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Introductory

SPECIES: Cornus sericea | Red-Osier Dogwood
ABBREVIATION : CORSER SYNONYMS : Cornus stolonifera Cornus alba var. occidentalis Cornus occidentalis Cornus alba var. baileyi Cornus alba var. californica Cornus alba var. interior Cornus alba var. coloradense Cornus baileyi Cornus instolonea Cornus interior Cornus sanguinea Svida stolonifera Cornus pubescens Cornus californica Suida interior Ossea interior Suida stolonifera var. riparia Cornus instoloneus Cornus nelsoni Cornus amomum SCS PLANT CODE : COST4 COMMON NAMES : red-osier dogwood western dogwood American dogwood redstem dogwood red dogwood kinnikinnik squawbush creek dogwood California dogwood red-stemmed cornel redbrush gutter tree red willow harts rouges poison dogwood shoemack waxberry cornel dogberry tree redosier dogwood TAXONOMY : The currently accepted scientific name of red-osier dogwood is Cornus sericea L. Although the name "Cornus stolonifera Michx." is in wide current use [45,60] and the merits of both names have been debated [40,112], a recent, extensive review of work with the genus gives the correct name as C. sericea [34]. The discussion of how closely related forms in the Cornaceae should be segregated has gone on for years and is summarized by Eyde [33,34] and Ferguson [36,37]. Two main groups within Cornus are red-line dogwoods, with showy bracts below the flowers and red fruit, and blue-line dogwoods, without bracts and blue or white fruit [34]. Other authors have placed the bractless dogwoods, including red-osier dogwood, in a separate genus called Thleycrania, Ossea, or Svida (originally Swjda, also spelled Swida and Suida) [33,36,37]. Red-osier dogwood is very widespread and variable; thus similar populations have been considered variously as separate but interbreeding species, subspecies, and varieties [40,60,69]. Recognized subspecies of Cornus sericea are as follows [69]: ssp. occidentalis ssp. sericea The two subspecies interbreed quite freely and C. X californica is the name given resultant plants. C. X acadiensis is a hybrid of C. sericea and C. alternifolia [112]. C. X slavini is a hybrid of C. sericea and C. rugosa [143]. There are relatively slight differences between the two subspecies and this paper considers the entire complex as red-osier dogwood. LIFE FORM : Tree, Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : M. F. Crane, August 1989 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Crane, M. F. 1989. Cornus sericea. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Cornus sericea | Red-Osier Dogwood
GENERAL DISTRIBUTION : Red-osier dogwood occurs from Alaska and the Yukon Territory east to Labrador and Newfoundland. It extends south to Virginia in the East, to Kansas in the Great Plains, to northern Mexico in the Rocky Mountains, and through California on the West Coast [52,53,54,71,100]. In the northeastern and midwestern United States it is common in previously glaciated areas; south of these areas it occurs locally where site conditions are favorable [142]. ECOSYSTEMS : FRES10 White - red - jack pine FRES11 Spruce - fir 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 FRES27 Redwood FRES28 Western hardwoods FRES29 Sagebrush FRES34 Chaparral - mountain shrub FRES35 Pinyon - juniper FRES36 Mountain grasslands FRES37 Mountain meadows FRES38 Plains grasslands FRES39 Prairie STATES : AL AZ CA CO CT ID IL IN IA ME MA MI MN MT NE NV NH NJ NM NY ND OH OR PA RI SD UT VT VA WA WV WI WY AB BC LB MB NB NF NT NS ON PE PQ SK ADMINISTRATIVE UNITS : ACAD ALPO APIS BAND BICA BLCA BLRI BRCA CACH CANY CARE CODA CRLA CRMO CUVA DETO DINO EFMO FLFO GATE GWMP GLAC GRCA GRTE GRBA GRKO GRSA INDU ISRO JODA KICA LACL LAVO MORA MORU NOCA PIRO ROMO SAGU SEQU SLBE THRO TICA PIPE VOYA WACA WICA WRST YELL YUCH ZION BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains 3 Southern Pacific Border 4 Sierra Mountains 5 Columbia Plateau 6 Upper Basin and Range 8 Northern Rocky Mountains 9 Middle Rocky Mountains 10 Wyoming Basin 11 Southern Rocky Mountains 12 Colorado Plateau 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 K003 Silver fir - Douglas-fir forest K004 Fir - hemlock forest K005 Mixed conifer forest K006 Redwood forest K008 Lodgepole pine - subalpine forest K010 Ponderosa shrub forest