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Gaylussacia baccata

• INTRODUCTORY
• DISTRIBUTION AND OCCURRENCE
• BOTANICAL AND ECOLOGICAL CHARACTERISTICS
• FIRE ECOLOGY
• FIRE EFFECTS
• MANAGEMENT CONSIDERATIONS
• REFERENCES

INTRODUCTORY

AUTHORSHIP AND CITATION FEIS ABBREVIATION SYNONYMS NRCS PLANT CODE COMMON NAMES TAXONOMY LIFE FORM FEDERAL LEGAL STATUS OTHER STATUS
	©Kerry Barringer Brooklyn Botanical Gardens

DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• ECOSYSTEMS
• STATES/PROVINCES
• BLM PHYSIOGRAPHIC REGIONS
• KUCHLER PLANT ASSOCIATIONS
• SAF COVER TYPES
• SRM (RANGELAND) COVER TYPES
• HABITAT TYPES AND PLANT COMMUNITIES

UNITED STATES
Connecticut: mountain-laurel (Kalmia latifolia)-black huckleberry-pink azalea (Rhododendron periclymenoides) thickets in the Colebrook Forest [101]

Kentucky: mountain summit and southeastern slope communities with pine-oak (Pinus-Quercus spp.) and oak overstories [13]

chestnut oak (Q. prinus) woodlands near Kentucky Lake in Kentucky and Tennessee [48]

Maine: black huckleberry dwarf shrub heath community with eastern white pine (P. strobus) on the driest inland bog sites in the Bay of Fundy region [27]

Massachusetts: huckleberry-bear oak (Gaylussacia spp.-Q. ilicifolia) heathland communities and mixed maritime shrublands with low sweet blueberry (Vaccinium angustifolium) on Nantucket, Cape Cod, and Martha's Vineyard [39]

pine-oak/huckleberry community with pitch pine (P. rigida), white oak (Q. alba), and black oak (Q. velutina) on Cape Cod [40]

dwarf tree/shrub communities with stunted tamarack (Larix laricina) and scattered black spruce (Picea mariana) on Acadia bogs in central Massachusetts [96]

Michigan: open black oak stands in the southeast [2]

New Hampshire: huckleberry-highbush blueberry (V. corymbosum) vegetation type in the Mud Pond Bog [35]

New Jersey: plains vegetation type dominated by stunted pitch pine with post oak (Q. stellata) and sassafras (Sassafras albidum) as subdominants and pitch pine barrens dominated by shortleaf pine (Pinus echinata), Virginia pine (P. virginiana), and white, chestnut, scarlet, and black oak possible [83]

New York: ericaceous dry mesic communities dominated by pitch pine in the northeast [7]

sedge/bryophyte meadow community in Adirondack Park's Panther Bog [26]

dwarf shrub bogs, highbush blueberry bogs, maritime heathlands, dwarf pine plains, pitch pine-scrub oak barrens, pitch pine-oak/heath woodlands, pitch pine/heath barrens, maritime oak/holly (Ilex spp.) forests, Allegheny oak forests, chestnut oak forests, and Appalachian oak-pine forests [114]

North Carolina: Pitch pine-oak vegetation dominated by chestnut oak in the Wine Spring Creek watershed [92]

North Carolina and Tennessee: pitch pine/heath, Table Mountain pine (P. pungens)/heath, and heath bald communities of the Great Smoky Mountains [20,144]

Vermont: beech (Fagus spp.)-oak/blueberry (Vaccinium spp.) vegetation in the Green Mountain National Forest [124]

Virginia: longleaf pine-loblolly pine-turkey oak (P. palustris-P. taeda-Q. laevis) barrens in the southeast [109]

Wisconsin: eastern white pine/blueberry-huckleberry forests in the southern and central portions of the state [73]

Blue Ridge Province: mountain uplands with black oak and scarlet oak [91]

Northeastern U.S.: black huckleberry/sphagnum (Sphagnum fuscum) vegetation type [29]

raised bogs [53]

Southern Appalachians: Table Mountain pine-pitch pine forests [146]

Upper Midwest: dry sand savannahs with northern pin oak (Q. ellipsoidalis) and black oak [145]

CANADA
New Brunswick: black huckleberry dwarf shrub heath community with eastern white pine (P. strobus) on the driest inland bog sites in the Bay of Fundy region [27]

Nova Scotia: black huckleberry barrens in western portion of the province [129,130]

Ontario: eastern white pine/black huckleberry habitat type in the Chalk River region [18]

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

GENERAL BOTANICAL CHARACTERISTICS RAUNKIAER LIFE FORM REGENERATION PROCESSES SITE CHARACTERISTICS SUCCESSIONAL STATUS SEASONAL DEVELOPMENT
	Dr. Robert W. Freckmann Robert W. Freckmann Herbarium

Aboveground description: Black huckleberry is a low-growing, freely branched, deciduous shrub. It is rigid and erect, generally growing to 3 feet (1 m) tall. Shrubs are often found in clumps due to dense clonal spread [22,34,54,110,117,121,125,131]. Site conditions can affect the growth form. Black huckleberry shrubs grown in the shade are typically taller and more open, while those in open conditions are often shorter and more compact [112].

New branches are minutely hairy, and older wood often has peeling bark [125]. Leaves are simple, alternate, and measure 0.9 to 2.2 inches (2-5.5 cm) long by 0.4 to 1 inch (1-2.5 cm) wide. The firm, shiny, hairless leaves have resinous dots [22,34,54,110,121,125,131].

