lost East Asian cousins, which generally exhibit gregarious flowering
over many 100s or 1000s of acres or even whole regions, after non-flowering periods of several
decades. Flowering is generally rare and sporadic in Arundinaria, with no evidence of such
widespread gregarious events.

The following review is based on literature, meetings and conversations with growers,
plus 20 years of personal experience in Kentucky trying to grow and establish cane, especially
transplants into restoration sites and, more recently, seedlings. My associates at Roundstone
Native Seed Inc. (Bonnieville, Kentucky), John and Randy Seymour, have also become much
involved, and they are also working with Jeremy Hamlington and Mark Smith of Auburn
University in Alabama (School of Forestry and Wildlife Sciences). I do not deal here with
tissue culture and micro-propagation, which is being studied by Baldwin et al. (2009), Margaret
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Figure 1. Mason Brock helping the author to collect cane seed (Arundinaria gigantea) in June
2009. This is the only patch of cane known in Harrison County, Kentucky, growing along
Wornall Lane about 4 miles north of Griffith Woods. During 2009-2012, there have been
several flowerings of cane in central Kentucky.

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Cirtain (Dept. of Biological Sciences, Univ. of South Carolina, pers. comm.), Sharon Kester
(Dept. of Horticulture, Univ. of Kentucky, pers. comm.) and others. Moreover, Susanne Lucas
(Pioneer Plants LLC, Plymouth, Massachusetts), in partnership with Oprins Plant NV in
Belgium, is developing the market for mass-production of selected bamboo clones.

FLOWERING, SEEDING AND GERMINATION

Essential sources of information on flowering of Arundinaria are the many herbaria, with
dried specimens dating back to the earliest periods of botanical exploration (Campbell 1985). In
recent decades, there has also been some useful accumulation of records by the American
Bamboo Society (Betty Shor, La Jolla, California, pers. comm.), Brian Baldwin at Missississipi
State Univ. (Dept. of Plant and Soil Sciences, pers. comm.; Baldwin et al. 2009), and other
researchers (e.g. Marsh 1977, Gagnon and Platt 2008, Mathews et al. 2009). There appears to
be some tendency for more frequent flowering in some years in some regions, with clusters of
records covering 100s or 1000s of square miles. But even this clustering generally includes no
more than 1-10% of the plants within those regions. There may be weak association between
flowering frequency and years with sun-spot maxima and wetter periods, but deeper analysis is
needed (Figures 1 and 2).

There is virtually no definitive documentation of life-span for individual clones of

Arundinaria. A few horticultural observations indicate that as little as 3-15 years can
sometimes elapse between seed germination and flowering, but these plants are probably
aberrant individuals within seed lots (observations of myself, G. Cooper, G. Lundquist, and
others compiled by Betty Shor). Long-term observations of particular cane patches in the wild
suggest that the life-cycle is usually at least several decades. In an early account, Neisler (1860)
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Figure 2. Numbers of flowering records for Arundinaria spp. from the southeastern U.S.A.,
in relation to date and the sun-spot cycle. This is an updated version of the presentation in
Campbell (1985), but still a ‘work in progress’ that needs further analysis.
Records per year for flowering by Arundinaria spp. in SE U.S.A. Orange shows total records.
Red points to somewhat gregarious events at least in a region (higher points for widespread).
Yellow shows sun-spot maxima (to be investigated with climate etc.).
30
Next maximum
will be 2012-14

25

20

15

10

0
1870

1875

1880

1885

1890

1895

1900

1905

1910

1915

1920

1925

1930

1935

1940

1945

1950

1955

1960

1965

1970

1975

1980

1985

1990

1995

2000

2005

2010
Initial analysis (Campbell 1985) showed:
Flowering is about twice as frequent at sunspot maxima plus the two adjoining years, versus minima plus adjoining.
There is also a weak correlation with May+June+July precipitation.
But direct association between sunspots and precipitation (above/below mean) may be weak.

