© Dr. Donald J. Leopold
18 July 2013: DeGraaf, Richard M.; Rudis, Deborah D. 2001 citation corrected to DeGraaf, Richard M.; Yamasaki, Mariko. 2001.
When habitats overlap, bear oak hybridizes with several species within the red oak group (Erythrobalanus) . The following hybrids are recognized [48,50,77,90,129]:
Quercus × brittonii W.T. Davis, a cross with blackjack oak (Q. marilandica)
Quercus × caesariensis Moldenke, a cross with southern red oak (Q. falcata)
Quercus × fernaldii Trel., a cross with northern red oak (Q. rubra)
Quercus × giffordii Trel., a cross with willow oak (Q. phellos)
Quercus × rehderi Trel., a cross with black oak (Q. velutina)
Quercus × robbinsii Trel., a cross with scarlet oak (Q. coccinea)
Q × brittonii hybrids studied in New Jersey had morphological characteristics intermediate between bear oak and blackjack oak . In New Jersey, a hybridization study revealed that although occupying a common area, bear oak and blackjack oak were hybridizing but bear oak and black oak were not .LIFE FORM:
Bear oak is a dominant species in the following vegetation classifications:
mixed deciduous forests dominated by red maple (Acer rubrum), gray birch (Betula populifolia), and quaking aspen (Populus tremuloides) with a lesser amount of pitch pine (Pinus rigida)
pitch pine/bear oak vegetation
bear oak communities with dense bear oak thickets and scattered pitch pine 
chestnut oak (Q. prinus)/bear oak vegetation in the Appalachian Province of Allegheny County 
pitch pine-bear oak communities dominate the area immediately surrounding Halfway Pond in Plymouth County 
black huckleberry (Gaylussacia baccata)-bear oak tall shrublands on Nantucket Island and Martha's Vineyard, less common on Martha's Vineyard 
pitch pine/bear oak/broom crowberry (Corema conradii) and pitch pine/bear oak-northern bayberry (Morella pensylvanica) communities of the Cape Cod National Seashore 
pitch pine/bear oak, manzanita (Arctostaphylos spp.)-bear oak, mixed oak-pine/huckleberry (Quercus-Pinus-Gaylussacia spp.) vegetation types on Cape Cod 
pitch pine woodlands in the central part of the state with shortleaf pine (Pinus echinata) and Virginia pine (P. virginiana) possible 
pitch pine-post oak (Q. stellata) and chestnut oak forests of the central pine barrens 
pitch pine/bear oak communities in the pine barrens region 
plains community type with a pitch pine overstory and post oak and sassafras (Sassafras spp.) occasionally present, all vegetation is less than 11 feet (3.4 m) tall
transition community dominated by same species as the plains community, but trees are larger with "poor" forms or shapes
barrens community with an open canopy dominated by pitch pine but with some shortleaf pine, Virginia pine, white oak (Q. alba), chestnut oak, scarlet oak, and black oak possible 
open-canopy community dominated by stunted pitch pine, shrub oaks, and a dense layer of ericaceous shrubs in the Hudson Valley 
barrens communities dominated by pitch pine with some scattered white oak and chestnut oak on Long Island
transition communities dominated by bear oak; plains communities
with dwarfed pitch pine, bear oak, and low-growing huckleberry and blueberry (Vaccinium spp.) on Long Island 
woodlands with a mixed canopy of pitch pine, northern red oak, and eastern
white pine (P. strobus) in the northeast 
woodlands with a mixed canopy of pitch pine, northern red oak, and eastern white pine (P. strobus) in the northeast 
Table Mountain pine (P. pungens)/bear oak/heath woodlands in the Blue Ridge Province 
dry slope and ridge forests of the valley and ridge Appalachian provinces 
granite and sand barrens in the southern part of the province 
© 2004 Dr. George P. Chamuris
© 2002 Dr. Donald J. House
GENERAL BOTANICAL CHARACTERISTICS:
This description provides characteristics that may be relevant to fire ecology, and is not meant for identification. Keys for identification are available (e.g. [30,31,48,107,116,129]).
Aboveground description: Bear oak is a deciduous gangly shrub or small tree that often forms dense thickets. It ranges from 3 to 30 feet (1-8 m) tall and 2 to 6 inches (5-15 cm) in diameter [30,31,48,107,116,129]. Stems are short-lived and grow slowly [109,134]. The typical stem life span is 20 to 30 years; however, the bear oak root system is reportedly long lived and may support several generations of short-lived stems [99,134].
Leaves are arranged alternately. Leaf length measures 2 to 6 inches (5-15 cm), and the width is 1 to 4 inches (3-10 cm) [30,31,48,101,116,129]. Bear oak is monoecious, with both male and female flowers on the same plant. Male flowers are catkins and female flowers are borne in clusters or singly. Female flowers are produced on current-year's growth [31,107,116,135]. Acorns are egg-shaped and measure 0.4 to 0.8 inch (1-2 cm) long. The acorn cup is saucer shaped with large scales and covers about 1/2 the nut. Bear oak requires 2 growing seasons to produce fruit [31,48,55,107,116].
Belowground description: Underground bear oak structures carefully excavated from the New Jersey pine barrens revealed that bear oak stems arise from a "massive" irregularly shaped taproot. The taproot is commonly 2 to 4 inches (5-10 cm) thick but may measure as thick as 7 inches (20 cm). Three common root structures were observed. One type had a single main taproot that reached 20 to 36 inches (51-91 cm) deep and lateral roots that lay just below the soil surface. Another had 2 main taproots and several well-developed lateral roots. Root systems with 2 or more main roots growing vertically or obliquely were also common. Two other root types were observed but were less common. One had a single main taproot that extended obliquely, and the other had 1 taproot that forked within a few inches of the soil surface allowing 2 taproot branches to extend horizontally at a depth of 4 to 8 inches (10-20 cm) .
Bear oak roots did not reach into the water table even when it was shallow (1 to 2 feet (0.3-0.6 m), and bear oak was absent or scarce on imperfectly to poorly drained soils. Bear oak roots are intolerant of saturated conditions, which may explain some of the variation in root structure in the New Jersey pine barrens .RAUNKIAER  LIFE FORM:
Pollination: Flowers are wind pollinated .
Breeding system: Bear oak is monoecious , but male and female flowers produced on the same plant are not compatible . Female flowers are borne only on current-year's growth .
Seed production: Mature bear oak acorn production requires 3 growing seasons. Flower buds are produced in the 1st growing season, flowering occurs in the 2nd growing season, and immature acorns are produced in the fall but are not mature until the following growing season . The 1st bear oak sprouts to produce acorns were 3 years old on a clearcut area of New Jersey's Coastal Plain. The 1st bear oak sprouts to flower were 2 years old. Female flower production was greatest in the 5-year-old bear oak age class, lowest in the 13-year-old age class, and intermediate in the 9-year-old age class when the 3 age classes were compared [132,135].
Numerous studies have found that acorn production can be affected by stem age, weather, site condition, and/or herbivory. Cumming  indicates that bear oak acorn production is sporadic. However, in a review, Halls  reports that bear oak produces good acorn crops annually and that yields are greatest for bear oak stems between 4 and 9 years old, although 24-year-old stems may still produce acorns. Wolgast  found that acorn production decreased with age in 5-, 9-, and 13-year age classes in the pine barrens of New Jersey. Acorn number and acorn weight were greatest in the 5-year-old age class. Below are the acorn weights and production by age class. Bear oak abundance on the 0.01-ha plots was not provided .
|Age class (years)||5||9||13|
|Average acorn number/0.01-ha plot||4,599||2,574||1,356|
|Average acorn weight (kg)/0.01-ha plot||4.57||3.16||1.53|
In southern New Jersey pitch pine forests, however, 30-year-old bear oak stems produced more acorns than 7-year-old stems. Researchers collected acorns from 0.4-ha quadrats. No acorns were collected from sites dominated by 3-year-old bear oak stems. Regardless of stem age, however, acorn production was greatest on sites free of pitch pine shading. Data are summarized below .
|7-year-old bear oak stems||30-year-old bear oak stems|
|# acorns found||2||97||36||140|
|# sound acorns||0||64||25||98|
On the University of Rhode Island campus, bear oak plants entirely defoliated by gypsy moths produced a 2nd flush of leaves 2 to 3 weeks following defoliation but had "strongly reduced" acorn production .
In controlled experiments, randomly selected bear oaks from the New Jersey pine barrens were subjected to high (61%-71%) and low (38%-50%) humidity levels at the time of flowering. Significantly (p<0.01) fewer immature acorns were produced under high humidity conditions [133,135].
Measurements of stem height, flowering, and acorn production along a 574-foot (175-m) slope in Plymouth County, Massachusetts, revealed that generally the greatest number of flowers and mature acorns were produced on the tallest stems, which grew at the top of the gradient. The lowest point along the gradient was at the bottom of a 70-foot (20-m) deep depression. See the table below for a summary of the data .
|Relative elevation||Female flowers||Male inflorescences||Proportion of stems producing mature acorns||Stem height|
(average number ± s)
(cm ± s)
Based on the results of 2 separate studies, Wolgast [131,135] concluded that genetic factors were more important than soil site conditions in determining acorn productivity of bear oak. The 1st study measured soil and moisture characteristics beneath 9-year-old open-grown plants that were producing acorns at high, intermediate, and low levels. Site quality was not significantly (p=0.1) different beneath the plants. To better evaluate site influences, 18 of the highest, 18 of the lowest, and 18 of the intermediate acorn producers were transplanted onto a common site in Ocean County, New Jersey. Yields remained relatively the same for the high, low, and intermediate producers, and a significant (p<0.05) genotype difference was observed [131,135].
