Arundinaria gigantea



INTRODUCTORY


  Ted Bodner. Southern Weed Society. www.forestryimages.org
AUTHORSHIP AND CITATION:
Taylor, Jane E. 2006. Arundinaria gigantea. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].

FEIS ABBREVIATION:
ARUGIG

SYNONYMS:
Species synonyms―
Arundinaria macrosperma (Michx.) [8,18,38,39,113]
Arundo gigantea Walter [18,64]
   = Arundinaria gigantea (Walt.) Muhl. [38,39,48,58,64,97,110,114]

Subspecies synonyms―
A. tecta (Walt.) Muhl. [38,39,48,51,77,114]
Arundo tecta Walt. [18,58,64,101]
   = Arundinaria gigantea (Walt.) Muhl. subsp. tecta (Walt.) McClure [16,42,58,114]

A. gigantea (Walt.) Muhl ssp. macrosperma (Michx.) McClure [16,42,64]
   = A. gigantea (Walt.) Muhl. subsp. gigantea [58,64,114]

NRCS PLANT CODE [100]:
ARGI
ARGIG
ARGIT8

COMMON NAMES:
cane
giant cane
switch cane

TAXONOMY:
The scientific name of cane is Arundinaria gigantea (Walt.) Muhl. (Poaceae). Some systematists recognize 2 subspecies of cane [58,64,114]:

A. gigantea subsp. gigantea (Walt.) Muhl., giant cane
A. gigantea subsp. tecta (Walt.) McClure, switch cane

The taxonomy of Arundinaria species in the United States has been confusing and poorly understood. Switchcane and giant cane are sometimes considered distinct species [8,18,38,39,46,48,51]. Plant height and the height, position of the seed heads, and rhizome structure (air canals) are sometimes used to differentiate the infrataxa of cane in the United States. However, plants often exhibit wide ranges in height growth on different sites, and flower and seed production tends to be sporadic or rare. The presence or absence of air canals in the rhizomes is another primary factor used to differentiate between subspecies; however, this criterion is also somewhat inconclusive [8,46,64]. Hughes [53] commented on the difficulty of cane taxonomy by saying, "it seems that the criteria used to differentiate A. gigantea from A. tecta are of questionable validity". Gilly [38] was 1 of the early taxonomists to suggest that only 1 species of Arundinaria was native to North America. In much of the literature, both A. gigantea and A. gigantea subsp. gigantea are called "giant cane", and A. gigantea subsp. tecta is usually referred to as "switch cane". For this review, the species in general is called cane, A. gigantea subsp. gigantea is called giant cane, and A. gigantea subsp. tecta is called switch cane.

LIFE FORM:
Graminoid

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
Information on state-level protected status of plants in the United States is available at Plants Database.

DISTRIBUTION AND OCCURRENCE

SPECIES: Arundinaria gigantea
GENERAL DISTRIBUTION:
Cane occurs from southern New York south to central Florida and west to Texas, Oklahoma, Kansas, Missouri, and Illinois [18,58]. Plants Database provides a distributional map of cane and its infrataxa.

Infrataxa: Giant cane has a distribution similar to cane in general, but does not occur in New York. Switch cane has a distribution similar to that of cane throughout the Atlantic and Southern Coastal Plains, but it does not occur in Delaware, Illinois, Indiana, Kansas, Kentucky, Missouri, Ohio, Texas, or West Virginia [100].

ECOSYSTEMS [35]:
FRES12 Longleaf-slash pine
FRES13 Loblolly-shortleaf pine
FRES14 Oak-pine
FRES15 Oak-hickory
FRES16 Oak-gum-cypress
FRES17 Elm-ash-cottonwood
FRES41 Wet grasslands

STATES/PROVINCES: (key to state/province abbreviations)
UNITED STATES

AL AR DE FL GA IL IN KS KY LA
MD MS MO NJ NY NC OH OK SC TN
TX VA WV              

BLM PHYSIOGRAPHIC REGIONS [3]:
None

KUCHLER [61] PLANT ASSOCIATIONS:
K079 Palmetto prairie
K089 Black Belt
K090 Live oak-sea oats
K099 Maple-basswood forest
K100 Oak-hickory forest
K101 Elm-ash forest
K102 Beech-maple forest
K103 Mixed mesophytic forest
K104 Appalachian oak forest
K109 Transition between K104 and K106
K111 Oak-hickory-pine
K112 Southern mixed forest
K113 Southern floodplain forest
K114 Pocosin