Effects of solar irradiance on climate are controversial, but seem significant over 3-15 year periods and longer.
J-L. Le Mouel, V. Courtillot, E. Blanter & M. Shnirman. 2008. Comptes Rendus Geoscience 340: 421-430.
J.L. Lean. 2010. WIRESs Climate Change 1: 111-122.

5
indicated that ca. 25 years was typical in gigantea, but longer periods are generally suspected
today. There are only two flowering records for appalachiana (1956 and 2006). Whole clones
of Arundinaria usually die after flowering and seeding but sometimes death is delayed for 1-3
years. Stephen Breyer (Tripple Brook Farm, Southampton, Mass., pers. comm.) has reported
recovery of gigantea without producing seed at a site well north of its native range.

Potential problems from inbreeding might occur if flowering is somehow decoupled from
regular gregarious behavior. Research of Franklin (2004) on a bamboo species in northern
Australia has shown much higher rates of cross-pollination, seed-set, and regeneration among
plants in peak flowering years, versus precocious or straggling plants. Baldwin et al. (2009, and
pers. comm.) suspect similar problems based on initial observations of low seed set in
Mississippi, with much more viability after artificial cross-pollination. However, I have got
good germination (ca. 80-100% of sound fresh seed) with several batches of seed that were
probably self-pollinated in Kentucky.

Bamboo seed generally dies after drying out at ca. 60-80oF (15-25oC) for 2-4 months, and
is thus considered ‘recalcitrant’ (McClure 1966, Stapleton 1987, 1994, Bellairs et al. 2008).
Drying in some sun for one day is often useful, in order to slow immediate germination and
reduce microbial attack, but several days may be damaging. Germination usually occurs within
a few days if seed are kept in a continually moist state after shedding. However, a few reports
indicate that germination by some East Asian species of colder zones may occur after 1-5 years
of ‘dormancy’ on the forest floor or similar storage (Qin 1985, Taylor & Qin 1988, Stapleton
1994, Wang et al. 2007). Stapleton stated: “Seed of the smaller subtropical and temperate
bamboos may have substantial dormancy, and it might germinate more quickly after a period of
cold pre-treatment, such as stratification or refrigeration at 5oC [41oF].”
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Figure 3. Decline of germination by seed stored under cold humid conditions. Six seeds lots (3
dates at 2 sites) were stored under various conditions; full explanation of methods and analysis
will be provided in a complete publication.
Decline of germination by seed stored in cold humid conditions:
green = Harrison Co. lots (H); red = Bourbon Co. lots (B).
Regression lines show declines for separate lots; the legend
shows equations and variance explained for pooled data.
100
Harrison Co. seed:
90 G = 89 - 0.24xD
H1 R-squared = 0.82 (n = 45)
80
G: germination percentage

H2 Bourbon Co. seed:

70 G = 40 - 0.12xD
H3 R-squared = 0.49 (n = 36)
60
B3
50

40

30 B1
20 B2
10

0
0 50 100 150 200 250 300 350 400
D: days after May 15th

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 Recent research on seed of Arundinaria by myself at the Univ. of Kentucky (in the lab of
Carol & Jerry Baskin) and Mississippi State University (Baldwin et al. 2009; D. Neal et al., in
prep.) has confirmed that even in cold storage at ca. 32-35oF [0-2oC], germination of seed
(when warmed-up and wetted for 14 days) gradually declines to zero after 1-2 years, with
considerable variation between seed lots (Figure 3). Zero germination is observed after seeds
drop below ca. 6-8% moisture content (Figure 4). To date, no freezing treatment has been
found to preserve viability for longer periods. But after cold damp storage, with or without
slight freezing, there was faster germination in some seed from Kentucky, as compared to cold
dry storage. After wetting and maintaining sound seed at ca. 70-80oF [25-30oC], germination
generally starts within a few days, and the first leaves appear at 1-4 weeks. Older seed tends to
be slower, with leaves sometimes not appearing until 2 or even 3 months. If seed does not start
to germinate within a few weeks, it is almost certainly dead—or perhaps fatally infected by
microbial growth.