Seed dispersal: Small mammals and birds may aid in the dispersal of oak acorns by abandoning caches. American and European jays often cache acorns a few meters apart in open environments and cover them with debris or soil .
Seed banking: Bird and small mammal caches suggest that a bear oak seed bank may exist, but to date (2006), information regarding the longevity and viability of bear oak seed banks is lacking. However, Lanner  reports that American and European jays are "vital" to oak establishment, as acorns in their unused winter caches have a high probability of germinating.
Germination: Cold stratification followed by warm temperatures encourages bear oak germination. Bear oak seed collected in the fall from southwestern Virginia was treated to 6 weeks of cold temperatures. The cold temperatures successfully broke bear oak shoot dormancy that exists after emergence of the radicle. Cold stratification also increased the range of temperatures required for successful germination. Below are the germination percentages, shoot lengths, and number of leaves produced per shoot of stratified and unstratified seed under different temperature regimes .
|45-61 °F||64-75 °F||45-61 °F||64-75 °F|
|Average percent germination with shoot elongation||3%||17%||47%||63%|
|Average shoot length (inches)||3.7||2.7||2.8||3.5|
|Number of leaves/shoot||2.8||4.1||3.7||6.2|
Seedling establishment/growth: Bear oak seedling growth, closely monitored for 3 years, revealed rapid 1st season growth that slowed in subsequent years. Seed collected in North Carolina and Virginia was planted in nursery beds in Tennessee. By the end of the 1st growing season, average seedling height was 16 inches (41 cm), mean leaf number was 40, leaf size averaged 9 cm², and an average of 4.5 growth flushes was produced. Seedlings were fertilized . The 1-year-old seedlings were planted onto clearcut or old-field sites in the southern Appalachians. The average initial height of bear oak seedlings planted on the sites ranged from 12 to 14 inches (30-36 cm), and after 3 years the average height was 16 to 23 inches (41-58 cm). Survival of bear oak seedlings ranged from 74% to 86%. A severe late frost in the winter of 1977 to 1978 may have affected seedling survival and/or growth .
Asexual regeneration: Bear oak produces vegetative sprouts from the root crown [2,75]. While asexual regeneration is abundant following top-kill [2,57,60,75], bear oak reproduces vegetatively in the absence of aboveground damage as well.
Bear oak sprout densities can be as high as 50,000 to 100,000 stems/acre following fire or cutting. In Pennsylvania bear oak stems were persistent following consistent cutting. Sprouts were clear cut on two 25 ft² plots twice a year between early June and late September. Twelve cuttings were required to eliminate sprouting. The researcher did not indicate whether sprouting ever resumed following the 12 cuttings .
As sexual reproduction reaches its peak at 6 to 7 years of age, sprout production declines "sharply." A switch in reproductive strategies was observed between 5 and 13 years of age in bear oak plants of the New Jersey pine barrens. The average number of acorns produced per 0.2-acre (0.01-ha) plot was 4,599 for 5-year-old bear oak, and the average fresh weight of sprouts was 0.25 kg/0.01 ha. Thirteen-year-old bear oak plants produced an average of 1,356 acorns/0.01 ha plot, while the average fresh weight of sprouts was 12.29 kg/0.01 ha . However, Buchholz  reported that in the dwarf pitch pine-mixed oak forests of New Jersey's pine barrens plains, vegetative reproduction dominates, and seedling establishment is rare even in the absence of fire.SITE CHARACTERISTICS:
Climate: Bear oak habitats have very similar climate regimes; however, temperature extremes are typically less pronounced in more coastal habitats. On Martha's Vineyard, Massachusetts, the climate is mild. Average winter and summer temperatures are 30 °F (0 °C) and 70 °F (20 °C), respectively. Average annual precipitation is 47 inches (1,200 mm), and thunderstorms commonly occur about 20 times a year . In southern New Jersey, average temperatures are similar. The coldest month is typically February when temperatures average 29 to 34 °F (-1.7-1.1 °C). The warmest month is July when temperatures average 73 to 77 °F (23-25 °C). However, the record high as of 1934 was 107 °F (42 °C), and the record low was -16 °F (-27 °C) . In Maryland's Appalachian Province, there are commonly 120 to 150 days with below freezing temperatures, and the average annual precipitation is 43 inches (1,100 mm), most of which falls in June . In central Pennsylvania, late spring frosts are common .
Elevation: The bear oak forest cover type occupies elevations from near sea level to 3,000 feet (1,000 m) throughout its range . Bear oak is restricted to elevations of less than 1,000 feet (300 m) in the Adirondacks of New York  and from 1,500 to 2,500 feet (460-760 m) in North Carolina .
Soils: Bear oak grows on sandy, rocky, well-drained, nutrient-poor soils [30,31,48,60,127]. In northern Virginia's Blue Ridge Province, Table Mountain pine/bear oak/heath woodlands are found in infertile soils with low pH, calcium, and magnesium levels but high iron and aluminum levels . In the Appalachians, bear oak is common on shallow soils .
Bear oak's persistence depends on periodic disturbances. Bear oak is intolerant of dense shade, rapidly recolonizes clearcut or burned sites, and is most typical of early seral communities.
General: The bear oak forest cover type is maintained by frequent top-killing disturbances . In New York's Albany Pine Bush, bear oak was most common on recently disturbed sites. The recent disturbances included fire and/or land clearing . Pine barren habitats with high bear oak abundance are considered early seral communities in southeastern Massachusetts .
Other researchers describe bear oak habitats as fire-maintained "subclimax" types [91,124]. Buchholz and Good  consider dwarf pitch pine-mixed oak forests of New Jersey steady state, early seral communities that resist change following disturbances.
Observations of the succession of old-field sites in southern New Jersey indicate that recolonization is more rapid on sites where bear oak was established prior to the disturbance than on sites without bear oak coverage at the time of disturbance. Old-field sites, in the beginning, are sedge (Carex spp.) and forb dominated, but soon shortleaf pine, bear oak, and pitch pine invade. When pines are 15 to 20 years old, tree oak species such as white oak, scarlet oak, black oak, and/or chestnut oak begin to invade the understory. As the oaks mature the site is considered a mixed oak-pine vegetation type. Once the pines mature and die, hardwood species dominate the stand. The climax vegetation for these old fields is thought to be hardwood dominated. Frequent "killing" fires arrest successional advancement of the vegetation and maintain subclimax pitch pine/bear oak stands [80,81].
On disturbed sites, bear oak abundance is related to disturbance type and connectivity of suitable habitat. In the Connecticut Valley of Massachusetts, bear oak has not colonized previously plowed old-field sites that were abandoned 50 to 100 years earlier. The researchers suggest that seed dispersal opportunities may be limited in the highly fragmented landscape . However, on the Montague Plain of central Massachusetts, bear oak frequency was equal on plowed and unplowed sites, although abundance was greater on unplowed sites. Agriculture was abandoned 50 to more than 100 years prior to the study . On the Cape Cod National Seashore, bear oak frequency was equal on old woodlots, areas where vegetation was logged but never completely eliminated, and old-field sites that were plowed in the past .
A single study of vegetation change in the Lebanon State Forest of New Jersey revealed a decrease in bear oak coverage as succession progressed. There were 8 years, free of major disturbances, between the 2 vegetation surveys. The largest decrease in bear oak cover, 13.1%, occurred in the pitch pine-mixed oak/scrub oak vegetation type, and the smallest decrease, 2%, occurred in mixed oak-pine/scrub oak vegetation .
Shade relationships: Bear oak is described as extremely shade intolerant and "light demanding" [57,110]. In a Table Mountain pine stand in North Carolina, shaded bear oak shrubs were less than 3 feet (1 m) tall and produced no flowers or acorns during the year observed .
Early postdisturbance responses: Disturbances normally increase bear oak frequency, cover, and/or density. Relative bear oak cover increased on an often-disturbed power line right-of-way in the Hudson Valley Sand Belt of New York. Treatments to the right-of-way included mowing, herbicide applications, and combinations of the 2 treatments. Bear oak average relative cover before treatments was 2.9% and after treatments averaged 4.5%. Frequency was unchanged .
In the New Jersey pine barrens, bear oak was present in 88% of continuous forest stands and 53% of fragmented forests. A significantly (p<0.025) greater proportion, 50%, of continuous forests than fragmented stands, 10.5%, were burned since 1979. Increased burned area may explain the greater bear oak abundance in continuous stands. However, researchers considered fragments more disturbed than continuous forest stands .
Bear oak coverage decreased slightly on repeatedly mowed heathland sites on Nantucket Island, Massachusetts. Coverage increased slightly on untreated sites. Mowing left only 2 inches (5 cm) of aboveground vegetation. Sites were mowed in the fall and winter 4 times in 13 years on a nearly biennial schedule and were evaluated 2 years after the last mowing. Bear oak frequency and cover on mowed and control sites are provided below . For information on bear oak recovery on repeatedly burned sites in Nantucket see Effects of repeated fire.
|Years since treatment||NA||NA||Prefire||2|
|North and South Carolina||May to June |
|Massachusetts (eastern)||mid- to late May |
|West Virginia||May |
|Atlantic Coast||May |
|Blue Ridge Province||April to May |
|North and South Carolina||August |
Flowers are produced prior to leaf out . A single flush of leaves is produced over a 7- to 10-day period in mid- to late May in eastern Massachusetts . Leaves persist through the winter .
