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| Understory infestation. Photo ©John M. Randall/The Nature Conservancy |
| State | Status |
| Alabama | Noxious weed |
| Georgia | Noxious weed [37] |
| Kentucky | Rank 1, a severe threat [53] |
| Tennessee | Rank 1, a severe threat [89] |
| Virginia | Highly invasive [104] |
The following lists give biogeographic classifications where Japanese stiltgrass is known to be present or invasive, or is likely to be invasive based upon current knowledge of Japanese stiltgrass's habitat preferences. Precise distribution information is unavailable for all locations where Japanese stiltgrass may be invasive. The following lists are therefore speculative and not exhaustive, and Japanese stiltgrass may be present and possibly invasive in other vegetation types.
ECOSYSTEMS [36]:| AL | AR | CT | DE | FL | GA | IL |
| IN | IA | KY | LA | MD | MS | MO |
| NJ | NY | NC | OH | PA | SC | TN |
| TX | VA | WV | DC | PR | VI |
In recently burned, mixed-mesophytic woodland of southern Illinois, Japanese stiltgrass has associated with Philadelphia fleabane (Erigeron philadelphicus), clammy groundcherry (Physalis heterophylla), fragrant bedstraw (Galium triflorum) and drooping woodreed (Cinna latifolia) in the understory. Overstory associates included river birch (Betula nigra), black walnut (Juglans nigra), sycamore (Platanus occidentalis), black cherry (Prunus serotina), and winged elm (Ulmus alata) [3]. Overstory associates in a southern Illinois black oak-post oak (Quercus velutina-Q. stellata) forest in early old-field succession included eastern redcedar (Juniperus virginiana), flowering dogwood (Cornus florida), sassafras (Sassafras albidum), and common persimmon (Diospyros virginiana). Coralberry (Symphoricarpos orbiculatus), Japanese honeysuckle (Lonicera japonica), and poison-ivy (Toxicodendron radicans) were common woody understory associates. Herbaceous associates included big bluestem (Andropogon gerardii), golden alexanders (Zizia aurea), and blunt-lobe woodsia (Woodsia obtusa) [38].
In New Jersey, Japanese stiltgrass occurred in mixed red-black-chestnut-white oak (Q. rubra-Q. velutina-Q. prinus-Q. alba) and white ash-sweet birch-American beech (Fraxinus americana-Betula lenta-Fagus grandifolia) forests, but was less common on sites with high cover of overstory oaks and understory blueberries (Vaccinium spp.) [55]. Overstory associates of Japanese stiltgrass in a sugar maple-red maple (Acer saccharum-A. rubrum)-sweet birch forest in New Jersey included shagbark hickory (Carya ovata), bitternut hickory (C. cordiformis), and American elm (U. americana). The most common shrubs included multiflora rose (Rosa multiflora), black haw (Viburnum prunifolium), and spicebush (Lindera benzoin). Jack-in-the-pulpit (Arisaema vimineum) frequently co-occurred in the ground layer, although Japanese stiltgrass was the most common ground layer species [103].
Japanese stiltgrass occurs in the understories of Virginia pine-southern red oak (Pinus virginiana-Q. falcata) communities in Maryland. Yellow-poplar (Liriodendron tulipifera), red maple, hickories (Carya spp.), and black cherry are associates [16]. Japanese stiltgrass is also a component of mixed oak-sweetgum-swamp tupelo (Quercus spp-Liquidambar styraciflua-Nyssa sylvatica var. biflora) communities on in inland coastal plain of Chesapeake Bay, off Maryland and Virginia [79]. On the George Washington Memorial Parkway in Virginia, Japanese stiltgrass occurs in the ground layer of old-growth oak-hickory (Quercus-Carya spp.) forest. Dominant trees include white, scarlet (Q. coccinea), and chestnut oaks, shagbark hickory, and mockernut hickory (C. tomentosa). Shrub associates include mountain-laurel (Kalmia latifolia), pink azalea (Rhododendron periclymenoides), and black huckleberry (Gaylussacia baccata). Ground-layer herbaceous associates are winter bent grass (Agrostis hyemalis), broomsedge bluestem (Andropogon virginicus), common velvet grass (Holcus lanatus), and white clover (Trifolium repens). Vines are common in the forest and include trumpet-creeper (Campsis radicans), Oriental bittersweet (Celastrus orbiculatus), Japanese honeysuckle, and summer grape (Vitis aestivalis) [107].
Japanese stiltgrass is very common in low-elevation oak-pine (Quercus-Pinus spp.) forests of the Piedmont [49,80,81], where it is an indicator of red clay soils [49]. Romagosa and Robinson [81] provide a comprehensive list of shrub, vine, and herbaceous associates of Japanese stiltgrass in upland loblolly pine (P. taeda)-mixed oak forest on Piedmont sites in Pennsylvania.
In mixed-hardwood forest in the Cumberland Mountains of Kentucky, overstory associates of Japanese stiltgrass have included northern red oak (Q. rubra), white oak, yellow-poplar, Virginia pine, sugar maple (Acer saccharum), basswood (Tilia heterophylla), American beech, and yellow buckeye (Aesculus octandra). Common shrubs and vines were strawberry-bush (Euonymus americana), hillside blueberry (Vaccinium pallidum), Virginia creeper (Parthenocissus quinquefolia), and common greenbrier (Smilax rotundifolia). At 9% to 35% cover, Japanese stiltgrass was the most common graminoid. Associated grasses and forbs included mannagrass (Glyceria spp.), slender muhly (Muhlenbergia tenuiflora), white snakeroot (Ageratina altissima), and panicledleaf ticktrefoil (Desmodium paniculatum) [76].
