United States Department of Agriculture

Forest 

Service

Pacific Southwest 

Research Station

General Technical Report

PSW-GTR-252 May

2016

Restoring California Black Oak

Ecosystems to Promote Tribal

Values and Wildlife 

Jonathan W. Long, M. Kat Anderson, Lenya Quinn-Davidson, Ron W. Goode, 

Frank K. Lake, and Carl N. Skinner 
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usda.gov. 

USDA is an equal opportunity provider, employer, and lender.

Authors 

Jonathan W. Long is an ecologist, Pacific Southwest Research Station, 1731 

Research Park Dr., Davis, CA 95618; M. Kat Anderson is an ethnoecologist, U.S. 

Department of Agriculture, Natural Resources Conservation Service, 430 G St., 

Davis, CA 95616; Lenya Quinn-Davidson is a staff research associate, University 

of California Cooperative Extension, 5630 South Broadway, Eureka, CA 95503; 

Ron W. Goode is Chairman, North Fork Mono Tribe, 13396 Tollhouse Rd, Clovis, CA 93619; Frank K. Lake is an ecologist and Carl Skinner is a geographer 

(retired), Pacific Southwest Research Station, 3644 Avtech Parkway, Redding, CA 

96002. 

Cover photograph: Tribal acorn gatherers and scientists gather underneath an 

actively tended black oak tree near North Fork, California. Photo by Jonathan Long. 
Abstract
 

Long, Jonathan W.; Anderson, M. Kat; Quinn-Davidson, Lenya; Goode, 

Ron W.; Lake, Frank K.; Skinner, Carl N. 2016. Restoring California black 

oak ecosystems to promote tribal values and wildlife. Gen. Tech. Rep. PSWGTR-252. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific 

Southwest Research Station. 110 p. 

This report synthesizes information to help promote the distinctive ecological 

and cultural benefits provided by California black oak. Production of abundant, 

high-quality acorns desired by Native Americans in California, as well as other 

valued services, requires the presence of mature, broad-crowned trees with low fuel 

levels and low pest levels. Although black oaks are vulnerable to intense fires, they 

depend on low-intensity, more frequent fires to reduce competition from conifers, 

pest loads, and build-up of fuels that promote intense fires. Traditional burning 

by Native Americans helped to promote these conditions historically; however, 

in many areas that have become overly dense, thinning, out-of-season burns, or 

relatively severe fires may be needed to reopen the forest and reduce fuel levels 

before a more customary use of fire can maintain desired outcomes. Applying a 

landscape-scale approach to black oak restoration can help sustain tribal values 

and wildlife habitat, as well as promote greater ecological resilience to drought and 

wildfire during this time of a warming climate. 

Keywords: Forest management, cultural burn, ecosystem services, landscape 

restoration, prescribed burning, resilience, traditional ecological knowledge, 

woodlands, acorns. 
Preface
 

To better understand how to restore California black oak ecosystems and the 

reasons for doing so, we convened a forum at which people with indigenous, local, 

and Western scientific perspectives were brought together in one room and across 

various field sites to enable us to learn together. We held a 21⁄2 day meeting in July 

2013, coincident with the nearby Aspen Fire. In attendance were members of several tribes from in and around the community of North Fork, California; scientists 

from the U.S. Forest Service Pacific Southwest Research Station (PSW) and the 

USDA Natural Resources Conservation Service; managers from the Sierra National 

Forest, particularly its Bass Lake Ranger District; and academic researchers (see 

“Acknowledgments”). At the start of that meeting, Bass Lake District Ranger 

Dave Martin (now retired) noted that he and many of his fellow forest managers 

had come to recognize that California black oak was an important indicator of 

forest health, and that they were eager to work with tribes in collaborative forest 

restoration. Participants at the workshop offered insights into the dynamics of 

forest ecosystems with black oak, including both ecological and social elements. 

From those discussions emerged key topics and ideas, many of which are reviewed 

in this report. At a subsequent meeting of the Northern California Prescribed Fire 

Council, Danny Manning from the Greenville Rancheria delivered a presentation 

on the importance of black oaks to Maidu people, which has been condensed into 

a short section in this report. Also included are references that reinforce, expand 

upon, or contextualize many of the observations and understandings that were 

shared at these meetings. This report focuses on the Sierra Nevada, where black 

oak’s role is particularly distinctive and important. Because many of the specific 

relationships between tending practices and ecological outcomes have not been 

validated or quantified in experimental settings, they represent hypotheses that may 

warrant further exploration, particularly through field experiments where feasible. 

Further monitoring and research partnerships among scientists, land managers, and 

tribal peoples will improve understanding of roles, values, and prospective restoration strategies across the full range of black oak, and ultimately help to secure the 

benefits of black oak for future generations. 

ii 
Summary 

California black oak provides distinctive benefits to wildlife and people throughout 

its wide range in California and southern Oregon, making it both an ecological 

keystone and a cultural keystone for many California tribes. Production of abundant, high-quality acorns as well as other valued services requires the presence of 

mature, broad-crowning trees with low fuel levels and low pest levels. Although 

black oaks are vulnerable to intense fires, they depend on low-intensity, more 

frequent fires to reduce competition from conifers, pest loads, and build-up of 

fuels that promote intense fires. Native American tending of black oak trees not 

only facilitated such conditions, but it also exerted effects on the broader ecosystem through an array of food webs and fire-related interactions. Restoring the 

desired condition of mature, broad-crowning, productive oaks will depend upon 

reestablishing a more frequent fire regime. However, in many areas that have 

become overly dense, thinning, out-of-season burns, or relatively severe fires may 

be needed to reopen the forest and reduce fuel levels before a more customary 

use of fire can maintain desired outcomes. In addition, in some stands, treatments 

intended to enhance acorn production may conflict with policies that require maintaining high forest canopy cover and decadent structures to support some sensitive 

wildlife species. Despite this apparent conflict, a landscape-scale approach may 

support both acorn gathering and old forest-associated wildlife species as management objectives. Developing strategies to reconcile multiple objectives for black 

oak ecosystems would likely benefit from an adaptive management approach that 

engages tribes and other land managers. Restoration of California black oak would 

not only sustain tribal values and wildlife habitat, but it would also promote greater 

ecological resilience to drought and wildfire during this time of a warming climate. 

iii 
 

 

 

 

 

 

 

Contents
 

1 Introduction 

1 Black Oak Characteristics 

1 Description 

4 Hybrids 

6 Distribution 

7 Associated Tree Species 

7 Distinctive Qualities 

11 Values for Biological Diversity and Food Webs 

11 Ecological Keystone 

11 Plant Diversity 

12 Fungal Diversity 

12 Animal Diversity 

14 Values for Humans 

14 Wood Products and Other Material Goods 

17 Game 

18 Food 

21 Cultural Keystone 

26 Relationships Between Fire and Black Oak 

26 Goldilocks Principle 

27 Cultural Burns 

29 Studies of Past Fire Frequency in Forests With Black Oak 

30 Trends 

30 Declines of Black Oak 

34 Acorn Production on Public Lands 

35 Regeneration Concerns 

37 Concerns Regarding Future Climate 

38 Desired Conditions 

40 Conditions Associated With Tribal Values 

46 Stand, Community, and Landscape Scales 

48 Conditions Associated With California Spotted Owls and Fisher 

iv 
 

51 Impacts of Fires and Tending Practices 

51 Wildfires 

52 Recovery Following Wildfires 

54 Prescribed Burns 

55 Fire Susceptibility of Encroached Oaks 

55 Effects of Season of Prescribed Burning 

56 Effects of Cultural Burns 

62 Production of Epicormic Branches or Root Sprouts 

63 Effects of Other Native American Tending Practices 

64 Approaches for Reclaiming Degraded Stands 

64 Spring Burns and Multiple Burns 

65 Corrective Fires 

65 Thinning Conifers to Release Oaks 

68 Thinning Resprouts 

68 Promoting Black Oak When Replanting Conifers After Wildfires 

69 Protecting Legacy Black Oaks 

70 Other Treatments 

70 A Landscape Strategy for Intensive Black Oak Restoration 

70 Considering the Landscape Context for Acorn Gathering 

74 Avoiding Conflicts With Sensitive Species 

75 Conclusions: Opportunities to Increase Socioecological Resilience 

75 Promoting Ecological Diversity 

76 Promoting Resilience to Climate-Related Stressors 

76 Promoting Adaptive Management 

78 Supporting Community Engagement in Restoration 

80 Acknowledgments 

81 List of Organisms With Common, Scientific, and Western Mono 

(Nium) Names and Uses 

85 English Equivalents 

85 References 

108 Glossary 

v 
Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Introduction
 

California black oak (Quercus kelloggii) is one of the most widespread and important oaks of California forests. This report synthesizes information needed to 

develop strategies for restoring black oak to support tribal values while considering 

other benefits such as habitat for diverse wildlife species. Much scientific information has been generated about black oak ecology and the species’ importance 

to California native peoples (Anderson 1993, 2005; McDonald 1969, 1990). This 

report1 builds upon that foundation by laying out a strategy to restore desired 

conditions for black oaks within the range where native peoples have traditionally 

used fire to tend this important species. Most research on this species has been 

conducted in the Sierra Nevada and southern Cascade Range, although much of the 

information should generally apply across its distribution (fig. 1). 