Bear oak is capable of producing a 2nd growth flush if defoliated severely by herbivores or if plants experience a killing frost . May and Killingbeck  studied the 2nd set of bear oak leaves produced following severe defoliation by gypsy moths in Rhode Island. They found that the 2nd flush of leaves had increased specific mass. This may indicate an increase in tannins and phenols that may result in "tougher" leaves and decreased subsequent herbivory.Microclimate and site conditions may also affect the seasonal development of bear oak. Along a north-facing slope gradient in Plymouth County, Massachusetts, bear oak flowering and leaf out was delay by nearly a month in the bottom of a depression as compared to the top of a slope .
Fire regimes: Frequent fires are characteristic of bear oak habitats. Dwarf mixed pine-shrub oak vegetation in the New Jersey pine barrens are consistently recognized as fire-maintained communities [79,83,91,136]. A fire frequency of 6 to 8 years is suggested by Bucholz and Gallagher  to preserve dwarf pitch pine-bear oak and blackjack oak forests on New Jersey's pine barren plains. Reasons for frequent fires in bear oak habitats are described for southern New Jersey's pitch pine forests. Bear oak communities are highly flammable because of long growing seasons, high maximum temperatures, strong dry winds, and level to rolling topography that favor ignition and extensive fire spread .
Fire occurrence and return intervals: Bear oak communities may burn as frequently as every 5 years, and the longest mean fire return interval reported for bear oak habitat is 60 years . Wade and others  suggest that the bear oak cover type burns in understory fires at intervals of 35 years or less in eastern forests. Likely the type of vegetation that invades bear oak communities and the speed at which it comes to dominate the canopy affects bear oak's longevity in fire-free communities.
In pitch pine forests of southern New Jersey, Lutz  estimated that prior to 1923, fires burned in plains vegetation an average of once every 8 years. "Transition" communities burned every 12 years, and barren vegetation types burned every 16 to 26 years, on average. For a brief description of these community types, see the New Jersey: section of Habitat Types and Plant Communities.
In the New Jersey pine plains, bear oak is present in several community types with mean fire return intervals ranging from 5 to 60 years. In open shrublands dominated by low-growing blackjack oak, bear oak, and pitch pine, surface and crown fires are common at 5- to 15-year intervals. In mature closed shrublands where vegetation composition is the same as in open pine plain shrublands but grows slightly taller, the mean fire return interval is 15 to 60 years. Fires are sporadic, and a mix of crown and surface fires is common. In pitch pine-shrub oak barrens that support tree-sized pitch pine burn, crown fires are more common than surface fires, and the average fire frequency is 15 to 25 years. Pitch pine-post oak/shrub oak woodlands typically burn in crown fires that occur every 25 to 30 years, on average. Pitch pine-tree oak/shrub oak and pitch pine-shortleaf pine-tree oak/shrub oak vegetation types support a mixture of crown and surface fires at 30- to 40-year intervals .
In Cape Cod, eastern Massachusetts, signs of past fire were evident in most bear oak habitats. Fire scars and/or macroscopic charcoal were found in over 70% of pitch pine/bear oak and oak-pine/huckleberry communities and in nearly 50% of manzanita (Arctostaphylos spp.)-bear oak communities. Because of the methods used in this study, the evidence of past fire has likely been underestimated. Underestimates were likely greatest for shrub-dominated communities, which lack trees .
Fire behavior: Severe fires were typical in bear oak communities. In the Connecticut Valley of Massachusetts, large, severe, summer fires were predominant in pitch pine/scrub oak vegetation early in and through the middle of the 20th century . Spring fires before leaf out were most common in Suffolk County, New York, between 1938 and 1995. The "high-intensity, top-killing" fires burned when winds were high, humidity was low, and litter and fine fuels were readily combustible. However, the deep duff layer was normally moist, so deep-penetrating fires were unlikely. Researchers suggest mimicking historical fire regimes in the Connecticut Valley and Suffolk County today is difficult given the highly fragmented nature of the vegetation and its proximity to wildland urban interfaces .
Changes in fire frequency over time: Fire frequency changes in bear oak habitats are often the result of anthropogenic influences. In the pitch pine forests of southern New Jersey, fires were common, but natural fire ignitions were rare. Anthropogenic fire starts were common, and since these woodlands were not valued for timber production large areas were allowed to burn. However, successful fire exclusion in the area led to reduced fire frequencies beginning in 1932 . Historical fire records indicate that fire size and area burned annually has decreased since 1940 in the 1,400,000-acre (550,000-ha) New Jersey pine barrens. From 1906 to 1939, an average of 55,275 acres (22,369 ha) burned per year, and the average fire size was 112 acres (45.5 ha). During the early 1900s, the fire return interval for the area was an estimated 20 years. From 1940 to 1980, an average of 20,050 acres (8,115 ha) burned annually, fire size averaged 16 acres (6.4 ha), and the estimated fire return interval was 65 years. Most fires in the pine barrens were stand replacing .
The species composition of pitch pine barrens in Suffolk County, New York, has changed with fire exclusion practices. Aerial photographs revealed that from 1938 to 1995 average fire size has significantly (p=0.001) decreased. From 1938 to 1996, approximately 55% of the study area was unburned. Barren communities including dwarf pine plains, pitch pine/scrub oak, heath, pitch pine/heath, and scrub oak shrublands made up 87% of the study area in 1938 but just 36% in 1994. Bear oak shrublands decreased by 84% (857 acres (347 ha)) from 1938 to 1994, and pitch pine/bear oak woodlands decreased by 72% (5,100 acres (2,065 ha)) from 1938 to 1994. A severe wildfire in 1995, however, mitigated the loss of bear oak shrublands by creating 2,000 acres (820 ha) of this vegetation type. Fire return intervals of more than 30 years allow tree oaks to establish and persist .
In Suffolk County, New York, European settlement increased the amount of area occupied by pine barrens vegetation through logging and burning in the 17th, 18th, and 19th centuries. Logging typically involved the selective removal of deciduous hardwoods for building material and fuel. Fires were used to clear areas for farming, came from railroad engine sparks, and occurred through human carelessness. Fire exclusion in the 20th century has led to the conversion of a lot of once barren vegetation to tree oak-hardwood forests in north-central Suffolk County and tree oak-pitch pine woodlands in south-central Suffolk County . However, research in the Pocono till barrens of Pennsylvania indicates that areas were bear oak-dominated prior to settlement. These areas are thought to have been maintained through periodic fire, fire-adapted species dominance, low-nutrient soils, and microclimates .
On Martha's Vineyard Island of Massachusetts, historical records indicate that fire size has decreased since the late 1940s. Between 1855 and 1990, over 20 fires burned that were 990 acres (400 ha) or larger, and 9 or more fires exceeded 5,000 acres (2,000 ha) in size. More than 90% of fires burned in the spring. However, since the late 1940s there was only 1 fire larger than 1,000 acres (600 ha), and since 1965 no fires have exceeded 100 acres (40 ha) in size .
The following table provides fire return intervals for plant communities and ecosystems where bear oak is important. Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".
|Community or ecosystem||Dominant species||Fire return interval range (years)|
|shortleaf pine||Pinus echinata||2-15|
|shortleaf pine-oak||Pinus echinata-Quercus spp.||<10|
|Table Mountain pine||Pinus pungens||<35 to 200 |
|pitch pine||Pinus rigida||6-25 [17,59]|
|eastern white pine||Pinus strobus||35-200 [117,121]|
|eastern white pine-northern red oak-red maple||Pinus strobus-Quercus rubra-Acer rubrum||35-200|
|Virginia pine||Pinus virginiana||10 to <35|
|Virginia pine-oak||Pinus virginiana-Quercus spp.||10 to <35 |
|quaking aspen-paper birch||Populus tremuloides-Betula papyrifera||35-200 [28,121]|
|northeastern oak-pine||Quercus-Pinus spp.||10 to <35|
|southeastern oak-pine||Quercus-Pinus spp.||<10|
|white oak-black oak-northern red oak||Quercus alba-Q. velutina-Q. rubra||<35 |
|bear oak||Quercus ilicifolia||5-60 [83,128]|
|chestnut oak||Quercus prinus||3-8|
|northern red oak||Quercus rubra||10 to <35|
|post oak-blackjack oak||Quercus stellata-Q. marilandica||<10|
|black oak||Quercus velutina||<35 |
On a burned site in eastern Massachusetts, an estimated 50% of bear oak stems were produced within 2 years of the fire. Fire characteristics were not described . Bear oak sprout density following fire can be 50,000 to 100,000 stems/acre . While repeated severe fires are tolerated, a fire frequency of less than 5 years reportedly decreases bear oak size and "vigor." When plots in Massachusetts burned in a severe drought-season fire and then reburned for several successive years in summer fires, bear oak was eliminated .
Vegetative regeneration predominates on burned sites. Following a May wildfire in dwarf pitch pine-mixed oak forests of New Jersey, just 2 oak (bear or blackjack oak) seedlings occurred in a 40-m² area in the 3rd postfire month . Bear oak's established root system may be more effective than seedlings in capturing the short-lived flush of nutrients that occurs following fire and may explain the lack of seedling emergence . Following an "intense crown fire" in mixed oak-red maple forests of north-central Pennsylvania, bear oak sprout and seedling density averaged 883±883 (sx)/ha and 16±16/ha, respectively on 12 burned stands. The 6 unburned stands had no sprouts or seedlings .DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Bear oak populations in North Carolina, the southern limit of bear oak's range, have declined by 24% from 1984 to 1997 due to advancing succession brought about by fire exclusion . In western Rhode Island, bear oak was much more important on sites that had burned 25 to 30 years prior to the study than on sites that had been free of fire for 200 years or more. The relative density of bear oak in the understory on past burned sites was 93.4% and was 4.7% on unburned sites. Bear oak was absent from the overstory on unburned sites and had a relative density of 0.9% on burned sites .
