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Introductory

SPECIES: Rubus idaeus | Red Raspberry
ABBREVIATION : RUBIDA SYNONYMS : R. idaeus ssp. sachalinensis (Levl.) Focke = R. i. ssp. strigosus SCS PLANT CODE : RUID RUIDI RUIDS2 COMMON NAMES : red raspberry raspberry black-haired red raspberry smoothleaf red raspberry American red raspberry wild raspberry brilliant red raspberry wild red raspberry TAXONOMY : The currently accepted scientific name of red raspberry is Rubus idaeus L. There are two subspecies [53]: R. i. ssp. idaeus R. i. ssp. strigosus (Michx.) Focke Numerous red raspberry hybrids have been reported, although many are infertile [43,104]. This shrub hybridizes with many species in the Rubus genus including R. arcticus, R. ursinus, R. occidentalis, R. rubrisetus, and R. odoratus [49,55,68,104]. Red raspberry has hybridized with thimbleberry (R. parviflorus) in the laboratory [49]. LIFE FORM : Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : D. Tirmenstein, December 1989. LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Tirmenstein, D. 1990. Rubus idaeus. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Rubus idaeus | Red Raspberry
GENERAL DISTRIBUTION : Red raspberry occurs throughout most of the temperate regions of the world [20]. In North America it grows from Alaska through Canada to Newfoundland, southward to North Carolina and Tennessee in the East, and to Arizona, California, and northern Mexico in the West [36,93,98]. The native North American red raspberry is Rubus idaeus ssp. strigosus [36]. R. i. ssp. idaeus grows across northern Europe to northwestern Asia [36]. It is cultivated in Hawaii [109] and throughout much of North America and has naturalized in many locations [36]. ECOSYSTEMS : FRES10 White - red - jack pine FRES11 Spruce - fir FRES15 Oak - hickory FRES17 Elm - ash - cottonwood FRES18 Maple - beech - birch FRES19 Aspen - birch FRES20 Douglas-fir FRES21 Ponderosa pine FRES22 Western white pine FRES23 Fir - spruce FRES24 Hemlock - Sitka spruce FRES25 Larch FRES26 Lodgepole pine FRES28 Western hardwoods FRES34 Chaparral - mountain shrub FRES35 Pinyon - juniper FRES37 Mountain meadows FRES38 Plains grasslands FRES39 Prairie FRES44 Alpine STATES : AK AL AZ AR CA CO CT DE GA HI ID IL IN IA KY ME MD MA MI MN MO MT NV NH NJ NM NY NC ND OH OR PA RI SC SD TN UT VA WA WV WI WY AB BC MB NB NF ON PQ SK MEXICO ADMINISTRATIVE UNITS : ACAD BLCA BRCA CACO CEBR CRMO DEWA GLAC GRTE GRSA ISRO LACL MORU NOCA ROMO THRO WICA YELL BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains 3 Southern Pacific Border 5 Columbia Plateau 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 KUCHLER PLANT ASSOCIATIONS : K001 Spruce - cedar - hemlock forest K002 Cedar - hemlock - Douglas-fir forest K004 Fir - hemlock forest K011 Western ponderosa forest K012 Douglas-fir 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 K020 Spruce - fir - Douglas-fir forest K021 Southwestern spruce - fir forest K023 Juniper - pinyon woodland K025 Alder - ash forest K037 Mountain mahogany - oak scrub K052 Alpine meadows and barren K064 Grama - needlegrass - wheatgrass K067 Wheatgrass - bluestem - needlegrass K074 Bluestem prairie K081 Oak savanna K093 Great Lakes spruce - fir forest K095 Great Lakes pine forest K096 Northeastern spruce - fir forest K097 Southeastern spruce - fir forest K098 Northern floodplain forest K102 Beech - maple forest K106 Northern hardwoods K107 Northern hardwoods - fir forest K108 Northern hardwoods - spruce forest SAF COVER TYPES : 1 Jack pine 5 Balsam fir 12 Black spruce 13 Black spruce - tamarack 15 Red pine 16 Aspen 17 Pin cherry 18 Paper birch 21 Eastern white pine 22 White pine - hemlock 25 Sugar maple - beech - yellow birch 30 Red spruce - yellow birch 31 Red spruce - sugar maple - beech 32 Red spruce 33 Red spruce - balsam fir 35 Paper birch - red spruce - balsam fir 37 Northern white cedar 39 Black ash - American elm - red maple 42 Bur oak 60 Beech - sugar maple 107 White spruce 108 Red maple 109 Hawthorn 201 White spruce 202 White spruce - paper birch 204 Black spruce 210 Interior Douglas-fir 211 White fir 212 Western larch 213 Grand fir 215 Western white pine 217 Aspen 218 Lodgepole 222 Black cottonwood - willow 224 Western hemlock 226 Coastal true fir - hemlock 227 Western redcedar - western hemlock 228 Western redcedar 229 Pacific Douglas-fir 230 Douglas-fir - western hemlock 235 Cottonwood - willow 236 Bur oak 237 Interior ponderosa pine 239 Pinyon - juniper 252 Paper birch 253 Black spruce - white spruce 254 Black spruce - paper birch SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Red raspberry is well represented in many plant communities throughout North America. It grows within the understory of many quaking aspen (Populus tremuloides), mixed conifer, cottonwood (Populus spp.), cedar (Thuga spp.)-hemlock (Tsuga spp.), ponderosa pine (Pinus ponderosa), spruce (Picea spp.)-fir (Abies spp.), and Douglas-fir (Pseudotsuga menziesii) forests of the West [6,23,37]. In the Lake States and Northeast, red raspberry frequently grows in oldfield communities, or in association with jack pine (Pinus banksiana), white spruce (Picea glauca), black spruce (P. mariana), red spruce (P. rubens), Atlantic white-cedar (Thuja occidentalis), balsam fir (Abies balsamea), aspen (Populus spp.), beech (Fagus spp.), maple (Acer spp.), red pine (Pinus resinosa), and eastern white pine (P. strobus) [1,3,29,30,34,41,40,77]. It is a common component of northern hardwood forests and often assumes dominance on sites which have been subject to windthrow, fire, or timber harvest [100]. Red raspberry is a prominent component of many taiga communities in Alaska [25] and the Canadian North. Associated species: Red raspberry grows with a wide variety of plants across its extensive geographic range. The following species are particularly common plant associates [1,40,12,13,95]: Canada beadruby (Maianthemum canadense), thimbleberry, bunchberry (Cornus canadensis), huckleberry (Vaccinium spp.), fireweed (Epilobium angustifolium), bluejoint reedgrass (Calamagrostis