Index of Species Information

SPECIES:  Rubus ursinus


SPECIES: Rubus ursinus
AUTHORSHIP AND CITATION : Tirmenstein, D. 1989. Rubus ursinus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: [].
ABBREVIATION : RUBURS SYNONYMS : Rubus macropetalus Rubus ursinus var. macropetalus Rubus vitifolius Rubus ursinus var. medusae Rubus ursinus var. sirbenus Rubus ursinus var. ursinus Rubus sirbenus Rubus vitifolius ssp. ursinus SCS PLANT CODE : RUUR RUURM RUURU RUURS RUURU2 COMMON NAMES : California blackberry California dewberry California grapeleaf dewberry Douglasberry Pacific blackberry TAXONOMY : The scientific name of California blackberry is Rubus ursinus Cham. and Schlecht. Recognized subspecies and varieties are as follows [43]: Rubus ursinus subsp. macropetalus (Dougl. ex Hook.) Taylor and MacBryde Rubus ursinus subsp. ursinus Many commercially grown cultivars have been derived from the California blackberry [54]. LIFE FORM : Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY


SPECIES: Rubus ursinus
GENERAL DISTRIBUTION : California blackberry grows from British Columbia to northern California and eastward to central Idaho [6,27,36]. It is particularly common from the Cascades to the Pacific Coast [27]. California blackberry extends through southern California into Mexico [9,55]. The subspecies macropetalus occurs from British Columbia and Idaho southward into northern California [36]. ECOSYSTEMS : FRES20 Douglas-fir FRES23 Fir - spruce FRES24 Hemlock - Sitka spruce FRES25 Larch FRES27 Redwood FRES28 Western hardwoods STATES : AZ CA ID OR BC MEXICO BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains 3 Southern Pacific Border 5 Columbia Plateau 8 Northern Rocky Mountains KUCHLER PLANT ASSOCIATIONS : K001 Spruce - cedar - hemlock forest K002 Cedar - hemlock - Douglas-fir forest K003 Silver fir - Douglas-fir forest K004 Fir - hemlock forest K006 Redwood forest K012 Douglas-fir forest K014 Grand fir - Douglas-fir forest K029 California mixed evergreen forest SAF COVER TYPES : 210 Interior Douglas-fir 211 White fir 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 234 Douglas-fir - tanoak - Pacific madrone 246 California black oak 249 Canyon live oak SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : California blackberry is well represented in a wide variety of plant communities [9,54,55]. It has been identified as a codominant with Columbia brome (Bromus vulgaris) and Shasta red fir (Abies magnifica var. shastensis) in certain plant communities of the Cascades [8]. California blackberry is listed as a codominant in the following publication: Natural vegetation of Oregon and Washington [21] Associated species: California blackberry grows as an understory species with Shasta red fir, Pacific silver fir (A. amabilis), spruce (Picea spp.), Douglas-fir (Pseudotsuga menziesii), white fir (A. concolor), grand fir (A. grandis), western redcedar (Thuja plicata), western hemlock (Tsuga heterophylla), bigleaf maple (Acer macrophyllum), and red alder (Alnus rubra) [1,19,21,32,56]. California blackberry also occurs in many West Coast riparian communities dominated by willows (Salix spp.) or cottonwoods (Populus spp.) [23.42,62], as a codominant with salmonberry (Rubus spectabilis) and thimbleberry (R. parviflorus), and in baccharis (Baccharis spp.) shrub communities of the northern California coast [31]. Common understory associates include Oregon oxalis (Oxalis oregana), sweetscented bedstraw (Galium triflorum), elderberry (Sambucus spp.), and other blackberries, raspberries, or brambles (Rubus spp.) [21].


