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Introductory

SPECIES: Rubus parviflorus | Thimbleberry
ABBREVIATION : RUBPAR SYNONYMS : NO-ENTRY SCS PLANT CODE : RUPA RUPAP2 RUPAV COMMON NAMES : thimbleberry western thimbleberry salmonberry mountain sorrel white-flowering raspberry western thimble raspberry TAXONOMY : The currently accepted scientific name of thimbleberry is Rubus parviflorus Nutt. [72]. Many formerly recognized varieties are no longer accepted by most taxonomists. Varieties are as follows [72,103]: R. p. var. parviflorus R. p. var. velutinus (Hook and Arn.) Greene Hybrids between thimbleberry and red raspberry (R. idaeus) have been reported, as have hybrids between thimbleberry and evergreen blackberry (R. laciniatus) [71]. However, these hybrids are frequently sterile. LIFE FORM : Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : D. Tirmenstein, October 1989 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Tirmenstein, D. 1989. Rubus parviflorus. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Rubus parviflorus | Thimbleberry
GENERAL DISTRIBUTION : Thimbleberry grows from Alaska to California and the mountains of New Mexico, and into northern Mexico [64,133]. It extends eastward through the Dakotas to the Great Lakes States [64,133]. ECOSYSTEMS : FRES11 Spruce - fir 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 FRES27 Redwood FRES28 Western hardwoods FRES34 Chaparral - mountain shrub FRES44 Alpine STATES : AK AZ CA CO ID MI MN MT NM ND OR SD UT WA WY AB BC ON MEXICO ADMINISTRATIVE UNITS : APIS BAND CRLA CRMO DENA GLAC GRCA GRTE ISRO KICA LAVO MORA NOCA OLYM PIRO PORE REDW ROMO SAJH SEQU TICA VOYA WHIS YELL YOSE BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains 3 Southern Pacific Border 4 Sierra Mountains 5 Columbia Plateau 8 Northern Rocky Mountains 9 Middle Rocky Mountains 10 Wyoming Basin 11 Southern Rocky Mountains 12 Colorado Plateau KUCHLER PLANT ASSOCIATIONS : K001 Spruce - cedar - hemlock forest K002 Cedar - hemlock - Douglas-fir forest K003 Silver fir - Douglas-fir forest K004 Fir - hemlock forest K005 Mixed conifer forest K006 Redwood forest K007 Red fir forest K011 Western ponderosa forest K012 Douglas-fir forest K013 Cedar - hemlock - pine forest K014 Grand fir - Douglas-fir forest K015 Western spruce - fir forest K018 Pine - Douglas-fir forest K020 Spruce - fir - Douglas-fir forest K021 Southwestern spruce - fir forest K025 Alder - ash forest K029 California mixed evergreen forest K052 Alpine meadows and barren K093 Great Lakes spruce - fir forest K095 Great Lakes pine forest K106 Northern hardwoods K107 Northern hardwoods - fir forest K108 Northern hardwoods - spruce forest SAF COVER TYPES : 5 Balsam fir 16 Aspen 18 Paper birch 25 Sugar maple - beech - yellow birch 27 Sugar maple 37 Northern white cedar 107 White spruce 201 White spruce 202 White spruce - paper birch 206 Engelmann spruce - subalpine fir 207 Red fir 210 Interior Douglas-fir 211 White fir 212 Western larch 213 Grand fir 215 Western white pine 217 Aspen 218 Lodgepole pine 221 Red alder 222 Black cottonwood - willow 223 Sitka spruce 224 Western hemlock 225 Western hemlock - Sitka spruce 226 Coastal true fir - hemlock 227 Western redcedar - western hemlock 228 Western redcedar 229 Pacific Douglas-fir 230 Douglas-fir western hemlock 232 Redwood 233 Oregon white oak 234 Douglas-fir - tanoak - Pacific madrone 237 Interior ponderosa pine 243 Sierra Nevada mixed conifer 244 Pacific ponderosa pine - Douglas-fir 245 Pacific ponderosa pine 251 White spruce - aspen 252 Paper birch SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Thimbleberry is dominant or subdominant in a variety of forest and shrub communities. It has been included as a codominant in published classifications with grand fir (Abies grandis), Shasta red fir (Abies magnifica var. shastensis), red alder (Alnus rubra), Sitka spruce (Picea sitchensis), Rocky Mountain maple (Acer glabrum), quaking aspen (Populus tremuloides), trailing blackberry (Rubus ursinus), salmonberry (R. spectabilis), black twinberry (Lonicera involucrata), dwarf bilberry (Vaccinium myrtillus), twinflower (Linnaea borealis), western swordfern (Polystichum munitum), bearded wheatgrass (Elymus trachycaulus ssp. subsecundum), wild sarsaparilla (Aralia nudicaulis), brome (Bromus spp.), pokeweed fleeceflower (Polygonum phytolaccaefolium), fireweed (Epilobium angustifolium), fleabane (Erigeron eximus), and broadleaf starflower (Trientalis latifolia). Thimbleberry is listed as an indicator or dominant in the following publications: 1. Classification of the forest vegetation of Colorado by habitat type and community type [2] 2. Forest vegetation on National Forests in the Rocky Mountain and Intermountain Regions: habitat and community types [3] 3. Plant communities and environmental