New York State Invasive Species Information
Origin and Introduction:
Lonicera tatarica is native to Central Asia and Southern Russia and is believed to have been introduced into North America for ornamental purposes as early as the 1750s. Lonicera japonica, – a native of China, Japan and Korea – was introduced for horticultural purposes in 1806 on Long Island; it was widely distributed as a garden plant through the early-1900s when it was finally recognized as a weed. Lonicera maackii, also native to China, Japan and Korea, was introduced as seeds to arboreta throughout the U.S. in the late-1800s to determine whether the plant would grow in North America. This species of honeysuckle was utilized as a soil stabilization and wildlife planning until the mid-1980s and is still available for sale on-line. Lonicera morrowii, a native of Japan, was imported to Massachusetts in the 1860s and was later released as an ornamental. All four species have escaped cultivation and are easily spread by birds.
Lonicera morrowii, Lonicera tatarica, and Lonicera maackii, are perennial shrubs; Lonicera japonica is a perennial woody vine (although its leaves can remain green throughout mild winters). The shrub forms range from 6 to 15 feet in height, while vines can reach 30 feet in length. The egg-shaped leaves range from 1 to 3 inches in length and are arranged oppositely along stems. Invasive honeysuckles begin flowering from May to June and bear small (less than 1 inch long), very fragrant tubular flowers ranging from creamy white through various shades of pink to crimson. Lonicera morrowii and Lonicera tatarica produce ¼ inch red berries from mid-summer through early-fall; Lonicera maackii’s dark-red berries don’t ripen until late-fall; Lonicera japonica produces dark-purple or black berries in the fall. Stems of all four are hollow.
All three species can form very dense populations that can outcompete and suppress the growth of native plant species. These dense stands suppress the growth of other native species. Lonicera maackii leafs out very early in spring, giving it a competitive advantage over native plants. Lonicera japonica leaves are semi-evergreen allowing the plant to grow longer into the winter and giving it a competitive advantage over native vegetation. It shades out understory growth preventing the success of native understory plants and tree seedlings. Its vigorous vine growth covers native trees; the weight of the vine growth can bring down weak trees. By decreasing light availability to the understory, these invasive honeysuckles can alter habitats by depleting soil moisture and nutrients. The invasive honeysuckle berries do not contain the amount of fat and nutrients present in native honeysuckle berries; eating large amounts of the less nutritious invasive berries rather than native berries can have negative impacts on migrating.
Prevention and Control:
Because these plants spread rapidly via birds eating seeds, control should be started in late-summer or early-fall before seeds are ready to be dispersed. In early stages of invasion, or in cases where populations are at low levels, hand removal of honeysuckle seedlings or young plants is a viable option when repeated annually. Systemic herbicides can be utilized in cases of heavy infestation. Specific state rules should be followed and the appropriate (low environment impact, legally labeled for control of these plants) herbicides should be used. For invasive honeysuckles growing in open habitats, prescribed burning may be an effective control alternative.
History in the United States
Exotic bush honeysuckles have been introduced for use as ornamentals, for wildlife cover and for soil erosion control.
History in the United States
Range and Habitat in Illinois
Regularity: Regularly occurring
Regularity: Regularly occurring
Global Range: Lonicera maackii is native to northeastern Asia (Luken and Mattimiro, 1991) including China, Manchuria, Korea, and, less commonly, Japan (Batcher and Stiles, 2000).
(In North America, this species has escaped from cultivation in at least 25 states east of the Rocky Mountains (Batcher and Stiles, 2000; Hutchinson et al., 1998; Luken and Thieret, 1996) and is considered a widespread invasive almost everywhere it occurs.)
Winter honeysuckle was introduced to the U.S. from eastern China in 1845 [36,74,191]. It is distributed in the eastern U.S. from New York, west to Ohio and south to Alabama and Georgia. It is not reported in New Jersey and Delaware, but does occur in Louisiana, Texas, and Utah [36,83,133,161,182]. According to Virginia Department of Conservation and Recreation  winter honeysuckle occurs in the Piedmont region of that state.
Amur honeysuckle is native to central and northeastern China, Manchuria, the Amur and Ussuri river valleys, Korea, and isolated parts of Japan [98,106]. It was first introduced into the U.S. in 1897/98 [74,106], and by 1931 was available from at least 8 commercial nurseries . For a thorough review of the historical cultivation and dissemination of Amur honeysuckle, from its apparent origins in China to its cultivation in Russia, Europe, and North America, see Luken and Thieret .
Amur honeysuckle is distributed in the eastern U.S. from Massachusetts west to North Dakota and south to Texas. However, there are no specific reports of occurrence in Minnesota, South Dakota, or Florida [18,36,43,54,59,83,99,100,122,171,178,186,186]. Amur honeysuckle also occurs in Idaho  and southern Ontario . Lorenz and others  indicate Amur honeysuckle is climatically adapted to all but the coldest areas in this range, such as northern Maine, New Hampshire, and Vermont, the Adirondack area of New York, and southwestern portions of Michigan's Upper Peninsula (see Site Characteristics). According to Sharp and Belcher  the Amur honeysuckle cultivar 'Rem-Red' is "climatically adapted" from Massachusetts to South Carolina and west to Missouri. Based on a survey of herbaria in eastern North America, Trisel  described distribution of "naturalized" Amur honeysuckle from "New Hampshire south to Augusta, Georgia, west to Greenville, Mississippi and Tulsa, Oklahoma, north to Ames, Iowa, and Madison, Wisconsin." Rolfsmeier and others  reported Amur honeysuckle growing outside cultivation in Nebraska and Kansas, but indicate it may not be spreading rapidly in this area. According to Virginia Department of Conservation and Recreation , Amur honeysuckle occurs in the Mountain and Piedmont regions of that state.
Morrow's honeysuckle is native to Japan [54,161,186], and occurs throughout the Japanese archipelago (Talewaki 1969 as cited in ). According to Hidayati and others  it was introduced to the U.S. from Japan in 1875. Morrow's honeysuckle is distributed in the eastern U.S. from Maine west to Minnesota and south to Arkansas, Tennessee, and the Carolinas, as well as in Alabama [18,44,68,76,83,108,122,149,171,172,178,186,197]. It also occurs in Colorado and Wyoming [83,135,189,190], and in the Canadian provinces of New Brunswick, Quebec, Ontario, and Saskatchewan . According to Virginia Department of Conservation and Recreation  Morrow's honeysuckle occurs in the Mountain and Piedmont regions of that state.
Most sources indicate Tatarian honeysuckle is native to eastern Europe and adjacent Asia [18,54,59,92,149,157,186,191], although according to Strausbaugh and Core  it was introduced from western Asia. In the U.S., Tatarian honeysuckle is reported from Maine south to Virginia and west to Washington, Oregon and California, but not in Missouri or Nevada [13,18,20,35,38,45,59,68,69,83,83,92,93,108,122,135,149,157,161,178,186,190,191,198]. It is also reported in Georgia , Alaska  and in Canada from Nova Scotia west to Alberta [83,121]. Lorenz and others  indicate Tatarian honeysuckle is climatically adapted throughout the northeastern U.S., from Maine south to Virginia and west to Kentucky, Ohio, and Michigan (see Site Characteristics). According to Virginia Department of Conservation and Recreation , Tatarian honeysuckle occurs in the Mountain and Piedmont regions of that state.
European fly honeysuckle was introduced from Eurasia [18,54,149,186]. European fly honeysuckle is distributed in the eastern U.S. from Maine west to Wisconsin, and south to Virginia, but it is not reported in West Virginia or Kentucky [18,83,122,149,178]. It is also reported in Oregon, and in the Canadian provinces of New Brunswick, Quebec, and Ontario .
Since it appears the native ranges of Morrow's honeysuckle and Tatarian honeysuckle do not overlap, Bell's honeysuckle probably only occurs outside cultivation in North America . According to a review by Barnes , initial reports of Bell's honeysuckle in North America date to around the late 1800s to early 1900s, while its parent species, Tatarian honeysuckle and Morrow's honeysuckle, were introduced sometime in the mid-1700s and mid-1800s, respectively. Determining the distribution of Bell's honeysuckle is especially problematic, since it is a hybrid of 2 nonnative parent species and is often difficult to identify  (see Taxonomy). A distribution map provided by Barnes and Cottam  shows the U.S. distribution of Bell's honeysuckle from Maine west to northeastern Montana, most of the Dakotas, eastern Nebraska and northeastern Kansas, and south to North Carolina, Tennessee, and northern Missouri. There are also records of its occurrence in Colorado east of the continental divide , in South Carolina and Wyoming [83,135], and in Washington . In Canada, Barnes and Cottam  indicate that Bell's honeysuckle occurs in far southern Quebec and adjacent eastern Ontario, and while less common, is also established in southwestern Manitoba, southern Saskatchewan, and southeastern Alberta. There is also a record of its occurrence in New Brunswick . According to Barnes , Bell's honeysuckle is most widespread in New England and around the southern Great Lakes.
USDA Plants database provides state distribution maps of bush honeysuckles, although additional information may have led to more extensive descriptions of distribution in this review.
The following biogeographic classification systems demonstrate where bush honeysuckles could potentially be found based on floras and other literature, herbarium samples, and confirmed observations. Precise distribution information is unavailable. Therefore, these lists are speculative and may be imprecise.
Regional Distribution in the Western United States
This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):
BLM PHYSIOGRAPHIC REGIONS :
1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
5 Columbia Plateau
6 Upper Basin and Range
7 Lower Basin and Range
8 Northern Rocky Mountains
9 Middle Rocky Mountains
10 Wyoming Basin
11 Southern Rocky Mountains
12 Colorado Plateau
13 Rocky Mountain Piedmont
14 Great Plains
15 Black Hills Uplift
16 Upper Missouri Basin and Broken Lands
States or Provinces
European fly honeysuckle:
Distribution in the United States
Amur, Tartarian, Morrow's, and pretty honeysuckle generally range from the central Great Plains to southern New England and south to Tennessee and North Carolina. The remaining species are sporadically distributed.
Distribution and Habitat in the United States
The following descriptions of bush honeysuckles provide characteristics that may be relevant to fire ecology, and are not meant for identification. Keys for identification are available (e.g. [18,36,54,59,133,161,186,189,190,191]).
|Winter honeysuckle is a deciduous or semievergreen shrub, mainly 3.2 to 10 feet (1-3 m), occasionally to 15 feet (4.6 m) tall [36,37,127,133,191]. The crown is an erect but wide-spreading, irregularly rounded, tangled mass of slender recurved branches [37,127,133]. Leaves are 0.6 to 3.5 inches (1.5-9 cm) long and 0.4 to 1.8 inches (1-4.5 cm) wide [36,37,127,133,191]. Flowers are 0.4 to 0.5 inches (1-1.2 cm) long, borne in pairs on short peduncles . Fruit is a berry, 0.25 to 0.4 inches (6.4-10 mm) in diameter [36,37,127,133], with seeds 0.05 to 0.08 inches (1.3-2 mm) long .|
|Winter honeysuckle. ©Univ. Connecticut Plant Database.|
|Amur honeysuckle is an upright, spreading, deciduous shrub, 12 to 20 feet (3.7-6.1 m) tall with hollow branches [36,37,54,131]. Leaves are 1.4 to 3.5 inches (3.5-9 cm) long and 0.5 to 1.5 inches (1.3-3.8 cm) wide [37,54,131]. Fruit is a 0.08 to 0.25 inch (2-6.4 mm) diameter berry [36,37].|
|Amur honeysuckle. © John M. Randall / The Nature Conservancy.|
|Morrow's honeysuckle is a deciduous shrub, 4.9 to 8 feet (1.5-2.4 m) tall and 6 to 10 feet (1.8-3 m) wide [37,152,161,175], "forming a broad, rounded, dense, tangled mound with foliage and branches to the ground" . Leaves are 1 to 2.5 inches (2.5-6.4 cm) long and 0.5 to 1.25 inches (1.3-3.2 cm) wide [37,161]. The fruit is a 0.25 inch (6 mm) diameter berry .|
|Morrow's honeysuckle. © John M. Randall/The Nature Conservancy.|
|Tatarian honeysuckle is an upright deciduous shrub, 3.3 to 12 feet (1-3 m) tall and 10 feet (3 m) wide [18,37,54,59,93,155,157,161,176,191], often dense with fine branches . Dirr  describes the Tatarian honeysuckle crown as "strongly multi-stemmed with the upper branches arching and the overall effect one of a dense, twiggy mass." Twigs are hollow , and 0.03 to 0.04 inches (0.8-1 mm) in diameter . Bark has long, flat, thin scales and not much shredding , although older stems have shredding bark . Leaves are 0.6 to 2.5 inches (1.5-6.4 cm) long and 0.2 to 1.5 inches (0.5-3.8 cm) wide [37,54,59,157,191]. Flowers are pedunculate and borne in sessile pairs in leaf axils [59,157,191]. Fruits are berries, 0.2 to 0.3 inch (4-8 mm) in diameter, borne singly or in pairs with the bases fused, with 3-6 seeds per fruit [37,59,157,191]. Seeds are about 0.1 inch (2.5-3 mm) long and 0.08 to 0.1 inch (2-2.5 mm) wide [59,157].|
|Tatarian honeysuckle. © John M. Randall/The Nature Conservancy.|
|European fly honeysuckle is a rounded deciduous shrub with spreading arching branches, 3.3 to 10 feet (1-3 m) tall, 10 to 12 feet (3-3.7 m) wide, with hollow twigs [18,37,54,177].|
|European fly honeysuckle. ©Univ. Connecticut Plant Database.|
Bell's honeysuckle is an erect shrub, 4 to 10 feet (1.2-3 m) tall and often at least as wide as it is tall [25,37,133]. It has a round growth habit with spreading, somewhat arching branches . Leaves are 1 to 3 inches (2.5-7.6 cm) long . Fruits are 0.25 to nearly 0.5 inch (6.5-13 mm) diameter round berries [25,37], with 2 to 6 seeds per fruit .