SAF COVER TYPES [30]:
23 Eastern hemlock
26 Sugar maple-basswood
27 Sugar maple
53 White oak
57 Yellow-poplar
59 Yellow-poplar-white oak-northern red oak
60 Beech-sugar maple
61 River birch-sycamore
70 Longleaf pine
71 Longleaf pine-scrub oak
72 Southern scrub oak
74 Cabbage palmetto
75 Shortleaf pine
76 Shortleaf pine-oak
80 Loblolly pine-shortleaf pine
81 Loblolly pine
82 Loblolly pine-hardwood
83 Longleaf pine-slash pine
84 Slash pine
85 Slash pine-hardwood
87 Sweetgum-yellow-poplar
88 Willow oak-water oak-diamondleaf (laurel) oak
89 Live oak
91 Swamp chestnut oak-cherrybark oak
92 Sweetgum-willow oak
93 Sugarberry-American elm-green ash
94 Sycamore-sweetgum-American elm
96 Overcup oak-water hickory
97 Atlantic white-cedar
98 Pond pine
100 Pondcypress
101 Baldcypress
102 Baldcypress-tupelo
103 Water tupelo-swamp tupelo
104 Sweetbay-swamp tupelo-redbay
110 Black oak

SRM (RANGELAND) COVER TYPES [93]:
805 Riparian
809 Mixed hardwood and pine
812 North Florida flatwoods
813 Cutthroat seeps
814 Cabbage palm flatwoods
815 Upland hardwood hammocks
816 Cabbage palm hammocks
817 Oak hammocks
821 Pitcher plant bogs
822 Slough

HABITAT TYPES AND PLANT COMMUNITIES:
Extensive monotypic stands of cane known as canebrakes were a dominant landscape feature in the southeastern United States at the time of European settlement. Historical accounts indicated that hundreds of thousands of acres were characterized by this ecosystem. Canebrakes disappeared rapidly following European settlement because of a combination of overgrazing, altered burning regimes, and agricultural land clearing [7,73,74,88]. It is estimated that there has been a 98% decline in canebrakes communities [14,69]. Today cane exists as an important understory component in a variety of deciduous and evergreen forest and shrub types.

Schafale and Weakley [84] describe 2 plant communities in the wet pine flatwood forests of North Carolina in which cane is a codominant: longleaf pine (Pinus palustris)/cane and loblolly pine (P. taeda)/cane. These communities are similar in composition with a sparse canopy of pines and a mid-story dominated by cane. The understory is typically a mixture of shrubs, including inkberry (Ilex glabra), creeping blueberry (Vaccinium crassifolium), wax myrtle (Morella cerifera), and blue huckleberry (Gaylussacia frondosa); and grasses, including pineland threeawn (Aristida stricta), cutover muhly (Muhlenbergia expansa), little bluestem (Schizachyrium scoparium), and toothache grass (Ctenium aromaticum).

A cane shrubland alliance occurs on floodplains and alluvial soils in eastern Oklahoma [52]. Common associates in this alliance include boxelder (Acer negundo), river birch (Betula nigra), smallspike false nettle (Boehmeria cylindrica), jewelweed (Impatiens capensis), northern spicebush (Lindera benzoin), and eastern poison-ivy (Toxicodendron radicans).

Kologski [60] describes a longleaf pine/cane community type in the Green Swamp of the North Carolina coastal plain. This type is described as a wetter pine savanna community.

In Missouri cane is a component of the swamp chestnut oak (Quercus michauxii)-Shumard's oak (Q. shumardii)- sweetgum (Liquidambar styraciflua)/cane mesic floodplain forest alliance [86].

Switch cane: Glitzenstein and others [40] describe a "globally rare" woodland association in South Carolina of longleaf pine-switch cane-sweetgum-bushy bluestem (Andropogon glomeratus)-hooded pitcher plant (Sarracenia minor). In this association switch cane and bushy bluestem usually comprise the majority of the plant cover, and the tree canopy cover is generally less than 10% [40].

