SPECIES: Prunus virginiana


INTRODUCTORY

SPECIES: Prunus virginiana
AUTHORSHIP AND CITATION:
Johnson, Kathleen A. 2000. Prunus virginiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].

ABBREVIATION:
PRUVIR

SYNONYMS:
No entry

NRCS PLANT CODE:
PRVI
PRVID
PRVIM
PRVIV

COMMON NAMES:
chokecherry
western chokecherry
common chokecherry
black chokecherry

TAXONOMY:
The fully documented scientific name of chokecherry is Prunus virginiana L. (Rosaceae) [58,72,73,80]. Recognized varieties are:

Prunus virginiana var. demissa (Nutt.) Torr. - western chokecherry [73,80]
Prunus virginiana var. melanocarpa (A. Nels.) Sarg. - black chokecherry [80]
Prunus virginiana var. virginiana L. - common chokecherry [73,80]

LIFE FORM:
Tree-shrub

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
No entry


DISTRIBUTION AND OCCURRENCE

SPECIES: Prunus virginiana
GENERAL DISTRIBUTION:
Chokecherry is widely distributed throughout southern Canada and much of the United States. It occurs from Newfoundland to British Columbia and south to North Carolina, Tennessee, Missouri, Kansas, Oklahoma, Texas, New Mexico, California, and northern Mexico. Plants showing a gradation from black chokecherry to common chokecherry occur in Kansas and Nebraska. The three varieties are distributed as follows [59,72,73]:

common chokecherry - eastern variety; occurs from Saskatchewan to Newfoundland southward to Kansas, Missouri, Tennessee and North Carolina

black chokecherry - restricted to the western portion of North America; occurs in southern Canada from eastern British Columbia to Alberta and the Dakotas; southward throughout the Rocky Mountains to New Mexico; and along the east slope of the Cascade Range to northern California

western chokecherry - occurs from British Columbia southward into northern Mexico, Texas, and California (except the coast and Central Valley)

ECOSYSTEMS:
FRES10 White-red-jack pine
FRES11 Spruce-fir
FRES13 Loblolly-shortleaf pine
FRES14 Oak-pine
FRES15 Oak-hickory
FRES17 Elm-ash-cottonwood
FRES18 Maple-beech-birch
FRES19 Aspen-birch
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES23 Fir-spruce
FRES26 Lodgepole pine
FRES28 Western hardwoods
FRES29 Sagebrush
FRES30 Desert shrub
FRES34 Chaparral-mountain shrub
FRES35 Pinyon-juniper
FRES36 Mountain grasslands
FRES38 Plains grasslands
FRES39 Prairie

STATES:
AK AZ AR CA CO CT DE GA ID IL
IN IA KS KY ME MD MA MI MN MO
MT NE NV NH NJ NM NY NC ND OH
OK OR PA RI SD TN TX UT VT VA
WA WV WI WY DC

AB BC MB ON PQ SK

MEXICO

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
7 Lower Basin and Range
8 Northern Rocky Mountains
9 Middle Rocky Mountains
10 Wyoming Basin
11 Southern Rocky Mountains
12 Colorado Plateau
13 Rocky Mountain Piedmont
14 Great Plains
15 Black Hills Uplift
16 Upper Missouri Basin and Broken Lands

KUCHLER PLANT ASSOCIATIONS:
K002 Cedar-hemlock-Douglas-fir forest
K003 Silver fir-Douglas-fir forest
K005 Mixed conifer 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
K018 Pine-Douglas-fir forest
K019 Arizona pine forest
K020 Spruce-fir-Douglas-fir forest
K021 Southwestern spruce-fir forest
K023 Juniper-pinyon woodland
K024 Juniper steppe woodland
K026 Oregon oakwoods
K028 Mosaic of K002 and K026
K029 California mixed evergreen forest
K030 California oakwoods
K033 Chaparral
K034 Montane chaparral
K037 Mountain-mahogany-oak scrub
K038 Great Basin sagebrush
K051 Wheatgrass-bluegrass
K055 Sagebrush steppe
K063 Foothills prairie
K064 Grama-needlegrass-wheatgrass
K065 Grama-buffalo grass
K066 Wheatgrass-needlegrass
K067 Wheatgrass-bluestem-needlegrass
K068 Wheatgrass-grama-buffalo grass
K074 Bluestem prairie
K081 Oak savanna
K095 Great Lakes pine forest
K097 Southeastern spruce-fir forest
K098 Northern floodplain forest
K099 Maple-basswood forest
K100 Oak-hickory
K101 Elm-ash forest
K102 Beech-maple forest
K103 Mixed mesophytic forest
K106 Northern hardwoods
K107 Northern hardwoods-fir forest
K108 Northern hardwoods-spruce forest
K111 Oak-hickory-pine
K112 Southern mixed forest