The report has four main areas of emphasis: 

•	 Explaining why black oak can be considered both an ecological and cultural keystone species. 

•	 Describing its historical relationship with fire, current trends, and a desired 

condition for black oaks and their forest habitat to support tribal values and 

wildlife. 

•	 Highlighting the effects of fire and traditional Native American management practices on forest ecosystems with black oak. 

•	 Outlining strategies for achieving desired conditions for black oaks and 

their habitat given current forest conditions. 

Black Oak Characteristics 

Description 

In 1857, California black oak was given the scientific name Quercus kelloggii by 

John Strong Newberry in honor of Albert Kellogg, who prepared botanical illustrations of this and other oaks of western North America (fig. 2). California black oak 

is a “red” or “black” oak belonging to the section Lobatae, whose members are 

characterized by bitter acorns owing to their high tannic acid levels; relatively dark 

bark, jagged leaves; and reddish-brown wood (Messner 2011). Black oak is noted for 

its deeply lobed, alternate leaves that turn from green to yellow and then brown in 

1

 A much abbreviated version of this report was presented at the 7th California Oak 

Symposium: Managing Oak Woodlands in a Dynamic World, Nov. 3–6, 2014, Visalia, 

California, and published in the conference proceedings as Long, J.W.; Quinn-Davidson, 

L.N.; Goode, R.W.; Lake, F.K.; Skinner, C.N. (2015). Restoring California black oak to 

support tribal values and wildlife. In: Standiford, R.B., ed. Proceedings of the 7th California Oak Symposium. Gen. Tech. Rep. PSW-GTR-251 Albany, CA: U.S. Department of 

Agriculture, Forest Service, Pacific Southwest Research Station: 113–122 

1 
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GENERAL TECHICAL REPORT PSW-GTR-252 #*

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Observations of California black oak 

Oracle oak without black oak present 

Collection of black oak in Sierra de Juarez, Mexico 

Forest Inventory and Analysis record for black oak 

State boundaries 

Predominant range of California black oak 

Map compiled by Jonathan Long from sources in the text 

Figure 1—Range of California black oak, including populations of oracle oak that were observed 

without black oaks. 
3 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Figure 2—Illustration of California black oak by Albert Kellogg, from Greene (1889). 
4 

GENERAL TECHICAL REPORT PSW-GTR-252

fall. The bark on young trees is thin and smooth, but it becomes moderately thick, 

deeply fissured, and platy as the tree grows old (Fryer 2007). Mature trees produce 

large acorns (up to 4.4 cm long) that typically take 2 years to develop (McDonald 

1969). The acorns are held within deeply cupped caps that have fine, light-brown 

scales (McDonald 1969). 

Black oaks are the tallest true oaks in the mountains of California and Oregon, 

with individual trees typically growing about 23 m tall (McDonald 1990). The 

current “national champion” black oak, in Groveland, California, in the heart of the 

Sierra Nevada, is 28 m tall with a girth of 726 cm (UFEI 2015). The current record 

tree in Oregon is reported as 32 m tall but only 602 cm in girth (Ascending the 

Giants 2015). A previous national champion from the Siskiyou National Forest of 

Oregon was 38 m tall and 859 cm in circumference (Fryer 2007) before it died, so 

black oaks larger than the current record trees may well occur elsewhere.2 

Hybrids 

Black oak, which is completely deciduous, hybridizes with several evergreen oaks 

in the section Lobatae that overlap with it in the lower elevation of its range. The 

resulting crosses retain sparse foliage in the winter. For example, a cross with interior live oak (Quercus wislizeni) creates oracle oak (Quercus × morehus). Oracle 

oaks are typically smaller than black oaks, as the largest oracle oak listed in the 

big tree registry is only 18 m tall (UFEI 2015); however, Phil McDonald and Carl 

Skinner have identified an oracle oak over 27 m tall in a city park at Shasta Lake 

city (fig. 3). A cross with coast live oak (Quercus agrifolia) forms Chase oak 

(Quercus × chasei), as reported in Santa Clara County from a tree that was 

much larger than an oracle oak (McMinn et al. 1949). A cross with the interior form 

of coast live oak (Quercus agrifolia var. oxyadenia) results in Gander oak (Quercus 

× ganderi ), reported from San Diego County (Wolf 1944). A hybrid black oak 

reported from Santa Cruz island has an unknown parent, which could possibly 

be coast live oak, or even Santa Cruz island oak (Quercus parvula var. parvula), 

which, as a variety of Shreve’s oak (Quercus parvula), is closely related to interior 

live oak (Dodd et al. 2002). 

Although these hybrid black oaks are generally considered oddities of minor 

significance, they have special qualities owing to their semi-evergreen nature, 

and the wood of oracle oak has been described as superior to that of either parent 

(Jepson 1910). Furthermore, in parts of California, the oracle oak has been reported 

2 Ritter, M. 2015. Personal communication. Professor, California Polytechnic State 

University, San Luis Obispo, CA 93407. 
5 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife John Duckett 

Figure 3—This tall oracle oak in Claire Engle Park in the city of Shasta Lake could be 

nominated the state champion. Note the persistent leaves in the middle of winter. 
6 

GENERAL TECHICAL REPORT PSW-GTR-252

from locations without well-established observations of the pure parent black oak 

(fig. 1). This unusual distribution warrants attention when considering restoration 

in outlying locations such as Santa Cruz Island, where the local population of either 

black oak or oracle oak may have been extirpated. Therefore, restoration strategies 

for black oak should consider the importance of these hybrids. 

Distribution 

Black oak has a wide range across California and into south-central Oregon, with 

an isolated population reported from Baja California, Mexico. The map in figure 

1 incorporates range data from the U.S. Geological Survey (1999), isolated occurrences of oracle oak from Griffin and Critchfield (1976), a collection of California 

black oak in 1987 at La Matanza Ranch in the Sierra de Juarez of Baja California 

(Carrillo N.d.), and forest inventory plots containing California black oak (USDA 

FS 2014). The hybrid oracle oak is widespread in California and south-central 

Oregon. A black oak hybrid was collected on Santa Cruz Island in the 1930s, 

although the identity of the other parent species needs to be confirmed, and that 

particular specimen is reported to have died. 

The distribution of black oak reflects interacting influences of precipitation, 

elevation, aspect, and soil qualities. Specifically, McDonald (1990) reported that the 

tree occurs at elevations ranging from 61 m to over 2438 m; in north-central California, black oak occurs primarily in draws or on north slopes at elevations below 

305 m, on all aspects between 762 to 915 m elevation, and not on north- and eastfacing slopes above 1067 m. He also reported that black oak is found on west-facing 

slopes with sufficient moisture in the southernmost mountains. Since the Neogene 

period (23 to 2.6 million years ago), the range of black oak has apparently contracted as the climate has become drier in areas formerly occupied by its presumed 

ancestor, Quercus pseudolyrata, which is well represented in high-elevation fossil 

floras of California, Oregon, Nevada, and Idaho (Chaney 1918, McElwain 2004, 

Smith 1941). 

In recent decades, black oaks were estimated to occur on 2.9 million ha of 

forest land, which is approximately 7 percent of all lands in California (Waddell 

and Barrett 2005). That area was evenly divided between private lands and national 

forest land (Waddell and Barrett 2005). The species is most abundant in woodlands, 

ponderosa pine (Pinus ponderosa) forests, and mixed-conifer forests along the west 

side of the Sierra Nevada Mountains and along lower slopes in fairly dry sections of 

the Klamath and Cascade Mountains (McDonald 1990). 
7 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Associated Tree Species 

Throughout that range, black oak commonly grows in stands that are dominated 

by other species. As a result, only 506 000 ha in California were classified as the 

black oak forest type in a 1990 inventory (Waddell and Barrett 2005), indicating 

that the species was the predominant tree in less than one-fifth of the areas where 

it occurred. 

