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|Groundlayer infestation. Photo ©John M. Randall, The Nature Conservancy.|
In the United States, it is sporadically distributed throughout most of the East and in the Caribbean, from New York south to Texas, Florida, Puerto Rico, and the Virgin Islands [58,71,97]. Nepalese browntop was first noted in North America around 1918 in Tennessee [15,51], where it was probably introduced accidentally . It was formerly used as packing material for imported Chinese porcelain, and discarded packaging material containing seeds might have been the source of introduction . Nepalese browntop is rare in Florida and other parts of the Southeast [164,230] but is rapidly increasing in Maryland, New York, and other northern states [15,90,169]. It was introduced in New Jersey around 1959 and spread rapidly in that state in the 1990s and 2000s (review by ). Roads and waterways appear to be the primary corridors for population expansion ; see Site Characteristics and Impacts for information. Plants database provides a map of Nepalese browntop distribution in the United States.HABITAT TYPES AND PLANT COMMUNITIES:
Nepalese browntop is often associated with several other nonnative species in the United States. It is frequently found with garlic mustard (Alliaria petiolata) in the East and Southeast (; also see the Vegetation classifications list below). Japanese honeysuckle (Lonicera japonica) is often consistently associated with Nepalese browntop in the Great Lakes and eastern regions of the United States. In a southern Illinois oak-hickory forest, for example, Nepalese browntop cooccurred with Japanese honeysuckle and was also associated with nonnative sericea lespedeza (Lespedeza cuneata) and multiflora rose (Rosa multiflora) . Japanese barberry (Berberis thunbergii) commonly cooccurs with Nepalese browntop across Nepalese browntop's distributional range . In New Jersey, Nepalese browntop and Japanese barberry grew together in a bottomland oak-American beech-sweet birch (Quercus spp.-Fagus grandifolia-Betula lenta) forest . Nepalese browntop is sometimes associated with Norway maple. In red maple forests of New Jersey, Nepalese browntop dominated the ground layer of sites where Norway maple had replaced red maple as the overstory dominant .
The following descriptions provide information on where Nepalese browntop is known to be present, invasive, or likely to be invasive based upon current knowledge of Nepalese browntop's habitat preferences. Nepalese browntop is likely invasive or dominant in more plant communities than those described below
Great Lakes and Northeast: Nepalese browntop occurs in pine (Pinus), oak (Quercus)-pine, oak-hickory (Carya), and mixed-hardwood woodlands and forests in these regions. In recently burned, mixed-mesophytic woodlands of southern Illinois, overstory codominants of Nepalese browntop-infested sites included river birch (Betula nigra), black walnut (Juglans nigra), sycamore (Platanus occidentalis), black cherry (Prunus serotina), and winged elm (Ulmus alata). Philadelphia fleabane (Erigeron philadelphicus), clammy groundcherry (Physalis heterophylla), fragrant bedstraw (Galium triflorum) and drooping woodreed (Cinna latifolia) cooccurred with Nepalese browntop in the ground layer . Overstory codominants in a southern Illinois black oak-post oak (Q. 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), poison-ivy (Toxicodendron radicans), and nonnative Japanese honeysuckle were commonly associated understory species. Herbs associated with Nepalese browntop in the ground layer included big bluestem (Andropogon gerardii), golden alexanders (Zizia aurea), and blunt-lobe woodsia (Woodsia obtusa) .
In New Jersey, Nepalese browntop occurred in red oak-black oak-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. It was less common on sites with high cover of overstory oaks and understory blueberries (Vaccinium spp.) than in other hardwood forest types . Overstory associates of Nepalese browntop in a sugar maple-red maple (Acer saccharum-A. rubrum)-sweet birch forest in New Jersey included shagbark hickory (C. ovata), bitternut hickory (C. cordiformis), and American elm (U. americana). The most common shrubs included black haw (Viburnum prunifolium), spicebush (Lindera benzoin), and multiflora rose. Although Nepalese browntop was the most common groundlayer species, jack-in-the-pulpit (Arisaema vimineum) frequently cooccurred in the ground layer .
In Maryland, Nepalese browntop occurred in the ground layers of Virginia pine-southern red oak (Pinus virginiana-Q. falcata) communities. Yellow-poplar (Liriodendron tulipifera), red maple, hickory (Carya spp.), and black cherry were associated in the overstory . In Maryland and Virginia, Nepalese browntop was a component of mixed oak-sweetgum-swamp tupelo (Quercus spp.-Liquidambar styraciflua-Nyssa sylvatica var. biflora) communities on the inland coastal plain of Chesapeake Bay .
Appalachians: Nepalese browntop is common in low-elevation oak-pine forests of the Piedmont [90,171,172]. In Cumberland County, Pennsylvania, Nepalese browntop occurred in a red maple/spicebush/skunk cabbage-sphagnum (Symplocarpus foetidus-Sphagum spp.) swamp . Romagosa and Robinson  provide a comprehensive list of shrub, vine, and herbaceous associates of Nepalese browntop in an upland loblolly pine (P. taeda)-mixed oak forest on piedmont sites in Pennsylvania. The federally endangered  glade spurge (Euphorbia purpurea) cooccurred with Nepalese browntop in the forest's groundlayer vegetation .
Nepalese browntop is reported in mixed-hardwood and riparian communities in Kentucky. In mixed-hardwood forest in the Cumberland Mountains, overstory species associated with Nepalese browntop included northern red oak (Q. rubra), white oak, yellow-poplar (Liriodendron tulipifera), 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, Nepalese browntop 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) . Along the Blue River of Kentucky, Nepalese browntop occurred in a big bluestem-indiangrass (Sorghastrum nutans) prairie on gravel wash ; the federally endangered  Short's goldenrod (Solidago shortii) also occurred in the gravel-wash prairie community .
Southeast and South: In the Southeast, Nepalese browntop often occurs upland from or in dry portions of wet grasslands . On a North Carolina floodplain, Nepalese browntop and Japanese honeysuckle comprised nearly 100% of the ground layer and understory of a boxelder-green ash (Acer negundo-Fraxinus pennsylvanica)-sycamore forest .
On the George Washington Memorial Parkway in Virginia, Nepalese browntop occurred in the ground layer of old-growth oak-hickory forest. Dominant trees include white oak, scarlet oak (Q. coccinea), and chestnut oak, shagbark hickory, and mockernut hickory (C. tomentosa). Shrub associates included mountain-laurel (Kalmia latifolia), pink azalea (Rhododendron periclymenoides), and black huckleberry (Gaylussacia baccata). Groundlayer herbaceous associates were winter bent grass (Agrostis hyemalis), broomsedge bluestem (Andropogon virginicus), common velvet grass (Holcus lanatus), and white clover (Trifolium repens). Lianas were common in the forest and included trumpet-creeper (Campsis radicans), Oriental bittersweet (Celastrus orbiculatus), Japanese honeysuckle, and summer grape (Vitis aestivalis) .
Nepalese browntop dominates some deciduous forests of the South. In the Whitehall Experimental Forest, Georgia, Nepalese browntop formed a continuous lawn in the ground layer of a red maple-white oak-sycamore forest. The understory was depauperate . In surveys across west-central Georgia, Nepalese browntop was detected in 15 of 18 watersheds. Nepalese browntop and nonnative species in general were more common in or near urban-rural interfaces, but Nepalese browntop was also common in rural locations. Cover of Nepalese browntop and Chinese privet (Ligustrum sinense) was negatively correlated with overall species richness and overstory reproduction (r= -0.18, P=0.003) for both variables) .
Vegetation classifications describing plant communities in which Nepalese browntop dominates the groundlayer are listed below alphabetically.Arkansas
|Cured Nepalese browntop in a riparian area. Photo by James H Miller, USDA Forest Service, www.ipmimages.org.|
Morphology: Nepalese browntop is an annual. It has a straggling to decumbent, loosely branched habit. Aerial culms are 3 to 5 feet (1-1.5 m) long [34,59,71,164]. They may be "wiry" and multibranched . Nepalese browntop produces short to long (depending upon shading), spreading stolons. Intertwined stolons often form dense lawns. The leaves are cauline, with 0.5-inch (1 cm) wide and 3- to 4-inch (8-10 cm) long blades. The inflorescence is a 4.5 to 6 mm, terminal or axillary raceme bearing paired spikelets [34,59,71,164]. Terminal racemes bear chasmogamous flowers, while axillary racemes bear cleistogamous flowers . The fruit is a 2.8- to 3.0-mm, ellipsoid caryopsis. Fruits often have twisted awns, although some fruits are awnless [34,59,71,164]. In New England collections, presence of awns varied within and among populations . When present, awns are 3 to 8.5 mm long . Root biomass of Nepalese browntop is "remarkably small" compared to its aboveground biomass [51,53], and its roots are shallow [43,200]. A greenhouse study found that at the end of the growing season, Nepalese browntop roots were longest in dry (x=46 inches (18 cm)) soils compared to roots in soils of moderate (5 inches (13 cm)) and saturated (5.5 inches 14 cm)) water content. Lateral roots were few, averaging from 3 to 5 per plant . Another greenhouse study found Nepalese browntop's roots were shallow and its root biomass was significantly less than its aboveground biomass (P<0.001), so the authors concluded Nepalese browntop in unlikely to access moisture in deep soil layers .
There has been confusion as to whether Nepalese browntop is sometimes perennial [50,51,124], but it is not. Mehrhoff  states that this confusion arose from misidentification of white grass—a morphologically similar native perennial—as Nepalese browntop. Nepalese browntop is distinguished from white grass, with which it often cooccurs, by its ciliate leaf sheath collar and paired spikelets (vs. white grass's glabrous to pubescent leaf sheath and one-flowered spikelets) .
Physiology: Nepalese browntop is adapted to low-light conditions [37,83,201]. Nepalese browntop uses C4 pathway photosynthesis. It is unusual for a C4 grass to photosynthesize efficiently under low light conditions, but Nepalese browntop is very shade tolerant [12,14,25,83,228] (see Successional Status). In the greenhouse, Winter and others  found Nepalese browntop 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. Nepalese browntop 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 . Best Nepalese browntop growth occurs on forest-grassland ecotones, where mean photosynthetically active radiation (PAR) is 35% . Ueno  provides a description of Nepalese browntop's leaf physiology and cellular anatomy.
There are apparently genetic differences in shade tolerance among Nepalese browntop populations. Among 3 Nepalese browntop populations from Indiana grown in a growth chamber, 2 populations increased specific leaf area in response to shade, while the other did not .