Japanese stiltgrass often associates with other invasive, nonnative species in the understory, with native species in the overstory. Japanese honeysuckle is a consistent understory associate of Japanese stiltgrass. On a North Carolina floodplain, Japanese stiltgrass and Japanese honeysuckle comprised nearly 100% of the understory of a boxelder-green ash (Acer negundo-Fraxinus pennsylvanica)-sycamore forest [5]. Japanese barberry (Berberis thunbergii) is another nonnative shrub that commonly co-occurs with Japanese stiltgrass across Japanese stiltgrass's distributional range [82]. In New Jersey, Japanese stiltgrass and Japanese barberry co-occurred in bottomland oak (Quercus spp.)-American beech-sweet birch forest. Hillside blueberry and black huckleberry were commonly associated native shrubs [27]. In southern Illinois oak-hickory forest, Japanese stiltgrass co-occurred with Japanese honeysuckle, sericea lespedeza (Lespedeza cuneata), and multiflora rose [38].
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| James H Miller, USDA Forest Service, www.ipmimages,org |
Morphology: Japanese stiltgrass is a nonnative annual grass. It has a straggling to decumbent, loosely branched habit. Aerial culms are 3 to 5 feet (1-1.5 m) long. Japanese stiltgrass also produces short to long (depending upon shading), spreading stolons. Japanese stiltgrass populations often form dense lawns of intertwined stolons. Japanese stiltgrass leaves are cauline, with 0.5-inch- (1-cm) wide and 3- to 4-inch- (8- to 10-cm) long blades. The inflorescence is a 4.5 to 6 mm, terminal or axillary raceme bearing paired spikelets. The upper lemma is fertile; the lower lemma is sterile and often bears a 4- to 8-mm, twisted awn, although some fruits are awnless. The fruit is a 2.8- to 3.0-mm, ellipsoid caryopsis [18,31,39,75]. Root biomass of Japanese stiltgrass is "remarkably small" compared to its aboveground biomass [26,27].
There has been confusion as to whether Japanese stiltgrass is sometimes perennial [25,26,63]. Mehrhoff [63] states that this confusion arose from misidentification of white grass (Leersia virginica), a morphologically similar native perennial, as Japanese stiltgrass. Japanese stiltgrass is distinguished from white grass, which it often co-occurs with, by its ciliate leaf sheath collar and paired spikelets (vs. white grass' glabrous to pubescent leaf sheath and 1-flowered spikelets) [63].
Physiology: Japanese stiltgrass is adapted to low-light conditions [20,47,96]. Japanese stiltgrass uses C4 pathway photosynthesis. It is highly unusual for a C4 grass to photosynthesize efficiently under low light conditions, but Japanese stiltgrass is very shade tolerant [5,8,13,47,108]. In the greenhouse, Winter and others [108] found Japanese stiltgrass grew well under 5% of full sunlight, and the photosynthetic rate of individual leaves was fully saturated at 25% of full sunlight. Dry-matter biomass production was similar under 18% to 100% of full sunlight. Japanese stiltgrass in the understory of a closed-canopy yellow-poplar-white oak forest in Great Smoky Mountains National Park took advantage of occasional, high-intensity sunflecks for optimal photosynthesis [47]. Best Japanese stiltgrass growth occurs on forest-grassland ecotones, where mean photosynthetically active radiation (PAR) is 35% [20]. Ueno [102] provides a description of Japanese stiltgrass's leaf physiology and cellular anatomy.
Species response to increased levels of atmospheric CO2 can affect plant community composition. High CO2 levels may negatively affect Japanese stiltgrass compared to other plants species that are better able to assimilate extra CO2. In field experiments in Tennessee, Belote and others [11] found that in a wet year, Japanese stiltgrass produced twice as much biomass under ambient CO2 levels compared to elevated CO2 levels. In a dry year, there was no significant difference (P=0.07) in Japanese stiltgrass biomass between CO2 treatments. In contrast, Japanese honeysuckle, a common associate of Japanese stiltgrass, produced 3 times as much biomass under elevated CO2 levels in both wet and dry years.
RAUNKIAER [77] LIFE FORM:Breeding system: In Japan and Taiwan, both cleistogamy and chasmogamy have been observed in Japanese stiltgrass, with cleistogamous flowers growing on axillary racemes and chasmogamous flowers growing on terminal racemes [60,94]. Japanese stiltgrass in the United States apparently also utilizes both breeding systems, although mechanisms controlling which breeding system is utilized are understudied. In a Japanese stiltgrass population near Charlotte, North Carolina, Barden [5] found about 10% of plants had chasmogamous flowers, with chasmogamous plants mostly growing in moist, open sites. All Japanese stiltgrass growing in heavy shade had cleistogamous flowers. This suggests that light conditions may affect flower development and breeding. In southern Illinois population with 80% overstory cover, flowers were mostly cleistogamous [38]. In New York, chasmogamous flowers were most common in shady forests interiors. The ratio of cleistogamous:chasmogamous flowers increased in the greenhouse; tillers were also larger in the greenhouse compared to closed-canopy forest [17]. Further studies are needed regarding breeding systems of Japanese stiltgrass.
Pollination: Japanese stiltgrass is both self- and cross-pollinated [48].