Black oak grows with a wide variety of conifer trees, hardwood trees, and 

shrubs, although its most common associates are ponderosa pine, Pacific madrone 

(Arbutus menziesii), and tanoak (Notholithocarpus densiflorus) (McDonald 1990). 

Particularly in the northern and coastal part of its range, black oak often grows 

with Douglas-fir (Pseudotsuga menziesii var. menziesii) and with Oregon white 

oak (Quercus garryana). Within lower elevation woodlands, black oak occurs with 

other oaks, leading to several of the previously mentioned hybrids. Canyon live oak 

(Quercus chrysolepis) also grows within much of the range of black oak, although it 

tolerates drier and harsher sites. 

Tanoak is a distantly related hardwood species that also produces acorns and 

supports diverse wildlife species and tribal cultural uses, but tanoak is more concentrated along the coast than black oak. Although there are more tanoak trees than 

black oaks in California, their stem volumes are similar (Christensen et al. 2008), 

and the two species are tied for third in area of hardwood forest types in California 

behind blue oak (Quercus douglasii) and canyon live oak. However, black oak had 

the most large trees (≥28 cm diameter at breast height [dbh]) of any hardwood species in 1990 inventories (Waddell and Barrett 2005). 

This report devotes particular attention to the role of black oak as a component 

within forests dominated by conifers. Many of the management strategies discussed 

in the report apply to woodland areas, but the challenges are more complex within 

the mixed-conifer forests where black oaks, and especially mature trees, appear 

vulnerable to long-term declines. Within the typical range of mixed-conifer forests, 

black oaks occur as isolated trees, in clumps, or in small groves, but very rarely in 

large open-grown stands (McDonald 1969). However, historical data on size, crown 

attributes, and spatial pattern of black oak are more limited than for conifers. The 

section on “trends” considers the extent and condition of black oak in historical and 

recent periods, as well as concerns regarding future climate. 

Distinctive Qualities 

Black oak has several unusual adaptations and other qualities that afford it a 

distinctive role among the many conifer species with which it grows. Many of these 
8 

GENERAL TECHICAL REPORT PSW-GTR-252

Black oaks form large, 

often well-insulated 

cavities within their 

trunks and dead or 

broken branches, 

and these cavities 

support many animals, 

including Pacific fisher. 

qualities are also important for understanding how and why Native Americans 

tended this important tree, as will be discussed further. 

Hard mast— 

Acorns are an important hard-shelled food in landscapes that are dominated by 

conifers that produce soft seeds in cones. The hard shell allows the nut meat to be 

available longer to foraging people or wildlife, although acorns may quickly germinate or become too moldy for people to eat if left in moist conditions (Anderson 

2005, McDonald 1969). Other than tanoak, sources of hard mast that are commonly 

found within the range of black oak are less productive smaller trees and shrubs, 

including chinquapins (Chrysolepis chrysophylla and Chrysolepis sempervirens), 

California hazelnut (Corlyus cornuta ssp. californica), California nutmeg (Torreya 

californica), and California buckeye (Aesculus californica). Within the lower elevations of the range of black oak are many other important masting species, including other oaks as well as native walnut (Juglans hindsii), which has a very limited 

distribution. 

Maturation time— 

Black oak trees produce acorns starting as soon as age 30, with sporadic production until they enter prime production between 80 and 100 years; prime production 

can continue until at least 200 years (McDonald 1969). According to the USDA 

woody plant seed manual (Bonner and Karrfalt 2008), the minimum age of 30 years 

needed for black oaks to produce seeds is generally much longer than competing 

conifers, including Douglas-fir (10 years), Jeffrey pine (Pinus jeffreyi) (8 years), and 

ponderosa pine (16 to 20 years), although it is faster than the time for white fir to 

produce “commercial” quantities of seed (40 years) or for sugar pine (Pinus lambertiana) to produce seeds (40 to 80 years). Note that the time needed for fire-killed 

resprouts of mature trees to produce acorns may be much shorter given that such 

trees have well-developed root systems; however, research is needed to study the 

relationship between age of resprouts and acorn production, and how management 

practices can influence production. 

Cavities— 

Black oaks form large, often well-insulated cavities within their trunks and dead 

or broken branches, and these cavities support many animals, including Pacific 

fisher (Pekania pennanti spp. pacifica), as discussed further below. Black oaks become decadent sooner and provide more cavities than do conifers of similar sizes 

(Garrison et al. 2002b), and they reportedly have a higher incidence of decay and 

breakage than other oaks with which they occur (Bolsinger 1980).
 
9 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Flammable litter— 

Black oaks produce large, curling leaves that form the most flammable litter of the 

California oaks, which helps them to carry frequent, low-intensity fire (Engber and 

Varner 2012). In a comparison with seven common conifer trees, black oak litter 

was the fastest to ignite and the quickest to spread, producing tall flames that were 

not sustained as long as most of the conifers; these qualities could facilitate frequent 

fires that cause less injury to mature trees (de Magalhães and Schwilk 2012). 

Variable fire resistance— 

Whereas young stems are highly susceptible to fire, stems of mature trees are somewhat resistant to fire, although fires may cause open scars that can make trees more 

vulnerable to future fires. Because of their relatively thin bark, black oaks are more 

sensitive to fire than the mature forms of associated conifers, such as Douglas-fir, 

ponderosa pine, sugar pine, and white fir (Abies concolor) (Skinner et al. 2006); 

however, black oaks do have thicker bark and are more resistant to fire than many 

other hardwood trees (Plumb 1980). 

Resprouting of mature trees and seedlings— 

When the aboveground parts of black oaks have been killed or even merely injured 

or exposed to increased sunlight, the trees can resprout, often rapidly, from the 

base of their trunks, forming “root-crown sprouts.” Black oaks also have the ability 

to resprout from seedlings that have been top-killed, forming “seedling-sprouts.” 

McDonald and Tappeiner (2002) noted that black oak seedlings may take more than 

7 years before they begin to grow tall stems, during which time they develop a long 

tap root. The black oak seedlings may be repeatedly top-killed and then resprout, 

and they may remain only a few inches tall for decades, especially in shaded environments. The ability of seedlings to resprout following dieback demonstrates 

an adaptation that can maintain the species until better conditions for growth arise 

(McDonald and Tappeiner 2002). Because of the capacity to resprout vigorously 

following intense stand-replacing fires or other major disturbances that remove 

the overstory, black oaks can outgrow coniferous trees and shrubs (McDonald 

1969). For example, at a productive site on the Challenge Experimental Forest, 

black oak resprouts reached 6 m tall in 10 years following a clearcutting treatment 

that increased sunlight (McDonald and Tappeiner 2002). This rapid growth helps 

to reduce soil erosion in the short-term, because quickly established shoots and 

leaves break the impact of rainwater (McDonald 1969). Over longer periods, black 

oaks can provide a nursery environment for reestablishing conifers (Barr 1946, 

McDonald and Tappeiner 2002). 
10 

GENERAL TECHICAL REPORT PSW-GTR-252

 

When black oaks are 

retained in fuelbreaks 

and other fuels 

treatment areas, they 

maintain canopy cover 

while reducing fire 

hazard. 

Drought tolerant— 

Black oaks, like many other oaks, have numerous physiological adaptations to harsh 

conditions, especially drought, including an ability to tap deep water supplies in 

fractured bedrock and to tolerate moisture stress at levels that are lethal to competing species (McDonald 1969, McDonald and Huber 1995b). Studies have shown that 

large trees, including coast live oak, can lift deep water into shallower soil mycorrhizal networks, which in turn can help seedlings withstand droughts (Bingham and 

Simard 2012, Egerton-Warburton et al. 2007). Royer et al. (2008) reported that leaf 

shapes on black oak vary with elevation, possibly because of wind speed or water 

availability, but not with mean annual temperature. This adaptation suggests that 

black oak as a species can vary its leaves over time to take advantage of differing evapotranspiration conditions. Related studies (Royer 2012, Royer et al. 2009) 

found that red maples (Acer rubrum) had the ability to alter their leaf shapes over 

short periods (not only through long-term genetic selection); however, that researcher has not studied whether black oak’s ability to change was also so fast.3 

Open crown and canopy— 

Black oaks have an open, low-density crown structure, which allows more light 

in the understory compared with conifers (Fralish 2004). This quality also makes 

them less likely to promote or carry crown fires than evergreen and densely leaved 

forest types (Skinner et al. 2006). Thus, when black oaks are retained in fuelbreaks 

and other fuels treatment areas, they maintain canopy cover while reducing fire 

hazard.
 