A pond pine (P. serotina)/switchcane forest type occurs in the North Carolina coastal plain where the pine overstory is typically scattered and inkberry is an abundant shrub [54].

In addition to the plant communities discussed above, where cane is a dominant or codominant, there are a variety of other communities in which cane occurs in various levels of importance. Publications that discuss plant communities in which cane and switch may occur are listed below. The list is neither restrictive nor all inclusive.

AL: FL: GA: LA: MS: NC: OK: SC: VA: Atlantic and Gulf Coastal Plains:

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Arundinaria gigantea
GENERAL BOTANICAL CHARACTERISTICS:
This description provides characteristics that may be relevant to fire ecology, and is not meant for identification. Keys for identification of cane are available (e.g. [48,64,77,97]).

Cane is a native, perennial, evergreen grass that grows to a height of 6.6 to 32.8 feet (2-10 m). The coarse stems are round and hollow, 0.7 to 3 inches (2-7.6 cm) thick, and generally survive for about 10 years. Leaves range from 3.9 to 11.8 inches (10-30 cm) in length and from 0.8 to 1.6 inches (2-4 cm) wide. The flowers are racemes or simple panicles with several spikelets 1.6 to 2.8 inches (4-7 cm) long and 0.3 inch (8 mm) wide. The fruit is a caryopsis, 0.3 inch (8 mm) long and 0.1 inch (3 mm) wide. Cane forms an extensive system of tough, thick rhizomes [18,48,51,64,77]. Rhizomes vary in size but rarely are larger than 0.75 inch (1.9 cm) in diameter [55].

Physiology:
Flooding―Cane has high flood tolerance and is well adapted to waterlogged soils and frequently flooded sites [14,18,62].

RAUNKIAER [78] LIFE FORM:
Phanerophyte
Geophyte

REGENERATION PROCESSES:
Cane primarily reproduces vegetatively from rhizomes [18,62,64]. Seed is produced sporadically [88], and cane regeneration from seed is considered rare [54].

Pollination: Cane is pollinated by wind [57].

Breeding system: Cane is monoecious [64].

Seed production: Flowering and subsequent seed production in cane are sporadic, infrequent, and unpredictable [18,88,91]. Cane may flower in response to burning [49,63,88]. Hughes [54] reported that seedlings that establish following fire rarely develop into mature plants. Additional information on postfire seedling establishment is lacking, and more studies are needed.

Seed dispersal: Seeds fall or are shaken to the ground beneath the parent plant [53]. Although there is evidence that some birds and small mammals will feed on cane seed [74], no information is available on the role of animal dissemination in seed dispersal.

Seed banking: Cohen and others [17] examined the seed banks in longleaf pine, slash pine, and loblolly pine stands in North Carolina. Likewise, Schneider and Sharitz [85] examined the seed banks in floodplain hardwood forest communities of the Savannah River in South Carolina. No evidence of a cane seed bank was found in either study.

Germination: Germination information specific to seeds of cane is limited; however, it is reported that cane seed is characterized by low viability [53,88,115]. When seed is produced, much of it may be destroyed by insect predation [53].

Seedling establishment/growth: Cane seedlings develop quite slowly. Observations of natural seedlings in a canebrake in North Carolina found that 3-year-old seedlings were less than 1 foot (30.5 cm) tall. Present-day cane stands rarely regenerate from seed under natural conditions [55].

Asexual regeneration: Cane sprouts prolifically from rhizomes, and aboveground stem regeneration is rapid following disturbance [18,53,74]. Historically, fire has been chiefly responsible for stimulating cane regeneration from sprouts [21]. Good stand recovery will not occur if the rhizome and root systems are not healthy and vigorous and if there are not ample food reserves to support new shoots. The vigor of cane stands tends to decline over time and is especially harmed by heavy summer grazing of cattle and the rooting of wild and/or domestic swine [18,19,54,56,62,91].