SAF COVER TYPES:
1 Jack pine
5 Balsam fir
14 Northern pin oak
15 Red pine
16 Aspen
17 Pin cherry
18 Paper birch
19 Gray birch-red maple
20 White pine-northern red oak-red maple
21 Eastern white pine
22 White pine-hemlock
23 Eastern hemlock
24 Hemlock-yellow birch
25 Sugar maple-beech-yellow birch
26 Sugar maple-basswood
27 Sugar maple
28 Black cherry-maple
40 Post oak-blackjack oak
42 Bur oak
60 Beech-sugar maple
63 Cottonwood
80 Loblolly pine-shortleaf pine
81 Loblolly pine
82 Loblolly pine-hardwood
93 Sugarberry-American elm-green ash
108 Red maple
206 Engelmann spruce-subalpine fir
210 Interior Douglas-fir
211 White fir
212 Western larch
213 Grand fir
216 Blue spruce
217 Aspen
218 Lodgepole pine
219 Limber pine
220 Rocky Mountain juniper
222 Black cottonwood-willow
229 Pacific Douglas-fir
230 Douglas-fir-western hemlock
231 Port-Orford-cedar
233 Oregon white oak
234 Douglas-fir-tanoak-Pacific madrone
235 Cottonwood-willow
236 Bur oak
237 Interior ponderosa pine
238 Western juniper
239 Pinyon-juniper
243 Sierra Nevada mixed conifer
245 Pacific ponderosa pine
244 Pacific ponderosa pine-Douglas-fir
247 Jeffrey pine

SRM (RANGELAND) COVER TYPES:
107 Western juniper/big sagebrush/bluebunch wheatgrass
109 Ponderosa pine shrubland
201 Blue oak woodland
203 Riparian woodland
206 Chamise chaparral
207 Scrub oak mixed chaparral
208 Ceanothus mixed chaparral
209 Montane shrubland
210 Bitterbrush
302 Bluebunch wheatgrass-Sandberg bluegrass
303 Bluebunch wheatgrass-western wheatgrass
306 Idaho fescue-slender wheatgrass
312 Rough fescue-Idaho fescue
314 Big sagebrush-bluebunch wheatgrass
315 Big sagebrush-Idaho fescue
317 Bitterbrush-bluebunch wheatgrass
322 Curlleaf mountain-mahogany-bluebunch wheatgrass
401 Basin big sagebrush
402 Mountain big sagebrush
403 Wyoming big sagebrush
404 Black sagebrush
405 Low sagebrush
408 Other sagebrush types
409 Tall forb
411 Aspen woodland
412 Juniper-pinyon woodland
413 Gambel oak
415 Curlleaf mountain-mahogany
418 Bigtooth maple
419 Bittercherry
420 Snowbrush
421 Chokecherry-serviceberry-rose
422 Riparian
503 Arizona chaparral
504 Juniper-pinyon pine woodland
603 Prairie sandreed-needlegrass
606 Wheatgrass-bluestem-needlegrass
607 Wheatgrass-needlegrass
608 Wheatgrass-grama-needlegrass
710 Bluestem prairie

HABITAT TYPES AND PLANT COMMUNITIES:
The 3 chokecherry varieties occur in numerous habitat types and plant communities. Chokecherry often forms mixed stands with other tall shrubs. Common plant associates of chokecherry in some areas are listed below by state or province.

Idaho: Associated tall shrubs on logged sites in a northern Idaho western redcedar-western hemlock (Thuja plicata-Tsuga heterophylla) zone include Rocky Mountain maple (Acer glabrum), Saskatoon serviceberry (Amelanchier alnifolia), redstem ceanothus (Ceanothus sanguineus), snowbrush ceanothus (C. velutinus), dogwood (Cornus spp.), oceanspray (Holodiscus discolor), mockorange (Philadelphus lewisii), ninebark (Physocarpus malvaceus), bitter cherry (Prunus emarginata), cascara (Rhamnus purshiana), Scouler willow (Salix scouleriana), and red elderberry (Sambucus racemosa) [173]. In southern and central Idaho, chokecherry occurs in a number of Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca) habitat types, along with Pacific ponderosa pine (Pinus ponderosa var. ponderosa), Rocky Mountain maple and quaking aspen (Populus tremuloides) [143].

Michigan: Chokecherry occurs in northern Lake Michigan coastal sand dune communities that range from 175 to 835 years old, but is most prevalent in communities 225 to 400 years old. Associated overstory dominants in mixed-pine forests in a similar 200-400 year range include balsam fir (Abies balsamifera), paper birch (Betula papyrifera), red pine (Pinus resinosa), white pine (Pinus strobus), white spruce (Picea strobus), and northern white-cedar (Thuja occidentalis) [99]. In northern white-cedar forests in lower northern Michigan, important plant associates are sugar maple (Acer saccharum), white ash (Fraxinus americana), mountain maple (Acer spicatum), paper birch, basswood (Tilia americana), alternate-leaved dogwood (C. alternifolia), ironwood (Ostrya virginiana) and balsam fir [3].

Minnesota: Northeastern forests: overstory species include quaking aspen, bigtooth aspen (Populus grandidentata), red pine, and jack pine (Pinus banksiana). Common tall shrub species include mountain maple, American green alder (Alnus viridis ssp. crispa), alternate-leaf dogwood, round-leaved dogwood (C. rugosa) and serviceberry (Amelanchier spp.) [13].

In the northwestern forest and transition zones of Minnesota, overstory associates include bur oak (Quercus macrocarpa), American elm (Ulmus americana), basswood, sugar maple, green ash, quaking aspen, paper birch, ironwood, and balsam poplar (Populus balsamifera). Common shrub associates include smooth sumac (Rhus glabra), dogwood (Cornus spp.), black cherry (Prunus serotina), and sugar maple. [31,47].