Seed production is reportedly high for Japanese stiltgrass [18,105]. A single Japanese stiltgrass culm may produce 100 to 1,000 seeds [105]. A southern Illinois study found a mean of 81.7 spikelets/Japanese stiltgrass culm. However, pooled seed viability for 4 Japanese stiltgrass populations was low (33%). Spikelet production varied significantly (p<0.001) among populations. Japanese stiltgrass populations may fail to flower in drought years [38].
Seed dispersal: Reviews indicate that Japanese stiltgrass seed is dispersed by wind, water, animals, and humans [10,92,96]. Japanese stiltgrass fruits are light and easily float on water, and Japanese stiltgrass seeds may survive and germinate after "extended periods" of inundation [105], so flooding is a likely means of seed dispersal. Japanese stiltgrass often occurs on floodplains [5,38]. Its highest cover typically occurs on disturbed floodplains (e.g., developed or frequently mowed) [5]; however, frequent, severe flood scouring can limit Japanese stiltgrass establishment and spread [38]. The seed awns (of awned fruits) catch on animal hides, feathers, and human clothing [10,92,96]. Japanese stiltgrass fruits are small, so even awnless fruits can work their way into fur or clothing [96]. Machinery, fill dirt, and contaminated hay are also potential dispersal vectors for Japanese stiltgrass seed [10,92,96]. A Japanese stiltgrass population along a hiking trail in southern Illinois was thought to have established from seed dispersed by tractors used to grade the trail, and/or hikers [38].
Seed banking: Japanese stiltgrass builds a soil seed bank [5,38]. Seed longevity is estimated at 3 to 5 years [5,38,96]. On a North Carolina site, soil-stored Japanese stiltgrass seed remained viable for at least 3 years. Barden [5] estimated the number of plants produced from 1983 to 1986 on a 2-m˛ study plot was 1,000 (in 1983), 256 (1984), 44 (1985), and 0 (1986), respectively. Another 2-m˛ study plot on the North Carolina site, measured from 1984 to 1986, produced 857, 47, and 29 Japanese stiltgrass plants, respectively [5].
Germination: Although seed production is reportedly high [18,105], few seed germination and viability studies have been conducted as of this writing (2005). A greenhouse study found that fresh seed was not immediately germinable, while seeds grown for 90 days showed 100% germination rates [51]. A southern Illinois study found poor seed germinability during a drought year (see Seed production above). Even so, mean seed rain was 24.6 seeds/m˛ (n=34 seed traps). It is difficult to conclude that occasional seed failure is limiting for this species. Given a persistent seed bank, Japanese stiltgrass may establish in high densities the year following seed crop failure [38].
Seedling establishment/growth: In the southern Illinois study [38], Japanese stiltgrass seedlings established at a mean density of 43/m˛. Plant mortality was a greatest (50% or more) in early seedling establishment (mid-March), dropping to about 20% by July [38]. Japanese stiltgrass shows plasticity to its environment, allocating carbon resources for best growth and reproduction. On shaded sites, more carbon is allocated to leaves and aerial stems compared to stolons, maximizing photosynthetic tissues and output [18]. Japanese stiltgrass is well adapted to shady conditions. It can establish, grow, and produce seed in as little as 5% of full sunlight [105].
Barriers to reproduction: In North Carolina, Japanese stiltgrass regeneration was negatively correlated with higher soil pH (5.5 vs. a median of 5.1); higher levels of soil potassium, zinc, and calcium; higher percent silt (18% vs. 10%); deeper litter (8.6 vs. 5.5 cm); greater cumulative PAR on an overcast day (0.72 vs. 0.57 mol/m˛/day); and greater leaf area index (LAI) of other species (1.3 vs. 0.7) (see Japanese stiltgrass and Japanese honeysuckle for further information) [5]. In southern Illinois, reproductive success was correlated with soil conditions and canopy cover. Reproduction increased with increasing availability of soil cations and soil sand content, and decreased with increased soil silt content and canopy cover. Late-season drought can greatly reduce or eliminate Japanese stiltgrass seed production for a cohort [38].
Deep litter, especially oak litter, may retard Japanese stiltgrass establishment [5,26]. In a landscape-level study of 3 white oak-sweet birch forests in New Jersey, sites with Japanese stiltgrass had lower amounts of litter compared to adjacent uninvaded sites. Over 2 years, the on-site decay rate of white oak litter was slower (30% mass loss) than decay rates for sweet birch and Japanese stiltgrass litter (40%-50% for both spp.). Japanese stiltgrass may alter site conditions once it has established, impairing the ability of native plant species to establish. Other differences on Japanese stiltgrass-invaded sites included thinner organic soil horizons, higher soil pH values (µ=6.0), and higher levels of available soil nitrate compared to adjacent uninvaded sites [26]. Other researchers have had similar findings (see Impacts for further discussion).
Asexual regeneration: During a growing season, Japanese stiltgrass increases vegetatively by tillers and stolons [49]. Because Japanese stiltgrass is an annual, these vegetative shoots do not survive through the next growing season [38]. Large vegetative biomass does, however, increase the likelihood of reproductive success by increasing photosynthate and thus, the potential for greater seed production.