canadensis), kinnikinnick (Arctostaphylos uva-ursi), Virginia strawberry (Fragaria virginiana), green alder (Alnus virida ssp. crispa), twinflower (Linnaea borealis), sedges (Carex spp.), prickly rose (Rosa acicularis), twinberry (Lonicera spp.), lowbush blueberry (Vaccinium angustifolium), bog Labrador tea (Ledum groenlandica), red currant (Ribes triste), highbush cranberry (Viburnum edule), and red-osier dogwood (Cornus sericea). Red raspberry occurs as a dominant in a number of plant communities. It has been included as a codominant in rocky, high elevation alpine scree communities with Colorado columbine (Aquilegia coerulea) and littleflower alumroot (Heuchera parvifolia). Red raspberry has been listed as an indicator or dominant member of a plant community in the following publications: Plant associations of Region Two: Potential plant communities of Wyoming, South Dakota, Nebraska, Colorado, and Kansas [51] Habitat types on selected parts of the Gunnison and Uncompahgre National Forests [56]

VALUE AND USE

SPECIES: Rubus idaeus | Red Raspberry
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Raspberries provide food and cover for a wide range of wildlife species [10,100]. Some herbivores browse raspberry, but in general, it offers relatively poor forage. Red raspberry is browsed by moose in Alaska but is not considered to be of primary importance [79]. In some locations, deer, rabbits, mountain beaver, and elk eat the foliage of raspberries [14,91]. Porcupine and beaver occasionally consume buds, twigs, or cambium of species within the genus Rubus [91]. However, thorns generally prevent excessive wildlife use of red raspberry [95]. In general, raspberries have little forage value for domestic livestock [91]. Fruits of many species within the genus Rubus are eaten by ruffed grouse, blue grouse, sharp-tailed grouse, ring-necked pheasant, greater prairie chicken, California quail, northern bobwhite, gray catbird, northern cardinal, yellow-breasted chat, American robin, thrushes, towhees, brown thrasher, orchard oriole, summer tanager, pine grosbeak, gray (Hungarian) partridge, and band-tailed pigeon [14,91]. Mammals such as the coyote, raccoon, black bear, common opossum, squirrels, Townsend's chipmunk, skunks, red fox, and gray fox also seek out the fruits of many raspberries [14,91]. The eastern chipmunk, western chipmunk, deer mice, and grizzly bear consume red raspberry fruit where available [59,105]. Flowers of red raspberry provide nutritious food for bees [40]. PALATABILITY : Red raspberry browse appears to be relatively unpalatable to most ungulates. However, the fruits are highly palatable to many birds and mammals. The degree of use shown by livestock and wildlife species for red raspberry is rated as follows [23]: CO MT ND UT WY Cattle poor poor poor fair poor Sheep poor fair fair good fair Horses poor poor poor poor poor Pronghorn poor ---- ---- poor poor Elk ---- poor ---- fair fair Mule deer ---- fair ---- good fair White-tailed deer fair ---- ---- ---- ---- Small mammals good ---- ---- good fair Small nongame birds poor ---- ---- good fair Upland game birds ---- ---- ---- good fair Waterfowl ---- ---- ---- poor poor NUTRITIONAL VALUE : Browse: Red raspberry browse is rated as poor in both energy and protein value [23]. Nitrogen, phosphorus, and potassium concentrations are highest in young leaves but decrease as leaves mature [46]. Conversely, calcium and magnesium concentrations are generally highest in mature leaves but lowest in young, developing leaves [46]. Zinc typically increases through the growing season whereas manganese decreases [46]. Levels of nitrogen, phosphorus, potassium, and calcium generally decline as the growing season progresses but may increase in the fall if additional rainfall allows plants to resume growth [46]. Fruit: Raspberry fruits are sweet and contain relatively high amounts of both mono and disaccharides [88]. Relative glucose, starch, and sugar content has been documented for a number of red raspberry cultivars [16]. COVER VALUE : Dense red raspberry thickets serve as favorable nesting habitat for many small birds [14]. Small mammals such as rabbits and squirrels also find shelter in raspberry thickets [91]. The degree to which red raspberry provides environmental protection during one or more seasons is rated as follows [23]: CO UT WY Pronghorn ---- poor poor Elk ---- poor poor Mule deer ---- poor poor White-tailed deer ---- ---- poor Small mammals fair fair fair Small nongame birds ---- fair fair Upland game birds ---- good fair Waterfowl ---- poor poor VALUE FOR REHABILITATION OF DISTURBED SITES : Some ecotypes of red raspberry have value in reclamation [95]. Suitable ecotypes are rated as having low to moderate value for short-term revegetation, and at least moderate value for long-term revegetation projects [23]. Red raspberry exhibits good potential for erosion control on some sites [10,91,95]. It has been successfully used to stabilize roadcuts and other disturbed sites in Utah and to revegetate bare soils in subalpine zones of Colorado [95]. Red raspberry is recommended for revegetation projects on well-drained sites in interior Alaska where maximum spacing of 3.3 feet by 3.3 feet (1 meter x 1 meter) is suggested [95]. Natural seedling establishment has been observed on many types of harsh sites, such as on tailings and surface soil of oil sand extraction plants in northern Alberta [95]. Red raspberry is capable of establishing on acidic tailings which have been treated with lime and on tar sands [95]. Propagation: Red raspberry can be propagated through leaf bud cuttings, "rooted handles," stem cuttings, or root cuttings (suckers) [24,67,89,95]. Success of establishment through root cuttings varies according to the cultivar and planting date [89]. However, root cutting success has ranged up to 60 percent in experimental tests [89]. Correct choice of planting dates and techniques are important and significantly influence subsequent growth and establishment [14,89]. In vitro micropropagation techniques have also been