Species response to increased levels of atmospheric carbon dioxide can affect plant community composition. High carbon dioxide levels may negatively affect Nepalese browntop compared to plant species better able to assimilate extra carbon dioxide. In field experiments in Tennessee, Belote and others  found that in a wet year, Nepalese browntop produced twice as much biomass under ambient carbon dioxide levels compared to elevated carbon dioxide levels (P=0.07). In a dry year, there was no significant difference in Nepalese browntop biomass between carbon dioxide treatments. In contrast, Japanese honeysuckle, a common nonnative associate of Nepalese browntop, produced 3 times as much biomass under elevated carbon dioxide levels in both wet and dry years .Raunkiaer  life form:
Nepalese browntop phenology
|Illinois||seedlings establish||May |
|disperses seed and dies||October-November |
|New Jersey||seedlings emerge||late March-late April [28,30]|
|New York, Central Park||flowers||early September |
|disperses seed and dies||October [12,200]|
|Ohio, southern||germinates||mid-June |
|Eastern United States||fruits||September-October [15,125,201]|
|disperses seed and dies||September-December |
|greenhouse||seedlings emerge||early May|
|plants die||September-early October |
Pollination and breeding system: Nepalese browntop is both self- and cross-pollinated . Chasmogamy and cleistogamy are noted in nonnative Nepalese browntop populations in United States populations  and native populations in Asia [105,193]. Soil moisture and light intensity may affect flower development and breeding. In a population near Charlotte, North Carolina, Barden  found about 10% of plants had chasmogamous flowers, with chasmogamous plants mostly growing in moist, open sites. All Nepalese browntop plants growing in heavy shade had cleistogamous flowers. In a southern Illinois population with 80% overstory cover, flowers were mostly cleistogamous . In New York, chasmogamous flowers were most common in shady forests interiors. The ratio of cleistogamous:chasmogamous flowers increased in the greenhouse .
Genetic studies in the James River Basin of Virginia suggest considerable gene flow among populations, although other studies show interpopulation differences. In the Virginia study, genetic diversity was higher than expected for an introduced species that can self-pollinate, and the author speculated that cross-pollination within and among populations is common in Nepalese browntop. There was genetic evidence of long-distance dispersal of Nepalese browntop outside the James River Basin . In a greenhouse study, Nepalese browntop showed significant differences among families in the number of tillers produced (P<0.0001) but not in growth rates . Genetic differences in specific leaf area have been noted among populations  (see Physiology).
Seed production is generally high for Nepalese browntop [34,215]. Each Nepalese browntop tiller typically produces 1 terminal raceme and 2 to 7 axillary racemes . Consequently, a single tiller can bear many flowers, and a single Nepalese browntop tiller may produce 100 to 1,000 seeds [56,215]. Seed production varies between years and among populations, however. Based on a study in Great Smoky Mountains National Park, Williams  estimated that a single Nepalese browntop plant averages 77 seeds of 80% to 90% viability. A southern Illinois study found a mean of 81.7 spikelets/Nepalese browntop culm. However, spikelet production for 4 Nepalese browntop populations varied significantly among populations (P<0.001), and seed viability was generally low (33%). The study was conducted during a drought year (1999); even so, seed rain averaged 24.6 seeds/m² (n=34 seed traps) . In New Jersey, Cheplick [29,31] found seed production (both cleistogamous and chasmogamous) averaged about 72 seeds/tiller. The number of tillers produced varied among family lines .
Late-season drought can greatly reduce or eliminate Nepalese browntop seed production for a cohort , and seed production is reduced in low light. In West Virginia, Nepalese browntop production of chasmogamous flowers in a dry year was significantly higher in a mesic mixed-hardwood forest than in a dry oak-hickory forest (P=0.03), but there was no significant difference in a year of normal precipitation . In the greenhouse, Nepalese browntop produced significantly more fertile spikelets with full sunlight than with 21% or 10% full sunlight (P<0.05). There was no significant difference in Nepalese browntop fecundity between the 2 lower levels of sunlight . In oak-hickory forests of West Virginia, Nepalese browntop seed production was significantly higher along roadsides than in forest interiors .
Greenhouse and field experiments showed that Nepalese browntop produces some seed in shade. In the greenhouse, Nepalese browntop in 2% to 8% of full sunlight produced fewer chasmogamous and cleistogamous flowers and allocated more biomass to leaves compared to plants raised in full sunlight. Field trials produced similar results. In sweetgum-red maple-pin oak (Quercus palustris) forests in New Jersey, Nepalese browntop plants under the forest canopy produced fewer flowers (P≤0.002) and more leaves (P<0.003) than Nepalese browntop plants on forest edges. Relative percent of chasmogamous and cleistogamous flowers was similar under the canopy and on forest edges (16% and 11% vs. 6% and 7% of total aboveground biomass for under-canopy and edge locations, respectively) .
Seed dispersal: Reviews indicate that wind, water, animals, and humans disperse Nepalese browntop seed [17,173,191,201,220]. Nepalese browntop often occurs on floodplains [12,68]. Its close association with sites disturbed by heavy machinery implicates machines, fill dirt, and contaminated hay as potential dispersal agents of Nepalese browntop seed [17,191,201]. Rivers, ditches, and roads appear to be primary corridors for population expansion . Nepalese browntop fruits are light and float easily and the seeds may survive and germinate after "extended periods" of inundation , so flooding is a likely means of seed dispersal (see photo above). Nepalese browntop cover was greatest on disturbed (developed or frequently mowed) floodplains near the Mississippi River ; however, frequent, severe flood scouring can limit Nepalese browntop establishment and spread . Awned fruits can catch on fur, feathers, and clothing [17,191,201], but because the fruits are small, even awnless fruits can work their way into fur and clothing . A review reports that Nepalese browntop seeds often attach to hikers' clothing .
Nepalese browntop spreads from roads and trails into wildlands [32,117,130,194], but it usually disperses poorly without dispersal agents. Several studies show that roadways function as both corridors of dispersal and favorable germination sites for Nepalese browntop. On multiple sites in southern Ohio, Nepalese browntop was locally abundant in small gravel piles left by road graders and along streams and other water channels . In oak-hickory forests of southern Ohio, Nepalese browntop established along roadsides. Dispersal apparently occurred when contaminated gravel was spread; further dispersal occurred from water running through spread gravel. The author speculated that running water disperses Nepalese browntop from roadsides into forests. Nepalese browntop seeds, which in this experiment had no awns and therefore no apparent adaptations for dispersal, dispersed better than seeds of nonnative multiflora rose (Rosa multiflora), which has animal-dispersed seeds, and nonnative coltsfoot (Tussilago farfara), which has wind-dispersed seeds . A Nepalese browntop 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 by hikers . In mixed-hardwood communities in the Blue Ridge Mountains of North Carolina, Nepalese browntop presence was positively correlated with proximity to streams, closed-canopy sites, and developed sites (P<0.05) . In the Green Ridge State Forest, Maryland, Nepalese browntop presence was positively associated (P<0.001) with sites 30 to 490 feet (10-150 m) from roads . On the Daniel Boone National Forest, Kemtucky, Nepalese browntop spread onto new roads, onto old roads after road grading, and along streambanks after stream restoration. In all cases, Nepalese browntop spread from small source populations on sites where heavy equipment was used .
In closed-canopy forests on the Monongahela National Forest, West Virginia, soil-stored Nepalese browntop seed was found only within 30 feet (10 m) of roadsides, although patches of Nepalese browntop were found in forest interiors. The author surmised that secondary seed dispersal accounted for Nepalese browntop establishment in interior locations. Average seed spread rate across locations was 0.95 foot (0.29 m)/year ; possible methods of dispersal were not investigated.
Seed banking: Nepalese browntop seeds apparently have short-term persistence in soil [12,68,181,212]. Longevity of soil-stored seed is usually estimated at 3 to 5 years [12,68,201], although one author suggests that seeds may live less than 1 year . On a North Carolina site, soil-stored Nepalese browntop seed remained viable for at least 3 years. On a sloped seep in Pennsylvania, most viable Nepalese browntop seeds were collected at 0- to 4-inch (10 cm) depths. Most often, Nepalese browntop seed was found in the soil when Nepalese browntop plants were present in aboveground vegetation . In the greenhouse, a mean of 87.5 Nepalese browntop seedlings emerged from 400-cm² soil samples, which were collected in a red maple forest in Arkansas .
Nepalese browntop seed occurs in waterlogged soils and along waterways as well as in soils beneath upland plant communities. Nepalese browntop seed was collected from the seed bank of a tidal freshwater marsh along the Delaware River in New Jersey [109,110]. In swamplands of the Delaware River, Nepalese browntop appeared to be replacing native sedges (Cyperaceae) in the ground layer . By the Potomac River in Virginia, Nepalese browntop seeds were collected during spring from the seed bank of the high-drift shoreline. Nepalese browntop seeds were not found on the driftline in other seasons, and Nepalese browntop seeds were not found in any season by trawling along the water surface. The plant community above driftline was a narrowleaf cattail-arrow arum (Typha angustifolia-Peltandra virginica) tidal freshwater marsh .
Occasional seed crop failure is probably not limiting for this species. Given a persistent seed bank, Nepalese browntop may establish in high densities the year following poor seed production .
Germination: Although seed production can be high [34,215], few seed germination studies had been conducted as of this writing (2010), so Nepalese browntop germination requirements are unclear. On some sites, Nepalese browntop appears to require cold stratification (review by ), which is accomplished in the field by overwintering. A greenhouse study using seed from North Carolina found that fresh seed was not immediately germinable, while seeds stratified for 90 days showed 100% germination . However, Williams  reported immediate germination of Nepalese browntop seed collected in Great Smoky Mountains National Park.
Open sites and little to no litter favor Nepalese browntop germination. In oak-hickory forests of southern Ohio, Nepalese browntop germination in general was higher on open than on closed-canopy sites. Roadsides were particularly favorable for germination; Nepalese browntop seed sown along roadsides showed significantly better germination than seed sown in closed-canopy forest (P<0.05) [32,120]. Matlack  found that Nepalese browntop "completely saturates the roadsides in which it occurs". In a white oak-yellow-poplar forest in Tennessee, litter removal down to mineral soil or litter removal to mineral soil plus mineral soil disturbance significantly increased Nepalese browntop spread compared to plots with undisturbed litter (P=0.05) . On the Wayne National Forest, Ohio, seedlings rarely occurred on plots with deep litter; they were concentrated on microsites with exposed mineral soil (P<0.003) . In another experiment on the Wayne National Forest, forest floor disturbances that reduced litter were the most important factor in successful Nepalese browntop germination (abstract by ).
Seedling establishment and plant growth: Nepalese browntop may initially establish in large numbers, experience high seedling mortality, then form thick lawns via vegetative expansion of remaining plants. In southern Illinois, Nepalese browntop established at a mean density of 43 seedlings/m². Plant mortality was greatest (≥50%) during seedling establishment (mid-March), dropping to about 20% by July . In central New Jersey, Nepalese browntop seedling density in March and April averaged 1,963 seedlings (SD 652)/m², and the seedlings averaged 2 to 6 inches (5-15 cm) in height . Barden  estimated the number of plants produced from 1983 to 1986 on a 2-m² plot in North Carolina averaged 1,000 (in 1983), 256 (1984), 44 (1985), and 0 (1986), respectively. Density of Nepalese browntop on another 2-m² study plot on the North Carolina site averaged 857 (in 1984), 47 (1985), and 29 (1986) .