Flowers are small, cylindrical to bell shaped, and arranged in one-sided racemes [22,125]. Black huckleberry produces berrylike drupe fruits that are generally 0.25 inch (0.63 cm) in diameter. Ten seeds approximately 2 mm long are produced per drupe [12,22,34,76,105,131]. In a review, an average of 22,100 clean seeds weighed an ounce and 780 weighed a gram [12]. One hundred "plump" seeds collected from Maryland weighed 136 mg [30].

Belowground description: Black huckleberry is shallowly rooted below slender scaly rhizomes. It lacks a taproot [15,112]. In the New Jersey pine barrens, complete underground structures of 5 black huckleberry shrubs were exposed by careful hand digging. The researcher found that rhizomes were predominantly in the A₀ and A₁ soil horizons. In soils without these layers, rhizomes are normally concentrated in the top 2 to 3 inches (5-8 cm) of mineral soil. Long rhizomes, while typically confined to the upper soil horizons, may reach as deep as 8 inches (20 cm). Black huckleberry roots and rhizomes often reach the water table in lowland areas but rarely reach the water table in upland sites. Rhizome diameters were generally 0.25 to 0.75 inch (0.6-2 cm) but on occasion were as large as 2 inches (5 cm). Short roots were present along all rhizomes. Longer roots, sometimes as long as 2 feet (0.6 m), arose at rhizome forks or stem bases [79].

Pollination: Pollination is chiefly insect mediated. Flowers are self fertile, but cross pollination by insects, especially bees, is common [125]. In a review, black huckleberry was recognized as a nectar source for 1st and 2nd generations of endangered Karner blue butterfly populations in Wisconsin. The 1st and 2nd adult generations emerge from the pupa in late spring and mid-summer, respectively [58].

Breeding system: Flowers are perfect [125].

Seed production: Information on the quantity of black huckleberry seed produced is lacking.

Seed dispersal: A number of wildlife species eat black huckleberries; however, the viability of seed passed through the digestive tract is unknown [112]. A study of vegetation change on the Montague Plain of central Massachusetts suggests that black huckleberry dispersal by seed is limited. Black huckleberry occurred on 80% of sites that had not been plowed in the past and just 21% of old-field sites. Researchers suspected that black huckleberry's poor recolonization of abandoned agricultural sites was due to poor seed dispersal and/or seedling establishment [98].

Seed banking: Several studies suggest that black huckleberry has a limited seed bank or that germination requirements may be difficult to reproduce in controlled environments. From 34 oak-pine stands in southern New Jersey, soil was collected in June and July of 1986. The total 5,000 cm³ was unstratified and dark, cold greenhouse stratified and only 1 black huckleberry seedling/m² emerged, although it was a dominant understory shrub in the stands sampled [89].

No black huckleberry seedlings germinated from soil samples taken from regenerating eastern white pine-red pine (Pinus resinosa) forests in northern Lower Michigan. Sites burned 1 to 70 years prior, and soil from a total area of 0.25 m² was collected from each site for 3 years. Soil was watered and fertilized in a greenhouse to encourage seedling germination. Germination was monitored until no germination was observed for an 8-week period [119].

Neither black huckleberry seeds nor seedlings were recovered from soil samples taken from Brush Mountain on southwestern Virginia's Jefferson National Forest. Soil was collected in mature oak woodlands dominated by chestnut oak, scarlet oak, black huckleberry, and Blue Ridge blueberry (Vaccinium vacillans). Litter, humus, and surface soil to 2 inch (5 cm) depths were collected. Seventy-five percent of soil samples were encouraged to germinate in greenhouse conditions (40 to 100 °F (5-40 °C)) for a year. The other 25% of soil samples were sifted and visually searched for seed. Researchers acknowledged that some black huckleberry seed may have been missed as it resembled root fragments [120].

A review reported that several black huckleberry seed samples had fewer than 50% viable seeds. When seeds were stored at 41 °F (5 °C) in sealed bottles for more than 2 years, they retained their viability [12].

Germination: Black huckleberry seed germination can be slow, and often germination percentages are low. Fifty black huckleberry seeds taken from ripe fruits were cleaned and sown outdoors. Just 2 seeds germinated after 623 days [1]. Another researcher collected 200 black huckleberry seeds in northern Michigan. Thirteen seeds germinated in soil kept outdoors for a period of 83 days beginning in the late fall. The 1st seeds to germinate took 42 days; the last seeds to germinate took 169 days. Forty-four seeds germinated without cold treatments. The 1st noncold treated seeds germinated after 111 days in the greenhouse and the last seeds after 330 days [101]. Young and Young [150] suggested that a 30-day warm stratification followed by cold seed treatments "enhanced" black huckleberry germination.

Seedling establishment/growth: To date (2006), field studies documenting black huckleberry seedling establishment or growth rate are lacking, suggesting a heavy reliance on vegetative reproduction.

Asexual regeneration: Vegetative regeneration is through rhizome, root crown, and epicormic sprouting. Sprout growth triggered by stem damage typically arises from buds nearest the apex stem [15]. Black huckleberry stems that are merely bent may also produce epicormic shoots from preventitious buds [147].

Black huckleberry clonal growth is normally extensive [21]. Clonal growth is by rhizome extension and upward growth that produces ramets every 10 to 20 inches (30-50 cm). Through the excavation of 5 intact black huckleberry rhizome systems in the pitch pine barrens of New Jersey, researchers revealed a multibranched (mean 6.93 inches (17.6 cm)) between rhizome branches) system with a large number of buds (average 36.2 buds/10 cm). Few buds, however, were ever used for growth [86]. First growing season stems from rhizomes were unbranched and up to 8 inches (20 cm) tall. From ring counts of 69 stems, the average ramet age was 5.7 years, and the maximum age was 15 years [112].