developed for mass production of red raspberry [97]. Red raspberry seedlings may be transplanted, or seed may be sown directly onto disturbed sites. Seed which has been scarified can be successfully planted in the late summer or fall [10]. Cold treatment is not required for fall seedings. Previously stratified and scarified seed can be planted in the spring [10]. Good results have been obtained after seeds were planted with a drill and covered with 1/8 to 3/16 inch (0.3-0.5 cm) of soil. Cleaned seed averages approximately 328,000 per pound (722,467/kg) [10]. Detailed information is available on appropriate methods to obtain and plant red raspberry seed [95]. OTHER USES AND VALUES : The red raspberry was traditionally an important food of many Native American peoples. It was eaten fresh or preserved for winter use [66]. Approximately 0.27 quarts (250 ml) of wild red raspberry fruit can be hand-harvested within 30 minutes in good stands [66]. The fruit, bark of roots, and stems of raspberries have been used to make various medicinal preparations [10]. The unique edible fruit of the red raspberry is delicious fresh or preserved. Raspberries make excellent jams and jellies [93] and provide flavorful additions to pies and other baked goods, candies, and dairy products such as yogurt or ice cream. Raspberry tea is commercially available and good although mild in flavor. The raspberry industry in North America is a growing, multimillion dollar business [63]. Five primary regions produce most of the raspberries grown commercially in North America [63]: 1) Northeast-Atlantic Provinces: southern Quebec through Pennsylvania 2) Central Atlantic Region: Maryland to South Carolina, eastern Kentucky and Tennessee, northern Georgia and Alabama 3) Central Great Lakes Region: Michigan, southern Ontario, Indiana, Illinois, Ohio to Iowa, Missouri, western Kentucky and Tennessee 4) Prairie States Region: Minnesota, southern Manitoba, eastern North and South Dakota, and Wisconsin 5) Pacific Northwest: southern British Columbia, western Washington and Oregon Cultivars: Many cultivars have been developed to meet the needs of raspberry growers in a variety of climatic situations. Most are derived from the European subspecies idaeus [98]. Desirable traits for red raspberry cultivars include spinelessness, winter hardiness, high fruit yields, resistance to disease, perennial stems, and primocane (or autumn) fruiting [49]. Reviews of particular cultivars document the extreme plasticity of this species and consider the suitability of each to various geographic locations [20,20,22,69,83,67,63,97,16,48]. Cultivars exhibit great genetic variation in time of flowerbud initiation, number of drupelets produced per fruit, time of fruit ripening, amount and timing of root suckering, length of dormancy, winter hardiness, fruit yield, and disease resistance [19,22,48,67,69,71,83,92]. Consequently, care should be taken to select cultivars with desirable traits which would enhance suitability for growth in a specific location [20]. The commonly cultivated loganberry may have been derived from a red raspberry-trailing blackberry hybrid [17]. Commercial cultivation: A wide array of studies detail commercial propagation of the red raspberry. Traditional techniques include hill culture of canes (stems), removal of weeds, and elimination of intercane suckers to increase fruit yield [64]. Older and weaker canes may be mowed or otherwise pruned annually to improve yield, enhance access to fruit, and to maintain the general health of the cane [67,102]. Trends in red raspberry propagation include increasing mechanization [63]. Various cultivation techniques have been shown to improve fruit yields [16,65,72]. In some instances, application of nitrogen fertilizers can increase both cane growth and the number of flowers produced per node [67]. However, in other situations fertilizers appear to be of little benefit [33]. Following the addition of nitrogen fertilizer, Lawson and Waister [65] observed increased yields for two years, little effect during the third year, and decreased yields during the next two years. Similarly, irrigation appears to increase yields in some locations while having little effect elsewhere [72]. MANAGEMENT CONSIDERATIONS : Competition: Red raspberry typically increases dramatically after fire or timber harvest [27,39]. In many areas this shrub can compete vigorously with conifer seedlings for light, moisture, nutrients, and space [30,34,62,74]. Dense thickets of red raspberry reportedly suppress the growth of balsam fir (Abies balsamea) and spruce (Picea spp.) seedlings after spruce-fir forests of northern Maine are clearcut [30] and after timber harvest in the boreal forests of Ontario [82]. Raspberries also compete effectively with jack pine (Pinus banksiana) and red pine (P. resinosa) following timber harvest in northeastern Minnesota and Manitoba [3,74]. Graber and Thompson [34] observed that relatively few red raspberry seeds are present within the soil of northeastern hardwood forest harvested at 100-year intervals. However, in forests harvested at more frequent intervals, large numbers of red raspberry seed are present and massive simultaneous germination results in intense competition with conifer seedlings [34]. Chemical control: Red raspberry is susceptible to a number of herbicides [9]. Glyphosate is commonly used as a mid-to-late summer foliar spray [82]. A number of herbicides have been suggested for use in reducing weeds in cultivated red raspberry patches [8].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Rubus idaeus | Red Raspberry