In Minnesota oak (Quercus spp.) savanna overstory dominants include bur oak and pin oak (Q. ellipsoidalis). Shrubs commonly associated with chokecherry include smooth sumac and American hazel (Corylus americana) [169].

Montana: Plant associates in riparian sites in western and central Montana include Rocky Mountain Douglas-fir, Rocky Mountain juniper (Juniperus scopulorum), Pacific ponderosa pine, big sagebrush (Artemisia tridentata), western snowberry (Symphoricarpos occidentalis) and ninebark [46,66,119]. In eastern Montana hardwood forests that extend into the Dakotas, chokecherry occurs commonly with green ash, plains cottonwood (Populus deltoides var. monilifera), American elm, and box elder (Acer negundo) [64,66,98,119].

Nevada and Utah; Dominant associated shrubs in sagebrush (Artemisia spp.) rangelands in northeastern Nevada and mountain brush communities in Utah include Saskatoon serviceberry, shadscale (Atriplex confertifolia), rubber rabbitbrush (Chrysothamnus nauseosus), green rabbitbrush (C. viscidiflorus), antelope bitterbrush (Purshia tridentata), mountain snowberry (Symphoricarpos oreophilus), Gambel oak (Quercus gambelii), Wood's rose (Rosa woodsii), ninebark, curlleaf mountain-mahogany (Cercocarpus ledifolius), Rocky Mountain juniper, Rocky Mountain Douglas-fir, and white fir (Abies concolor). [11,33,79,90].

North Carolina: In a red spruce-Fraser fir (Picea rubens-Abies fraseri) forest in the Plott Balsam Mountains, chokecherry occurs with pin cherry, American mountain-ash (Sorbus americana), mountain maple, alternate-leaf dogwood, red maple (Acer rubrum), and willow (Salix spp.) species [128].

North Dakota: Chokecherry occurs in Missouri river slopes, floodplains, and also western woodlands. Tree associates are green ash (Fraxinus pennsylvanica), box elder, American elm, bur oak, basswood, quaking aspen, paper birch, Rocky Mountain juniper, ponderosa pine, and limber pine (Pinus flexilis). Common shrubs and woody vines in the floodplains include peach-leaved willow (S. amygdaloides), dogwood, western snowberry (Symphoricarpos occidentalis), eastern poison-ivy (Toxicodendron radicans), Saskatoon serviceberry, woodbine (Parthenocissus inserta), and frost grape (Vitis vulpina) [56,64,82,98,171].

Pennsylvania: In central Pennsylvania mixed-oak valley floor forests, dominant species associated with chokecherry are white oak (Q. alba), red oak (Q. rubra), black oak (Q. velutina) and black cherry [2].

South Dakota and Wyoming: Hoffman and Alexander [77] describe a ponderosa pine (P. ponderosa var. scopulorum/chokecherry community type occurring on the Black Hills National Forest. Important associates in this community are Saskatoon serviceberry, white spirea (Spirea betulifolia), and Oregon-grape (Mahonia repens). The authors list chokecherry as an important component in bur oak, ponderosa pine and quaking aspen series of habitat types. River drainage species in green ash/chokecherry habitats are similar to those listed for North Dakota.

Vermont: In the Green Mountains chokecherry occurs where common overstory trees are sugar maple, American beech (Fagus grandifolia), yellow birch (Betula alleghaniensis), red spruce, and balsam fir. Abundant understory shrub associates include striped maple (Acer pensylvanicum), mountain maple, pin cherry (P. pensylvanica), and American mountain-ash [136].

Wisconsin: In southwestern oak-hickory (Quercus-Carya spp.) forests (where replacement of oaks by others species is a recognized problem), chokecherry occurs where dominant overstory species include sugar maple, green ash, and slippery elm (Ulmus rubra) [100].

Chokecherry is listed as a dominant or indicator species in the following published classifications:

A preliminary classification of the natural vegetation of Colorado [12]
Vegetation and soils of the Rock Springs Watershed [21]
Native woodland ecology and habitat classification of southwestern North Dakota [57]
The vegetation of the Grand River/Cedar River, Sioux, and Ashland Districts of the Custer National Forest: a habitat type classification [64]
The vegetation of Theodore Roosevelt National Park, North Dakota: a habitat type classification [65]
Classification and Management of Montana's riparian and wetland sites [66]
Grassland, shrubland, and forestland habitat types of the White River-Arapaho National Forest [71]
Habitat types on selected parts of the Gunnison and Uncompahgre National Forests [85]
Aspen community types on the Caribou and Targhee National Forests in southeastern Idaho [115]
Forest habitat types of Montana [119]
Rangeland cover types of the United States [135]
Plant associations (habitat types) of Region 2., 3rd ed. [157]
Aspen community types on the Bridger-Teton National Forest in western Wyoming [178]

MANAGEMENT CONSIDERATIONS

SPECIES: Prunus virginiana
WOOD PRODUCTS VALUE:
Chokecherry wood is heavy, hard and close-grained, but rarely develops a trunk large enough to be commercially useful [116].

IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Chokecherry is widely regarded as an important wildlife food plant and provides habitat, watershed protection, and species diversity [36,92,121,140,147]. Fruits, leaves, and twigs are utilized. Large mammals including bears [34,83,124], moose [69,118,120,144], coyotes [38], bighorn sheep [154,160], pronghorn [41], elk [32,49,74,88,94], and deer use chokecherry as browse [7,8,26,132,145,146,147,148,149,150,156]. Chokecherry is also a food source for small mammals [37,38,62,70]. The fruits are important food for many birds [15,20,54,103]. Cattle and domestic sheep also eat chokecherry, and because of its toxicity (see below), poisoning sometimes occurs. Livestock normally do not eat fatal quantities except when other forage is scarce [66,116,166].

PALATABILITY:
Chokecherry is moderately palatable to all classes of livestock, although it is more heavily browsed by domestic sheep than by cattle [36,140]. It is a preferred mule deer browse on many winter ranges throughout the Intermountain West and Northern Great Plains [39,42].

The palatability of chokecherry to livestock and wildlife has been rated as follows [42]:
                     CO     MT     ND     UT     WY
Cattle               Fair   Fair   Fair   Fair   Fair
Domestic sheep       Good   Good   Good   Fair   Good
Horses               Poor   Poor   ----   Poor   Poor
Pronghorn            ----   ----   Good   Poor   Poor
Elk                  Poor   Fair   ----   Good   Fair
Mule deer            Fair   Fair   Good   Good   Good
Small mammals        Good   Good   ----   Good   Good
Small nongame birds  Good   Good   ----   Good   Good
Upland game birds    Fair   Good   Good   Good   Good
Waterfowl            ----   ----   ----   Poor   Poor

NUTRITIONAL VALUE:
The nutritional value of chokecherry is relatively high in comparison with that of other western browse species [19,88,168]. It has 38.8% dry matter digestibility and is 8.7% crude protein [35]. Crude protein levels do not appreciably decrease as winter progresses [39]. Dittberner and Olson [42] rate chokecherry good in energy value and poor in protein value. Seasonal trends in the nutritive content of chokecherry leaves and stems in the Black Hills of South Dakota are presented below. Units are in percent, on an oven-dry basis [39].
                Spring          Summer         Fall       
                leaves/stems    leaves/stems   leaves/stems
crude protein   21.9/17.4       15.2/9.5        6.6/8.8     
cellulose       12.3/19.7       12.6/22.8      14.7/24.2     
ash              5.9/5.4         6.2/4.3        6.1/3.0    
calcium         1.12/0.9         1.8/1.5       2.33/1.66     
phosphorus      0.51/0.41       0.39/0.21      0.37/0.21     
COVER VALUE:
Chokecherry provides important cover and habitat for many bird species [115,116,117,118,119,120,121,122,123,124], small mammals [4,44,55,139,146,165], large mammals, and livestock [16,24,45,65,66,159]. Chokecherry is an excellent shrub for providing thermal cover and erosion control in fisheries [66]. The degree to which chokecherry provides cover for wildlife species is as follows [42]:
                     CO     MT     ND     UT     WY
Pronghorn            ----   Fair   Good   Poor   Fair 
Elk                  Good   Fair   ----   Good   Good
Mule deer            Good   Good   Good   Good   Good
White-tailed deer    Good   Good   Good   ----   Good
Small mammals        Good   Good   ----   Good   Good
Small nongame birds  Good   Good   Good   Good   Good
Upland game birds    Good   Good   Good   Good   Good
Waterfowl            ----   ----   ----   Poor   Poor
VALUE FOR REHABILITATION OF DISTURBED SITES:
Chokecherry has been selected as a revegetation species for wildlife habitat [110,127,166], shelterbelts [131], mine spoils [109], and soil stabilization [102,104,131]. Chokecherry exhibited salt tolerance in a greenhouse study [155].

Chokecherry can be propagated from seed or rhizome cuttings [60,91,102,108,129,134,166]. About half of unstratified seed germinates within 60 days of collection; delayed germination can occur up to 120 days of sowing [166]. More consistent germination is achieved following cool, moist stratification lasting from 120 to 160 days at 36 to 41 degrees Fahrenheit (2.2-5 oC) [60,108,109]. In a heat-treatment field study, using thermocouples inserted into seedcoats, chokecherry germination was doubled to quadrupled by temperatures ranging from 180 to 280 degrees Fahrenheit (82-138 oC)[127].

Nursery-grown seedlings of chokecherry establish satisfactorily if planted free of competition on sites with at least 15 inches (38 cm) of annual precipitation. Young plants are not tolerant of competing vegetation for 2 to 3 years following planting [110].

OTHER USES AND VALUES:
Chokecherry plants are widely used as ornamentals. Chokecherry produces an abundance of attractive white flowers characterized by a strong, sweet, almondlike fragrance. This species is also valued for its fruit. Plantings increase habitat and natural food supplies for birds frequenting residential areas. Chokecherry is extensively planted for windbreaks in the prairie, plains, and western mountains [166]. Chokecherries are edible and, although somewhat astringent, are relatively sweet when fully ripe. Fruits are used to make wines, syrups, jellies, and jams. Indigenous peoples gathered chokecherries and used them to make pemmican and treat cold sores [68,84,147]. The Piutes made a medicinal tea from the leaves and twigs to treat colds and rheumatism [113].