Early fire effects: Commonly bear oak abundance is greater on burned than unburned sites as early as the 1st postfire growing season. Bear oak aboveground biomass was much greater on severely burned wildfire sites than on prescribed fire sites in pitch pine-mixed oak woodlands of New Jersey. Wildfires were effective in temporarily removing the canopy, while prescription fires left a scattering of large pitch pines (up to 56 feet (17 m)) and a thick oak subcanopy (10 to 30 feet (4-10 m)). Bear oak was absent from the unburned reference site. Bear oak aboveground biomass on unburned, wildfire, and prescribed fire sites is summarized below .
|Time since fire (years)||NA||2||3||1||3|
|Biomass (kg/ha ± sx)||0||471±216||1,396±202||37±37||160±91|
Bear oak sprout density in the 1st postfire growing season exceeded that of the 2nd postfire growing season following low-severity, March prescription fires in a variety of community types in Centre County, Pennsylvania. The density of sprouts in the 1st and 2nd postfire years was much greater than the density of sprouts on unburned sites. Bear oak sprouts on the 1-year-old burned site averaged a low of 11 inches (28 cm) in the scrub oak/mixed oak community and a high of 19 inches (47 cm) in the scrub oak community. On 2-year-old burned sites, average sprout height ranged from a low of 15 inches (37 cm) in scrub oak/mixed oak vegetation to a high of 25 inches (63 cm) in scrub oak/quaking aspen vegetation. No acorns were produced on either the 1- or 2-year-old burned sites, but they were produced on unburned sites. Ninety to ninety-seven percent of bear oak was top-killed by fires in the scrub oak communities. The average density (sprouts/m²) of bear oak on burned and unburned plots in scrub oak communities is summarized below .
|Time since fire||Scrub oak||Scrub oak/quaking aspen||Scrub oak/pitch pine||Scrub oak/mixed oak|
Effects of repeated fire: Bear oak abundance on burned plots following repeated fires is more variable. This variability is likely related to the number of fires, recovery time, seasonality, and severity of fires.
Bear oak coverage decreased slightly on repeatedly burned heathland sites on Nantucket Island, Massachusetts. Coverage increased slightly on untreated sites. Decreases were similar on fall- and spring-burned sites, although fall-burned sites burned 8 times in 13 years, and spring-burned sites burned 3 times in 13 years. Prescribed fires were predominantly strip headfires, and flame lengths, at times, exceeded 30 feet (10 m). Postfire measurements were made 4 years after the last fire on fall-burned sites and 2 years following fire on spring-burned sites. Spring fires were considered more severe than the moderate-severity fall fires. Bear oak frequency and cover on spring-burned, fall-burned, and unburned sites are given below . For additional information on this study, see Vegetation change in grasslands and heathlands following multiple spring, summer, and fall prescription fires in Massachusetts. For information on bear oak recovery on sites repeatedly mowed in the dormant season, see Early postdisturbance responses.
|Years since fire||NA||NA||prefire||4||prefire||2|
In New Jersey's Lebanon State Forest, bear oak importance increased with increased fire frequency in mixed oak-pine woodlands. Sites were burned under prescription every 1, 2, 3, 4, 5, 10, and 15 years in the winter. Annually burned sites were evaluated in postfire year 1; sites burned 2, 3, and 4 times were evaluated 2 years after the last fire; sites burned at 5 and 10 year intervals were visited in the 4th postfire year; and the 15-year site was evaluated 14 years following fire .
Total bear oak production increased with increased recovery time on once-burned and twice-burned plots in a Pennsylvania bear oak-dominated woodland. A site in Centre County was burned in April 1969 by an "intense" headfire when winds were 3 to 5 miles/hour (5-8 km/h), temperatures were 68 to 70 °F (20-21 °C), fuel moistures were low, and soils were wet. This fire top-killed 100% of the shrubs and trees less than 8 inches (20 cm) dbh. Portions of the Centre County site were burned again in April 1971 or April 1972 in low severity fires that top-killed sprouts and seedlings. Bear oak shoot production was significantly (p<0.05) greater on burned than unburned plots. Production of bear oak is summarized below for unburned, once-burned, and twice-burned plots . See Palatability/nutritional value for information regarding the nutrient concentration of bear oak foliage on burned and unburned plots.
|Treatment||Time since fire
|Total aboveground production
Fire in conjunction with other disturbances: Bear oak was more affected by lack of fire than by pollution when burned and unburned sites located close to a zinc smelter near Palmerton, Pennsylvania, were compared. Burned and unburned chestnut oak woodland sites located at various distances from the smelter were evaluated. Unburned sites had the lowest bear oak sapling density regardless of their distance from the smelter. Fourteen-year-old burned sites located between 9.9 and 21 miles (16-33 km) from the smelter supported 2,467 bear oak saplings/ha, and fifteen-year-old burned sites located approximately 1 mile (2 km) from Lehigh Gap's zinc smelter supported 2,140 saplings/ha. On unburned sites located between 9.9 and 21 miles (16-33 km) from the smelter, bear oak sapling density was 222/ha but was less, 8/ha, on sites nearest to the smelter .FIRE MANAGEMENT CONSIDERATIONS:
|Common name||Scientific name|
|pitch pine||Pinus rigida|
|bear oak||Quercus ilicifolia|
Historic fire regime characteristics for eastern pine barrens and oak-pine (Quercus-Pinus spp.) communities are summarized below:
|Fire regime information on the vegetation community studied in this Research Project Summary. Fire regime characteristics are taken from the LANDFIRE Rapid Assessment Vegetation Model . This vegetation model was developed by local experts using available literature and expert opinion as documented in the .pdf file linked from the name of the Potential Natural Vegetation Groups listed below.|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
Percent of fires
|Surface or low||65%||12|
|Oak-pine (eastern dry-xeric)||Replacement||4%||185|
|Surface or low||90%||8|
|Fire Severities: Replacement=Any fire that causes greater than 75% top removal of a vegetation-fuel type, resulting in general replacement of existing vegetation; may or may not cause a lethal effect on the plants. Surface or low=Any fire that causes less than 25% upper layer replacement and/or removal in a vegetation-fuel class but burns 5% or more of the area. Mixed=Any fire burning more than 5% of an area that does not qualify as a replacement, surface, or low-severity fire; includes mosaic and other fires that are intermediate in effects [58,70].|
|Fire severity by stand type|
|Dwarf pine plains 1||17.2|
|Dwarf pine plains 2||14.2|
|Dwarf pine plains 3||14.4|
|Dwarf pine plains 4||11.8|
|Dwarf pine plains 5||10.0|
|Tall stand 1||13.8|
|Tall stand 2||13.3|
Survival: Top-kill of pitch pine was nearly 100% on all sites except for the least severely burned dwarf stand. Postfire sprouting of pitch pine was rare, and of the 255 trees with basal sprouts in the dwarf pine plains in the first postfire spring, just 15 had live basal or epicormic sprouts 6 years after the fire. Observations in the intermediate and tall stands suggested that most postfire pitch pine sprouts died in those stands as well. Adult pitch pine tree survival was very low, and regeneration depended on seedling establishment.
Seedling recruitment: Over 85% of pitch pine trees in the dwarf pine plains produced serotinous cones, whereas serotinous cone production was about 20% in intermediate stands and less than 10% in tall stands. In some of the most severely burned dwarf pine sites, serotinous cones were consumed by the fire. There was a significant (P<0.05) negative relationship between fire severity and number of cones remaining on live or dead trees within the dwarf pine plains. An average of 4.4 pitch pine seedlings were recruited/prefire adult tree in the most severely burned dwarf pine site. In intermediate and tall pine stands, recruitment was also low due to a lack of mature cones at the time of the fire. Very few pitch pine seedlings occurred in nearby unburned stands. Most seedling recruitment (>90%) occurred in the first postfire growing season. However, pitch pine seedling establishment was observed in all stands until the fourth postfire year and in the dwarf pine plains until the sixth postfire year. There were no new pitch pine seedlings found in any stand in the eighth postfire year. Pitch pine recruitment results are summarized below.
|Pitch pine recruitment by stand type (values represent 10 months after fire unless otherwise indicated)|
|Site||Total pitch pine density (alive and dead)||Mean proportion of trees with live crowns||Mean proportion of sprouting trees/plot||Seedling density||Seedling density (8th postfire year)||Seedlings/prefire adult||Seedlings/prefire adult (8th postfire year)|
|Dwarf pine plains 1||0.27||0||0.03||1.0||0.33||4.4||1.5|
|Dwarf pine plains 2||0.43||0.01||0||6.8||1.5||24||5.0|
|Dwarf pine plains 3||0.37||0||0.01||4.6||1.5||20||6.0|
|Dwarf pine plains 4||0.30||0.02||0.02||11.1||3.6||43||14|
|Dwarf pine plains 5||0.55||0.10||0.06||27.5||5.0||75||14|
|Tall stand 1||0.09||0||0||1.4||0.7||25||13|
|Tall stand 2||0.02||0||0.01||1.5||0.7||30||11|
Seedling survival: Through the first postfire summer, pitch pine seedling survival was over 85% in all stands. In the second postfire summer, pitch pine seedling survival in the dwarf pine plains dropped to 44%; most seedlings died during a 2-week dry period. Survival was 82% in the tall stands and 90% in the intermediate stands in the second postfire year. In the eighth postfire year, pitch pine seedling survival in the dwarf pine plains was 20% to 40%, between 60% and 80% in the intermediate stands, and between 40% and 60% in the tall stands.