relationships in a portion of the Tilmoook Burn, northwestern Oregon [6] 4. Synecological features of a natural headland prairie on the Oregon coast [23] 5. Climax forest series of northern New Mexico and southern Colorado [26] 6. A classification of forest habitat types of northern New Mexico and southern Colorado [27] 7. Forest habitat types in the Apache, Gila, and part of the Cibola National Forests, Arizona and New Mexico [35] 8. Natural Vegetation of Oregon and Washington [39] 9. Ecoclass coding for the Pacific Northwest plant associations [49] 10. Plant association and management guide: Willamette National Forest [60] 11. A classification of spruce-fir and mixed conifer habitat types of Arizona and New Mexico [97] 12. Aspen community types of the Intermountain Region [101] 13. A forest habitat type classification of southern Arizona and its relationship to forests of the Sierra Madre Occidental of Mexico [102] 14. Classification of quaking aspen stands in the Black Hills and Bear Lodge Mountains [115] 15. The grand fir/mountain maple habitat type in central Idaho: succession and management [116] 16. A classification of the Cercocarpus montanus, Quercus macrocarpa, Populus deltoides, & Pices glasuca habitat types of the Black Hills NF [117] Common understory associates of thimbleberry include ceanothus (Ceanothus spp.), snowberry (Symphoricarpos spp.), elderberry (Sambucus spp.), gooseberry (Ribes spp.), willow (Salix spp.), tall Oregon-grape (Berberis aquifolium), salal (Gautheria shallon), red huckleberry (Vaccinium parvifolium), bigleaf maple (Acer macrophyllum), bittercherry (Prunus emarginata), oceanspray (Holodiscus discolor), ninebark (Physocarpus malvaceus), bracken, western swordfern (Polystichum munitum), lupine (Lupinus spp.), common cow-parsnip (Heracleum lanatum), and fireweed (Epilobium angustifolium) [4,13,16,29].

VALUE AND USE

SPECIES: Rubus parviflorus | Thimbleberry
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Wildlife: Thimbleberry provides food and cover for a variety of wildlife species. Browse: In many areas, thimbleberry browse is reportedly of little value to most ungulates [17,105]. However, in parts of Oregon, thimbleberry is considered one of the most important summer browse species of the black-tailed deer [61]. In a western Oregon study, deer ate foliage until leaf drop in autumn, but rarely fed on the twigs during winter [62]. Where deer winter at elevations below the range of thimbleberry, browse becomes unavailable [62]. Thimbleberry leaves, stems, blossoms, and fruit made up 1.82 percent of the total black-tailed deer browse in a western Washington study [12]. In parts of the West, thimbleberry is regarded as an outstanding mule deer food [127]. However, in some areas deer winter at elevations below the range of thimbleberry, and browse becomes unavailable [62]. This shrub is an important summer moose browse in parts of northwestern Wyoming [106]. Bighorn sheep feed on the foliage of many species within the Rubus genus [127]. Summer elk utilization in the Selway River Drainage of Idaho was reported as follows [137]: June 1-July 15 20 percent July 15-September 15 50 percent September 15-October 15 30 percent On particularly moist, sites thimbleberry foliage occasionally grows beyond the reach of deer and other large ungulates [61]. Numerous small rodents eat limited quantities of bark [24]. Thimbleberry leaves may represent as much as 3 percent of the annual diet of white-footed voles in parts of Oregon. White-footed vole utilization may be particularly heavy during the fall when thimbleberry leaves may comprise up to 14 percent of the rodent's total diet [132]. The leaves of many species of Rubus provide food for rabbits and mountain beaver [18,127]. The porcupine and beaver occasionally consume the twigs, buds, and cambium of Rubus [127]. Fruit: Thimbleberry fruit represents an important dietary item for many birds and mammals [17,105]. Fruit is eaten by black and grizzly bears [116,138], and numerous smaller mammals. The coyote, common opossum, Townsend chipmunk, raccoon, red fox, gray fox, pika, red squirrel, golden-mantled ground squirrel, skunks, squirrels, and chipmunks also eat the fruit of Rubus. "Berries" of species within the Rubus genus are eaten by many birds, including the ruffed grouse, blue grouse, sharp-tailed grouse, ring-necked pheasant, California quail, greater prairie chicken, gray (Hungarian) partridge, American robin, thrushes, thrashers, towhees, northern bobwhite, gray catbird, band-tailed pigeon, northern cardinal, yellow-breasted chat, pine grosbeak, and brown thrasher [18,127]. Livestock: Thimbleberry is of relatively little value as livestock forage [68,105,127]. It is occasionally eaten by domestic sheep, particularly in the Southwest, but is rarely used by