Aboveground damage can affect clonal growth. Experimental manipulations of black huckleberry shrubs in woodlands of New Jersey's Lebanon Forest showed that clipped ramets produced greater stem length and more stems than unclipped ramets. Ramets attached to a greater length of rhizome were more likely to sprout than those with shorter rhizome length. Sprouts were produced close to the severed end of rhizomes [87].

Rates of black huckleberry clonal expansion were studied in a little bluestem (Schizachyrium scoparium) grassland-shrubland community on Nantucket Island, Massachusetts. Clonal patches with discrete edges were compared in aerial photos from 1975 and ground surveys in 1990. Of 40 clones, 28 had significant (p<0.05) increases in size from 1975 to 1990. From all clones studied, the average increase in diameter was 6.9 feet (2.1 m) or 35% in 15 years. Clonal expansion averaged 29.5 m² or 89% from 1975 to 1990. Stem density of clones ranged from a high of 424 stems/ha in the clone center to a low of 4 stems/ha on the clone edge [59].

Black huckleberry is sensitive to salt spray in areas with low water availability and is likely restricted from extreme coastal locations [57].

Climate: Maritime and moist continental climates are common in black huckleberry habitats. The Atlantic coast of Newfoundland, Nova Scotia, and New Brunswick, where black huckleberry is common, has a maritime climate. Evenly distributed annual precipitation ranges from 30 to 59.8 inches (760-1,520 mm) [53]. On Long Island, New York, the climate is mild because of coastal influences. The minimum January and maximum July temperatures based on a 12-year record were 28 °F (-2.2 °C) and 82 °F (28 °C), respectively. Long Island receives an average of 48.7 inches of precipitation (1,240 mm) per year, but from June to September the average is just 3.3 to 5.3 inches (83.3-135 mm)/month [112]. In central Vermont, black huckleberry occurs in a humid continental climate where the minimum and maximum January temperatures are -0.4 °F (-18 °C) and 30 °F (-1 °C), respectively. In July the typical minimum temperature is 57 °F (14 °C) and the maximum temperature is 63 °F (17 °C). The average annual precipitation for this area is 42.1 inches (1,070 mm) [124].

In more western portions of black huckleberry's range, precipitation is slightly lower and the temperature range greater. In the lower Great Lakes region, the climate is continental and the average precipitation is typically 27 to 41.7 inches (680 to 1,060 mm)/year, and annual distribution is relatively even. In northern lower Michigan between 1941 and 1970, the average temperature was 17 °F (-8.6 °C) in January and 65.1 °F (18.4 °C) in July. Mean annual precipitation was 31.3 inches (794 mm) and was evenly distributed [116].

In eastern Kentucky's southern Appalachians, the climate is temperate continental. Spring and fall are mild and are often the fire seasons. Growing seasons typically last 180 to 190 days. Temperatures in January and July average 32°F (0 °C) and 75 °F (24 °C), respectively. Average annual precipitation is 44.5 inches (1,130 mm), and lightning storms are possible in the spring and summer [85].

Elevation: Few areas report elevational tolerances for black huckleberry. In the Adirondack uplands of New York, black huckleberry occurs from 200 to 2,040 feet (60-620 m) [76]. In the Great Smoky Mountains of North Carolina and Tennessee, black huckleberry occupies elevations of 2,200 to over 4,000 feet (670 to >1,200 m)[144].

Soils: Black huckleberry is most typical of course-textured, acidic, nutrient-poor soils throughout its range [14,54,76,112]. In southeastern Michigan, black huckleberry occurs in open black oak stands with infertile, very dry to moist, and moderate to highly acidic (4.5-6.0) soils [2]. In Indiana barrens vegetation with chestnut oak, black jack oak, and Virginia pine occurs on shaley siltstone substrates with sandstone and shale fragments. The soils are acidic with very little fertility [65].

In a survey of pitch pine barrens vegetation in New Jersey, black huckleberry was common or abundant on well-drained and imperfectly drained soils but was absent from poorly drained sites [79].

SUCCESSIONAL STATUS:
Black huckleberry is typically present in late seral communities that result from primary succession. However, following disturbances in areas where black huckleberry is established, it rapidly recolonizes the site.

General: Late seral and climax communities are typical black huckleberry habitat. Black huckleberry was common in the heath-dominated shrub layer of climax oak-hickory forests at historic Mount Vernon, Virginia [143]. In northern lower Michigan, black huckleberry was more characteristic of mature 2nd-growth stands between 54 and 83 years old dominated by northern red oak, red pine, and red maple when compared to disturbed young stands between 2 and 13 years old on similar sites [115].

On the Cape Cod peninsula of Massachusetts, black huckleberry was much more common on sites that had not been disturbed by plowing or farming in the past than on old-field sites [97]. Similarly on the Montague Plain of central Massachusetts black huckleberry occurred on 80% of sites that had not been plowed in the past and just 21% of old-field sites. Researchers suspected that black huckleberry's poor recolonization of abandoned agricultural sites was due to poor dispersal and/or establishment. Land once cleared for agriculture was still, 50 to 100 years later, compositionally different from relatively undisturbed areas [98].

Shade relationships: Black huckleberry tolerates some shade and likely prefers diffuse lighting over full sun conditions. Kudish [76] considered black huckleberry slightly shade tolerant, and Martin [85] indicated an intermediate shade tolerance. In oak-hickory forests of Pennsylvania, black huckleberry "persisted" under dense shade but failed to produce flowers or fruits [122].