Age of Bell's honeysuckle shrubs studied in southern Wisconsin ranged from 12 to 34 years, with a mean of 20.4 years .
By excavating numerous Bell's honeysuckle shrubs in Wisconsin, it was determined that most roots occurred at a depth of 0.98 to 5.9 inches (2.5-15 cm), and in many cases extended well beyond crown width .
Physiology: Barnes and Cottam  found no difference in photosynthetic response of Bell's honeysuckle shrubs originating from wet and dry sites, and subjected to shaded and exposed light conditions. Similarly, they detected no difference in photosynthetic response of shrubs from either site to induced water stress.
Exotic bush honeysuckles are upright, generally deciduous shrubs that range from 6 to 15 feet in height. The 1-2 ½ inch, egg-shaped leaves are opposite along the stem and short-stalked. Older stems are often hollow. Pairs of fragrant, tubular flowers less than an inch long are borne along the stem in the leaf axils. Flower color varies from creamy white to pink or crimson in some varieties of Tartarian honeysuckle. Flowering generally occurs from early to late spring, but varies for each species and cultivar. The fruits are red to orange, many-seeded berries. Native bush honeysuckles may be confused with these exotic species and cultivars, so proper identification is necessary. Unlike the exotics, most of our native bush honeysuckles have solid stems.
Description and Biology
- Plant: upright, deciduous shrub up to 15-20 ft. high; pith of mature stems is hollow and white or tan (in contrast to solid white pith of native shrub honeysuckle species).
- Leaves: opposite, ovate with a tapered tip, lightly pubescent, and up to 3½ in. long.
- Flowers, fruits and seeds: flowers paired, tubular, white to pinkish, fading to yellow, less than 1 in. long, borne from leaf axils, five petals, upper 4 fused; fruits are red to orange-red berries produced in late summer and persist through the winter.
- Spreads: by fruits which are abundant and highly attractive to birds that consume them and defecate the seeds in new locations. Vegetative sprouting aids in the local spread and persistence.
- Look-alikes: other exotic bush honeysuckles including Tartarian (L. tatarica) and Standish’s honeysuckle (L. standishii); native species of Lonicera like American fly honeysuckle (L. canadensis), swamp fly-honeysuckle (L. oblongifolia) and mountain fly honeysuckle (L. villosa); northern bush-honeysuckle (Diervilla lonicera) which has leaves with toothed margins.
Range and Habitat in Illinois
It is likely that sites characteristics where bush honeysuckles occur in North America are generally similar for the 6 species considered in this summary. However, it is difficult to definitively ascertain which species share affinities for which site characteristics, especially when considered across the entire North American range of bush honeysuckle distribution. Information about site characteristics that favor bush honeysuckle establishment, persistence, and spread is often anecdotal. Consequently, the following information describes site characteristics where bush honeysuckles are likely to be found, but should not be considered a comprehensive assessment. More research is needed to determine relationships between various site characteristics and bush honeysuckle invasion.
|In the North Carolina Piedmont and Coastal Plain, and in the South Carolina Piedmont, winter honeysuckle is found in woodlands and "waste places" . In north-central Texas it escapes to "forest margins" .|
|Winter honeysuckle. |
Horticulture Department/Auburn University.
|According to Luken (personal observation cited in ) and Luken and others , in its native range, Amur honeysuckle commonly grows on sites with some type of canopy cover (open forests, flood plain forests, periodically disturbed floodplains, riparian habitats and scrub communities). In North America, it is found in both open and wooded habitats [99,131]. In southern Wisconsin, Cochrane  described Amur honeysuckle occurrence as mostly in partially shaded fencerows, weedy thickets, and brushy groves, and less frequently in woods . In north-central Texas Amur honeysuckle escapes to "forest margins" , in Michigan it is found in "woods (upland and swampy), thickets, banks, fencerows, and often near a landscaped source" , and in southwestern Ohio it is mentioned as occurring in pastures and woodlands . Hutchinson and Vankat  examined Amur honeysuckle distribution in southwestern Ohio along northerly and westerly transects, emanating from a supposed central population source from which invasive populations have subsequently dispersed. Their results suggest Amur honeysuckle population spread is closely linked to forest cover and forest connectivity across the landscape. They propose that large expanses of agricultural land act as a barrier to dispersal, perhaps due to habitat constraints on frugivorous birds that disperse seeds. Medley  found that Amur honeysuckle density was significantly (p = 0.001) correlated with proximity to the edge of a 13 acre (5.2 ha) mature deciduous forest stand in southwestern Ohio. However, Amur honeysuckle stem basal area was also significantly (p<0.05) correlated with proximity to stream channels, with some of the largest individuals located near the center of the stand along streambanks.|
|Amur honeysuckle. © John M. Randall/The Nature Conservancy.|
|Morrow's honeysuckle is found in woodlands of the Blue Ridge mountains in Tennessee and Virginia , in thickets, fields, and hedgerows in New England , and along streambanks in Wyoming . In lower Michigan, it escapes to roadsides, railroads, thickets, lakeshores, riverbanks, and woods .|
|Morrow's honeysuckle. © John M. Randall/The Nature Conservancy.|
|In Michigan, Tatarian honeysuckle escapes to roadsides, railroads, thickets, lakeshores, riverbanks, woods, fields, "waste places" and swamps , and in the northern Great Plains it escapes to open woods, stream banks, or brushy pastures [59,157]. It is found in riverbank thickets, along roadsides, and in "waste land" in New England , along fence rows and stream banks in Montana and Wyoming , and in riparian areas along the Big Sioux River in eastern South Dakota . According to White , Tatarian honeysuckle is present, though not common, on poorly-drained shrub-dominated sites in southeastern Wisconsin . Moffatt and McLachlan  included Tatarian honeysuckle among indicator species found in disturbed riparian forest in southern Manitoba. It was 1 of 2 nonnative plant species that was significantly (p<0.05) more likely to occur within "urban" or "suburban" sites compared with "rural" (agricultural) or "reference" (undisturbed) sites.|
|Tatarian honeysuckle. Gary Fewless/UW-Green Bay Herbarium.|
|European fly honeysuckle is found in thickets and woods in New England . It is apparently tolerant of "difficult" growing sites .|
|European fly honeysuckle. © Henriette Kress.|
In Michigan, Bell's honeysuckle is found in habitats similar to those of Tatarian honeysuckle and Morrow's honeysuckle . It is found along roadsides and "scrub areas" bordering human habitation in the Adirondacks , and in thickets and "waste places" in New England and the Piedmont of North Carolina [133,149]. A review by Barnes  indicates a wide range of sites may support Bell's honeysuckle populations in Wisconsin, including roadsides, fencerows, pastures or fields, railroad rights-of-way, lake, river, or stream banks, and wooded areas, particularly within openings or edges of woods. Barnes  showed that Bell's honeysuckle distribution within the University of Wisconsin Madison Arboretum was aggregated. He reasoned that the observed patterns of distribution were strongly influenced by site characteristics favorable to seedling establishment (see above).
Soils and topography: Soil requirements and tolerances vary among bush honeysuckles, but most taxa seem to grow best on well-drained sites.
Winter honeysuckle prefers moist but well-drained, loamy soil .
Amur honeysuckle performs best on moist, well-drained sites, but is adaptable to "poor" soils, compacted soils, various soil pHs, restricted root zones, drought and salt spray . According to Vogel  the lower pH limit for Amur honeysuckle is 5.0. It escapes to calcareous slopes in north-central Texas , and grows in thin prairie soils over dolomite in southern Wisconsin . Amur honeysuckle generally occurs in mesic habitats in Virginia . According to Sharp and Belcher  the Amur honeysuckle cultivar 'Rem-Red' is "adapted" to deep, well-drained, fertile, sandy loam to clay loam soils, and is not "adapted" to droughty or wet soils. Lorenz and others  indicate that 'Rem-Red' "grows in medium-fertility, acid, clayey, loamy, and sandy soils, and tolerates somewhat poorly drained soil."
Morrow's honeysuckle prefers loamy, well-drained, moist soil . In Virginia, it generally occurs in mesic habitats . Vogel  reports that the lower pH limit for Morrow's honeysuckle is 5.0.
Barnes  indicates Tatarian honeysuckle occurs on a wide variety of soil types in central Asia. According to Lorenz and others , it "grows in medium-fertility, acid, clayey, loamy, and sandy soils, and tolerates moderately well-drained soil." Tatarian honeysuckle grows on peat and muck soils . White  reports that it is present, though not common, on poorly-drained shrub-dominated sites in southeastern Wisconsin . Tatarian honeysuckle generally occurs in mesic habitats in Virginia . According to Vogel  the lower pH limit for Tatarian honeysuckle is 5.0. Tatarian honeysuckle is considered salt sensitive .
European fly honeysuckle performs best on moist, well-drained sites, but is adaptable to "poor" soils, various soil pHs, restricted root zones, drought, and salt spray, but is not tolerant of wet sites or poorly drained sites .
Mature Bell's honeysuckle shrubs were found growing over a variety of soils at the University of Wisconsin Madison Arboretum including: a) a droughty, infertile, loamy sand, b) a well- to moderately well drained, moderately fertile, silt loam, c) an imperfectly- to poorly drained silt loam, and d) a muck soil where the water table was at or near the surface in spring . In a reciprocal transplant common garden experiment in southern Wisconsin, Barnes and Cottam  successfully transplanted Bell's honeysuckle shrubs at 2 sites with quite different soils. The wet site had muck soils with water-retaining capacity of 260% dry weight and organic matter content 64%, while the dry site was a loamy sand soil with water-retaining capacity of 50% dry weight and 2% organic matter. Survival of transplants, 50% at the dry site and 68% at the wet site, was not significantly different. Transplant origin (wet site or dry site) also did not affect performance at either common garden, indicating no evidence of ecotypic differentiation between populations based on soil type. Bell's honeysuckle generally occurs in mesic habitats in Virginia .
Amur honeysuckle distribution appears to be limited by drought and cold. Trisel  observed that during summer droughts in southwestern Ohio Amur honeysuckle leaves can become severely wilted, while native trees remain unaffected. However, based on unpublished data, it was further indicated that affected shrubs can fully recover from wilting with no apparent damage or mortality following rehydration. Trisel  also hypothesized that susceptibility to drought may result from its shallow root system, and that drought intolerance may be more pronounced in unshaded areas. Lorenz and others  characterize the Amur honeysuckle cultivar 'Rem-Red' as having "fair drought tolerance."