OTHER MANAGEMENT CONSIDERATIONS:
Grazing: Chokecherry is moderately tolerant of browsing [66], but heavy grazing by livestock and wild ungulates has impacted populations in many areas, especially the northern Great Plains [64,65,81,98,179].

Chokecherry foliage can be poisonous to grazing livestock [78,106,116,153]. Research has identified the toxic compound in chokecherry as the cyanogenic glycoside prunasin [101]. One-half gram of prunasin can produce approximately 46 mg of hydrogen cyanide (HCN); daily doses of 50 mg HCN/1 kg body weight are considered dangerous. Hydrogen cyanide is liberated either in the plant as a result of frost damage or in the animal during digestion. Results of that study indicated that prunasin concentrations are highest (5%) in the new stems and newly initiated leaves of chokecherry. Elevated levels (greater than 2.5%) are maintained in the leaves throughout the summer, but prunasin content of new twigs gradually diminishes over the season. The previous season's growth is generally not as toxic (1.2 to 2.2%) [166].

Pest management concerns: Chokecherry can be controlled by herbicides or plowing [116]. In the northeastern United States chokecherry is a primary host of the eastern tent caterpillar [164].

Chokecherry is susceptible to attack by the fungus Plowrightia stansburiana, which causes knotlike cankers to develop on stems. This condition eventually kills infected stems [92]. Afflicted plants usually have a shortened life span [166].


BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Prunus virginiana
GENERAL BOTANICAL CHARACTERISTICS:
Chokecherry is a native, deciduous, thicket-forming erect shrub or small tree. Stems are numerous and slender, either branching from the base or with main branches upright and spreading [166]. Heights vary considerably according to variety and site quality, ranging from 3 to 19.5 feet (1-6 m) [73]. In the Great Basin, chokecherry may grow to almost 40 feet (12 m) with trunk diameters of approximately 8 inches (20 cm) [92]. Perfect flowers are borne on leafy twigs of the season. Fruits are drupes, each containing a small stone [73]. Chokecherries have a network of rhizomes and a deep root system established at intervals along the rhizomes [129,140,175]. Roots may extend laterally more than 35 feet (10.6 m) and vertically more than 6 feet (1.8 m) [175]. Rhizomes range from 0.4 to 0.8 inch (1-2 cm) in diameter [129].

RAUNKIAER LIFE FORM:
Phanerophyte

REGENERATION PROCESSES:
Chokecherry reproduces sexually and vegetatively. In laboratory experiments involving excised rhizomes which were approximately 11 years of age, Schier [129] observed that chokecherry rhizomes sprouted at a faster rate and had higher sprouting percentages than Gambel oak (Quercus gambelii) rhizomes.

Seed crops are typically regular and viable [111], with seed-producing capacity higher in plants on open sites [166]. Seeds are surrounded by a stony endocarp that may offer some resistance to germination but is permeable to moisture. Chokecherry has seed dormancy; an after-ripening period in the presence of oxygen and moisture is necessary for adequate germination [60].

Rogers and Applegate [124] reported significantly (P <0.01) enhanced germination in chokecherry seeds ingested by black bears in Minnesota and attributed this to acid and mechanical scarification of seeds in the digestive tract.

Although large numbers of chokecherry seeds may be deposited beneath parent plants, long-distance dispersal also occurs via frugivorous birds and mammals [86,167,170,172]. Meyer and Witmer [107] studied the effect of gut-scarification on chokecherry seeds fed to captive frugivorous birds. Removal of fruit pulp was critical for germination, but they found no differences in germination success between seeds manually cleaned of pulp and bird-passed seeds lacking pulp. Seeds of chokecherry that were defecated and planted with feces, mimicking natural deposition, had reduced germination relative to manually cleaned seeds. Artificial seed treatments to enhance germination are discussed in the Value and Use section of this report.

Viable seed persists in the soil seedbank. In a closed-canopy forest in northern Idaho, chokecherry seeds were found in soil samples taken at depths of up to 4 inches (10 cm); overall seed viability equaled approximately 27% [86].

SITE CHARACTERISTICS:
In the western United States, chokecherry grows at low to mid-elevations in positions in the landscape where combinations of soil and topography permit greater than average accumulation of moisture. These sites include riparian areas, wooded draws, and steep ravines [43,64,65,66,149,163]. Soils supporting chokecherry are variable, ranging from Entisols to Mollisols, and soil texture ranges from silt to sandy loam. Chokecherry can tolerate weakly saline soils but is intolerant of poor drainage and prolonged flooding [66].

Chokecherry grows in very acid to moderately alkaline soils. In the green ash/chokecherry habitat type in the northern Great Plains, pH ranged from 6.0 to 7.6 in loam, clay, and clay loam [65,163]. In deciduous forests in Vermont, glacial till soils supporting chokecherry had a pH ranging from 3.5 to 5.5 [136]. Where chokecherry occurred in forests growing on coastal Lake Michigan sand dunes, pH ranged from approximately 4.0 to 6.0 [99]. In Pennsylvania, soils in oak-pine supporting chokecherry grew on well-drained limestone residuum soil and had a pH range from 4.8 to 5.4 [2].