Comparisons of black oak woodlands along the southern shores of Lake Michigan in northwestern Indiana revealed greater black huckleberry coverage in forests with reduced canopy cover. Black huckleberry coverage was 0.11%, and frequency was 4%, in black oak woodlands with 77% canopy cover. Coverage was 1.4%, and frequency was 28%, in woodlands with 33% canopy cover. Researchers noted that fire frequency was higher in woodlands with greater canopy cover and may have influenced black huckleberry coverage and frequency [63].

Black huckleberry biomass was much greater in open than dense coniferous forests in western Nova Scotia. Coniferous forests were a mixture of red spruce, black spruce, balsam fir, white pine, and eastern hemlock. Black huckleberry biomass was 4 kg/ha in dense forests with an average canopy cover of 75% and 348 kg/ha in open forests where the average canopy coverage was 44%. In mixed wood forests dominated by red maple, bigtooth aspen (Populus grandidentata), and red oak, however, black huckleberry biomass was 15 kg/ha where the average canopy coverage was 73% but was absent from mixed wood forests with 61% canopy cover [135].

In the Appalachian Plateau and ridge provinces of Pennsylvania, black huckleberry frequency and cover were reduced by harvesting (clearcuts and shelterwoods). Black huckleberry coverage 1 year before harvest and 1 year after harvest were 8% and 3%, respectively. Preharvest frequency was 26%, and frequency was 21% one year after harvest [45].

Primary succession: Black huckleberry is absent from the early stages of primary succession. Black huckleberry is not typically present on sand dunes until 6 to 10 centuries following their formation in eastern Gary, Indiana's, Marquette Park [104]. In the Wilderness State Park of northern Lower Michigan, black huckleberry occurred on dune ridges that were greater than 440 years old. Dune ridge ages were 25 to 2,375 years [80].

Secondary succession: Many disturbances are tolerated by black huckleberry, and postdisturbance recovery is normally quick. Black huckleberry coverage was reduced but frequency increased on spring-burned sites, and coverage and frequency were unchanged on summer burned and mowed plots. In grasslands of Nantucket Island, Massachusetts, sites were burned twice in April over a 2-year period. Vegetation was measured at postfire year 1. The 2nd April fire burned when wind speeds were 10 to 13 miles/hour (16-21 km/h), temperatures were 40 °F (7 °C), and relative humidity was 66%. Backfire flame lengths were 1 to 10 feet (0.3-4 m), and most litter was consumed. Another site burned in an August fire, but fire conditions were not described. Vegetation was measured 2 years following fire. Other sites were mowed biennially in August. Mowing removed vegetation to 2 inches (5 cm) above ground, and posttreatment vegetation changes were measured the 2nd posttreatment year. Changes in black huckleberry frequency and cover are provided below [36].

For information on black huckleberry burned and polluted sites see the discussion in Fire in conjunction with other disturbances.

Grazing: There was almost no difference in black huckleberry frequency and coverage on sites grazed 44 years prior and sites ungrazed for approximately 100 years prior to the study of coastal grasslands on Nantucket Island, Massachusetts. On grazed sites black huckleberry coverage and frequency were 33.3% and 50.5%, respectively. On ungrazed sites, black huckleberry coverage and frequency were 33.6% and 54.8%, respectively. The author noted that domestic sheep only "lightly" browsed black huckleberry [37].

In mixed pine-oak vegetation in southern New Jersey, black huckleberry was clipped in successive years. Sprout "vigor" diminished after the 2nd year of clipping. This finding suggests that black huckleberry may not be tolerant of continued browsing [88].

SEASONAL DEVELOPMENT:
Black huckleberry produced flowers and fruits earlier in more coastal and more southern portions of its range compared to inland and northern sites. Fruits typically persist for several weeks [12].

FIRE ECOLOGY

• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

Fire regimes: Black huckleberry occupies dry, open woodland sites that burn frequently. The pitch pine barrens and plains of southern New Jersey have long growing seasons, hot summer temperatures, strong winds that dry fuels, and level to rolling terrain that easily spread fires [83]. However, in coastal fir (Abies spp.) forests of northeastern North America, conditions are humid and vegetation rarely burns. Fires occur only during periods of extreme or prolonged drought or following disturbances that allow sunlight to penetrate and dry fuels. When fires kill fir trees, black huckleberry-dominated shrublands may persist for several decades following fire [50].

Fire history and return intervals: The coniferous, deciduous, and mixed overstories of black huckleberry habitats burned frequently in presettlement times. In most black huckleberry habitats, an anthropogenic influence on fire frequencies is evident.

Red pine forests: On Pictured Rocks National Seashore in Upper Michigan, data from living tree wedges, stump cross sections, and increment bores suggest that fires in the red, white, and jack pine forests occurred on average once every 21.8 years in presettlement times, but fire occurred just once since the early 1900s. The author indicated that Native people likely contributed to the frequent presettlement fires [82].

In red pine stands of northern Vermont, fire scar data from trees on Resin Ridge were used to reconstruct the fire history. Researchers estimated that from 17 to more than 20 fires occurred in the study area between 1815 and 1987; however, none occurred after 1921. Most were surface fires, indicated by the many red pine trees that survived their 1st fire when under 4 inch (10 cm) dbh. The calculated fire return interval for the area was 37+ years, which likely represents a minimum estimate as it includes the unburned time period of 1921 to 1987. Red pine "recruitment" fires, those that were stand replacing and exposed mineral soil, occurred at 50- to over 100-year intervals. European settlement of the area in all likelihood affected the fire frequency in all years. Likely anthropogenic fires were common in the late 1800s, when some stands burned at 3- to 5-year intervals, and fire suppression efforts were responsible for the fire-free period of 1922 to 1987 [43].