cattle [24]. PALATABILITY : Palatability of thimbleberry browse to most large ungulates is described as "fair" or "intermediate" [28,137]. Palatability of thimbleberry browse to elk in Idaho appears to be somewhat greater in late summer than earlier in the season [137]. Thimbleberry fruit is highly palatable to many birds and mammals. However, limited evidence suggests that palatability may vary geographically [130]. The degree of use shown by livestock and wildlife species for thimbleberry is rated as follows [28,113,116]: CA CO MT UT WY ID Cattle useless fair poor poor ---- low Domestic sheep poor fair fair fair ---- moderate Horses useless fair fair poor ---- ---- Pronghorn ---- ---- ---- poor poor ---- Bighorn ---- ---- ---- ---- ---- ---- Elk ---- ---- poor fair fair summer-high winter-low Moose ---- ---- ---- ---- ---- ---- Mule deer ---- ---- poor fair fair ---- White-tailed deer ---- fair poor ---- ---- ---- Small mammals ---- ---- ---- good good ---- Small nongame birds ---- ---- ---- good good ---- Upland game birds ---- ---- ---- good fair ---- Waterfowl ---- ---- ---- poor poor ---- Domestic goats fair-poor ---- ---- ---- ---- ---- Deer fair-poor ---- ---- ---- ----summer-mod. winter-low Black bear ---- ---- ---- ---- ----spring,fall-low summer-mod. NUTRITIONAL VALUE : Thimbleberry is rated as poor in energy and protein value [28]. Crude protein value of the foliage varies both seasonally and annually. Protein levels are generally highest during the spring and summer and then decline markedly in winter [32,108]. Specific values of foliage protein taken from plants in Oregon Coast Ranges are documented below [108]: crude protein - percent date grazed plots ungrazed plots October 8.1 7.8 March 22.5 25.2 October 9.0 8.2 In Oregon seasonal protein values varied from 8.58 percent in June to 4.25 percent in December [32]. COVER VALUE : Thimbleberry provides important cover for a variety of wildlife species. Dense thickets of thimbleberry form good nesting habitat for many small birds [18]. During summer, deer frequently bed down during the hottest part of the day in dense, cool stands of thimbleberry growing beneath a red alder canopy [61,62]. Coastal brushfields dominated by thimbleberry or salmonberry serve as favorable habitat for small mammals, such as deer mice, chipmunks, voles, shrews, hares, and mountain beaver [67]. Species within the Rubus genus provide cover for rabbits, red squirrel, pika, black bear, and beaver [127]. Subalpine fir (Abies lasiocarpa)/thimbleberry habitat types of the Southwest offer good thermal protection for big game during the hot summer months [35]. The degree to which thimbleberry provides environmental protection during one or more seasons is rated as follows [28]: UT WY Pronghorn poor poor Elk poor poor White-tailed deer ---- poor Small mammals good fair Small nongame birds good fair Upland game birds ---- fair Waterfowl poor poor VALUE FOR REHABILITATION OF DISTURBED SITES : Thimbleberry reportedly has only low to moderate value in preventing soil erosion [28], although it has shown good potential for revegetating many types of disturbed sites. It has the ability to form clumps which expand greatly once established [8]. Thimbleberry has proven useful in rehabilitation projects in Yosemite National Park, the coastal mountains and foothills of California, and on roadcuts in the northern Rocky Mountains of Idaho and Montana [8,68]. This shrub exhibited good survival (73 percent) as long as 4 years after plantings were made in western Montana and northern Idaho [68]. Initial propagation is reportedly more difficult than for other species within the Rubus genus [8]. Seed may be difficult to obtain commercially [34]. Thimbleberry may be propagated vegetatively by planting stem cuttings or rhizome fragments [15]. Best results have been obtained from starting dormant rhizome segments [8]. Several cultivars are now commercially available [8]. OTHER USES AND VALUES : The fruits of thimbleberry are edible, although palatability tends to be greater towards the eastern portion of this plant's range where rainfall is greater [130]. "Berries" make excellent jelly but are too seedy for jam [128]. Approximately 0.27 quart (250 ml) of fruit can be harvested within 10 minutes [87]. Young shoots may be eaten as greens and leaves have been used in making teas [75]. Thimbleberry was traditionally used by indigenous peoples throughout its range. The fruit was eaten fresh in summer and dried for winter use. The bark was boiled and made into soap, and leaves were used to make a medicinal tea. Leaves were powdered and applied to burns to minimize scarring [55]. Thimbleberry is occasionally planted as an ornamental because of its attractive fragrant flowers and colorful fall foliage [71,130]. Several cultivars have