Sunlight and moist soil increase the chances of Nepalese browntop establishment and favor its growth (review by ). Establishment and spread are limited in shaded environments . On shaded sites, more carbon is allocated to leaves and aerial stems than to stolons  and flowers . However, Nepalese browntop is well adapted to shady conditions. It can establish, grow, and produce some seed in as little as 5% of full sunlight .
In oak-hickory forests of West Virginia, Nepalese browntop was significantly taller along roadsides than within forest interiors; Nepalese browntop cover and spread were also higher along roadsides than in forest interiors . On rural and wildland sites in New Jersey, seedling emergence, growth, and seed production of sown Nepalese browntop seed was significantly greater on an open lawn than in an interior red maple-shagbark hickory-sweetgum woodland (P<0.05). Nepalese browntop density on the lawn averaged 1,573 plants/m², while density in the interior woodland averaged 709 plants/m². Seed production was positively correlated with light (r²=0.06, P>0.05) but not with Nepalese browntop density or soil moisture .
Litter apparently impairs Japanese seedling establishment [51,137,145]. In a landscape-level study of 3 white oak-sweet birch forests in New Jersey, sites with Nepalese browntop had less litter than adjacent uninvaded sites . In an oak-yellow-poplar plantation in southwestern Tennessee, plots where litter was removed in winter experienced 4.5 times the invasion of Nepalese browntop compared to plots where winter litter was left intact (P<0.001). At the end of the growing season, Nepalese browntop on plots without litter had spread an average 5.45 feet (1.66 m), while Nepalese browntop spread on plots with litter averaged 1.20 feet (0.37 m). Nepalese browntop cover averaged 48% and 5% on plots without and with litter, respectively. The authors suggested that increased light as a result of litter removal favored Nepalese browntop germination and growth . In a harvested white oak-yellow-poplar forest in Tennessee, Nepalese browntop spread was greater with litter removal or soil disturbance than on undisturbed sites [118,119]. Measured from plot edges, the distance at which 90% of Nepalese browntop plants occurred (P=0.02) and overall mean distance of Nepalese browntop spread (P=0.04) were significantly farther with litter removal than without. Outlier Nepalese browntop plants (those farthest from the population center) may be of greatest concern in terms of Nepalese browntop spread. The distance of outlier Nepalese browntop plants was significantly farther in litter-removal and soil-disturbance plots than control plots (P=0.02) The authors suggest that disturbing litter may increase Nepalese browntop invasion and spread in eastern hardwood forests, while leaving litter layers intact may slow Nepalese browntop invasion .
While litter may inhibit Nepalese browntop establishment, a greenhouse study suggests litter may not impede growth after seedlings establish. Using soils from oak-hickory and red maple forests of New Jersey, Ross  found that regardless of soil origin, leaf litter additions did not significantly decrease growth of established Nepalese browntop plants compared to soils without added litter. Additional greenhouse studies using soil from the 2 forests showed arbuscular mycorrhizae had no effect on Nepalese browntop growth. Nepalese browntop roots were susceptible to arbuscular mycorrhizal colonization, but Nepalese browntop height growth was similar with and without arbuscular mycorrhizal colonization .
Based on shade and litter manipulations in white oak-red oak-shagbark hickory, red maple-American elm, and white ash-yellow-poplar forests in New Jersey, Schramm and Ehrenfeld [179,180] suggested that deep litter, shade, or their interactions may limit Nepalese browntop spread (P=0.05 for all variables). Only seedlings with no litter or a litter layer one-half of average (~0.8 inch (2.2 cm)) showed "substantial" survivorship. There was a trend towards decreasing Nepalese browntop cover with increasing successional stage. Nepalese browntop was "effectively excluded" where American beech, a late-successional species that casts deep shade at maturity, dominated the canopy, while open, successional red maple- and white ash-dominated forests had 22% to 30% Nepalese browntop cover. Oak-hickory forest supported intermediate levels of Nepalese browntop (5-8% cover). Regardless of successional stage, there was a trend toward decreasing Nepalese browntop invasion with increasing stand size (r²=0.33) . The authors suggested that generally, loss of the shrub layer due to heavy white-tailed deer browsing could accelerate Nepalese browntop spread [179,180]. Interactive effects of white-tailed deer and Nepalese browntop on stand structure and plant species composition are discussed further in Impacts; see Successional Status for further information on Nepalese browntop and shade.
Rauschert and others  present a model of Nepalese browntop population growth based on broadcast seeding experiments in an oak-hickory-eastern white pine forest in Pennsylvania.
Several other site characteristics, and stand structure, apparently affect Nepalese browntop regeneration. In North Carolina, Nepalese browntop regeneration was negatively correlated with high soil pH (5.5 vs. a median of 5.1); high levels of soil potassium, zinc, and calcium; high percent silt (18% vs. 10%); deep litter (8.6 vs. 5.5 cm); high cumulative PAR on an overcast day (0.72 vs. 0.57 mol/m²/day); and high leaf area index (LAI) of other species (1.3 vs. 0.7) . In southern Illinois, reproductive success was correlated with soil conditions and canopy cover. Reproduction increased with increasing availability of soil cations and sand content and decreased with increased soil silt content and canopy cover (P<0.05 for all variables) .
Vegetative regeneration: Within a growing season, Nepalese browntop increases vegetatively by tillering [30,90] and by stolons , sometimes forming dense, monospecific stands through vegetative spread . Because Nepalese browntop is an annual, the vegetative shoots do not survive through the next growing season . High vegetative biomass does, however, increase the likelihood of reproductive success by increasing photosynthate gain and thus the potential for high seed production. High light and other favorable conditions maximize vegetative growth .SITE CHARACTERISTICS:
Nepalese browntop is strongly associated with disturbed forest sites, especially roads. The Virginia Department of Conservation and Recreation  stated that Nepalese browntop is common on disturbed soils and can rapidly spread onto undisturbed soils once established nearby. In white oak-eastern hemlock forests of Pennsylvania, Nepalese browntop was about 7 times more likely to occur on disturbed than on undisturbed sites . In the Green Ridge State Forest, Maryland, Nepalese browntop presence was positively associated with disturbed soils (P<0.001) . In sweetgum-sycamore and loblolly pine-white oak-sweetgum forests of Mississippi, Nepalese browntop was positively associated with canopy gaps and flooding (P<0.001 for both variables) . On 2,000 sites within oak-hickory forests of western Virginia, Nepalese browntop cover was positively related to road length (P=0.04) and length of the road relative to total area of the watershed in which it occurred (P<0.001). Nepalese browntop was rare in forest interiors relative to its abundance on roadsides, and Nepalese browntop by roads gained more biomass than Nepalese browntop growing in forest interiors (P<0.001) .
In a seeding experiment in an oak-hickory-eastern white pine community in Pennsylvania, Nord and others  concluded that disturbance and soil properties were more important to successful Nepalese browntop invasion of a site than the plant community type. They found that litter disturbance increased Nepalese browntop population expansion for the first 2 years of Nepalese browntop invasion compared to sites with undisturbed litter (P<0.02) and that populations consistently declined on closed-canopy sites. Disturbance × environment interactions were not significant for Nepalese browntop population growth . See Nutrients for more information on this study.
Soils: Nepalese browntop prefers damp or wet soils (, review by ), although it does not tolerate standing water for "extended periods" of time (review by ). It also establishes on dry upland soils . On the Jefferson National Forest and in Mountain Lake Wilderness, Virginia, Nepalese browntop occupied either damp sites without standing water or sites with "highly disturbed" soils such as gravel and dirt mounds by roadsides . In southern Ohio, Nepalese browntop was "particularly dense and vigorous" in swales and moist soil . In a yellow-poplar-common persimmon-sweetgum forest in North Carolina, Nepalese browntop successfully "outcompeted" native understory species on floodplains and midslopes but not on upland sites . In Florida, Nepalese browntop is common on wet hammocks . In the greenhouse, Nepalese browntop's relative growth rate was fastest in soil with 30% water content (P<0.05), but it persisted and produced some seed in flooded soils and in soils with <10% water content. The authors attributed Nepalese browntop ability to invade a site, in part, on its ability to tolerate "contrasting and extreme soil water conditions" .
Nepalese browntop is common on silty to sandy loams [12,56,68,165] and on clays [56,90]. In deciduous wetlands of New Jersey, Nepalese browntop was positively correlated with percent clay in soil (P<0.05) . Nepalese browntop an indicator of red clay soils in the Piedmont region .
Soil pH is usually mildly acidic to basic on sites with Nepalese browntop [56,68,201]. A survey in Maryland and Washington, DC, found that sites with Nepalese browntop ranged from pH 4.8 to 5.8 . On mine spoils in Kentucky, Nepalese browntop grew on loamy soils with pH ranging from 4.6 to 6.3. It was absent from an extremely acidic site (pH 4.4) . In an Illinois study, soils supporting Nepalese browntop were generally acidic and nutrient poor .
Some studies have found that Nepalese browntop was positively associated with basic soils [35,137] or that it raises soil pH . In deciduous wetlands of New Jersey, Nepalese browntop was positively correlated with nonacidic soils (P<0.05) . In white oak-eastern hemlock forests of Pennsylvania, sites most likely to support Nepalese browntop had basic soils and low understory cover . Studies in Tennessee oak-pine  and New Jersey oak-hickory  forests showed high soil pH favors Nepalese browntop, while a study in a oak-hickory forest of southeastern Ohio showed no significant increases in Nepalese browntop abundance with lime additions to soil . In mixed-hardwood forests of New Jersey, there was no significant relationship between Nepalese browntop invasion and soil pH .
Nutrients: Based on limited studies, Nepalese browntop may prefer soils with high mineral content. In an oak-hickory-eastern white pine community in Pennsylvania, phosphorus level (P=0.01), potassium level (P=0.01) moist soil (P<0.001), and high pH (P=0.002) were positively associated with Nepalese browntop abundance, while ammonium was negatively associated with Nepalese browntop abundance and seed production (P<0001) . Studies in Maryland and Washington, DC, found higher levels of nitrogen and average levels of potassium and phosphorus on Nepalese browntop-infested soils compared to soils without Nepalese browntop . In red maple forests of Arkansas, Nepalese browntop was positively correlated with high concentrations of soil boron (r=0.3) and zinc (r=0.5). In mixed-hardwood and oak-hickory forests of West Virginia, soils of interior plots with Nepalese browntop had significantly lower total carbon levels than plots without Nepalese browntop (P=0.07) . In mixed-hardwood forests of New Jersey, however, sites where soils had high organic matter content were more susceptible to Nepalese browntop invasion than sites with low organic matter content .
Elevation and aspect: Nepalese browntop occurs from sea level up to 4,000 feet (1,000 m) elevation [56,125]. It is most common in low-elevation woodlands in the mid-Atlantic states and in the Piedmont and Appalachian mountains . As of this writing (2010), it was not reported from high-elevation red spruce-Fraser fir (Picea rubens-Abies fraseri) forests. In mixed-hardwood communities in the Blue Ridge Mountains of North Carolina, Nepalese browntop was negatively correlated with high elevation (P<0.05) .