SITE CHARACTERISTICS:
Cane inhabits low-lying, moist to wet sites, including low woodlands of various mixtures, woodlands on mesic and sub-mesic slopes and uplands, river and stream banks, floodplains, levees, shrub-tree bogs and bays, swamplands, sloughs, bayous and pocosins, and mesic to wet savannahs [18,41,42,84,104,109]. Cane will grow on xeric and sub-xeric sites, but it thrives best on wetter sites that are typically seasonally flooded or saturated [75]. The water level often remains at or near the soil surface for extended periods during the wet season but falls well below the soil surface later in the growing season [84,91,102].

Although cane thrives best on well-drained loams or silt loams [43,90], it grows in a variety of soil types ranging from clay to sand and has a wide tolerance of soil nutrient conditions [14]. Soils are often poorly drained, highly acidic, and organic, peaty, or mucky [84,92,105]. On some sites, sandy surface soils overlie loamy or clayey subsoils. The heavier subsoil tends to retain moisture and nutrients during dry periods [6,40]. Cane has been observed growing on sandy soils with a mildly alkaline pH of 7.8 [31].

Cane is found at elevations ranging from sea level in southern floodplains to 2,000 feet (610 m) in the Appalachian Mountains. It has a broad climate tolerance and can withstand temperatures ranging from -9.4 to 106 F (-23 to 41 C) [18].

SUCCESSIONAL STATUS:
The canebrake community is fire dependent [34,40]. Historically, fire probably maintained canebrakes in a secondary successional sere [7]. Cane sprouts so prolifically following fire that it quickly achieves dominance after a burn, and the dense thickets suppress the growth of other vegetation for many years [32]. In the Southeast, canebrakes can form an ecotone transitional between savannas and wetlands such as pocosin, bay-gall, bay forest, or swamp forest. With different fire frequencies, canebrakes may alternate with these types over time. In the pine pocosins and shrub bogs of the Atlantic Coastal Plain, fire maintains cane dominance over evergreen shrubs such as inkberry, swamp titi, sweetbay, and redbay. Frequent fire can eliminate cane and favor a transition to a grass-sedge bog community. In the absence of fire, cane is gradually replaced by shrubs and trees [21,33,60,74,89,105,106]. Canebrakes succeed to multistoried wooded communities such as bottomland hardwood, pocosin, pond pine forest, red maple forest, and bay forest [34].

Cane is fairly shade tolerant. It thrives best in the open or under light tree cover, but can persist under dense canopies of up to 80% cover [27,54,62]. The ability of cane to survive under tree cover allows it to expand readily if the trees are removed [32]. For example, a cane stand expanded "readily" following logging of the tree overstory in blackgum and Atlantic white-cedar swamps in the Great Dismal Swamp, North Carolina [71].

Cane does not spread rapidly into either early or late successional forest types. It is hypothesized that cane was formerly concentrated in ecotones, between frequently disturbed areas and less disturbed forests of sugar maples, hickories, ashes, and oaks. The ecotonal vegetation may have been relatively stable, being maintained by small-scale oscillations of forest boundaries rather than long-term directional succession [9].

SEASONAL DEVELOPMENT:
Foliage production occurs from April to early July, and green foliage is held well into winter and even until the following spring on protected sites [91]. Flowering in cane is rare, but it may occur from March to May in Florida [16,48] and from April to July in the northern extent of its range [77,110]. The flowering period may continue for a year [51]. Seeds mature about 1 month following flowering, and seed germination may occur within a few days of the seed reaching the ground [53,55]. Aerial stems, and the rhizomes attached to them, die after flowering [42,53,64,77]. New stems arise from rhizomes from spring to mid-summer, and have been observed to elongate as much as 1.5 inches (3.8 cm) in 24 hours [53]. Stands usually decline in 3 to 4 years because of gradual mortality and replacement [55].

FIRE ECOLOGY

SPECIES: Arundinaria gigantea
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: Cane is adapted to fire by sprouting quickly and prolifically from rhizomes [18,53,74].

Fire regimes: Canebrakes are fire-dependent ecosystems [34,40]. Prior to European settlement, fire was the primary factor that maintained monotypic canebrakes on hundreds of thousands of acres across the mid-Atlantic and southeastern U.S. It is estimated that the historical fire frequency of canebrakes in the southeastern U.S. ranged from 2 to 8 years [34]. The dense growth creates heavy fuel loads and makes canebrakes highly flammable [55,91]. Canebrakes on peatlands historically experienced landscape-scale fires that burned for weeks or months, creeping through swamps, smoldering in peat, and flaring up when flammable vegetation was reached or when conditions of humidity and wind reached critical thresholds [32]. In canebrakes of bottomland hardwood ecosystems, fire intensity in the cane stands was much higher than in the adjacent hardwood forest, although the fire severity was low except during drought. Large fires only occurred after an extended drought, usually a dry fall followed by a dry spring [103].