SPECIES: Rubus ursinus
IMPORTANCE TO LIVESTOCK AND WILDLIFE : Wildlife: California blackberry provides food and cover for many wildlife species [9,14]. Blackberries are eaten by numerous birds, including the ruffed grouse, northern bobwhite, sharp-tailed grouse, California quail, ring-necked pheasant, blue grouse, gray (Hungarian) partridge, band-tailed pigeon, American robin, yellow-breasted chat, pine grosbeak, gray catbird, and summer tanager [3,10,64]. Jays, pigeons, northern mockingbird, sparrows, tanagers, thrashers, and towhees, consume the fruit of California blackberry and nest in its tangled branches [13]. Mammals, such as the coyote, common opossum, skunks, gray fox, red fox, raccoon, squirrels, chipmunks, and black bear, consume the fruit of blackberries [10,64]. Black-tailed deer feed on the stems and foliage of California blackberry [13], and in some parts of California it is considered a preferred browse [14]. In the Coast Range of western Oregon, leaves are selected by deer in all seasons except summer, when a wide variety of other foods are present [34]. In many areas California blackberry is particularly important to deer during the fall and winter [12,35]. Deer often feed heavily on the foliage until the leaves are covered by snow [12]. The young leaves, which develop earlier than those of most other associated shrubs, provide an important food source when forage supplies are lowest and deer are threatened with malnutrition [34]. Hines and Land [35] report that California blackberry browse is a preferred winter food of black-tailed deer inhabiting Douglas-fir (Pseudotsuga menziesii) forests of the Oregon Coast Ranges. In this area it supplied nearly 50 percent of the total deer forage at the beginning of winter. In other winter feeding trials, deer reduced the leaves and twigs of California blackberry by as much as 80 to 89 percent [34]. Elk feed on California blackberry through much of the year in parts of California, although utilization appears to be highest during the fall and winter [30]. Rabbits, porcupines, mountain beaver, and beaver occasionally consume the stems, leaves, and cambium of blackberries [10,64]. Livestock: Blackberries, in general, provide only minimal browse for domestic livestock. In some locations, California blackberry is moderately grazed by domestic sheep but is rarely used by cattle [37]. PALATABILITY : Berries: Fruits of the California blackberry are sweet and succulent at maturity [9]. Berries are palatable to many species of birds and mammals. Browse: California blackberry has been described as a relatively unpalatable shrub [49]. However, in parts of California and presumably elsewhere, deer exhibit a marked preference for the stem and foliage of this plant. Most blackberries are relatively unpalatable to domestic livestock. NUTRITIONAL VALUE : The specific food value of California blackberry browse has not been documented, but it is considered to be good nutritionally [35]. In feeding trials conducted by Hines [34], it was the only native browse species capable of maintaining the weight of deer over winter. The food value of California blackberry was found to vary seasonally, with the crude fat content of the leaves peaking in the fall. Only slight seasonal changes were noted in the crude fiber content of the leaves [34]. COVER VALUE : California blackberry provides important cover for a wide variety of wildlife species [14]. Dense thickets of blackberries form good nesting sites for many small birds including, thrashers, jays, pigeons, northern mockingbird, sparrows, tanagers, and towhees [10,13]. The endangered least Bell's vireo frequently nests in California blackberry thickets along willow (Salix spp.)-cottonwood (Populus spp.)-oak (Quercus spp.) ecotones in certain riparian areas of California [23]. Mammals such as rabbits, red squirrel, black bear, and beaver utilize blackberry thickets for cover in many areas [64]. VALUE FOR REHABILITATION OF DISTURBED SITES : Blackberries, because of their ability to grow well on infertile soils, may be valuable in preventing soil erosion on some sites [6,64]. California blackberry has been used to at least a limited extent in rehabilitation projects in the West. Expected mortality was estimated at approximately 50 percent following plantings in southern California. Planting densities of 66 per acre (163/ha) were recommended for best results [29]. Plants may be propagated vegetatively, transplanted, or seeded onto disturbed sites. Seed which has been scarified can be successfully planted in late summer or early fall [6]. Brinkman [6] reports that cold treatment is not required for fall plantings. Previously stratified and scarified seed can be planted in spring. 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. OTHER USES AND VALUES : Fruits of the California blackberry are sweet and edible [27]. The commercially grown loganberry, youngberry, and boysenberry were originally derived from this species [54]. Native Americans historically ate fresh blackberries in summer. Fruit was dried and combined with meat to make cakes which were eaten in winter [13]. Unripened berries were soaked in water to make a cool refreshing drink, and leaves or vines were used in making teas. Roots were boiled in water to make various medicinal preparations [9]. The fruit and stems of many blackberries have also been used to produce various tonics or medicines [6]. OTHER MANAGEMENT CONSIDERATIONS : Competition: California blackberry quickly assumes prominence on burned or logged sites. After disturbance it can compete aggressively with conifer seedlings in many locations. California blackberry frequently makes up a large proportion of the shrub cover on scarified plots in brushfields of coastal Oregon [43] and has been described as a principal understory species in recently clearcut Douglas-fir forests of the Olympic Mountains of Washington [19]. This highly competitive shrub also becomes prominent on cutover sites in many coniferous forests of both northeastern Oregon and northern Idaho [8]. Chemical control: Many chemicals including glyphosate, triclopyr, and roundup, have proven effective in controlling California blackberry [8,49]. Fifty to 80 percent control has been achieved with roundup in some locations [49].