Few studies had been conducted on possible aspect preferences of Nepalese browntop as of 2010. In the Green Ridge State Forest, Maryland, Nepalese browntop presence was significantly positively associated with southwest (P<0.001) and northwest (P<0.05) aspects . Nepalese browntop transplanted into canopy gaps in a New Jersey boxelder-green ash-sycamore forest showed better growth on the west side of the gaps compared to the east side .
Climate: Nepalese browntop grows in temperate to warm continental climates. In North America, the coldest reported winter temperatures that Nepalese browntop survives are approximately -5.8 to -9.4 °F (-21 to -23 °C) .SUCCESSIONAL STATUS:
Little English-language literature on succession in plant communities where Nepalese browntop is native was available as of 2010. In Japan, Nepalese browntop and other annual grasses typically dominate warm-temperate Chino bamboo (Pleioblastus chino) grasslands that are in early succession .
The following discussion applies only to plant communities in the eastern and southeastern United States.
Early succession: Nepalese browntop generally obtains greatest cover on open, seral sites or in canopy gaps [56,173] (see Late succession for information on canopy gaps). Open, early-seral sites in which it has established include old fields [7,68], active floodplains , minespoils , hurricane-disturbed sites , plantations , thinned  or clearcut  forests, burned woodlands and forests ([7,12], Shimp 2002 personal communication in ), and especially, forest edges [28,169]. In Great Smoky Mountains National Park, for example, Nepalese browntop was most invasive on forest edges . On abandoned surface coal mines in Kentucky, Nepalese browntop was the most important understory herb in early succession of a mixed-hardwood, mesophytic forest, forming 9% to 35% cover. It formed thick swards in open areas . In mixed-hardwood and oak-hickory forests of West Virginia, Nepalese browntop presence was associated with several indicators of early forest succession, including open canopies (P<0.001), high moss (Bryopsida) cover (P<0.001)), shallow litter (P=0.15) cover, and low levels of coarse woody debris (P=0.003) . In the Oak Ridge National Environmental Research Park, Tennessee, Nepalese browntop seedling survivorship averaged 100% in full sunlight; 90% in 40% sunlight; 30% in 16% sunlight; and 5% in 6% sunlight. Biomass gain over the May to October growing season was significantly greater at 100% sunlight than at lower light levels (P<0.001). The forest overstory was dominated by sycamore, boxelder, and black walnut .
Dry climate may favor Nepalese browntop invasion on old fields of the eastern Unites States. On old fields in the Hutcheson Memorial Forest, New Jersey, Nepalese browntop cover increased after a severe drought in 1999, when April and May rainfall was less than half of normal. Across plots, Nepalese browntop increased in total cover from a predrought level of 0.01% in 1997 to a postdrought level of 646.6% in 2001. During that time, Nepalese browntop increases in cover and frequency were greater than those of any other species in the old fields . Since then, Nepalese browntop has become the dominant groundlayer species in Hutcheson Memorial Forest (Yurkonis 2006 personal observation cited in ).
Disturbance ecology: Nepalese browntop 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 [191,215]. A survey (based on herbaria collections and remote-sensing data) of weed invasion patterns in West Virginia showed that Nepalese browntop was most common in roadside and streamside vegetation .
Nepalese browntop can recover rapidly—and may increase—after flooding (but see Gibson and others ). The input of silt and nutrients that accompanies short-term flooding can promote Nepalese browntop growth. For example, a study was initiated in 1982 on the Big Cross Creek floodplain of North Carolina. Big Cross Creek flooded in 1983, temporarily reducing Nepalese browntop cover, but Nepalese browntop exceeded preflood cover within 2 years .
|Nepalese browntop cover before and after flooding in North Carolina |
|1982 (preflood)||1983 (postflood)||1985 (postflood)|
By studying a boxelder-white ash-sycamore floodplain community in North Carolina, Barden  concluded that a history of disturbance was likely to improve Nepalese browntop's ability to invade a site. A relatively deep litter layer, greater LAI of other ground-dwelling species compared to Nepalese browntop, and high levels of sunlight reduced reproductive success of Nepalese browntop. He found that soil fertility was relatively unimportant in determining invasive ability of Nepalese browntop. Nepalese browntop failed to regenerate on undisturbed, fertile plots (high levels of soil nitrogen, potassium, calcium, and zinc). It showed greater biomass gain on plots treated with fertilizer compared to unfertilized control plots, but seed spikelet production was similar on fertilized vs. unfertilized plots .
It is unclear how vulnerable undisturbed sites are to Nepalese browntop invasion, and what factors, if any, contribute to a site's invasibility. Anecdotal evidence suggests that Nepalese browntop may not invade, or is slow to invade, undisturbed sites. However, long-term studies are needed to document Nepalese browntop's rate of colonization and expansion onto disturbed sites. A fact sheet suggests that Nepalese browntop may slowly spread onto undisturbed lands unless control measures are taken . Nepalese browntop was absent from unmowed land next to a sewer line right-of-way in North Carolina, but invaded annually mowed land near the right-of-way . An inventory of Land Between the Lakes National Recreation Area, Kentucky and Tennessee, showed Nepalese browntop 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 Nepalese browntop seed dispersal and invasion . A review suggests that Nepalese browntop can spread rapidly onto undisturbed sites from adjacent disturbed sites where it is well established . In a New Jersey survey, Nepalese browntop and garlic mustard were the only 2 nonnative species that invaded undisturbed chestnut oak-red oak-pitch pine stands .
Midsuccession: Nepalese browntop is common in midsuccessional forests. In the Green Ridge State Forest, Maryland, Nepalese browntop presence was positively associated (P<0.05) with moderate (26-50%) canopy openings . In Great Smoky Mountains National Park, mean height of Nepalese browntop stands peaked at 30% to 40% sunlight and decreased slightly after that. However, biomass of individual Nepalese browntop plants increased linearly with percent sunlight (P<0.001) . In Maryland, Nepalese browntop infestations were common on shaded roadsides but not open roadsides .
Late succession: Nepalese browntop is shade tolerant [45,56,83] and can persist in late-successional forests as the canopies close [5,83,169,181]). In mixed-hardwood communities in the Blue Ridge Mountains of North Carolina, Nepalese browntop presence was positively correlated with forest cover (P<0.05) . Nepalese browntop may form patches or dense, continuous lawns in late-successional forests . An invasive species guide reports Nepalese browntop can persist in <5% sunlight . Cheplick  reported Nepalese browntop on edges and under completely closed canopies of sweetgum-sycamore and loblolly pine-white oak-sweetgum forests of Mississippi, and Nepalese browntop was an understory component in old-growth sweetgum-overcup oak (Quercus lyrata)-river birch bottomland forests of Tennessee .
In late succession, Nepalese browntop usually occurs in canopy gaps. In a bottomland box elder-yellow-poplar-sycamore forest in Indiana, Nepalese browntop was positively correlated with light availability (r²=0.49, P=0.04) . Hemlock wooly adelgid infestations [54,55] or other canopy-opening events may provide favorable open sites for Nepalese browntop invasion. In eastern hemlock forests with high mortality from hemlock wooly adelgids in Connecticut, several nonnative species showed high cover including Nepalese browntop, Oriental bittersweet, Japanese barberry, and tree-of-heaven . In mixed-hardwood forests of North Carolina, vegetation frequency was surveyed on 107 permanent plots established in 1977 and resurveyed in 2000. Nepalese browntop had the second-highest increase in overall frequency during those 23 years; native American pokeweed (Phytolacca americana) increased most in frequency. Nepalese browntop was particularly abundant in open forest patches created by Hurricane Fran . Based on field experiments, Cheplick  reported that Nepalese browntop may persist or spread in late-successional hardwood communities where sunflecks reach photosynthetic Nepalese browntop tissue.
Apparently, Nepalese browntop does not typically invade closed-canopy forests lacking canopy gaps . Field experiments in Kentucky showed Nepalese browntop was unable to establish under the subcanopy, which consisted of juvenile red maple and spicebush. The oak-hickory forest was in late succession, and PAR was 1% to 2.5% of full sunlight beneath red maple and spicebush (abstract by ).
Nepalese browntop may alter successional pathways of forests in mid- to late succession. It grows quickly; Nepalese browntop is soon taller than the seedlings of most associated woody species and likely outcompetes young, native woody species and herbs for light . In an oak-beech-maple forest of Lilly-Dickey Woods, Indiana, Nepalese browntop gained significantly more aboveground biomass than the native grass deertongue (Dichanthelium clandestinum) in fully shaded sites, while deertongue gained more biomass than Nepalese browntop in full sunlight (P<0.001). Biomass of the 2 grasses was similar in partial shade . Nepalese browntop may establish in gaps that were historically colonized by oaks, hickories, ashes, and other early-seral tree species . Aronson  speculated that in young-secondary oak forests, environmental changes associated with Nepalese browntop invasion, such as increased soil pH and soil nitrogen, may facilitate invasion of other nonnatives. On the Hutcheson Memorial Forest, New Jersey, Nepalese browntop was negatively correlated with cover of other nonnative species in young-secondary white oak-black oak-red oak forests. However, in mature-secondary and old-growth white oak-black oak-red oak forests, Nepalese browntop presence was positively correlated with cover of other nonnative species (P<0.05 for all variables). Young-secondary, mature-secondary, and old-growth forests were 50, 150, and ~300 years old, respectively. Overall, Nepalese browntop and garlic mustard were the dominant groundcover species at Hutcheson Memorial Forest. From 1950 to 1979, importance value of Nepalese browntop was 0, but it jumped to 32 by 2003 . See the Impacts section for more information on other examples of Nepalese browntop's potential to alter forest succession.White-tailed deer and Nepalese browntop may synergistically alter successional pathways in eastern deciduous forests with dense white-tailed deer populations . See Impacts for more information on this relationship.
Immediate fire effect on plant:
As of this writing (2010) there were few accounts in the literature regarding the effects of fire on Nepalese browntop, and available information was mostly
anecdotal. As an annual, Nepalese browntop is likely killed by late-season fires , although spring [56,201] and summer  fire may only top-kill Nepalese browntop. Accounts of postfire establishment provided by
Barden  and Shimp (personal communication cited in ) suggest that Nepalese browntop seeds in the soil seed bank are likely to
survive fire. However, information on the fire ecology of Nepalese browntop is limited, and research is needed to clarify fire effects on Nepalese browntop.
Postfire regeneration strategy :
Caudex or an herbaceous root crown, growing points in soil
Stolons in organic soil or on soil surface
Ground residual colonizer (on site, initial community)
Initial off-site colonizer (off site, initial community)
Secondary colonizer (on- or off-site seed sources)
Fire adaptations and plant response to fire:
Fire adaptations: As an annual, Nepalese browntop likely relies mostly on postfire establishment from either on-site, soil-banked seed or off-site, transported seed. As of 2010, there were limited studies [6,7,69] and anecdotal accounts [12,68,189] of postfire Nepalese browntop establishment; however, details were few. Nepalese browntop may establish from seed on mineral soil after fire . It spread after either litter removal down to mineral soil or litter removal and mineral soil disturbance in Tennessee . In at least one account, Nepalese browntop likely established from soil-stored seed following a "hot" surface fire  (see Plant response to fire). Given its ability to store seed in the soil seed bank, effectively disperse seed, and establish on open, disturbed sites (see Successional Status), Nepalese browntop is likely to persist or invade after fire.