been developed, including `Golden' and `Colonel' [8]. MANAGEMENT CONSIDERATIONS : Timber harvest: Thimbleberry often assumes prominence on recent clearcuts in Douglas-fir (Pseudotsuga menziesii) forests of the Northwest [13,36,96]. Thimbleberry can establish as early as the first growing season after clearcutting [139]. In some areas it dominates the understory within 5 years after logging and slash disposal [96]. Thimbleberry is also an initial increaser after logging in western redcedar (Thuja plicata)-western hemlock (Tsuga heterophylla) forests of northern Idaho [100,135] and in grand fir forests of northwestern Montana [5]. Evidence suggests that thimbleberry is often most favored by relatively severe treatments. Cover is up to ten times greater on plots with little overstory than on sites where the tree cover is 55 percent or more [100]. Species such as thimbleberry and salmonberry compete vigorously with Douglas-fir, western hemock (Tsuga heterophylla) and Sitka spruce seedlings in parts of the Northwest [4,55,84,111,112], and conifer regeneration may be significantly impaired in some areas. Thimbleberry may compete with conifer seedlings for water during summer dry periods in parts of British Columbia, [15]. Conifer regeneration: Although competing with conifer seedlings in some areas, light to moderate thimbleberry cover may also provide "safe" microsites for developing seedlings on some disturbed sites [15,42,116]. Ponderosa pine (Pinus ponderosa), Engelmann spruce (Picea engelmannii), lodgepole pine (Pinus contorta), and grand fir regenerate well under thimbleberry in grand fir/Rocky Mountain maple or grand fir/blue huckleberry (Vaccinium membranaceum) habitat types of central Idaho [42]. Natural regeneration of western larch (Larix occidentalis) is generally good on sites with thimbleberry [116]. Chemical control: Thimbleberry is reported to be moderately susceptible to herbicides [44,125]. Herbicides such as picloram + 2,4-D, glyphosate, and 2,4,5-T have proven effective in controlling thimbleberry [15,16,44,90,91,94]. Late foliar application of 2,4,5-T can be particularly useful in releasing Douglas-fir and Sitka spruce (Picea sitchensis) seedlings [44]. Guidelines detailing the relative effectiveness of various herbicides on thimbleberry and the proper method of application are available [16,43,44,90,91,92,93,94,95,118,119,125]. Recreational impacts: Thimbleberry reportedly exhibits relatively low resistance to trampling in recreational use studies but has high short- and long-term resilience [14]. Browsing: Thimbleberry typically decreases in response to heavy domestic sheep use [101]. Mechanical removal: Thimbleberry generally resprouts rapidly following mechanical removal and can attain pretreatment height and cover within 1 year after cutting [15].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Rubus parviflorus | Thimbleberry
GENERAL BOTANICAL CHARACTERISTICS : Thimbleberry is a low, scrambling or erect, unarmed deciduous shrub which grows from 1.6 to 8.2 feet (0.5-2.5 m) in height [15,17,45,103,133]. The perennial stems are few, erect, and simple [75,128]. The perennial stems, or canes, typically live for 2 to 3 years [15]. Sterile first-year stems, known as primocanes, develop from buds at or below the ground surface and generally bear only leaves. During the second year, lateral branches, or floricanes, develop in the axils of the primocanes produce flowers and fruit [45]. Growth is rapid under favorable growing conditions. Plants may reach heights of 6.6 to 8.2 feet (2 to 2.5 m) within 10 years. Maximum rates of height growth often occur during the second and third years after establishment [15]. Twigs are greenish and finely hairy [128]. Grayish bark becomes flaky or shreddy with age [103]. Large, simple, palmately-lobed leaves are green above but pale beneath [133]. Leaves are unequally serrate [103] and turn brilliant orange to maroon in fall [8]. Showy white or, rarely, pink flowers occur in clusters of two to seven [133]. Loosely cymose, perfect flowers are borne in a flat-topped terminal panicle or corymb [45,64]. The fruit of thimbleberry is made up of a "thimblelike" aggregate of numerous hairy, red or scarlet drupelets [45,103,128,133]. These drupelets are nearly dry at maturity and fall apart readily when picked [133]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Thimbleberry reproduces through seed but also regenerates vegetatively, even in the absence of disturbance. It is capable of forming dense thickets through vegetative sprouting [112]. Establishment from seed appears to be the primary mode of colonization in newly disturbed areas [15]. Abundant seedling establishment typically occurs during the first year after disturbance [78]. Vegetative regeneration: Thimbleberry, a strongly rhizomatous