Seedling growth and reproduction: Pitch pine seedling height, measured in the eighth postfire year, revealed that seedlings grew most in the tall stands and least in the dwarf pine plains.
The first immature pistillate cones occurred on 2.5-year-old pitch pine seedlings. Pitch pine seedlings in the dwarf pine plains were reproductive at a significantly (P<0.05) shorter height than those in intermediate and tall stands. Pitch pine cone-bearing seedlings were significantly (P<0.05) younger in the tall than in the intermediate and dwarf pine plains. A summary of reproductive pitch pine seedlings on burned sites is presented below.
|Number, age, and size of reproductive seedlings by burned stand type|
|Stand||Number reproductive 8 years after fire||Proportion reproductive||Age at first cone production
(minimum age, in years)
|Height at first cone production
(minimum height, in cm)
|Dwarf pine plains||52||0.038||5.9a||58a|
|Different subscripts within a column are significantly (P<0.05) different.|
Bear oak sprouts were observed weeks following the fire. There were very few bear oak seedlings. Cover of bear oak increased with time since fire. By the fifth postfire year, bear oak cover was 40% to 50% in the dwarf pine plains, 75% in the intermediate stands, and 40% to 60% in the tall stands.
|Cover of bear oak in burned stands (postfire month 10)|
|Site||Bear oak cover (%)|
|Dwarf pine plains 1||28|
|Dwarf pine plains 2||16|
|Dwarf pine plains 3||29|
|Dwarf pine plains 4||20|
|Dwarf pine plains 5||21|
|Tall stand 1||9|
|Tall stand 2||18|
Effects of bear oak on pitch pine: Pitch pine seedling survival was significantly lower (P=0.0001) on plots where bear oak was clipped than on unclipped plots. However, pitch pine seedling growth was reduced under the bear oak canopy. In the eighth postfire year, pitch pine seedlings under bear oak canopies has smaller stem diameters and smaller projected crown areas in clipped than unclipped plots.
Soils: Characteristics of the soils on each site in the fifth postfire year are provided in Landis and others .FIRE MANAGEMENT IMPLICATIONS:
Bear: Acorns provide important fall weight gain for prehibernating black bears .
Deer: Bear oak stems and acorns are eaten by white-tailed deer [23,57]. Browse utilization may be related to season and stem age. White-tailed deer browsing was greater on burned than unburned bear oak-dominated woodlands in central Pennsylvania. In the summer of 1971, white-tailed deer browsed 43% of bear oak stems on plots burned the previous spring but browsed fewer stems, 23% to 26%, on unburned and 3-year-old burned plots. In 1972 white-tailed deer browsed 57% of bear oak stems on 1-year-old burned plots and browsed 25% or less on unburned and 2-year-old burned plots . Following low-severity, early spring, prescription fires in scrub oak communities of Centre County, Pennsylvania, white-tailed deer browsed 3.6% of all new sprouts .
In central Pennsylvania, white-tailed deer utilization of bear oak was heaviest in the late winter months. White-tailed deer browsed bear oak stems rarely (trace-0.7%) from April through November. Utilization in December, January, February, and March was 2.7%, 1.8%, 5.7%, and 9.9%, respectively. Percent utilization was averaged over 3.5 years of data collection .
Little and others  observed few white-tailed deer browsing on 1-year-old bear oak stems in southern New Jersey pitch pine forests. However, white-tailed deer did feed on the acorns of low-growing scrub oaks. The researchers predicted that scrub oak acorns likely fed deer for just 6 weeks of the year, but in good mast years acorns may be available for several months. For information on acorn production in southern New Jersey, see Seed production. Several other researchers indicate that acorns are palatable to white-tailed deer and an important source of fat through the fall and winter [23,25].
Small mammals: A number of small mammals feed on acorns. Small mammal browsing occurs but is likely affected by bear oak stem age and seasonality. A 2-year study of eastern and New England cottontail feeding habits in Connecticut revealed bear oak utilization was low. Based on 170 winter (January-March) observations, researchers observed feeding on bear oak 5 times. In the summer (April-October), cottontails did not feed on bear oak in 27 total observations .
Eastern chipmunks, eastern gray squirrels, red squirrels, southern flying squirrels, and northern flying squirrels all feed on acorns and commonly cache nuts for overwinter use .
Birds: Bear oak habitats and acorns are important for a diversity of game and songbird species. Great crested flycatchers are common in dry pitch pine barrens, pine warblers utilize pitch pine woodland habitats, and brown thrashers and eastern towhees are common in pitch pine/scrub oak forests of New England . In the Barrens Grouse Habitat Management Study Area in Centre County, Pennsylvania, Yahner  found that species richness, species diversity, and total density of the ground-shrub foraging bird guild were positively correlated with bear oak, dwarf chinkapin oak, and black cherry (Prunus serotina) densities. Birds characteristic of this guild included gray catbirds, golden-winged warblers, chestnut-sided warblers, and field sparrows . The number of bird species observed in bear oak was 17 in the spring and 2 in the winter from 1981 to 1984 . In Plymouth County, Massachusetts, ruffed grouse were associated with sites having high bear oak coverage in pitch pine/bear oak communities .
In pitch pine/bear oak barrens in southeastern Massachusetts, 31 breeding bird species were observed in the study area. Rufous-sided towhees, common yellowthroats, and prairie warblers made up 49% to 70% of the total breeding bird density. Rufous-sided towhees used bear oak vegetation when foraging above ground, and common yellowthroats and prairie warblers had a "pronounced preference" for bear oak when foraging .
A number of birds feed on acorns, so it is likely that when bear oak and bird habitats overlap, bear oak acorns would be utilized. Red-bellied and other woodpeckers, jays, white-breasted nuthatches, and brown thrashers feed on acorns. Acorns are also eaten by wild turkeys [25,57] and are preferred over other available natural foods . In the Nathaniel Mountain Refuge of southern Hampshire County, West Virginia, bear oak together with dogwood (Cornus spp.) and fox grape (Vitis labrusca) made up the bulk of wild turkey diets. The researcher indicated that wild turkeys were competing with white-tailed deer and squirrels for bear oak acorns .
Insects: Bear oak supports a rich insect fauna that includes several rare moth species. A diverse community of Coleoptera and Lepidoptera insect herbivores feeds on bear oak , and planthoppers of the Homoptera order also utilize bear oak as a host . Surveys of pitch pine/bear oak barrens from Maine to Virginia revealed that 44 Miridae insect species were associated with bear oak. The researcher noted that insect associations in bear oak were more diverse than insect associations in other northeastern oak communities .
Bear oak is the only or primary larval host for 16 of 56, or 29%, of the rare or endangered Lepidoptera species in southern New England and southeastern New York. However, this likely underestimates bear oak's importance, as 12 polyphagous species utilized bear oak and/or low sweet blueberry (Vaccinium angustifolium) as hosts, as well . In southeastern Massachusetts, bear oak is host for Lepidoptera species that include daggermoths, pine barrens itames, Melsheimer's sack-bearers, spiny oakworms, and Gerhard's underwing moths. These moths were associated with early successional pine barrens habitats with a high abundance of bear oak .
In bear oak, black jack oak, and pitch pine pygmy forests of New Jersey's Burlington and Ocean counties, pitfall trapping of 2 tiger beetles (Cicindelidae unipuncata and Megacephala virginica) was more successful than expected, suggesting that these species may be more common than realized .
Palatability/nutritional value: Foliar and acorn nutrient concentrations of bear oak in Pennsylvania are provided below. The nutrients available in dry and fresh bear oak acorns that were collected in late September from Pennsylvania are as follows :
|Percent content (dry)||Percent content (fresh)|
In Centre County, Pennsylvania, the foliar nutrient concentrations of bear oak on unburned, once-burned, and twice burned plots were compared. For information on the fires see Effects of repeated fire. For the data below, values in the same column that are followed by different letters were significantly (p<0.05) different :
|Treatment||Years since fire||P (%)||Crude protein (%)||K (%)||Ca (%)||Mg (%)|
Cover value: Bear oak thickets provide "excellent" cover for many wildlife species year round . For additional information on the importance of bear oak in wildlife habitats, see the species group of interest in Importance to livestock and wildlife.VALUE FOR REHABILITATION OF DISTURBED SITES:
In a review, bear oak was reportedly used by menstruating Iroquois women .OTHER MANAGEMENT CONSIDERATIONS:
1. Aizen, Marcelo A.; Kenigsten, Alejandra. 1990. Floral sex ratios in scrub oak (Quercus ilicifolia) vary with microtopography and stem height. Canadian Journal of Botany. 68: 1364-1368. 
2. Aizen, Marcelo A.; Patterson, William A., III. 1995. Leaf phenology and herbivory along a temperature gradient: a spatial test of the phenological window hypothesis. Journal of Vegetation Science. 6(4): 543-550. 
3. Allen, Roosevelt; Farmer, Robert E., Jr. 1977. Germination characteristics of bear oak. Southern Journal of Applied Forestry. 1(1): 19-20. 
4. Barden, Lawrence S. 1985. Bear oak (Quercus ilicifolia) in North Carolina. Castanea. 50(2): 121-123. 
5. Barden, Lawrence S. 2000. A common species at the edge of its range: conservation of bear oak (Quercus ilicifolia) and its low elevation rocky summit community in North Carolina. Natural Areas Journal. 20(1): 85-89. 
