Botanical description: This description provides characteristics that may be relevant to fire ecology and is not meant for identification. Keys for identification are available (e.g., [19,23,60,65,78,93]). Since there are many native raspberries that resemble and co-occur with wineberry (see Habitat Types and Plant Communities), it is recommended that readers seek out these keys for positive identification before any control methods are undertaken.
Wineberry is a deciduous, thicket-forming shrub that produces upright and arching biennial canes from a perennial root system [42]. Canes average 1.6 to 4.9 feet (0.5-1.5 m) in length and may reach 9 feet (2.7 m) tall [23,42,65,73,78]. Canes are bristly and thorny and covered with distinctive glandular red hairs that are 0.1 to 0.2 inch (3-5 mm) long [23,39,65,73,78]. The hairs give the canes a reddish color when seen from a distance ([73], a fact sheet).
Wineberry leaves are compound [65] and consist of 3 serrated, blunt-tipped leaflets with purple veins that are densely white-tomentose underneath ([73], a fact sheet). Petioles are densely hairy [23]. The terminal leaflet is 1.6 to 3.9 inches (4-10 cm) long and about as wide [65]. Lateral leaflets are 1.0 to 3.1 inches (2.5-8.0 cm) long [65]. Wineberry has small greenish flowers with white petals that occur in a terminal panicle on glandular short-hairy pedicels [73,78]. The glandular-hairy calyx lobes envelop the developing fruits and keep them covered until almost ripe [18,39,78].Photograph courtesy of Leslie J. Merhhoff, University of Connecticut, Bugwood.org.
Wineberry fruit is 0.4 inch (1 cm) thick and shiny red [39,65,78]. Each fruit is composed of an aggregate of large succulent drupelets commonly referred to as a "berry". Each fruit contains numerous seeds that are 0.1 to 0.2 inch (2-4 mm) long [6,42].
Other: Field experiments in mixed-hardwood forest in Maryland suggest that arbuscular mycorrhizal fungi may have no impact or a negative effect on wineberry. Wineberry inoculated with arbuscular mycorrhizal fungi had similar survival rates and lower leaf weight, root weight, and total biomass than noninoculated control seedlings. The author concluded that "an absence of arbuscular mycorrhizal fungi would not limit the establishment of wineberry in new habitats" [42].
SEASONAL DEVELOPMENT:
Growth and development of wineberry is typical of blackberries. Wineberry produces biennial canes from a perennial root system or from underground rhizomes (see Vegetative regeneration) [42]. First year canes (primocane) are unbranched, sterile, entirely vegetative, and develop from rhizominous buds at or below the ground surface [25,42]. In the 1st year, carbon allocation is primarily into leaf production and cane elongation [42]. In the 2nd year, lateral branches develop in the axils of the primocanes and produce leaves, flowers, and fruits [42], but "do not have extensive growth" [25]. Second-year canes are referred to as "floricanes". Unlike primocanes, floricanes are woody [25].
In April, floricanes produce new leaves. In early May, new primocanes originate from the perennial root system [42]. In late May, floricanes undergo lateral branching and may produce flowers and fruit; fruit production occurs in late June to August. Fruits of wineberry ripen together [17]. After producing fruit in late summer, the leaves of floricanes senesce and the cane gradually dies. In Maryland, wineberry loses its leaves in late November [42]. Generalized fruiting and flowering dates are as follows:
Location Flowering Fruiting Maryland late May-early June late June-July [42] Arkansas May-June June-July [39] New England --- 16 July-30 August [70] New York 20 May-31 May [16] --- North Carolina --- July-October, peak in August [27]The information available (2009) provides no clear direction for using fire as a management tool for wineberry. Because wineberry frequently invades after disturbance, prescribed fire and fuels management activities may increase its abundance [12]. The mechanisms by which wineberry establishes after fire are not completely understood, but establishment of wineberry through sprouting and/or seedling establishment from on- and off-site sources after fire is likely.
Preventing invasive plants from establishing in weed-free burned areas is the most effective and least costly management method. This may be accomplished through early detection and eradication, careful monitoring and follow-up, and limiting dispersal of invasive plant seed into burned areas. Specific recommendations include:
For more detailed information on these topics see the following publications: [3,7,24,84].
Fire adaptations and plant response to fire: Blackberries frequently respond to fire by rapidly increasing in abundance, but the response of blackberries to fire differs among species. Little information is available regarding wineberry's response to fire, but wineberry is often found on disturbed sites and, like some other blackberries, is likely to quickly occupy postfire habitat and persist for decades after fire (see Successional Status). In clearcut and burned sub-boreal spruce (Picea spp.) forest in northern British Columbia, red raspberry established rapidly after fire, peaking in cover during postfire year 3 (27.5%). Its cover declined over time, but mean percent cover 10 years after fire (0.95%) was higher than prefire cover (0.02%). The fire was low to moderate severity and consumed 22% of the forest floor [32]. In the Superior National Forest in Minnesota, vegetation changes were observed 11 and 14 years after fire in jack pine-black spruce (Pinus banksiana-Picea mariana) forest and jack pine plantation forest, respectively. These fires were "patchy", "hot", and resulted in "little or no soil burn". In this study, frequency of red raspberry at burned areas 11 and 14 years after fire (range: 53-87%) was greater than at unburned control areas (range: 20-23%). In contrast, dwarf raspberry (Rubus pubescens) responded differently at different areas of the burn, and no consistent response was detected for this species [50]. In white oak-bur oak (Quercus marcocarpa) woodlands in southwestern Wisconsin, stem densities of blackberries (red raspberry, black raspberry, and Allegheny blackberry) were not changed after 2 consecutive years of prescribed fire; however, no information was provided on fire severity [35]. Allegheny blackberry cover increased from nearly 10% before fire to over 50% after a low-severity surface fire in northern pin oak (Quercus ellipsoidalis) forest in Stevens Point, Wisconsin. In this study, mean flame height was <1 foot (0.3 m) and mean rate of spread for the headfire was 3.3 m/min [67]. Areas with annual (burned each year from 1995 to 1999) and periodic (burned in 1996 and 1999) spring prescribed fires in mixed-oak forest in Ohio typically had higher mean frequencies (approximate range: 22-45%) of blackberries than unburned sites (approximate range: 18-22%). Frequency of blackberries in annually and periodically burned areas were similar and tended to increase over time, while frequency at unburned sites remained relatively stable during the same time period. In this study, blackberries were a significant indicator of burned sites (P<0.01); however, the species of blackberry were not specified. Flame lengths were typically <0.5 m and fuel consumption was generally limited to unconsolidated leaf litter and small woody debris (1-hr fuels). Over 80% of the sites were burned, resulting in "relatively minor" reductions in overstory density. Other characteristics of the fires are provided in [40]. In a chronosequence study in mixed-coniferous forest on the western redcedar/queencup beadlily (Thuja plicata/Clintonia uniflora) habitat type in Idaho, cover and density of Pacific blackberry (Rubus ursinus) and blackcap raspberry (Rubus leucodermis) did not differ with burn severity ("high severity" and "low severity") or burn age (postfire year 1, 2, 3, 4, 5, and 15) [61].