According to Trisel , Amur honeysuckle range expansion to the west and north may be limited by moisture and winter temperature. Several sources indicate that it is adapted to USDA zones 2-8 [17,37], or 3-8 [115,174]. Analysis of herbaria records by Trisel  indicates Amur honeysuckle "escapes" become limited in USDA zones 4-5 (winter low temperatures of -20.2 to -31 ◦F (-29 to -35 ◦C)). Lorenz and others  provide a map of plant hardiness zones of the northeastern U.S., based on average annual minimum temperature, and refer to these as "areas of climatic adaptation" where various plant species are recommended for planting for "conservation" purposes. They indicate that while some species "may grow in zones other than the ones indicated, maximum conservation effectiveness occurs within these zones." Based on this information, Amur honeysuckle is climatically adapted throughout the northeastern U.S. where average annual minimum temperature ranges from 20 to -30 ◦F (-6.7 to -34 ◦C).
Several sources indicate that Tatarian honeysuckle is adapted to USDA zones 3-8 [117,128,176], or 2-8 . Lorenz and others  provide a map of plant hardiness zones of the northeastern U.S., based on average annual minimum temperature, and refer to these as "areas of climatic adaptation" where various plant species are recommended for planting for "conservation" purposes. They indicate that while some species "may grow in zones other than the ones indicated, maximum conservation effectiveness occurs within these zones." Based on this information, Tatarian honeysuckle is climatically adapted throughout the northeastern U.S. where average annual minimum temperature ranges from 20 to -40 ◦F (-6.7 to -40 ◦C). Tatarian honeysuckle is apparently winter hardy in valleys of western Montana . Lorenz and others  characterize Tatarian honeysuckle as having "fair drought tolerance."
Bell's honeysuckle is adapted to USDA zones 4-7 . Average annual precipitation throughout the North American range of Bell's honeysuckle is between 9.8 and 59 inches (250-1500 mm), and average frost-free period is 80 to 240 days .
Key Plant Community Associations
Precise information about habitat types and plant communities where bush honeysuckles may
occur is sparse. Information presented below is mostly gleaned from site descriptions where
research involving bush honeysuckles took place. It is not likely to be a representative
sample, much less an exhaustive review, of bush honeysuckle-invaded habitat types and
plant communities in North America. A review by Nyboer  indicates that, collectively,
bush honeysuckles can colonize a wide array of native habitats within their North American
range, including wetland, prairie, and forest communities. It is further asserted that,
although individual bush honeysuckle species may differ in their tolerances to various site characteristics, most natural communities within this range are susceptible to invasion by 1 or more
Luken , Luken and Goessling , and Luken and Mattimiro  studied Amur honeysuckle
populations in northern Kentucky growing along roadsides (open-grown) and within forest stands
variously dominated by black locust (Robinia pseudoacacia), American elm (Ulmus americana),
slippery elm (U. rubra), sugar maple (Acer saccharum), hackberry
(Celtis occidentalis), and white ash (Fraxinus americana). Gould and Gorchov
, Miller and Gorchov , and Swanson and Vankat  described Amur honeysuckle as the
dominant shrub species growing in the understory of 3 southwestern Ohio hardwood forests.
Important overstory species in these forests were a) shagbark hickory (Carya ovata),
shellbark hickory (C. laciniosa), elms (Ulmus spp.), and northern red oak
(Quercus rubra), b) northern red oak, elms, sugar maple, and American beech
(Fagus grandifolia), and c) oaks (Quercus spp.), sugar maple, and hickories
(Carya spp.). Amur honeysuckle was also mentioned as occurring in another southwestern
Ohio forest dominated by sugar maple along with subdominants American beech, black cherry
(Prunus serotina), bitternut hickory (C. cordiformis), yellow-poplar
(Liriodendron tulipifera), and ash (Fraxinus spp.) 
Tatarian honeysuckle is found in mesic sugar maple- and red maple (Acer saccharinum)-dominated
forests in Vermont and Massachusetts .
Bell's honeysuckle is found in habitats similar to those where Tatarian honeysuckle and
Morrow's honeysuckle occur in Michigan . It occurs in mesic sugar maple- and red
maple-dominated forests in Vermont and Massachusetts . Bell's honeysuckle
and common buckthorn (Rhamnus cathartica) were the dominant shrub species in an oak (Q.
Ã palaeolithicola) -dominated forest in southern Wisconsin . Bell's honeysuckle is
present, but not common, in southeastern Wisconsin shrub-carr communities, which
are wet-ground plant communities dominated by tall shrubs other than alder (Alnus
spp.), with an understory intermediate between meadow and lowland forest .
Barnes and Cottam  noted Bell's honeysuckle in all of the 30 terrestrial plant
communities located within the University of Wisconsin Madison Arboretum.
Habitat: Rangeland Cover Types
This species is known to occur in association with the following Rangeland Cover Types (as classified by the Society for Range Management, SRM):
More info for the terms: cover, forb, fresh, mesic, shrub
SRM (RANGELAND) COVER TYPES :
101 Bluebunch wheatgrass
102 Idaho fescue
103 Green fescue
104 Antelope bitterbrush-bluebunch wheatgrass
105 Antelope bitterbrush-Idaho fescue
106 Bluegrass scabland
107 Western juniper/big sagebrush/bluebunch wheatgrass
109 Ponderosa pine shrubland
110 Ponderosa pine-grassland
201 Blue oak woodland
202 Coast live oak woodland
203 Riparian woodland
204 North coastal shrub
205 Coastal sage shrub
206 Chamise chaparral
207 Scrub oak mixed chaparral
208 Ceanothus mixed chaparral
209 Montane shrubland
211 Creosote bush scrub
213 Alpine grassland
214 Coastal prairie
215 Valley grassland
301 Bluebunch wheatgrass-blue grama
302 Bluebunch wheatgrass-Sandberg bluegrass
303 Bluebunch wheatgrass-western wheatgrass
304 Idaho fescue-bluebunch wheatgrass
305 Idaho fescue-Richardson needlegrass
306 Idaho fescue-slender wheatgrass
307 Idaho fescue-threadleaf sedge
308 Idaho fescue-tufted hairgrass
309 Idaho fescue-western wheatgrass
310 Needle-and-thread-blue grama
311 Rough fescue-bluebunch wheatgrass
312 Rough fescue-Idaho fescue
313 Tufted hairgrass-sedge
314 Big sagebrush-bluebunch wheatgrass
315 Big sagebrush-Idaho fescue
316 Big sagebrush-rough fescue
317 Bitterbrush-bluebunch wheatgrass
318 Bitterbrush-Idaho fescue
319 Bitterbrush-rough fescue
320 Black sagebrush-bluebunch wheatgrass
321 Black sagebrush-Idaho fescue
322 Curlleaf mountain-mahogany-bluebunch wheatgrass
323 Shrubby cinquefoil-rough fescue
324 Threetip sagebrush-Idaho fescue
401 Basin big sagebrush
402 Mountain big sagebrush
403 Wyoming big sagebrush
404 Threetip sagebrush
405 Black sagebrush
406 Low sagebrush
407 Stiff sagebrush
408 Other sagebrush types
409 Tall forb
411 Aspen woodland
412 Juniper-pinyon woodland
413 Gambel oak
414 Salt desert shrub
415 Curlleaf mountain-mahogany
416 True mountain-mahogany
417 Littleleaf mountain-mahogany
418 Bigtooth maple
601 Bluestem prairie
602 Bluestem-prairie sandreed
603 Prairie sandreed-needlegrass
604 Bluestem-grama prairie
605 Sandsage prairie
611 Blue grama-buffalo grass
613 Fescue grassland
614 Crested wheatgrass
704 Blue grama-western wheatgrass
710 Bluestem prairie
711 Bluestem-sacahuista prairie
715 Grama-buffalo grass
717 Little bluestem-Indiangrass-Texas wintergrass
722 Sand sagebrush-mixed prairie
727 Mesquite-buffalo grass
730 Sand shinnery oak
731 Cross timbers-Oklahoma
732 Cross timbers-Texas (little bluestem-post oak)
735 Sideoats grama-sumac-juniper
802 Missouri prairie
803 Missouri glades
804 Tall fescue
806 Gulf Coast salt marsh
807 Gulf Coast fresh marsh
903 Beach wildrye-mixed forb
904 Black spruce-lichen
905 Bluejoint reedgrass
906 Broadleaf forest
909 Freshwater marsh
912 Low scrub shrub birch-ericaceous
913 Low scrub swamp
914 Mesic sedge-grass-herb meadow tundra
915 Mixed herb-herbaceous
916 Sedge-shrub tundra
917 Tall shrub swamp
920 White spruce-paper birch
Habitat: Cover Types
This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):
More info for the terms: cover, swamp
SAF COVER TYPES :
1 Jack pine
5 Balsam fir
12 Black spruce
13 Black spruce-tamarack
14 Northern pin oak
15 Red pine
17 Pin cherry
18 Paper birch
19 Gray birch-red maple
20 White pine-northern red oak-red maple
21 Eastern white pine
22 White pine-hemlock
23 Eastern hemlock
24 Hemlock-yellow birch
25 Sugar maple-beech-yellow birch
26 Sugar maple-basswood
27 Sugar maple
28 Black cherry-maple
30 Red spruce-yellow birch
31 Red spruce-sugar maple-beech
32 Red spruce
33 Red spruce-balsam fir
34 Red spruce-Fraser fir
35 Paper birch-red spruce-balsam fir
37 Northern white-cedar
39 Black ash-American elm-red maple
40 Post oak-blackjack oak
42 Bur oak
43 Bear oak
44 Chestnut oak
45 Pitch pine
46 Eastern redcedar
50 Black locust
51 White pine-chestnut oak
52 White oak-black oak-northern red oak
53 White oak
55 Northern red oak
58 Yellow-poplar-eastern hemlock
59 Yellow-poplar-white oak-northern red oak
60 Beech-sugar maple
61 River birch-sycamore
62 Silver maple-American elm
65 Pin oak-sweetgum
66 Ashe juniper-redberry (Pinchot) juniper
67 Mohrs (shin) oak
69 Sand pine
70 Longleaf pine
71 Longleaf pine-scrub oak
72 Southern scrub oak
73 Southern redcedar
74 Cabbage palmetto
75 Shortleaf pine
76 Shortleaf pine-oak
78 Virginia pine-oak
79 Virginia pine
80 Loblolly pine-shortleaf pine
81 Loblolly pine
82 Loblolly pine-hardwood
83 Longleaf pine-slash pine
84 Slash pine
85 Slash pine-hardwood
88 Willow oak-water oak-diamondleaf (laurel) oak
89 Live oak
91 Swamp chestnut oak-cherrybark oak
92 Sweetgum-willow oak
93 Sugarberry-American elm-green ash
94 Sycamore-sweetgum-American elm
95 Black willow