SITE CHARACTERISTICS:Soils: Japanese stiltgrass is common on silty to sandy loams [5,38,76]. It is also reported on red clays [49]. Soil pH is generally acidic, although some populations occur on neutral, limestone- or marble-derived soils [96]. A survey in Maryland and Washington, DC, found that Japanese stiltgrass sites ranged from 4.8 to 5.8 in pH [78]. On mine spoils in Kentucky, Japanese stiltgrass grew on loamy soils ranging from 4.6 to 6.3 pH. It was absent from an extremely acidic site (pH 4.4) [76]. Nutrient content may vary. In an Illinois study, soils supporting Japanese stiltgrass were generally acidic and nutrient poor [38]. Other studies report high levels of nitrogen, and average levels of potassium and phosphorus, on Japanese stiltgrass-infested soils [78].
Japanese stiltgrass prefers damp to wet soils, although it does not tolerate standing water for "extended periods" of time (review by [105]). It may also establish on drier, upland soils [92]. The Virginia Department of Conservation and Recreation [105] states that Japanese stiltgrass is common on disturbed soils and can rapidly spread onto undisturbed soils once established. Characteristics for undisturbed sites most vulnerable to invasion were not given.
Elevation: Japanese stiltgrass occurs from sea level up to 4,000 feet (1,000 m) elevation [64]. It is most common in low-elevation woodlands in the Mid-Atlantic states and the Piedmont and Appalachian mountains [75]. As of this writing (2005), it is not reported from higher-elevation red spruce-Fraser fir (Picea rubens-Abies fraseri) forests.
Climate: Information on optimal growing temperature and temperature limits for Japanese stiltgrass were not available as of this writing (2005). The coldest reported winter temperatures that Japanese stiltgrass survived were approximately -5.8 to -9.4 °F (-21 to -23 °C) [78].
SUCCESSIONAL STATUS:Japanese stiltgrass can invade old growth forest understories. For example, it is a component of the understory in old-growth sweetgum-overcup oak (Quercus lyrata)-river birch bottomland forests of Tennessee [85]. It can form patches or dense, continuous lawns in late-successional forests [26], but may not establish in very shady sites. Field experiments in an oak-hickory forest in Kentucky showed that Japanese stiltgrass was unable to establish under the understory canopy, which consisted of juvenile red maples and spice bushes (Lindera bensoin). The oak-hickory forest was in late succession, and PAR was 1% to 2.5% of full sunlight beneath the red maple and spice bush subcanopy [20].
Japanese stiltgrass can recover rapidly after flooding (but see [38]). A North Carolina study was initiated in 1982 on the Big Cross Creek floodplain. Big Cross Creek flooded the study plots in 1983, reducing Japanese stiltgrass cover. Japanese stiltgrass cover during the study period was [5]:
| 1982 (preflood) | 1983 (postflood) | 1985 (postflood) |
| 48% | 23% | 55% |
The input of silt and nutrients that accompanies short-term flooding can increase Japanese stiltgrass growth rate [5].
SEASONAL DEVELOPMENT:| Area | Event | Season |
| Carolinas | flowers | September-October [75] |
| Florida | flowers | Fall [109] |
| Illinois | seedlings establish | May [38] |
| flowers | September-October [66] | |
| disperses seed and dies | October-November [38] | |
| North Carolina | germinates | March |
| stem expands | April | |
| flowers | September | |
| disperses seed and dies | October [5] | |
| Virginia | germinates | March |
| flowers | October [22] | |
| Eastern U.S. | fruits | September-October [64,96] |
| disperses seed and dies | September-December [64] |
Fuels: As an annual, mat-forming grass, Japanese stiltgrass produces large amounts of fine fuels. Japanese stiltgrass stems lodge soon after they die in autumn [5,16], creating a continuous fuelbed of matted straw that can fuel surface fire [5]. In a New Jersey study, Japanese stiltgrass litter decayed more slowly than litter of native hillside blueberry [27].
Fire regimes vary across Japanese stiltgrass's range. In northeastern maple-birch-beech (Acer-Betula-Fagus spp.) forests, historic fire return intervals were highly variable, depending upon microclimate, topography, and soil. Fires were mostly of mixed severity. Stand-replacing, medium-interval (~80-yr) fires were most common in forests dominated by birches, while long-interval (≥300 years), mixed-severity or stand-replacing fires occurred in forests dominated by maple and/or beech [29,33,45,83,106]. Oak-hickory (Quercus-Carya spp.), oak-pine (Quercus-Pinus spp.), and pine (Pinus spp.) forests of the Northeast and Southeast had mostly short-return interval, understory surface fires [91,106]. Japanese stiltgrass was not present in these forests while historic fire regimes were still operating. It is unclear how Japanese stiltgrass may affect or alter fire regimes in plant communities where it is present because as of this writing (2005), fire ecology studies are lacking for Japanese stiltgrass. Given Japanese stiltgrass's proclivity to invade early seral, disturbed sites [3,38], and its ability to produce abundant litter [5,16,27], Japanese stiltgrass probably alters fuel characteristics on invaded sites and may increase fire frequency and severity on sites where it is abundant. Fire studies are needed on Japanese stiltgrass.