Like other bamboos (Janzen 1976), cane seed is consumed by a wide range of pathogens,
pests and herbivores. For example, much fungus and other microbial growth often occurred in
petri dishes used for 14-day germination tests by myself. However, there was significantly less
microbial growth after seed had been stored in cold damp versus cold dry treatment; seeds lost
weight during cold damp storage, presumably exuding anti-microbial compounds. Sharon
Kester (pers. comm.) has had great difficulty extracting sterile material from seeds for tissue
culture. It is likely that cane seed in the wild is generally threatened with excessive drying and
microbial attack. Cane seed often does not mature due to attack of flowers by weevils, which
need to be identified. Due to such insects, several patches of flowering cane in Hart County
have generally failed to produce any mature seed within the past decade, before they die (R.
Seymour, pers. comm.). However, an initial flowering patch in 2000 did produce much seed,
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Figure 4. Relationship of germination to moisture content of seed, dried slowly in varied types
of storage. Data from cold damp treatments and frozen treatments are excluded. Full
explanation of methods and analysis will be provided in a complete publication.
Seed germination in relation to water content, excluding cold wet
treatments. Trend lines are for each collection lot (2 sites x 3 dates);
legend includes explained variance (R-squared) for pooled data
100

90
H2 H3
80
H1

70
Germination (%)

60

50
H = Harrison Co.
40 (R-squared = 0.61)

30 B = Bourbon Co.
B3 B1 (R-squared = 0.32)
20
B2
10

0
0 5 10 15 20 25 30 35 40
Water content (%)

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which was used by Cirtain et al. (2004, 2009) to grow seedlings for their experiments. Small
mammals are avid eaters of cane seed—it is usually essential to protect flats of germinating
seeds using wire mesh or other means.

GROWTH, PROPAGATION AND ENVIRONMENTAL FACTORS

`Based on general horticultural experience, bamboos in general are known to be

particularly sensitive to interruptions in moisture supply, poor aeration of roots, and other
physical stresses. For example, Stapleton (1987) found that division and repotting in hot dry
nurseries was risky, “thus it seems division of seedlings is only suited to cooler or more humid
nursery locations.” Thanks especially to experienced growers like Bill Hendricks (Kline
Nursery, Perry, Ohio), Ned Jacquith (Bamboo Garden, North Plains, Oregon) and Nevin Smith
(Suncrest Nurseries, Watsonville, California), it is possible to gain some general insights into
the best physical conditions for growing temperate species. From varied successes and failures,
it appears to me that stresses often cause plants to go into physiological ‘shock’—presumably
involving growth-suppressing hormones, and sometimes aggravated by microbial problems.
With return of good conditions, it can take up to a year for plants to resume rapid growth. In
containers, much more stress can occur with smaller sizes, especially if exposed to extremes of
temperature and moisture during daily or seasonal cycles. Study of hormones in bamboos is a
promising field that will eventually help in understanding of how these plants deal with stress
(e.g. Zhang et al. 2011).

The obvious unusual feature of Arundinaria, compared to associated plants in North

America, is its long spreading rhizome system, allowing clonal growth for 100-1000 meters or
more during its sexual life cycle. Brian Baldwin (pers. comm.) has evidence from DNA
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Figure 5. Survival of cane plantings at the ‘front-central’ field of Silver Lake Farm (Griffith
Woods), in relation to an index of cool aspect. Data from ca. 300 plants (mostly 10 year old
seedlings) are grouped here into seven vegetation types as the plotted observations, each the
mean of many plants. The index of cool aspect = [[difference of site aspect from 200o azimuth
(SSW)] × [slope in degrees] / 90]. Ongoing research indicates that growth (instead of survival),
shows a weak positive relationship with drainage.
R squared = 0.747; P = 0.012 (2 tailed Pearson)
70