96 Overcup oak-water hickory
97 Atlantic white-cedar
98 Pond pine
103 Water tupelo-swamp tupelo
104 Sweetbay-swamp tupelo-redbay
107 White spruce
108 Red maple
110 Black oak
201 White spruce
202 White spruce-paper birch
203 Balsam poplar
204 Black spruce
206 Engelmann spruce-subalpine fir
207 Red fir
210 Interior Douglas-fir
211 White fir
212 Western larch
213 Grand fir
215 Western white pine
216 Blue spruce
218 Lodgepole pine
219 Limber pine
220 Rocky Mountain juniper
221 Red alder
222 Black cottonwood-willow
223 Sitka spruce
224 Western hemlock
225 Western hemlock-Sitka spruce
226 Coastal true fir-hemlock
227 Western redcedar-western hemlock
228 Western redcedar
229 Pacific Douglas-fir
230 Douglas-fir-western hemlock
233 Oregon white oak
234 Douglas-fir-tanoak-Pacific madrone
236 Bur oak
237 Interior ponderosa pine
238 Western juniper
243 Sierra Nevada mixed conifer
244 Pacific ponderosa pine-Douglas-fir
245 Pacific ponderosa pine
246 California black oak
247 Jeffrey pine
248 Knobcone pine
249 Canyon live oak
250 Blue oak-foothills pine
251 White spruce-aspen
252 Paper birch
253 Black spruce-white spruce
254 Black spruce-paper birch
255 California coast live oak
Habitat: Plant Associations
This species is known to occur in association with the following plant community types (as classified by Küchler 1964):
More info for the terms: bog, shrub
KUCHLER  PLANT ASSOCIATIONS:
K001 Spruce-cedar-hemlock forest
K002 Cedar-hemlock-Douglas-fir forest
K003 Silver fir-Douglas-fir forest
K004 Fir-hemlock forest
K005 Mixed conifer forest
K006 Redwood forest
K007 Red fir forest
K008 Lodgepole pine-subalpine forest
K009 Pine-cypress forest
K010 Ponderosa shrub forest
K011 Western ponderosa forest
K012 Douglas-fir forest
K013 Cedar-hemlock-pine forest
K014 Grand fir-Douglas-fir forest
K015 Western spruce-fir forest
K016 Eastern ponderosa forest
K017 Black Hills pine forest
K018 Pine-Douglas-fir forest
K019 Arizona pine forest
K020 Spruce-fir-Douglas-fir forest
K021 Southwestern spruce-fir forest
K022 Great Basin pine forest
K023 Juniper-pinyon woodland
K024 Juniper steppe woodland
K025 Alder-ash forest
K026 Oregon oakwoods
K027 Mesquite bosques
K028 Mosaic of K002 and K026
K029 California mixed evergreen forest
K030 California oakwoods
K034 Montane chaparral
K035 Coastal sagebrush
K036 Mosaic of K030 and K035
K037 Mountain-mahogany-oak scrub
K038 Great Basin sagebrush
K041 Creosote bush
K048 California steppe
K053 Grama-galleta steppe
K055 Sagebrush steppe
K056 Wheatgrass-needlegrass shrubsteppe
K057 Galleta-threeawn shrubsteppe
K063 Foothills prairie
K065 Grama-buffalo grass
K068 Wheatgrass-grama-buffalo grass
K069 Bluestem-grama prairie
K070 Sandsage-bluestem prairie
K073 Northern cordgrass prairie
K074 Bluestem prairie
K075 Nebraska Sandhills prairie
K076 Blackland prairie
K077 Bluestem-sacahuista prairie
K078 Southern cordgrass prairie
K081 Oak savanna
K082 Mosaic of K074 and K100
K083 Cedar glades
K084 Cross Timbers
K085 Mesquite-buffalo grass
K086 Juniper-oak savanna
K089 Black Belt
K090 Live oak-sea oats
K093 Great Lakes spruce-fir forest
K094 Conifer bog
K095 Great Lakes pine forest
K096 Northeastern spruce-fir forest
K097 Southeastern spruce-fir forest
K098 Northern floodplain forest
K099 Maple-basswood forest
K100 Oak-hickory forest
K101 Elm-ash forest
K102 Beech-maple forest
K103 Mixed mesophytic forest
K104 Appalachian oak forest
K106 Northern hardwoods
K107 Northern hardwoods-fir forest
K108 Northern hardwoods-spruce forest
K109 Transition between K104 and K106
K110 Northeastern oak-pine forest
K112 Southern mixed forest
K113 Southern floodplain forest
This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):
FRES10 White-red-jack pine
FRES12 Longleaf-slash pine
FRES13 Loblolly-shortleaf pine
FRES21 Ponderosa pine
FRES22 Western white pine
FRES24 Hemlock-Sitka spruce
FRES26 Lodgepole pine
FRES28 Western hardwoods
FRES30 Desert shrub
FRES32 Texas savanna
FRES33 Southwestern shrubsteppe
FRES34 Chaparral-mountain shrub
FRES36 Mountain grasslands
FRES37 Mountain meadows
FRES38 Plains grasslands
FRES41 Wet grasslands
FRES42 Annual grasslands
Habitat in the United States
Exotic bush honeysuckles are relatively shade-intolerant and most often occur in forest edge, abandoned field, pasture, roadsides and other open, upland habitats. Woodlands, especially those that have been grazed or otherwise disturbed, may also be invaded by exotic bush honeysuckles. Morrow's honeysuckle and pretty honeysuckle have the greatest habitat breadth and are capable of invading bogs, fens, lakeshores, sandplains and other uncommon habitat types.
Broad-scale Impacts of Plant Response to Fire
Kline and McClintock  observed Bell's honeysuckle sprouting from the base following
prescribed fire, but "resprouts were not very vigorous". Barnes  observed
fire effects on 2 populations of Bell's honeysuckle at the University of Wisconsin Madison
Arboretum. All 12 shrubs observed at 1 site resprouted from "the rootstock".
At the other site, a late-summer surface fire burned through a Bell's honeysuckle thicket,
leaving scorched bark on basal stems and killing but not consuming leaves. All plants subsequently
sampled had survived and had produced "basal sprouts" by early the following summer.
The Research Paper by Bowles and others 2007 provides information on postfire responses of several plant species, including Amur honeysuckle, that was not available when this review was originally written.
Plant Response to Fire
Although information about asexual regeneration is relatively sparse, it is apparent that sprouting is a common response to mechanical stem damage in bush honeysuckles (see physical/mechanical control). Similarly, bush honeysuckles can also produce sprouts in response to damage from fire [7,75,87,102,126,152].
Bush honeysuckles may establish from bird-dispersed seed after fire. Since snags, surviving trees, or tall shrubs are often present in postfire environments where bush honeysuckles are likely to found (see habitat types and plant communities) and provide perches for frugivorous birds, bush honeysuckle postfire seedling establishment may occur in this environment.
Broad-scale Impacts of Fire
Barnes  observed the effects of fire on 2 populations of Bell's honeysuckle at the University of Wisconsin Madison Arboretum. At 1 site, all leaves and buds on 9 of 12 shrubs were "apparently" killed by a fire in early May. Some "dormant buds" did survive on 3 individuals and were actively growing by late May, although location of these buds was not discussed. No further information was provided on fire effects or burn characteristics. At another site, a late-summer surface fire burned through a Bell's honeysuckle thicket, leaving scorched bark on basal stems and killing but not consuming leaves. Basal and aerial sprouts began to appear within 3 weeks postfire, and some plants produced new leaves that summer. Although it appears that some plants were top-killed, all plants survived. Only 2 of 30 sampled plants produced fruit 1 year postfire.
Kline and McClintock  conducted 2 consecutive annual mid-April prescribed burns in an oak (Q.Ã palaeolithicola) -dominated forest in southern Wisconsin, where Bell's honeysuckle was common in the shrub layer. "Most of the individuals resprouted from the base, but the resprouts were not very vigorous. Some completely dead honeysuckles were observed" in the 1st postfire year.
Immediate Effect of Fire
Fire may top-kill bush honeysuckle plants, and is likely to kill seedlings and unhealthy plants [75,126,152]. However, perennating tissues on roots and root crowns are often protected from fire damage by soil. By excavating numerous Bell's honeysuckle shrubs in Wisconsin, it was determined that most roots occurred at a depth of 0.98 to 5.9 inches (2.5-15 cm) and in many cases extended well beyond crown width .
POSTFIRE REGENERATION STRATEGY :
Tall shrub, adventitious bud/root crown
Small shrub, adventitious bud/root crown
Geophyte, growing points deep in soil
Ground residual colonizer (on-site, initial community)
Crown residual colonizer (on-site, initial community)
Initial off-site colonizer (off-site, initial community)
Secondary colonizer (on-site or off-site seed sources)
Information about the fire ecology of bush honeysuckles is lacking.
Fire adaptations: Although no information could be found regarding the evolutionary relationship between fire and bush honeysuckles in their native ranges, it appears that bush honeysuckles are adapted to survive fire by shielding perennating buds below the soil surface on roots and/or the root crown. Postfire sprouting has been documented [7,75,87,102,126,152] (see Fire Effects), although it is unclear if all bush honeysuckle taxa discussed possess similar abilities.
FIRE REGIMES: Information about bush honeysuckles and FIRE REGIMES is lacking. Research is needed that examines the interactions of fire and bush honeysuckles, effects these interactions may have on native communities and ecosystems, and effects on their respective FIRE REGIMES. For example, bush honeysuckles are present in oak-dominated communities in the eastern U.S. [57,87,119,163]. Historically, fire has been an important ecological influence in oak forests, woodlands, and savannas . Understanding the response of bush honeysuckles (and other nonnative species) to periodic fire could be critical for management and restoration efforts in these and other areas.
The following table lists fire return intervals for communities or ecosystems throughout North America where bush honeysuckles may occur. This list is presented as a guideline to illustrate historic FIRE REGIMES and is not to be interpreted as a strict description of FIRE REGIMES for bush honeysuckles. For further information on FIRE REGIMES in these communities or ecosystems see the corresponding FEIS summary for the dominant taxa listed below.