Deciduous— 

In contrast to evergreen oaks and most conifers in California, black oaks are dormant in winter, which reduces both transpiration and interception of precipitation 

in winter months. This trait also allows sunlight to dry fuels in the fall and winter, 

which facilitates fires when more shaded areas are too wet to burn (Knapp et al. 

2009). Furthermore, the absence of leaves on oaks during those seasons allows 

heat from fires to dissipate easily and alleviates the potential for crown scorch 

(Skinner 1995).
 

Shade intolerance— 

Mature black oaks are one of the least tolerant trees to shading within the mixedconifer forest, although very young trees may be more tolerant than pines (Bigelow 

et al. 2011, McDonald 1990). 

3 Royer, D. 2015. Personal communication. Professor, Wesleyan University, Department 

of Earth and Environmental Sciences, Exley Science Center 445 (265 Church St.), Middletown, CT 06459-0139. 
11 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Values for Biological Diversity and Food Webs 

Ecological Keystone 

Scientists who study the ecology of California black oak, and Native Americans 

who depend on its products, both regard it as a critically important species. Some 

ecologists use the term “ecological keystone” to describe species whose removal 

would cause dramatic changes in the structure and function of its biological community (De Leo and Levin 1997). Despite some historical efforts to curtail black 

oak to favor conifers in some forests, the species has proved highly resilient, and 

there do not appear to have been studies that tested effects of its removal.4 

Other scientists have proposed somewhat narrower criteria for a keystone 

species, including having disproportionately large ecosystem impacts relative to 

their abundance (Paine 1995, Power et al. 1996) and performing unique roles within 

the ecosystem (Kotliar 2000). Although the abundance of oaks makes it difficult to 

qualify them as keystones in many ecosystems (McShea and Healy 2002), Fralish 

(2004) asserted that various kinds of black oaks and hickory in hardwood forests of 

the central United States qualified as keystones because they performed disproportionate and unique roles in producing mast and high-light environments that 

facilitated persistence of other species in the community. California black oak 

similarly meets these criteria within landscapes such as the Sierra Nevada that are 

dominated by conifer trees because of hard mast, open crowns, flammable litter, 

rapid resprouting, and other distinctive qualities described in the previous section. 

Although some scientists have suggested that the concept of keystone species is 

problematic to apply to plants, there is general agreement that it is important to 

understand the complex interactions within food webs or communities (Menge and 

Freidenburg 2001, Mills et al. 1993). 

Plant Diversity 

In areas dominated by deciduous oaks, understory plant diversity can be very 

high relative to other vegetation types (Jimerson and Carothers 2002, Thysell and 

Carey 2001). Both fire and thinning treatments can promote structural and biological diversity and counteract the effects of conifer encroachment by maintaining a 

patchy array of open and closed areas (Engber et al. 2011, Wayman and North 

 2007, Webster and Halpern 2010). The high-light environment afforded by the 

relatively open crown and leaf structure of California black oaks are likely to favor 

4

 However, the spread of sudden oak death (caused by the pathogen Phytophthora ramorum) may provide a test of the importance of black oak and tanoak in parts of California 

where the disease is rampant (see Waring and O’Hara 2008). 
12 

GENERAL TECHICAL REPORT PSW-GTR-252

Acorns provide a 

unique food resource 

that can either be 

consumed directly 

by wildlife to build fat 

reserves, or stored 

for use during leaner 

times. 

understory plants in ways similar to oaks in other parts of the United States (Fralish 

2004). Although burning in these ecosystems can volatize nitrogen, an important 

nutrient, it can also promote understory plants that can fix nitrogen, including big 

deervetch (Hosackia crassifolia), deerbrush (Ceanothus integerrimus), whitethorn, 

lupines, and others (Kilgore 1973, St. John and Rundel 1976). 

Fungal Diversity 

Fungi are an important component of the biodiversity in forests with black oak, 

especially as food sources for wildlife (North et al. 1997), and because many fungi 

form mycorrhizal associations that help trees to exchange carbon, nitrogen, and 

water (Southworth et al. 2011). Although it is apparently not known whether black 

oaks support unique fungi, the frequent fires that maintain mature black oaks may 

encourage various kinds of fungi to fruit. More studies are needed to understand 

the responses to season of burn and burn severity for various species (Pilz et al. 

2007, 2004). Morels (Morchella sp.) and the charcoal-loving elf cup (Geopyxis 

carbonaria) are two examples of fungi from a group known as “Pezizales,” which 

often flourish between 6 weeks and 2 years following fires; insect disturbances that 

damage trees can also increase production of fruiting bodies (Greene et al. 2010, 

Kuo et al. 2012, Pilz et al. 2004). A study in a moist forest from British Columbia 

that experienced high-severity fire suggested that both consumption of duff (with 

remaining depths less than 4 to 5 cm) and damage to nearby trees may be necessary 

to trigger an unusually large abundance of mushrooms (Greene et al. 2010, Kuo et 

al. 2012). Some of the known fire-following fungi such as morels may form mycorrhizal associations in the absence of fire (Fujimura et al. 2005). 

Animal Diversity 

Black oaks provide a food source and habitat for a variety of wildlife species. 

Acorns provide a unique food resource that can either be consumed directly by 

wildlife to build fat reserves, or stored for use during leaner times (McShea and 

Healy 2002). Black oaks support black bear (Ursus americanus), mule/black-tail 

deer (Odocoileus hemionus), and many small mammals and their associated predators in the fall, winter, and other times when other food sources are not available 

(Mazur et al. 2013, McShea and Healy 2002). In a burned area on the Plumas 

National Forest, where the current crop of acorn and other tree nuts had failed and 

fungi were rare, Tevis (1952) reported long-eared chipmunks (Tamias quadrimaculatus) surviving on year-old black oak acorns. Dusky-footed woodrats (Neotoma 

fuscipes), an important prey species for California spotted owl (Strix occidentalis 

ssp. occidentalis), appeared in higher densities in areas with large (>33 cm dbh) 
13 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

black oaks in a mixed-conifer forest with few other sources of hard mast (Innes et 

al. 2007). Black oak acorns are also important for mountain quail (Oreortyx pictus) 

and valley quail (Callipepla californica) (Potter and Johnston 1980). The acorn 

woodpecker (Melanerpes formicivorus) depends on black oak acorns, and good 

crops have demonstrated potential to expand the bird’s breeding season (Koenig 

and Knops 1995). These woodpeckers also use mature black oaks, along with other 

large trees (fig. 4), as granaries to store acorns within productive stands (Johnson 

and Rosenberg 2006, MacRoberts and MacRoberts 1976). Abundance of bandtailed pigeons (Columba fasciata) may be an important indicator of the condition 

of black oaks because of their strong association with those trees for roosting 

and acorns (Bottorff 2007, McDonald 1969). The western gray squirrel (Sciurus 

griseus), western scrub jay (Aphelocoma californica), and Steller’s jay (Cyanocitta 

stelleri) are important consumers of acorns, but they also disperse acorns into new 

areas (Kauffman and Martin 1987, McDonald 1990). Consequently, these wildlife 

species demonstrate a strong mutualism by both eating black oak acorns and 

benefitting the trees. 

Jonathan Long 

Figure 4—Acorn woodpeckers consume black oak acorns they store in granary trees, including 

these dead ponderosa pines on the Power Fire, Eldorado National Forest. 
14 

GENERAL TECHICAL REPORT PSW-GTR-252

An 8-year study of ponderosa pine forests on the Sierra National Forest identified 19 bird species that nest in California black oak, with a greater use of live trees 

than snags (Purcell and Drynan 2008). It found that black oaks were used for 21 

percent of the nests even though they represented only 10 percent of all trees >2 m 

tall. Black oaks used for nesting were larger than other available trees in terms of 

both height (mean was 21 m) and dbh (mean was 64 cm), which points to the importance of mature trees as a resource for wildlife. Black oak provides important den 

sites for fisher (Yaeger 2005, Zielinski 2014), which also uses black oaks as resting 

sites (fig. 5) out of proportion to their availability in the southern Sierra Nevada 

(Purcell et al. 2009). Many bird species, including northern pygmy-owl (Glaucidum 

gnoma), downy woodpecker (Picoides pubescens), mountain chickadee (Poecile 

gambeli), red-breasted nuthatch (Sitta canadensis), white-breasted nuthatch (Sitta 

carolinensis), and brown creeper (Certhia americana), use dead black oak branches 

for foraging, nesting, or roosting (Garrison et al. 2002b). 