60
Percent Survival in Cover Type

50
Initial results from cane plantation at
Griffith Woods: 2003/4 to 2006/7
40

30

20

10

0
3 3.5 4 4.5 5 5.5 6 6.5 7
Index of Cool Aspect
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markers that one clone of gigantea has spread over 1-2 km at Dahomey National Wildlife
Refuge in Mississippi. Another unusual feature is the concentration of extension growth by new
culms during just 1-2 months in the summer, usually when soils are still damp after spring
rains. It is likely that the extensive rhizome system allows rapid supply of moisture for such
growth. Moreover, on less well-drained ground, the air-canals in rhizomes of tecta (and
sometimes appalachiana) are presumed to enhance oxygen delivery for the rapid extension of
culms (McClure 1963, Triplett et al., 2006). Similar air-canals are known in some East Asian
species (e.g. Phyllostachys atrovaginata, heteroclada, nidularia). New culms are protected by
leathery sheaths, which are shed within a year or so, especially when upper nodes develop
branches. There is one dominant branch per node, like most temperate bamboos of eastern
China and Japan, but in marked contrast to most Sino-Himalayan and tropical bamboos.

Arundinaria is sometimes assumed to be a plant of wetlands—the USDA has misleadingly

listed it a “facultative wetland” species (Griffith et al. 2009). Although sometimes flooded for
short periods in the wild, these bamboos do not have optimal growth on saturated soils, but
tecta is more tolerant (Baldwin et al. 2009; with R. Jolley, M.C. Mills, and D. Russell, in
prep.). Like most bamboos, Arundinaria is moderately ‘mesophytic’—easily stressed by
droughts or floods during the growing season, especially if the rhizome system is reduced or
cut. Cirtain et al. (2004) found that gigantea seedlings on well-watered but well-drained soil
had more one-year growth (ca. 28 cm shoots) than periodically dried or periodically flooded
(ca. 20 cm shoots). However, transplants or cut rhizome sections do best when humidity is
maintained at high levels. Adam & Sue Turtle (Summertown, Tenn., pers. comm.) recommend
that during the growing season, transplanted bamboos in general should be regularly soaked
and kept in shade for a month or so before planting out. Baldwin et al. showed that the vascular
system of gigantea is sensitive to embolism (cavitation) when rhizomes cut. These researchers
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also recommend soaking root-rhizome systems of larger plants for a month or more, or
enclosing plants with leafy tops in plastic bags, before they are set out in the field.

With care, at least 50% success rate is expected with transplants of gigantea from existing
stands. The best season for transplanting is probably Feb-Mar, based on much experience in
Kentucky. When digging plugs from the wild, it is important to select 1-3 good culms for each
unit and to retain an approximate cylinder of at least 6-9 inches of soil—in depth and width—
together with the rhizome and root system. It is important to dig straight down with a long
heavy sharp spade (e.g. the King of Spades™ made by W.W. Manufacturing, Bridgeton, New
Jersey)—and to dig all around the culms, not angling down into the plug or otherwise reducing
or damaging the transplanted rhizome sections. However, even with much care, loose soil often
falls off roots, and tying plants up with burlap (or similar material) could be useful in some
contexts. To reduce transpiration, it is often important to cut off the top 30-70% of leafy
material when digging transplants, especially if plants are large and soil falls off. It is also
important to keep transplants cool and cover them with wet blankets (or similar material) when
transporting them, then settle them gently into their new homes within a day or so. And pray
for as much rain as possible, but without severe floods or winds, for the next few months.

Propagation from rhizome sections is somewhat erratic, but reliable methods can probably
be developed and have been actively sought in several studies (Sexton et al. 2003, Zaczek et al.
2003, 2009, Hartleb & Zaczek 2007, Brendecke and Zaczek 2008, Baldwin et al. 2009,
Schoonover et al. 2011). Based especially on Baldwin et al. (2009 and pers, comm.), the source
and initial condition of material can be a significant factor. Larger containers or trays with at
least 2-3 nodes are recommended, especially sections closer to culms of origin (proximal);
diameter of rhizomes appears to have little or no effect on success. A misting system can
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probably enhance success rates, but rooting hormones may not be particularly useful. Zaczek et
al. (2003) achieved more success when rhizomes were planted shallow, exposed to sunlight.
Jane Harrod (Jane’s Native Plants, Lawrenceburg, Ky., pers. comm.) and Bill Hendricks (pers.
comm.) have successfully grown many rhizome sections in early spring to summer by
enclosing them in humid poly-houses with root temperatures ca. 40-70o F and regular watering.
Chuck Rhodes (when at Dept. of Forestry, Univ. of Ky., pers. comm.) has observed reliable
rooting by floating rhizome sections in full sunlight. Paul Capiello (Yew Dell Garden,
Crestwood, Ky., pers. comm.) is currently comparing rhizome cuttings in fall versus spring,
since other bamboos have show more successful rooting in the fall. Rhizomes simply cut and
transplanted during the growing season tend to die much more readily than if some leafy tops
are retained, even if watered regularly. Containerized stock is clearly superior for field
plantings than just rhizome sections—but there is much potential for mortality of rhizomes in
the greenhouse while developing that stock (Schoonover et al. 2001).