Fire regimes in the various woodland and shrubland communities where cane may occur can be variable. The southern pine forests and pine savannas typically have fire return intervals of less than 10 years [68,103]. In the southeastern U.S., adiabatic thunderstorms can occur almost daily during the summer, and this region has 1 of the highest annual lightning frequencies in the world [68,75]. Although the number of lightning fires is highest from June to August, the majority of acreage burns in May and June in Florida and south Georgia, when the time between thunderstorms is longer. In the late summer, thunderstorms and associated rainfall are more frequent and humidities are higher. Historically, fires associated with dry frontal systems probably were quite large and may have burned for weeks or months, particularly in organic/peaty soil. Such fires likely spread into adjacent upland communities. The historic high fire frequency resulted in a frequent low-severity fire regime. Exceptions occurred when catastrophic events, such as hurricanes, tornados, and severe drought, were precursors to fires of much higher intensity and severity [75,103].

Pond pine pocosins burn on a 20- to 50-year cycle, but on highly productive sites, fire-return intervals of 3 to 10 years can be common. The shorter interval fires may produce a pine savanna with a grass understory. Mesic sites have a shrub layer comprised of many ericaceous evergreen shrubs that tend to burn intensely, resulting in the top-kill or death of all vegetation except pond pine. Pond pine has the ability to sprout from its base as well as along its stem and branches; thus, its aboveground stem survives higher severity fires than stems of most other pine species. This trait allows the species to dominate wet areas such as pocosins. Summer fires during severe droughts can eliminate pond pine and cane, because the underlying organic soil burns, destroying root systems [103].

Cane grows in hardwood communities with a wide range of fire frequencies, from the short return interval of 3 to 8 years for chestnut oak, to the moderate-return intervals of 35 years for yellow-poplar and oak-hickory communities, and the 1000+ years possible for some maple, beech, and birch communities. On bottomland hardwood sites, low-severity fires are the norm because fuel loads are generally light due to rapid decomposition on these moist, humid sites. Insect- and disease-related mortality and windthrow can result in heavy loadings of large woody fuels which, in times of drought, will support stand-replacement fires [103].

Evergreen bay forests of loblolly bay, sweetbay, and redbay are characterized by a stand replacement fire regime. This type now burns on about a 20- to 100-year cycle, but the historic fire frequency is not well documented [65]. Shrub bogs are bay forests that burn every 2 to 5 decades. More frequent burning, at least once a decade, removes the shrub layer, resulting in an herb bog. If the underlying organic soils are completely consumed, both pocosins and bays will revert to marsh [103].

Before European settlers harvested Atlantic white-cedar, it was generally perpetuated by major disturbances, probably stand-replacing crown fires that occurred at 25 to 300 year intervals [103].

Embedded within pine and floodplain hardwood ecosystems are numerous other ecosystems such as depressional wetlands, including bays, lime sinks, cypress ponds and savannas, gum ponds, bay swamps, pitcher plant bogs, shrub bogs, and spring seeps. Fires in these wetland communities are typically stand-replacing. Fire return intervals can be variable: 3 to 9 years in herb bogs and shrub bogs; 20 to 30 years in gum ponds and bog swamps; 20 to 50 years in titi shrub bogs, and 20 to 150 years in many cypress ponds and bay swamps [103].

Wet grassland ecosystems are characterized by a presettlement fire frequency of 1 to 3 years. These ecosystems typically contain large quantities of herbaceous vegetation and are considered highly flammable. The coastal grassland landscapes are often quite extensive, a factor that aids in the propagation of an individual fire. Depending on fuel and wind speeds, fire may either bridge small to moderate-sized natural breaks, such as stream channels, or be stopped by them [103]. Lightning-strike fires are common in coastal wetlands, and often fire from adjacent uplands can spread into the wetlands [32].