Elevational ranges for chokecherry are:

Idaho: 3,100 to 8,000 feet (945-2440 m) [143,173]
Michigan: 580 to 738 feet (177-225 m) [99]
Montana: 580 to 738 feet (177-225 m) [46,66,119]
Nevada and Utah: 4,986 to 10,170 feet (1520-3100 m) [11,33,79,90]
North Dakota: 1,023 to 1,095 feet (312-334 m) [171]
South Dakota: 3,002 to 3,494 feet (915-1065 m) [77]
Vermont: 1,797 to 2,798 feet (548-853 m) [136]

SUCCESSIONAL STATUS:
Because chokecherry occurs so widely, it is reported in numerous habitat types and plant associations that range from post-disturbance invaders to early-successional to climax or stable. It grows in sparse stands, dense thickets, and under open forest canopies [64,66,116,119]. It is shade tolerant [31,64,99,100], but reaches its greatest density near forest edges [64,65]. Plant association descriptions for most the studies discussed below appear in the Distribution and Occurrence section of this species report.

EASTERN AND CENTRAL REGION-
In the eastern and central region of the U.S. and Canada, chokecherry occurs in a broad range of successional habitats. It been studied in both seral and climax or stable communities. It sprouts readily and also persists under open and closed forest canopies.

Chokecherry was characterized as early-successional following logging and (or) burning in northern white-cedar and jack pine forests in Michigan [1]. Chokecherry was mid-successional in coastal Lake Michigan chronosequences that focused on long-term vegetation succession on sand dunes, reaching its greatest abundance after pine and oak cover was well developed [99,117].

In Pennsylvania mixed-oak forests chokecherry was described as a later-successional understory species where the overstory is dominated by oak and pine [2].

In a 1924 study of succession in northwestern Minnesota chokecherry was prominent in non-climax brush stands in ecotones between prairie and deciduous forests [47]. In a 1951 study in northern Minnesota chokecherry was noted as a major component in the sparse shrub layer of a climax maple-basswood forest. The maple-basswood forest was characterized as having light penetration of less than 5%, indicating shade tolerance, at least in mature chokecherry [31].

In a 1964 study in Vermont, chokecherry was one of 14 shrub species documented in the understory of an old, undisturbed remnant of northern hardwood forest. Sugar maple, beech, and white ash were overstory dominants. The authors predicted that this assemblage replaces itself in forest succession, creating a stable overstory community [23].

In Ontario, following forest clearcutting and brush removal for a utility right-of-way, chokecherry was a prominent initial colonizer. The original forest cover was dominated by sugar maple, white ash, quaking aspen and black cherry. In addition to abundant chokecherry seedlings, raspberry (Rubus spp.) seedlings and sprouts, and sprouts from quaking aspen were also prolific initial colonizers. After 6 years chokecherry and quaking aspen stems had declined and white ash stems had become most numerous. The author indicated that rapid invasion by chokecherry and raspberry, followed by replacement with more shade-tolerant species was a common pattern of secondary succession [27].

In southern and western North Dakota chokecherry is an indicator species for the green ash/chokecherry woodland habitat type [64,65]. This habitat type is characterized as a topographic climax. Where disturbance from livestock grazing is heavy, shrub cover is greatly reduced and unpalatable western snowberry becomes dominant. Tree seedlings and saplings decline, leaving only older trees and an open understory [65]. In southern North Dakota chokecherry is an indicator species for the interior ponderosa pine/chokecherry habitat type, determined to be an edaphic climax. In undisturbed vegetation of this type, ponderosa pine forms a closed overstory and chokecherry shrubs reach about 3.28 feet (1 m) in height. Chokecherry responds to fire in these stands by sprouting vegetatively, and as succession advances following fire, chokecherry gradually increases while other shrubs decrease [64]. The green ash/chokecherry and interior ponderosa pine/chokecherry habitat types also occur in South Dakota and eastern Montana [64,66,77,119].

WESTERN STATES-
In the western United States, chokecherry is usually identified as seral but persists under closed canopies in mature conifer forests and in riparian areas. Central Rocky Mountain quaking aspen stands, where chokecherry is prevalent in the tall shrub layer, are thought to be a regional transition zone between sporadic groves and extensive forests. The quaking aspen/chokecherry community is categorized as seral [114].

In a classification of forest habitats of Montana [119], chokecherry is documented in numerous habitat types within forest climax series for limber pine, ponderosa pine, Douglas-fir, and Engelmann spruce. In that classification it is also an indicator species for a ponderosa pine/chokecherry type present in eastern Montana (see discussion in North Dakota section above). In river drainages of central Montana, Eichorn and Watts [46] studied succession following wildfire. In burned north-slope sites characterized by a Douglas-fir/common juniper association, chokecherry, snowberry (Symphoricarpos spp.) and rose (Rosa spp.) were predominant among shrubs that increased significantly (P<0.05) in years 5 through 28 following burning. In Douglas-fir habitat types in western Montana, chokecherry becomes common after stand-replacing wildfires and clearcuts with or without subsequent broadcast burns [8].

In central Idaho chokecherry is classified by Steele and Geier-Hayes [143] as mid-seral in 7 Douglas-fir habitat types. Though seral to Douglas-fir climax forests, these sites may be dominated by open-canopy stands of fire-maintained ponderosa pine. Chokecherry may regenerate vegetatively or by seed following logging or burning, depending on the type and severity of the disturbance.