shrub [52,105,131], is also capable of vigorous sprouting from rootcrowns [105] and roots [17,118]. A single thimbleberry seedling can spread and occupy a relatively large area as rhizomes develop and spread [80]. Most local expansion of this shrub is attributable to rhizome sprouting [123]. Seed: Most species within the genus Rubus produce good seed crops nearly every year [11]. However, thimbleberry seed production can be limited at higher elevations. Thimbleberry fruits, or "berries," are made up of an aggregate of numerous small red drupelets [128,133] which fall to the ground when ripe [15]. Fruit is almost dry at maturity and readily breaks apart when picked [133]. Seed averages 0.8 inches (2 mm) in length [80]. Germination: Thimbleberry seeds have a hard, impermeable endocarp and dormant embryo [80]. Consequently, germination is often slow [105]. Seed has exhibited moderate viability in germination tests, with maximum germination estimated at 62 percent [15]. Specific germination requirements have not been documented for thimbleberry, but both warm and cold stratification are probably required [105]. Most Rubus 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. These conditions are frequently encountered naturally as seeds mature in summer and remain in the soil throughout the cold winter months. Laboratory tests indicate that exposure to sulfuric acid solutions or sodium hyperchlorite prior to cold stratification can enhance germination [11]. Allelopathic compounds produced by bracken (Pteridium aquilinum) can apparently inhibit germination and subsequent growth of thimbleberry [15]. Seedbanking: Thimbleberry seed is noted for its ability to remain viable for long periods of time [105,116]. Seedbanking is believed to represent an important postdisturbance regenerative strategy in this species [77,80,98,116]. Kramer [80] observed buried thimbleberry seed in 50 percent of all plots examined in several Douglas-fir and grand fir habitat types of central Idaho. Kramer and Johnson [82] observed maximum seed densities of 17.6 per foot square (189/m sq) in central Idaho. Approximately 91 percent of the thimbleberry seed was found buried at a depth of 0 to 2 inches (0-5 cm), with 9 percent at 2 to 4 inches (5-10 cm) [82]. Kramer [80] found that 22 percent of all seeds were buried at depths of 2 to 4 inches (5 to 10 cm). Approximately 14 percent of all buried seed was viable, although the total number of viable seeds from lower soil layers (2 to 4 inches [5-10 cm]) was less than one half of that of the more shallowly buried seed [80]. Seed dispersal: Thimbleberry seed is readily dispersed by birds and mammals [1,52,80]. Groups of seedlings occasionally germinate from rodent caches. However, small mammals generally play only a local role in thimbleberry seed dispersal [98]. Birds often effect wider dispersal [15]. Gravity may also aid in seed dispersal [105]. SITE CHARACTERISTICS : Thimbleberry occurs in a variety of moist-to-dry and wooded-to-open sites [64]. It commonly grows on open, wooded hillsides, in subalpine meadows, along streambanks and canyons, on borders, and roadsides, and on dry exposed sites at higher elevations [17,39,45,60,128,130]. Thimbleberry is often well represented in areas of snow creep, such as in avalanche chutes or on steep slopes [29,48]. It is particularly common on cool moist sites and predominates on north aspects throughout much of northern Idaho [69]. Thimbleberry frequently occurs as scattered individuals, but in some areas occurs in dense contiguous patches [15]. This moderately shade-tolerant shrub [76,116] is occasionally found at low light levels but is typically most abundant at 60 to 100 percent of full sunlight in Oregon forests [116]. In some areas, thimbleberry can survive within the understory of forests receiving only 3.5 percent of full sunlight [15]. Although it persists in closed stands, it is typically most abundant on disturbed sites within the forest canopy, such as cutover or windthrown areas [16,30,80,87,100]. Soil: Species within the genus Rubus grow well on a variety of barren infertile soil types [11]. These shrubs tolerate a wide range of soil temperature and pH but do require adequate soil moisture for good growth [18]. Thimbleberry grows well on dry, rocky soil and deep well-drained loam [29,68]. Growth of thimbleberry is reported as "good" on loam or clay-loam, "fair-to-good" on sandy loams, but only "fair to poor" on gravel, sand, or clay [28]. Thimbleberry grows well on soils derived from a variety of parent materials [15]. Elevation: Thimbleberry grows across a wide elevational gradient from sea level to subalpine zones [55,64]. Generalized elevational ranges by state are as follows [28,73,103,133]: from 8,000 to 9,500 feet (2,438 to 2,896 m) in AZ < 8,000 feet (2,438 