Values for Humans 

Many cultures have identified oaks as “trees of life” (Cristancho and Vining 2004). 

Acorns have been central to indigenous cultures in California for millennia (Baumhoff 1963, Stevens 2002), serving as both a direct food source and an indirect food 

source by sustaining game animals. Black oaks also have served as important 

sources of raw materials, primarily various wood products, for a wide variety of 

household goods, and for cultural inspiration (figs. 6 and 7; box 1). In addition to 

their material significance, mature oak trees have intangible sociocultural values as 

large and old denizens of forests (Blicharska and Mikusiński 2014). These values 

include the scenic appeal of fall color and the diverse form that oaks, especially 

large trees, add to conifer-dominated forests (McDonald and Huber 1995b). Favorable water flows are another benefit of restoring black oak, as the leafless crowns of 

deciduous trees reduce evapotranspiration and allow winter rain and snow to enter 

the litter and soil; consequently, conversion of stands from deciduous hardwoods to 

conifers can reduce water yields (McDonald and Huber 1995b). 

Wood Products and Other Material Goods 

California black oak has long been a source of a wide variety of household goods 

used by Native Americans (fig. 6). Black oak has also provided considerable 

economic values for more recent settlers of California, who use the wood as fuel 

and building materials, although forests have not typically been managed to produce oak lumber (Rossi 1980). McDonald and Huber (1995a) reported that there 

was potential to develop a value-added hardwood industry within the central Sierra 
15 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife Zane Miller 

Figure 5—Fisher resting in a black oak cavity formed from a dead and broken branch. 

Figure 6—Native American uses of California black oak (modified from Anderson 2007). 
16 

GENERAL TECHICAL REPORT PSW-GTR-252

A B 

C D 

E 

Figure 7—(A) Danny 

Manning cutting a black 

oak sapling, (B) bending 

the sapling into a frame, 

and then (C) the finished 

cradleboard; (D) a basket 

of black oak acorns with 

beaded necklace by Mike 

Savala, and (E) a drawing 

by Paul Cason further 

illustrate the importance of 

black oak to Maidu people. 

(Photos courtesy of Danny 

Manning) 
17 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Box 1 

Significance of California Black Oak to Maidu 

Contributed by Danny Manning, Greenville Rancheria 

Black oaks are significant to Maidu because they were/are one of our primary 

food sources. We make baskets for storing acorns, for winnowing and processing acorns, and for cooking acorns. Different families or clan groups have 

responsibility for and rights to certain trees. When babies are first born, they 

are placed in a cradleboard with a frame made from an oak branch. In Mountain Maidu country, traditionally the split rock or o’witchono at the top of the 

Feather River Canyon must be filled in each year to stop the wind from blowing in and knocking off the acorns too early. In addition, the ponderosa pine 

tree, bu’bum ch’am or wind-lessening tree, is named for its role in catching the 

wind so that it does not reach the shorter oak tree and blow the acorns down. 

Although we no longer depend on acorns the way we did in the past, Maidu 

elders warn of a future time in which we will need to know how to steward 

oaks and process acorns because “we will be happy to have acorns to eat.” 

Nevada, with black oak and canyon live oak contributing a “moderate” volume for 

potential utilization. Maximizing the economic value of hardwood lumber requires 

changes in typical methods used to process conifer lumber, in order to accommodate shorter and more variable logs, the need for slower drying (seasoning), and 

different milling requirements (Huber and McDonald 1992). 

Game 

Black oak acorns support a variety of game species, including deer, black bear, 

band-tailed pigeons, wild hogs (Sus scrofa), and wild turkeys (Meleagris gallopavo). The economic value of deer hunting associated with black oak has resulted 

in state and federal efforts to promote black oak (Skinner 1995). One study found 

that black oak contributed substantial economic values by enhancing the quality of 

deer hunting experiences (Loomis et al. 1995). Indeed, within an area that overlaps 

the Shasta-Trinity National Forest, that study found that proposed efforts to reduce 

oak basal area from an average of 3.9 m2/ha to only 2.3 or 1.1 m2/ha would reduce 

annual hunting values by $3.4 million and $5.5 million, respectively, whereas 

increasing basal area of oaks to 6.9 m2/ha would increase hunting benefits by $8 

million. Within the San Jacinto Ranger District on the San Bernardino National 
18 

GENERAL TECHICAL REPORT PSW-GTR-252

Many Native Americans 

and others are “oak 

enthusiasts” who 

continue to cultivate, 

process, and consume 

acorns today, and some 

share their skills with 

the general public. 

Forest, economists estimated that burning in habitats that include California black 

oak would generate approximately $8.65 to $17.80 per hectare of increased deer hunting benefits for the first 445 ha burned, or about 3.4 percent of the total costs of such a 

burn, with diminishing returns for larger burns (González-Cabán et al. 2003). 

Food 

Acorns have long been an important part of the Native American diet in most places 

where oaks grow. In addition to eating wild game that consumed acorns, Native 

American families would gather and store thousands of pounds of acorns. Acorns 

were sometimes used right away, but they could be stored for at least 4 years and 

possibly over a decade (Anderson 1993). Many Native Americans and others are 

“oak enthusiasts” who continue to cultivate, process, and consume acorns today, and 

some share their skills with the general public (fig. 8) (Ortiz 1991, Stevenot 2008). 

Processing acorns involves several time-consuming steps: removing the shell (fig. 9), 

winnowing to remove the skins on the shelled acorns, pounding (fig. 10) and sifting 

(fig. 11) the nuts into a fine meal, leaching the meal to remove tannins, and then 

cooking it to create an excellent acorn porridge, mush, or soup (fig. 12). Frank Lake 

Figure 8—Lois Conner Bohna 

with acorns on a tree tended by 

her grandmother, Lilly Harris, at a 

ranch near North Fork.
19 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife Kat Anderson 

Figure 9—Lois Conner Bohna cracking an acorn with a hammer stone to remove the shell. Kat Anderson 

Figure 10—Lois Conner Bohna pounding the shelled acorns into flour 

in a traditional mortar hole. 
20 

GENERAL TECHICAL REPORT PSW-GTR-252

Figure 11—Lois Conner Bohna sifting the acorn flour to separate 

the coarse grains from the fine flour. 

Kat Anderson 

Kat Anderson 

Figure 12—Black oak acorn mush or porridge, a traditional food prepared by Lois Conner
 

Bohna, North Fork Mono, in a basket made by her grandmother, Lilly Harris, circa 1920.
 
21 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Interviews with tribal elders indicate that black oak was the most favored acorn 

among many of the tribes in California, and second only to tanoak for many others 

(Appendix J in Anderson 1993) (fig. 13). A comparison with the range map (fig. 1) 

indicates that this preference even extends beyond the range of black oak in southern California. However, tribes in northern California often have preferred tanoak 

over black oak where they had access to both species, either within their territory 

or through trade (Baumhoff 1978, Holt 1946, Kroeber 1925, Schenck and Gifford 

1952). However, even in these instances, black oak may have a distinctive value, 

such as serving as a ceremonial food among the Karuk (Heffner 1984). In southern 

Oregon, black oak acorn paste was reportedly traded from California to the Rogue 

River Valley (Dixon 1907), and Sapir (1907) specifically described black oak 

acorns as the chief of the staple foods for the Takelma Native Americans. Among 

coastal tribes in southern Oregon, Barnett (1937) reported that acorns (species 

unspecified) were commonly used, but that they were especially important for the 

Tolowa (along the California border) and became less important farther north. 

Food preferences depend on a variety of influences, including individual tastes, 

accessibility, ease of processing, and traditions. California natives used acorns from 

a variety of oak species that occur within the range of black oak, including blue 

oak, interior live oak, canyon live oak, and Oregon white oak (Anderson 2009). 

Oregon white oak is widely found in southern Oregon and northern California, and 

scattered in the Sierra Nevada, but it was generally not a preferred food because 

of its sticky consistency (Dixon 1907). Desirable properties of black oak acorns 

include a large nut that is easier to deshell, pound, and grind than other nuts; a 

distinctive taste and texture; high oil content; and storability (Anderson 1993, Lee 

1998). That tribes would prefer such a tannic-rich food, which requires more time 

to process (Bettinger et al. 1997), may seem counterintuitive; however, the same 

preference for acorns from red or black oaks (section Lobatae) appears to hold for 

Native Americans from eastern North America (Messner 2011). The higher tannin 

and oil content of black oak may account for enhanced texture and flavor, better 

storage, and dietary benefits. In addition to being an important source of energy, 

acorns have been shown to have desirable qualities for controlling blood sugar 

levels (Brand et al. 1990). 