Plant response to fire: Details of Nepalese browntop postfire establishment were lacking in available literature (2010). Because it is an annual, this grass must establish from soil-stored seed and/or off-site seed transported onto burned sites after late-season fire ([7,12,68], review by ). A review by Tu  suggests that following early-season fire, top-killed Nepalese browntop may sprout and set seed later in the year (see Seasonal Development). According to a management guide for the southern United States  and Tu , Nepalese browntop that has not yet flowered may sprout from tillers and stolons following top-kill by fire . A second crop of seedlings may establish after spring fire . A review indicated that exposed mineral soils, such as those occurring after fire, provide a favorable seedbed for Nepalese browntop germination and establishment .
Nepalese browntop benefits from disturbances that open the canopy (see Successional Status); this likely includes fire . A few studies demonstrate Nepalese browntop's ability to establish in postfire environments.
In oak-hickory and sugar maple-sweetgum-yellow-poplar communities of the Vinton Furnace Experimental Forest, Ohio, either mechanical litter removal or prescribed fires (both low and moderate severity) increased Nepalese browntop seedling establishment and growth compared to control plots (P<0.05 for all variables) . Burned plots were sown with Nepalese browntop seeds in postfire year 1; litter-disturbed plots were also sown at that time. Nepalese browntop was removed prior to seed set to prevent invasion beyond study plots. In postfire year 2, seeds were sown in different burned plots that had previously been sown with multiflora rose but not Nepalese browntop. On burned plots, Nepalese browntop stem height and leaf number were greatest in canopy gaps on moderate-severity plots (P<0.05). August surveys revealed year and site interactions in Nepalese browntop's response to prescribed fire. In postfire year 1, Nepalese browntop seedling establishment was greatest on burned or litter-removed plots (P<0.0001). In postfire year 2, seedling establishment was greater in valley plots, where sugar maple tended to dominate, than on ridges, where oaks tended to dominate (P<0.01) . The authors concluded that prescribed fire created a disturbance suitable to Nepalese browntop invasion (, abstract by Glasgow and Matlack ), and litter removal was the mechanism by which fire enhanced Nepalese browntop seedling recruitment . See the Research Project Summary of this study for details on the fire prescription, fire behavior, and postfire responses of Nepalese browntop and multiflora rose.
Nepalese browntop invaded a remnant prairie after thinning and prescribed burning on the LaRue-Pine Hills Research Natural Area, Illinois [6,7]. See Preventing postfire establishment and spread for details.
There are several anecdotal accounts of postfire Nepalese browntop recruitment. In a boxelder-white ash-sycamore floodplain community in North Carolina, a 9 April 1982 prescribed fire entered a dense upland stand of Nepalese browntop seedlings. The previous year's cohorts had left a dense mat of Nepalese browntop litter that fueled "a hot ground fire" that killed the seedlings. By mid-June, a second cohort of Nepalese browntop had established, presumably from soil-stored seed, and provided dense ground cover . Gibson and others  reported "increased recruitment" of Nepalese browntop following prescribed fire in a xeric, early-successional oak-hickory woodland that established on old fields abandoned in the 1960s (Shimp personal communication cited in ). In black oak-blackjack oak-post oak forests of northern Mississippi and western Tennessee, Surrette  found that Nepalese browntop was more abundant on spring-burned (March-April) plots compared to unburned plots. The author speculated that Nepalese browntop cover increased because the prescribed burning immediately preceded the time of Nepalese browntop germination .FUELS AND FIRE REGIMES:
Fuels: Live Nepalese browntop may be difficult to burn. Its low flammability and relative unpalatability suggest that it has high silica content, which could reduce its ability to carry fire when green .
As of 2010, measurements of Nepalese browntop fuel loads in northeastern or southeastern forests were not available in the literature. Nepalese browntop's ability to exclude woody species and form thick ground cover suggest that it may increase fine fuels and reduce woody debris from historical levels. However, Kourtev and others  reported that in New Jersey, sites invaded by Nepalese browntop had thinner litter and organic soil layers than sites without Nepalese browntop, which they attributed to high densities of nonnative earthworms on sites with Nepalese browntop (see Soil and soil microfauna changes for more information). Similarly, in white oak forests of New York, Nepalese browntop-invaded sites had thinner organic soil horizons than adjacent uninvaded sites . Nepalese browntop litter tends to decay slowly, which may increase fine fuels compared to sites with litter of faster-decaying native species.
As an annual, mat-forming grass, Nepalese browntop often produces large amounts of fine litter that may remain on the forest floor longer than litter of some native plant species. Nepalese browntop stems lodge soon after they die in autumn [12,27]. When thick, they create a continuous fuelbed of matted straw that could potentially fuel a surface fire . Nepalese browntop litter apparently decays more slowly than litter of some associated species . In a New Jersey study, Nepalese browntop litter decayed more slowly than litter of native hillside blueberry . In a North Carolina field experiment, litter of nonnative Oriental lady's-thumb (Polygonum caespitosum) was about 30% decayed after 120 days, while Nepalese browntop was only about 5% decayed [41,43]. However, in a landscape-level study of 3 white oak-sweet birch forests in New Jersey, sites with Nepalese browntop had less litter than adjacent uninvaded sites. Over 2 years, the on-site decay rate of white oak litter was slower (30% mass loss) than decay rates for Nepalese browntop litter (40%-50%) .
Dibble and others  reported that standing dead and down litter of Nepalese browntop and other nonnative invasive grasses may present a fuel hazard in drought years. Flammability of live Nepalese browntop, however, may be low. In the laboratory, Nepalese browntop's heat of combustion was among the lowest of 42 native and nonnative species in the Northeast . A management guide for the southern United States reports that Nepalese browntop is not a fire hazard .
In mixed-hardwood and oak-hickory forests of West Virginia, interior forest plots with Nepalese browntop had significantly lower coarse woody debris cover than plots without Nepalese browntop (P<0.003) .
Fire regimes: Across Nepalese browntop's US distribution, fire regimes vary from frequent surface fires to long-interval, stand-replacement fires. 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 [57,65,77,177,216]. Oak-hickory, oak-pine, and pine forests of the Northeast and Southeast had mostly frequent understory surface fires [190,216]. See the Fire Regime Table for further information on fire regimes of vegetation communities in which Nepalese browntop may occur. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".
Nepalese browntop was not present in these forests while historic fire regimes were still operating, and it is unclear how Nepalese browntop may affect or alter fire regimes in plant communities where it is present. Nepalese browntop's tendency to invade disturbed forests (see Successional Status), its ability to produce abundant litter that decays slowly, and its potential to reduce establishment of woody species and form monocultures—thereby altering stand structure (see Impacts)—make it likely that Nepalese browntop alters fuel loads and fire behavior from historic patterns. Further fire studies on Nepalese browntop and observations of fire behavior where Nepalese browntop is present are needed.FIRE MANAGEMENT CONSIDERATIONS:
Potential for postfire establishment and spread: Nepalese browntop's autecology suggests that it is likely to invade burns. It favors disturbed, open sites and mineral soil for establishment (see Regeneration Processes) and once present, tends to displace native vegetation (see Impacts). Postfire establishment is especially likely on burns subject to foot, motor vehicle, and other traffic that transports Nepalese browntop seeds onto the burn (see Seed dispersal). Romanello  reported that Nepalese browntop was most likely to establish from the soil seed bank if present before disturbance, suggesting postfire Nepalese browntop establishment can be expected where Nepalese browntop was present before fire. Based on reports to date (2010), groundlayer dominance of Nepalese browntop has been greatest in yellow-poplar-sweetgum communities; however, given Nepalese browntop invasion and spread in a wide range of forest and some shrubland and grassland types in the eastern and southern United States (see Habitat Types and Plant Communities), most low- to midelevation sites can be considered vulnerable to postfire Nepalese browntop invasion.
Preventing postfire establishment and spread: Preventing Nepalese browntop and other invasive plants from establishing in weed-free burned areas is the most effective and least costly management method. This may be accomplished through early detection and eradication, careful monitoring and follow-up, and limiting dispersal of invasive plant propagules into burned areas. General recommendations for preventing postfire establishment and spread of invasive plants include:
Nepalese browntop may require postfire control on sites where thinning and prescribed fire promoted its germination and spread. The LaRue-Pine Hills Research Natural Area of southern Illinois is a remnant little bluestem-indiangrass prairie barren that was historically maintained by frequent fires. The fires, probably intentionally set by Native Americans [1,40,66,67,192], maintained the barren by pruning woody vegetation to a bushy, scrub form. Forest Service personnel intermittently managed the Research Natural Area with fire from 1969 to 1993. That period included 16 years of fire exclusion (1974-1989), during which woody vegetation began invading the barrens. Restoration thinnings of white oak, southern red oak, common persimmon, and other woody species began in 1988. Annual prescribed burning was resumed in 1990. Nepalese browntop was first noted on woodland study plots in 1992 but was not found on similarly treated barren or woodland-barren transition area plots. The authors suggest that Nepalese browntop "was likely favored by the disturbance associated with mechanical removal of woody species and the reintroduction of prescribed burning" in the woodland [6,7].
Use of prescribed fire as a control agent: To date (2010), the available literature provided no accounts of successful control of Nepalese browntop using prescribed fire; however, there may potential for using prescribed fire to control Nepalese browntop under some circumstances and in combination with other treatments. For example, burning might be used to help reduce litter and standing plant biomass prior to herbicide application for Nepalese browntop control , although there is some question about whether Nepalese browntop will carry fire when green (see Fuels). Early-season fire does not control Nepalese browntop (Barden 1991 as cited by ); burned plants may sprout and seedlings may establish from soil-stored seed and produce new seed by the end of the growing season. Fall fire, when Nepalese browntop is flowering but before seed set (see Seasonal Development), may help control Nepalese browntop .
In Big Oaks National Wildlife Refuge, Indiana, late summer prescribed fire, spring prescribed fire, hand-pulling, and fall mowing were compared as control treatments for Nepalese browntop. Study sites were in second-growth American beech-black walnut-Virginia pine/northern spicebush forest with a history of prescribed fire. Late summer fires were ignited and mowing was conducted in early September after Nepalese browntop had set seed. Spring fires were ignited and hand-pulling started in June, when Nepalese browntop seedlings were 4 to 8 inches (10-20 cm) tall. Compared to untreated control plots, fall fire and mowing caused significant reductions in Nepalese browntop cover and biomass. Compared to controls, fall fires reduced Nepalese browntop cover by 79% and biomass by 90%, while mowing reduced cover by 70% and biomass by 95%. Spring fire significantly reduced Nepalese browntop cover but not its biomass (P<0.05 for all variables). Hand-pulling in spring did not significantly change Nepalese browntop cover or biomass. Native understory species showed no significant difference in cover or biomass on treated compared to control plots .Altered fuel characteristics: Nepalese browntop has the potential to increase litter, reduce woody debris, and alter stand structure where it is present. See Fuels and Impacts for further details.