SPECIES: Rubus ursinus
GENERAL BOTANICAL CHARACTERISTICS : California blackberry is a low-growing, California or climbing, native evergreen shrub [9,27,55]. This mound-building shrub can grow to 15 or 20 feet (5-6 m) in length [6,13,55]. The densely prickled stems are greenish when young but turn brown at maturity [9]. The somewhat prickly, deeply-lobed, alternate leaves are palmate and a lighter green color beneath [9,13,27]. The stems of most blackberries are biennial. Sterile first-year stems, known as primocanes, develop from buds at or below the ground surface and produce only leaves. Lateral branches, or floricanes, develop in the axils of the primocanes during the second year and bear both leaves and flowers [24]. Perfect flowers of California blackberry develop in clusters of 2 to 15 near the ends of leafy branches [9,13,55]. Fruit is red and hard when immature but shiny black when ripe [6]. Fruit is oblong or conical, somewhat bristly, and up to 0.8 inches (2 cm) in length [9,55]. Aggregates of druplets, commonly referred to as "berries," are sweet and flavorful at maturity [9,27]. RAUNKIAER LIFE FORM : Hemicryptophyte REGENERATION PROCESSES : California blackberry exhibits vigorous vegetative regeneration but also commonly reproduces through seed. Reproductive versatility is common 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 fruits without fertilization) occurring widely. The following types of reproduction have been documented in blackberries: (1) sexual reproduction, (2) nonreduction at meiosis on the female, male, or both sides, (3) apomixis (seed contains embryo of maternal rather than sexual origin) with segregation, (4) apomixis without segregation, and (5) haploid parthenogenesis [11]. These modes of asexual reproduction contribute to the aggressive, vigorous spread of blackberries. Vegetative regeneration: Most species within the Rubus genus are capable of vigorous sprouting from root or stem suckers, or rooting stem tips [24]. California blackberry sprouts readily from "suckers" (presumably root suckers), or "nonrhizomatous sprouts" after fire or mechanical disturbance [9,52,61]. It is also capable of spreading rapidly from trailing stems which root at the nodes [37,65]. These modes of vegetative spread occur even in the absence of disturbance. Seed production: Most blackberries produce good seed crops nearly every year [6]. During the first year of development, blackberries grow from perennial rootstocks or creeping stems and produce sterile vegetative shoots known as primocanes [24]. Lateral branches which produce both leaves and flowers (floricanes) develop in the axils during the second year [24]. Black shiny drupelets are produced on the floricanes of California blackberry [6]. Fruit is oblong to conical, and up to 0.8 inches (2 cm) in length [55]. 384,000 per pound (845,814/kg) [6]. Germination: Blackberry seeds have a hard, impermeable coat and dormant embryo; consequently, germination is often slow. Most blackberries require, as a minimum, warm stratification at 86 to 68 degrees F (30 to 20 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 [6]. 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 [3]. Seed dispersal: Fruit of the California blackberry is readily dispersed by many small birds and mammals [27]. After they mature, the sweet, succulent berries rarely remain on the plant for long [6]. Seedbanking: The seeds of most blackberries remain viable for at least several years after being buried in the soil or duff [6]. Although the precise length of viability has not been determined for the California blackberry, Morgan and Neuenschwander [52] regard it as a species which relies heavily on seedbanking for postfire regeneration. Average seed densities of 290 per foot square (27 per m sq) have been reported in western redcedar (Thuja plicata)/pachistima and western redcedar/ queencup beadlily (Clintonia uniflora) habitat types in coniferous forests of northern Idaho [55]. SITE CHARACTERISTICS : The California blackberry occurs across a wide range of sites from warm, open areas to dense woodlands [27,36]. It is particularly common in prairies, clearings, waste places, and canyons [36,55]. California blackberry frequently assumes prominence on sites which have been burned or logged [16,27,36] and on river terraces or gravel bars dominated by red alder (Alnus rubra) [19]. Soils: Blackberries (Rubus spp.) grow well on a variety of barren, infertile soils [6]. These shrubs tolerate a wide range of soil texture and pH but require adequate soil moisture for good growth [10]. California blackberry appears to