|Community or Ecosystem||Dominant Species||Fire Return Interval Range (years)|
|silver fir-Douglas-fir||Abies amabilis-Pseudotsuga menziesii var. menziesii||> 200|
|grand fir||Abies grandis||35-200 |
|silver maple-American elm||Acer saccharinum-Ulmus americana||< 35 to 200|
|sugar maple||Acer saccharum||> 1,000|
|sugar maple-basswood||Acer saccharum-Tilia americana||> 1,000 |
|California chaparral||Adenostoma and/or Arctostaphylos spp.||129]|
|bluestem prairie||Andropogon gerardii var. gerardii-Schizachyrium scoparium||90,129]|
|Nebraska sandhills prairie||Andropogon gerardii var. paucipilus-Schizachyrium scoparium||< 10|
|bluestem-Sacahuista prairie||Andropogon littoralis-Spartina spartinae||129]|
|silver sagebrush steppe||Artemisia cana||5-45 [73,132,199]|
|sagebrush steppe||Artemisia tridentata/Pseudoroegneria spicata||20-70 |
|basin big sagebrush||Artemisia tridentata var. tridentata||12-43 |
|mountain big sagebrush||Artemisia tridentata var. vaseyana||15-40 [3,22,120]|
|Wyoming big sagebrush||Artemisia tridentata var. wyomingensis||10-70 (40**) [181,201]|
|coastal sagebrush||Artemisia californica||< 35 to < 100|
|saltbush-greasewood||Atriplex confertifolia-Sarcobatus vermiculatus||129]|
|plains grasslands||Bouteloua spp.||129,199]|
|blue grama-needle-and-thread grass-western wheatgrass||Bouteloua gracilis-Hesperostipa comata-Pascopyrum smithii||129,138,199]|
|blue grama-buffalo grass||Bouteloua gracilis-Buchloe dactyloides||129,199]|
|California montane chaparral||Ceanothus and/or Arctostaphylos spp.||50-100 |
|sugarberry-America elm-green ash||Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica||187]|
|curlleaf mountain-mahogany*||Cercocarpus ledifolius||13-1,000 [5,143]|
|mountain-mahogany-Gambel oak scrub||Cercocarpus ledifolius-Quercus gambelii||129]|
|Atlantic white-cedar||Chamaecyparis thyoides||35 to > 200 |
|blackbrush||Coleogyne ramosissima||< 35 to < 100|
|northern cordgrass prairie||Distichlis spicata-Spartina spp.||1-3 |
|beech-sugar maple||Fagus spp.-Acer saccharum||> 1,000 |
|California steppe||Festuca-Danthonia spp.||129,160]|
|black ash||Fraxinus nigra||187]|
|Ashe juniper||Juniperus ashei||< 35|
|western juniper||Juniperus occidentalis||20-70|
|Rocky Mountain juniper||Juniperus scopulorum||129]|
|cedar glades||Juniperus virginiana||3-22 [63,129]|
|tamarack||Larix laricina||35-200 |
|western larch||Larix occidentalis||25-350 [2,10,31]|
|wheatgrass plains grasslands||Pascopyrum smithii||129,132,199]|
|Great Lakes spruce-fir||Picea-Abies spp.||35 to > 200|
|northeastern spruce-fir||Picea-Abies spp.||35-200 |
|southeastern spruce-fir||Picea-Abies spp.||35 to > 200 |
|Engelmann spruce-subalpine fir||Picea engelmannii-Abies lasiocarpa||35 to > 200 |
|black spruce||Picea mariana||35-200|
|conifer bog*||Picea mariana-Larix laricina||35-200 |
|blue spruce*||Picea pungens||35-200 |
|red spruce*||Picea rubens||35-200 |
|pine-cypress forest||Pinus-Cupressus spp.||1]|
|jack pine||Pinus banksiana||40]|
|Rocky Mountain lodgepole pine*||Pinus contorta var. latifolia||25-340 [9,10,164]|
|Sierra lodgepole pine*||Pinus contorta var. murrayana||35-200 |
|shortleaf pine||Pinus echinata||2-15|
|shortleaf pine-oak||Pinus echinata-Quercus spp.||187]|
|Colorado pinyon||Pinus edulis||10-400+ [49,56,84,129]|
|slash pine||Pinus elliottii||3-8 |
|Jeffrey pine||Pinus jeffreyi||5-30|
|western white pine*||Pinus monticola||50-200 |
|longleaf-slash pine||Pinus palustris-P. elliottii||1-4 [124,187]|
|longleaf pine-scrub oak||Pinus palustris-Quercus spp.||6-10 |
|Pacific ponderosa pine*||Pinus ponderosa var. ponderosa||1-47 |
|interior ponderosa pine*||Pinus ponderosa var. scopulorum||2-30 [1,6,94]|
|Arizona pine||Pinus ponderosa var. arizonica||2-15 [6,30,148]|
|Table Mountain pine||Pinus pungens||187]|
|red pine (Great Lakes region)||Pinus resinosa||10-200 (10**) [40,51]|
|red-white-jack pine*||Pinus resinosa-P. strobus-P. banksiana||10-300 [40,70]|
|pitch pine||Pinus rigida||6-25 [21,71]|
|pond pine||Pinus serotina||3-8|
|eastern white pine||Pinus strobus||35-200|
|eastern white pine-eastern hemlock||Pinus strobus-Tsuga canadensis||35-200|
|eastern white pine-northern red oak-red maple||Pinus strobus-Quercus rubra-Acer rubrum||35-200|
|loblolly pine||Pinus taeda||3-8|
|loblolly-shortleaf pine||Pinus taeda-P. echinata||10 to < 35|
|Virginia pine||Pinus virginiana||10 to < 35|
|Virginia pine-oak||Pinus virginiana-Quercus spp.||10 to < 35|
|sycamore-sweetgum-American elm||Platanus occidentalis-Liquidambar styraciflua-Ulmus americana||187]|
|galleta-threeawn shrubsteppe||Pleuraphis jamesii-Aristida purpurea||< 35 to < 100|
|eastern cottonwood||Populus deltoides||129]|
|aspen-birch||Populus tremuloides-Betula papyrifera||35-200 [40,187]|
|quaking aspen (west of the Great Plains)||Populus tremuloides||7-120 [1,62,113]|
|Texas savanna||Prosopis glandulosa var. glandulosa||129]|
|black cherry-sugar maple||Prunus serotina-Acer saccharum||> 1,000 |
|Rocky Mountain Douglas-fir*||Pseudotsuga menziesii var. glauca||25-100 [1,3,4]|
|coastal Douglas-fir*||Pseudotsuga menziesii var. menziesii||40-240 [1,123,136]|
|California mixed evergreen||Pseudotsuga menziesii var. menziesii-Lithocarpus densiflorus-Arbutus menziesii||< 35|
|California oakwoods||Quercus spp.||1]|
|oak-hickory||Quercus-Carya spp.||< 35|
|northeastern oak-pine||Quercus-Pinus spp.||10 to 187]|
|oak-gum-cypress||Quercus-Nyssa-spp.-Taxodium distichum||35 to > 200 |
|southeastern oak-pine||Quercus-Pinus spp.||187]|
|coast live oak||Quercus agrifolia||2-75 |
|white oak-black oak-northern red oak||Quercus alba-Q. velutina-Q. rubra||187]|
|canyon live oak||Quercus chrysolepis||<35 to 200|
|blue oak-foothills pine||Quercus douglasii-P. sabiniana||1]|
|northern pin oak||Quercus ellipsoidalis||187]|
|Oregon white oak||Quercus garryana||1]|
|bear oak||Quercus ilicifolia||187]|
|California black oak||Quercus kelloggii||5-30 |
|bur oak||Quercus macrocarpa||187]|
|oak savanna||Quercus macrocarpa/Andropogon gerardii-Schizachyrium scoparium||2-14 [129,187]|
|chestnut oak||Quercus prinus||3-8|
|northern red oak||Quercus rubra||10 to < 35|
|post oak-blackjack oak||Quercus stellata-Q. marilandica||< 10|
|black oak||Quercus velutina||187]|
|interior live oak||Quercus wislizenii||1]|
|blackland prairie||Schizachyrium scoparium-Nassella leucotricha||187]|
|little bluestem-grama prairie||Schizachyrium scoparium-Bouteloua spp.||129]|
|redwood||Sequoia sempervirens||5-200 [1,48,162]|
|southern cordgrass prairie||Spartina alterniflora||1-3 |
|baldcypress||Taxodium distichum var. distichum||100 to > 300|
|pondcypress||Taxodium distichum var. nutans||124]|
|western redcedar-western hemlock||Thuja plicata-Tsuga heterophylla||> 200 |
|eastern hemlock-yellow birch||Tsuga canadensis-Betula alleghaniensis||> 200 |
|western hemlock-Sitka spruce||Tsuga heterophylla-Picea sitchensis||> 200 |
More info for the terms: codominant, cover, density, formation, hardwood, presence, shrub, shrubs, succession, tree, vines
Bush honeysuckles are likely to occur across a variety of successional habitats within their North American range . Luken and McKnight  suggest that Amur honeysuckle can dominate habitats ranging from recently disturbed areas to mature forest.
Several studies document the influence of bush honeysuckles in old-field succession. Foster and Gross  examined woody plant establishment following abandonment of cultivation in a southwestern Michigan old-field habitat. Tatarian honeysuckle established in the 3rd season following abandonment. Average density increased steadily from 6.7 stems/acre in the year of establishment, to 91.1 stems/acre 4 years later (7 years following abandonment of agriculture). Fike and Niering  documented nearly 40 years of vegetation change in an old-field habitat in southeastern Connecticut. This site was cultivated until about 1945 and grazed until 1951. The initial survey in 1954 indicated a continuous perennial herbaceous cover with scattered woody plants <3.3 feet (1 m) tall. By 1960 the herbaceous community had decreased slightly and tree species were becoming more established. Presence of Morrow's honeysuckle (<1% cover) was first recorded in 1973, in conjunction with development of a thicket community of small trees, shrubs, and woody vines. This woody stratum was 13 to 20 feet (4-6 m) in height and had >40% cover. By 1983 Morrow's honeysuckle cover increased to 10% in a community characterized as "a young hardwood forest", with tree cover of 90%, shrub/vine cover of 85%, and herbaceous cover decreasing to about 40%. By the end of this study in 1993, there was a relatively continuous tree canopy 40 to 59 feet (12-18 m) in height, a 3.3 to 6.6 feet (1-2 m) high shrub stratum contributing about 50% cover, and 10% herbaceous cover. Morrow's honeysuckle was the dominant shrub species at 24% cover. Vankat and Snyder  examined floristics of a chronosequence of nearby stands corresponding to old-field/deciduous-forest succession in southwestern Ohio. Amur honeysuckle was present, but not common, in a 10-year-old goldenrod (Solidago spp.)- and fescue (Festuca spp.)-dominated old field, and in an approximately 50-year-old goldenrod-dominated old field with a sparse white ash-black cherry tree stratum. Amur honeysuckle was codominant in the understory (3.3 to 9.8 feet (1-3 m) tall) with sugar maple, and codominant in the ground layer (<3.3 feet (1 m)) with jewelweed (Impatiens capensis), in an approximately 90-year-old sugar maple-slippery elm forest. Vankat and Snyder  concede that conclusions about successional status of Amur honeysuckle, based on the above data, are limited by the study design. Nevertheless, Amur honeysuckle was common in a young closed-canopy forest stand, sparse in 2 old fields with some woody plant composition but no closed canopy, and absent from a 2-year-old abandoned agricultural field and an old-growth American beech-sugar maple forest.
It appears that bush honeysuckle establishment is often facilitated by some form of habitat disturbance . In forested habitats, Amur honeysuckle performs best near edges and in canopy gaps, where light levels are favorable .
Once established, the ability of bush honeysuckles to persist and spread within various successional habitats is less clear. Hutchinson and Vankat  assert that late successional forests dominated by shade-tolerant tree species such as sugar maple and American beech are more resistant to Amur honeysuckle invasion, probably due to low light levels near the forest floor. Luken  suggested forest patches having complete canopy closure can resist Amur honeysuckle invasion, but if canopy gaps are created, Amur honeysuckle can establish and persist. Bush honeysuckle populations, once established, can persist for many years. Age of Bell's honeysuckle shrubs studied in southern Wisconsin ranged from 12 to 34 years, with a mean of 20.4 years .
There are suggestions that bush honeysuckles could alter successional trajectories in ways that favor their persistence. Collier and others  hypothesized that Amur honeysuckle invasion may alter patterns of forest succession in southwestern Ohio. If development of a dense Amur honeysuckle shrub layer suppresses establishment of shade-tolerant tree seedlings, recruitment of mid- and late successional tree species may be inhibited. Hypothetically then, as older canopy trees die, closed-canopy forests could change to open-canopy woodlands or even Amur honeysuckle-dominated shrublands. Luken  demonstrated that Amur honeysuckle dominance in the shrub layer of northern Kentucky hardwood forests can suppress advance regeneration of overstory species. Woods  came to a similar conclusion after finding a significant (p<0.01) negative correlation between Tatarian honeysuckle cover and tree seedling (<3.3 feet (1 m tall)) density (in this study Tatarian honeysuckle and Bell's honeysuckle were not distinguished, although the text referred only to Tatarian honeysuckle (see Taxonomy)). In contrast, tree seedlings 3.3 to 6.6 feet (1-2 m) tall were not significantly (p<0.05) related to Tatarian honeysuckle cover. Woody seedlings in this larger size class frequently overtopped Tatarian honeysuckle. Examination of growth rings revealed that establishment of these larger seedlings predated Tatarian honeysuckle invasion. It was speculated that persistent Tatarian honeysuckle cover could suppress advance regeneration of overstory species, possibly leading to changes in canopy composition or even conversion of forests to more open canopies or shrublands. Gorchov and Trisel  provide direct evidence that Amur honeysuckle invasion can inhibit tree seedling establishment in southwestern Ohio. For further discussion, see Impacts.
Shade tolerance: Most sources characterize bush honeysuckles as intermediate in shade tolerance, adapted to grow in full sun to partial shade conditions [17,95,127,173,174,175,177,185]. Descriptions of Amur honeysuckle shade tolerance range from shade intolerant  to intermediate  to tolerant  to "amazing" shade tolerance .
Virginia Department of Conservation and Recreation  indicates the following light regimes for bush honeysuckles occurring in Virginia:
|Full Sun||Part Sun||Shade|
One reason often cited for an apparent competitive advantage of bush honeysuckles over native shrubs (see Impacts) is the ability of bush honeysuckles to respond rapidly to changes in light availability. While bush honeysuckles have some ability to establish and persist in relatively low light environments, growth is typically greatest under high light availability. For example, Harrington and others  found Bell's honeysuckle aboveground growth rates were significantly (p=0.0008) higher in open habitat than under a mature closed-canopy hardwood forest. Amur honeysuckle can respond to gap formation in otherwise shaded habitats via phenotypic plasticity in photosynthetic capability of shade-grown leaves, as well as by rapid stem elongation and production of new leaves with even greater photosynthetic capacity . Luken and others  compared performance of Amur honeysuckle shrubs with the indigenous shade-tolerant northern spicebush (Lindera benzoin), under shade and full sun. They transplanted forest-grown plants into pots and grew them in a greenhouse under full sun, 25% full sun, or 1% full sun treatments. Relative stem growth rates (integrated over an 11-week period) under the 1% full sun treatment were similar for both species. Amur honeysuckle stem growth was greatest under full sun, while maximum stem growth for northern spicebush occurred under the 25% full sun treatment. Amur honeysuckle stem growth was substantially greater than spicebush under both the 25% and 100% full sun treatments. Observed differences in leaf-level morphology and physiology, and biomass allocation indicate that Amur honeysuckle has a much greater ability than spicebush to acclimate and respond to enhanced light levels that might occur following forest canopy disturbance. This trait, coupled with an ability to tolerate heavy shade comparable to native shade-tolerant shrubs such as spicebush, indicates Amur honeysuckle may be highly competitive under a range of conditions, and may persist under a variety of successional stages and pathways in eastern deciduous forests.