Wineberry may occupy postfire habitat by sprouting and/or seedling establishment as do many blackberries (e.g., [1,31,32,50,61,76]). For example, red raspberry in clearcut and burned sub-boreal spruce forest in northern British Columbia established after fire from buried seeds and from sprouting of plants present before the fire [31,32]. Presence of wineberry after prescribed fire was reported in Prentice Cooper State Forest and Wildlife Management Area in Tennessee [5]; in this study it was unclear whether wineberry established through on- or off-site sources. Wineberry seeds may accumulate in soil seed banks, so establishment of wineberry from the seed bank may be possible (see Seed banking). Wineberry may also establish after fire from seed brought on site by animals (see Seed dispersal). Fire may favor wineberry, like other blackberries, by increasing available nutrients [13,67]. Many blackberries require exposed mineral soil and light for germination [71], and fire may create a favorable seedbed for blackberries by creating these conditions (see [62] for a review).
Fuels: Little information is available on the fuel characteristics of wineberry invaded sites as of 2009. Like some other blackberries, the canes and foliage of wineberry are likely highly flammable (see Immediate fire effect on plant). In addition, wineberry may form dense thickets ([73], a fact sheet), leading to complete change of physical structure in invaded communities. Thus, wineberry has the potential to substantially alter fuel loads and fire behavior. More information is needed on these topics.
FIRE REGIMES: Little information is available on the FIRE REGIMES of plant communities in wineberry's native habitat. Its ability to sprout from rhizomes and the possibility of establishment from on-site seeds stored in the soil seed bank suggest that wineberry may be favored by fires of low severity and short duration that remove little of the surface organic layer [68]. In addition, the possibility of establishment from off-site, animal-dispersed seeds, its ability to grow rapidly in high light and on exposed mineral soil, and its appearance in early-successional plant communities in North America (see Habitat Types and Plant Communities) suggest that the species would be tolerant of short fire-return intervals and stand-replacing disturbances. However, because wineberry may not reach adequate size for fruiting or tip-rooting until 3 years of age or more [25], fire-return intervals >3 years are likely most favorable to wineberry persistence. Persistence into midsuccessional stages and probable longevity in the soil seed bank suggest that moderate to long fire-return intervals may be tolerated. The Fire Regime Table summarizes characteristics of FIRE REGIMES for vegetation communities in which wineberry may occur. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find FIRE REGIMES".
As of this writing (2009), little English-language literature is available on wineberry's native habitats. What information is available indicates that wineberry grows at low to medium elevations in montane valleys and along roadsides in China [94]. In Japan, wineberry occurs in lowland and mountainous regions in clearings associated with spruce (Picea spp.), fir (Abies spp.), and birch (Betula spp.) [82]. In South Korea, wineberry occurs at elevations ranging from 70 to 460 feet (20-140 m) along streambanks [49].
In the eastern United States, wineberry occupies a wide range of habitats including early- to midsuccessional forest, floodplain forest, herbaceous and shrub wetland, wet meadows, riparian corridors, old fields, open disturbed areas, burned areas, trailsides, roadsides, ditches, and vacant lots, as well as ecotones between these habitats [4,5,22,28,42,58,59,64,65,70,80,91,93].
According to reviews, wineberry prefers open, mesic conditions with rich soils but tolerates a wide range of soil types, textures, and pH values [15,22,73,87]. At Great Falls Park in northeastern Virginia, wineberry occurred on soils ranging from "relatively fertile", with basic pH, and silt loam to silty clay loam textures to dry, "extremely acidic, infertile" silty clay loams. At this site, wineberry occurred on very dry upper slopes and ridge crests with "high solar exposure and low moisture potential" as well as seasonally flooded swamps [74]. Wineberry was found in Sussex County, New Jersey on trails and roadsides where soils were thin and rocky though moist [4]. In Chittenden County, Vermont, wineberry established on a limy talus slope in the dense shade of northern whitecedar (Thuja occidentalis) [99]. Wineberry occurred relatively infrequently in sweetgum (Liquidambar styraciflua) -sycamore streambank habitat with sandy soils in Newton County, Arkansas; this site was regularly disturbed by spring and fall flooding and anthropogenic influences [81]. In New Jersey, wineberry occurred in constructed wetlands with coarse soil [53]. In Inwood Hill Park in New York, wineberry occurred on some sites with "deep soils" [54]. Wineberry occurred on wet, seasonally flooded and mesic soils at the Piscataway and Fort Washington National Parks in Maryland [75]. Along a 250-mile (402 km) reach of the New River Gorge in West Virginia, wineberry was found at a variety of sites including regularly flooded streambeds, riverside beach areas, and wooded upper beach areas with soils ranging from cobblestone and gravel to sand and mudflats. Additional sites occupied by wineberry in this study included rocky summits and cliff faces and woodlands with shallow and sandy soils [79].