The following table provides fire return intervals for plant communities and ecosystems where Japanese stiltgrass is important. For further information, see the FEIS review of the dominant species listed below. This list may not be inclusive for all plant communities in which Japanese stiltgrass occurs. If you are interested in plant communities or ecosystems that are not listed below, see the complete FEIS Fire Regime Table.
| Community or Ecosystem | Dominant Species | Fire Return Interval Range (years) |
| maple-beech | Acer-Fagus spp. | 684-1,385 [19,106] |
| silver maple-American elm | A. saccharinum-Ulmus americana | < 35 to 200 |
| sugar maple | A. saccharum | > 1,000 |
| sugar maple-basswood | A. saccharum-Tilia americana | > 1,000 [106] |
| birch | Betula spp. | 80-230 [91] |
| sugarberry-America elm-green ash | Celtis laevigata-U. americana-Fraxinus pennsylvanica | < 35 to 200 |
| beech-sugar maple | Fagus spp.-A. saccharum | > 1,000 |
| black ash | Fraxinus nigra | < 35 to 200 [106] |
| cedar glades | Juniperus virginiana | 3-22 [44,73] |
| yellow-poplar | Liriodendron tulipifera | < 35 |
| shortleaf pine | Pinus echinata | 2-15 |
| shortleaf pine-oak | P. echinata-Quercus spp. | < 10 |
| slash pine | P. elliottii | 3-8 |
| slash pine-hardwood | P. elliottii-variable | < 35 |
| sand pine | P. elliottii var. elliottii | 25-45 [106] |
| longleaf-slash pine | P. palustris-P. elliottii | 1-4 [67,106] |
| longleaf pine-scrub oak | P. palustris-Quercus spp. | 6-10 [106] |
| pitch pine | P. rigida | 6-25 [14,46] |
| pocosin | P. serotina | 3-8 |
| pond pine | P. serotina | 3-8 |
| eastern white pine-northern red oak-red maple | P. strobus-Q. rubra-A. rubrum | 35-200 |
| loblolly pine | P. taeda | 3-8 |
| loblolly-shortleaf pine | P. taeda-P. echinata | 10 to < 35 |
| Virginia pine | P. virginiana | 10 to < 35 |
| Virginia pine-oak | P. virginiana-Quercus spp. | 10 to < 35 |
| sycamore-sweetgum-American elm | Platanus occidentalis-Liquidambar styraciflua-U. americana | < 35 to 200 [106] |
| eastern cottonwood | Populus deltoides | < 35 to 200 [73] |
| black cherry-sugar maple | Prunus serotina-A. saccharum | > 1,000 |
| oak-hickory | Quercus-Carya spp. | < 35 |
| northeastern oak-pine | Quercus-Pinus spp. | 10 to < 35 [106] |
| oak-gum-cypress | Quercus-Nyssa-spp.-Taxodium distichum | 35 to > 200 [67] |
| southeastern oak-pine | Quercus-Pinus spp. | < 10 |
| white oak-black oak-northern red oak | Q. alba-Q. velutina-Q. rubra | < 35 |
| bear oak | Q. ilicifolia | < 35 [106] |
| chestnut oak | Q. prinus | 3-8 |
| northern red oak | Q. rubra | 10 to < 35 |
| post oak-blackjack oak | Q. stellata-Q. marilandica | < 10 |
| black oak | Q. velutina | < 35 |
| eastern hemlock-yellow birch | Tsuga canadensis-Betula alleghaniensis | > 200 [106] |
| elm-ash-cottonwood | Ulmus-Fraxinus-Populus spp. | < 35 to 200 [24,106] |
In North Carolina, a 9 April 1982 controlled burn from an unrelated study killed a dense upland stand of Japanese stiltgrass seedlings. The previous year's cohorts left a dense mat of Japanese stiltgrass straw on the ground that fueled what was described as "a hot ground fire" that killed the Japanese stiltgrass seedlings. By mid-June, a 2nd cohort of Japanese stiltgrass had established, covering the study site with a dense stand of grass. Since the study site was above the floodplain, the seedlings presumably established from soil-stored seed rather than water-transported seed [5].
Gibson and others [38] provide and anecdotal account of "increased recruitment" of Japanese stiltgrass following prescribed fire in a xeric, early successional oak-hickory woodland established on old fields abandoned from agriculture in the 1960s (Shimp, personal communication cited in [38]). No additional information is given.
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:Preventing postfire establishment and spread: More research is needed specific to fire tolerance and response of Japanese stiltgrass in specific sites and ecosystems in which it occurs. The USDA Forest Service's "Guide to Noxious Weed Prevention Practices" [100] provides several fire management considerations for weed prevention in general that apply to Japanese stiltgrass. Guidelines for determining burn severity, revegetation necessity, and establishing and managing competitive plants are available [41,42]. The following paragraphs provide some general guidelines for invasive species management after fire. See Integrated Noxious Weed Management after Wildfires for a more detailed source of this information.
When planning a prescribed burn, preinventory the project area to evaluate cover and phenology of any Japanese stiltgrass or other invasive plants present on or adjacent to the site, and avoid ignition and burning in areas at high risk for Japanese stiltgrass establishment or spread due to fire effects. Avoid creating soil conditions that promote weed germination and establishment. Areas of soil disturbance (e.g. those brought about by fire suppression activities) are especially susceptible to invasive plant establishment. Weed status, risks, and prevention must be incorporated in fire rehabilitation plans. Also, wildfire managers might consider including weed prevention education and providing weed identification aids during fire training; avoiding known weed infestations when locating fire lines; monitoring camps, staging areas, helibases, etc., to be sure they are kept weed free; taking care that equipment is weed free; and acquiring restoration funding. Additional guidelines and specific recommendations and requirements are available [42,100].
