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Euonymus alatus

INTRODUCTORY

AUTHORSHIP AND CITATION:
Fryer, Janet L. 2009. Euonymus alatus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.usda.gov/database/feis/plants/shrub/euoala/all.html [].

FEIS ABBREVIATION:
EUOALA

NRCS PLANT CODE [46]:
EUAL13

COMMON NAMES:
burningbush
winged burning bush
winged spindletree
winged euonymous
winged wahoo


TAXONOMY:
The scientific name of burningbush is Euonymus alatus (Thunb.) Sieb. (Celastraceae) [4,15,16,19,27,42]. Worldwide, 5 poorly differentiated varieties are recognized by various systematists based on leaf color and relative hairiness (review by [4]). These varieties occur in North America [46,51]:

Euonymus alatus (Thunb.) Sieb. var. alatus
Euonymus alatus (Thunb.) Sieb. var. apterus Regel

SYNONYMS:
Euonymus alata (Thub.) Sieb. [34,49]

LIFE FORM:
Shrub

FEDERAL LEGAL STATUS:
None

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

DISTRIBUTION AND OCCURRENCE

SPECIES: Euonymus alatus
GENERAL DISTRIBUTION:
North America: Burningbush is nonnative in North America, where it occurs from Ontario south to Missouri, Kentucky, and South Carolina and east to New Hampshire [27]. Booth and Wright [5] noted its presence in west-central Montana in 1962, but it was not listed in a 1991 flora of the same area [28]. Plants Database provides a distributional map of burningbush and varieties in North America.

Other continents: Burningbush's native distribution extends from central China to northeastern Asia [5,19,33,42,49]. Besides central and eastern China, burningbush occurs in Korea, the Sakhalin islands of eastern Russia, and Japan [15]. It was introduced in Europe and North America as an ornamental [10,15].

Invasion history in the United States: Burningbush was first introduced in the United States in the 1860s [10]. It has escaped cultivation or become invasive mostly in the northeastern United States.

Northeast: Burningbush was well established in some parts of the Northeast by the late 20th century. It had established near Palestine in Wirt County, West Virginia, by 1977 [42]. In a 1985 Michigan flora, burningbush was noted as rarely escaping cultivation but spreading "occasionally" into woodlands, thickets, and uncultivated urban and rural areas [49]. In 1991, Gleason and Cronquist [19] described it as widely cultivated but only "locally escaped from cultivation" in the Northeast. By the turn of the 21st century, burningbush was locally invasive in many northeastern states. A 2002 fact sheet describes burningbush as most invasive in Connecticut, Virginia, Pennsylvania, and Illinois [33]. A 2006 review describes 21 states as "invaded" [9].

Midwest: As of 2009, burningbush was noted as invasive in the Midwest only in Illinois, where it was first reported as "naturalized" in Coles County in 1973 [11]. By 1984, burningbush was reported from 13 Illinois counties [12]. In 1989, burningbush was the dominant woody species invading the Waterworks Ravine Hill Prairie, where woody species were historically rare. Most other invading woody species were native. Mean density (and frequency) of burningbush was 7,809 stem/ha (66%) [2].

HABITAT TYPES AND PLANT COMMUNITIES:
In the Northeast, burningbush is invasive in pastures, glacial drift hill prairies, woodlands, and mature second-growth forests [20,22,33]. It may be common in late-successional oak-hickory (Quercus-Carya spp.), maple-beech (Acer-Fagus spp.), and other mixed-hardwood forests [22,33,51]. It is noted in mixed-oak forests and swamp chestnut oak-swamp white oak-Shumard's oak (Q. michauxii-Q bicolor-Q. shumardii) swamps of Great Falls National Park, Virginia [23] and in oak-hickory-sugar maple (Acer saccharum) forests of Illinois. It also occurs in little bluestem-Indian grass (Schizachyrium scoparium-Sorghastrum nutans) prairie in Illinois [2]. In southern Connecticut, burningbush occurred in a mixed-hardwood forest with high densities of white-tailed deer. Due to browsing, woody understory vegetation was sparse except for sugar maple seedlings, burningbush, and other nonnative shrubs [51]. See Palatability and/or nutritional value for more information on this study.