Insects graze Nepalese browntop, although the extent of their use was largely unstudied as of 2010. In a red maple-white oak-sycamore forest in the Whitehall Experimental Forest, Georgia, some genera of short-horned grasshoppers, katydids, crickets, and bugs obtained a substantial fraction (35-100%) of their diet from Nepalese browntop. Sample sizes ranged from 1 to 10 individuals per insect genus. Insect guilds using early-successional forests may be more likely to use Nepalese browntop than insects using forests in later seres. In this study, invertebrates in canopy gaps (where Nepalese browntop forage is usually most abundant) tended to actively avoid capture and were mostly green, while invertebrates under closed canopies tended to remain still when detected and had cryptic coloration .
Nutritional value: No information was available on the nutritional content of fresh Nepalese browntop forage. Strickland and others  provide information on the nutritional content of Nepalese browntop litter.
Cover value: Nepalese browntop may provide important cover for white-footed mice. In loblolly pine-Virginia pine forests of Virginia, white-footed mice were more abundant on plots with than without Nepalese browntop. The author suggested that sites with Nepalese browntop may provide more nesting sites, nesting materials, and/or have decreased predation rates than sites without Nepalese browntop. White-footed mice were observed navigating through dense Nepalese browntop culms without difficulty, although they avoided areas with dense cover of native little bluestem. Among 6 other small mammal species, none were either positively or negatively associated with Nepalese browntop .
Nepalese browntop may reduce suitable cover and habitat quality for the federally threatened  bog turtle on old-field or resting pastures. In surveys of potential bog turtle habitats in New Jersey and New York, Nepalese browntop was present in <10% of wetland plots with bog turtles. On those plots, Nepalese browntop was significantly taller (3 feet (0.9 m)) in wetlands that dairy cattle had formerly grazed compared to its height (1 foot (0.3 m)) in ungrazed wetlands (P<0.01). Its cover was also greater in formerly grazed (3.9%) than in ungrazed (2.0%) wetlands, although the difference was not statistically significant. Overall, height of herbaceous species was lower and native species diversity higher on formerly grazed than ungrazed wetlands, and significantly more bog turtles were captured on formerly grazed than ungrazed wetlands (P=0.001) .
Nepalese browntop may reduce habitat quality of some tick species. In Indiana, experimentally introduced lone star ticks (Amblyomma americanum) and dog ticks (Dermacentor variabilis) showed higher mortality rates in Nepalese browntop-invaded plots than in plots without Nepalese browntop. In Nepalese browntop plots, mortality of lone star ticks and dog ticks increased 173% and 70%, respectively, compared to mortality in uninvaded plots. The authors attributed the higher death rates in Nepalese browntop plots to increased temperatures and decreased humidity at the soil surface and in litter compared to uninvaded plots .OTHER USES:
A 2003 review of vegetation surveys in the eastern United States revealed that Nepalese browntop was among the most commonly reported invasive species, and it was the most common invasive annual grass. It was most frequent on floodplains and in mesic forests . It was ranked a high invasive threat in deciduous, coniferous, and mixed forests, grasslands, old fields, riparian zones, and freshwater wetlands of the Northeast , and it was ranked a high to moderately-high threat in red oak and eastern hemlock forests of Delaware Water Gap National Recreation Area . As of 2000, the density of Nepalese browntop infestations in Dixon State Park, Illinois, ranged from 2.3 stems/m² to 16,706 stems/m² .
Surveys show that as of 2008, Nepalese browntop occupied about 650,000 acres (260,000 ha) in the Southeast , and it was most invasive in Tennessee, North Carolina, and northwestern South Carolina . It is ranked a high invasive threat in upland grasslands and oak-hickory woodlands and a potentially high threat in wet grasslands and palmetto (Arecacae) prairies . In the southern Appalachian region, 8 of 35 federal, state, and private agencies ranked Nepalese browntop among their greatest ongoing or potential management problems (behind kudzu (Pueraria montana var. lobata) and multiflora rose) . It was the most frequent (23%) of any nonnative species found in a 2006 survey of riparian forests in North Carolina . Surveys in mixed-hardwood communities in the Blue Ridge Mountains of North Carolina also found Nepalese browntop was the most frequent nonnative invasive species, occurring in 100% of watersheds and 84% of plots . In Oak Ridge National Environmental Research Park, Tennessee, Nepalese browntop 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 were ranked 2nd and 3rd, respectively . Nepalese browntop reportedly replaced existing ground vegetation in 3 to 5 years on sites in Great Smoky National Park , and it has formed "extensive and dense" infestations in Natural Areas and Parks, managed forests, wetlands, riparian areas, and rights-of-way in Alabama and adjacent states .
Because Nepalese browntop is an annual, its productivity is more closely tied to yearly climate fluctuations than that of perennial herbaceous species. Annual variations in Nepalese browntop productivity can have important effects on forest understory species composition and diversity. On a sweetgum site on the Oak Ridge National Environmental Research Park, Nepalese browntop produced 64% as much biomass in a wet year compared to a dry year . Using a model, Holcombe  predicts a gain of 51,400 miles² (133,000 km²) in Nepalese browntop cover in North America due to climate change.
Ecosystem function: Nepalese browntop is associated with changes in ecosystem function, including altered soil characteristics, changes in soil microfaunal composition, lowered plant and animal species diversity, and altered stand structure. These changes may interfere with growth and establishment of native and other invasive nonnative species. Nepalese browntop has also been implicated as being allelopathic. Sites with Nepalese browntop may also have less coarse woody debris and more fine fuels than uninvaded sites; this is discussed in Fuels.
Nepalese browntop may alter soil nutrient cycling [42,43,43,188], although some claim the already altered nutrient status of disturbed sites favors Nepalese browntop establishment . In a North Carolina wetland undergoing restoration, sites dominated by Nepalese browntop appeared to have decreased nitrogen cycling compared to sites where Nepalese browntop was removed. Decomposition and nitrogen release from Nepalese browntop litter was about half that of litter of native groundlayer species, and species richness was significantly less on invaded plots than on plots where Nepalese browntop was controlled [42,43]. DeMeester  concluded that compared to native species, Nepalese browntop "is clearly superior in capitalizing resources and suppressing other vegetation". In oak-pine forest in Whitehall Experimental Forest, Georgia, carbon apparently cycled more quickly sites with Nepalese browntop than on sites without Nepalese browntop. Plots with Nepalese browntop showed reduced total organic carbon (24% decline, P<0.09), particulate organic matter (34% decline, P<0.08), mineralizable carbon (a measure of microbially-available carbon; 36% decline, P<0.01), and microbial-biomass carbon (72% decline, P<0.05). The authors suggested that Nepalese browntop may accelerate carbon cycling and deplete carbon levels in southern oak-pine forests . In mixed-hardwood and oak-hickory forests of West Virginia, interior forest plots with Nepalese browntop had significantly lower soil carbon levels than plots without Nepalese browntop (P=0.07) .
Changes in soil chemistry and microfaunal composition associated with soil disturbances tend to favor Nepalese browntop. Across Fairfax County, Virginia, riparian sites in zones changing from rural to urban had increased sediment deposition, increased available soil phosphorus, and decreased soil nitrogen compared to rural riparian zones. In aboveground Nepalese browntop tissues, phosphorus content increased with urbanization, while the nitrogen:phosphorus ratio decreased. The authors suggested that disturbances and changes in soil nutrient levels enhanced the suitability of urbanizing riparian zones as Nepalese browntop habitat . Nonnative earthworms may also favor Nepalese browntop invasion. In sugar maple and oak-hickory forests of New York and Pennsylvania, biomass of nonnative earthworm species was positively associated with Nepalese browntop and 2 other nonnative species, garlic mustard and Japanese barberry. Nonnative earthworm biomass was negatively correlated with leaf litter volume (r= -0.58, P<0.001) . Several studies show that deep litter, which is more typical of early- than late-successional forests, discourages Nepalese browntop establishment [32,120,194] (see Germination and Seedling establishment and plant growth). Nuzzo and others  suggest that nonnative earthworm species, rather than Nepalese browntop, may be driving changes in ecosystem function—such as reduced native plant diversity—in forest communities of the eastern United States, and that nonnative earthworms may facilitate establishment of nonnative plant species.
Nepalese browntop may favor insect guilds that use the ground layer as habitat. In a harvested white oak-yellow-poplar forest in Tennessee, there was significantly greater cover of all insect guilds (herbivores, omnivores, carnivores, and scavengers) on sites with than without Nepalese browntop (P≤0.05), probably because there was 2.5 times more plant cover on sites with Nepalese browntop. Measurements were taken at the end of the growing season (mid-October) .
Diversity and stand structure:
Plant species diversity: Sites with Nepalese browntop tend to have lower native and total plant species diversity than sites without Nepalese browntop [2,3,21,41,68,87,223]. In an oak-yellow-poplar forest in Tennessee, density (r²=0.80, P<0.001) and diversity (r²=0.31, P=0.02) of native woody species was less in Nepalese browntop-infested compared to uninfested sites. The authors suggested that regeneration of woody species in southern forests will likely be reduced with Nepalese browntop invasion . In a bottomland box elder-yellow-poplar-sycamore forest in Indiana, plots tilled and sown with native herbs and Nepalese browntop had significantly different groundlayer species composition than plots tilled and sown with only native herbs. Nepalese browntop plots showed 43% lower groundlayer species richness and 38% lower diversity than plots without Nepalese browntop. There was a strong negative correlation between Nepalese browntop presence and biomass of the sown native herbs (P<0.0001 for all variables) [61,63]. In urban riparian forests of North Carolina, Nepalese browntop presence was negatively correlated with presence of white oak, hickories, flowering dogwood, and mapleleaf viburnum (Viburnum acerifolium) (P<0.05). The authors found that light and high soil nutrient levels were positively associated with cover of nonnative species in general (P<0.05), and they suggested that Nepalese browntop is competitively excluding woody species in urban riparian forests of the eastern United States . In sweetgum-sycamore and loblolly pine-white oak-sweetgum forests of Mississippi, Nepalese browntop presence was significantly associated with low species richness, and Nepalese browntop production was less in species-rich plant communities than in species-poor communities (P<0.001) . In mixed hardwood and oak-hickory forests of West Virginia, interior forest plots with Nepalese browntop had significantly lower herb, liana, and shrub diversity (P=0.03) and tree seedling richness (P=0.02) and diversity (P=0.07) than plots without Nepalese browntop . In surveys within Chesapeake and Ohio Canal National Historic Park, Maryland, plots with Nepalese browntop had greater native species diversity than plots without Nepalese browntop until August, when Nepalese browntop overtopped associated groundlayer species. After that, native species diversity was greater on plots without than with Nepalese browntop [2,3].