Preventing invasive plants from establishing in weed-free burned areas is the most effective and least costly management method. This can be accomplished through early detection and eradication, careful monitoring, and by limiting invasive plant seed dispersal into burned areas by [4,42,100]:
re-establishing vegetation on bare ground as soon as possible
using only certified weed-free seed mixes when revegetation is necessary
cleaning equipment and vehicles prior to entering burned areas
regulating or preventing human and livestock entry into burned areas until desirable site vegetation has recovered sufficiently to resist invasion by undesirable vegetation
detecting weeds early and eradicating before vegetative spread and/or seed dispersal
eradicating small patches and containing or controlling large infestations within or adjacent to the burned area
In general, early detection is critical for preventing establishment of large populations of invasive plants. Monitoring in spring, summer, and fall is imperative. Eradicate established Japanese stiltgrass plants and small patches adjacent to burned areas to prevent or limit seed dispersal into the site [4,42,100].
The need for revegetation after fire can be assessed on the basis of the degree of desirable vegetation displaced by invasive plants prior to burning, and on postfire survival of desirable vegetation. Revegetation necessity can also be related to invasive plant survival as viable seeds, root crowns, or rhizomes capable of reproduction. In general, postfire revegetation should be considered when desirable vegetation cover is less than about 30% [42].Palatability: Japanese stiltgrass is unpalatable to deer and all classes of livestock (A. Houston, personal communication cited in [6]), [64]. White-tailed deer do not graze it, and may indirectly encourage Japanese stiltgrass spread by avoiding Japanese stiltgrass and foraging on more palatable species [92].
Nutritional value: No information is available on this subject.
Cover value: Japanese stiltgrass mats create good hiding and nesting cover for rats, especially cotton rats. Conversely, Japanese stiltgrass degrades nesting habitat for northern bobwhite (A. Houston, personal communication cited in [6]). Since cotton rats are predators of the northern bobwhite, Japanese stiltgrass must doubly impact northern bobwhite by reducing high-quality native cover and increasing predation. No information is available on how Japanese stiltgrass affects cover value for other wildlife species.
OTHER USES:
IMPACTS AND CONTROL:
Impacts:
Japanese stiltgrass is a serious weed on
disturbed soils [9]. It is capable of altering ecosystem function
[26,27,54,55,56,57,58] and reducing site diversity [38]. For
example, Kourtev and others reported that Japanese stiltgrass-invaded areas in
New Jersey had
thinner litter and organic soil layers [55]; lower levels of soil carbon,
nitrogen, and net ammonification [57]; dissimilar soil enzymes; and
significantly (p<0.001) higher soil pH compared to uninvaded areas [54,56,58].
Additionally, soil microbial communities differed in species composition in
Japanese stiltgrass and uninvaded areas, and nonnative earthworms were more
common on Japanese stiltgrass sites compared to uninvaded sites [55,57].
Such drastic changes to soils are likely to be long term, and may encourage
reinvasion of Japanese stiltgrass or new invasions of other nonnative species
[54]. Characteristics that contribute to Japanese stiltgrass invasion include [18,96]:
rapid invasion of disturbed habitats
reproductive plasticity to environmental conditions
annual life history
high seed production
rapid clonal growth
In the southern Appalachian region, 8 of 35 federal, state, and private agencies ranked Japanese stiltgrass among their greatest ongoing or potential management problems (behind kudzu and multiflora rose) [61]. On the Oak Ridge National Environmental Research Park, Japanese stiltgrass was ranked the most "aggressively invasive" nonnative species based on distribution, abundance, relative difficulty of control, and ability to exclude native plant species. Japanese honeysuckle and Chinese privet (Ligustrum sinense) were ranked 2nd and 3rd, respectively [23]. Japanese stiltgrass reportedly replaced existing ground vegetation in 3 to 5 years on sites in Great Smoky National Park [89]. As of 2000, extent of infestations in Dixon State Park, Illinois, ranged from 2.3/m˛ to 16,706/m˛ in size [38].
Forestry: Japanese stiltgrass is identified as a potentially serious competitor on productive timber sites in the Southeast [5,81,88]. It interferes with forage production and reduces growth of silvicultural species. On an oak plantation in southwestern Tennessee, Japanese stiltgrass presence was negatively correlated (r=-0.82) with growth of northern red oak seedlings. There was a strong negative correlation (r=-0.74) between Japanese stiltgrass biomass and mean northern red oak seedling height growth. Four silvicultural treatments were tested: clearcut (all stems >6 inches diameter removed); 2-aged cut (harvest to retain a stand basal area of 15-20 ft˛/ac of residual oaks, hickories, and yellow-poplar); high-grade cut (all stems >14 inches dbh removed); and a control no-cut treatment. Mean biomass gain of Japanese stiltgrass was greatest with a 2-aged selection cut and least with the no-cut control [71]:
| 2-aged | Clearcut | High-grade | No cut |
| 1,400 kg/ac | 800 kg/ac | 250 kg/ac | 100 kg/ac |
As an annual herb, Japanese stiltgrass productivity is more closely tied to yearly climate fluctuations compared to other perennial understory species. Annual variations in Japanese stiltgrass productivity can have important effects on forest understory species composition and diversity. On a sweetgum site on the Oak Ridge National Environmental Research Park, Tennessee, Japanese stiltgrass produced 64% as much biomass in a wet year compared to a dry year [11].