Bush honeysuckles regenerate from seeds, as well as vegetatively following disturbance.
Pollination: According to Hauser  Morrow's honeysuckle, Tatarian honeysuckle, and Bell's honeysuckle are pollinated by bumblebees. Graenicher states  Bell's honeysuckle is pollinated by a variety of bees and perhaps by hummingbirds.
Seed production: Information about seed production is sparse, but it is apparent that some bush honeysuckles are capable of producing substantial numbers of seeds. Barnes  indicates Bell's honeysuckle produces consistent annual seed crops. A single "typical" Bell's honeysuckle shrub, about 6.6 feet (2 m) tall, growing in southern Wisconsin, produced 3,554 berries in 1 year. Numbers of seeds/fruit, sampled from several shrubs at this site, averaged 5 to 7, indicating that a "typical" plant may produce >20,000 seeds annually .
Estimates of annual fruit production for Amur honeysuckle and European fly honeysuckle in southwestern Ohio ranged from 0 to 1.2 million berries per plant, and approximately 400 million berries ha-1 . According to Welsh and others , winter honeysuckle fruits are "seldom formed", although no further explanation was provided.
There is some evidence for shrub age and size as determinants of reproductive ability. According to Sharp and Belcher , Amur honeysuckle plants begin flowering in the 3rd or 4th year, after which flowers appear on stems 2 years old and older. Deering and Vankat  compared reproductive state with shrub age and height within an Amur honeysuckle population in southwestern Ohio. Established shrubs took 3-8 years to reach reproductive age. At age 3 only 5.7% of individuals were reproductive, while >50% were reproductive by age 5. All shrubs ≥8.2 feet (2.5 m) tall were reproductive, while none <3.3 feet (1 m) tall were reproductive. Bell's honeysuckle shrubs may also produce fruit at as young as 3 years of age .
Site characteristics may also affect seed production. Amur honeysuckle flowering and fruiting were significantly (p=0.001 and p=0.03, respectively) correlated with light availability in southern Vermont .
Seed dispersal: Several sources indicate bush honeysuckle seeds are dispersed primarily by frugivorous birds [7,80,93,126,186]. Bartuszevige and Gorchov  showed that a wide variety of bird species consumed Amur honeysuckle fruit in southwestern Ohio. They also confirmed that American robins dispersed viable Amur honeysuckle seed, usually into woodlot edge and fencerow habitats. White-tailed deer may also consume and disperse viable seeds of Tatarian honeysuckle, Morrow's honeysuckle, Bell's honeysuckle, and Amur honeysuckle . Barnes  suggests that "many, if not most" fruits fall near the parent plant. For more information see Importance to Livestock and Wildlife.
Seed banking: It appears the potential for bush honeysuckles to form seed banks is low, but more research is needed to confirm this assertion and to determine interspecific differences. According to Luken and Mattimiro , seeds of Amur honeysuckle are "not long-lived in the soil." Hidayati and others  concluded that neither winter honeysuckle, Amur honeysuckle, or Morrow's honeysuckle have the potential to form persistent seed banks. However, Stevens and Jorgensen  found 12-year-old Tatarian honeysuckle seed to be still viable. They compared germination in seed stored for 12 years with fresh seed. Stored seeds were kept in a dry, open warehouse where temperatures over a 25-year study period ranged from -21.8 to 100.9 Â°F (-29.9 to 38.3 Â°C). Seeds from both lots were germinated over a 16-month period in a refrigerator (34 to 38 Â°F (1.1-3.3Â°C)). Germination rates were 57% for fresh seed and 31% for 12-year-old seed.
Germination: Bush honeysuckle germination requirements are variable between species.
Winter honeysuckle seeds require warm plus cold stratification prior to breaking dormancy. Although seeds mature in late spring/early summer, they generally will not germinate until late winter/early spring of the following year. A greenhouse study by Hidayati and others  indicated that winter honeysuckle seeds germinated while buried under 2 inches (5 cm) of leaf litter or 2.8 inches (7 cm) of soil.
Stratification requirements for Amur honeysuckle seed germination are unclear. According to Luken and Goessling , seeds are released in a nondormant condition, and germinate easily in warm, moist conditions. According to Hidayati and others , Amur honeysuckle seeds require a period of either warm- or cold stratification. Nevertheless, they are dispersed in fall and may germinate in fall or spring [74,103]. According to Hidayati and others , if seeds mature early enough and are subjected to a sufficiently long warm stratification period prior to onset of cold winter temperatures, they may germinate in fall. Late-maturing seeds are cold-stratified over winter, and will germinate in early spring when warm temperatures induce embryo growth.
Light seems to enhance Amur honeysuckle seed germination, but it is not obligatory. In a laboratory experiment, Amur honeysuckle germination was significantly (p<0.01) higher in light (35 Âµmol m-2 s-1, 14/10 hour light/dark photoperiod) than in dark (light excluded). Nevertheless, after 88 days, mean cumulative germination ranged from 53.7% to 81.3% in light, and from 31.3% to 55.0% in dark . Hidayati and others  found that Amur honeysuckle seeds were not inhibited by burial under 2 inches (5 cm) of leaf litter or 2.8 inches (7 cm) of soil in a greenhouse.
Germination may be enhanced when seeds are separated from the fruit pulp. Bartuszevige and Gorchov  found that seeds within intact fruit were significantly less viable (44% germination) than either seeds that were separated from pulp by hand (76% germination) or seeds that had passed through the guts of American robins (86% germination), after 12 weeks of favorable laboratory germination conditions.
Morrow's honeysuckle seeds, which are dispersed in summer, require warm stratification only and typically germinate prior to winter . Germination will occur in light or dark. Hidayati and others  found that, while Morrow's honeysuckle seeds germinated more readily under light than in dark under laboratory conditions, they were not inhibited by burial under 2 inches (5 cm) of leaf litter or 2.8 inches (7 cm) of soil in a greenhouse. A greenhouse study by Ruesink  demonstrated no effect of shading (25% vs. full light) on germination.
Tatarian honeysuckle seed germination is affected by scarification and cold stratification. Krefting and Roe  tested the effects of cold stratification, and ingestion and passage by American robins, on Tatarian honeysuckle seed germination. Unstratified seeds recovered from bird droppings germinated more readily (46%) than unstratified controls (24.5%). Stratification (90 days at 41 to 50 Â°F (5-10 Â°C) prior to feeding to birds) resulted in substantial improvement in germination, regardless of whether seeds had passed through bird guts (95% for bird ingested seeds, 92% for stratified controls). Although unstratified seed germinated more rapidly if passed through bird guts, this effect was not detected with stratified seed. Apparently avian frugivory has some positive effect on Tatarian honeysuckle seeds with seedcoat dormancy, while simultaneously, cold stratification is effective for breaking internal dormancy.
Seeds of Tatarian honeysuckle, Morrow's honeysuckle, Bell's honeysuckle, and Amur honeysuckle remain germinable following passage through the guts of white-tailed deer. Vellend  measured 76% germination for seeds collected from deer feces, compared with 81% for fresh-collected seeds.
Seedling establishment/growth: Bush honeysuckle seedling establishment appears most successful where litter cover and herbaceous competition are sparse [126,185]. Luken  found that after clipping established Amur honeysuckle plants in forested and pasture habitats, Amur honeysuckle seedlings established in forested plots at approximately twice the rate of those in pastures. In pasture plots, grasses and forbs were relatively undisturbed, and probably continued suppression of Amur honeysuckle seedlings. Barnes  sampled Bell's honeysuckle seedling density and frequency at 4 sites in southern Wisconsin. The site with the highest seedling frequency (39%) was characterized as a red pine (Pinus resinosa)- and eastern white pine (P. strobus)-dominated overstory and a sparse understory. This site had a primarily pine straw litter layer of variable depth over sandy loam and loamy sand soils. Within this site, Bell's honeysuckle seedlings were found within microsites having little to no litter cover. The site with the greatest seedling density (5,280 seedlings acre-1 ) contained a "very dense" population of mature Bell's honeysuckle shrubs, with near-continuous cover in some places. Observations indicate that at this site, seedlings occurred mainly under mature Bell's honeysuckle, where litter accumulation and herbaceous competition were sparse. Two other sites each had only 1 and 2 Bell's honeysuckle seedlings total. Among reasons provided for the paucity of seedlings at these sites were lack of soil disturbance, a thick layer of leaf litter from the oak overstory, and strong herbaceous and vine competition. A subsample was obtained from another section within 1 of these seedling-poor sites, where a dense population of mature Bell's honeysuckle shrubs had been eradicated during the previous year. Because of eradication treatments, plant litter and herbaceous competition were sparse. Consistent with other observations, substantial numbers of seedlings (26% occurrence, 2,560 seedlings acre-1) were found where litter cover and herbaceous competition were sparse and a seed source had been present .
However, the relationship between canopy cover and bush honeysuckle seedling establishment and growth is not straightforward. According to a review by Nyboer , bush honeysuckles commonly establish under tall shrubs or trees that serve as perch areas for seed-dispersing birds. As discussed above, canopy shading may also suppress strong herbaceous competition and permit greater bush honeysuckle seedling establishment. However, too much shading may result in reduced seedling establishment and growth . Luken and Goessling  studied Amur honeysuckle seedling establishment in forest patches dominated by sugar maple, white ash, and American elm in northern Kentucky. Seedling densities were greatest near the edges of forest patches and declined steadily toward their interior. While they were unable to establish a firm causal link between light levels and seedling densities, light levels and seedling densities were significantly positively correlated (p<0.05; r = 0.88) along transects from forest edge to interior. Ruesink  compared Morrow's honeysuckle greenhouse-grown seedlings under full-sun conditions with identical seedlings grown under 25% of full sun. After 50 days, full-sun seedlings were twice as tall and produced 6 times more aboveground biomass. It is likely that in most habitats where seeds are present, such as under the canopy of preexisting bush honeysuckle shrubs or where frugivorous birds find perch sites proximate to fruiting bush honeysuckles, any disturbance that increases light at ground level is likely to release bush honeysuckle seedlings .
Deering and Vankat  described the age structure and allometric development of a relatively isolated Amur honeysuckle population of recent origin (colonized ~ 1979) in southwestern Ohio. First-year shrubs averaged 1.3 feet (0.4 m) tall and 2 basal stems per shrub. Most individuals were >3.3 feet (1 m) tall by their 3rd year. For the first 4 years of development, numbers of 1-year-old stems averaged 2.2 to 2.6 per shrub. Individual plants averaged 4.3 total stems per plant by age 3. As shrubs reached reproductive age (beginning at 3-8 years), height growth continued, but recruitment of new stems ceased. With time, resource allocation shifted from new basal stem production and height growth in young shrubs to a balance of height growth, radial growth of existing stems, and reproduction in older shrubs .
Asexual regeneration: Information on asexual regeneration in bush honeysuckles is generally sparse. Studies cited below are specific to Amur honeysuckle and Bell's honeysuckle. Although it seems likely that these traits are shared by other bush honeysuckles, applicability to other taxa is not confirmed. Since Bell's honeysuckle is a hybrid of Morrow's honeysuckle and Tatarian honeysuckle (see Taxonomy), it is likely that either or both of the parent species share the traits discussed for Bell's honeysuckle below. More information is needed to determine similarities and differences in the biology of asexual regeneration in bush honeysuckles.
Amur honeysuckle will sprout from adventitious buds on the root crown in response to stem damage [105,168]. Repeated cutting throughout the growing season results in continued but diminished sprouting (see Physical/mechanical control) . The sprouting response of Amur honeysuckle to any particular stem damage event does not appear to diminish with stem age .