According to reviews, wineberry tolerates a range of light levels, with light availability in suitable habitat ranging from full sun to partial shade [22,64,73,87]. Although established plants may persist in low light, wineberry germination and survival appear best in moderate to high light environments (see Seedling establishment and plant growth) [25]. In field experiments in mixed-hardwood forest in Maryland, wineberry seedling survival was significantly reduced under leaf litter (P<0.001); this was attributed to a lack of light and increased potential for root rot as a result of increased moisture levels [42]. Although wineberry tolerates a variety of light levels and soil conditions, like other blackberries, adequate soil moisture and light appear important for best growth and fruit production (see Successional Status) [12,64].Photograph courtesy of John M. Randall, The Nature Conservancy, Bugwood.org
Elevation/Topography: Wineberry occurs in lowlands and mountainous terrain on slopes ranging from 0% to 60%. At Evansburg State Park, wineberry occurred between 98 and 397 feet (30-121 m) elevation. Wineberry occurred at Fernow Experimental Forest in north-central West Virginia at elevations ranging from 1,749 to 3,648 feet (533-1,112 m) and slopes ranging from 10% to 60% [56]. In Inwood Hill Park in southern New York, wineberry occurred on sites with slopes >10% [54]. Wineberry occurred on sites where slopes averaged >30% in Great Falls Park in Virginia .
Climate: Wineberry is hardy to USDA hardiness zone 5, where average annual minimum temperatures are as low as -20 °F (-26 °C) [46,64]; although some damage may be caused to the plant at this temperature, the plants "usually recover well" (Davis 1990, as cited in [64]).
Precipitation may affect wineberry density. In Maryland, drought reduced the density of wineberry and sawtooth blackberry, with greater mortality in forest edge sites than in intact forest (P=0.034). The subsequent year's precipitation was average; although wineberry density increased during that year, its density did not return to pre-drought levels [42].
Wineberry occurred throughout Great Falls Park in Virginia and Piscataway and Fort Washington National Parks in Maryland where there is no distinct dry season, summers are hot, and winters are mild [74,75]. Mean annual precipitation at these Parks was approximately 45 inches (115 cm), and mean annual temperature was approximate 56 °F (13 °C) [74]. At Evansburg State Park in Pennsylvania, annual rainfall averaged 40 inches (103 cm) and annual temperatures averaged 51 °F (10 °C) [48]. Wineberry occurred in Baltimore City, Maryland, where average annual precipitation was 42 inches (1060 mm) [29].
Immediate fire effect on plant: Like some other blackberries, wineberry is probably top-killed by fire, while some portion of the roots and rhizomes are likely to remain unharmed and enable wineberry to sprout after fire. Depth of wineberry's regenerative structures within the soil profile has not been reported as of this writing (2009), but regenerative structures of other blackberries occur within the mineral soil where they would "probably survive fire" [21]. In Elk Island National Park, Alberta, the root system of red raspberry growing in a trembling aspen-balsam poplar (Populus tremuloides-Populus balsamifera) forest appeared to be well protected from the damaging effects of heat. In this study, red raspberry was experimentally subjected to 5 levels of fire severity by adjusting fuel load (range: 0-9.65 kg/m²) such that flame lengths ranged from 1.6 to 8.2 feet (0.5-2.5 m), frontal fire intensity ranged from 57 to 1905 kW/m, and residence time ranged from 1.5 to 10 minutes. Other characteristics of the fire are provided in [45]. In this study, all red raspberry canes and foliage were extremely susceptible to fire-induced mortality and were partially or completely killed at all fire severity levels; no aboveground biomass remained with fuel loadings >3.94 kg/m². Mortality of underground regenerative structures occurred only in areas of relatively high surface fuel loading (>3.9 kg/m²). At these sites, tissue mortality extended as far as 0.4 to 1.2 inches (1-3 cm) below the duff surface, but red raspberry rhizomes extended as deep as 2.0 inches (5 cm) below the duff surface so many rhizomes were protected. Sprouting occurred from the more deeply buried rhizomes that survived the fire. High duff moisture content (120%) likely contributed to protection of underground structures in this study [45].
Under certain environmental conditions, seeds of some blackberries may be protected from fire. Although no studies have been conducted on wineberry seeds, blackberry seeds subjected to a simulated prescribed summer burn in southeastern Arkansas were likely to remain unharmed by fire when protected by soil but unlikely to survive if they were located within the portion of the litter layer consumed by fire. Air-dried blackberry seeds of unspecified species were placed at 3 depths in a reconstructed forest floor within a loblolly pine (Pinus taeda) forest and subjected to fire. Mean fireline intensity was 6.4 Btu/ft-sec and rate of spread was 3.2 feet/minute. Fire consumed all of the litter (L) and upper fermentation (upper-F) layers and a portion of the lower fermentation (lower-F) layer. Other characteristics of the fire are provided in [10]. Postfire seed viability was assessed by germinating seeds in a greenhouse. Germination rates of seeds from the L layer (0.03%) and the upper-F/lower-F interface (0.33%) were low, and seeds tended to be charred. Germination rates of seeds from the lower-F/mineral soil interface were significantly higher (23.43%) than at the upper 2 layers (P<0.01) and did not differ from germination rates of seeds from unburned control plots. These results suggest that survival of blackberry seeds increases as depth of burial in the soil profile increases. The authors caution that fresh blackberry seeds or those consumed by animals may have a different response to fire than the air-dried seeds used in this study [10].
Impacts: The range of wineberry has expanded considerably since its introduction in the 1890s (see General Distribution). Despite its long history in North America, Innis [42] commented that it was not until the 1970s that it became a problem in Maryland. In Inwood Hill Park, Manhattan, New York, populations of wineberry, as well as 14 other nonnative invasive species were said to be expanding as of 2008 and wineberry was described as a "problem species" there [20]. Currently, wineberry is considered invasive in the Appalachian Mountain and Coastal regions of the east-central United States ([73], a fact sheet).
Where infestations are dense, wineberry is capable of limiting regeneration of forests, pastures, and croplands [42,80]. Wineberry is considered a threat to native flora in parts of the eastern United States largely because of its rapid growth, which allows it to crowd out native plants and establish extensive patches. In field experiments in Maryland, fewer individuals (P=0.040) and fewer ramets/m² (P=0.034) of nonnative Indian strawberry (Duchesnea indica) in plots with wineberry than without suggested that wineberry excluded Indian strawberry from the understory. There was no difference in Indian strawberry density in plots with or without native sawtooth blackberry [42].