Animal species diversity and stand structure: In areas with dense white-tailed deer populations, Nepalese browntop and white-tailed deer interactions may be altering forest structure, with attendant changes to wildlife populations. White-tailed deer avoid grazing Nepalese browntop because it is unpalatable (see Importance to Wildlife and Livestock). Heavy white-tailed deer browsing of palatable woody species can result in dense cover of Nepalese browntop and little woody species regeneration [10,75,221]. Royo and Carson  termed this phenomenon a "recalcitrant understory"; such understories can persist for decades, altering forest structure and successional pathways. Baiser and others  postulated that in eastern deciduous forests, decreases in bird guilds that nest on the ground, the understory, or the midstory may be partially due to decline of under- and midstory woody species that are subject to heavy white-tailed deer browsing and replacement of the woody species by Nepalese browntop. The authors found that from 1980 to 2005, breeding bird guilds using lower forest layers averaged greater population declines than bird species using the canopy for breeding, and the only bird species with increased populations were those nesting in the canopy. This general decline occurred for both resident and neotropical bird species that nest below the canopy. Among these guilds, eastern wood-pewees (midstory nester) and black-billed cuckoos (ground or understory nester) showed greatest declines in abundance .
Interference: Nepalese browntop may negatively impact establishment and growth of native species. For example, in hardwood floodplain forests of north-central Mississippi, Nepalese browntop interfered with growth of native slender woodoats (Chasmanthium laxum), whitegrass, and white oak seedlings. Density of the native species was negatively correlated with that of Nepalese browntop (P≤0.03) . Nepalese browntop may interfere with production of forage species on rangelands .
Nepalese browntop may competitively exclude midstory species from germination and establishment sites. Based on germination and shade manipulation experiments conducted in a loblolly pine-red oak-black oak/flowering dogwood/mayapple (Cornus florida/Podophyllum peltatum) forest in Virginia, Shaw  suggested that Nepalese browntop may interfere with recruitment of midstory species such as eastern redbud (Cercis canadensis) and flowering dogwood (Cornus florida). There were significantly more eastern redbud (Cercis canadensis) germinants on plots without Nepalese browntop than on plots with Nepalese browntop (P<0.001). There were also more flowering dogwood germinants on plots without Nepalese browntop, but on all plots, recruitment of flowering dogwood was too scant for statistical analyses .
Silvicultural implications: Nepalese browntop is identified as a potentially serious competitor on productive timber sites in the Southeast [12,172,184]. It is implicated in reducing growth of timber species and associated species growing under the canopy. Because it is a tall grass that can form thick lawns, it often overtops and excludes native species. On the Hutcheson Memorial Forest, height of Nepalese browntop ranges from 10 to 40 inches (30-100 cm), far taller than most tree seedlings and forest herbs . In red oak-green ash forests of New Jersey, survival of planted red oak and American ash seedlings was less on sites with Nepalese browntop than on sites where Nepalese browntop was removed (P<0.0001), but survival of associated red maple was not significantly affected by Nepalese browntop. Relative growth rates of red oak and American ash were significantly reduced on plots with Nepalese browntop (P<0.0001). Overall herbaceous species richness was less on plots with than on plots without Nepalese browntop (P=0.02). The author speculated that Nepalese browntop interference and white-tailed deer browsing (deer density range: 58-77/km²) have a synergistic, negative effect on oak and ash regeneration in New Jersey forests  (see Animal species diversity for more information). On an oak plantation in southwestern Tennessee, Nepalese browntop presence was negatively correlated (r= -0.82) with growth of northern red oak seedlings. Four silvicultural treatments were tested: clearcut (all stems >6 inches (20 cm) diameter removed); 2-aged selection cut (harvest to retain a stand basal area of 15 to 20 feet²/acre of residual oaks, hickories, and yellow-poplar); high-grade cut (all stems >14 inches (36 cm) DBH removed); and a control no-cut treatment. Mean biomass gain of Nepalese browntop was greatest with a 2-aged selection cut and least with the no-cut control :
|Nepalese browntop productivity (lb/acre) by silvicultural treatment in a Tennessee oak plantation |
In a harvested white oak-yellow-poplar forest in Tennessee, Nepalese browntop mean stem length and number of nodes increased as canopy cover decreased, while soil temperature and moisture increased as Nepalese browntop cover increased. Leaf area of red maple and yellow-poplar was less in plots with than without Nepalese browntop, likely because Nepalese browntop outcompeted the hardwoods for soil moisture. Measurements were made at the end of the growing season (mid-October) .
Other nonnative species: Nepalese browntop may outcompete other nonnative herbs and woody species. Miller and others  compared the relative competitive abilities of Nepalese browntop and garlic mustard in greenhouse and field experiments. In the greenhouse, they found that in both shaded conditions and open sunlight, Nepalese browntop gained more aboveground biomass and had higher rates of photosynthesis than garlic mustard. In the field, Nepalese browntop seedlings also gained more biomass and had higher rates of photosynthesis than garlic mustard; additionally, it suffered less mortality and insect herbivory (P<0.001 for all variables). The authors concluded that in eastern forests, Nepalese browntop has greater potential than garlic mustard for spread on both open and shaded sites .
In a sweetgum plantation in Tennessee, Nepalese browntop outcompeted Japanese honeysuckle for light, gaining more height growth and biomass than and shading out Japanese honeysuckle when the 2 species were grown together. Watering increased Nepalese browntop's interference with Japanese honeysuckle growth. Since Nepalese browntop is an annual, Nepalese browntop's negative effect on Japanese honeysuckle growth may decrease as Japanese honeysuckle matures and gains height .
Allelopathy: In the laboratory, the inhibitory effect of Nepalese browntop extracts on germination of radish (Raphanus sativus) seed was strong enough (β= -0.37) that the authors suspected Nepalese browntop may be allelopathic. They called for field studies testing Nepalese browntop's possible allelopathy .
Control: Control of Nepalese browntop is difficult and requires multiple treatments . In order to locally control this annual, seed-banking grass, repeated annual efforts must be made to prevent flowering and seed set until the seed bank is exhausted . Nepalese browntop resembles native white grass, so proper identification of Nepalese browntop before control measures are undertaken is advised . Shaw  writes that "M. vimineum is proving to be an enigma for scientists because it can grow and succeed in a wide range of habitats. This plasticity makes M. vimineum a difficult weed (in terms of preventing) its invasion and/or (controlling the) spread of existing patches".
Several researchers stress the importance of controlling Nepalese browntop along roadsides and trails in order to prevent its invasion into forest interiors [36,117,131]. Because Nepalese browntop seed production, cover, and rate of spread were significantly greater along roadsides than within oak-hickory and maple-beech-birch forest interiors of West Virginia, Huebner  also recommended making control of Nepalese browntop along roadsides a priority.
In all cases where invasive species are targeted for control, no matter what method is employed, the potential for other invasive species to fill their void must be considered . Control of biotic invasions is most effective when it employs a long-term, ecosystem-wide strategy rather than a tactical approach focused on battling individual invaders .
Prevention: It is commonly argued that the most cost-efficient and effective method of managing invasive species is to prevent their establishment and spread by maintaining "healthy" natural communities [104,183,183] (for example, avoid road building in wildlands ) and monitoring several times each year . Managing to maintain the integrity of the native plant community and mitigate the factors enhancing ecosystem invasibility is likely to be more effective than managing solely to control the invader . Monitoring efforts are best concentrated on the most likely sites of invasion, particularly along potential pathways for Nepalese browntop invasion: waterways, roadsides, and adjacent old fields and woodlands. Periodically surveying to detect new invasions is recommended . The Center for Invasive Plant Management provides an online guide to noxious weed prevention practices.
Weed prevention and control can be incorporated into many types of management plans, including those for logging and site preparation, grazing allotments, recreation management, research projects, road building and maintenance, and fire management . Nord and others  suggested that Nepalese browntop invasion may be prevented if disturbed sites are kept free of Nepalese browntop seed and stolons (by, for example, cleaning logging or other equipment coming into disturbed sites), and that disturbed plant communities are likely to become less vulnerable to Nepalese browntop over time. The rate of Nepalese browntop population expansion decreased with time since disturbance on their Pennsylvanian oak-hickory-eastern white pine forest study sites . See the Guide to noxious weed prevention practices  for specific guidelines in preventing the spread of weed seeds and propagules under different management conditions.
Swearingen  stresses that preventing the introduction of Nepalese browntop into uninfested areas, and early control of small infestations, should be a priority. Removing Nepalese browntop plants late in the growing season, before Nepalese browntop seed set but after seed set of most associated species, is recommended [68,215]. Once established, Nepalese browntop requires major, long-term eradication and restoration efforts. The Nature Conservancy  reports high potential for successful control and management of Nepalese browntop if it is detected and controlled in the early stages of invasion, but they report only moderate potential for Nepalese browntop control and large-scale wildland restoration in areas where Nepalese browntop is already well established. Tu  provides a contact list of managers who have used control measures (successful or not) on Nepalese browntop in Natural Areas.
Fire: For information on the use of prescribed fire to control this species, see Fire Management Considerations.These methods of Nepalese browntop control are discussed below:
Mowing is recommended late in the growing season (August-September), when plants are flowering but before seed set. Because Nepalese browntop is an annual, late-season mowing curtails growth. Early-season mowing does not control Nepalese browntop 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 [44,191,215].
Tilling also reduces Nepalese browntop . Soil must be tilled late in the growing season to avoid establishment of soil-stored seed. Tilling may not be appropriate in Natural Areas and may damage desirable plants.
Flooding for 3 straight months, or intermittent inundation, may kill Nepalese browntop plants. It may not kill soil-stored seed .
Biological control: Nepalese browntop has few natural predators and pathogens in North America . No biological control agents were available for Nepalese browntop control as of 2010 [191,201]. Biological control of invasive species has a long history that indicates many factors must be considered before using biological controls. Refer to these sources: [211,227] and the Weed control methods handbook  for background information and important considerations for developing and implementing biological control programs.
Cultural control: Little information was available on cultural control of Nepalese browntop as of 2010, but one study demonstrates how native-species planting after control treatment helped control Nepalese browntop. In a 3-year study in a native cane (Arundinaria gigantea) wetland in Palo Verde National Park, Costa Rica, Nepalese browntop became dominant on plots where nonnative Chinese privet had been removed and cane was not planted. However, cane became dominant on plots where it was planted after Chinese privet removal, and overall plant species diversity increased compared to plots where Chinese privet was removed but cane was not planted (P≤0.05 for all variables) .
Chemical control: 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 . 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 (for example, ). 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 Nepalese browntop can be controlled with systemic herbicides . Herbicides may be the only practical method to effectively control large infestations. Glyphosate may control Nepalese browntop , but since glyphosate is a nonselective herbicide, care must be taken to avoid drift onto desirable native species. The University of Tennessee reported good control of Nepalese browntop on their Ames Plantation, but they also reported that managing for a desirable plant community after Nepalese browntop was controlled was "difficult". The University found good control of Nepalese browntop with imazameth . Because imazameth is selective for only a few plant species, it killed Nepalese browntop plants without killing associated native herbaceous species. Sethoxydim and fluazifop are grass-specific herbicides reported as giving some control for Nepalese browntop (Tu 2005 personal communication cited in ). See these references for further information on using herbicides to control Nepalese browntop: [56,74,95,121,163,163,201,229].