Cultural Keystone 

The opportunity to gather traditional foods and maintain associated lifeways are 

important in maintaining the biological, economic, and cultural well-being of tribal 

communities (Lynn et al. 2013). Garibaldi and Turner (2004: 4) described cultural 

keystone species as “culturally salient species that shape in a major way the cultural 

Interviews with tribal 

elders indicate that 

black oak was the most 

favored acorn among 

many of the tribes in 

California, and second 

only to tanoak for many 

others. 
22 

GENERAL TECHICAL REPORT PSW-GTR-252

Figure 13—Preferences for black oak acorns among tribal language groups. 
23 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

identity of a people, as reflected in the fundamental roles these species have in 

diet, materials, medicine, and/or spiritual practice.” Garibaldi and Turner (2004) 

proposed a list of criteria to determine the relative value of different species according to this definition (table 1). California black oak clearly meets the criteria for this 

distinction based upon its multiplicity of uses; significance in names, stories, and 

ceremonies; value for trade; and lack of substitutability for Native Americans who 

live in the heart of its range, such as the different Mono cultural groups (Aginsky 

1943, Anderson 2007). 

Cristancho and Vining (2004: 155) proposed the term “culturally defined 

keystone species” as “plant and animal species whose existence and symbolic value 

are essential to the stability of a culture over time” (table 2). The questions they 

proposed to identify these species focus on spiritual values that are harder to objectively evaluate, and it is worth noting that Garibaldi (2009) later suggested dropping 

the question about ceremonial uses owing to the sensitivity of such information. 

However, Ortiz (2008) shared several quotes from Native American acorn gatherers 

and other examples that illustrate the spiritual significance of oaks. 

Acorns have had an important role in cultural transmission and rituals, including dances, festivals, and ceremonies. At the July 2013 workshop, Ron Goode 

recounted a story explaining how acorns were used as a currency. Tribal members 

also noted that social activities often centered on acorn gathering and processing, 

with women gathering and processing while children would play games of tag 

in the trees. The trees provided shade under which women worked long hours. 

The relationships that were established to facilitate acorn harvest often governed 

broader social interactions among families and across the landscape. For example, 

people would consult one another to determine the optimum gathering sites in a 

given year, and they would often work together to complete gathering and processing activities (Anderson 1993, Bettinger et al. 1997). 

During the 2013 workshop, Ron Goode presented a representation of a food 

web associated with black oak (fig. 14). He noted a particularly strong relationship 

between oaks and squirrels, chipmunks, deer, and their predators, and he stressed 

the cascading effects associated with compromised oak health: “If trees are not 

producing abundant acorns, then squirrels, chipmunks, and deer aren’t doing as 

well. Predators aren’t going to do as well either.” This position is consistent with the 

claim by a wildlife biologist that acorn production “reigns supreme as the greatest 

contribution to the welfare of forest wildlife” (Kerns 1980: 359); evidence that deer 

populations are strongly correlated with acorn production (Leach and Hiehle 1957); 

and an assertion by a prominent wildlife researcher (Barrett 1980) that several 

mammals, especially bears, deer, pigs, and squirrels, depend on regular supplies 

If trees are not 

producing abundant 

acorns, then squirrels, 

chipmunks, and deer 

aren’t doing as well. 

Predators aren’t going 

to do as well either. 
24 

GENERAL TECHICAL REPORT PSW-GTR-252

Table 1—Criteria suggested by Garibaldi (2009) for indexing cultural keystone species, with responses and 

citations for California black oak 

Category Question Response and citation 

Intensity, type, and multiplicity of use 

Naming and terminology in a language, including use as seasonal or 

phenological indicators, names of 

months or seasons, place names 

Role in narratives, ceremonies, or 

symbolism 

Persistence and memory of use in 

relationship to cultural change 

Level of unique position in culture 

Extent to which it provides opportunities for resource acquisition from 

beyond the territory 

Is the species used intensively (routinely 

and/or in large quantities)? 

Does the species have multiple uses? 

Does the language incorporate names 

and specialized vocabulary relating to 

the species? 

Is it prominently featured in narratives 

or ceremonies, dances, songs, or as a 

major crest, totem, or symbol? 

Is the species ubiquitous in the collective cultural consciousness and 

frequently discussed? 

Would it be hard to replace this species 

with another available native species? 

Is this species used as a trade item for 

other groups? 

Black oak acorns have been intensively 

used in very large quantities (Anderson 1993). 

Black oak has been an important source 

of a wide array of material items 

(Anderson 1993). 

The species name is commonly used in 

place names and names of months and 

seasons (Anderson 1993, Pavlik et al. 

1995). 

Black oaks are prominent in stories, 

personal names, songs, festivals, and 

ceremonies of native Californians, 

including first acorn feast ceremonies 

(Anderson 1993, Kroeber 1925, Ortiz 

2008, Pavlik et al. 1995). 

Black oak is one of the most important 

species utilized by tribes in the area 

(Anderson 1993). 

Gatherers noted that black oak has a 

large nut, it is easier to grind, and it 

has more oil; it was highly valued 

and grows at elevations above other 

oaks in the Sierra Nevada. Therefore, 

substitution with other species is not 

desirable, although it may be possible 

through trade and gathering at different locations (Anderson 1993, Ortiz 

2008). 

Trade networks involving black oak 

were extensive in California and 

beyond; for example, Miwok traded 

black oak acorns for piñon nuts from 

the Great Basin (Anderson 1993). 
25 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Table 2—Criteria proposed for “culturally-defined” keystone species by 

Cristancho and Vining (2004), with responses and citations for California 

black oak 

Question Black oak example 

The story of the species’ origin is tied to 

the myths, the ancestors, or the origin 

of the culture. 

The species is central to the transmission 

of cultural knowledge. 

The species is indispensable in the major 

rituals on which the community’s stability depends. 

The species is either related to or used in 

activities intended to supply the basic 

needs of the community, such as getting food, constructing shelters, curing 

illnesses, etc. 

The species has significant spiritual or 

religious value for the culture in which 

it is embedded. 

The species exists physically within the 

territory that the cultural group inhabits 

or to which it has access. 

The cultural group refers to the species 

as one of the most important species. 

Myths among Mono (Gifford 1923:366) and 

Karuk (Harrington 1932) personify black oaks 

as young women. 

Harvest and tending practices are a central, intergenerational cultural activity (Ortiz 2008). 

Fall festival, bear dance, and other rituals 

(Ortiz 2008). 

As depicted in figure 6 and supporting 

references (Anderson 1993, Goode 1992). 

Provides habitat for spiritually important animals, such as owls and bears. 

The range of black oak (fig. 1) overlaps 

with cultural groups who prefer it as a food 

(fig. 13). 

Although groups might have concerns 

about ranking species in terms of importance, 

they regard black oak as a notably important 

species. 
26 

GENERAL TECHICAL REPORT PSW-GTR-252

Ron Goode 

Figure 14—Elements of black oak food web. The quote at the bottom by Rebecca Bliege Bird 

reads, “People affect all other species, to manage is a religious philosophy of dominion. An alternative is the Aborigine way of thinking—everything has an inter-relationship, a web of positive 

and negative species.” 

of acorns to maintain their normal densities. Many carnivores, including bears, are 

particularly important symbolic animals for tribes. 