K011 Western ponderosa forest K012 Douglas-fir forest K013 Cedar - hemlock - pine forest K014 Grand fir - Douglas-fir forest K015 Western spruce - fir forest K016 Eastern ponderosa forest K017 Black Hills pine forest K020 Spruce - fir - Douglas-fir forest K023 Juniper - pinyon woodland K029 California mixed evergreen forest K037 Mountain mahogany - oak scrub K038 Great Basin sagebrush K055 Sagebrush steppe K056 Wheatgrass - needlegrass shrubsteppe K063 Foothills prairie K066 Wheatgrass - needlegrass K067 wheatgrass - bluestem - needlegrass K070 Sandsage - bluestem prairie K074 Bluestem prairie K081 Oak savanna K093 Great Lakes spruce - fir forest K094 Conifer bog K095 Great Lakes pine forest K096 Northeastern spruce - fir forest K098 Northern floodplain forest K099 Maple - basswood 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 16 Aspen 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 62 Silver maple - American elm 107 White spruce 109 Hawthorn 201 White spruce 202 White spruce - paper birch 204 Black spruce 205 Mountain hemlock 206 Engelmann spruce - subalpine fir 210 Interior Douglas-fir 211 White fir 212 Western larch 213 Grand fir 215 Western white pine 216 Blue spruce 217 Aspen 218 Lodgepole pine 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 229 Pacific Douglas-fir 232 Redwood 235 Cottonwood - willow 236 Bur oak 237 Interior ponderosa pine 239 Pinyon - juniper 244 Pacific ponderosa pine - Douglas-fir 251 White spruce - aspen 253 Black spruce - white spruce 254 Black spruce - paper birch 256 California mixed subalpine SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Red-osier dogwood occurs most frequently as a riparian species within various forest, woodland, and grassland habitat types. It is a seral species in moist forest habitats and persists in openings. Published classification schemes listing red-osier dogwood as an indicator species or a dominant part of the vegetation in community types (cts), habitat types (hts), plant associations (pas), riparian zone associations (rzas), forest ecosystem associations (eas), dominance types (dts), or riparian site types (rst) are presented below. Area Classification Authority e ID, w WY riparian cts Youngblood & others 1985a CO: White River, grassland, shrubland Hess & Wasser 1982 Arapaho NFs and forest hts NM: Cibola NF forest hts Alexander & others 1987 n CO, s WY forest pas Johnston 1985 wc AB forest eas Corns & Annas 1986 MT riparian dts Hansen & others 1988 sw MT riparian rst, cts, hts Hansen & others 1989 c,e MT riparian rst, cts, hts Hansen & others 1989 central MT wetland cts Pierce and Johnson 1986 WY rst Olson and Gerhart 1982 WY shrubland cts Collins 1984a Region 2: WY, gen. veg. hts, pas Wasser and Hess 1982 SD, NE, CO, KS Region 2: WY, gen. veg. pas Johnston 1987 SD, NE, SO, KS n UT, s ID riparian cts Youngblood and others 1985b s UT riparian cts Padgett & Youngblood 1986 sw MT riparian rst, cts, hts Hanson and others 1989 nNM, nAZ forest and woodland Larson & Moir 1987 hts, pas n NM, s CO forest hts DeVelice and others 1986 CO: Gunnison and general veg. hts, cts Komarkova 1986 Uncompahgre NFs MI, WI forest hts Coffman and others 1980 nw BC forest eas Haeussler and others 1985 c ID riparian cts, hts Tuhy and Jensen 1982 Saint Lawrence general veg. pas Dansereau 1959 Valley

VALUE AND USE

SPECIES: Cornus sericea | Red-Osier Dogwood
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Red-osier dogwood is used for food and cover by white-tailed deer, mule deer, elk, moose, mountain goats, cottontail rabbits, snowshoe hares, and numerous birds, including the bobwhite, ring-necked pheasant, wild turkey, and grouse [41,48,82,89,105,138,155]. Red-osier dogwood fruit is low in sugar so it is initially less attractive to wildlife and less inclined to rot than other fruits. Consequently, the fruit stays on the plant through the winter and is availabile when fruits of other plants are gone [133]. In the northern Rockies, its fruit is a key grizzly [156] and black bear food [115]. Fruit of red-osier dogwood is also eaten by songbirds, grouse, quail, partridge, cutthroat trout, ducks, crows, mice and other mammals [34,143]. Deer mice, meadow voles, and other small rodents feed on the young stems and bark [104]. Beavers use it for food and to build dams and lodges [23,54,88,103]. Red-osier dogwood is particularly important to moose in the winter; it is also used in the summer and in the fall when leaves that have escaped frost are particularly favored [61,73,135]. In a Minnesota study, moose used some red-osier dogwood during the summer, but primary use was in the fall after the stems had reddened [65,105]. Although it remained important, it was not used as much in the winter, possibly because some plants were covered by deep snow [65,105]. Red-osier dogwood is valuable winter forage for elk [43,82. In Minnesota, white-tailed deer browse it in April and May [65]. In the western United States and Canada, mule deer use is heavy in the summer and moderate in the fall and winter [83]. Livestock eat it, but it is not a preferred species [28,118]. A 2-year study of summer cattle use of a riparian zone in Oregon found use of red-osier dogwood was relatively heavy the first year and light the second year [114]. PALATABILITY : Leaves of red-osier dogwood are relatively unpalatable to livestock, but the young sprouts are palatable to livestock and deer [93,118]. A study of foliage use by captive mule deer found red-osier dogwood to be second only to Scouler willow (Salix scouleriana) in palability, with the highest use from August through September [124]. In Montana, mule deer apparently prefer red-osier dogwood, since its occurrence in the diet exceeds its abundance [30]. It is palatable to elk in northern Idaho, although not abundant in the area [89]. For white-tailed deer in the Black Hills palatability varies by season [59]: Palatability January to March high April to June low July to September high October to December medium Red-osier dogwood is a preferred moose browse in central and eastern North America [51,79,105,155]. In the Intermountain West, it is highly palatable to moose [61,105,132]. In Quebec, red-osier dogwood, willows (Salix spp.), and mountain ash (Sorbus scopulina) are given the highest palatability ratings for moose [105], and in Manitoba it is a preferred browse species [156]. The palatability of red-osier dogwood for wildlife in several western states is rated as follows [28,118]: CA CO MT ND WY Cattle poor poor fair poor poor Sheep poor fair fair poor fair Horses useless poor poor poor fair Goats good-fair ---- ---- ---- ---- Pronghorn ---- ---- ---- poor poor Elk ---- good poor ---- good Mule deer ---- fair good poor good White-tailed deer ---- ---- fair good good Deer good-fair ---- ---- ---- ---- Small mammals ---- ---- ---- poor good Small nongame birds ---- ---- fair fair good Upland game birds ---- ---- fair ---- good Waterfowl ---- ---- ---- ---- fair NUTRITIONAL VALUE : Red-osier dogwood has been rated fair in energy value and poor in protein value in some western states [28]. The fruit is low in lipids and sugars and classified as "low quality" [132]. Tannins in the leaves appear to inhibit protein availability, but there appears to be little or no inhibition of protein availability in the stems browsed in the winter [114]. The following crude protein content and digestibilities were found in red-osier leaves fed to mule deer [114]: Crude protein content Apparent protein Digestible protein (% of dry matter) digestibility (%) in feed (g/100 g) 13.44 16.27 2.19 COVER VALUE : Red-osier dogwood provides valuable cover for birds and other small animals, especially where it grows in thickets [124,149]. In Arizona red-osier dogwood, along with willows, blueberry elder (Sambucus cerulea), Rocky Mountain maple (Acer glabrum), and thin-leaf alder (Alnus incana ssp. tenuifolia), provides nesting habitat for the dusky flycatcher, MacGillivary warbler, orange-crowned warbler, broad-tailed hummingbird, white-crowned sparrow, and Lincoln sparrow [10]. Red-osier dogwood provides cover and shade that cools water temperatures in streams for trout [124]. In the Pacific Northwest, red-osier dogwood and other riparian species provide good mule deer fawning and fawn-rearing areas in addition to good year-round security and thermal cover [13,83]. The degree to which red-osier dogwood provides environmental protection during one or more seasons for wildlife species is as follows [28]: CO MT ND WY Pronghorn ---- ---- poor poor Elk ---- fair ---- fair Mule deer ---- fair poor good White-tailed deer ---- fair good good