be tolerant of periodic flooding by brackish or fresh water [65]. Elevation: California blackberry grows from sea level along the Pacific Coast to middle elevations farther inland [27,36]. Generalized elevational ranges for given locations are as follows [9,13]: < 2,000 feet (610 m) in the Santa Monica Mtns., CA < 3,000 feet (914 m) in southern California SUCCESSIONAL STATUS : California blackberry is a vigorous competitor which commonly invades disturbed sites created by logging, fire, or other types of disturbance [16,39]. It is particularly well represented following "catastrophic disturbance" in Douglas-fir forests of the Pacific Northwest [25], and readily established on mudflows and other harsh microsites following the eruption of Mount St. Helens. California blackberry typically increases rapidly on disturbed sites, persisting until suppressed by canopy closure [26]. It occurs in stands of all ages but reaches greatest abundance in early seral communities [16,37,44]. Although primarily an early seral species, California blackberry can sometimes persist in low densities as a residual species in mature forest communities [16,59]. California blackberry was observed in initial postdisturbance, early immature, late immature, mature, and old growth stands in coniferous forests of southwestern British Columbia [44]. This shrub increases rapidly and can dominate the herbaceous layer as early as 2 to 5 years after disturbance [25]. In many western hemlock-western redcedar or Douglas-fir forests of the Pacific Northwest, this shrub remains dominant for at least 20 years after disturbance [2,25]. Bailey [2] found that California blackberry increased to 50 percent cover 4 years after disturbance, fluctuated between 25 and 50 percent cover for 20 years, and declined to 1 percent cover in climax stands. Maximum cover values were reached 15 to 30 years after logging and fire in Douglas-fir plantations in western hemlock-Douglas-fir habitats of western Oregon [59]. California blackberry is present in red alder communities, which on certain upland sites, appear to represent early seral stages of western hemlock forests. Where these communities occur along streambanks, periodic flooding can maintain species such as salmonberry and red alder in long-lived, disclimax situations. California blackberry is considered a major dominant in early successional stages of these communities [33]. SEASONAL DEVELOPMENT : Seasonal development of the California blackberry varies according to geographic and climatic factors. Phenology has been documented as follows [6,9,13,27]: location flowering fruit ripening seed dispersal -- April-June June-August July-September -- April-Aug. -- -- Santa Monica Mts.,CA Feb.-June -- -- s CA March-July -- -- California blackberry remains dormant during the winter [12].


SPECIES: Rubus ursinus
FIRE ECOLOGY OR ADAPTATIONS : California blackberry is a common invader on recently burned sites in the Pacific Northwest [16,39,53]. Populations are capable of dramatic and rapid expansion on disturbed sites [25,66] through sprouting or seedling establishment [9,16]. Belowground regenerative structures are generally well protected from the harmful effects of heat and permit rapid recovery where California blackberry plants were present in the preburn community. Seedbanking is also an important postfire regenerative strategy [52]. Seeds accumulate in the soil or duff, remaining viable long after this seral species has been eliminated from mature forest communities. Seeds commonly germinate in great abundance after fire. The relatively large, sweet, succulent fruit of blackberries amply "reward" animal dispersers [40], and some postfire dispersal of seed from off-site is probable. FIRE REGIMES : Find 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". 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)


SPECIES: Rubus ursinus
IMMEDIATE FIRE EFFECT ON PLANT : California blackberry is described as "rather tolerant" of fire [25]. Although it may be top-killed [52], underground regenerative portions of this shrub generally survive [9,52,61]. Fires of relatively high severity or intensity, with the potential to harm belowground regenerative structures, appear to be the most damaging to California blackberry [52]. Most California blackberry seed stored on-site in the soil or duff is probably unharmed by fire [52]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Vegetative response: California blackberry is capable of vegetative regeneration following fire through nonrhizomatous basal sprouts or root "suckers" [9,52]. Basal sprouting is believed to