Relationships Between Fire and Black Oak 

Goldilocks Principle 

McDonald (1969: 15) described fire as “black oak’s worst enemy,” reflecting a 

concern that individual black oak stems are highly vulnerable to fire damage, 

even from low-intensity prescribed fires. However, he and other researchers later 

acknowledged that in the long-term, fire can also be a “blessing” for black oak 

by providing favorable conditions against competitors (McDonald and Tappeiner 

2002). Furthermore, traditional practitioners and researchers both have shared the 

perspective that large oaks frequently tolerated fire well and that frequent, lowintensity fires maintained the more open structures and low fuel levels that helped 

avoid top-kill of oaks (Jack 1916, Skinner et al. 2006). For example, in reviewing 

effects of winter prescribed burning around Shasta Lake, Skinner (1995) observed 

complete top-kill of black oaks less than 5 cm dbh, mixed impacts to trees between 

5 and 20 cm dbh, and no cambial damage to black oaks larger than 20 cm dbh that 

did not have any existing open fire scars. Similarly, Kauffman and Martin (1990) 
27 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

found high rates of top-kill from prescribed burning of small black oaks (<3 m tall), 

with no top-kill to taller trees. Furthermore, Douglas-fir, ponderosa pine, white 

fir, and incense cedar (Calocedrus decurrens) tend to establish underneath black 

oaks and eventually overtop them, which reduces the amount of light and other 

resources available to the oaks and which, over time, may reduce their resilience to 

fire and other disturbances (Cocking et al. 2012b, McDonald 1969). Frequent fires 

provide shade-intolerant black oak with openings and resources needed to thrive 

within forest landscapes dominated by conifers. That black oak both needs fire 

and is threatened by it suggests a “Goldilocks” effect in which fuels and structural 

conditions need to fall within particular margins for fire to favor development and 

persistence of large oak trees. This complex relationship with fire helps explain why 

the pros and cons of fire in forests with black oak have been debated for at least a 

century. Klamath River Jack wrote, “Indian burn every year just same, so keep all 

ground clean, no bark, no dead leaf, no old wood on ground, no old wood on brush, 

so no bug can stay to eat leaf and no worm can stay to eat berry and acorn. Not 

much on ground to make hot fire so never hurt big trees, where fire burn” (Jack 

1916). In response, forest ranger Jim Casey said, “If that place had been left alone 

there would be big timber all over it and you could ride anywhere and there would 

be lots of grass on the ground for deer and cattle....And I have seen the best kind of 

acorn crops in places that never felt a fire” (Jack 1916). 

Cultural Burns 

Cultural burning is the traditional use of fire to stimulate desired conditions for 

specific cultural purposes (Goode 2014, Lake 2013). Intended outcomes of cultural 

burning may differ according to their diverse objectives, which also include making 

lands more favorable for particular activities (fig. 15). Accordingly, the historical effects of cultural burning would have been most pronounced surrounding 

frequently used areas such as villages, acorn processing camps, and trade or travel 

routes (Bettinger et al. 1997, Morgan 2008). The early appeal for cultural burning 

in oak forests by Klamath River Jack, cited above, explained that regular fires had 

been used to reduce pests in acorns and to sustain deer and berry production. 

Goode (2014) explained how burning practices have been taught among the 

Mono through a system of lifelong learning. His description emphasizes that 

frequent understory burning was not practiced everywhere on the landscape; 

instead, it would have been concentrated along trails, around camp sites, and at 

meadows. Some areas that had been overgrown would be targeted with three burns 

in a decade. Subsequent burns were not necessarily so frequent, with two to three 

maintenance burns spread across the following two decades. However, actively 

That black oak 

both needs fire 

and is threatened 

by it suggests a 

“Goldilocks” effect 

in which fuels and 

structural conditions 

need to fall within 

particular margins 

for fire to favor 

development and 

persistence of large 

oak trees. 
28 

GENERAL TECHICAL REPORT PSW-GTR-252

A 

B 

Danny Manning 

C 

Danny Manning Danny Manning 

Figure 15—(A and B) During and (C) after a cultural burn 

under black oaks to prepare the grounds at a Native AmericanMission site for the traditional bear dance. 

harvested oak groves have been burned more regularly to consume litter and 

remnant acorns under the trees. Consistent with these descriptions, Anderson (1993) 

noted that indigenous fires were set every year in the Sierra Nevada wherever the 

land “needed it,” as indicated by abundant growth of brush, with the result that 

tended areas were customarily burned every 2 to 10 years. To curtail pests, tribal 

burning in black oak forests managed for acorn production may have been conducted as frequently as surface fuels and weather conditions allowed for a contiguous understory burn. 
29 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Studies of Past Fire Frequency in Forests with Black Oak 

Research to infer historical fire patterns based upon fire scars confirms that forests 

with black oak experienced frequent fires, and such studies have concluded that 

native peoples likely increased that frequency in many areas. According to a review 

by Van de Water and Safford (2011), median fire return intervals for dry mixedconifer and yellow pine (ponderosa pine and/or Jeffrey pine) forests that commonly 

have a California black oak component were 9 and 7 years, respectively. Historical 

fire return intervals can vary considerably, with a reported range of 2 to 23 years in 

the yellow pine/black oak forest type (Skinner and Chang 1996). Fires were more 

frequent in the southern and central Sierra Nevada and throughout the black oak 

range in drier sites associated with south aspects, ridges, and lower elevations (van 

Wagtendonk and Fites-Kaufman 2006). At a more northerly site in the Klamath 

Mountains, black oak appeared more common on mesic, north-facing slopes at 

lower elevations with a median fire return interval of about 11 years (Taylor and 

Skinner 2003). 

Researchers generally agree that Native Americans used fire and other practices to promote desired forest conditions, although there has been considerable 

debate over the extent of such modifications (Lake 2013). In many parts of North 

America, oaks and other trees producing fruits were most accessible in landscapes 

maintained by Native Americans using fire and other tools to cultivate these 

resources (Abrams and Nowacki 2008). Within California, Anderson and Moratto 

(1996) reviewed information about the population densities and likely ecological 

impacts of Native Americans in the proto-historical period, noting that frequent 

burning was an insurance policy against loss of important resources by promoting 

the black oak–ponderosa pine forest type and other highly valued habitats. Anderson and Carpenter (1991) reviewed pollen and other evidence in suggesting that 

Yosemite Valley was a specific example of a landscape modified by Native Americans to favor black oaks and other food resources. 

Researchers have identified burning by Native Americans as a probable contributor to the high frequency of fires in many areas prior to settlement or active 

fire suppression by Euro-Americans. For example, at a mixed-conifer forest at 

Black Mountain in the San Jacinto Mountains where black oak was a co-dominant 

tree, Everett (2008) reported a median fire return interval of 2 years between 1700 

and 1900, and 5 years in the 20th century; he suggested that burning by Native 

American groups, including the Cahuilla, Serrano, and Kamia, likely contributed to 

the frequent fires recorded in the pre-settlement period. Most indigenous landscape 

burning ended by the early 20th century, coinciding with the beginning of the fire 

suppression era, although in some cases it was curtailed or ended even earlier 
30 

GENERAL TECHICAL REPORT PSW-GTR-252

because of settlement impacts on Native American populations (Fry and Stephens 

2006, Swetnam and Baisan 2003). Alterations to traditional burning practices 

associated with Native American population declines as early as the late 18th 

century may have contributed to a reported shift from frequent small fires to larger 

fires, owing to increased continuity of heavier fuels across landscapes (Taylor and 

Beaty 2005). Stephens (1997) reported a median fire return interval of 7 to 8 years 

for a site in the Sierra Nevada foothills in El Dorado County from 1850 to 1952; 

he speculated that early range managers accounted for the frequent fires, as Native 

Americans had already been displaced by the start of that record. At a more northerly site with black oak near Whiskeytown Lake, Fry and Stephens (2006) suggested that Native American ignitions likely contributed to the frequent fire regime 

prior to 1850; however, fire frequency after 1850 declined dramatically with EuroAmerican settlements that greatly affected the local Wintu population. However, 

at a nearby site west of Hayfork, California, Taylor and Skinner (2003) found that 

active fire suppression and declines in fire frequency did not become evident until 

after 1920, and they concluded that the longer active fire regime likely reflected 

continued fire use by Native Americans to promote acorns and other desired values. 

Trends 

Declines of Black Oak 

Competing ecological trends over the past century have caused black oak to 

increase in some areas while decreasing in others, as reported in comparisons to 

historical vegetation surveys from the 1930s (Thorne et al. 2007, 2008). Those 

researchers reported that, between 1936 and 1996, the area where black oak 

was present as a subdominant under conifers declined by 25 percent, but that it 

expanded by a similar amount in areas classified as dominated by montane hardwoods. They attributed the increase in oak-dominated area to removal of overstory 

conifers by logging and wildfires. Historical logging often selectively removed 

conifer species, especially large pines and Douglas-fir. For example, Stephens 

(2000) reported that 25 percent of black oaks were cut in four “average” mixed 

conifer stands, which was a lower percentage than white fir (33 percent), sugar 

pine (88 percent), ponderosa pine (58 percent), and Douglas-fir (100 percent), and a 

higher cut rate than incense cedar (13 percent), which had low value. This selective 

timber cutting may have caused black oak to become more dominant; however, 

oaks were also cut to fuel logging engines in the late 19th and early 20th centuries 

(Merriam 2013). Furthermore, in the mid-20th century black oaks were systematically killed to reduce competition and improve growth of conifers (Pavlik et al. 