Small mammals fair fair fair good Small nongame birds fair fair fair good Upland game birds ---- fair ---- good Waterfowl ---- ---- ---- fair VALUE FOR REHABILITATION OF DISTURBED SITES : Red-osier dogwood is recommended for rehabilitating moist sites within its range. It is well adapted to disturbed sites, excellent at stabilizing soil, easy to establish, and grows rapidly [130,158]. It need fresh, aerated water to establish and may be particularly useful in stabilizing eroding streambanks [52,86]. Its high tolerance for oil could make it useful on oil-damaged sites [148]. Rooted cuttings or nursery-grown seedlings are easily established on moist, well-drained soils and grows rapidly [54,148]. Seed production and handling are described as "very good" [158]. On Intermountain sites, it is moderately easy to establish from seed and spreads from seed at a moderate rate; it also establishes readily from transplants and spreads rapidly [158]]. Based on reports from Colorado, Wyoming, Montana, and North Dakota, red-osier dogwood's growth is best on gentle slopes [28]. Its establishment requirements are moderate to high, and its potential biomass production is moderate. It has moderate potential for use in erosion control and greater potential for long-term revegetation than for short-term revegetation [28]. Studies of rehabilitation along a Columbia River impoundment in Washington found 5-year-old plantings of red-osier dogwood to be very successful based on survival, growth, and use by wildlife for forage or cover [12]. In the Tahoe Basin, establishment was fair to good and long-term survival was good; red-osier dogwood did best on moist sites with better soils [123]. On northwestern Montana roadcuts, red-osier dogwood's survival and growth was good. Although plants on drier south-facing slopes did not develop as well as those on moister sites, they grew and survived much better than expected [64]. Cuttings of red-osier dogwood root easily without treatment and can be directly planted providing sufficient moisture is available [29]. One hundred percent of cuttings taken in early August were rooted in 5 weeks and 90 percent of those taken in mid-April were rooted in 8 weeks [29]. Red-osier dogwood also readily layers [126]. Rooted stock was recommended for a Utah forest where land managers had difficulty stabilizing areas damaged by mudslides and flooding [146]. Rooting of red-osier dogwood cuttings was slowed slightly by endomycorrhizal infection in one study, but overall rooting was not affected [95]. Grasses decrease red-osier dogwood growth in containers by as much as 72 percent [145]. OTHER USES AND VALUES : Red-osier dogwood is an attractive landscaping plant [7,134]. In the winter its deep red stems and twigs provide color, in the spring it produces many creamy white flowers followed by attractive white fruits, and in the fall its leaves turn a spectacular maroon [7]. Once established, it is drought tolerant [7] and, for gardeners in rural areas, it is less palatable to white-tailed deer and mule deer than many other ornamental shrubs [5,22]. The long slim stems were used by Indians for basket weaving and are still used by present-day crafters [72]. Indians and early settlers smoked the inner bark, stem scrapings, and leaves, which have a slightly narcotic effect [72,93]. In Southern California the inner green cambium layers were peeled, dried, and smoked ceremonially [23]. Indians also used an extract from red-osier dogwood as an emetic for treating fevers and coughs and obtained dyes from the bark and roots [93]. MANAGEMENT CONSIDERATIONS : Red-osier dogwood is especially valuable for protecting and stabilizing streambanks because of its thick, extensive root system [52,54,153]. This is particularly important to management of higher gradient stream channels where scouring by seasonal flooding occurs. In a Lake States study, red-osier dogwood did not hold up under heavy use but did well with light clipping [2]. Other studies indicate that red-osier dogwood may increase with some browsing [107]. Following 61 percent leader use by cattle in Oregon, red-osier dogwood responded with exceptional growth the following year when it had light use, and should continue to do well with alternating years of moderate and light use [114]. However, heavy and