Pitch pine forests and barrens: Pitch pine vegetation types in which black huckleberry is common burn at 10- to 30-year intervals [70,90]. The New Jersey pine plains, pitch pine-shrub oak barrens, and pitch pine-post oak-shrub oak woodlands burn often. The pine plains vegetation dominated by dwarf pitch pine, blackjack oak (Quercus marilandica), and scrub oak burn at 5- to 15-year intervals in a mixture of crown and surface fires. Pitch pine/scrub oak barrens dominated by tree-sized pitch pines burn at 15- to 25-year intervals, and crown fires are more typical than surface fires. Pitch pine-post oak/scrub oak woodlands burn at 25- to 30-year intervals in what are normally crown fires [148].

Much of the research in pitch pine forests indicates a change in composition since the fire exclusion era. Pollen records, historical maps, and historical accounts of central Suffolk County, New York, indicate that settlement of the area increased the fire frequency and likely the coverage of pine barrens vegetation from the 17th to early in the 20th century. Following 20th century fire suppression practices, however, much of the pitch pine-oak/heath woodlands and pitch pine-scrub oak barrens have changed to oak-hardwood dominated forests [77]. In a review, Williams [146] reports that southern Appalachian Table Mountain pine-pitch pine forests burned every 10 to 12 years from the mid-1800s to approximately 1940 in low-severity surface fires. After about 1940, suppression of fires decreased the fire frequency allowing forests to burn only rarely. Decreased fire frequency is linked to decreased pine reproduction, and forests are becoming oak and hardwood dominated [146].

The species composition of pine barrens vegetation has changed since the exclusion of fire in central Suffolk County. Using aerial photographs, researchers found that from 1938 to present, fire size has significantly (p=0.001) decreased, and average area burned per year has generally decreased. From 1938 to 1996, approximately 55% of the study area was unburned. Barrens communities including dwarf pine plains, pitch pine-scrub oak, heath, pitch pine-heath, and scrub oak shrublands vegetation made up 87% of study area in 1938 but just 36% in 1994. Loss of the vegetation was to development or to conversion into pitch pine-oak forests. Fire return intervals of more than 30 years allow oaks to establish and persist. Black huckleberry generally remains in oak forests; however, growth and berry production are less in oak forests than in barren communities [70].

For 177 fires that occurred between 1938 and 1995 in Suffolk County, spring fires occurring prior to plant leaf out were most common. These "high-intensity, top-killing" fires burned when winds were high, humidity was low, and litter and fine fuels were readily combustible. However, deep duff was normally moist, so deep-penetrating fires were unlikely. Researchers suggest that mimicking this type of fire today is difficult because of fragmentation and proximity to wildland urban interfaces [70]. Pitch pine-dominated stands in the Connecticut Valley lowlands of Massachusetts are also highly fragmented, often less than 20 acres (10 ha) in size, and cannot support an occurrence or intensity of fires that likely occurred prior to European settlement of the area [95].

Mixed pine/hardwood woodlands and shrublands: Like the above forests, the fire frequency in mixed woodlands has been highly influenced by settlement and fire suppression. Historical records, photos, and ground surveys revealed that fires on the Cape Cod National Seashore were concentrated in pine-oak woodlands where black huckleberry is a typical understory species. Between 1896 and 1963 there were 31 fires, many of which were greater than 70 acres (30 ha) in size; however, since 1961 there have been no large fires [40]. In what are now oak-dominated woodlands in the central uplands of north-central Massachusetts, researchers used sediment core and dendrological methods to reveal that "infrequent or occasional" fire was important from 1500 to 1900. Since the early 1900s, however, no fire has occurred in the study area [47].

In a fire history review of Pennsylvania's Pocono Plateau, Latham and others [78] indicated that berry pickers set large fires frequently in the late 19th and early 20th centuries. This practice, made illegal by the state in 1897, did not deter berry pickers. In 1942 the state designated the area as unprotected in an attempt to encourage local land owners to stop the burning practice. Based on a single pitch pine tree cut down in 1946, the fire frequency was 1 in 23 years for the vicinity of this tree from the 1820s to 1918 and increased to 1 in 4.5 years from 1919 to 1946 [78].

In the coastal forests of Acadia National Park, Maine, black huckleberry is common, and forest canopies are a mixture of red spruce-Atlantic white-cedar (Picea rubens-Chamaecyparis thyoides), mixed pine, paper birch-mixed aspen, red oak, and northern hardwood stands. From a 24-year period with complete fire records, an average of 9 fires and 371 acres (50 ha) burned, and the fire return interval was an estimated 94 years. When researchers excluded a single year in which a large fire occurred, an average of 7 acres (3 ha) burned per year, and the fire return interval was an estimated 5,000 years. Researchers predicted that the fire return interval for the Park is 100 to 150 years. Park visitors caused over 50% of the fires, and from 1937 to 1979 only 5 of 204 fires were lightning ignited [107].

The following table provides fire return intervals for plant communities and ecosystems where black huckleberry is important. Find further 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".