be of primary importance, and often results in a large number of sprouts [51]. This low-growing shrub can also root at the stem nodes [37,65] and can presumably spread quickly where portions of the aboveground stem remain undamaged. Vegetative spread is generally both vigorous and rapid. Stewart [61] observed an average of approximately 1,520 sprouts (plus some surviving original stems) per acre (3,762/ha) within 2 year after fire. Prior to the fire, an average of only 40 original California blackberry stems had been counted within the same area. Evidence suggests that all forms of sprouting may be favored after fires of relatively low severity or intensity which are unlikely to damage belowground regenerative structures [52]. Expansion of California blackberry may be delayed on heavily burned sites [25,50]. Comparative cover and density values of sprouts on a 2-year-old burn in western redcedar/pachistima and western redcedar/queencup beadlily habitat types in northern Idaho are as follows [50]: low severity high severity % cover 32.5 16.0 density 1.7 1.8 Seedling establishment: Seedbanking is reportedly an important means of postfire reestablishment for the California blackberry [50]. High-severity fires, which burn to mineral soil, frequently create a favorable seedbed for buried blackberry seed, and seedlings sometimes germinate in abundance [52]. However in several instances, researchers have observed decreased seedling establishment after unusually hot fires with "much fuel consumption" [52,64]. Other factors, such as site characteristics or climatic conditions, may have contributed to the variable responses. Seedling establishment of California blackberry can also occur through seed transported from off-site by birds and mammals. Rate of postfire recovery: Sprouting produces the most rapid early growth, as plants draw upon portions of previously established root systems [52]. In many locations California blackberry has exhibited the most rapid postfire expansion of any residual species [25]. This shrub can dominate the herbaceous layer within 2 to 5 years after fire [25,53,66]. Peak cover values have been reported from 0 to 5 years after fire [25,51]. California blackberry is characterized by a relatively long (> 5 years) period of postfire abundance and generally persists until suppressed by canopy closure [25]. California blackberry cover occasionally exhibits a temporary decline after rapid early growth as one or a few stems attain dominance over many initial sprouts [50]. In western redcedar habitat types of northern Idaho, Morgan and Neuenschwander [52] observed highest cover values in the third and fifth years after fire, with California blackberry disappearing by the fifteenth year. However, California blackberry frequently remains abundant for 11 to 16 years or more after fire in the Cascade Mountains of western Oregon [59,60]. Halpern [25] reported that it remained prominent for at least 20 years after fire in seral Douglas-fir forests of western Oregon. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : Fire severity and intensity can influence the rate of postfire recovery in California blackberry regardless of whether regeneration occurs vegetatively or through seedling establishment. The following response has been observed in western redcedar/queencup beadlily habitat types of Idaho [52]: mean frequency of occurrence burn age in years severity 1 2 3 4 5 15 low 33 82 71 74 98 0 high 71 80 68 74 100 0 FIRE MANAGEMENT CONSIDERATIONS : Timber harvest and slash burns: California blackberry commonly invades logged and slash burned sites in the Douglas-fir zone of the Pacific Northwest [39,61]. Reestablishment is rapid and can occur by the second season after treatment [53]. In early postfire years, cover of California blackberry is frequently as much as 3 times higher on slash burned sites than in adjacent undisturbed stands [15]. California blackberry remains prominent until suppressed by the closure of the forest canopy [25]. Schoonmaker and McKee [59] reported the following cover values after clearcutting and broadcast burning in the Cascades of western Oregon: yrs. since tmt. 2 5 10 15 20 30 40 old growth cover(%) 2.28 2.18 1.11 9.7 20.52 7.35 0.66 0.18 Stewart [61] observed similar increases after clearcutting and broadcast burns in Coastal Oregon: years preburn 1 3 4 cover (%) .20 .90 2.40 .20 Competition: California blackberry is favored by fire and can aggressively compete with conifer seedlings in some postfire communities. Wildlife: Species which consume large amounts of blackberries are often benefited by fire [45].