GENERAL BOTANICAL CHARACTERISTICS : Red raspberry is a deciduous, erect or arching, thicket-forming shrub which grows from 1.6 to 9.8 feet (0.5-3 m) in height [36,80,86,93]. The total height and extent of growth is largely attributable to climatic factors [101]. Woody stems are bristly or prickly with shreddy, exfoliating yellow-brown bark [36,93]. Leaves are alternate and pinnately compound in leaflets of three to five [86,93]. Leaves are green and glabrous to hairy above but white or gray, hairy to glabrate and greenish beneath [98]. Small showy perfect white flowers are borne in clusters of one to four in a compound cyme [36,55,80,93]. Fruit of the red raspberry is made up of many to several, red or pinkish-purple drupelets [80,98]. Aggregates of drupelets are commonly referred to as a "berry." Red raspberry is made up of mostly biennial canes (stems) on a long-lived perennial rootstock [45,99]. This rootstock initially forms from seedling establishment or the production of root suckers [99], which gradually separate from parent plants as the connecting root tissue dies [45]. Sterile first-year stems, or primocanes, develop from buds at or below the ground surface and generally bear only leaves [36,99]. During the second year, lateral branches, known as floricanes, develop in the axils of the primocanes which produce both leaves and fruit [36,100]. A "typical" raspberry rootstock system is made up of at least one floricane and several primocanes [99]. It should be noted that some commercially grown strains of red raspberry are primocane-fruiting; that is, they are capable of bearing fruit during the fall of the first year of development [21,92]. Primocane-fruiting appears to be absent entirely or represents an atypical situation in native-growing populations of red raspberry. RAUNKIAER LIFE FORM : Hemicryptophyte REGENERATION PROCESSES : Red raspberry reproduces through seed and also regenerates vegetatively. It is capable of forming dense thickets through sprouting. Reproductive versatility is well represented in the Rubus genus, with sexual reproduction, parthenogenesis (development of the egg without fertilization), pseudogamy (a form of apomixis in which pollination is required), and parthenocarpy (production of fruit without fertilization) occurring widely [17]. The following types of reproduction have been documented within the genus: (1) sexual reproduction, (2) nonreduction at meiosis on the female, male, or both sides, (3) apomixis with segregation, (4) apomixis without segregation, and (5) haploid parthenogenesis [17]. These modes of asexual reproduction are important because they help contribute to the vigorous, aggressive spread of red raspberry. Red raspberry is capable of vigorous sprouting after disturbance [18] but also expands in clonal area through vegetative regeneration [95,100]. Natural vegetative regeneration occurs through root sprouts or "suckers" [95,100,101], "stolons" [95], "rhizomes" [39,52], and basal stem buds or rootcrowns [45,95,101]. The precise mode of vegetative regeneration depends on the type and severity of disturbance. Dense raspberry thickets form from the roots or stems of parent plants which separate to form individual plants with the deterioration of connecting tissue [45]. Red raspberry allocates most energy to vegetative regeneration on recently disturbed sites with favorable growing conditions [99]. With time, initially elevated nutrient levels decline, and shading increases. As growing conditions deteriorate, red raspberry shifts its reproductive effort to the production of large numbers of seed [39,100]. Red raspberry sprouts readily from portions of aboveground stems which survive disturbance [52]. Many raspberry species are capable of rooting from the stem nodes, and layering has been widely reported in the red raspberry [95]. This shrub is also capable of sprouting from axillary buds located "well above the ground level" [45]. Rootcrown or stembase sprouting is an important regenerative mode, which in the raspberry gives rise to biennial stems even in the absence of disturbance [36,45]. Red raspberry typically sprouts from the rootcrown if aerial foliage is cut late in the growing season [95]. In related species such as salmonberry (R. spectabilis), apical dominance exerted by extant rootcrowns inhibits sprouting from belowground structures such as roots or rhizomes [106]. Root "suckering" is a normal, on-going process in red raspberry stands [45,101]. However, particularly vigorous root suckering is often observed after the aboveground vegetation is damaged or destroyed. This shrub regenerates from buds located on the larger main roots as well as those present on lateral roots which are often located fairly close to the soil surface [52,101]. The mean depth of these underground regenerative structures (root buds) was estimated at 2.4 inches (6 cm) in a New Brunswick study [28]. During the first 2 to 3 years after establishment, root suckers fill in spatial gaps in the clone [100]. Root sucker mortality is generally high during the third and fourth years because of intense intraspecific competition for sunlight, space, and nutrients which result in "self thinning" of stands [45,100]. Suckering ability declines with age, with production decreasing from an average of 1.5 per square foot (16.0/sq m) in 3-year-old stands to 0.77 per square foot (8.25/sq m) in 4-year-old stands [100]. Although relatively few root suckers actually reach the canopy, survival rates of those that do is high [100]. Most root suckers live for only 1 or 2 months [100]. Several researchers report that red raspberry is capable of sprouting from rhizomes after fire or other disturbance [39,52]. However, others have observed that red raspberry lacks rhizomes with any regenerative capability [28]. The term "rhizome" may have been loosely applied to rhizomelike roots which do possess the ability to sprout. Geographic or genetic differences in red raspberry morphology and physiology are also possible. Seed: Immature fruit, commonly referred to as "berries," are pink and hard [10]. Ripe fruit is generally red, but less commonly white or yellow [43]. Several to many small individual drupelets form an aggregate fruit [10,98]. Fruit size appears to be related to soil moisture [72], although significant genotypic variation has also been noted in the size and number of fruits produced annually [22]. Decreased stored nutrient availability and water stress can influence overall fruit production [16]. It is estimated that 70 to 90 percent