Wineberry may occur at higher densities than its native congenerics. For example, in Inwood Hill Park in southern New York, wineberry was consistently recorded at higher densities than Allegheny blackberry or black raspberry where these species were found together [20]:
Density/ha of wineberry and 2 native blackberries in 3 forest site types in Inwood Hill Park, New York [20] Forest site type Species East ridge and slopes East and west ridgetops West ridge and slopes Allegheny blackberry 38 0 117 black raspberry 0 113 211 wineberry 469 469 972Wineberry's growth habit may contribute to its establishment and spread. Wineberry may form longer and stouter canes than some native raspberries, such as red raspberry (e.g., [18,41]). Comparison of wineberry growth and that of 9 other blackberries in field experiments in Japan found that wineberry produced the longest primocanes. Wineberry produced the 3rd largest diameter primocane and the 5th largest number of floricanes [41]:
Growth of field-planted wineberry, red raspberry, and black raspberry in Japan [41] Species Primocane length (cm) Primocane diameter (mm) Number of floricanes wineberry 370.4 22.1 17.7 red raspberry 272.7 15.6 25.0 black raspberry 309.0 21.5 4.0Wineberry's physiological efficiency may enhance its establishment and spread. Wineberry exhibited a higher ratio of maximum photosynthetic rates to dark respiration (P=0.10), higher leaf nitrogen concentration (P=0.02), and higher specific leaf area (P<0.01) than native sawtooth blackberry in the coastal plain region of Maryland. These results indicated a greater rate of leaf-level photosynthesis and higher resource use efficiency in wineberry than sawtooth blackberry. The manner in which these characteristics varied across habitats indicated greater phenotypic plasticity in wineberry relative to sawtooth blackberry. High phenotypic plasticity, low tissue costs, ability to utilize high resource levels for rapid growth, and high seed production may partially explain wineberry's ability to be an "aggressive" invader in some areas [42].
Control: Wineberry may be controlled through mechanical and chemical means [80]. In all cases where invasive species are targeted for control, no matter what method is employed, the potential for other invasive species to fill their void must be considered [8]. For example, removal of nonnative Norway maple (Acer platanoides) from the canopy of an even-aged sugar maple-Norway maple forest in New Jersey resulted in the establishment of wineberry and other nonnative species including tree-of-heaven, Japanese barberry (Berberis thunbergii), winged burning bush (Euonymous alata), Japanese honeysuckle, and black locust 2 years after treatment; it was unclear whether these species established from the seed bank or from off-site sources [88]. Wineberry and other nonnative invasive species including tree-of-heaven and oriental bittersweet invaded large, herbicide-treated areas on the western ridge of Inwood Hill Park, New York 3 years after invasive species control efforts were abandoned [20]. These examples underscore the importance of long-term maintenance and monitoring of treatment areas to restore native communities and reduce nonnative species in the long term. Control efforts that keep disturbed areas small and native plants available to colonize openings may help prevent the establishment and spread of wineberry and other nonnative species [88]. Ultimately, management of biotic invasions is most effective when it employs a long-term, ecosystem-wide strategy rather than a tactical approach focused on battling individual invaders [55].
Fire: For information on the use of prescribed fire to control this species see Fire Management Considerations.
Prevention: It is commonly argued that the most cost-efficient and effective method of managing invasive species is to prevent their establishment and spread by maintaining "healthy" natural communities [55,72], for example, by avoiding road building in wildlands [83] and by conducting monitoring several times each year [44]. Managing to maintain the integrity of the native plant community and mitigate the factors enhancing ecosystem invasibility are likely to be more effective than managing solely to control the invader [37]. Weed prevention and control can be incorporated into many types of management plans including those for logging and site preparation, grazing allotments, recreation management, research projects, road building and maintenance, and fire management [84]. See the "Guide to noxious weed prevention practices" [84] for specific guidelines in preventing the spread of weed seeds and propagules under different management conditions.
Cultural control: No information is available on this topic.
Physical or mechanical control: Removal of plants by hand-pulling or use of a spading fork can be an effective means of controlling wineberry, especially if the soil is moist and the roots and any cane fragments are completely removed. Removal and destruction of branches with fruits is recommended to reduce the number of seeds in the seed bank ([73], a fact sheet).
Like other blackberries, wineberry is likely encouraged by practices such as mowing or deep cultivation; thus, these methods are not recommended for wineberry control, and are not usually appropriate for wildlands and natural areas. In general, mowing of raspberries stimulates sprouting and reduces interference from neighboring vegetation. Deep cultivation (6-9 inches (15-23 cm)) cuts the roots of existing blackberry plants and causes the formation of large numbers of "sucker" plants [12]. However, if mowing is conducted 2 to 3 times per season for 2 or more years, eradication may be accomplished by exhausting the plant's carbohydrate reserves [86].
Biological control: Numerous diseases and insects affect wineberry, including wineberry latent virus. See Ellis and others [17] for a review.
Chemical control: A review states that wineberry can be controlled with a systemic herbicide like glyphosate or triclopyr [80]. Herbicides may be effective in gaining initial control of a new invasion or a severe infestation, but they are rarely a complete or long-term solution to weed management [9]. Herbicides are more effective on large infestations when incorporated into long-term management plans that include replacement of weeds with desirable species, careful land use management, and prevention of new infestations. Control with herbicides is temporary, as it does not change conditions that allow infestations to occur [97]. See the Weed Control Methods Handbook for considerations on the use of herbicides in natural areas and detailed information on specific chemicals.
Integrated management: Increased effectiveness generally occurs when multiple approaches are combined to control an invasive species. For wineberry, mowing or cutting prior to herbicide application may be more effective than either method alone [80]. Integrated management should include considerations of not only killing the target plant but also of establishing desirable species and maintaining weed-free systems over the long term.
Wineberry may be of limited importance to domestic livestock, but the fruit, foliage, and stems of wineberry provide food and cover for many wildlife species.