Like many bamboos, Arundinaria can grow well in full sun, if soil conditions are suitable,
but some shade is tolerated well and may be beneficial for reducing temperatures and resulting
moisture stresses. Cirtain et al. (2009) found that gigantea—in growth chamber and woods—
did best in full sun, and there was a positive interaction with N level in their growth chamber.
Baldwin et al. (2009; and in prep.) grew gigantea with a shaded pot-in-pot system, regular
watering and NPK amendments; they found that total growth increased in full sun, but above-
ground growth alone was maximal under 60% shade. Smith (2011) has shown that gigantea is a
relatively light-demanding bamboo, compared to some of its East Asian relatives, including
smaller bamboos like Sasa species which did not increase photosynthetic rates when grown in
less shade. But she also showed that gigantea is relatively sensitive to moderate drought,
displaying signs of wilting or cavitation before any of the Asian species that were compared.
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Figures 6, 7 and 8. General illustration of typical habitats for the three species of Arundinaria.

Arundinaria tecta (photos from Virginia at dnr.virginia.gov) xeric

Upper left: deeper non-riverine swampy woods with oaks
(laurifolia, michauxii, pagoda) and relatively thin cane.
Upper center: transitional non-riverine pine-hardwoods and
denser cane. mesic
Upper right: more open longleaf pine woods, with frequent
seasonal drought and fire; cane grades into grass on the
ground.
Lower right: soils, red = strongly acid; green = average hydric

15
 At Roundstone Native Seed, we suspect that hot summer temperatures can be highly
detrimental, especially when black pots are exposed to full sun or within trays on greenhouse
benches. In several cases, we have observed much better survival and growth, above ground at
least, where containers remain below ca. 75-85o F (25-30oC)—experimental work is needed to
determine the exact response. For example, large freshly transplanted, potted plants did much
better when placed along the north side of a barn but still receiving skylight from above. Even
with drip-fed irrigation and a sunken pot-in-pot system, similar plants in full sun mostly died
above ground during 2011. In an experimental planting of 150 cane seedlings at Griffith Woods
(Harrison Co., Kentucky), survival after 3 years was correlated with an index of cool (N/NE-
facing) aspect (Figure 5).

Bamboos are generally considered nutrient-demanding plants (Lawson 1968, Lucas 2008).
Relationships of nutrient levels to Arundinaria—especially nitrogen (N)—have been studied in
a few, varied contexts. Cirtain et al. (2004, 2009) found that N amendment did not improve
growth of gigantea seedlings until after their first year. With transplants of gigantea into an old
field, Datillo & Rhoades (2005) found that fertilizer and manure both increased culm numbers
by ca. 10-40% after 2 years, but there was less effect on height. With transplanted rhizomes of
gigantea, Zaczek et al. (2010) found that NPK increased survival after 2 years, but it did not
offset short-term reductions of above-ground growth due to fire. Blattel et al. (2009) surveyed
soils across riparian buffers with native (unplanted) gigantea at three sites, and found 80-95%
decreases in nitrate from field to interior (downslope) soils at one site, in ammonium at another
site, and no significant trends at the third sites. Griffith et al. (2009) found that gigantea in
western North Carolina is associated with well-drained sandy soils, relatively low nutrient
levels, but low C:N ratios and pH of 5-6.6. A partner of Roundstone Native Seed grew
seedlings of gigantea in 2 × 2 × 5 inch cells on acid soils with unusually low Ca level, using
16
Figure 7 (see also Figures 6 and 8).