m) in CA from 7,000 to 10,000 feet (2,134 to 3,048 m) in CO 3,500 to 8,000 feet (1,067 to 2,438 m) in MT 4,691 to 9,004 feet (1,435-2,745 m) in UT 6,700 to 7,900 feet (2,042-2,408 m) in WY SUCCESSIONAL STATUS : Thimbleberry is a persistent seral species which frequently dominates the understory during the first several decades after disturbance [89,116,123]. It readily invades disturbed sites through rhizomes or seedling establishment [15]. Thimbleberry quickly established on mudflows and other harsh microsites after the eruption of Mount St. Helens [54]. This shrub is an important seral species in many cedar-hemlock, western hemlock, grand fir, Douglas-fir, redwood, and other coniferous forests of the West [25,39,59,116,140]. This shrub generally occurs in greatest abundance in seral stands throughout its range [59] but can persist in trace amounts in many climax forests [13,80,135]. In many mature forest stands, thimbleberry may be restricted to disturbed sites such as clearings or areas of windthrow [30]. On certain disturbed sites perpetuated by snow creep, thimbleberry is functionally both a pioneer and climax species [29]. Thimbleberry is a nitrogen-demanding species and in some areas, begins to decline within 2 to 5 years after timber harvesting as soil nutrient levels decrease [79]. Plant vigor and canopy cover generally decline as tree cover increases [69,100]. Douglas-fir: Thimbleberry is an important seral species in Douglas-fir forests of the Pacific Northwest and Rocky Mountain regions [25,51,52,53]. During the first few years after disturbance, weedy annual colonizers become dominant but are soon replaced by perennial invaders such as thimbleberry [51]. This shrub reaches peak abundance more than 10 years after disturbance in Douglas-fir forests of Oregon's western Cascades [52]. However, it reportedly becomes dominant as soon as 5 years after logging and slash disposal in northwestern Douglas-fir forests [96] and assumes prominence during the first 3 years after disturbance in northwestern Montana [21]. Grand fir: Thimbleberry is also a major seral species in grand fir habitat types of Idaho where it quickly increases following reduction of the overstory [116,139]. This shrub remained dominant in grand fir/pachistima (Pachistima myrsinites) stands which had been clearcut and burned 12 years earlier [139]. In northern Idaho it is particularly abundant after timber harvest on sites which were severely burned by wildfires 40 to 60 years earlier and frequently persists for relatively long periods of time. Consequently, some researchers note that the relative successional amplitude of thimbleberry in grand fir types of northern Idaho is uncertain [116]. In some grand fir-Sitka spruce-Douglas-fir forests of northern California, thimbleberry is abundant 5 to 10 years after disturbance. Thimbleberry is overtopped at approximately 30 years in redwood (Sequoia sempervirens)-grand fir forests but generally persists in reduced abundance [140]. Cedar-hemlock: Thimbleberry occurs in cedar-hemlock communities from newly disturbed sites to stands of 400 years or older [47] but is much more prevalent in younger stands [47]. In cedar-hemlock forests of northern Idaho, thimbleberry is an initial increaser after logging but begins to decrease by approximately 25 years after disturbance [135]. It has reached maximum cover in western redcedar/queencup beadlily (Clintonia uniflora) habitat types by 3 to 5 years after fire [98]. SEASONAL DEVELOPMENT : Phenological development of thimbleberry varies according to elevation and weather conditions [15]. It generally leafs out in mid to late spring. However, buds may become active by late February in parts of Oregon and Washington [119]. Leaves begin to fall in late summer to autumn [114]. Leaf fall may be early in dry years. In Oregon, leaves were shed by late August in particularly dry years [61]. Phenological development of thimbleberry foliage in the northern Rocky Mountains is detailed below [114]: leaf buds leaves full leaves start leaves begin leaves burst grown to color to fall fallen earliest April 5 May 12 August 1 August 21 August 28 latest June 3 July 20 Sept. 30 October 1 October 23 Similar phenological development has been reported from the central coast of British Columbia [15]. Generalized fruiting and flowering dates by location are as follows [15,28,73,87,104,128]: location flowering dates fruiting dates AK June - July August - September AZ July - September ---- BC ---- early June to September CA March - August ---- CO June - August August - later Great Plains May - July ---- MT June - August ---- n ID, w MT May - August late June - September UT May - July ---- WY June - August ---- In northern Idaho and western Montana, seed dispersal generally begins in July or August [114].