Distribution of nonnative species is often difficult to accurately describe due to a lack of details survey information, gaps in understanding of nonnative species' ecological characteristics, and because nonnative species may still be expanding their North American range. Therefore, burningbush may occur in plant communities other than those discussed here and listed in the Fire Regime Table.

As of 2009, little English-language literature was available on burningbush's native habitats. In China, burningbush grows in forests, woodlands, and scrublands [15].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Euonymus alatus
Burningbush stem wings.
Photo courtesy of Duke University.		 Split burningbush capsules, showing orange arils within. Photo © Steve Baskauf.

GENERAL BOTANICAL CHARACTERISTICS:
Botanical description: This description provides characteristics that may be relevant to fire ecology and is not meant for identification. Keys for identification are available (for example, [19,34,42,49]).

Burningbush is a rounded shrub that grows to 8.2 feet (2.5 m) tall [5,19]. The branches typically have prominent, corky wings [5,19,34,42,49], although stem wings are reduced to ridges or absent on some plants [4,16,33,38]. Burningbush is deciduous [5,19,49], with opposite leaves [34]. The inflorescence is a long-peduncled cyme with perfect flowers [34]. The fruit is a dehiscent capsule [19] containing up to to 4 seeds [33] that are enclosed in an aril [19] (see above right photo). A fact sheet describes burningbush's root system as deep and fibrous [38].

 

Stand structure: Based on limited studies, burningbush may form shrublands or dense forest understories; it is uncertain whether this pattern is typical on burningbush-invaded sites. Burningbush has formed dense thickets in Pennsylvania [33]. In New York, burningbush was most common in the tall shrub layer of a yellow-poplar-red oak-American beech (Liriodendron tulipifera-Quercus rubra-Fagus grandifolia) forest [22]:

Cover (%) of burningbush in a mixed-hardwood forest in Long Island, New York [22]
Tall shrub layer (2-3.5 m) Medium shrub layer (1-2 m) Herb layer (0-1 m) All layers
1.86 0.60 0.53 0.53

Raunkiaer [37] life form:
Phanerophyte

SEASONAL DEVELOPMENT:
In general, burningbush flowers from May to June in the Northeast [19]. Late April to late June flowering is noted in Pennsylvania [33,38], while burningbush flowers in June and July in Illinois [34]. Fruit capsules ripen in September and October in the Northeast [33]. The capsules dehisce and seeds disperse in September and October in Pennsylvania [38]. Leaves turn a "brilliant" purplish red to scarlet before dropping in autumn [33]. Fall color is most intense in plants growing in sun [38].
Fall color; burningbush seedlings in foreground. Photo © Barry A. Rice, The Nature Conservancy.  

REGENERATION PROCESSES:

Burningbush regenerates from seed and vegetatively [33]. Little is known of burningbush's regeneration requirements in wildland ecosystems. Research is needed on all aspects of burningbush regeneration.

Pollination and breeding system: No information is available on this topic.

Seed production: A fact sheet described burningbush seed production as "prodigious" [33].

Seed dispersal: Seeds are dispersed by frugivorous birds [20,33].

Seed banking: No information is available on this topic.

Germination: Passage through a bird's digestive system may increase germination rates [33], although this has not been confirmed experimentally. Commercial burningbush seeds are stratified to increase germination rates in the nursery and in the field [48].

Seedling establishment and plant growth: As of 2009, no published information was available on requirements for burningbush establishment and growth. The photo on the right illustrates burningbush establishing in oak litter; the photo in Seasonal Development shows burningbush seedlings establishing beneath parent plants. Seedlings are shade tolerant (see Successional Status), but it is uncertain whether growth rates differ between open than with closed canopies. Fact sheets variously describe burningbush as a fast- [38] or slow-growing [13] species.