of red raspberry flowers eventually mature into fruit which results in an abundance of seed [99]. Whitney [100] observed that 77 percent of all plants flowered, with 85 percent of those flowering producing seed. Most species of raspberry produce good seed crops nearly every year [10], but seed production does vary annually in the red raspberry according to climatic factors and the age of the cane. Whitney [100] observed average seed production of 65 seeds per square foot (700 seeds/ sq m) in 2-year-old canes, with maximum production of 1,301 seeds per square foot (14,000 seeds/sq m) in 4-year-old canes. Annual seed production averaged 604 seeds per square foot (6,500 seeds/sq m) over a 4-year period [100]. Pollination: Red raspberry is primarily pollinated by bees, although flies and beetles also pollinate some flowers [40]. Under natural conditions, it is almost exclusively self-incompatible [55] which contributes to morphological variability. Germination: Seed of the red raspberry is relatively large [31], with viabilty averaging up to 92 to 99 percent in laboratory tests [95]. Red raspberry seeds have a hard, thick, impermeable coat and dormant embryo [10]. Seeds have the ability to become dormant a second time in response to environmental factors [50]. Consequently, germination is often slow. Most raspberry seeds require, as a minimum, warm stratification at 68 to 86 degrees F (20 to 30 degrees C) for 90 days, followed by cold stratification at 36 to 41 degrees F (2 to 5 degrees C) for an additional 90 days [10]. Cold stratification alone is insufficient to induce germination in red raspberry [59]. Laboratory tests indicate that exposure to sulfuric acid solutions or sodium hyperchlorite prior to cold stratification can improve germination [10,43,50,95]. Evidence suggests that the digestive enzymes of mammals can also enhance germination, with seeds eaten by chipmunks and deer mice exhibiting better germination than untreated seeds [59]. Sowing seeds at greater depths with subsequent exposure to light can produce better germination than shallow plantings, presumably because of greater soil moisture [50]. Results of specific germination tests have been documented in a number of studies [10,50,59]. Seedbanking: Red raspberry amasses large numbers of seed which persist in the soil until favorable germination conditions are encountered [31,35,100]. Often, many seeds remain buried in the soil of stands which lack any sign of the parent plants [31]. Red raspberry seed can remain viable for 60 to 100 years or more [62,73,100]. Seeds are less likely to germinate when fresh [50,62], and may reach maximum viability at 50 to 100 years of age [34]. In a New Hampshire study, approximately 90 percent of Rubus (R. idaeus and R. allegheniensis) seed germinated during the first summer after disturbance in 38-, 95-, and > 200-year-old stands, whereas only 60 percent of those in 5-year-old stands germinated [34]. More than 4,048,583 Rubus seeds per acre (10 million/ha) have been found in the soil of 5-year-old beech (Fagus spp.)-birch (Betula spp.)-maple (Acer spp.) stands [34]. Numbers declined to 48,588 per acre (120,000/ha) in 200-year-old stands [34]. Annual reductions in stored seed have been attributed to: (1) degeneration resulting in death, (2) fungi or animal predation, and (3) annual germination of some seeds. Fyles [31] reported 237 to 1,883 seeds per foot square (22-175/m sq) in organic soil and 0 to 2,582 per foot square (0-240 m sq) in mineral soil of upland coniferous forests of central Alberta. Distribution of germinating seeds by stand age in beech-birch-maple forests of New Hampshire were as follows [34]: stand age in years 5 38 95 200 + #seeds/m sq. 1,016 286 68 12 Seed dispersal: Red raspberry seed is readily dispersed by birds and mammals [87,100]. After they mature, the highly sought-after fruit rarely remains on the plants for long [10]. Birds have been observed to deposit 2,429 to 2,834 viable seeds per acre (6,000 to 7,000/ha) annually in beech-birch-maple forests of New Hampshire [34]. Mammals such as mice and chipmunks may be important dispersal agents in some areas [59]. Seedling establishment: Most seedlings germinate during the first year after disturbance [99,100] and produce stands which are primarily even aged. In many instances, as much as 70 to 90 percent of all individuals establish during the first year after disturbance [100]. Researchers have observed minimal recruitment in the second, third, and fourth years after fire [100]. Little seedling establishment occurs beneath the shade of a closed forest canopy [100]. SITE CHARACTERISTICS : Red raspberry grows across a wide range of sites throughout most of the world's temperate regions [20]. It commonly occurs in clearings or borders in boreal forests, in ravines, on bluffs and streambanks of prairie regions, and on talus or scree above timberline [39,86,93,95]. Soil: Raspberries are tolerant of a wide range of soil pH and texture but do require adequate soil moisture [14]. Red raspberry grows on imperfectly to well-drained sandy loam to silty clay loam, but best growth occurs on moderately well-drained soils [95]. Although red raspberry grows well on barren and infertile soils, it reportedly has a relatively high demand for soil nutrients and is most abundant on nutrient-rich soils [39]. This shrub is moderately tolerant of acidic soils [95]. Elevation: Generalized elevational ranges for selected locations are as follows [23,98]: from 6,500 to 11,700 feet (1,981 to 3,569 m) in CO 2,400 to 7,000 feet (732 to 2,134 m) in MT 5,500 to 9,600 feet (1,676 to 3,420 m) in UT 6,500 to 11,000 feet (1,981 to 3,355 m) in WY SUCCESSIONAL STATUS : Red raspberry vigorously invades and colonizes many types of disturbed sites [62,95,100]. It is generally considered a pioneer or early seral species [35] which flourishes and completes its life cycle during the first years after disturbance [100]. This shade-intolerant species often dominates sites during early successional stages but decreases as the canopy closes [62,100]. Although