FIRE ECOLOGY

SPECIES: Rubus parviflorus | Thimbleberry
FIRE ECOLOGY OR ADAPTATIONS : Thimbleberry is resistant to and generally enhanced by fire [56,57,131]. In areas of rigorous fire suppression, thimbleberry fruit production and plant vigor has declined [87]. This species is well-adapted to vigorously invade many types of burned sites through rhizomes or seed [53,100,136]. Rhizome sprouting is an important postfire strategy which enables rapid reestablishment and spread [22,89,105,123]. Thimbleberry, a seed banker [77,80,82], also reestablishes through viable seed stored in the soil or duff [22]. Birds and mammals add to seedling establishment by transporting seed to the site [22,74]. Thimbleberry typically becomes abundant within the first few years after fire and remains prominent during the early postfire decades [22,98,123]. Hamilton and Yearsley [57] note that thimbleberry "may be well-adapted to the high nutrient availability and low competition from other species found immediately after burning, but is less successful once other species have reestablished." This fire-adapted species typically declines as the overstory develops in postfire communities. POSTFIRE REGENERATION STRATEGY : Tall shrub, adventitious-bud root crown Rhizomatous shrub, rhizome in soil Ground residual colonizer (on-site, initial community) Initial-offsite colonizer (off-site, initial community)

FIRE EFFECTS

SPECIES: Rubus parviflorus | Thimbleberry
IMMEDIATE FIRE EFFECT ON PLANT : Thimbleberry is described as resistant to fire [57,131]. Although often top-killed, underground rhizomes generally survive. High severity fires which damage belowground regenerative structures may be most damaging to rhizomes [98]. The aerial portions of relatively few plants actually survive fire, except for those on unusual microsites [52]. Most seed stored on-site is probably unharmed by fire. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Thimbleberry cover and vigor is generally enhanced by fire [20,57,100]. Vegetative response: Thimbleberry is capable of rapid, vigorous postfire spread through an extensive network of often deeply-buried rhizomes [22,74,116]. Sprouting through surviving rootcrowns is also possible, although rhizome sprouting probably represents the primary mode of postfire regeneration [105]. Multiple sprouts often replace the single stems observed in preburn communities [20]. Limited evidence suggests that sprouting of rhizomes is favored by fires of low intensity and severity [20,98]. Hot fires presumably offer greater potential for damaging underground regenerative structures. In western redcedar habitat types of northern Idaho, thimbleberry cover was estimated at 10.9 percent 2 years after low severity fire. However, following a high severity fire, thimbleberry cover reached only 4.9 percent [98]. Seedling establishment: Seed banking is an important postfire regenerative strategy in thimbleberry, a prolific seed producer [57]. Birds and mammals also transport some viable seed to burned sites [22,74]. Some seedlings have been observed to germinate from rodent caches present on-site [99]. Thimbleberry seed is stimulated by fire [52,120] and subsequently germinates in great numbers [116,120]. Researchers have observed up to 7.5 times more seedlings on plots which have been burned than on adjacent untreated plots [120]. Most seedling establishment occurs immediately after fire [57]. Some researchers have suggested that fires of high severity and intensity can expose mineral soil, which serves as a favorable seedbed for thimbleberry [98,105]. However, Morgan and Neuenschwander [99] observed greatest seedling establishment on low severity burns where duff reduction was incomplete. The capacity to regenerate through seed may be reduced by extremely hot, duff-reducing burns [99,127]. Rate of postfire recovery: Both sprouting [56] and seedling establishment occur soon after fire. Growth is rapid and some plants occasionally bear fruit during the first postfire year [22]. Thimbleberry generally reaches greatest abundance during the first years after fire and decreases as the overstory canopy closes [57]. Dwindling soil nutrients may also contribute to its decline [79]. On coastal sites in British Columbia, thimbleberry typically attains greatest height growth during the first few years after fire and remains abundant for approximately 5 years [57]. Rhizome sprouting may produce