  
Photo by Leslie J. Mehrhoff, University of Connecticut, Bugwood.org.

Vegetative regeneration: According to expert opinion [33] and fact sheets [13,38], burningbush sprouts from the root crown after top-kill from herbicides, so it is likely that it also sprouts following other top-killing events. Burningbush tolerates severe pruning [13].

SITE CHARACTERISTICS:
Burningbush appears adaptable to a variety of site conditions. As of 2009, few details were available regarding site conditions preferred by this species. Burningbush tolerates a variety of soil textures [33,36] and pH levels [33], and grows in dry to moist soils [10]. A fact sheet described burningbush as drought tolerant but intolerant of wet soils [13]. In oak-hickory-sugar maple forests in Illinois, burningbush dominated forest understories most often within small ravines. It also dominated shady valley floors and shady microsites on north-facing hillslopes [11]. Research is needed on site conditions that may increase vulnerability of native plant communities to burningbush invasion.

In its native China, burningbush occurs in forests, woodlands, and scrublands from sea level to 8,900 feet (2,700 m) elevation [15]. Elevational ranges for burningbush in North America were not available of as 2009.

Burningbush may tolerate elevated levels of soil methane. In New York, it was planted on landfill sites, where methane concentration was 0.9%; and on control sites, where methane was not detected in the soil. Burningbush persisted in landfill soils, although its growth was significantly greater in control soils (P<0.05) [18].

SUCCESSIONAL STATUS:
Burningbush tolerates full sun [10] to nearly full shade [10,11,33] and has invaded closed-canopy, "relatively undisturbed" forest communities in Illinois [12]. Little research had been conducted on successional patterns in plant communities with burningbush as of 2009, and research is needed on burningbush's successional role in eastern mixed-hardwood and other plant communities where it is invasive. In the Northeast, burningbush may reach greatest coverage in late-successional hardwood forests. According to a fact sheet, burningbush has overgrown native shrub species and become dominant in the understories of closed-canopy, mixed-hardwood forests in Pennsylvania [38].

FIRE EFFECTS AND MANAGEMENT

SPECIES: Euonymus alatus

FIRE EFFECTS
Immediate fire effect on plant: There was no information on the direct effects of fire on burningbush as of 2009. According to expert opinion [33] and a fact sheet [38], herbicides may only top-kill burningbush, so it is likely that fire may also only top-kill burningbush.

Postfire regeneration strategy [41]:
Tall shrub, adventitious buds and/or a sprouting root crown
Small shrub, adventitious buds and/or a sprouting root crown
Initial off-site colonizer (off site, initial community)
Secondary colonizer (on- or off-site seed sources)

Fire adaptations and plant response to fire: Documentation of burningbush's adaptations and response to fire were lacking as of 2009. Since burningbush sprouts from the root crown after top-kill from herbicides [33,38], it is likely that it also sprouts from the root crown after top-kill by fire. Because birds disperse burningbush seed [20,33], postfire seedling establishment from off-site seed sources may occur. Research is needed on all aspects of burningbush's fire ecology.

FUELS AND FIRE REGIMES:
Fuels: According to a fact sheet, burningbush can successionally replace native shrubs and form a dense forest understory on some sites in the Northeast [38], so burningbush may increase fuel loads in plant communities it has invaded. However, data are lacking on how burningbush may alter horizontal and/or vertical fuel continuity and fuel loads from historical conditions.