In a central Utah study, Christensen [33] reported that although mountain brush stands are often interpreted as stable, chokecherry was among 10 shrub species in a mountain brush stand undergoing transition to a conifer stand dominated by white fir and Douglas-fir. The author interpreted the transition as primary succession because no evidence was found of disturbance by livestock use, fire or logging. Chokecherry is common in Gambel oak communities in Utah, which Kunzler and others [90] predicted might succeed to ponderosa pine, bigtooth maple (Acer grandidentatum), or white fir and Douglas-fir, depending on site conditions. Chokecherry is common in northern Utah quaking aspen stands, which are reported to succeed to conifer stands [14]. In a study of early succession following clearcutting of quaking aspen, chokecherry and snowberry were "by far" the most dominant shrubs in uncut control plots. In the 4 years following clearcutting, percent composition of chokecherry in the undergrowth increased by much as 5 times over that in the control plots. The authors did not indicate the method of chokecherry regeneration in the clearcuts [14].

SEASONAL DEVELOPMENT:
Generally chokecherry plants leaf out in spring to early summer and flower 1 to 3 weeks later, with fruits maturing in late summer to fall [166]. Fruits dehisce soon after maturity [59]. Average date of phenological stages for chokecherry east and west of the Continental Divide in Montana from 1928 to 1937 are presented below [130]:
                           East Divide      West Divide
leaf buds burst            May 2            April 29
leaves full grown          June 11          May 17
flowers start              June 4           May 19 
flowers end                June 17          June 11
fruits ripe                August 22        August 14
leaves start to color      August 31        September 15
leaves begin to fall       September 10     September 28
seed fall starts           September 12     September 19
leaves fallen/withered     September 30     October 14

FIRE ECOLOGY

SPECIES: Prunus virginiana
FIRE ECOLOGY OR ADAPTATIONS:
Chokecherry is well adapted to disturbance by fire [5,25,52,97,105,174,177]. Although susceptible to to top-kill by fire, it resprouts rapidly and prolifically from surviving root crowns and rhizomes [51,97,105,162,166]. Several studies reporting chokecherry recovery by sprouting are discussed in the Fire Effects section of this report. Seed germination improves with heat treatment, suggesting scarification by fire is an important adaptation [127]. Postfire regeneration probably also involves the germination of off-site seed dispersed by mammals and birds [162].

No data were found for natural intervals of fire in stands that consist mainly of chokecherry. Gartner [52] provides a description of pre and post-settlement accounts of fire in the grasslands and ponderosa forests of the Black Hills of South Dakota. The historical information is detailed, but fire return intervals are not given. Hansen [63] provides limited historical and fire interval information for Minnesota forests in Isle Royale National Park, Itasca State Park, and the Boundary Waters Canoe Area. Twenty-six lighting fires were recorded on Isle Royale from 1965 to 1949. In Itasca State Park the incidence of fires caused by lighting or set by Indians before 1859 averaged about one fire every 12 years. In the Boundary Waters Canoe Area, major fires recurred at 5- to 50-year intervals from 1600 to 1920.

Fire regimes for other plant communities in which chokecherry occurs are summarized below. For further information regarding fire regimes and fire ecology of communities where chokecherry is found, see the Fire Ecology and Adaptations section of the FEIS species summary for the plant community or ecosystem dominants.

Community or Ecosystem Scientific Name of Dominant Species Mean Fire Return Interval
Pacific ponderosa pine* Pinus ponderosa var. ponderosa 1-47 yrs
Rocky Mountain ponderosa pine* P. ponderosa var. scopulorum 2-10 yrs [30]
Rocky Mountain lodgepole pine* P. contorta var. latifolia 50-300+ yrs [6,125]
Colorado pinyon P. edulis 10-49 yrs
Rocky Mountain Douglas-fir* Pseudotsuga menziesii var. glauca 40-140 yrs [30]
Wyoming big sagebrush Artemisia tridentata var. wyomingensis 10-70 yrs [161,176]
mountain big sagebrush A. tridentata var. vaseyana 5-15 yrs
curlleaf mountain-mahogany* Cercocarpus ledifolius 13-1,350 yrs [9,132]
quaking aspen (west of the Great Plains) Populus tremuloides 7-100 yrs [61,106]
*fire return intervals vary widely; trends in variation are noted in the FEIS species summary

POSTFIRE REGENERATION STRATEGY:
Tree with adventitious-bud root crown/soboliferous species root sucker
Tall shrub, adventitious-bud root crown
Small shrub, adventitious-bud root crown
Ground residual colonizer (on-site, initial community)
Initial-offsite colonizer (off-site, initial community)

FIRE EFFECTS

SPECIES: Prunus virginiana
IMMEDIATE FIRE EFFECT ON PLANT:
Fire often kills aboveground chokecherry stems and foliage, but it quickly sprouts, either the same year following a spring burn, or by the next growing season [97,105,162,166,177]. In the South Dakota Black Hills chokecherry sprouts were double the preburn numbers within 2 months of an early May burn [51]. Conversely, in an early May prescribed burn in central Alberta quaking aspen parkland, chokecherry shrubs did not sprout within the first 3 months following burning [63]. Fire intensity was not described for either study.