prolonged grazing in Montana ponderosa pine (Pinus ponderosa) or cottonwood (Populus spp.) and red-osier dogwood riparian communities may eliminate the dogwood and convert the area to a Kentucky bluegrass (Poa pratensis) understory [54]. Heavy grazing by Wisconsin dairy cows eliminated red-osier dogwood from streambanks [149]. In southwestern Montana, after moose hedging of red-osier dogwood increased 80 to 100 percent in 2 years, forage production and plant vigor became very low [132]. Moose browsing on Isle Royale has reduced red-osier dogwood abundance and height [113,128]. Red-osier dogwood is most valuable to beaver in its early growth stages when it is resprouting after fire or other disturbance [152]. Managers find that marsh sites are excellent for beaver production but that the beaver impoundments raise the water level to a point where red-osier dogwood and other food plants are killed. The problem is solved naturally when the beaver abandon the site and the plants regenerate, but to hasten succession and provide suitable beaver habitat, red-osier dogwood cuttings can be planted near the new pond edge [103]. Herbicides designed to weed ornamental crops also injured up to 20 percent of the tissue of red-osier dogwood plants [145]. Red-osier dogwood can be controlled by spraying with mixtures of 2,4-D and 2,4,5-T or dicamba and either 2,4-D or 2,3,4-T [125]. Treatment with 2,4-D or paraquat causes chlorosis and necrosis of leaves and stems and delays spring growth [25]. Plants treated with 2,4-D and paraquat were affected much less when they were treated after vegetative maturity (the stage of dormancy preceding visible leaf senescence) than when treated earlier in the year [25].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Cornus sericea | Red-Osier Dogwood
GENERAL BOTANICAL CHARACTERISTICS : Red-osier dogwood is a deciduous, many-stemmed shrub which varies in height from 3 to 19 feet (1-6 m) [60,142]. The young stems and twigs are dark red, gradually fading to gray-green, and becoming red again in the fall and winter [60,142]. The leaves are opposite with prominent lateral veins that curve toward the tip and smooth edges [93]. Many, small white flowers are borne in a flat-topped cyme and, unlike many dogwoods, there are no large, showy bracts. The flowers are followed by berrylike fruits that are white or lead colored at maturity [126,143]. Red-osier dogwood's wide range and ability to tolerate extremely cold temperatures (laboratory temperatures as low as -320 degrees F [-196 degrees C]) [96,126] have prompted a number of studies of its physiology and cold acclimation [14,15,16,17,42,56,63,74,75,76,90,91,102,119, 120,139]. Red-osier dogwood avoids freezing injury caused by ice forming within living protoplasm by having freezable water frozen extracellularly [96]. The factors that seem to affect cold acclimation the most are low temperatures, short days, water stress, and the developmental stage of the plant [15,75,76,159]. Key points in the developmental cycle, chilling requirements, and temperature effects have been described and modeled [75,76,119,120,159]. Far red light, characteristic of the long twilights at high latitudes, and short day length promote cold acclimation [90]. Water-stressed plants have an increased tolerance of freezing and increase their freezing point [16]. In plants exposed to short days, tissue changes occur that reduce the plant's ability to take up water and simultaneously increase water loss so that the plant partially dehydrates even when water is plentiful [91,102]. A study of different geographic races found that stem pubescence and leaf size and form vary, and plant form can vary from very upright to decumbent between races, although there are no clear patterns [126]. Growth rates of more northern races are slower under shorter photoperiods than those of southern races growing in their own geographic areas. Total growth of northern races was reduced because they stopped growth and started cold acclimation earlier than more southern races [126]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Natural regeneration of red-osier dogwood is both sexual and asexual [60,126]. The flowers of red-osier dogwood are self-sterile and outcrossing is obligate [34,47]. This is controlled by a single, multiallelic