Fire regimes: Burningbush grows in plant communities that historically experienced both long and short fire-return intervals. The northeastern beech-maple communities that burningbush has invaded [33,51] historically experienced stand-replacement fires at long fire-return intervals, with an estimated range of 230 to 4,970 years in New Hampshire [14]. Oak-hickory communities historically had mostly short fire-return intervals. Some burningbush-invaded oak-hickory communities in Pennsylvania historically experienced low-severity surface fires at 4- to 7-year intervals. Short fire-return intervals kept stand structure open, usually maintaining the communities as woodlands [29]. Several authors have noted cooccurrence of sugar maple and burningbush in oak-hickory forests [2,11]. On many sites, presence of sugar maple in oak-hickory communities may indicate that these burningbush-invaded forests now have longer fire-return intervals than were typical in the past ([39], review by Pallardy and others [35]). Research on how burningbush affects fire regimes of plant communities it has invaded was lacking as of 2009. See the Fire Regime Table for further information on fire regimes of vegetation communities in which burningbush 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".

FIRE MANAGEMENT CONSIDERATIONS:
Without information on how burningbush responds to fire or how it may affect fuel loads and fire regimes in plant communities it has invaded, it is impossible to make recommendations for its fire management. Studies on the sprouting ability of burningbush after fire; longevity of soil-stored seed; effects (if any) of fire and passage through animal digestive tracts on seed germination; and possible changes in fuels due to burningbush invasion are critically needed to understand the fire ecology burningbush. Postfire monitoring of burningbush can help managers access whether control measures are needed on burned sites.

MANAGEMENT CONSIDERATIONS

SPECIES: Euonymus alatus
IMPORTANCE TO WILDLIFE AND LIVESTOCK:
There was little information on wildlife or livestock use of burningbush as of 2009. Rabbits browse burningbush [3], and birds eat burningbush arils [20,38].

Palatability and/or nutritional value: A southern Connecticut study suggests burningbush is not palatable to white-tailed deer. Although burningbush was plentiful in the understory of a mixed-hardwood forest, white-tailed deer selected other browse species over burningbush. In a related greenhouse study, seedlings of 17 browse species emerged from white-tailed deer fecal samples collected in the forest. None of the samples contained burningbush seedlings [51].

Cover value: No information is available on this topic.

OTHER USES:
Burningbush is widely cultivated for its brilliant autumn foliage and distinctive branches [49].

IMPACTS AND CONTROL:
Impacts: According to a Pennsylvania fact sheet, burningbush may become invasive when planted near pastures, woodlands, and mature second-growth forests, but does not usually escape urban plantings. It is most invasive in moist forest habitats [38]. Burningbush is often used in highway plantings [10], which may contribute to its seed spread.

Limited evidence suggests that in eastern mixed-hardwood forests, heavy white-tailed deer browsing of native shrubs may favor growth of burningbush and other nonnative shrubs [8] (see Palatability and/or nutritional value).		  
Burningbush infestation.
Photo by Leslie J. Mehrhoff, University of Connecticut, Bugwood.org.

Control: Little published information on controlling burningbush was available as of 2009, and further research is needed on this topic. See The Nature Conservancy's Weed Control Methods Handbook for general information on controlling nonnative invasive species.

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 [7]. 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 [32].

Fire: There was no published information on prescribed fire use for controlling burningbush as of 2009.

Prevention:
General prevention of invasives: Preventing invasive plants from establishing in weed-free burned areas is the most effective and least costly management method. This can be accomplished through early detection and eradication, careful monitoring, and by limiting invasive plant seed dispersal into burned areas by:

For more detailed information on these topics see the following publications: [1,6,21,45].

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 [32,40] (for example, avoid road building in wildlands [44]) and by conducting monitoring several times each year [26]. Managing to maintain the integrity of native plant communities and identifying causal factors enhancing ecosystem invasibility are likely to be more effective than an emphasis on controlling the invader [25].

Weed prevention and control can be incorporated into all types of management plans, including logging and site preparation, management of grazing allotments, recreation management, research projects, road building and maintenance, and fire management [45]. See the "Guide to noxious weed prevention practices" [45] for specific guidelines on preventing the spread of weed seeds and propagules under various management conditions.