FIRE EFFECTS

• IMMEDIATE FIRE EFFECT ON PLANT
• DISCUSSION AND QUALIFICATION OF FIRE EFFECT
• PLANT RESPONSE TO FIRE
• DISCUSSION AND QUALIFICATION OF PLANT RESPONSE
• FIRE MANAGEMENT CONSIDERATIONS

Multivariate analysis of burned areas in New Jersey's pine plains indicates that black huckleberry may suffer high mortality and may be the slowest dominant species to recover following severe fires that remove humus layers. Following less severe fires, black huckleberry coverage is typically decreased for the 1st few postfire years, but black huckleberry regains dominance 5 to 7 years following fire. Recovery of black huckleberry is slower on severely burned sites [148]. Cape Cod's pine-oak forests were burned under prescription at 1- to 4-year intervals during both dormant and growing seasons. Frequent fires (exact rotation not provided) that burned when sites were damp and vegetation was dormant encouraged sprouting but decreased black huckleberry biomass. However, a single severe fire during the growing season "destroyed" the black huckleberry root and rhizome system [106].

Fire effects related to seasonality/severity: Typically dormant-season and/or low-severity fires are better tolerated by black huckleberry than growing-season and/or high-severity fires. Black huckleberry regrowth was evident by early summer following an early March low-severity surface fire in pine barren pine-oak vegetation dominated by white, chestnut, black, and scarlet oak with lesser amounts of pitch and shortleaf pine in southern New Jersey. The fire killed all aboveground black huckleberries, and no seedlings occurred on either the burned or unburned sites. Fewer new stems were produced in the 2nd postfire year than in the 1st [88].

In rocky barrens of western Nova Scotia, a "cool" night fire burned the vegetation, and within 2 years black huckleberry and other shrubs had recovered such that burned and unburned sites were undistinguishable [129]. Black huckleberry frequency was exactly the same in 10- to 26-month-old spring-burned and unburned oak woodlands of south-central New York [133].

Black huckleberry coverage was much greater on burned than unburned sites and tended to increase with increased time since fire in the westernmost portion of Great Smoky Mountains National Park, Tennessee. Permanent plots were burned in several different fires from 1976 to 1977. Fires were set in summer, fall, and winter, and postfire litter depths ranged from 0.04 to 1 inch (0.1-3.0 cm). Postfire coverage of black huckleberry was recorded periodically between 0 and 19 years following the fires. Data are summarized below [60]:

Black huckleberry showed a short-lived increase in cover, increased density, and decreased frequency and basal area following a spring fire in mixed oak woodlands of western North Carolina's Nantahala National Forest. The April fire was stand replacing. The prefire coverage of black huckleberry was 1.03%, coverage at postfire month 3 was 1.88%, and 0.96% at postfire month 15. Researchers reported that burning increased berry production and that density increased due to prolific sprouting [41]. For further information on fire responses of black huckleberry and other plants in this community, see the Fire Research Project Summary Early postfire effects of a prescribed fire in the southern Appalachians of North Carolina.

Density increased but frequency and basal area of black huckleberry decreased in the 1st postfire year in mixed oak woodlands in the Nantahala National Forest. The prescription fire burned in April in an overstory of red maple, chestnut oak, red oak, yellow-poplar (Liriodendron tulipifera), and hickory (Carya spp.). A combination of backing and strip fires burned when air temperatures were 46 to 65 °F (8-18 °C), and relative humidity was 30% to 40%. Flame lengths were 10 to 20 inches (30-60 cm), and fire temperature averaged 370 °F (188 °C). Litter was reduced by 92%, fermentation and humus layers by 48%, and small wood (<3 inches (7.5 cm) diameter) by 36%. Black huckleberry was not detected on burned sites sampled at postfire month 3. Findings are summarized below [42].

Black huckleberry was more productive on sites burned in spring wildfires than in spring prescription fires in the pitch pine-oak barrens of New Jersey. One- to three-year-old burned sites were compared to unburned sites, and black huckleberry biomass was lower on all burned sites. Control and wildfire sites had not burned for 51 years prior to the recent fires, and prescription sites burned 14 years previous to these fires [10]. Wildfires were considered more severe than prescription fires, and calcium, magnesium, and potassium in vegetation biomass and in the litter were lower on wildfire than prescription sites. Possibly black huckleberry, a low-nutrient adapted species, was affected more by nutrient levels than burn severity [11]. Aboveground black huckleberry biomass on burned and control sites is summarized below [10]:

In the Red River Gorge Geological Area of eastern Kentucky, black huckleberry presence after fire may have been related to fire severity. The prefire vegetation was dominated by scarlet and chestnut oak in the 4-inch (10 cm) or greater dbh size class and red maple and eastern white pine in the 0.8- to 4-inch (2-10 cm) dbh class size. Fires burned on 2 similar ridge sites. Presence was evaluated for 3 postfire growing seasons. The Pinch-Em-Tight Ridge burned when temperatures averaged 68 °F (20 °C), mean relative humidity was 29%, and winds were between 0 and 1.1 miles/hour (0-1.7 km/h). Black huckleberry was present on unburned reference sites but was absent from burned sites. The Klaber Ridge site burned when the mean temperature was slightly lower, 63 °F (17 °C), relative humidity was greater, 46%, and winds were stronger, 0.99 to 2 miles/hour (1.6-3.2 km/h). Black huckleberry was present on both burned and unburned Klaber Ridge sites [75].

Black huckleberry coverage decreased considerably in the New Jersey pine barrens burned in July 1955. Fire behavior and/or conditions were not described. Prefire black huckleberry coverage was 38.2%, 3 months after fire was 4.8%, over 1 year following fire was 11.7%, and over 2 years after fire was 7.7%. The mortality rate was several times that of Blue Ridge huckleberry (Vaccinium pallidum), a typical associate species with deeper perennating buds [79].