gene that inhibits full pollen tube growth in the style of the plant [62]. Pollinators include the honey bee, bumble bee, solitary bee [47] and possibly beetles, flies, and butterflies [34]. Seeds are dispersed primarily by songbirds, although other animals including bears, mice, grouse, quail, partridges, and even ducks and cutthroat trout may eat the fruit and disperse seeds [34,126,143]. The seeds may be stored in seedbanks [117]. Individual plants generally first bear fruit at 3 to 4 years of age, but older plants are more prolific [125]. Red-osier dogwood seeds have dormant embryos and need cold stratification for 1 to 3 months [9,55,121]. Sometimes hard seed coats are also present and scarification is then necessary [125]. Germination rates increased after passage through a black bear's digestive tract [115] but were inconsistant after passage through a pheasant's digestive tract [80]. The seeds will remain viable in cold storage for 4 to 8 years [9]. Details of seed collection and nursery germination can be found in several studies [9,80,121,144,148]. On good sites red-osier dogwood can form dense thickets through vegetative reproduction [48,52]. Red-osier dogwood spreads by layering when the lower stems touch or lie along the ground and root at the nodes [48,60]. In the northeastern United States, production of new plants from stolons is most likely for plants in very moist situations and wet meadows [125]. Plants may also produce new shoots from the roots and new branches from the bases of dying branches [125]. If 27 percent or more of the stem is girdled by small rodents, the stem will die back to the injury, and new growth begins below that point on the stem [104]. Red-osier dogwood resprouted promptly from the roots in riparian zones in the airfall area and more slowly in the devastated zone following the eruption of Mount St. Helens [92]. SITE CHARACTERISTICS : Red-osier dogwood is a characteristic species of swamps, low meadows, and riparian zones; it is also found in forest openings, open forest understories, and along forest margins [48,142]. It is a facultative wetland plant [111]. It is found on warmer, more productive sites in the taiga [141], in rich swamps [57], and generally on very rich, very moist sites [66,116]. Red-osier dogwood's southern limits appear to be determined by high temperatures [125]. Near its southern limit in New Mexico, it is only a riparian species [27], and in other southwestern states and the southern Great Plains, it is primarily a riparian species [46,70,85,143,147]. In the Sacramento-San Joaquin Delta of California, it grows just above the tidal zone and on hummocks and higher areas of marsh in oxbows and overflow basins [21,151]. In bluestem (Andropogon spp.) prairie in Minnesota, red-osier dogwood grows in sedge (Carex spp.) dominated swales that are inundated during the spring [31]. Although red-osier dogwood grows on a variety of soils, it prefers rich, moist soils [48,56,125,150]. Ratings in several western states give growth on gravel as fair to poor; growth on sand, sandy-loam, and loam as good; growth on clay-loam growth as fair to good; and growth on clay or dense clay as poor [28]. Opinions differ on its pH preference which is given as 5.5 to 7.5 [149] or 7.0 to 8.0 [138]. It needs high levels of mineral nutrients for vigorous growth [57,149]. Growth on acidic soils is rated as fair in Montana and Wyoming but poor in Colorado [28]. Growth on organic soils, saline soils, sodic soils, and sodic-saline soils in Montana, Wyoming, and Colorado is rated as poor, and optimum soil depth is given as over 20 inches (51 cm) [28]. Soil temperatures can change root morphology and the mineral content of the stem and leaves of red-osier dogwood [6]. The best root growth occurs at lower soil temperatures than the best shoot growth [6]. Red-osier dogwood can tolerate flooding and, consequently, is found on floodplains and wetlands and is often one of the first shrubs to invade wet meadows [48,125,129]. Its seeds germinate above water level, but after several years growth, the plants can live with the roots submerged in water for most of the growing season [129]. Plants on such wet sites are found in mineral rich swamps or fens and not in ombrotrophic, sphagnum bogs [57]. Elevation: Elevational ranges in several