The following table provides fire return intervals for plant communities and ecosystems where cane is important. For further information, see the FEIS review of the dominant species listed below.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
maple-beech Acer-Fagus spp. 684-1,385 [15,103]
maple-beech-birch Acer-Fagus-Betula spp. >1,000
sugar maple Acer saccharum >1,000
sugar maple-basswood Acer saccharum-Tilia americana >1,000
sugarberry-America elm-green ash Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica <35 to 200
Atlantic white-cedar Chamaecyparis thyoides 35 to >200
beech-sugar maple Fagus spp.-Acer saccharum >1,000 [103]
green ash Fraxinus pennsylvanica <35 to >300 [28,103]
yellow-poplar Liriodendron tulipifera <35
shortleaf pine Pinus echinata 2-15
shortleaf pine-oak Pinus echinata-Quercus spp. <10
slash pine Pinus elliottii 3-8
slash pine-hardwood Pinus elliottii-variable <35 [103]
longleaf-slash pine Pinus palustris-P. elliottii 1-4 [68,103]
longleaf pine-scrub oak Pinus palustris-Quercus spp. 6-10
pocosin Pinus serotina 3-8
pond pine Pinus serotina 3-8
loblolly pine Pinus taeda 3-8
loblolly-shortleaf pine Pinus taeda-P. echinata 10 to <35
sycamore-sweetgum-American elm Platanus occidentalis-Liquidambar styraciflua-Ulmus americana <35 to 200
oak-hickory Quercus-Carya spp. <35 [103]
oak-gum-cypress Quercus-Nyssa-spp.-Taxodium distichum 35 to >200 [68]
southeastern oak-pine Quercus-Pinus spp. <10
white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra <35
chestnut oak Quercus prinus 3-8
black oak Quercus velutina <35
live oak Quercus virginiana 10 to <100 [103]
cabbage palmetto-slash pine Sabal palmetto-Pinus elliottii <10 [68,103]
baldcypress Taxodium distichum var. distichum 100 to >300
pondcypress Taxodium distichum var. nutans <35 [68]

POSTFIRE REGENERATION STRATEGY [96]:
Rhizomatous herb, rhizome in soil
Surface rhizome/chamaephytic root crown in organic mantle or on soil surface

FIRE EFFECTS

SPECIES: Arundinaria gigantea
IMMEDIATE FIRE EFFECT ON PLANT:
The coarse stems and leaves of cane are readily killed by fire, but the rhizomes usually survive [55].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
Although fire will kill all aboveground plant parts, it maintains cane stands by stimulating the production of new sprouts and eliminating other vegetation that would compete with the sprouts for water and nutrients [55].

PLANT RESPONSE TO FIRE:
Fire stimulates cane sprouting [21]. Cane will start to sprout soon following a spring burn, and stem density may return to prefire levels by mid-summer of the same season [91]. Following a winter fire in North Carolina, cane stems grew as much as 1.5 inches in 24 hours in the following spring [55].>

Cane may flower in response to burning [49,63,88]. Seedlings occasionally establish after a fire, but the seedlings rarely develop into full-sized plants [54].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Cane stands tend to remain even-aged for 2 to 3 years following fire with many sprouts emerging within the 1st year, and few new shoots in the 2nd and 3rd years. Thereafter, new sprouts again start to appear, and the stands become uneven-aged [54].

Cane may not respond to burning if overall stand vigor is extremely poor. If cane stands of low vigor are burned, other plant species may regenerate more quickly, and the cane may never recover [54].

A spring prescribed burn promoted cane in a pond pine/cane community in the North Carolina coastal plain. On sites without tree cover, cane stem numbers increased 88% in the first year following the burn. On sites with pond pine tree cover, cane stem numbers increased 40% in the first year [54].

Fire favored switch cane in longleaf and loblolly pine communities in the South Carolina coastal plain. Prescribed burns were carried out in the winter over a 12-year period at intervals of 1, 2, 3, and 4 years. Prior to burning, the understory was predominantly shrubs, with a minor to moderate component of switch cane. Burning resulted in a general conversion of the understory from shrubs to grasses, primarily switch cane [82].