6. Baskin, Carol C.; Baskin, Jerry M. 2001. Seeds: ecology, biogeography, and evolution of dormancy and germination. San Diego, CA: Academic Press. 666 p. 
7. Bernard, John M.; Seischab, Franz K. 1995. Pitch pine (Pinus rigida Mill.) communities in northeastern New York State. The American Midland Naturalist. 134(2): 294-306. 
8. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. 
9. Boerner, Ralph E. J. 1981. Forest structure dynamics following wildfire and prescribed burning in the New Jersey Pine Barrens. The American Midland Naturalist. 105(2): 321-333. 
10. Boyd, Howard P. 1985. Pitfall trapping Cicindelidae (Coleoptera) and abundance of Megacephala virginica and Cicindela unipunctata in the pine barrens of New Jersey. Entomological News. 96(3): 105-108. 
11. Bramble, W. C.; Goddard, M. K. 1943. Seasonal browsing of woody plants by white-tailed deer in the bear oak forest type. Journal of Forestry. 41(7): 471-475. 
12. Bramble, William C. 1947. Indicator types for Virginia pine stands in central Pennsylvania. Research Paper No. 8. University Park, PA: The Pennsylvania State Forestry School. 8 p. 
13. Brown, James H., Jr. 1960. The role of fire in altering the species composition of forests in Rhode Island. Ecology. 41(2): 310-316. 
14. Brush, Grace S.; Lenk, Cecilia; Smith, Joanne. 1980. The natural forests of Maryland: an explanation of the vegetation map of Maryland. Ecological Monographs. 50(1): 77-92. 
15. Buchholz, Kenneth. 1983. Initial responses of pine and oak to wildfire in the New Jersey Pine Barren plains. Bulletin of the Torrey Botanical Club. 110(1): 91-96. 
16. Buchholz, Kenneth; Gallagher, Mark. 1982. Initial ectomycorrhizal density response to wildfire in the New Jersey Pine Barren Plains. Bulletin of the Torrey Botanical Club. 109(1): 396-400. 
17. Buchholz, Kenneth; Good, Ralph E. 1982. Density, age structure, biomass and net annual aboveground productivity of dwarfed Pinus rigida Moll. from the New Jersey Pine Barren Plains. Bulletin of the Torrey Botanical Club. 109(1): 24-34. 
18. Buell, Murray F.; Cantlon, John E. 1953. Effects of prescribed burning on ground cover in the New Jersey pine region. Ecology. 34: 520-528. 
19. Burnham, C. F.; Ferree, M. J.; Cunningham, F. E. 1947. The scrub oak forests of the Anthracite Region. Station Paper No. 4. [Philadelphia, PA]: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 9 p. 
20. Catling, Paul M; Brownell, Vivian R. 1999. The flora and ecology of southern Ontario granite barrens. In: Anderson, Roger C.; Fralish, James S.; Baskin, Jerry M., eds. Savannas, barrens, and rock outcrop plant communities of North America. New York: Cambridge University Press: 392-405. 
21. Christensen, Norman L. 1988. Vegetation of the southeastern Coastal Plain. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. Cambridge: Cambridge University Press: 317-363. 
22. Copenheaver, Carolyn A.; White, Alan S.; Patterson, William A., III. 2000. Vegetation development in a southern Maine pitch pine - scrub oak barren. Journal of the Torrey Botanical Society. 127(1): 19-32. 
23. Cumming, James A. 1969. Prescribed burning on recreation areas in New Jersey: history, objectives, influence, and technique. In: Proceedings, annual Tall Timbers fire ecology conference; 1969 April 10-11; Tallahassee, FL. No. 9. Tallahassee, FL: Tall Timbers Research Station: 251-269. 
24. Dalke, Paul D.; Sime, Palmer R. 1941. Food habits of the eastern and New England cottontails. Journal of Wildlife Management. 5: 216-228. 
25. DeGraaf, Richard M.; Yamasaki, Mariko. 2001. New England wildlife: habitat, natural history, and distribution. Hanover, NH: University Press of New England. 467 p. 
26. Dey, Daniel. 2002. The ecological basis for oak silviculture in eastern North America. In: McShea, William J.; Healy, William M., eds. Oak forest ecosystems: Ecology and management for wildlife. Baltimore, MD: The Johns Hopkins University Press: 60-79. 
27. Donohue, Kathleen; Foster, David R.; Motzkin, Glenn. 2000. Effects of the past and the present on species distribution: land-use history and demography of wintergreen. Journal of Ecology. 88(2): 303-316. 
28. Duchesne, Luc C.; Hawkes, Brad C. 2000. Fire in northern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 35-51. 
29. Ducousso, A.; Michaud, H.; Lumaret, R. 1993. Reproduction and gene flow in the genus Quercus L. Annales des Sciences Forestieres. 50(Suppl. 1): 91s-106s. 
30. Duncan, Wilbur H.; Duncan, Marion B. 1987. The Smithsonian guide to seaside plants of the Gulf and Atlantic coasts from Louisiana to Massachusetts, exclusive of lower peninsular Florida. Washington, DC: Smithsonian Institution Press. 409 p. 
31. Duncan, Wilbur H.; Duncan, Marion B. 1988. Trees of the southeastern United States. Athens, GA: The University of Georgia Press. 322 p. 
32. Dunwiddie, Peter W. 1990. Postglacial vegetation history of coastal islands in southeastern New England. National Geographic Research. 6(2): 178-195. 
33. Dunwiddie, Peter W. 1991. Forest history and composition of Halfway Pond Island, Plymouth County, Massachusetts. Rhodora. 93(876): 347-360. 
34. Dunwiddie, Peter W. 1998. Ecological management of sandplain grasslands and coastal heathlands in southeastern Massachusetts. In: Pruden, Teresa L.; Brennan, Leonard A., eds. Fire in ecosystem management: shifting the paradigm from suppression to prescription: Proceedings, Tall Timbers fire ecology conference; 1996 May 7-10; Boise, ID. No. 20. Tallahassee, FL: Tall Timbers Research Station: 83-93. 
35. Dunwiddie, Peter W.; Zaremba, Robert E.; Harper, Karen A. 1996. A classification of coastal heathlands and sandplain grasslands in Massachusetts. Rhodora. 98(894): 117-145. 
36. Eberhardt, Robert W.; Foster, David E.; Motzkin, Glenn; Hall, Brian. 2003. Conservation of changing landscapes: vegetation and land-use history of Cape Cod National Seashore. Ecological Applications. 13(1): 68-84. 
37. Elias, Thomas S.; Dykeman, Peter A. 1982. Field guide to North American edible wild plants. New York: Outdoor Life Books. 286 p. 
38. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. 
39. Farmer, R. E., Jr. 1980. Comparative analysis of 1st-year growth in six deciduous tree species. Journal of Forest Research. 10: 35-41. 
40. Farmer, R. E., Jr. 1981. Early growth of black cherry, oaks, and yellow-poplar in southern Appalachian plantings. Tree Planters' Notes. 32(3): 12-14. 
41. Fleming, Gary P. 2003. Ecological communities of the northern Virginia Blue Ridge, [Online]. Richmond, VA: Virginia Department of Conservation and Recreation, Division of Natural Heritage (Producer). Available: http://www.dcr.virginia.gov/dnh/Noblueridge.pdf [2005, November 21]. 
42. Forman, Richard T. T.; Boerner, Ralph E. 1981. Fire frequency and the pine barrens of New Jersey. Bulletin of the Torrey Botanical Club. 108(1): 34-50. 
43. Forrester, Jodi A.; Leopold, Donald J.; Hafner, Sasha D. 2005. Maintaining critical habitat in a heavily managed landscape: effects of power line corridor management on Karner blue butterfly (Lycaeides melissa samuelis) habitat. Restoration Ecology. 13(3): 488-498. 
44. Foster, David R.; Hall, Brian; Barry, Sylvia; Clayden, Susan; Parshall, Tim. 2002. Cultural, environmental and historical controls of vegetation patterns and the modern conservation setting on the island of Martha's Vineyard, USA. Journal of Biogeography. 29(10/11): 1381-1400. 
45. Franklin, Misty A.; Finnegan, John T., ed. 2004. Natural Heritage Program list of rare plant species of North Carolina, [Online]. Raleigh, NC: Department of Environment and Natural Resources, Office of Conservation and Community Affairs (Producer). Available: http://www.ncnhp.org/Images/Other%20Publications/2004%20Rare%20Plant%20List.pdf [2006, March 20]. 
46. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. 
47. Gibson, David J.; Collins, Scott L.; Good, Ralph E. 1988. Ecosystem fragmentation of oak-pine forest in the New Jersey pinelands. Forest Ecology and Management. 25: 105-122. 
48. Gleason, Henry A.; Cronquist, Arthur. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York: New York Botanical Garden. 910 p. 
49. Glover, Fred A. 1948. Winter activities of wild turkey in West Virginia. Journal of Wildlife Management. 12(4): 416-427. 
50. Govaerts, Rafael; Frodin, David G. 1998. World checklist and bibliography of Fagales (Betulaceae, Corylaceae, Fagaceae and Tricodendraceae). Kew, England: The Royal Botanic Gardens. 497 p. 
51. Grand, Joanna; Cushman, Samuel A. 2003. A multi-scale analysis of species-environment relationships: breeding birds in a pitch pine--scrub oak (Pinus rigida--Quercus ilicifolia) community. Biological Conservation. 112(3): 307-317. 