Palatability/nutritional value: Wineberry produces fruits that are readily consumed by birds, reptiles, and mammals ([73], a fact sheet). Wineberry was documented in fecal droppings of white-tailed deer in southern Connecticut [90] and was considered a preferred food of box turtles in the laboratory [6]. Although not reported for wineberry specifically, fruits of raspberry are eaten by many eastern birds including ruffed grouse, American woodcock, ring-necked pheasant, northern bobwhite, wild turkey, gray catbird, northern cardinal, brown thrasher, American robin, thrushes, and towhees. Mammals such as coyote, raccoon, black bear, white-tailed deer, common opossum, squirrels, chipmunks, skunks, and foxes also eat the fruits of raspberries [12,30,57,86].
Palatability of wineberry browse has not been determined. According to a review, raspberries generally have little forage value for domestic livestock [86]. However, stem densities and heights of blackberries (red raspberry, black raspberry, and Allegheny blackberry) in paddocks grazed by cattle in white oak-bur oak woodlands in southwestern Wisconsin were significantly lower than in ungrazed paddocks (P<0.03 for all variables), suggesting that blackberries were a preferred forage species there [35]. Forage value of raspberry fruit and browse to wildlife apparently varies among species [86]. Deer and rabbits eat the foliage and stems of raspberries, and porcupine and beaver occasionally consume the buds, twigs, or cambium of raspberries [12,57,86].
Cover value: According to reviews, many species of birds and mammals use the brambles of raspberries for protective cover and nesting [12,57,73]. Veery frequently placed nests on or near wineberry plants in mixed-hardwood forest in the middle-Atlantic Piedmont forest physiographic province in New Castle County, Delaware [36]. Similarly, crow tits nested in the brambles of wineberry along streambanks in wineberry's native range in South Korea [49].
Growth and development of wineberry is typical of blackberries. Wineberry produces biennial canes from a perennial root system or from underground rhizomes (see Vegetative regeneration) [42]. First year canes (primocane) are unbranched, sterile, entirely vegetative, and develop from rhizominous buds at or below the ground surface [25,42]. In the 1st year, carbon allocation is primarily into leaf production and cane elongation [42]. In the 2nd year, lateral branches develop in the axils of the primocanes and produce leaves, flowers, and fruits [42], but "do not have extensive growth" [25]. Second-year canes are referred to as "floricanes". Unlike primocanes, floricanes are woody [25].
In April, floricanes produce new leaves. In early May, new primocanes originate from the perennial root system [42]. In late May, floricanes undergo lateral branching and may produce flowers and fruit; fruit production occurs in late June to August. Fruits of wineberry ripen together [17]. After producing fruit in late summer, the leaves of floricanes senesce and the cane gradually dies. In Maryland, wineberry loses its leaves in late November [42]. Generalized fruiting and flowering dates are as follows:
Location Flowering Fruiting Maryland late May-early June late June-July [42] Arkansas May-June June-July [39] New England --- 16 July-30 August [70] New York 20 May-31 May [16] --- North Carolina --- July-October, peak in August [27]Pollination and breeding system: Wineberry flowers are hermaphroditic and pollinated by insects [64]. In field experiments in mixed-hardwood forest in Maryland, wineberry was self-compatible and less dependent upon cross-pollination by pollinators to set fruit than a coexisting native congener, sawtooth blackberry, suggesting that "wineberry could more easily establish itself in habitats with low pollinator service or a lack of mates" than sawtooth blackberry [42].
Seed production: A review of blackberries states that good seed crops occur nearly every year and that environmental factors affect the amount of flowering and fruit production in the genus (see Climate) [98]. As of this writing (2009), little information was available on seed production in wineberry but according to Swearington and others [80], wineberry is capable of producing fruits in "great abundance". Wineberry may not fruit until 3 years of age or more [25]. For example, in Pisgah National Forest, North Carolina, raspberries, including wineberry, did not produce fruit until 3 and 4 years after silvicultural treatments in upland hardwood and cove hardwood forest, respectively; it is unclear whether plants in this study established from seed or by sprouting. Upland hardwood forest was dominated by scarlet oak, chestnut oak, and black oak (Quercus velutina) and cove hardwood forest was dominated by yellow-poplar and northern red oak [27].
The number of seeds per fruit in wineberry ranges from 30 to 60 [6,42]. In mixed-hardwood forest in Maryland, the number of wineberry seeds per fruit and the number of fruits per plant were typically greater than those of sawtooth blackberry, a coexisting native congener. In addition, "local frugivores" consumed more wineberry fruits than sawtooth blackberry fruits (P<0.001). Wineberry fruits ripen together, are more abundant, and are displayed in tighter drupelets than fruits of sawtooth blackberry; this may partially explain preference for wineberry by frugivores in this study. These data suggest that seeds of wineberry may be more readily produced and more readily dispersed than those of native sawtooth blackberry [42], which may have important implications for the establishment and spread of wineberry in native communities.
Seed dispersal: Birds, reptiles, and mammals may contribute to the establishment and spread of wineberry by dispersing and scarifying seeds. Examination of fecal droppings of box turtles in the laboratory [6] and white-tailed deer in oak (Quercus spp.)-sugar maple-yellow- poplar-sweetbirch-American beech forest in southern Connecticut [90] suggest that these species may disperse viable wineberry seeds.
A review suggests that the action of avian gizzards and exposure to mammalian digestive acids may scarify and thus enhance germination of blackberry seeds [30]. However, the importance of ingestion to wineberry germination is unclear. Germination of box turtle-ingested and non-ingested wineberry seeds were similarly low (<10%), suggesting that wineberry seeds were not scarified by box turtle ingestion [6].
Seed banking: As of 2009, little information was available on seed banking of wineberry. Seeds of wineberry are dormant at maturity [63,96] and apparently long-lived [11]. Raspberries are capable of amassing large numbers of seeds in the seed bank that are capable of persisting for 100 years or more (see FEIS review for red raspberry). Seeds of blackberry that were cold-stratified and dry-stored for 22 to 26 years had germination rates as high as 84% in the laboratory; wineberry germination rates in this study were 8%. These data suggest that under some conditions some proportion of wineberry seeds may persist in the seed bank [11]; however, it is unclear how often conditions suitable for long-term storage of wineberry seeds in the seed bank are met in nature.