western states are as follows [28,71,143]: Minimum Maximum feet meters feet meters Arizona 5,000 1,524 9,000 2,743 Colorado 4,500 1,372 10,000 3,048 Montana 3,400 1,036 6,600 2,012 Utah 4,800 1,463 9,500 2,896 Wyoming 5,500 1,676 8,300 2,530 Associates: The plants most closely associated with red-osier dogwood are willows and alders (Alnus spp.). Other plants frequently found with red-osier dogwood include cottonwoods, aspen (Populus tremuloides), birch (Betula spp.), Wood's rose (Rosa woodsii), gooseberries (Ribes spp.), hawthorne (Crataegus spp.), horsetails (Equisetum spp.), thistle (Cirsium spp.), and Kentucky bluegrass (Poa pratensis). SUCCESSIONAL STATUS : Red-osier dogwood is an early to mid seral species [11,140,148] that is supressed in shade and is not normally found in the understory of closed canopy forests [125]. It is found in the understory of mixed open forests [48]. Red-osier dogwood needs moderate to full sunlight [11]. A shade-frame study found that red-osier dogwood grew best in 75 percent of full light intensity. Its natural occurrence in full sunlight may be facilitated by its growth in wet situations where it encounters no water stress [122]. Red-osier dogwood is a dominant understory shrub in the early successional willow and balsam poplar (Populus balsamifera) communities that follow oxbow lakes in Alberta, and it has 60 to 75 percent canopy cover in these communities [140]. It is also the most important species in the seral shrub-carrs of Wisconsin that succeed wet prairies, fens, or sedge meadows and are followed by lowland forest or conifer swamp [148]. Red-osier dogwood is not a primary invader in these areas. It enters the stand later in succession and is least common in greatly disturbed stands [148]. Red-osier dogwood is also a successional species in this area following fire or when a bog basin is partially drained by stream downcutting [21]. In Ontario sugar maple-beech (Acer saccharum-Fagus grandifolia) and eastern hemlock-yellow birch (Tsuga canadensis-Betula lutea) forests, red-osier dogwood is a pioneer in open and disturbed areas [11]. In New York wetland forest gaps caused by American elm (Ulmus americana) mortality, red-osier dogwood is frequent and abundant in multiple-tree gaps but rare beneath the closed canopy and in single-tree gaps [160]. SEASONAL DEVELOPMENT : One consequence of red-osier dogwood's wide geographic range has been the development of geographic races with different seasonal responses. A comparison of 21 clones from different areas grown together in a Minnesota plot found that spring events, such as bud break, flowering, and greening of the bark are, determined by temperature, and all of the clones responded to the same temperature [126]. In contrast fall bud set, red bark color, leaf abscission, and cold acclimation were determined by photoperiod, and the geographic races responded to different photoperiods [8,126]. This indicated genetic control and genetic differences between the races. Not all the races were able to acclimate in time to avoid winter cold injury in Minnesota, but once cold acclimated, all the races were able to withstand -130 degrees F (-90 degrees C) without injury. Another study [8], however, indicates that clones from Montana, Oregon, and Idaho have different hardiness levels than clones from North Dakota and Washington. Comparisons of the timing of fall events between different climatic races of red-osier dogwood growing at the same location in Minnesota are as follows (Onset of rest is the point in the growth cycle beyond which plants or cuttings moved into a favorable environment will not grow) [127]: Onset of Rest Red bark Color 50% leaf Abscission Seattle, Wash. Oct. 15 Nov. 2 Nov. 6 Cadillac, Mich. Aug. 18 Oct. 10 Oct. 31 Moscow, Idaho Aug. 31 Oct. 10 Oct. 31 Wayland, Mass. Sept. 17 Oct. 4 Oct. 31 Excelsior, Minn. Aug. 30 Oct. 10 Oct. 27 Ottawa, Ontario Aug. 18 Sept. 20 Oct. 27 Madison, Wisc. Sept. 2 Oct. 3 Oct. 27 Dickinson, ND Aug. 10 Sept. 20 Oct. 21 College, Alaska Aug. 10 Aug. 10 Oct. 17 Dropmore, Manitoba Aug. 10 Sept. 19 Oct. 14 Flowering times for red-osier dogwood in some western states are as follows [28]: Utah Colorado Wyoming Montana North Dakota Begining of Anthesis: May May June May May Anthesis: June June July July June End of Anthesis July July August July July Comparisons of spring phenology in different parts of the country include [9,47,99,131]: Leafing Out Blooming Fruit ripe Massachusetts May mid-July n Minnesota June-August n Idaho late April-May May-June late July-August Great Plains June August-September