Bell's honeysuckle reproduces asexually by root suckering and layering . Barnes  studied root suckering and layering in 4 populations of Bell's honeysuckle in Wisconsin. Between 4 and 7% of shrubs sampled exhibited suckers. Suckers were encountered primarily on small shrubs, and those found on large, mature plants were usually within 2 to 3 feet (60-90 cm) of the root crown. Frequency of layering was estimated by examining all branches of sampled shrubs in contact with the soil surface for evidence of root development. Layering frequency varied between sites, with 1 site having 3% of shrubs showing evidence of layering, 2 sites having 9%, and a 4th site 19%. Layering frequency appeared to be positively related to soil moisture and duration of contact between branch and soil, although there were no supporting data. Barnes  also indicated that suckering and layering occurred most frequently on sites where Bell's honeysuckle seedling establishment was poorest.
Growth Form (according to Raunkiær Life-form classification)
More info for the terms: geophyte, hemicryptophyte, phanerophyte
RAUNKIAER  LIFE FORM:
Life History and Behavior
More info for the terms: formation, phenology, shrub, shrubs, tree
A reason often cited for an apparent competitive advantage of bush honeysuckles over native shrubs (see Impacts) is the advantage of longer leaf display. Barnes  compared Bell's honeysuckle leaf lifespan with that of several native shrub species in southern Wisconsin. Average period for functional leaves on Bell's honeysuckle shrubs at this site was May 18 to October 30. The following table provides observed duration from fully expanded leaves to the time when 50% of leaves are no longer green for sympatric shrub species in southern Wisconsin.
|Bell's honeysuckle||gray dogwood (Cornus racemosa)||red-osier dogwood (Cornus sericea)||American hazel (Corylus americana)||chokecherry (Prunus virginiana)||common buckthorn||red raspberry (Rubus idaeus)|
Trisel  studied leaf phenology of Amur honeysuckle and some sympatric tree and shrub species in southwestern Ohio. During all 3 years studied, Amur honeysuckle displayed fully expanded leaves for significantly (p<0.001) longer than all native species (except northern red oak and slippery elm whose small sample size precluded comparison). Amur honeysuckle was always the 1st species to expand leaves and the last to lose them. During 1994, Amur honeysuckle began leaf expansion in March and retained leaves into late November. The following table provides, for each species studied, the mean number of days between the initial and last census date during which expanded green leaves accounted for >50% of that individual's total number of leaves.
|Year 1||Year 2||Year 3|
|Sugar maple|| |
|Ohio buckeye (Aesculus glabra)||111||136||122|
|pawpaw (Asimina triloba)||166||165||167|
|eastern redbud (Cercis canadensis)||151|
|blue ash (Fraxinus quadrangulata)||162||155|
|northern red oak||174||169|
|blackhaw (Viburnum prunifolium)||195||185||189|
Harrington and others  demonstrated phenological differences in carbon gain between Bell's honeysuckle and the native shrub gray dogwood in southern Wisconsin. Under the canopy of a mature deciduous forest where understory light was about 4-6% of ambient levels, Bell's honeysuckle shrubs made approximately 47% of their annual carbon gain (35% in spring and 12% in fall) during two 2-week periods when gray dogwood was leafless. Leaf emergence in Bell's honeysuckle was around April 10, approximately 2 weeks prior to gray dogwood. Bell's honeysuckle retained leaves until around November 10, about 2 weeks longer than gray dogwood.
Reported flowering dates for bush honeysuckles:
|Winter honeysuckle||Amur honeysuckle||Morrow's honeysuckle||Tatarian honeysuckle||European fly honeysuckle||Bell's honeysuckle|
|southern Appalachians [144,145,146,147]||May-June||May-early June||May-early June||May-June|
|Blue Ridge Mountains (TN, VA) ||May-June|
|Connecticut [173,174,175,176,177]||early spring||mid-spring||mid-spring||May||May|
|Great Plains ||May-June|
|north-central Great Plains ||late April-May|
|northern Illinois ||mid-May to mid-June|
|Michigan||spring ||May , May-June ||May ||May |
|Minnesota ||May-June |
|New England ||May||mid-May to mid-June||mid-May to mid-June||mid-May to early June|
|eastern Canada and northern New England ||June|
|southern New England ||May|
|North Carolina Piedmont ||April-May|
|North Carolina Piedmont and Coastal Plain; South Carolina Piedmont ||February-early April|
|northeastern U.S. ||May-June|
|northeastern U.S. and adjacent Canada ||May-June||May-June|
|Ohio||May-June ||May ||May ||early May ||May |
|southwestern Ohio ||early May||early May|
|western Oregon [127,128]||December-March||after leaves begin unfolding|
|southeastern U.S. ||January-February|
|north-central Texas ||January-April||March|
|West Virginia ||May-June||May-June|
|Wisconsin||May-June ||mid-May to early June |
|southeastern Wisconsin ||May-June|
Reported fruiting dates for bush honeysuckles (This information represents a general guideline. Fruiting dates vary among sites and years. Observations of "fruiting" are typically subjective and may refer to fruit formation or ripening):
|Winter honeysuckle||Amur honeysuckle||Morrow' honeysuckle||Tatarian honeysuckle||European fly honeysuckle||Bell's honeysuckle|
|southern Appalachians [144,145,146,147]||August-September||June-July||June-July||June to mid-summer|
|Connecticut [173,174,175,176,177]||mid-summer||late summer||late summer||July-August||August|
|Great Plains ||mid-summer|
|north-central Great Plains ||July-August|
|southern New Jersey ||October-November|
|North Carolina Piedmont and Coastal Plain; South Carolina Piedmont ||April-May|
|northeastern U.S.||late fall ||July , July-August ||August |
|southwestern Ohio ||early June-early September||early June-early September|
|Western Oregon ||late spring|
|Wisconsin ||late June-late September|
Amur honeysuckle and Bell's honeysuckle fruit can persist on the plant well into winter [12,97,150,185], while Morrow's honeysuckle and Tatarian honeysuckle fruit abscise soon after ripening [150,185].
Biology and Spread
Open-grown exotic bush honeysuckles fruit prolifically and are highly attractive to birds. In the eastern United States, over twenty species of birds feed on the persistent fruits and widely disseminate seeds across the landscape. In established populations, vegetative sprouting also aids in the persistence of these exotic shrubs.
Molecular Biology and Genetics
Barcode data: Lonicera maackii
Statistics of barcoding coverage: Lonicera maackii
Public Records: 3
Specimens with Barcodes: 7
Species With Barcodes: 1
National NatureServe Conservation Status
Rounded National Status Rank: NNA - Not Applicable
Rounded National Status Rank: NNA - Not Applicable
NatureServe Conservation Status
Rounded Global Status Rank: GNR - Not Yet Ranked
Winter honeysuckle, Amur honeysuckle, Morrow's honeysuckle, Tatarian honeysuckle, and Bell's honeysuckle are ranked as severe threats by the Tennessee Exotic Pest Plant Council , and Amur honeysuckle, Morrow's honeysuckle, and Tatarian honeysuckle are ranked as severe threats by the Kentucky Exotic Pest Plant Council . Morrow's honeysuckle is ranked highly invasive, Amur honeysuckle and Tatarian honeysuckle moderately invasive, and winter honeysuckle and Bell's honeysuckle occasionally invasive by the Virginia Department of Conservation and Recreation .
U.S. Forest Service Region 8 (Southern Region) lists winter honeysuckle, Amur honeysuckle, Morrow's honeysuckle, and Tatarian honeysuckle as category 1 weeds (exotic plant species that are known to be invasive and persistent throughout all or most of their range within the Southern Region and that can spread into and persist in native plant communities and displace native plant species and therefore pose a demonstrable threat to the integrity of the natural plant communities in the Region). The introduction of category 1 species is prohibited on National Forest System Lands .
Morrow's honeysuckle, Tatarian honeysuckle, and Bell's honeysuckle are "high-priority" nonnative invasive plants of the Ottawa National Forest (MI) .
Prevention and Control
Relevance to Humans and Ecosystems
Other uses and values
Amur honeysuckle has been cultivated as an ornamental in North America [106,131,150], and as of 1996, was still commercially available . Beginning in the 1960s, USDA Soil Conservation Service developed and distributed the Amur honeysuckle cultivar 'Rem-Red' for use as an ornamental shrub, promoted as valuable for wildlife and as useful for soil conservation and as a windbreak, border, hedge, or screen [95,150]. Amur honeysuckle, along with Tatarian honeysuckle and Morrow's honeysuckle, is among species recommended for use in strip mine site reclamation [77,185]. Amur honeysuckle makes a very productive honey plant . However, due to its invasive propensity in natural and seminatural woodlands, Clark  recommends against its use outside urban areas where it is already an established part of the flora.
Tatarian honeysuckle has been cultivated as an ornamental [18,150,191], and Dirr  provides a list of 14 available cultivars. It has been recommended as a windbreak, shelterbelt, or hedge species [28,53,95,110,157,169], especially in areas with extreme seasonal temperatures . Tatarian honeysuckle has been characterized as useful for range restoration and soil stabilization , and has been used for streambank reclamation .
Bell's honeysuckle has been used for landscape and ornamental purposes in the northern U.S. .
Importance to Livestock and Wildlife
Although it appears bush honeysuckles are typically planted for other purposes, they may provide some value for wildlife and are occasionally planted for this use. According to Sharp and Belcher , Tatarian honeysuckle has been planted for summer wildlife food in the eastern U.S., and Luken and Thieret  state Amur honeysuckle has been planted in the eastern U.S. for wildlife habitat "improvement."
White-tailed deer browse bush honeysuckle twigs and foliage . Vellend  confirmed the presence of Tatarian honeysuckle, Morrow's honeysuckle, Bell's honeysuckle, and Amur honeysuckle seeds in white-tailed deer scat in central New York, but it is unclear if fruits were eaten purposely or inadvertently. Bush honeysuckle fruits are borne in leaf axils, so white-tailed deer that are browsing on leaves and twigs are also likely to ingest fruits in season. While Velland  did not specify which plant parts are selected by deer, he inferred that leaves and twigs are purposely browsed, and it is obvious that fruits are at least not avoided.
Tatarian honeysuckle is browsed by eastern cottontail [64,150]. Small mammals eat fallen bush honeysuckle fruit in late winter and early spring . Deer mice extract and consume Amur honeysuckle seeds from intact fruits. However, it is speculated that relative presence or absence of Amur honeysuckle has little effect on small mammal forage habitat quality, and conversely, small mammal seed predation probably has little influence on Amur honeysuckle fecundity .
Bush honeysuckle fruits are eaten at least occasionally by songbirds, and avian frugivory is thought to be an important bush honeysuckle seed dispersal mechanism. Amur honeysuckle fruits are eaten at least occasionally by songbirds, especially in winter, and Tatarian honeysuckle fruits are eaten by songbirds in summer, soon after maturity (see Seasonal Development) [64,95,150]. Bartuszevige and Gorchov  studied avian Amur honeysuckle seed dispersal in southwestern Ohio. Out of 17 bird species captured near fruiting Amur honeysuckle shrubs, 12 were found to have consumed fruit. American robin, cedar waxwing, European starling, hermit thrush, and northern mockingbird defecated viable Amur honeysuckle seed. American tree sparrow, Carolina chickadee, dark-eyed junco, northern cardinal, song sparrow, tufted titmouse, and white-throated sparrow displayed evidence of consuming fruit, but without evidence of passing viable seed. Species that showed no evidence of Amur honeysuckle frugivory included brown creeper, Carolina wren, downy woodpecker, golden-crowned kinglet, and white-breasted nuthatch. Eastern bluebirds were also observed eating Amur honeysuckle fruit, but were not captured in the study . Wild turkey, ruffed grouse, northern bobwhite, and ring-necked pheasant use Amur honeysuckle for food .
Palatability/nutritional value: As of this writing (2005) very little information is available concerning palatability/nutritional value of bush honeysuckles. Analysis of Amur honeysuckle and European fly honeysuckle fruit in southwestern Ohio showed C:N ratios ranging from 29:1 to 56:1 (mean = 41.1, SD = 9.17) and percent lipid ranging from 4.53% to 5.01% (mean = 4.78%, SD = 0.20) .
Although avian frugivory is thought to be an important seed dispersal mechanism, preference for bush honeysuckle fruit as a food source is unclear. McRae  noted Tatarian honeysuckle as a major dietary component of northern bobwhite, especially after mid-February, at 2 northwestern Georgia piedmont upland forest sites. Apparently the raisin-like fruit of Tatarian honeysuckle is not necessarily a preferred late-season food, but provides available forage for songbirds when preferred foods are scarce . According to Dirr , birds will consume Amur honeysuckle fruit if other food is unavailable.