Effects of repeated fire: The number of studies reporting decreased black huckleberry coverage, frequency, and density with repeated fire is nearly equal to those that found increased black huckleberry abundance after several fires. Differences in black huckleberry survival and recovery from multiple fires may be a result of fire season, severity, and/or vegetation type burned. Black huckleberry coverage was often lower on sites burned in multiple fires when compared to preburn coverage in grasslands and coastal heathlands of southeastern Massachusetts. Typically black huckleberry was top-killed in all fires (spring and fall), but sprouted soon after. Black huckleberry was present even on sites burned 6 times in 15 years, and pre- and postfire coverages were only slightly different [38]. See the Research Project Summary Vegetation change in grasslands and heathlands following multiple spring, summer, and fall prescription fires in Massachusetts for further details. On the Necedah National Wildlife Refuge of central Wisconsin, prairie and savannah vegetation was burned under prescription every 1 to 5 years. On unburned marshes black huckleberry frequency was 9%, and frequency on burned marshes was 16% [9].

In red and white pine-dominated stands in northwestern Wisconsin, black huckleberry frequency was 7.8% less on sites burned an average of 3.5 times usually with spring fires than on unburned sites. Twenty burned sites were sampled the summer following the last spring burn, and 8 others were sampled 1 year following fire [139].

Black huckleberry coverage decreased with increasing fire frequency and decreasing time since fire on prescribed burn sites in oak-dominated forests with a "liberal scattering" of nonreproducing pines (species not identified) in New Jersey's Lebanon State Forest. Prescription fires were typically low severity and occurred in the winter annually, biennially, and on 3-year, 4-year, 5-year, 10-year, and 15-year rotations. Black huckleberry coverage was 40% on control sites and just slightly less on sites burned on 15-year rotations with 14 year recovery time. On 10-year-old burn sites with 4 years recovery time, black huckleberry coverage was about 30%. On sites burned in 2-, 3-, and 4-year rotations, black huckleberry coverage was approximately 10% in the 2nd postfire year. On annually burned sites, black huckleberry coverage was nearly 3% in the 1st postfire growing season [17].

Multiple prescribed fires in black oak sand savannahs of Indiana's Hoosier Prairie decreased black huckleberry density. One site burned in 1 fall and 3 spring prescribed fires in 8 years, another other plot burned in 3 spring and 2 fall fires in 8 years. No other fire information was provided [5,6].

Black huckleberry survived single fires with very little change in coverage but had not recovered by the 2nd postfire year following 2 fires in 3 years in the Daniel Boone National Forest of eastern Kentucky. Prescribed fires burned in 1993 and 1995 in scarlet, chestnut, and white oak-shortleaf, Virginia, and pitch pine forests. Both fires were in March and burned under similar prescription conditions that included wind speeds below 18 mph (29 km/hr), relative humidities greater than 25%, and air temperatures below 75 °F (24 °C). Flame lengths for the 1993 fire were 1 to 3 feet (0.3-1 m) and due to a head wind in 1995 flame lengths were longer, 4.3 to 5.2 feet (1.3-1.6 m). The 1995 fires were "hotter." Below are the coverage and frequency of black huckleberry on control, once-burned, and twice-burned sites as measured in August 1997 [4]:

Coverage and frequency of black huckleberry increased on annually and biennially burned sites in Connecticut's eastern uplands. Most fires burned in April from 1968 to 1985. Flame lengths were several meters in shrub thickets, but no other fire information was reported. Site 1 was burned annually from 1968 to 1976 and again in 1978, 1980, and 1983. Site 2 was burned in 1968 and 1969 and then biennially until 1980. Both sites had woody and herbaceous vegetation. Typical woody species included black huckleberry, winged sumac (Rhus copallinum), black cherry (Prunus serotina), and black oak, and the dominant herb was little bluestem. Prior to burning all trees greater than 3 feet (1 m) tall were cut, and stumps were treated with herbicide. Changes in black huckleberry coverage and frequency before and after fire are provided below [103].

Black huckleberry coverage was reduced from prefire values on spring biennial burned sites, relatively unchanged on summer biennial burned plots, but increased with time on control plots in little bluestem grasslands of Nantucket Island, Massachusetts. Spring sites burned in April, and postfire vegetation measurements were made in postfire year 1. The 2nd April fire burned when wind speeds were 10 to 13 mph (16-21 km/hr), the average temperature was 40 °F (7°C), and relative humidity was 66%. Backfire flame lengths were 1 to 10 feet (0.3-4 m), and most litter was consumed. The 2 August fires were not described, and postfire vegetation was measured in postfire year 2. Changes in black huckleberry frequency and cover are provided below [36].

Density and frequency of black huckleberry were greater than prefire measurements on twice- and once-burned sites in longleaf pine-loblolly pine-turkey oak barrens of southeast Virginia. The twice burned site burned once in February 1986 when conditions were warm (>60 °F (20 °C)) and fueled flame heights of 20 feet (6 m). The second fire burned in July 1987. The July fire burned at night when temperatures were lower, and humidity levels were higher. The single burn site burned in February 1988 under conditions similar to the other February fire. Increases in black huckleberry density and frequency were greater on the twice-burned area. Study results are summarized below [109]:

Fire in conjunction with other disturbances: Multiple stresses may decrease black huckleberry coverage. Coverage was lowest on burned sites located closest to a zinc smelter near Palmerton, Pennsylvania. Burned and unburned chestnut oak woodland sites located at various distances from the smelter were evaluated. Fifteen-year-old burned and unburned sites located approximately 1 mile (2 km) from Lehigh Gap's zinc smelter had reduced black huckleberry coverage. Black huckleberry coverage on unburned and burned sites at Lehigh gap was 1.5% and 0.6%, respectively. Black huckleberry coverage on sites located between 9.9 and 21 miles (16-33 km) from the smelter was 18.5% on 14 year old burned sites and 6.6% on unburned sites [68].