Germination: Raspberry seeds have a dormant embryo and a hard endocarp that inhibits germination [63,96]. Seeds of wineberry must be scarified and/or stratified for long periods (3-4 months) at cold temperatures (36-41 °F (2-5 °C)) for germination to occur [63]. Scarification using sulfuric acid is frequently performed in experimental studies to stimulate germination of wineberry seeds (e.g., [11]). Several studies provide reviews of treatments used to improve overall germination and rate of germination in blackberries in the laboratory [43,63,98]. In nature, seeds of wineberry may be scarified by passing through an animal's digestive system (see Seed dispersal). Like many blackberries, wineberry germination and seedling establishment may be favored by exposed mineral soil and high light (see Successional Status) [25].
Seedling establishment and plant growth: As of this writing (2009), little information is available regarding wineberry seedling establishment and growth. What information is available suggests that while wineberry is able to persist for decades in shaded areas, best survival and growth are obtained in moderate to high light. In a greenhouse study, leaf relative growth rate of 1-year-old wineberry seedlings was higher in high (22% photosynthetic photo flux density (PPFD)) than in medium (12% PPFD) or low (5-5.5% PPFD) light treatments (P<0.05), although "growth was high regardless of light treatment". Conversely, growth of primocanes of 2-year-old wineberry seedlings was greatest in medium light followed by high light and low light treatments (P=0.006). These results suggest that although high light is best for wineberry establishment, once established, wineberry is able to grow in medium or even low light [25]. This has important implications for persistence of wineberry in plant communities over time. See Successional Status for additional information.
Vegetative regeneration: Wineberry reproduces clonally from underground rhizomes and by tip-rooting [39,42]. Tip-rooting occurs when arching canes touch the ground and adventitious roots form at the tip, giving rise to new ramets. Only canes ≥3.3 feet (1 m) tall tip-rooted in mixed-hardwood forest in Maryland; at this site, tip-rooting was the predominant form of vegetative reproduction and typically occurred in large tree-fall gaps with high light. Wineberry may not reach adequate size for tip-rooting until 3 years of age or more [25].
Wineberry may reproduce more by seed than by vegetative regeneration. A lack of asexual reproduction by wineberry in mixed-hardwood forest in Maryland was attributed to advanced age (>5 years) of the perennial root system and to "extreme precipitation years with drought conditions followed by heavy precipitation" [42].
SITE CHARACTERISTICS:
As of this writing (2009), little English-language literature is available on wineberry's native habitats. What information is available indicates that wineberry grows at low to medium elevations in montane valleys and along roadsides in China [94]. In Japan, wineberry occurs in lowland and mountainous regions in clearings associated with spruce (Picea spp.), fir (Abies spp.), and birch (Betula spp.) [82]. In South Korea, wineberry occurs at elevations ranging from 70 to 460 feet (20-140 m) along streambanks [49].
In the eastern United States, wineberry occupies a wide range of habitats including early- to midsuccessional forest, floodplain forest, herbaceous and shrub wetland, wet meadows, riparian corridors, old fields, open disturbed areas, burned areas, trailsides, roadsides, ditches, and vacant lots, as well as ecotones between these habitats [4,5,22,28,42,58,59,64,65,70,80,91,93].
According to reviews, wineberry prefers open, mesic conditions with rich soils but tolerates a wide range of soil types, textures, and pH values [15,22,73,87]. At Great Falls Park in northeastern Virginia, wineberry occurred on soils ranging from "relatively fertile", with basic pH, and silt loam to silty clay loam textures to dry, "extremely acidic, infertile" silty clay loams. At this site, wineberry occurred on very dry upper slopes and ridge crests with "high solar exposure and low moisture potential" as well as seasonally flooded swamps [74]. Wineberry was found in Sussex County, New Jersey on trails and roadsides where soils were thin and rocky though moist [4]. In Chittenden County, Vermont, wineberry established on a limy talus slope in the dense shade of northern whitecedar (Thuja occidentalis) [99]. Wineberry occurred relatively infrequently in sweetgum (Liquidambar styraciflua) -sycamore streambank habitat with sandy soils in Newton County, Arkansas; this site was regularly disturbed by spring and fall flooding and anthropogenic influences [81]. In New Jersey, wineberry occurred in constructed wetlands with coarse soil [53]. In Inwood Hill Park in New York, wineberry occurred on some sites with "deep soils" [54]. Wineberry occurred on wet, seasonally flooded and mesic soils at the Piscataway and Fort Washington National Parks in Maryland [75]. Along a 250-mile (402 km) reach of the New River Gorge in West Virginia, wineberry was found at a variety of sites including regularly flooded streambeds, riverside beach areas, and wooded upper beach areas with soils ranging from cobblestone and gravel to sand and mudflats. Additional sites occupied by wineberry in this study included rocky summits and cliff faces and woodlands with shallow and sandy soils [79].
According to reviews, wineberry tolerates a range of light levels, with light availability in suitable habitat ranging from full sun to partial shade [22,64,73,87]. Although established plants may persist in low light, wineberry germination and survival appear best in moderate to high light environments (see Seedling establishment and plant growth) [25]. In field experiments in mixed-hardwood forest in Maryland, wineberry seedling survival was significantly reduced under leaf litter (P<0.001); this was attributed to a lack of light and increased potential for root rot as a result of increased moisture levels [42]. Although wineberry tolerates a variety of light levels and soil conditions, like other blackberries, adequate soil moisture and light appear important for best growth and fruit production (see Successional Status) [12,64].Photograph courtesy of John M. Randall, The Nature Conservancy, Bugwood.org
Elevation/Topography: Wineberry occurs in lowlands and mountainous terrain on slopes ranging from 0% to 60%. At Evansburg State Park, wineberry occurred between 98 and 397 feet (30-121 m) elevation. Wineberry occurred at Fernow Experimental Forest in north-central West Virginia at elevations ranging from 1,749 to 3,648 feet (533-1,112 m) and slopes ranging from 10% to 60% [56]. In Inwood Hill Park in southern New York, wineberry occurred on sites with slopes >10% [54]. Wineberry occurred on sites where slopes averaged >30% in Great Falls Park in Virginia .
Climate: Wineberry is hardy to USDA hardiness zone 5, where average annual minimum temperatures are as low as -20 °F (-26 °C) [46,64]; although some damage may be caused to the plant at this temperature, the plants "usually recover well" (Davis 1990, as cited in [64]).