Ingold and Raycraft  examined 115 individuals of 26 bird species for evidence of Amur honeysuckle and European fly honeysuckle frugivory in southwestern Ohio between mid-September and mid-November. Nine species (American robin, gray-cheeked thrush, Swainson's thrush, gray catbird, cedar waxwing, northern cardinal, purple finch, American goldfinch, and white-throated sparrow), and 21 of 82 individuals representing these species, showed evidence of feeding on fruits. Sixteen species (35 total individuals) exhibited no evidence of Amur honeysuckle/European fly honeysuckle fruit consumption (Carolina chickadee, tufted titmouse, brown creeper, white-eyed vireo, red-eyed vireo, Tennessee warbler, magnolia warbler, bay-breasted warbler, ovenbird, common yellowthroat, yellow-breasted chat, indigo bunting, American tree sparrow, field sparrow, fox sparrow, and song sparrow). The low proportion of species (and individuals within these species) found to be feeding on Amur honeysuckle and European fly honeysuckle fruit led to questions concerning forage quality, especially since these fruits were abundant and conspicuous in the study area.
Although information is sparse, in some cases bush honeysuckle frugivory may be harmful to birds. Casual observations indicate that Tatarian honeysuckle fruit can be toxic to birds , but this is not confirmed. There is some evidence to indicate birds that eat bush honeysuckle fruit may experience changes in feather coloration. Apparently Morrow's honeysuckle fruit contains the carotenoid rhodoxanthin, which causes normally yellow tail feather bands in cedar waxwings to appear orange. Similar yellow-to-orange changes in feather color have been described in Kentucky warblers and white-throated sparrows, perhaps also as a result of bush honeysuckle fruit consumption. While no definitive impact has yet been established as a result of this phenomenon, subtle differences in coloration within species may affect behavior such as mate selection .
Cover value: Bush honeysuckles probably provide some cover for wildlife. Amur honeysuckle provides nesting sites and protection for songbirds from late spring to late fall, and cover for rabbits . Tatarian honeysuckle provides year-round cover for birds and small mammals .
However, indirect effects of bush honeysuckle invasion on wildlife may be difficult to predict. Schmidt and Whelan  examined the effect of Amur honeysuckle invasion on nest predation of American robins in northern Illinois deciduous woodlands. Nests built in Amur honeysuckle had significantly (p<0.001) higher daily nest mortality rate compared with nests built in native species. Reasons offered for increased nest predation in Amur honeysuckle included lower nest height (compared with many native shrubs and trees), absence of sharp thorns (compared with native hawthorns (Crataegus spp.)), and branch architecture that may facilitate predator (e.g. raccoon) movement. Unfortunately, Amur honeysuckle may provide more attractive nest sites due to its early leaf flush (see Seasonal Development) and sturdy branches. In fact, American robins significantly (r2=0.912, p<0.01) increased their use of Amur honeysuckle over the 6-year study period. Wood thrush also nested in Amur honeysuckle, although use was apparently limited by competition from American robins.
Ecological Threat in the United States
Exotic bush honeysuckles can rapidly invade and overtake a site, forming a dense shrub layer that crowds and shades out native plant species. They alter habitats by decreasing light availability, by depleting soil moisture and nutrients, and possibly by releasing toxic chemicals that prevent other plant species from growing in the vicinity. Exotic bush honeysuckles may compete with native bush honeysuckles for pollinators, resulting in reduced seed set for native species. In addition, the fruits of exotic bush honeysuckles, while abundant and rich in carbohydrates, do not offer migrating birds the high-fat, nutrient-rich food sources needed for long flights, that are supplied by native plant species.
Ecological Threat in the United States
Lonicera maackii (Amur Honeysuckle or Bush Honeysuckle) is a species of honeysuckle in the family Caprifoliaceae, native to temperate Asia in northern and western China (south to Yunnan), Mongolia, Japan (central and northern Honshū, rare), Korea, and southeastern Russia (Primorsky Krai).
It is listed as an endangered species in Japan. It has escaped from cultivation and become naturalized in New Zealand and the eastern United States; in the forests of the latter, it has become an important invasive species.
It is a deciduous large shrub growing to 6 m tall with stems up to 10 centimeters diameter. The leaves are oppositely arranged, 5–9 cm long and 2–4 cm broad, with an entire margin, and with at least some rough hairs on them. The flowers are produced in pairs, commonly with several pairs grouped together in clusters; they are 2 cm long, two-lipped, white later turning yellow or light orange in color; flowering is from mid spring to early summer.
The fruit is a bright red to black semi-translucent berry 2–6 mm diameter containing numerous small seeds; they ripen in autumn, and are eaten by birds, which disperse the seeds in their droppings. It is fast growing and favours shady habitats such as the forest understory, neglected urban areas, and fence rows. It can form extremely dense thickets.
Cultivation and uses
It is grown as an ornamental plant for its attractive flowers, and also as a hedge. A number of cultivars have been selected for garden use, including 'Erubescens' with pink flowers, and 'Rem Red' with erect habit.
The flowers are sometimes used by children, who remove the blossom by hand, and pull off the bottom to suck out the sweet nectar in the center. The red berries are mildly poisonous to humans and should not be eaten.
As an invasive species
Spread of this plant is illegal or controlled in some areas of the United States due to its well documented invasive character. It is listed as a "invasive, banned" species in Connecticut, "prohibited" in Massachusetts, and a "Class B noxious weed" in Vermont. It is also officially listed as an invasive species by government agencies in Wisconsin and Tennessee.
This plant is adaptable and successful in a wide range of conditions. In the United States, Amur honeysuckle was once planted to control erosion, and as hedges. It spread quickly as birds eat the fruit and disperse the seeds, and was soon naturalized. Notably, in deciduous forest understories of the eastern United States it forms dense growths with thick canopies that shade out native shrubs, young trees, and wild flowers. Uncontrolled, these growths create a near monoculture of Amur Honeysuckle. This species poses a serious threat not only to the diversity of the ecosystems which they invade but also to forest regeneration itself. as the plant is known for reducing the growth and diversity of native seedlings. Moreover studies have shown that plant is responsible for having a negative impact on birds, and tadpoles. In 2010 a study in the St. Louis area showed that the plant increases the risk of tick-born diseases such as Erlichiosis and Lyme disease in suburban natural areas, by attracting deer activity and increasing the number of infected ticks. Furthermore, experimental removal of the plant was shown to reduce deer activity and number of infected ticks, through the shifting of ticks' blood meals away from deer.
Honeysuckle can be controlled by cutting, flaming, or burning the plant to root level and repeating on two-week increments until nutrient reserves in the roots are depleted. To ensure eradication newly cut stumps should be treated with herbicide. Control through prescribed burning has been found to be most effective during the seed dispersal phase (late summer, early fall). Honeysuckle can also be controlled through annual applications of glyphosate which thoroughly soak the leaves, or through grubbing of the shallowly rooted young plants. Both of these methods are only practical if high labor costs and soil damage are not of concern.
Due to the invasive nature of this species and the ecological threat it possess it may be inadvisable to cultivate this plant in climates similar to those found where this species has become invasive (e.g. eastern United States). It has been suggested that existing specimens found outside of their native range in east Asia should be removed and replaced with alternative non-invasive species. Possible alternative fast growing, shade tolerant, deciduous shrubs include Calycanthus floridus, Cornus mas, Cornus sericea, Forsythia hybrids, Hydrangea spp., Syringa vulgaris, Viburnum cassinoides, Viburnum dentatum, Viburnum dilatatum, Viburnum opulus, Viburnum prunifolium, Viburnum trilobum, Weigela florida.
The common name Amur Honeysuckle comes from the Amur River which is the world's eighth longest river. This river forms the border between the Russian Far East and Manchuria in China. L. maackii is native to the area surrounding this river. The species name maackii is derived from Richard Maack, a 19th-century Russian naturalist.
|Wikimedia Commons has media related to Lonicera maackii.|
- "The Plant List: A Working List of All Plant Species". Retrieved 7 December 2014.
- Germplasm Resources Information Network: Lonicera maackii
- Ministry of the Environment (Japan): Red List of Threatened Plants of Japan (excel spreadsheet)
- Red Data Book (Japan): Lonicera maackii (in Japanese, with map; google translation)
- USDA Forest Service Weed of the Week Fact Sheet (pdf file)
- Huxley, A., ed. (1992). New RHS Dictionary of Gardening. Macmillan ISBN 0-333-47494-5.
- Missouriplants: Lonicera maackii
- Invasive.org: Lonicera species (pdf file)
- Annotated Bibliography of primary research on invasive qualities of L. maackii in the USA: National Biological Information Infrastructure
- University of Connecticut Plant Database: L. maackii
- Ohio State University Pocket Gardener: L. maackii
- USDA PLANTS DATABASE: L. maackii
Names and Taxonomy
The currently accepted genus name for honeysuckle is Lonicera L.
(Caprifoliaceae) [18,36,54,59,82,83,93,133,161,189,190,191,197]. This report
summarizes information on 5 species and 1 hybrid of Lonicera:
Lonicera fragrantissima Lindl. & Paxt. [36,82,83,133,191] winter honeysuckle
Lonicera maackii Maxim. [18,27,36,54,59,82,83,131,137,186] Amur honeysuckle
Lonicera morrowii A. Gray [18,39,54,60,83,161,186,189,190,197] Morrow's honeysuckle
Lonicera tatarica L. [18,38,39,54,59,60,82,83,92,93,157,161,186,190,191] Tatarian honeysuckle
Lonicera xylosteum L. [18,54,60,83,186] European fly honeysuckle
Lonicera ÃÂ bella Zabel [54,60,83,133,186,190]
Bell's honeysuckle. This is a cross between L. tatarica and L. morrowii
that has arisen in cultivation and probably spontaneously in the wild [7,54,68,133,186].
According to Barnes , Bell's honeysuckle is intermediate to its parent species in most
characteristics, but "many of these characteristics vary between extremes so that often
detection of the hybrid nature of an individual can only be accomplished by looking at a large
number of characters." Field observations in Ohio and Michigan suggest that Tatarian
honeysuckle is the pollen parent of Bell's honeysuckle . Bush honeysuckle shrubs in
southern Vermont were described as a "hybrid complex", with most individuals
resembling the Morrow's honeysuckle type, but some apparently more influenced by Tatarian
honeysuckle . Barnes  illustrates the difficulty often encountered in distinguishing
between Bell's honeysuckle and its parent species, and asserts that in many instances
Bell's honeysuckle is misidentified in the literature as Tatarian honeysuckle or Morrow's
honeysuckle. Chapman and Bessette  describe Bell's honeysuckle as the dominant
"species" in Adirondack Park, New York, with pure specimens of either Tatarian
honeysuckle or Morrow's honeysuckle becoming difficult to find.
When discussing characteristics typical (or likely to be typical) of all 6 of the above
taxa, this report refers to them collectively as bush honeysuckle(s). When referring
to individual taxa, the common names listed above are used.
Although apparently not widely escaped, additional hybrids may be formed among the
abovementioned taxa and others, as described below [60,72]:
Lonicera ÃÂ minutiflora Zabel (bunchberry honeysuckle), a cross between L. morrowii
and L.ÃÂ xylosteoides.
Lonicera ÃÂ muendeniensis Rehd. (Muenden honeysuckle), a cross between L.ÃÂ bella
and L. ruprechtiana (Manchurian honeysuckle).
Lonicera ÃÂ muscaviensis Rehd. (Muscovy honeysuckle), a cross between L. morrowii and L. ruprechtiana.
Lonicera ÃÂ notha Zabel (Rutarian honeysuckle),
a cross between L. ruprechtiana and L. tatarica.
Lonicera ÃÂ salicifolia Dieck ex Zabel (willowleaf honeysuckle), a cross between
L. ruprechtiana and L.ÃÂ xylosteoides.
Lonicera ÃÂ xylosteoides Tausch (Vienna honeysuckle),
a cross between L. tatarica and L. xylosteum.
European fly honeysuckle
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