52. Grand, Joanna; Mello, Mark J. 2004. A multi-scale analysis of species-environment relationships: rare moths in a pitch pine--scrub oak (Pinus rigida--Quercus ilicifolia) community. Biological Conservation. 119(4): 495-506. 
53. Greller, Andrew M. 1988. Deciduous forest. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. Cambridge; New York: Cambridge University Press: 288-316. 
54. Griffiths, Megan E.; Orians, Colin M. 2003. Responses of common and successional heathland species to manipulated salt spray and water availability. American Journal of Botany. 90(12): 1720-1728. 
55. Grimm, William Cary. 1967. Recognizing native shrubs. Camping Journal. September: 49-61. 
56. Hallisey, Dennis M.; Wood, Gene W. 1976. Prescribed fire in scrub oak habitat in central Pennsylvania. Journal of Wildlife Management. 40(3): 507-516. 
57. Halls, Lowell K., ed. 1977. Southern fruit-producing woody plants used by wildlife. Gen. Tech. Rep. SO-16. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Region, Southern Forest Experiment Station, Southeastern Area, State and Private Forestry. 235 p. 
58. Hann, Wendel; Havlina, Doug; Shlisky, Ayn; [and others]. 2005. Interagency fire regime condition class guidebook. Version 1.2, [Online]. In: Interagency fire regime condition class website. U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior; The Nature Conservancy; Systems for Environmental Management (Producer). Variously paginated [+ appendices]. Available: http://www.frcc.gov/docs/126.96.36.199/Complete_Guidebook_V1.2.pdf [2007, May 23]. 
59. Hendrickson, William H. 1972. Perspective on fire and ecosystems in the United States. In: Fire in the environment: Symposium proceedings; 1972 May 1-5; Denver, CO. FS-276. [Washington, DC]: U.S. Department of Agriculture, Forest Service: 29-33. In cooperation with: Fire Services of Canada, Mexico, and the United States; Members of the Fire Management Study Group; North American Forestry Commission; FAO. 
60. Hutnik, Russell J. 1980. Bear oak. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 40-41. 
61. Jensen, Richard J.; Knops, Judith F. 1980. Patterns of morphological and phenolic variation in a hybridizing population of Quercus. Proceedings, Indiana Academy of Science. 89: 353. 
62. Jordan, Marilyn J. 1975. Effects of zinc smelter emissions and fire on a chestnut-oak woodland. Ecology. 56: 78-91. 
63. Jordan, Marilyn J.; Patterson, William A., III; Windisch, Andrew G. 2003. Conceptual ecological models for the Long Island pitch pine barrens: implications for managing rare plant communities. Forest Ecology and Management. 182(1-2): 151-168. 
64. Kartesz, John T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. 1st ed. In: Kartesz, John T.; Meacham, Christopher A. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Chapel Hill, NC: North Carolina Botanical Garden (Producer). In cooperation with: The Nature Conservancy; U.S. Department of Agriculture, Natural Resources Conservation Service; U.S. Department of the Interior, Fish and Wildlife Service. 
65. Kaufman, Herbert P. 1942. A study of forest-wildlife problems. Annual Progress Report: Pittmann-Robertson Project 9R; Federal Aid in Wildlife Restoration. Harrisburg, PA: Pennsylvania Game Commission; Washington, DC: U.S. Department of Interior, Fish and Wildlife Service. 38 p. 
66. Killingbeck, Keith T.; Costigan, Steve A. 1988. Element resorption in a guild of understory shrub species: niche differentiation and resorption thresholds. Oikos. 53: 366-374. 
67. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. 
68. Kudish, Michael. 1992. Adirondack upland flora: an ecological perspective. Saranac, NY: The Chauncy Press. 320 p. 
69. Kurczewski, Frank E.; Boyle, Hugh F. 2000. Historical changes in the pine barrens of central Suffolk County, New York. Northeastern Naturalist. 7(2): 95-112. 
70. LANDFIRE Rapid Assessment. 2005. Reference condition modeling manual (Version 2.1), [Online]. In: LANDFIRE. Cooperative Agreement 04-CA-11132543-189. Boulder, CO: The Nature Conservancy; U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior (Producers). 72 p. Available: http://www.landfire.gov/downloadfile.php?file=RA_Modeling_Manual_v2_1.pdf [2007, May 24]. 
71. LANDFIRE Rapid Assessment. 2007. Rapid assessment reference condition models. In: LANDFIRE. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Lab; U.S. Geological Survey; The Nature Conservancy (Producers). Available: http://www.landfire.gov/models_EW.php 
72. Landis, R. Matthew; Gurevitch, Jessica; Fox, Gordon A.; Fang, Wei; Taub, Daniel R. 2005. Variation in recruitment and early demography in Pinus rigida following crown fire in the pine barrens of Long Island, New York. Journal of Ecology. 93(3): 607-617. 
73. Lanner, Ronald M. 1996. The pine birds. In: Lanner, Ronald M. Made for each other: a symbiosis of birds and pines. New York: Oxford University Press: 32-37. 
74. Latham, Roger Earl; Thompson, John E.; Riley, Sarah A.; Wibiralske, Anne W. 1996. The Pocono till barrens: shrub savanna persisting on soils favoring forest. Bulletin of the Torrey Botanical Club. 123(4): 330-349. 
75. Laycock, William A. 1967. Distribution of roots and rhizomes in different soil types in the Pine Barrens of New Jersey. Geological Survey Professional Paper 563-C. Washington, DC: U.S. Department of the Interior, Geological Survey. 29 p. 
76. Little, Elbert L., Jr. 1977. Atlas of United States trees. Volume 4. Minor eastern hardwoods. Misc. Pub. No. 1342. Washington, DC: U.S. Department of Agriculture, Forest Service. 17 p. 
77. Little, Elbert L., Jr. 1979. Checklist of United States trees (native and naturalized). Agric. Handb. 541. Washington, DC: U.S. Department of Agriculture, Forest Service. 375 p. 
78. Little, S. 1946. The effects of forest fires on the stand history of New Jersey's pine region. Forest Management Paper No. 2. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 43 p. 
79. Little, S. 1964. Fire ecology and forest management in the New Jersey pine region. In: Proceedings, 3rd annual Tall Timbers fire ecology conference; 1964 April 9-10; Tallahassee, FL. No. 3. Tallahassee, FL: Tall Timbers Research Station: 35-59. 
80. Little, S.; Moore, E. B. 1949. The ecological role of prescribed burns in the pine-oak forests of southern New Jersey. Ecology. 30(2): 223-233. 
81. Little, Silas. 1952. Effects of forest fires on upland sites in the pine region of southern New Jersey. Leaflet 100. New Brunswick, NJ: The State University of New Jersey, College of Agriculture, Experiment Station. 8 p. 
82. Little, Silas; Moorhead, George R.; Somes, Horace A. 1958. Forestry and deer in the pine region of New Jersey. Stn. Pap. No. 109. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 33 p. 
83. Lutz, Harold J. 1934. Ecological relations in the pitch pine plains of southern New Jersey. Bulletin No. 38. New Haven, CT: Yale University, School of Forestry. 80 p. 
84. May, Jeffrey D.; Killingbeck, Keith T. 1995. Effects of herbivore-induced nutrient stress on correlates of fitness and nutrient resorption in scrub oak (Quercus ilicifolia). Canadian Journal of Forest Research. 25: 1858-1864. 
85. McCormick, Jack. 1998. The vegetation of the New Jersey Pine Barrens. In: Forman, Richard T. T., ed. Pine Barrens: ecosystem and landscape. New Brunswick, NJ: Rutgers University Press: 229-243. 
86. McNab, Henry W. 1988. Hardwoods and site quality. In: Smith, H. Clay; Perkey, Arlyn W.; Kidd, William E., Jr., eds. Workshop proceedings: Guidelines for regenerating Appalachian hardwood stands; 1988 May 24-26; Morgantown, WV. SAF Publ. 88-03. Morgantown, WV: West Virginia University Books: 226-240. 
87. McNamara, E. F. 1960. Sprouting capacity of scrub oak. Journal of Forestry. 58(7): 563-564. 
88. Milne, Bruce T. 1985. Upland vegetational gradients and post-fire succession in the Albany Pine Bush, New York. Bulletin of the Torrey Botanical Club. 112(1): 21-34. 
89. Moerman, Dan. 2003. Native American ethnobotany: A database of foods, drugs, dyes, and fibers of Native American peoples, derived from plants, [Online]. Dearborn, MI: University of Michigan (Producer). Available: http://www.umd.umich.edu/ [2006, April 14]. 
90. Moldenke, Harold N. 1953. Notes on new and noteworthy plants. XVI. Phytologia. 4(4): 285-295. 
91. Moore, E. B. 1939. Forest management in New Jersey. Trenton, NJ: Department of Conservation and Development, Division of Forests and Parks. 55 p. 
92. Morimoto, David C.; Wasserman, Fred E. 1991. Intersexual and interspecific differences in the foraging behavior of rufous-sided towhees, common yellowthroats and prairie warblers in the pine barrens of southeastern Massachusetts. Journal of Field Ornithology. 62(4): 436-449. 
93. Motzkin, G.; Patterson, W. A., III; Foster, D. R. 1999. A historical perspective on pitch pine - scrub oak communities in the Connecticut Valley of Massachusetts. Ecosystems. 2(3): 255-273. 
94. Motzkin, Glenn; Eberhardt, Robert; Hall, Brian; Foster, David R.; Harrod, Jonathan; MacDonald, Dana. 2002. Vegetation variation across Cape Cod, Massachusetts: environmental and historical determinants. Journal of Biogeography. 29: 1439-1454. 