Disturbance ecology: Japanese stiltgrass readily establishes following disturbances such as flooding, mowing, and tilling. Within 3 to 5 years it may form monotypic stands that crowd out native vegetation [92,105]. A survey (based on herbaria collections and remote-sensing data) of weed invasion patterns in West Virginia showed that Japanese stiltgrass was most likely to invade road- and streamside vegetation, although both Japanese stiltgrass and Oriental bittersweet showed relatively high occurrence in closed-canopy forests (23% and 31% of total collections, respectively) [48].
Barden [5] concluded that a history of disturbance was likely to improve Japanese stiltgrass's ability to invade a site. A relatively deep litter layer, greater LAI of other ground-dwelling species compared to Japanese stiltgrass, and high levels of sunlight (PAR) reduced reproductive success of Japanese stiltgrass. He found that soil fertility was relatively unimportant in determining invasive ability of Japanese stiltgrass. In Barden's North Carolina study, Japanese stiltgrass failed to regenerate on undisturbed, naturally fertile plots (high levels of soil nitrogen, potassium, calcium, and zinc; see Barriers to Reproduction for further details). On plots treated with a 15-16-17 N-P-K fertilizer, Japanese stiltgrass showed greater biomass gain compared to unfertilized control plots, but seed spike production was similar on fertilized vs. unfertilized plots [5].
Japanese stiltgrass may be less successful at invading undisturbed sites. Rates of Japanese stiltgrass spread onto undisturbed sites are unclear because study data are not available as of 2005. However, anecdotal evidence suggests that Japanese stiltgrass may not invade, or is slow to invade, undisturbed sites. For example, Japanese stiltgrass was absent from unmowed land next to a sewer line right-of-way in North Carolina, but did invade the annually mowed right-of-way [5]. Some sources suggest that Japanese stiltgrass may slowly spread onto undisturbed lands unless control measures are taken [96]. An inventory of Land Between the Lakes National Recreation Area, Kentucky and Tennessee, showed Japanese stiltgrass occurred both within and adjacent to the Recreation Area boundaries. It was more common on adjacent private lands than inside the Recreation Area, which has been protected from mining, logging, and grazing since 1963. The authors cautioned, however, that periodic flooding left the Recreation Area vulnerable to Japanese stiltgrass seed dispersal and invasion [62].
One review suggests that Japanese stiltgrass can spread rapidly onto undisturbed sites from adjacent disturbed sites where it is well established [105]. In a New Jersey survey, Japanese stiltgrass and garlic mustard (Alliaria petiolata) were the only 2 nonnative species that invaded undisturbed chestnut oak-red oak-pitch pine stands [9]. Because it was a presence-absence inventory, density and rates of spread were not available for Japanese stiltgrass. Long-term studies are needed to document Japanese stiltgrass's rate of colonization and expansion onto undisturbed sites.
Japanese stiltgrass and Japanese honeysuckle: Japanese stiltgrass spreads more readily into disturbed, shaded mesic sites compared to Japanese honeysuckle. Although the 2 species commonly co-occur, Japanese stiltgrass shows limited ability to invade established Japanese honeysuckle stands. In a North Carolina study, more than 2,400 Japanese stiltgrass seeds/plot were sown into 40 "undisturbed," well-established Japanese honeysuckle study plots (2-m˛ plots). A year later, less than 1% of the Japanese stiltgrass seeds had germinated. Two years after the seeds were sown, 12 plots had no Japanese stiltgrass plants; 20 plots contained 20 or fewer Japanese stiltgrass plants. Only 1 plot produced more than 2,400 plants (the approximate number of seeds originally cast). For all 40 plots, the ratio of Japanese stiltgrass seeds originally cast to Japanese stiltgrass plants established 2 years later was 6:1. Reciprocal transplant experiments on cleared sites showed no interference in Japanese honeysuckle growth when Japanese honeysuckle was planted on soils that previously supported Japanese stiltgrass compared to Japanese honeysuckle grown on soils that previously supported Japanese honeysuckle. Similarly, Japanese stiltgrass showed no interference in growth when planted on soils that had supported Japanese honeysuckle compared to soils that supported Japanese stiltgrass. Study plots were weeded during the 2-year experiment to eliminate other vegetation [5].
Japanese stiltgrass may be more invasive than Japanese honeysuckle on disturbed, shaded sites. After the reciprocal transplant experiment had ended and plots were no longer being weeded, Japanese stiltgrass rapidly invaded study sites that had been cleared and planted to Japanese honeysuckle. Two years after weeding was discontinued, Japanese stiltgrass showed 63% LAI in Japanese honeysuckle plots compared to adjacent undisturbed Japanese honeysuckle stands. Cumulative PAR at the plots was 3% to 12% of full sunlight. Japanese stiltgrass invaded some undisturbed Japanese honeysuckle floodplain stands after several extreme weather events coincided. Early April frosts in 3 of 4 study years and a record January 1985 low of -4 °F (-20 °C) killed most Japanese honeysuckle leaves. Japanese honeysuckle is evergreen, so leaf loss in successive years is highly detrimental. As an annual, Japanese stiltgrass is much less affected by low winter and early spring temperatures [5].
Control of Japanese stiltgrass is difficult, and requires multiple treatments [23]. In order to locally control this annual, seed-banking plant, repeated annual efforts must be made to prevent flowering and seed set until the seed bank is exhausted [38]. Japanese stiltgrass resembles native white grass, so Japanese stiltgrass should be properly identified before control measures are undertaken [64].