the plants themselves remain prominent for only a relatively brief period, viable seeds can persist for 60 years or more in the soil or duff [73]. Widespread germination after disturbance frequently leads to the development of even-aged stands [100]. In many areas, red raspberry is absent beneath the canopy of mature forests but persists in forest openings [39]. Whitney [99] reports that few stands of red raspberry persist for longer than 5 to 12 years. Red raspberry invades black and white spruce stands in Alaska during the first years after disturbance but declines as taller shrubs and trees become established [29,39]. In many northern black spruce forests, red raspberry is present only in early successional stages [29]. On mesic and submesic sites in sub-boreal forests of British Columbia it typically increases during the first 10 years after timber harvest or fire but is virtually eliminated within 14 years because of rapid increases in shade [39]. Red raspberry often dominates jack pine stands of Minnesota within 5 years after disturbance [2,39] and subsequently declines as the canopy develops. In parts of western Montana, red raspberry initially grows rapidly but begins to decline within 3 to 4 years after disturbance as nutrient levels decrease [18]. Red raspberry can persist for up to 4 or 5 years in northern hardwood forests as long as stands remain relatively open [47,73]. In birch-maple forests of New Hampshire, red raspberry reaches peak abundance in the second through fourth years after disturbance [100]. However, it rarely persists for more than 10 years [100]. Red raspberry is subsequently replaced by species such as aspen, chokecherry (Prunus spp.), and birch [100]. SEASONAL DEVELOPMENT : Red raspberry is typically biennial, with each shoot passing through well-defined phenological stages during its 2-year lifespan [45]. Vegetative shoots develop from the roots or stems of parent plants, or as seedlings, during the first year [45,99,100,101]. Lateral flowering stalks (floricanes) are produced during the second year [22,99,100]. Floricanes leaf out early and exhibit rapid growth [99]. After producing fruit in late summer, the leaves of floricanes senesce and the cane gradually dies [99]. Stages of the 2-year growth cycle of red raspberry are detailed below [45]: YEAR 1 Phases phase 1: initiation of root buds phase 2: subterranean suckering phase 3: emergent suckers; elongation slows or stops as sucker reaches surface; leaves form a rosette at or above the soil surface. phase 4: 1st winter dormancy-most leaves shed YEAR 2 phase 5: elongating shoot; rapid elongation. phase 6: initiation of flower buds; shoot stops elongation at end of growing season. secondary rosettes form; axillary meristems initiate flower primordia; dormant fruit buds; leaves become senescent and fall. phase 7: breaking dormancy of flower buds; require cold to break dormancy; buds grow in spring (some cultivars produce fruit before dormancy). phase 8: flowering and fruiting; basal buds elongate into a vegetative replacement shoot which repeats the biennial cycle. phase 9: senescence and death. (after fruiting the shoot dies back "to the position from which a replacement shoot has grown"). Flowering: Flowerbud initiation is influenced by temperature, genetics (cultivar), and geographic location [21,48,69,92]. Flowering is also related to the age and vigor of the plant and the date at which vegetative growth terminates [16,67]. Flowerbud initiation is triggered by low temperatures and short days and generally begins in late summer or autumn [16,21]. Flowerbud initiation can be induced by exposure to temperatures of 55 degrees F (12.8 degrees C) at 9 hour days or 50 degrees F (10.0 degrees C) at 16 hour days [21]. Although flowerbud initiation occurs over winter in most red raspberries, initiation in primocane-fruiting cultivars begins in summer [92]. Bud break typically occurs in early spring [16]. Evidence suggests that higher spring temperatures may promote earlier and more rapid flowering [67]. Fruiting: Fruit maturation begins soon after flowering [88]. Timing of flowerbud initiation largely determines fruiting season [21], although fruiting dates also vary according to cultivar and geographic location [21]. Annual variation in fruit ripening has also been reported [21]. Both flowering and fruiting proceeds from the top of the floricane downward [22]. After maturation, fruit spoils rapidly [88]. Generalized fruiting and flowering dates by geographic location are as follows [21,23,36,54,80,84,86,88,93]: location flowering fruiting AK June-July July-September AZ June-July ----- BC ----- July-August CO June-July ----- East ----- July-October Great Plains May-July ----- MT June-August ----- NC, SC June-August July-August (or later) ND June ----- New England ---- late June-August nc Plains June-July July-August UT May-July ----- WY June-August -----

FIRE ECOLOGY

SPECIES: Rubus idaeus | Red Raspberry
FIRE ECOLOGY OR ADAPTATIONS : The life cycle of red raspberry is integrally associated with disturbances such as fire. In many areas of vigorous fire suppression, both plant vigor and abundance have decreased [66]. Red raspberry typically flourishes, completes its life cycle and declines within the early years after disturbance [73]. As shade levels increase in the postfire community and soil nitrate levels drop (generally during the first 5 years after fire), red raspberry shifts resource allocation from vegetative growth to seed production [39,99]. Although the plants themselves soon senesce and die, viable seed persists for decades [62,73], germinating in great numbers after the next fire [100] creates favorable conditions for growth and establishment. Seed is effectively scarified by heat [78,94], and exposed mineral soil serves as a favorable substrate for early growth and development [26]. Underground regenerative structures appear to be well protected from the damaging effects of heat [28,52], and reestablishment is typically rapid where plants were present in the preburn community. POSTFIRE REGENERATION STRATEGY : Tall shrub, adventitious-bud root crown Rhizomatous shrub, rhizome in soil Geophyte, growing points deep in soil Ground residual colonizer (on-site, initial community) Initial-offsite colonizer (off-site, initial community)