SPECIES: Rubus ursinus
REFERENCES : 1. Atzet, Thomas. 1979. Description and classification of the forests of the upper Illinois River drainage of southwestern Oregon. Corvallis, OR: Oregon State University. 211 p. Dissertation. [6452] 2. Bailey, Arthur Wesley. 1966. Forest associations and secondary succession in the southern Oregon Coast Range. Corvallis, OR: Oregon State University. 166 p. Thesis. [5786] 3. Barber, William Hollis, Jr. 1976. An autecological study of salmonberry (Rubus spectabilis, Pursh) in western Washington. Seattle, WA: University of Washington. 154 p. Thesis. [7189] 4. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434] 5. Bolsinger, Charles L. 1988. The hardwoods of California's timberlands, woodlands, and savannas. Resour. Bull. PNW-RB-148. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 148 p. [5291] 6. Brinkman, Kenneth A. 1974. Rubus L. blackberry, raspberry. In: Schopmeyer, C. S., ed. Seeds of woody plants in the United States. Agriculture Handbook No. 450. Washington, DC: U.S. Department of Agriculture, Forest Service: 738-743. [7743] 7. Conard, Susan G. 1987. First year growth of canyon live oak sprouts following thinning and clearcutting. In: Plumb, Timothy R.; Pillsbury, Norman H., technical coordinators. Proceedings of the symposium on multiple-use management of California's hardwood resources; 1986 November 12-14; San Luis Obispo, CA. Gen. Tech. Rep. PSW-100. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 439. [5392] 8. Conard, Susan G.; Emmingham, W. H. 1983. Herbicides for shrub control on forest sites in northeastern Oregon and northern Idaho. Special Publication 5. Corvallis, OR: Oregon State University, College of Forestry, Forest Research Laboratory. 7 p. [3579] 9. Conrad, C. Eugene. 1987. Common shrubs of chaparral and associated ecosystems of southern California. Gen. Tech. Rep. PSW-99. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 86 p. [4209] 10. Core, Earl L. 1974. Brambles. In: Gill, John D.; Healy, William M., compilers. Shrubs and vines for Northeastern wildlife. Gen. Tech. Rep. NE-9. Broomall, PA: U.S. Department of Agriculture, Forest Service: 16-19. [8923] 11. Crane, M. B. 1940. Reproductive versatility in Rubus. I. Morphology and inheritance. Journal of Genetics. 40: 109-118. [8443] 12. Crouch, Glenn L. 1966. Preferences of black-tailed deer for native forage and Douglas-fir seedlings. Journal of Wildlife Management. 30(3): 471-475. [8881] 13. Dale, Nancy. 1986. Flowering plants: The Santa Monica Mountains, coastal and chaparral regions of southern California. Santa Barbara, CA: Capra Press. In cooperation with: The California Native Plant Society. 239 p. [7605] 14. Dimock, Edward J., II. 1974. Animal populations and damage. In: Cramer, Owen P., ed. Environmental effects of forest residues management in the Pacific Northwest: A state-of-knowledge compendium. Gen. Tech. Rep. PNW-24. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station: O-1 to O-28. [6394] 15. Dyrness, C. T. 1965. The effect of logging and slash burning on understory vegetation in the H. J. Andrews Experimental Forest. Res. Note PNW-31. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 13 p. [4939] 16. Dyrness, C. T. 1973. Early stages of plant succession following logging and burning in the western Cascades of Oregon. Ecology. 54(1): 57-69. [7345] 17. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 18. Fiedler, Peggy Lee; Leidy, Robert A. 1987. Plant communities of Ring Mountain Preserve, Marin County, California. Madrono. 34(3): 173-192. [4068] 19. Fonda, R. W. 1974. Forest succession in relation to river terrace development in Olympic National Park, Washington. Ecology. 55(5): 927-942. [6746] 20. Fonda, R. W. 1979. Fire resilient forests of Douglas-fir in Olympic National Park: a hypothesis. In: Linn, Robert M., ed. Proceedings, 1st conference on scientific research in the National Parks, Vol. 2; 1976 November 9-12; New Orleans, LA. NPS Transactions and Proceedings No. 5. Washington, DC: U.S. Department of the Interior, National Park Service: 1239-1242. [6698] 21. Franklin, Jerry F.; Dyrness, C. T. 1973. Natural vegetation of Oregon and Washington. Gen. Tech. Rep. PNW-8. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 417 p. [961] 22. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998] 23. Gray, M. Violet; Greaves, James M. 1984. Riparian forest as habitat for the least Bell's vireo. In: Warner, Richard E.; Hendrix, Kathleen M., eds. California riparian systems: Ecology, conservation, and productive management: Proceedings of a conference; 1981 September 17-19; Davis, CA. Berkeley, CA: University of California Press: 605-611. [5862] 24. Great Plains Flora Association. 1986. Flora of the Great Plains. Lawrence, KS: University Press of Kansas. 