Integrated management: A combination of complementary control methods may be helpful for rapid and effective control of Nepalese browntop. Integrated management includes not only killing the target plant, but also establishing desirable species and discouraging nonnative, invasive species over the long term. Nepalese browntop control is rarely successful with only one method of control , but a combination of control methods may be effective. Unfortunately, few studies on using integrated management to control Nepalese browntop had been reported as of 2010.
The best way to prevent large Nepalese browntop infestations is to control small patches. Small patches of Nepalese browntop in Great Smoky Mountains National Park have been controlled through a combination of herbicides, mowing, and hand-pulling (Johnson 2001 cited in ). Prescribed fire may be used in combination with other control methods for Nepalese browntop. For example, burning can be used to help reduce litter and standing plant biomass prior to herbicide application for Nepalese browntop control .
Comparisons of different control methods: A comparison of 5 Nepalese browntop control methods in North Carolina suggest hand-pulling or a grass-specific herbicide are good choices for Nepalese browntop control. The control treatments were: 1) season-long hand-pulling, 2) fall mowing, 3) a single application of glyphosate in fall, 4) selective hand-pulling of only Nepalese browntop, or 5) fenoxaprop (a grass herbicide) application once or twice a year as needed. Fall treatments were done before Nepalese browntop was flowering. These treatments were conducted for 3 consecutive years on 2 sites. On the Duke Forest site, Nepalese browntop dominated the ground layer of a loblolly pine plantation and was interfering with growth of loblolly pine regeneration. On the Schenck Memorial Forest site, Nepalese browntop and sweetgum seedlings dominated the ground layer of a white ash-American elm forest. After 3 years, all treatments reduced Nepalese browntop cover and presence in the seed bank compared to control plots. There were no significant differences in Nepalese browntop cover among treatments, but native plant recruitment and species richness were highest with selective hand-pulling of Nepalese browntop or fenoxaprop applications. Because it reduced recruitment of native woody species the most, glyphosate was considered the least effective for restoration purposes [93,96].Some Nepalese browntop control treatments serve overall restoration objectives better than others. On 3 mixed-hardwood forest sites in southern Indiana, hand-pulling Nepalese browntop promoted cover of native grasses better than a postemergent herbicide (fluazifop) the 1st year after treatments, while either hand-pulling or postemergent herbicide best promoted forb cover. However, Nepalese browntop invaded hand-pulled areas the spring after treatment. Both pre- and postemergent herbicide prevented Nepalese browntop reinvasion the spring after treatment, although postemergent herbicide promoted higher overall native plant diversity. Seeding with native species did not increase native plant diversity over that of unseeded plots in posttreatment year 2 (P<0.05 for all variables) [61,62].
|Fire regime information on vegetation communities in which Nepalese browntop may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models , which were developed by local experts using available literature, local data, and/or expert opinion. This table summarizes fire regime characteristics for each plant community listed. The PDF file linked from each plant community name describes the model and synthesizes the knowledge available on vegetation composition, structure, and dynamics in that community. Cells are blank where information is not available in the Rapid Assessment Vegetation Model.|
|Northern Great Plains|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|Northern Plains Grassland|
|Northern tallgrass prairie||Replacement||90%||6.5||1||25|
|Surface or low||2%||303|
|Surface or low||76%||4|
|Northern Plains Woodland|
|Surface or low||98%||7.5|
|Northern Great Plains wooded draws and ravines||Replacement||38%||45||30||100|
|Surface or low||43%||40||10|
|Great Plains floodplain||Replacement||100%||500|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|Great Lakes Grassland|
|Mosaic of bluestem prairie and oak-hickory||Replacement||79%||5||1||8|
|Surface or low||20%||2||33|
|Great Lakes Woodland|
|Great Lakes pine barrens||Replacement||8%||41||10||80|
|Surface or low||83%||4||1||20|
|Jack pine-open lands (frequent fire-return interval)||Replacement||83%||26||10||100|
|Northern oak savanna||Replacement||4%||110||50||500|
|Surface or low||87%||5||1||20|
|Great Lakes Forested|
|Northern hardwood maple-beech-eastern hemlock||Replacement||60%||>1,000|
|Conifer lowland (embedded in fire-prone ecosystem)||Replacement||45%||120||90||220|
|Conifer lowland (embedded in fire-resistant ecosystem)||Replacement||36%||540||220||>1,000|
|Great Lakes floodplain forest||Mixed||7%||833|
|Surface or low||93%||61|
|Surface or low||21%||300|
|Minnesota spruce-fir (adjacent to Lake Superior and Drift and Lake Plain)||Replacement||79%||80|
|Great Lakes pine forest, jack pine||Replacement||67%||50|
|Surface or low||10%||333|
|Surface or low||67%||500|
|Maple-basswood mesic hardwood forest (Great Lakes)||Replacement||100%||>1,000||>1,000||>1,000|
|Surface or low||89%||35|
|Northern hardwood-eastern hemlock forest (Great Lakes)||Replacement||99%||>1,000|
|Surface or low||76%||11||2||25|
|Surface or low||81%||85|
|Red pine-eastern white pine (frequent fire)||Replacement||38%||56|
|Surface or low||26%||84|
|Red pine-eastern white pine (less frequent fire)||Replacement||30%||166|
|Surface or low||23%||220|
|Great Lakes pine forest, eastern white pine-eastern hemlock (frequent fire)||Replacement||52%||260|
|Surface or low||35%||385|
|Eastern white pine-eastern hemlock||Replacement||54%||370|
|Surface or low||34%||588|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|Northern coastal marsh||Replacement||97%||7||2||50|
|Eastern woodland mosaic||Replacement||2%||200||100||300|
|Surface or low||89%||4||1||7|
|Surface or low||65%||12|
|Northern hardwoods (Northeast)||Replacement||39%||>1,000|
|Eastern white pine-northern hardwood||Replacement||72%||475|
|Surface or low||28%||>1,000|
|Northern hardwoods-eastern hemlock||Replacement||50%||>1,000|
|Surface or low||50%||>1,000|
|Appalachian oak forest (dry-mesic)||Replacement||2%||625||500||>1,000|
|Surface or low||92%||15||7||26|
|Northeast spruce-fir forest||Replacement||100%||265||150||300|
|Southeastern red spruce-Fraser fir||Replacement||100%||500||300||>1,000|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|South-central US Grassland|
|Southern tallgrass prairie||Replacement||91%||5|
|Surface or low||93%||3||1||4|
|South-central US Woodland|
|Oak-hickory savanna (East Texas)||Replacement||1%||227|
|Surface or low||99%||3.2|
|Interior Highlands dry oak/bluestem woodland and glade||Replacement||16%||25||10||100|
|Surface or low||80%||5||2||7|
|Oak woodland-shrubland-grassland mosaic||Replacement||11%||50|
|Surface or low||33%||17|
|Interior Highlands oak-hickory-pine||Replacement||3%||150||100||300|
|Surface or low||97%||4||2||10|
|Surface or low||96%||4|
|South-central US Forested|
|Interior Highlands dry-mesic forest and woodland||Replacement||7%||250||50||300|
|Surface or low||75%||22||5||35|
|Gulf Coastal Plain pine flatwoods||Replacement||2%||190|
|Surface or low||95%||5|
|West Gulf Coastal plain pine (uplands and flatwoods)||Replacement||4%||100||50||200|
|Surface or low||93%||4||4||10|
|West Gulf Coastal Plain pine-hardwood woodland or forest upland||Replacement||3%||100||20||200|
|Surface or low||94%||3||3||5|
|Surface or low||58%||100|
|Southern floodplain (rare fire)||Replacement||42%||>1,000|
|Surface or low||58%||714|
|Surface or low||94%||6|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|Southern Appalachians Grassland|
|Surface or low||44%||16|
|Eastern prairie-woodland mosaic||Replacement||50%||10|
|Surface or low||50%||10|
|Southern Appalachians Woodland|
|Appalachian shortleaf pine||Replacement||4%||125|
|Surface or low||92%||6|
|Table Mountain-pitch pine||Replacement||5%||100|
|Surface or low||92%||5|
|Surface or low||49%||55|
|Southern Appalachians Forested|
|Bottomland hardwood forest||Replacement||25%||435||200||>1,000|
|Surface or low||51%||210||50||250|
|Mixed mesophytic hardwood||Replacement||11%||665|
|Surface or low||79%||90|
|Surface or low||89%||6||3||10|
|Eastern hemlock-eastern white pine-hardwood||Replacement||17%||>1,000||500||>1,000|
|Surface or low||83%||210||100||>1,000|
|Red pine-eastern white pine (frequent fire)||Replacement||38%||56|
|Surface or low||26%||84|
|Eastern white pine-northern hardwood||Replacement||72%||475|
|Surface or low||28%||>1,000|
|Oak (eastern dry-xeric)||Replacement||6%||128||50||100|
|Surface or low||78%||10||1||10|
|Appalachian Virginia pine||Replacement||20%||110||25||125|
|Surface or low||64%||35||10||40|
|Appalachian oak forest (dry-mesic)||Replacement||6%||220|
|Surface or low||79%||17|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|Southeast Gulf Coastal Plain Blackland prairie and woodland||Replacement||22%||7|
|Surface or low||9%||20|
|Southern tidal brackish to freshwater marsh||Replacement||100%||5|
|Gulf Coast wet pine savanna||Replacement||2%||165||10||500|
|Surface or low||98%||3||1||10|
|Surface or low||97%||4||1||5|
|Longleaf pine (mesic uplands)||Replacement||3%||110||40||200|
|Surface or low||97%||3||1||5|
|Longleaf pine-Sandhills prairie||Replacement||3%||130||25||500|
|Surface or low||97%||4||1||10|
|Surface or low||10%||43||2||50|
|South Florida slash pine flatwoods||Replacement||6%||50||50||90|
|Surface or low||94%||3||1||6|
|Atlantic wet pine savanna||Replacement||4%||100|
|Surface or low||94%||4|
|Sand pine scrub||Replacement||90%||45||10||100|
|Coastal Plain pine-oak-hickory||Replacement||4%||200|
|Surface or low||89%||8|
|Atlantic white-cedar forest||Replacement||34%||200||25||350|
|Surface or low||59%||115||10||500|
|Surface or low||80%||9||3||50|
|Surface or low||97%||2||1||8|
|Surface or low||93%||63|
Replacement: Any fire that causes greater than 75% top removal of a vegetation-fuel type, resulting in general replacement of existing vegetation; may or may not cause a lethal effect on the plants.
Mixed: Any fire burning more than 5% of an area that does not qualify as a replacement, surface, or low-severity fire; includes mosaic and other fires that are intermediate in effects.
Surface or low: Any fire that causes less than 25% upper layer replacement and/or removal in a vegetation-fuel class but burns 5% or more of the area [76,107].
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