FIRE EFFECTS

SPECIES: Rubus idaeus | Red Raspberry
IMMEDIATE FIRE EFFECT ON PLANT : Red raspberry is described as "resistant" to fire [39,103]. However, foliage is extremely susceptible to fire-induced mortality [52]. In an Alberta study, all aboveground stems were completely killed wherever supplemental fuels contributed to relatively intense fires [52]. Where fuels were reduced and fires less intense, the stems of many plants were only partially killed [52]. However, all aerial stems experienced at least partial mortality, regardless of fire intensity. Belowground regenerative structures appear to be relatively resistant to fire [39]. Johnston and Woodard [52] observed belowground mortality only on plots with high surface fuel loadings (3.94 or 9.65 kg/m sq). Here, tissue mortality extended as far as 0.4 to 1.2 inches (1-3 cm) below the duff surface. Raspberry is capable of sprouting from lateral buds on relatively shallow roots. These roots, which are small and succulent, are poorly protected by duff and can be damaged by fires of high intensity and severity [52]. However, at least some regenerative structures typically grow to 2 inches (5 cm) below the soil surface, and many are apparently unharmed by fires of even high intensity and severity [52]. In general, the effects of fire on red raspberry are much less pronounced wherever nutrients and water are abundant [39]. The long-lived seed of red raspberry is generally unharmed by fire when protected by overlying soil [39,78,94]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : In an Alberta study, both dead and live woody stems remained where fuel loadings of 0.00, 0.17, and 0.87 kg/m sq were recorded [52]. However, all foliage was completely consumed on plots with fuel loadings of 3.94 and 9.65 kg/m sq [52]. PLANT RESPONSE TO FIRE : Red raspberry is well adapted to reoccupy a site quickly after fire. This common "fire follower" is favored by increased amounts of nitrates present on burned sites and generally exhibits rapid and vigorous postfire growth through sprouting and/or seedling establishment [4,95]. Vegetative response: Red raspberry typically sprouts readily after fire wherever present in preburn communities [18,95]. Most belowground regenerative structures appear to be well protected from the damaging effects of heat [52]. Postfire sprouting of root buds is commonly observed. Although more shallow root buds may be damaged or killed by heat, root bud depths can range from 1.9 to 2.4 inches (5-6 cm) or greater [28,52] and many escape serious damage. Postfire sprouting from rhizomes may also occur [39], although a number of researchers have reported no evidence of any rhizomes with regenerative capabilities in the red raspberry [28]. Where light fires damage but do not kill the aboveground foliage, aerial stems generally resprout and quickly resume growth [52]. Consequently, red raspberry is reported to be "rejuvenated" by fire [103]. Johnston and Woodard [52] reported that fire intensity and severity had little effect on the sprouting ability of red raspberry in aspen communities of east-central Alberta. Both the number of sprouts produced per plant and total biomass appeared unaffected by fire intensity and severity. However, the height growth of individual sprouts was greatest after fires of low severity. Thus, although high severity fires reduced the rate of sprout growth, they did not affect the number of sprouts produced by each plant [52]. Seedling establishment: Rapid postfire establishment through on-site seed is common in the red raspberry [95]. Long-lived seed, which is produced in abundance, accumulates in seedbanks in the soil or duff [35,38]. Germination is enhanced by exposure to heat [78,94], and large numbers of seed germinate soon after disturbance [100]. Mineral soil creates a favorable seedbed [26] and elevated nitrate levels enhance early seedling growth. Most germination occurs within the first year after fire [38]. Limited evidence suggests that fires of high intensity and severity may promote red raspberry seedling establishment more than light fires. Bock and Bock [6] observed vigorous seedling establishment after crown fires in ponderosa pine forest of the southern Black Hills. However, large increases in red raspberry did not occur after lighter, cooler ground fires in the same area [6,7]. Extremely light fires may provide insufficient heat scarification and do little to prepare a seedbed. Postfire recovery: Postfire recovery of red raspberry is generally rapid, with vigorous expansion in cover during early seral stages. This shade-intolerant species [95,100] declines as tree cover increases [29]. In many communities, red raspberry begins to decline within only 3 or 4 years after fire [18,103]. It is important to note that many variables can significantly influence the speed of postfire recovery and subsequent persistence within the community. Such variables may include season of burn, fire intensity and severity, site characteristics, genetic variation, and climatic factors. Specific postfire response of red raspberry by community is discussed in the 'Successional Status' slot. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : In some areas, significant differences in postfire recovery of red raspberry have been noted after fires of varying intensity and severity. The following data document postfire recovery in ponderosa pine stands of the southern Black Hills of South Dakota [7,6]: # of stems per subplot fire type preburn 1st year 2nd year 5th year light burn 0 0 0.07 --- crown fire --- --- 0.01 116.8 # of individuals fire type 1st year 2nd year 3rd year 5th year cool ground fire 0 0 3 --- hot crown fire --- --- --- 4,672 FIRE MANAGEMENT CONSIDERATIONS : Fire generally benefits animals that consume the fruits of species within the genus Rubus [58].