Precipitation may affect wineberry density. In Maryland, drought reduced the density of wineberry and sawtooth blackberry, with greater mortality in forest edge sites than in intact forest (P=0.034). The subsequent year's precipitation was average; although wineberry density increased during that year, its density did not return to pre-drought levels [42].
Wineberry occurred throughout Great Falls Park in Virginia and Piscataway and Fort Washington National Parks in Maryland where there is no distinct dry season, summers are hot, and winters are mild [74,75]. Mean annual precipitation at these Parks was approximately 45 inches (115 cm), and mean annual temperature was approximate 56 °F (13 °C) [74]. At Evansburg State Park in Pennsylvania, annual rainfall averaged 40 inches (103 cm) and annual temperatures averaged 51 °F (10 °C) [48]. Wineberry occurred in Baltimore City, Maryland, where average annual precipitation was 42 inches (1060 mm) [29].
SUCCESSIONAL STATUS:
According to reviews, blackberries in North America occur on a range of sites at all stages of succession, but the majority of blackberries are considered pioneers of open and disturbed habitats and are capable of invading and rapidly occupying burns, eroded areas, old fields, and logged areas [12,18,86,98]. A review states that dense stands of blackberries can prevent or greatly delay establishment of trees and other species (see Impacts and Control) [98].
Like many other blackberries, wineberry is generally considered a pioneer or early-successional species that flourishes after disturbance, often forming dense thickets and dominating sites ([73], a fact sheet). For example, in Inwood Hill Park in southern New York, wineberry dominated the understory of yellow-poplar-white oak-northern red oak forest [54]. Although wineberry frequently establishes after disturbance, stem density typically decreases over time as the canopy closes and shade increases. However, wineberry apparently tolerates shade and may persist in shaded environments for several decades after disturbance [12]. Wineberry's relatively high phenotypic plasticity (see Impacts) [42] may allow it to survive a wide range of environmental conditions and successional stages [68]. In its native Japan, wineberry cover ranged from 0.5% to 1.9% at 3 ski areas 7 to 20 years after clearcutting [82]. Wineberry was considered a "typical successional species in the more mesic sites of northern New Jersey"; at these sites, wineberry was a relatively common component of some upland forest stands that had been free of major disturbance for at least 60 years. Although present in shaded, undisturbed habitat, the authors considered the occurrence of wineberry at this site as "vegetative holdovers from earlier successional stages" and as "chance establishment in gaps formed by wind throw or other catastrophe" [15].
Treefall gaps and other local disturbances may play important roles in the establishment and persistence of wineberry. A field study at the Smithsonian Environmental Research Center in Maryland found wineberry ramets and seedlings occurred more frequently in 2-year-old, storm-created gaps than in random plots in 135-year-old ("old") forest dominated by yellow-poplar, oak, hickory, American beech, and sweetgum and 45-year-old ("young") forest dominated by yellow-poplar [26]:
Frequency (%) of wineberry ramets and seedlings in young and old forests in Maryland [26] Old forest Young forest Type Gaps (n=20) Random (n=19) Gaps (n=4) Random (n=5) Ramets 50 11 100 80 Seedlings 50 0 75 50Greater establishment of wineberry seedlings at sites with high light and exposed mineral soil (i.e., large gaps with uprooted trees) indicates that disturbance may be important for seedling establishment. In old forest gaps, density of wineberry ramets was 34 times greater and primocane length was 2 times greater in large gaps (size range: 290-939 m²) than in small gaps (size range: 38-200 m²). In addition, sexual and asexual reproduction were more common in large gaps than in small gaps. In old stands, fruits were present in 15% of large gaps but not in small gaps or random plots, and tip-rooting was most common in large gaps [25]. In young stands, fruits were found in 100% of all gaps and 20% of random plots, but tip-rooting was "extremely rare". Wineberry seedling density was 4 times greater in gaps associated with uprooted trees compared to gaps with "snapped" trees [26]. Once established, measures of survivorship indicated that wineberry individuals persisted despite canopy closure [25].
Treefall gaps appear less important for wineberry seedling establishment, vegetative reproduction, and fruiting in early than in late succession [25,26]. For example, in the young forest, seedling establishment and fruiting was not limited to gaps. Although wineberry ramets were more likely to occur at sites with high light and large gaps, ramets that occurred in low light were more likely to occur in the young forest than in the old forest. Greater proportion of bare mineral soil and fewer layers of leaf litter in the young forest compared to the old forest may partially explain seedling establishment and fruiting outside of gaps in the young forest. These data suggest that in young forest wineberry may establish and spread without canopy-opening disturbances [25].
Birds, reptiles, and mammals may contribute to the establishment and spread of wineberry by dispersing and scarifying seeds. Examination of fecal droppings of box turtles in the laboratory [6] and white-tailed deer in oak (Quercus spp.)-sugar maple-yellow- poplar-sweetbirch-American beech forest in southern Connecticut [90] suggest that these species may disperse viable wineberry seeds.
A review suggests that the action of avian gizzards and exposure to mammalian digestive acids may scarify and thus enhance germination of blackberry seeds [30]. However, the importance of ingestion to wineberry germination is unclear. Germination of box turtle-ingested and non-ingested wineberry seeds were similarly low (<10%), suggesting that wineberry seeds were not scarified by box turtle ingestion [6].
A review of blackberries states that good seed crops occur nearly every year and that environmental factors affect the amount of flowering and fruit production in the genus (see Climate) [98]. As of this writing (2009), little information was available on seed production in wineberry but according to Swearington and others [80], wineberry is capable of producing fruits in "great abundance". Wineberry may not fruit until 3 years of age or more [25]. For example, in Pisgah National Forest, North Carolina, raspberries, including wineberry, did not produce fruit until 3 and 4 years after silvicultural treatments in upland hardwood and cove hardwood forest, respectively; it is unclear whether plants in this study established from seed or by sprouting. Upland hardwood forest was dominated by scarlet oak, chestnut oak, and black oak (Quercus velutina) and cove hardwood forest was dominated by yellow-poplar and northern red oak [27].