The heat content of black huckleberry foliage was measured and found to vary only slightly in different-aged regenerating jack pine stands in Clinton County, New York [127].

MANAGEMENT CONSIDERATIONS

• IMPORTANCE TO LIVESTOCK AND WILDLIFE
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER USES
• OTHER MANAGEMENT CONSIDERATIONS

Domestic sheep: The coastal grassland vegetation of Nantucket Island, Massachusetts, was routinely grazed by sheep until 1948. The author noted that domestic sheep only "lightly" browsed black huckleberry [37].

Deer: White-tailed deer browsing of black huckleberry is variable. Likely white-tailed deer densities, availability of more palatable browse, and age of black huckleberry stems affect selection. Between 1% and 6% of white-tailed deer diets can be huckleberry browse in the eastern U.S. [86]. Little and others [81] suggest that new huckleberry growth is only "lightly" browsed when available.

However, white-tailed deer heavily browsed black huckleberry in red spruce-hemlock-pine forests of southwestern Nova Scotia. White-tailed deer population density was an estimated 5 deer/mi² in the area where black huckleberry shrubs made up 5.4% of the total available weight. White-tailed deer utilization was 6.7%, and the preference factor suggested heavy black huckleberry browsing [136].

Small mammals: Black huckleberry fruits are likely eaten by a variety of small mammals. Martin and others [86] report that huckleberry makes up to 2% of eastern gray squirrel and fox diets in the eastern United States.

Snowshoe hares heavily browsed black huckleberry in red spruce-hemlock-pine forests of southwestern Nova Scotia. Snowshoe hare populations were considered high, but no densities were reported. Black huckleberry shrubs made up 5.4% of the total available weight, and percent utilization by snowshoe hares was 8.0%. The preference factor suggested heavy black huckleberry browsing [136].

Birds: Gamebirds, songbirds, and raptors feed on black huckleberry fruit and/or utilize black huckleberry for cover. Up to 2% of ruffed grouse, northern bobwhite, wild turkey, catbird, white winged crossbill, pine grosbeak, Florida jay, orchard oriole, scarlet tanager, and red-eyed towhee diets can be huckleberries [86]. Reiners [112] indicates that birds of Long Island, New York, have purple-blue scat due to the high number of huckleberries consumed. The viability of seed once passed through the digestive tract is unknown.

In Wisconsin, 1.9% of the fall foods recovered from sharp-tailed grouse was black huckleberry, and 3 of 17 greater prairie-chicken stomach contents contained black huckleberry [138].

Northern harrier habits were studied from 19 February 1987 to 26 April 1988 on Barney's Joy Point, Massachusetts. Northern harrier roosting sites were chiefly in dense black huckleberry patches. The 2 nests found with egg shell pieces were found in dense shrublands dominated by black huckleberry. Barney's Joy Point represents the only known Massachusetts mainland northern harrier breeding site excluding Cape Cod [23].

Reptiles: In New York's Cicero Swamp, the eastern massasauga rattlesnake, listed as endangered in New York [100], was found primarily in a peatland dominated by mountain holly (Nemopanthus mucronatus), highbush blueberry, and black chokeberry, with some black huckleberry and leatherleaf (Chamaedaphne calyculata). The peatland is thought to be important in the gestation period and for overwintering [67].

Insects: Several insects feed on black huckleberry leaves [112], and in a review, black huckleberry is recognized as a nectar source for the 1st and 2nd generations of endangered Karner blue butterflies in Wisconsin. The 1st and 2nd adult generations emerge from the pupa at different times of the year, late spring and mid-summer, respectively [58].

Palatability/nutritional value: Several states report the nutritional value of black huckleberry. Black huckleberry fruits collected in the fall of 1970 and 1971 from several sites in Natchitoches Parish, Louisiana, averaged 4,324 cal/g in energy value and 68.5% in moisture content [19].

The nutrient content and concentrations in black huckleberry leaves collected in open white and northern red oak forests of west Greenwich, Rhode Island, were as follows [72]:

The nutrient and ash concentration of black huckleberry varied only slightly in different-aged jack pine stands regenerating after in Clinton County, New York. Data are summarized below [127]:

Cover value: Huckleberry shrubs provide important cover for birds and small mammals in dry open eastern woodlands [52].

In a review, Duke [33] indicated that several tribes utilized black huckleberry. The Cherokee chewed black huckleberry leaves like tobacco to treat dysentery and tender gums. A black huckleberry tea was drunk when suffering from Bright's disease, the common cold, dysentery, or indigestion. Iroquois smoked black huckleberry leaves, and drank a tea to purify the blood, and to treat arthritis, colds, kidney ailments, rheumatism, tapeworm infections, and venereal diseases. Members of the Chippewa, Delaware, Mohegans, Menominee, Ojibwatribe, Potawatomi, and Shinnecock tribes drank huckleberry teas as a blood tonic and to treat colds, rheumatism, kidney ailments, fevers, and lumbago [33].

Black huckleberry is adapted to nutrient-poor sites, and fertilization of these sites may decrease the abundance of black huckleberry. Researchers irrigated oak forests near the Falmouth Wastewater Treatment Plant in Cape Cod, Massachusetts, with nitrogen-enriched wastewater. Black huckleberry coverage before irrigation treatments was 72.0%. One, two, and three years following irrigation treatments black huckleberry coverage was 75.2%, 71.2%, and 6.0%, respectively. The dramatic decreases in black huckleberry coverage were likely due to extreme increases in weedy American pokeweed (Phytolacca americana) with increased nitrogen inputs [69].

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