FIRE CASE STUDIES

SPECIES: Rubus idaeus | Red Raspberry
CASE NAME : Elk Island Burn, Alberta REFERENCE : Johnston, M.; Woodard, P. 1985 [52] SEASON/SEVERITY CLASSIFICATION : May 5, 1980/variable STUDY LOCATION : The study site is located in Elk Island National Park, approximately 23 miles (37 km) east of Edmonton, Alberta. PREFIRE VEGETATIVE COMMUNITY : Preburn overstory vegetation was made up of quaking aspen (Populus tremuoides) and balsam poplar (P. balsamifera), which ranged from 50 to 70 years of age. Understory vegetation was dominated by red raspberry (Rubus idaeus), beaked hazelnut (Corylus cornuta), strawberry (Fragaria spp.), fleabane (Erigeron spp.), and reedgrass (Calamagrostis spp.). TARGET SPECIES PHENOLOGICAL STATE : not reported. SITE DESCRIPTION : Soil - orthic gray luvisol Elevation - not reported Topography - not reported Size of treated area - 9.9 acres (4 ha) Weather conditions - dry FIRE DESCRIPTION : Seven artificial fuel beds (24 x 24 inches [60 x 60 cm]) were constructed of varying amounts of excelsior, or excelsior mixed with white spruce slats. The prescribed head fire did not spread to the beds because of discontinuous fuels. The beds were ignited with matches after the passage of the flame front. Specific weather and fire behavior characteristics were as follows: Fire weather: Dry bulb temperature (C) - 14.5 Relative humidity (%) - 33 Wind speed at 10 m (kh/h) - 6 Fine fuel moisture code - 88 Duff moisture code - 57 Drought code - 106 Initial spread index - 4 Buildup index - 58 Fire weather index - 12 Estimated fire behavior characteristics recorded for the seven fuel beds ignited with matches - fuel loading flame length frontal fire residence time (kg/m sq.) (m) intensity (minutes) (kW/m) 0.17 0.5 57 1.5 0.87 1 258 2 0.87 1 258 2 3.94 1.5 622 4 3.94 1.5 622 4 9.65 1.5 1,162 10 9.65 2.5 1,905 10 FIRE EFFECTS ON TARGET SPECIES : All aboveground red raspberry stems were killed where fuels were added. On fuel-free plots, only portions of the aboveground stems were killed. Plants sprouted from underground regenerative structures where aboveground mortality was complete, but from both aboveground and belowground tissues where portions of the aerial stems were killed. Depth of underground regenerative structures ranged from 0 to 2 inches (0-5 cm), with mortality of tissues occasionally extending to 0.4 to 1.2 inches (1-3 cm) below the duff surface. Portions of most deeper "rhizomes" apparently survived. Red raspberry also reproduces through lateral buds located on small, shallow, succulent, poorly protected roots which can be damaged by fires of high intensity. The number of sprouts produced per plant did not vary according to burn treatment. Height growth and the number of leaves did vary by treatment, indicating that red raspberry recovery may be affected by fire intensity and severity. Specific recovery rate data for red raspberry were as follows: date response fuel loading (kg/m sq.) parameter 0.00 0.17 0.87 3.94 9.65 July 4 #sprouts/plot 5 4 4 4 5 ht. growth/sprout(cm) 52.0 32.7 17.4 12.2 8.8 avg. # leaves/sprout -- 8 6 5 3 July 22 #sprouts/plot 6 4 5 4 7 ht. growth/sprout(cm) 58.7 36.5 19.5 16.1 12.8 avg. # leaves/sprout -- 8 6 5 5 Aug. 1 #sprouts/plot 3 4 4 4 7 ht. growth/sprout(cm) 58.4 36.8 19.1 16.2 13.5 avg. # leaves/sprout -- 8 6 5 5 Aug. 16 #sprouts/plot 3 4 5 4 7 ht. growth/sprout(cm) 60.0 37.2 21.0 16.7 13.5 avg. # leaves/sprout -- 7 7 6 6 Aug. 30 #sprouts/plot 3 2 4 3 7 ht. growth/sprout(cm) 60.2 37.0 20.8 16.8 13.8 avg. # leaves/sprout -- 7 6 5 6 Mean abovegrd. ovendry biomass (grams/plot) 2.5 4.1 5.0 4.6 4.4 FIRE MANAGEMENT IMPLICATIONS : Red raspberry may be somewhat susceptible to high intensity fires because of shallow, fairly poorly protected root buds. This study suggests that hot prescribed fires may be useful in reducing red raspberry in some carefully selected instances.

REFERENCES

SPECIES: Rubus idaeus | Red Raspberry
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Related categories for Species: Rubus idaeus | Red Raspberry

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