Arundinaria gigantea (Hart Co., KY)

Lower left: thin cane in maturing woods on
stream terrace; much flowered & died here.
Upper left: dense cane at crest of adjacent
stream bank.
Upper right: scattered brakes in transitions
to upland fields; somewhat short; some
flowered.
xeric
Right: soil
gradients,
green =
average pH, m
blue =
base-rich
association
hydric

17
standard medium for loblolly pine seedlings. These developed much less rhizome growth after
6 months, with virtually none escaping the containers. In better soil with the same cell size
(PRO-MIX™ plus clay and nutrient amendments), leafy shoot growth was similar after the
same period but several rhizomes usually appeared out of the bottom of each cell. There is a
need for broader experimental studies of growth under a range of nutrient conditions.

COMPETITORS AND CONSUMERS

Some observations, including mulching studies, may indicate effects of competing plants
with similar or shorter stature. In their field of gigantea transplants, Datillo & Rhoades (2005)
found that mulch (with or without extra nutrients) increased culm numbers by ca. 40-60% after
two years, but there was less effect on height. Using various manipulations, Cirtain (2009),
Hartleb and Zaczek (2007), Osland et al. (2009) and Schoonover et al. (2011) found that
gigantea transplants were not much reduced by dense competition in the ground vegetation,
such as Japanese grass (Microstegium vimineum). However, Baldwin et al. (in prep.) found that
after one year of experiments, transplanted rhizomes of gigantea grew much less among
rhizomatous alien grasses (Johnson grass and Bermuda grass) than among native deep-rooted
clumpers (big blue-stem and Indian grass). Reduction of this competition by tilling or
herbiciding increased cane growth among the alien grasses, but reduced it among the natives—
might these grasses have protected the cane from hot dry air? I found that establishment of
gigantea transplants was virtually all prevented by the densely rhizomatous quackgrass
(Elymus repens) at Griffith Woods, in Harrison County, Kentucky, but it was often partially
successful with much taller but thinner competition including ironweed (Vernonia gigantea)
and even poison hemlock (Conium maculatum). Tall associates may be beneficial in some
cases, by reducing hot sun and drying out of the soil surface (see previous section).
18
Figure 8 (see also Figures 6 and 7).

xeric
Arundinaria appalachiana
Upper left: typical habit, from Triplett et al. (2006, Sida).
+
Upper right: denser growth along upland roadsides,
Rt. 76 in northern GA (photo from Paul Schneider). m + +

Lower right: typical position along soil gradients;

red = associated species of strongly acid soils;
green = associated species of average soil pH. hydric

19
 There has been virtually no systematic study of consumer relations—herbivores, pests and
pathogens—but there have been varied initial anecdotal observations. In petri dishes used for
my germination tests with gigantea seed, fungal growth became severe in several cases, but
was much less after cold moist storage of seeds (unpublished data). In the greenhouse at
Roundstone Native Seed, gigantea seedlings suffered greatly from fungal infection of leaves
during the hot humid conditions of summer 2010. In my garden, a patch of cane grew to 10-20
feet across in a decade then gradually declined in the subsequent decade without any flowering,
and with no obvious reduction in light or other resources—I suspect fungal accumulation in the
plants, as evidenced by blackened twigs and leaves. A new genus of rust-like fungus has been
discovered on “tecta” in Alabama (Olive 1945); see also Hyde et al. (2002). Rabbits caused
repeated significant damage to my planting of 7-year old gigantea seedlings at Cane Run
(Fayette County) during 2000-2002, but the plants finally prospered. Mammalian herbivores in
general can have significant effects on cane. Cattle have often browsed it back in Kentucky,
and continuous grazing appears to kill the plants after a decade or so. On the uplands of central
Kentucky, most remaining cane has survived in old fencerows, wherefrom it locally recovers
into rights-of-way (especially along interstate highways) and other abandoned land that is no
longer grazed or mowed.