dense cover up to 3 feet (1 m) tall within 3 years after disturbance on some particularly moist sites [15]. Thimbleberry reportedly dominates the understory of many Northwestern Douglas-fir forests within just five years after logging and slash disposal [96]. In Douglas-fir forests of western Montana, thimbleberry was well-represented on many sites during the first 2 to 3 years after fire [21,74]. Maximum cover values have been observed within three to 5 years after fire in western redcedar habitat types of Idaho [98]. In many areas, this shrub remains widely distributed during the first several decades after fire [123]. Dense early growth of aggressive species such as fireweed can slow thimbleberry establishment and growth [121]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : Rate of postfire recovery may be significantly related to fire severity and intensity. However, contradictory findings have been reported. Hooker and Tisdale [66] observed rapid growth rates on severely burned sites (0.25 inch/day [0.6 cm/day]) but slower growth on less severely burned plots [66]. Slower growth may be expected where cover (and thus increased intraspecific competition) is greatest, and these results do not necessarily indicate that actual cover was greatest on these severely burned plots. Other research has suggested that thimbleberry cover develops most rapidily after fires of low intensity [15]. Some researchers suggest that, in general, rhizome sprouting may be favored by fires of low intensity and severity, whereas seedling establishment may be more effectively promoted by fires of relatively high severity [98,105]. However, researchers in northern Idaho have occasionally observed greater seedling establishment after fires of low severity [99]. Factors other than fire severity and intensity can also contribute to the rate of postfire recovery. Differences in the season of burn, plant density and vigor in the preburn community, site differences, and climatic factors all contribute to postfire recovery. Fire severity and intensity are not consistently defined in the literature, which compounds the problems of interpreting plant response. The rate of postfire recovery is significantly related to climatic and topoedaphic factors. In coastal Oregon, 8,760 thimbleberry seedlings per acre (21,382/ha) were present 1 year after fire on north aspects where 4,450 seedlings per acre (10,988/ha) had been observed prior to the fire. On south aspects, increases were even more dramatic, with 10,988 seedlings per acre (24,640/ha) counted 1 year after fire where none had been observed prior to the fire [120]. Mean frequency of thimbleberry by burn age and severity was documented as follows after fire in a western redcedar/queencup beadlily habitat type of northern Idaho [98]: fire severity years 1 2 3 4 5 15 low 83 80 80 59 56 98 high 61 76 78 75 53 100 FIRE MANAGEMENT CONSIDERATIONS : Timber harvest: Thimbleberry typically increases dramatically after timber harvest and subsequent slash burns [44,86]. In many areas, it may provide formidable competition for regenerating conifer seedlings [57]. Mueggler [100] reports that thimbleberry often assumes dominance on broadcast burns in western redcedar and western hemlock habitat types of northern Idaho [100]. Response may be rapid. Maximum cover values were reached within 5 years after fire in western hemlock and western redcedar habitat types of northern Idaho [122]. This shrub can establish as early as the first growing season after clearcutting and broadcast burns in grand fir/pachistima habitat types of north-central Idaho. More serious disturbance may enhance the growth and recovery of thimbleberry more than lighter disturbance. In the Intermountain West, Laursen [86] reported greater increases in ground cover in stands that were clearcut and burned than in unburned shelterwood or selection cuts. Maximum cover was typically reached within 20 to 25 years after clearcutting and burning [86]. Thimbleberry commonly assumes importance soon after timber harvest and slash burning in Douglas-fir forests of the Oregon Cascades. Frequency of occurrence by years after treatment was documented as follows [136]: site # 1 4 5 (years) 1 15 32 36 2 4 14 14 Wildlife: Fire generally benefits animals that eat the fruits of Rubus [83].

REFERENCES

SPECIES: Rubus parviflorus | Thimbleberry
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