In the absence of fire, cane stands lose vigor, culms die, and succession by other plant species exceeds the rate of cane regeneration. In 1 study in a pond pine/cane forest in the coastal plain of North Carolina, cane stem density started to decline 10 years after a spring wildfire. From 10 to 13 years after fire, cane stem numbers declined 50%, and by year 14, there was a 65% reduction in density [54].

Repeated annual or semi-annual fires are detrimental to cane stands because the continuous removal of the stems and leaves depletes food reserves in the rhizomes, and new sprouts cannot be produced [7,56,106].

FIRE MANAGEMENT CONSIDERATIONS:
Prescribed fire can be used to renovate decadent cane stands. Hughes [54] recommends prescribed fire at 10-year-intervals to increase cane density. Leithead and others [62] caution that burns should not be conducted any more often than every 3 to 4 years. Low-severity fires limited to surface litter are adequate to stimulate new sprout growth, but aerial vegetation that was killed, but not consumed, by fire presents an increased fire hazard, and reburning may be warranted for fire hazard reduction. In 1 study in a North Carolina canebrake, a prescribed fire reduced fire hazard for the first 2 to 3 years, and fuels reached a peak of 5 to 7 tons/acre after 3 or 4 years of fire protection. Therefore, if a reduction of fire hazard is desired, a short burning cycle is preferable [54].

Prescribed fire may not help to promote the rapid spread of cane into adjacent areas. If soils are compacted, lateral penetration of roots and rhizomes is slow [54].

Grazing reduces the fire hazard in cane stands. In the pocosins of North Carolina, grazing reduced the total combustible material per acre by 43%. Three different fires were noticeably slowed down and/or stopped once they entered the grazed area. Although burning may be beneficial in some respects, burned cane range is particularly susceptible to grazing damage, and over-use of fresh burns must be avoided to maintain grazing values [91].

Hilmon and Hughes [50] cautioned that control of wild cane fires may be "difficult or impossible" because of their speed and intensity [50].

MANAGEMENT CONSIDERATIONS

SPECIES: Arundinaria gigantea
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Cane provides high quality forage for cattle, horses, swine, and domestic sheep [62]. Because it is evergreen, cane is good for grazing year-round [4]. Cane was once widely utilized as a forage plant for cattle and domestic sheep across much of the southeastern U.S. In Mississippi cane was once commonly called "mutton grass" because of its value as domestic sheep forage [62]. Because of the dramatic reduction in cane habitat, it is generally no longer considered a valuable range forage plant [46].

Cane is easily damaged by grazing and the rooting of swine, and stands may take years to recover from damage [20,51,91]. Overgrazing is considered 1 of the major factors involved in the decrease of cane habitat in the U.S. following European settlement [7]. Plants are most susceptible to grazing damage in the spring and summer [4]. Continuous summer grazing can cause a decline in cane stem density and a reduction in stem height [54]. According to a 1971 handbook, no more than 50% of the current year's growth should be grazed off in any season. It is also recommended that summer grazing be deferred for at least 90 days every 2 to 3 years. Controlled burns every 3 to 4 years can be used to maintain cane fields and improve forage value. Burned fields must be protected from grazing for the first growing season to allow the cane to recover [62].

Palatability/nutritional value: Where it occurs, cane is 1 of the most palatable and preferred forages by cattle, and it can comprise the bulk of the animal's diet when abundant [91]. The crude protein, calcium, and phosphorus content of cane average higher than other native southern grasses [46]. Digestible nutrients in cane foliage are highest in May and June and decline rapidly during the remainder of the summer and fall [55].

Cover value: Cane provides good cover for nesting birds, small mammals, and reptiles [5,74]. Canebrakes are critical nesting habitat for the Swainson's warbler [7,44,99]. In the South Carolina coastal plain, hooded warblers have a high nesting success rate in dense patches of cane, possibly because the nests are well protected from snake predation [67]. Bachman's warbler historically required extensive canebrakes for nesting, and the possible extinction of this bird is probably related to the disappearance of large canebrakes [72,81]. The white-eyed vireo and Kentucky warbler are also strongly associated with cane [83]. Cane growing in creek valleys provides desirable cover for northern bobwhite [19].