ECOLOGICAL NICHES, HABITATS AND RESTORATION

As outlined above, Arundinaria is generally typical of edges and other transitions from
deep woods to full sun, probably with repeated disturbance rather than a simple ‘successional’
niche after catastrophic disturbance. Within this broad zone, the three species have somewhat
distinct habitats along the gradient in moisture conditions (Figures 6, 7, 8): from subhydric
(tecta) to submesic (gigantea) to subxeric (appalachiana). In addition, gigantea tends to occur
20
on more base-rich soils, especially alluvial soils in the Mississippi Valley. The spread of
gigantea onto some calcareous uplands, such as the Bluegrass region of Kentucky, may be a
relatively recent phenomenon. Triplett et al. (2010) have not yet detected consistent genetic
differences between upland and lowland plants, but gigantea from the Mississippi Valley does
appears somewhat distinct in DNA from gigantea of Atlantic states. Hybridization does appear
to occur between gigantea and tecta—the geographic and ecological context of any
intergradation will deserve deeper study.

In nature, there has been little experimental work into what disturbance regimes are
optimal. Hughes et al. (1960, Hughes 1966) showed that burning or other intense disturbance at
intervals of about 10 years was probably optimal for tecta, and that the cane could be
successfully browsed by cattle in alternate years, especially during the winter. Gagnon & Platt
(2008a; Gagnon 2009) found that gigantea grew more in a blow-down area, compared to
deeper woods, and much more (× 2) with fire as well as blow-down; but growth was less with
fire alone. They also found (Gagnon & Platt 2008b) that sown seed did less well on bare burned
ground than with regular leafy litter. Zaczek et al. (2010) planted rows of gigantea rhizomes
then observed effects of prescribed fire, which increased culm density and rhizomatous spread
two years later but with reduced culm sizes and reduced overall leafy cover.

On relatively uniform base-rich soils, Figure 9 presents a conceptual model for the original
‘niche’ of gigantea within the dynamics of native woodland, based on much general
observation and historical data from the Bluegrass region of Kentucky. In addition to the
general gradient from deep shade to full sun (left to right), one can envisage an independent
gradient related to browsing by generalist herbivores. Before excessive human influence, large
animals such as giant bison and mastodons probably were significant browsers in woodland
21
with cane. Similar patterns do occur in modern vegetation, though they have been fragmented
and any ancient migrations are now of course lost. In the original woods, it is suggested that
there was a messy (highly stochastic) cyclical tendency, counterclockwise on the diagram. This
concept is allied with Vera’s (2000) hypothesis concerning the ancient role of herbivores in the
woodlands of central Europe.

Focus on cane in conservation deserves much more effort, given this plant’s historical
abundance in some regions, its potential role to counter invasive alien shrubs (Osland et al.
2009; David Brand, Warren Wilson College, 2010 abstract), its potential role in nutrient uptake
and reducing erosion, especially along riparian zones and headwater streams (Schoonover et al.
2005, 2011), and its potential role as a perennial forage for wildlife (McHargue 1941, Platt et
al. 2001) or even livestock in some contexts (Biswell 1941, Hughes et al. 1960, Smart et al.
1960, Halvorson et al. 2010).

Griffith Woods (Silver Lake Farm), covering 750 acres in Harrison County, provides an
excellent site for deeper study of cane in the Bluegrass region of Kentucky. In addition to
patches of ancient woodland, including the world champion chinquapin oak, there are many old
fields where cane can be established. Several general goals can be set: (1) genetic collection,
with varied uses; (2) comparative study of growth rates and responses to site types; (3)
experimental management—browsing, burning, cutting, chemical, competition, other
consumers; (4) studying effects of cane (compared to other vegetation) on soil, plants, animals,
etc. Now transferred from The Nature Conservancy and University of Kentucky to the
Kentucky Department of Fish and Wildlife Resources, this old farm should become the center
for restoration of Bluegrass woodlands, their canebrakes and their wildlife, together with
associated research and education.
22
Figure 9.