Prevention: The most efficient and effective method of managing invasive species such as Japanese stiltgrass is to prevent their invasion and spread [86]. Preventing the establishment of nonnative invasive plants in wildlands is achieved by maintaining native communities and conducting aggressive surveying, monitoring, and any needed control measures several times each year. Monitoring efforts are best concentrated on the most likely sites of invasion, particularly along potential pathways for Japanese stiltgrass invasion: waterways, roadsides, and adjacent old fields and woodlands. Uninvaded sites should be periodically surveyed to detect new invasions. The Center for Invasive Plant Management provides an online guide to noxious weed prevention practices.
Preventing the introduction of Japanese stiltgrass into uninfested areas, and early control of small infestations, should be a priority [92]. Removing Japanese stiltgrass plants late in the growing season, before Japanese stiltgrass seed set but after seed set of most associated species, is recommended [38,105]. Once established, Japanese stiltgrass requires major, long-term eradication and restoration efforts. The Nature Conservancy [6,96] reports high potential for successful control and management of Japanese stiltgrass if detected and controlled in the early stages of invasion, and moderate potential for Japanese stiltgrass control and large-scale wildland restoration in areas where Japanese stiltgrass is already well established.
Integrated management: A combination of complementary control methods may be helpful for rapid and effective control of Japanese stiltgrass. Integrated management includes not only killing the target plant, but establishing desirable species and discouraging nonnative, invasive species over the long term. Japanese stiltgrass control is rarely successful with only 1 method of control [72], but a combination of control methods can be effective.
The best way to prevent large infestations is to control small patches. Small patches of Japanese stiltgrass in Great Smoky Mountains National Park have been controlled through a combination of herbicides, mowing, and hand pulling (Johnson, K. 2001, cited in [23]). Tu [96] provides a contact list of managers who have used control measures (successful or not) on Japanese stiltgrass in Natural Areas.
Physical/mechanical: Hand-pulling, mowing, tilling, and flooding can control Japanese stiltgrass. Hand-pulling controls small Japanese stiltgrass infestations [23]. Japanese stiltgrass is shallow-rooted and prefers moist soils; hence, it is usually easy to pull. Hand-pulling is most effective in late summer (August-September) when plants are tall and branched. Plants pulled before seed set can be left on site; plants with fruits should be bagged and removed. Hand-pulling upturns soil, and is likely to create microsites favorable to seed-banked Japanese stiltgrass seed. Late summer pulling is advantageous in that seed-banked seed does not have a long enough growing season to establish. Pulling in July or earlier is not recommended. Hand-pulling needs to be continued until the seed bank is exhausted, which may take many years [92,96]. Floodplains and other sites subject to continual replenishment of the seed bank require hand-pulling treatments indefinitely [96].
Mowing is recommended late in the growing season (August-September), when plants are flowering but before seed set. Because Japanese stiltgrass is an annual, late-season mowing curtails regrowth. Early season mowing does not control Japanese stiltgrass because 1) seed-banked seeds can still establish and produce a new crop of seeds by the end of the growing season, and 2) plants cut in early summer respond with new growth and flower production soon after cutting [22,92,105].
Tilling also reduces Japanese stiltgrass [96]. Soil must be tilled late in the growing season to avoid establishment of soil-stored seed. However, tilling may not be appropriate in many Natural Areas and may damage desirable plants.
Flooding for 3 straight months, or intermittent inundation, may kill Japanese stiltgrass plants. It will not kill soil-stored seed [96].
Fire: See the Fire Management Considerations section of this summary.
Biological: Japanese stiltgrass has few natural predators and pathogens in North America [18]. No biological control agents were available for Japanese stiltgrass control as of 2005 [92,96].
Grazing is not recommended as a control measure, as deer and all classes of livestock avoid Japanese stiltgrass (A. Houston, personal communication cited in [6]).
Chemical: Herbicides may provide initial control of a new invasion or a severe infestation, but used alone, they are rarely a complete or long-term solution to invasive species management [15]. Herbicides are most effective on large infestations when incorporated into long-term management plans that include replacement of weeds with desirable species, careful land use management, and prevention of new infestations. Control with herbicides is temporary, as it does not change the conditions that allowed the invasion to occur (e.g. [110]). See The Nature Conservancy's Weed Control Methods Handbook for considerations on the use of herbicides in Natural Areas and detailed information on specific chemicals.
Extensive infestations of Japanese stiltgrass can be controlled with systemic herbicides [92]. Herbicides may be the only practical method to effectively control large infestations. Glyphosate can effectively control Japanese stiltgrass, but since glyphosate is a nonselective herbicide, care must be taken to avoid drift onto desirable native species. Effective use of herbicides requires appropriate herbicide concentrations, application techniques, and timing [68,105]. The University of Tennessee reported good control of Japanese stiltgrass on their Ames Plantation, but also reported that managing for a desirable plant community after Japanese stiltgrass was controlled was "difficult." The University found "good control" with imazapic [6]. Since imazapic is selective for only a few plant species, it killed Japanese stiltgrass plants without killing associated native herbaceous species. Sethoxydim and fluazifop are grass-specific herbicides reported as giving some control for Japanese stiltgrass (Tu, personal communication, [97]). Barden [6] provides information on application rates for Natural Areas and suggestions for herbicide use in wetland areas. Studies reporting the efficacy of various herbicides used for Japanese stiltgrass control in Pennsylvania [43] are available.
It may be necessary to reduce litter build-up before herbicide can be successfully applied (see Fire Management Considerations). Successful herbicide control requires several treatments [23].
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