Canebrakes formerly supported high population densities of white-tailed deer, bison, and wild turkeys in the southeastern U.S., and provided good denning cover and escape corridors for black bear and mountain lion [7]. Swamp rabbits utilize canebrakes for cover and browse the foliage and shoots. The rabbits appear to be restricted to canebrakes in southern Indiana and southeastern Missouri [74,98]. The disappearance of large canebrakes has been cited as a causal factor in population declines of bison, black bear, and swamp rabbit in the Southeast [7,74]. White-tailed deer forage switch cane stems only in the spring of the first year following a burn. Thereafter, the stems become too coarse and are no longer palatable [111]. Switch cane is an important summer food of black bears in the Great Dismal Swamp in Virginia and North Carolina [22,49]. Meadow voles, southern bog lemmings, and several species of shrew are frequently associated with cane in the Great Dismal Swamp [80]. Golden mice incorporate cane foliage into aboveground nests that are frequently supported by cane stems. American beaver consume living stems and foliage, particularly during late winter when other herbaceous vegetation is unavailable [74].

The southern subspecies of the timber rattlesnake is commonly referred to as the "canebrake rattlesnake" because of its affinity for cane habitats. Cottonmouths, copperheads, and pygmy rattlesnakes are also commonly found in canebrakes, presumably because of the abundance of birds and small rodents that are their prey. In a radio-telemetry study in Virginia, it was found that copperheads spent more time in small canebrakes than the adjacent lowland swamps [74].

At least 6 species of butterfly are considered obligate cane specialists: creole pearly eye, southern pearly eye, southern swamp skipper, cobweb little skipper, cane little skipper, and the yellow little skipper [7].

VALUE FOR REHABILITATION OF DISTURBED SITES:
High culm density, rapid lateral spread, and rapid height growth make cane a good choice for riparian buffer zones. Cane's compact network of rhizomes provides streambank stabilization, sediment retention, and bioaccumulation of nutrients and toxins [23]. Without the mediation effects of cane, there is an increased potential for damage to the riparian system. Research has shown that cane can significantly reduce nitrogen, phosphorus, and sediment in surface runoff and nitrogen and phosphorus in groundwater. In 1 study around row-crop fields in southern Illinois, cane was effective in reducing ground water nitrate levels by 90%, and dissolved reactive phosphorus concentrations by 28% [5,87].

Cirtain and others [14] conducted greenhouse studies on the germination and growth of cane seedlings. Seedlings were able to survive both flooding and drought, but grew better under well-drained conditions. Although cane can be propagated by seed, seed is sporadically produced and has low viability. Therefore, artificial propagation is best achieved by vegetative means including rhizome cuttings and clump division [88]. Transplanting stem clumps is often more successful than using individual stems [31]. The survival of transplanted cane varies widely, and slow growth is a common problem. Care should be taken to keep transplant stock from drying out. Amendments of hardwood mulch and composted manure may help increase the success of transplantings [23]. Because propagation of cane by digging and transplanting culms is labor intensive, cumbersome, and costly, research is being carried out to develop procedures for producing machine-plantable rhizome stock for use in canebrake restoration [115].

OTHER USES:
Native Americans utilized cane for a variety of purposes. The stems were used to make spears, arrows, blowguns, pipes, flutes, and fish traps. The leaves were woven into baskets and mats [94]. It is estimated that Native Americans burned cane every 7 to 10 years to maintain and expand canebrakes [7,26]. Cane has also been used as a potherb and for fishing poles [51].

OTHER MANAGEMENT CONSIDERATIONS:
More than 98% of all large canebrakes in the U.S. have been lost since the time of European settlement, and canebrakes are considered "critically endangered" ecosystems as defined by the National Biological Service. Large canebrakes historically performed a valuable role in protecting water quality by their ability to mediate sedimentation and nutrient pollution. They also provided a level of flood control in low-lying areas [69]. The loss of canebrakes has left many areas more vulnerable to damage from sedimentation, nutrient pollution, and flood damage, and the loss of cane habitat has been strongly tied to declines in several associated wildlife species [69,88]. In recent years there has been significant interest in the restoration of canebrakes through the use of prescribed fire and artificial propagation, and continued efforts are needed to assure the survival of this ecosystem [5].

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