23
 DISCUSSION

So what generally characterizes the growth form, life cycle and ecology of temperate
bamboos, and does Arundinaria have any special distinction? Bamboos in general have a
unique ‘punching and branching’ ability to send up rapidly growing culms through brush and
vines in transitions from shady woodland to more open vegetation. After escaping herbivory in
tender young stages, and growing through any competing thickets, culms then branch out into
spaces above. In species with running rhizomes, large areas can be colonized, especially on
gentle uniform slopes and plains without excessive droughts or floods. Disturbances of varied
kind—from tree-falls to wild fires, when repeated at intervals of ca. 5-25 years, generally
provide the optimal habitat for bamboos.

Arundinaria is similar in these respects to its East Asian cousins, with a somewhat similar
range of habitats from low, seasonally damp plains to drier, broad ridges in the mountains. The
loss of large canebrakes from more fertile lowland plains and some calcareous uplands presents
a significant problem for conservation and restoration, since the plant has not yet been
propagated in large numbers (Platt and Brantley 1997, Stewart 2007). East Asian people have
cultivated many species of temperate bamboo for millenia, but the many Native American uses
of cane were interrupted (Platt et al. 2009). There was little initial adoption of these plants by
the settlers from Europe or even by their slaves from Africa, except for some local uses as
fishing-poles, bean-stalks and the like.

Bamboos in general, especially running temperate species, tend to develop dense

competitive stands that can generally prevent seedling survival except after parental death. As
previously discussed (Campbell 1985), their ‘monocarpy’ (death after flowering) could have
24
been selected partly by such need for parental death, assuming that occasional sexual
reproduction is essential. Other evolutionary forces favoring infrequent or gregarious flowering
could include selection for ‘satiation’ of seed consumers when large seed crops are produced
(Janzen 1976), and selection for association with particular phases of environmental cycles—
perhaps allowing seedlings to renew the population in rainy periods after parents decline in
drier periods, or after fire (Keeley and Bond 1999). The rare flowering, poor dispersal, and
frequent self-pollination of bamboos pose special problems for maintaining genetic diversity.

The much reduced extent of observed gregarious flowering in Arundinaria, compared to

its East Asian cousins, suggests possible decoupling of a more regular ancestral life-cycle from
environmental cues. Such decoupling might have developed as climatic patterns became less
predictable during the Quaternary era, including severe disruptions during glacial periods.
Problems for cross-pollination and genetic conservation may be particularly acute in these
bamboos. Nevertheless, I am confident that more concentrated attention by horticulturalists and
biologists can refocus North American effort on this worthy cause (Figure 10).

ACKNOWLEDGEMENTS

Many people have directly or indirectly helped me to grow and study cane—or other
bamboos—over the past thirty years. These include Carol & Jerry Baskin, Frank & Melinda
Boyer, Adam Datillo, Ned Jacquith, Li De-zhu, Susanne Lucas, Will Overbeck, Tim Phillips,
Layton Register, Chuck Rhoades, Qin Ze-sheng, John and Randy Seymour, Nevin Smith, Chris
Stapleton, Adam & Sue Turtle, and Josh Wysor. Four other special people also provided initial
inspiration at local and global levels, but are now, sadly, deceased: Peter Addington, David
McClintock, Willem Meijer, Tom Soderstrom and Xiao Xue (son of Xue J.R.).
25
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Figure 10 (next page). The cane planting along Cane Run on north side of Lexington. In the
winter of 1999/2000, 100-150 seven-year old seedlings were planted here on the old university
farm, part of the first ‘Reforest the Bluegrass’ event of Lexington Fayette Urban Couinty
Government. Plants were mulched and manured initially, suffered much from rabbits in 2002-
2003, then exploded in growth during 2005-2010.

31
 Layton Register is 6 ft. tall This was first
‘Reforest-the-
Bluegrass’
site of LFUCG

Along Cane Run on north side of Lexington (by Citation Blvd. bridge): 100 seedlings planted in winter of
1999/2000, now covering an acre in remnant of old woodland with large bur oaks, blue ash, rue anemone

32