95. Motzkin, Glenn; Foster, David R. 2002. Grasslands, heathlands and shrublands in coastal New England: historical interpretations and approaches to conservation. Journal of Biogeography. 29(10-11): 1569-1590. 
96. Motzkin, Glenn; Foster, David; Allen, Arthur; [and others]. 1996. Controlling site to evaluate history: vegetation patterns of a New England sand plain. Ecological Monographs. 66(3): 345-365. 
97. Noss, Reed F.; LaRoe, Edward T., III; Scott, J. Michael. 1995. Endangered ecosystems of the United States: a preliminary assessment of loss and degradation. Biological Report 28. Washington, DC: U.S. Department of the Interior, National Biological Services. 58 p. 
98. Olsson, Hans. 1998. Vegetation of the New Jersey Pine Barrens: a phytosociological classification. In: Forman, Richard T. T., ed. Pine Barrens: ecosystem and landscape. New Brunswick, NJ: Rutgers University Press: 245-263. 
99. Olsvig, Linda S.; Cryan, John F.; Whittaker, Robert H. 1998. Vegetational gradients of the pine plains and barrens of Long Island, New York. In: Forman, Richard T. T., ed. Pine Barrens: ecosystem and landscape. New Brunswick, NJ: Rutgers University Press: 265-281. 
100. Preston, Richard J., Jr. 1948. North American trees. Ames, IA: The Iowa State College Press. 371 p. 
101. Radford, Albert E.; Ahles, Harry E.; Bell, C. Ritchie. 1968. Manual of the vascular flora of the Carolinas. Chapel Hill, NC: The University of North Carolina Press. 1183 p. 
102. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. 
103. Rice, Steven K.; Westerman, Bryant; Federici, Robert. 2004. Impacts of the exotic, nitrogen-fixing black locust (Robinia pseudoacacia) on nitrogen-cycling in a pine - oak ecosystem. Plant Ecology. 174(1): 97-107. 
104. Robinson, George R.; Handel, Steven N. 1993. Forest restoration on a closed landfill: rapid addition of new species by bird dispersal. Conservation Biology. 7(2): 271-278. 
105. Ruffner, Charles M. 1997. Early plant succession following wildfire in Pennsylvania's mixed-oak woodlands. In: Greenlee, Jason M., ed. Proceedings, 1st conference on fire effects on rare and endangered species and habitats; 1995 November 13-16; Coeur d'Alene, ID. Fairfield, WA: International Association of Wildland Fire: 239-244. 
106. Seischab, Franz K.; Bernard, John M. 1996. Pitch pine (Pinus rigida Mill.) communities in the Hudson Valley region of New York. The American Midland Naturalist. 136(1): 42-56. 
107. Seymour, Frank Conkling. 1982. The flora of New England. 2nd ed. Phytologia Memoirs 5. Plainfield, NJ: Harold N. Moldenke and Alma L. Moldenke. 611 p. 
108. Sidelinger, John E. 1977. Composition and structure of vegetation and wildlife utilization of a scrub oak forest following a prescribed burn. University Park, PA: Pennsylvania State University. 93 p. Thesis. 
109. Simon, Chris; Karban, Richard; Lloyd, Monte. 1981. Patchiness, density, and aggregative behavior in sympatric allochronic populations of 17-year cicadas. Ecology. 62(6): 1525-1535. 
110. Smith, David Wm. 1993. Oak regeneration: the scope of the problem. In: Loftis, David L.; McGee, Charles E., eds. Oak regeneration: Serious problems, practical recommendations: Symposium proceedings; 1992 September 8-10; Knoxville, TN. Gen. Tech. Rep. SE-84. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 40-52. 
111. Stanek, W.; State, D. 1978. Equations predicting primary productivity (biomass) of trees, shrubs and lesser vegetation based on current literature. [Ottawa]: Environment Canada, Forestry Service. 58 p. On file with: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 
112. Stebbins, G. L., Jr.; Matzke, E. B.; Epling, C. 1947. Hybridization in a population of Quercus marilandica and Quercus ilicifolia. Evolution. 1(1/2): 79-88. 
113. Steffen, David E.; Lafon, Nelson W.; Norman, Gary W. 2002. Turkeys, acorns, and oaks. In: McShea, William J.; Healy, William M., eds. Oak forest ecosystems: Ecology and management for wildlife. Baltimore, MD: The Johns Hopkins University Press: 241-255. 
114. Stephenson, S. N. 1965. Vegetation change in the pine barrens of New Jersey. Bulletin of the Torrey Botanical Club. 92(2): 102-114. 
115. Stickney, Peter F. 1989. Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. FEIS workshop: Postfire regeneration. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. 
116. Strausbaugh, P. D.; Core, Earl L. 1977. Flora of West Virginia. 2nd ed. Morgantown, WV: Seneca Books, Inc. 1079 p. 
117. Swain, Albert M. 1978. Environmental changes during the past 2000 years in north-central Wisconsin: analysis of pollen, charcoal, and seeds from varved lake sediments. Quaternary Research. 10: 55-68. 
118. U.S. Department of Agriculture, Natural Resources Conservation Service. 2007. PLANTS Database, [Online]. Available: http://plants.usda.gov/. 
119. Vermont Department of Fish and Wildlife. 2005. Rare and uncommon native plants of Vermont, [Online]. Montpelier, VT: Nongame and Natural Heritage Program (Producer). Available: http://www.anr.state.vt.us/FW/FWHOME/library/Reports_and_Documents/ NonGame_and_Natural_Heritage Rare_Threatened_and_Endangered_Species/ List_of_Rare_and_Uncommon_Native_Plants_of_Vermont.pdf [2006, March 22]. 
120. Vogt, Albert Ralph. 1962. The influence of auxin on basal sprouting of oak seedlings. Columbia, MO: University of Missouri. 81 p. Thesis. 
121. Wade, Dale D.; Brock, Brent L.; Brose, Patrick H.; Grace, James B.; Hoch, Greg A.; Patterson, William A., III. 2000. Fire in eastern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 53-96. 
122. Wagner, David L.; Nelson, Michael W.; Schweitzer, Dale F. 2003. Shrubland Lepidoptera of southern New England and southeastern New York: ecology, conservation, and management. Forest Ecology and Management. 185(1-2): 95-112. 
123. Wainio, Walter W.; Forbes, E. B. 1941. The chemical composition of forest fruits and nuts from Pennsylvania. Journal of Agricultural Research. 62(10): 627-635. 
124. Westveld, Marinus; Ashman, R. I.; Baldwin, H. I.; Holdsworth, R. P.; Johnson, R. S.; Lambert, J. H.; Lutz, H. J.; Swain, Louis; Standish, Myles. 1956. Natural forest vegetation zones of New England. Journal of Forestry. 54(5): 332-338. 
125. Wheeler, A. G., Jr. 1991. Plant bugs of Quercus ilicifolia: myriads of mirids (Heteroptera) in pitch pine-scrub oak barrens. Journal of the New York Entomological Society. 99(3): 405-440. 
126. Wheeler, A. G., Jr.; Wilson, Stephen W. 1996. Planthoppers of pitch pine and scrub oak in pine barrens communities (Homoptera: Fulgoroidea). Proceedings, Entolomological Society of Washington. 98(1): 100-108. 
127. Whitney, Gordon G. 1991. Relation of plant species to substrate, landscape position, and aspect in north central Massachusetts. Canadian Journal of Forest Research. 21(8): 1245-1252. 
128. Windisch, Andrew G. 1999. Fire ecology of the New Jersey pine plains and vicinity. New Brunswick, NJ: Rutgers, The State University of New Jersey. 327 p. Dissertation. 
129. Wofford, B. Eugene. 1989. Guide to the vascular plants of the Blue Ridge. Athens, GA: The University of Georgia Press. 384 p. 
130. Wolgast, Leonard J. 1974. Bear oak. In: Gill, John D.; Healy, William M. Shrubs and vines for northeastern wildlife. Gen. Tech. Rep. NE-9. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 108-110. 
131. Wolgast, Leonard J. 1978. Effects of site quality and genetics on bear oak mast production. American Journal of Botany. 65(4): 478-489. 
132. Wolgast, Leonard J.; Stout, Benjamin B. 1977. Effects of age, stand density, and fertilizer application on bear oak reproduction. Journal of Wildlife Management. 41(4): 685-691. 
133. Wolgast, Leonard J.; Stout, Benjamin B. 1977. The effects of relative humidity at the time of flowering on fruit set in bear oak (Quercus ilicifolia). American Journal of Botany. 64(2): 159-160. 
134. Wolgast, Leonard J.; Zeide, Boris. 1983. Reproduction of trees in a variable environment. Botanical Gazette. 144(2): 260-262. 
135. Wolgast, Leonard Joseph. 1972. Mast production in scrub oak (Quercus ilicifolia) on the coastal plain of New Jersey. New Brunswick, NJ: Rutgers University. 137 p. Dissertation. 
136. Woodwell, George M. 1998. Leaky ecosystems: nutrient fluxes and succession in the Pine Barrens vegetation. In: Forman, Richard T. T., ed. Pine Barrens: ecosystem and landscape. New Brunswick, NJ: Rutgers University Press: 333-343. 
137. Yahner, Richard H. 1986. Structure, seasonal dynamics, and habitat relationships of avian communities in small even-aged forest stands. The Wilson Bulletin. 98(1): 61-82. 
138. Yahner, Richard H. 1987. Use of even-aged stands by winter and spring bird communities. Wilson Bulletin. 99(2): 218-232.