1392 p. [1603] 25. Halpern, C. B. 1989. Early successional patterns of forest species: interactions of life history traits and disturbance. Ecology. 70(3): 704-720. [6829] 26. Halpern, Charles B.; Harmon, Mark E. 1983. Early plant succession on the Muddy River mudflow, Mount St. Helens, Washington. American Midland Naturalist. 110(1): 97-106. [8870] 27. Halverson, Nancy M., compiler. 1986. Major indicator shrubs and herbs on National Forests of western Oregon and southwestern Washington. R6-TM-229. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 180 p. [3233] 28. Halpern, Charles B.; Franklin, Jerry F. 1990. Physiognomic development of Pseudotsuga forests in relation to initial structure and disturbance intensity. Journal of Vegetation Science. 1(4): 475-482. [13288] 29. Harlacher, Richard A. 1987. Determination of planting densities for revegetation projects. In: Rieger, John P.; Williams, Bradford K., eds. Proceedings of the second native plant revegetation symposium; 1987 April 15-18; San Diego, CA. Madison, WI: University of Wisconsin Arboretum, Society for Ecological Restoration & Management: 22-27. [4090] 30. Harper, James A. 1962. Daytime feeding habits of Roosevelt elk on Boyes Prairie, California. Journal of Wildlife Management. 26(1): 97-100. [8876] 31. Heady, Harold F.; Foin, Theodore C.; Hektner, Mary M.; [and others]. 1977. Coastal prairie and northern coastal scrub. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 733-760. [7211] 32. Hemstrom, Miles A.; Logan, Sheila E.; Pavlat, Warren. 1987. Plant association and management guide: Willamette National Forest. R6-Ecol 257-B-86. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 312 p. [13402] 33. Henderson, Jan A. 1978. Plant succession on the Alnus rubra/Rubus spectabilis habitat type in western Oregon. Northwest Science. 52(3): 156-167. [6393] 34. Hines, William W. 1973. Black-tailed deer populations and Douglas-fir reforestation in the Tillamook Burn, Oregon. Game Research Report Number 3. Federal Aid to Wildlife Restoration, Project W-51-R, Final Report. Corvallis, OR: Oregon State Game Commission. 59 p. [8431] 35. Hines, William W.; Land, Charles E. 1974. Black-tailed deer and Douglas-fir regeneration in the Coast Range of Oregon. In: Black, Hugh C., ed. Wildlife and forest management in the Pacific Northwest: Proceedings of a symposium; 1973 September 11-12; Corvallis, OR. Corvallis, OR: Oregon State University, School of Forestry, Forest Research Laboratory: 121-132. [7999] 36. Hitchcock, C. Leo; Cronquist, Arthur. 1973. Flora of the Pacific Northwest. Seattle, WA: University of Washington Press. 730 p. [1168] 37. Ingram, Douglas C. 1931. Vegetative changes and grazing use on Douglas-fir cut-over land. Journal of Agricultural Research. 43(5): 387-417. [8877] 38. Isaac, L. A. 1930. Seedling survival on burned and unburned surfaces. Journal of Forestry. 28: 569-571. [16902] 39. Isaac, Leo A. 1940. Vegetative succession following logging in the Douglas-fir region with special reference to fire. Journal of Forestry. 38: 716-721. [4964] 40. Janzen, Daniel H. 1984. Dispersal of small seeds by big herbivores: foliage is the fruit. American Naturalist. 123(3): 338-353. [6901] 41. Kartesz, John T.; Kartesz, Rosemarie. 1980. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Volume II: The biota of North America. Chapel Hill, NC: The University of North Carolina Press; in confederation with Anne H. Lindsey and C. Richie Bell, North Carolina Botanical Garden. 500 p. [6954] 42. Katibah, Edwin F.; Nedeff, Nicole E.; Dummer, Kevin J. 1984. Summary of riparian vegetation aerial and linear extent measurements from the Central Valley Riparian Mapping Project. In: Warner, Richard E.; Hendrix, Kathleen M., eds. California riparian systems: Ecology, conservation, and productive management: Proceedings of the conference; 1981 September 17-19; Davis, CA. Berkeley, CA: University of California Press: 46-50. [5824] 43. Kelpsas, B. R. 1978. Comparative effects of chemical, fire, and machine site preparation in an Oregon coastal brushfield. Corvallis, OR: Oregon State University. 97 p. Thesis. [6986] 44. Klinka, K.; Scagel, A. M.; Courtin, P. J. 1985. Vegetation relationships among some seral ecosystems in southwestern British Columbia. Canadian Journal of Forestry. 15: 561-569. [5985] 45. Kramp, Betty A.; Patton, David R.; Brady, Ward W. 1983. The effects of fire on wildlife habitat and species. RUN WILD: Wildlife/ habitat relationships. Albuerque, NM: U.S. Department of Agriculture, Forest Service, Southwestern Region, Wildlife Unit Technical Report. 29 p. [152] 46. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384] 47. Laymon, Stephen A. 1984. Photodocumentation of vegetation and landform change on a riparian site, 1880-1980: Dog Island, Red Bluff, California. In: Warner, Richard E.; Hendrix, Kathleen M., eds. California riparian systems: Ecology, conservation, and productive management: Proceedings of a conference; 1981 September 17-19; Davis, CA. Berkeley, CA: University of California Press: 150-159. [5833] 48. Lyon, L. Jack; Stickney, Peter F. 1976. Early vegetal succession following large northern Rocky Mountain wildfires. In: Proceedings, Tall Timbers fire ecology conference and Intermountain Fire Research Council fire and land management symposium; 1974 October 8-10; Missoula, MT. No. 14. Tallahassee, FL: Tall Timbers Research Station: 355-373. [1496] 49. Miller, Daniel L.; Kidd, Frank A. 1982. How to write a herbicide prescription for shrub control. Forestry Technical Paper TP-82-6. Lewiston, ID: Potlatch Corporation, Wood Products, Western Division. 12 p. [3390] 50. Miller, Margaret M.; Miller, Joseph W. 1976. Succession after wildfire in the North Cascades National Park complex. In: Proceedings, annual Tall Timbers fire ecology conference: Pacific Northwest; 1974 October 16-17; Portland, OR. No. 15. Tallahassee, FL: Tall Timbers Research Station: 71-83. [6574] 51. Mitchell, John E. 1983. Overstory-understory relationships: Douglas-fir forests. In: Bartlett, E. T.; Betters, David R., eds. Overstory-understory relationships in western forests. Western Regional Res. Publ. No. 1. Fort Collins, CO: Colorado State University Experiment Station: 27-34. [3314] 52. Morgan, Penelope; Neuenschwander, Leon F. 1988. Shrub response to high and low severity burns following clearcutting in northern Idaho. Western Journal of Applied Forestry. 3(1): 5-9. [3895] 53. Morris, William G. 1958. Influence of slash burning on regeneration, other plant cover, and fire hazard in the Douglas-fir region (A progress report). Res. Pap. PNW-29. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 49 p. [4803] 54. Munz, Philip A. 1973. A California flora and supplement. Berkeley, CA: University of California Press. 1905 p. [6155] 55. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. [4924] 56. Pojar, J.; Klinka, K.; Meidinger, D. V. 1987. Biogeoclimatic ecosystem classification in British Columbia. Forest Ecology and Management. 22: 119-154. [7314] 57. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843] 58. Sawyer, John O.; Thornburgh, Dale A.; Griffin, James R. 1977. Mixed evergreen forest. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 359-381. [7218] 59. Schoonmaker, Peter; McKee, Arthur. 1988. Species composition and diversity during secondary succession of coniferous forests in the western Cascade Mountains of Oregon. Forest Science. 34(4): 960-979. [6214] 60. Steen, Harold K. 1966. Vegetation following slash fires in one western Oregon locality. Northwest Science. 40(3): 113-120. [5671] 61. Stewart, R. E. 1978. Origin and development of vegetation after spraying and burning in a coastal Oregon clearcut. Res. Note PNW-317. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 11 p. [6541] 62. Stromberg, Laurence P.; Katibah, Edwin F. 1984. An application of the spatial-aggregation method to the description of riparian vegetation. In: Warner, Richard E.; Hendrix, Kathleen M., eds. California riparian systems: Ecology, conservation, and productive management: Proceedings of a conference; 1981 September 17-19; Davis, CA. Berkeley, CA: University of California Press: 347-355. [5839] 63. U.S. Department of Agriculture, Soil Conservation Service. 1994. Plants of the U.S.--alphabetical listing. Washington, DC: U.S. Department of Agriculture, Soil Conservation Service. 954 p. [23104] 64. Van Dersal, William R. 1938. Native woody plants of the United States, their erosion-control and wildlife values. Washington, DC: U.S. Department of Agriculture. 362 p. [4240] 65. Willoughby, John W.; Davilla, William. 1984. Plant species composition and life form spectra of tidal streambanks and adjacent riparian woodlands along the lower Sacramento River. In: Warner, Richard E.; Hendrix, Kathleen M., eds. California riparian systems: Ecology, conservation, and productive management: Proceedings of a conference; 1981 September 17-19; Davis, CA. Berkeley, CA: University of California Press: 642-651. [5866] 66. Yerkes, Vern P. 1960. Occurrence of shrubs and herbaceous vegetation after clear cutting old-growth Douglas-fir. Res. Pap. PNW-34. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 12 p. [8937] 67. Stickney, Peter F. 1989. Seral origin of species originating in northern Rocky Mountain forests. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT; RWU 4403 files. 7 p. [20090] 68. U.S. Department of the Interior, National Biological Survey. [n.d.]. NP Flora [Data base]. Davis, CA: U.S. Department of the Interior, National Biological Survey. [23119]