FIRE ECOLOGY

SPECIES: Cornus sericea | Red-Osier Dogwood
FIRE ECOLOGY OR ADAPTATIONS : Red-osier dogwood is able to sprout from surviving roots or stolons and from the base of aerial stems following fire [38,39,101,149]. It can be killed by severe fires which cause extended heating of the upper soil [38]. Red-osier dogwood is considered to be a semi-fire-tolerant, seed-banking species [117]. Light fires which partially remove the duff stimulate germination of buried seed [48]. In a northern Idaho grand fir (Abies grandis) forest seed bank study, red-osier dogwood seed was found in the top 2 inches (5 cm) of soil; however, viability was low (4%) [77,78]. In another postfire soil germination study, all red-osier dogwood plants were sprouts from root fragments [1]. POSTFIRE REGENERATION STRATEGY : survivor species; on-site surviving roots ground-stored residual colonizer; fire-activated seed on-site in soil

FIRE EFFECTS

SPECIES: Cornus sericea | Red-Osier Dogwood
IMMEDIATE FIRE EFFECT ON PLANT : Aboveground foliage of red-osier dogwood is usually killed by fire. However, the roots will survive all but the most severe fires which remove the duff and heat the upper soil for extended periods [38]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Red-osier dogwood generally increases following fire [1,18,39,106], and it may invade a recently burned area from adjacent unburned areas [38]. It may take some time before resprouting red-osier dogwood regains its former cover and volume. A Montana study in aspen found that 2 years after prescribed burning red-osier dogwood resprouts had attained 72 percent of their prefire cover and 54 percent of their prefire volume, while density was back to prefire levels [161]. In moist forests of British Columbia, red-osier dogwood appears to increase in abundance following logging and burning [32,50]. It established in a logged and burned Manitoba black spruce (Picea mariana) stand by the fifth postfire year [18]. On Minnesota black spruce sites most fires stimulate sprouting of red-osier dogwood, although severe fires favor tree seedlings [1]. A study in the cedar-hemlock (Thuja spp.-Tsuga spp.) zone of northern Idaho found no red-osier dogwood in closed stands. It established with very slight frequency (1%) in logged stands without fire; somewhat higher frequency (5%) in areas with both single and multiple broadcast burns; and highest frequency (12%) in areas that were piled and burned [94]. In a northwestern Montana subalpine fir/queencup beadlily (Abies lasiocarpa/Clintonia uniflora) habitat type, red-osier dogwood cover was highest (15%) in stands that had burned 35 to 70 years ago and very low in clearcuts (0-1.4%) whether dozer piled or not [156,157]. In Wisconsin shrub-carrs (wet ground tall-shrub communities) light to medium fires cause resprouting in red-osier dogwood and serve to maintain the shrub-carr [149]. Following prescribed burning in central Wisconsin shrub-invaded sedge meadows, red-osier dogwood resprouted and was favored over other shrubs [162]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : In the northern Rockies red-osier dogwood is a common member of the seral brushfields which occur following fire and compete with tree seedlings [98,109]. In order to reduce brushfields on sites that are away from streams and floodplains, the use of logging methods which cause a relatively high amount of site disturbance followed by a fire which removes most of the soil organic horizons is recommended [109]. In northwestern Montana, clearcutting alone apparantly dislodged enough roots of red-osier dogwood so that few plants were able to sprout [156,157]. Postfire sprouts in the early stages of growth are the most valuable for beaver [103,152]. Following fire in Minnesota, red-osier dogwood became more important to moose and white-tailed deer, even though it grew only near streams and was not abundant within the burn [65].

REFERENCES

SPECIES: Cornus sericea | Red-Osier Dogwood
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