Preventing burningbush invasion: Burningbush's escape from cultivation could be slowed or prevented if fertile burningbush plants were no longer commercially available and/or if native species were substituted in landscaping projects. However, frugivorous birds may disperse fertile burningbush seed into uninvaded areas (see Seed dispersal). Studies to develop sterile burningbush cultivars were in progress in 2008 [8,17]. Lists of alternative native species that provide bright red or pruple fall color are available in these sources: [33,38].

Cultural: No information is available on this topic.

Physical and/or mechanical: Mechanical treatments are generally preferred over other methods for controlling small infestations of invasive species [43]. Burningbush seedlings up to 2 feet (0.6 m) tall can be hand-pulled, especially in moist soils. Larger plants require cutting and are likely to sprout unless the stumps are removed or painted with herbicide [33,38].

Biological: As of 2009, biological agents for burningbush control were not available [38]. A review identifies 13 potential biological control agents from China that may help control burningbush in North America [9]. Biological control of invasive species has a long history, and many factors must be considered before using biological controls. Refer to the Weed Control Methods Handbook [43] and these sources: [47,52] for background information and important considerations for developing and implementing biological control programs.

Chemical: Glyphosate may be effective on extensive stands, especially when applied in early summer [33,38].

Integrated management: As of 2009, published studies on using multiple control methods on burningbush were lacking. According to a fact sheet, glyphosate application to cut stumps can help prevent cut stumps from sprouting [38].

APPENDIX: FIRE REGIME TABLE

SPECIES: Euonymus alatus

The following table provides fire regime information that may be relevant to burningbush habitats. 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".

Fire regime information on vegetation communities in which burningbush may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models [30], 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 Great Lakes Northeast
Northern Great Plains
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)		 Minimum interval
(years)		 Maximum interval
(years)
Northern Plains Grassland
Northern tallgrass prairie Replacement 90% 6.5 1 25
Mixed 9% 63    
Surface or low 2% 303    
Oak savanna Replacement 7% 44    
Mixed 17% 18    
Surface or low 76% 4    
Northern Plains Woodland
Oak woodland Replacement 2% 450    
Surface or low 98% 7.5    
Northern Great Plains wooded draws and ravines Replacement 38% 45 30 100
Mixed 18% 94    
Surface or low 43% 40 10  
Great Lakes
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)		 Minimum interval
(years)		 Maximum interval
(years)
Great Lakes Grassland
Mosaic of bluestem prairie and oak-hickory Replacement 79% 5 1 8
Mixed 2% 260    
Surface or low 20% 2   33
Great Lakes Woodland
Northern oak savanna Replacement 4% 110 50 500
Mixed 9% 50 15 150
Surface or low 87% 5 1 20
Great Lakes Forested
Northern hardwood maple-beech-eastern hemlock Replacement 60% >1,000    
Mixed 40% >1,000    
Maple-basswood Replacement 33% >1,000    
Surface or low 67% 500    
Maple-basswood mesic hardwood forest (Great Lakes) Replacement 100% >1,000 >1,000 >1,000
Maple-basswood-oak-aspen Replacement 4% 769    
Mixed 7% 476    
Surface or low 89% 35    
Northern hardwood-eastern hemlock forest (Great Lakes) Replacement 99% >1,000    
Oak-hickory Replacement 13% 66 1  
Mixed 11% 77 5  
Surface or low 76% 11 2 25
Pine-oak Replacement 19% 357    
Surface or low 81% 85    
Northeast
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)		 Minimum interval
(years)		 Maximum interval
(years)
Northeast Woodland
Eastern woodland mosaic Replacement 2% 200 100 300
Mixed 9% 40 20 60
Surface or low 89% 4 1 7
Northeast Forested
Northern hardwoods (Northeast) Replacement 39% >1,000    
Mixed 61% 650    
Eastern white pine-northern hardwoods 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
Mixed 6% 250 200 500
Surface or low 92% 15 7 26
Beech-maple Replacement 100% >1,000    
*Fire Severities—
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 [24,30].

Euonymus alatus: REFERENCES

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