The number of seeds per fruit in wineberry ranges from 30 to 60 [6,42]. In mixed-hardwood forest in Maryland, the number of wineberry seeds per fruit and the number of fruits per plant were typically greater than those of sawtooth blackberry, a coexisting native congener. In addition, "local frugivores" consumed more wineberry fruits than sawtooth blackberry fruits (P<0.001). Wineberry fruits ripen together, are more abundant, and are displayed in tighter drupelets than fruits of sawtooth blackberry; this may partially explain preference for wineberry by frugivores in this study. These data suggest that seeds of wineberry may be more readily produced and more readily dispersed than those of native sawtooth blackberry [42], which may have important implications for the establishment and spread of wineberry in native communities.
The accepted scientific name of wineberry is Rubus phoenicolasius Maxim.
(Rosaceae) [47]. Wineberry is in the subgenus Idaeobatus, which
are raspberries in which the ripe fruit separates from the receptacle (Focke 1914, cited in [91]).
Hybridization within the Rubus genus occurs within and between subgenera [2].
Although natural hybrids between wineberry and native Rubus species have not
been reported as of this writing (2009), wineberry has been intentionally crossed with
red raspberry (R. idaeus) and black raspberry (R. occidentalis) in
breeding programs [14,38].
In this review, "blackberry" refers to species in the genus Rubus
and "raspberry" refers to species in the subgenus Idaeobatus.
Vegetative regeneration: Wineberry reproduces clonally from underground rhizomes and by tip-rooting [39,42]. Tip-rooting occurs when arching canes touch the ground and adventitious roots form at the tip, giving rise to new ramets. Only canes ≥3.3 feet (1 m) tall tip-rooted in mixed-hardwood forest in Maryland; at this site, tip-rooting was the predominant form of vegetative reproduction and typically occurred in large tree-fall gaps with high light. Wineberry may not reach adequate size for tip-rooting until 3 years of age or more [25].
Wineberry may reproduce more by seed than by vegetative regeneration. A lack of asexual reproduction by wineberry in mixed-hardwood forest in Maryland was attributed to advanced age (>5 years) of the perennial root system and to "extreme precipitation years with drought conditions followed by heavy precipitation" [42].
Rubus phoenicolasius (Japanese wineberry,[2] wine raspberry,[3] wineberry or dewberry) is an Asian species of raspberry (Rubus subgenus Idaeobatus) in the rose family, native to China, Japan, and Korea.[4]
The species was introduced to Europe and North America as an ornamental plant and for its potential in breeding hybrid raspberries. It has subsequently escaped from cultivation and become naturalized in parts of Europe and North America.[5][6][7]
The species is a perennial plant which bears biennial stems ("canes") from the perennial root system. In its first year, a new stem ("primocane") grows vigorously to its full height of 1–3 m, unbranched, and bearing large pinnate leaves with three or five leaflets; normally it does not produce any flowers the first year. In its second year, the stem ("floricane") does not grow taller, but produces several side shoots, which bear smaller leaves always with three leaflets; the leaves are white underneath.[4]
The flowers are produced in late spring on short, very bristly racemes on the tips of these side shoots, each flower 6–10 mm diameter with five purplish red to pink petals and a bristly calyx. The fruit is orange or red, about 1 cm diameter, edible, produced in summer or early autumn; in botanical terminology, it is not a berry at all, but an aggregate fruit of numerous drupelets around a central core. Ripening occurs from early summer.[4][5] The canes have red glandular hairs. These red hairs give the species its scientific name, from the Latin phoenicus, meaning red.[4]
In addition to seed propagation, new plants are formed from the tips of existing canes touching the ground. They enjoy moist soil and grow near and within wooded areas.[4]
As a fruit develops, it is surrounded by a protective calyx covered in hairs that exude tiny drops of sticky fluid. While the structure resembles those of carnivorous plants, the wineberry plant does not get nutrients from insects caught in the sap: the sticky mucilage contains no digestive enzymes, surrounding tissues cannot absorb nutrients, and there are no protein-storage tissues. Also, unlike carnivorous plants, wineberry grows in nutrient-rich soil, so it need not resort to insect proteins as a source of nitrogen.[4][8]
The plant's leaves and stems/branches are covered in spines. The leaves appear in sets of three, where the two side leaves are small, and the center leaf is large. The leaves are green on top, and white on the bottom because of a dense layer of woolly hairs.[4]
Wineberries grow in the wild in many parts of the United States, primarily the Appalachian Mountains.[9][10] They are common along the edges of fields and roadsides, and still are used as breeding stock for raspberry cultivars.[11] They are edible, with no poisonous look-a-likes in North America.[12] Other plants that may be mistaken as wineberries include red raspberry, salmonberry, black raspberry, and blackberry, all of which are edible. Sweet and tart with a raspberry-like flavor, wineberries are used similarly to raspberries to make pastries, such as pie or other sweet treats.[12]
The wineberry is native to eastern Asia and has been introduced into parts of North America and Europe. Wineberries grow vigorously and can form extensive, dense thickets that displace many native species.[13][14] Wineberries can be found in many habitats, such as forest, fields, stream banks, and wetland edges, as well as open woods.[11] The first years of growth for wineberries are longer (approximately 32%) than many other black and red raspberries.[13] Wineberries are a host to several viruses, such as raspberry yellow spot that can affect native species of raspberry.[15]
Wineberry vine growth may be controlled variously, such as by biological mechanisms increasing susceptibility to diseases, or mechanical effects by digging, excavating or repeatedly cutting back the plant until it dies.[13] Using a systemic herbicide, such as glyphosate, it can be controlled chemically.[14] Concentrations mixed to the manufacturers specifications can be applied to the leaves to kill the plants.[13] The wineberry is listed as a noxious weed by both Connecticut and New York where its possession and sale are prohibited by law.[16]
Rubus phoenicolasius (Japanese wineberry, wine raspberry, wineberry or dewberry) is an Asian species of raspberry (Rubus subgenus Idaeobatus) in the rose family, native to China, Japan, and Korea.
The species was introduced to Europe and North America as an ornamental plant and for its potential in breeding hybrid raspberries. It has subsequently escaped from cultivation and become naturalized in parts of Europe and North America.