A prescribed fire study was conducted in northern Idaho to test the effect of spring versus fall burning on elk browse. Measurements were made of crown diameter, crown height, number of basal sprouts, and sprout height before and after each burn. Postfire measurements were made on 11 shrubs the first growing season after the fall burns. Seasonal fire effects were similar for chokecherry crown diameter, crown height, and sprout height. Though not statistically significant (at P = 0.05), the number of chokecherry basal sprouts was somewhat higher after the spring burn, suggesting that spring burning may be more conducive to the rapid recovery of chokecherry than fall burning [97].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
No entry

PLANT RESPONSE TO FIRE:
Most studies report either an increase in chokecherry in the years following fire, or an increase followed by a return to prefire numbers. After wildfires in the oakbrush zone in Utah, McKell [105] reported twice as many chokecherry stems sprouting from root crowns on 1-year-old burns than on adjacent unburned sites. A reduction to prefire densities occurred within 18 years.

Following wildfires in Rocky Mountain Douglas-fir/Rocky Mountain juniper/Wyoming big sagebrush associations in the Missouri Breaks area of central Montana, chokecherry canopy cover increased consistently for 13 years, then stabilized [46].

Bock and Bock [22] compared data from prescribed October burns in 1974 and 1979 in the South Dakota Black Hills. The 1974 burn escaped and became a crown fire, killing ponderosa pines of all sizes. The 1979 fire remained a controlled understory fire. When measured in 1981, the 1974 burn site supported higher densities of all woody taxa except chokecherry and western poison-ivy (Toxicodendron rydbergii). There was no significant difference (P = 0.71) in numbers of chokecherry plants between the 2 burn sites. Measurements taken within the 1979 surface fire site (prefire, postfire yr 1, and postfire yr 2) showed that chokecherry stems were not significantly (P = 0.75) reduced by the fire.

Geier-Hayes [53] included chokecherry in a study of vegetation response to helicopter logging and broadcast burning in an Idaho Douglas-fir forest. Data were collected in 3 cutting units prior to burning and 1,2,5 and 10 years after. Fire severity was higher in 2 of the units and altered the vegetation from the original. Fires classed at a severity level of 2M [126] were less severe and had little or no impact on chokecherry percent cover and root frequency. 2M fires are characterized as having a flame length of 2 to 4 feet (0.6-1.2 m) and a corresponding crown scorch height of 9 to 24 feet (2.7-7.3 m), with moderate ground charring. The units that burned hotter, having a severity rating of 3M had markedly less chokecherry cover and root frequency during all postburn years measured. 3M fires have flame lengths of 8 to 12 feet (2.4-3.7 m), corresponding crown scorch to 64 feet (20 m), with moderate ground charring.

Following a September prescribed burn in a quaking aspen stand in Idaho, chokecherry biomass exceeded preburn biomass within 2 seasons and was double preburn biomass after 5 seasons [29]. Biomass was computed using weight versus stem diameter relationships [28]. Details of fuel conditions are provided: litter and woody material moisture content was 8 to 9% and herbaceous vegetation was 40 to 50% cured. Fire severity was rated as moderate to high [126].

After 24 years of annual early spring burning in quaking aspen parklands in Alberta, chokecherry percent cover had decreased but the number of stems increased in density from 6% to 15% [5].

In a 20-year study of the effects of fire frequency on Minnesota oak savanna herbs and shrubs, Tester [150,151] determined that increased fire frequency tended to increase the density of native, true prairie shrubs and decrease the density of native, non-prairie shrubs (including chokecherry). Chokecherry cover estimates were negatively correlated with burn frequency (r = -0.51, P = 0.09).

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
The Research Project Summaries Understory recovery after low- and high-intensity fires in northern Idaho ponderosa pine forests and Understory recovery after burning and reburning quaking aspen stands in central Alberta, and the Research Paper by Bowles and others 2007 provide information on prescribed fire and postfire response of several plant species including chokecherry.

FIRE MANAGEMENT CONSIDERATIONS:
Chokecherry is a component of persistent, fire-maintained seral shrubfields on steep slopes in Northern Idaho. Fuels in shrubfields differ in quantity and distribution from those on forested sites. Herbaceous and large woody fuels are relatively light. Live and dead shrub biomass, which includes chokecherry, can reach nearly 20 tons per acre. After fires, which are severe during summer drought conditions, dense shrub cover regenerates within 10 years. Trees regenerate slowly or not at all on these dry sites, because of erosion, depleted soil organic matter, high soil temperatures, and lack of seed [138].

Arno [7] hypothesized that relatively frequent fires set by Native Americans in western grassland and sagebrush communities, where chokecherry occurs, favored expansion of grasslands into adjacent shrub or tree communities. In recent times shrub and tree communities have developed in former grasslands due to fire exclusion and grazing. Arno argues that baseline information on Native American use of fire will aid land managers in predicting vegetative development under different fire regimes.

Morber and Miyanishi [112] studied fire as a tool for controlling chokecherry and black cherry in Ontario oak savanna. A controlled spring burn had no effect on chokecherry seedlings. Postfire seedling emergence was concluded to be largely dependent on postfire seed production or seed influx from adjacent unburned areas, because there was no viable soil seedbank.


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