1995, Rossi 1980). Fire exclusion has helped shade-tolerant conifers invade and 
31 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

outcompete black oak, reducing the vigor of remnant mature oaks (Cocking et al. 

2012b, Skinner et al. 2006) and making old-growth, open ponderosa pine-California 

black oak woodlands rare (Taylor 2010). Figure 16 depicts a historical example of 

a logged ponderosa pine site that had large black oak with a full crown. Increasing the density of shade-tolerant conifers promotes more severe fires that top-kill 

mature oaks. Though oaks usually survive as young sprouts, the large, mature 

trees are lost at least for decades (Cocking et al. 2012, 2014). Therefore, a common 

trend is a decline in the abundance of large, productive black oak trees, especially 

those embedded in a forest of conifers (fig. 17). Within Yosemite National Park, 

black oaks larger than 31 cm dbh were twice as dense in plots from the 1990s than 

they had been in Wieslander plots from the 1930s, although that difference was not 

statistically significant, and there were few very large trees (>61 cm dbh) in either 

period (Lutz et al. 2009). 

Quantitative data on historical black oak composition in mixed-conifer forests 

are limited and often problematic, as hardwoods were typically ignored in early 20th 

century timber inventories (Collins et al. 2011). The General Land Office (GLO) 

surveys in the Sierra Nevada showed that black oaks represented between 23 and 28 

percent of total stems per acre at that time of those surveys in the later 19th century 

(Fites-Kaufman et al. 2007). However, researchers have cautioned that certain tree 

species might be overrepresented in those surveys if they were perceived to better 

A common trend 

is a decline in the 

abundance of large, 

productive black oak 

trees, especially those 

embedded in a forest 

of conifers. 

Figure 16—Black oak within second-growth ponderosa pine forest in Antelope Creek, one-half 

mile east of Mount Grossman in Amador County, California. Photo taken June 1, 1934 by Albert 

Wieslander, courtesy of the Marian Koshland Bioscience and Natural Resources Library, University 

of California–Berkeley, http://www.lib.berkeley.edu/BIOS/vtm/. 
32 

GENERAL TECHICAL REPORT PSW-GTR-252

Jonathan Long 

Alteration of fire 

regimes has been 

associated with shifts 

to closed canopy 

forests that have less 

deciduous oak. Beyond 

the impacts to oaks 

themselves, these 

shifts have resulted 

in landscapes that 

are more sensitive to 

fire, have less diverse 

and abundant plant 

understories, and 

contribute less to diets 

of humans and wildlife. 

Figure 17—Shade-tolerant conifers encroaching upon a mature black oak at Blodgett Forest near 

Georgetown, California. 

serve as long-term monuments. An 1899 survey recorded black oak as having a 

basal area of 2.4 m2/ha and tree density of 13.3 trees/ha within “average” mixed 

conifer stands, but it did not report any black oak in “large” mixed conifer stands 

that had much higher densities of shade-tolerant trees (Stephens 2000). Another 

source of historical stand data and photographs are the “methods of cutting” plots 

available on various national forests; unfortunately, these reports offer only partial 

information regarding oaks (see box 2). 

In many areas, alteration of fire regimes has been associated with shifts to 

closed canopy forests that have less deciduous oak (Agee 1993; Cocking et al. 2014, 

2012b; Taylor 2010; Vankat and Major 1978). Beyond the impacts to oaks themselves, these shifts have resulted in landscapes that are more sensitive to fire, have 

less diverse and abundant plant understories, and contribute less to diets of humans 

and wildlife (Abrams and Nowacki 2008). Increased stand density and reductions in 

understory vegetation have reduced the fine fuels that carried lighter, more frequent 

surface fires (Engber et al. 2011). Research from the western part of Yosemite 

National Park has found that cessation of fire within the past century has resulted in 

much higher overall tree density and canopy cover, and shifted species dominance 

from pine to fir (Collins et al. 2011). In particular, that study remeasured historical belt transects and determined that mean canopy cover (only conifer trees were 
33 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Box 2 

A 1911 Plot with Large Black Oaks on the Sierra 

National Forest 

One historical “methods-of-cutting” plot on the Sierra National Forest, located 

on a gently sloping ridge at 1370 m in elevation in the Chowchilla Mountains, 

was mapped and described in a report by Dunning (1925). The report did 

not provide size or species information for individual oaks, but the author 

noted the “large size attained by the conifers and black oak,” and black oaks 

are evident in photo points (fig. 18). All conifers greater than 4 cm were 

recorded; it is not clear if the same threshold or a larger one was applied to 

oaks. Twenty-five oaks were mapped in the 2.4 ha plot (10.4 trees/ha), compared with 112 conifer trees/ha with dbh greater than 10 cm. The map shows 

that 48 percent of the oaks were widely separated (by more than 9 m), 24 

percent were pairs spaced between 2 m and 6 m apart, and the remaining five 

oaks were in a semi-circular clump with a 4 m radius. The stand had nearly 

16-percent cover of shrubs, dominated by mountain misery (also known as 

bearclover, Chamaebatia foliolosa) and dense pinemat (Ceanothus diversifolius), along with Mariposa manzanita (Arctostaphylos viscida ssp. mariposa) 

and whitethorn (Ceanothus cordulatus). The report also mentioned presence 

of grasses and herbs, including strawberry (Fragaria spp.), lupines (Lupinus 

spp.), and mariposa lily (Calochortus spp.). The plot was about 800 m from the 

Hogan Ranch, where a famous Maidu woman, Lucy Hite, passed away in the 

care of her niece in 1927; this proximity suggests that native tending and burning practices might have been practiced through the time when the plot was 

first sampled. Meyer et al. (2013) were not able to relocate and resample the 

plot, but they reported that recent Forest Inventory and Analysis plots in yellow pine forest in the area had nearly five times more trees, similar amounts 

of conifer basal area, and twice the cover of shrubs. 
34 

GENERAL TECHICAL REPORT PSW-GTR-252

Reducing canopy cover 

below a threshold value 

of 40 percent appears 

to be important to favor 

shade-intolerant yellow 

pines and California 

black oak over more 

shade-tolerant 

conifers. 

Figure 18—Photo point 1976A in a “methods-of-cutting” plot established on the Bass Lake Ranger 

District, Sierra National Forest, taken in 1911, 5 years after an initial timber harvest as reported by 

Dunning (1925). The plot was dominated by ponderosa pine, but it included large black oaks such as 

the one in the upper right corner. 

measured) in 1911 was typically below 30 percent, compared with modern stands 

with no fire that have mean canopy covers of more than 50 percent. Researchers 

have found similar results for the Stanislaus National Forest (Collins et al. 2015) and 

the Sequoia National Forest (Stephens et al. 2015). The authors also emphasized that 

historical stands were heterogeneous and that such variability should be promoted 

in restoration treatments. Reducing canopy cover below a threshold value of 40 

percent appears to be important to favor shade-intolerant yellow pines and California black oak over more shade-tolerant conifers (Bigelow et al. 2011). 

Acorn Production on Public Lands 

Gatherers attending the July 2013 workshop indicated that there used to be many 

more acorns, and because they find relatively few productive black oaks on public 

lands today, they mostly gather on private lands. The belief that acorn production 

has declined and is less reliable than it was in the early 20th century has been documented in interviews with elders from many different tribes (Anderson 2009). This 
35 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

perception of inadequate supply likely leads acorn gatherers to be reticent about 

sharing the locations of their preferred harvesting sites. National forests and other 

public lands are particularly important in providing gathering opportunities for 

Native Americans in California because most tribes do not control their traditional 

territories, and access to private lands is often limited. 

Regeneration Concerns 

Regeneration of black oak may be an important issue in some areas; however, 

McDonald and Tappeiner (2002) noted that environmental conditions typically pose 

more challenges to growing large trees rather than establishing young trees. Managers on the Sierra National Forest and tribal members participating in the July 2013 

workshop noted that young oaks remain abundant in the southern Sierra Nevada, 

especially following burns. Wildfires like the Rim Fire have stimulated extensive 

resprouting of oak trees (fig. 19). The extent to which young oaks are growing from 

acorns, seedling-sprouts, or root-crown sprouts is not consistently reported in studies, although it has important ramifications for growth rates because stems growing 

from larger root systems develop more quickly (McDonald and Tappeiner 2002). 

Reproduction from acorns might be particularly important for colonizing areas 

where oaks were completely killed by unusually severe fire, and for facilitating 

National forests and 

other public lands are 

particularly important 

in providing gathering 

opportunities for Native 

Americans in California 

because most tribes 

do not control their 

traditional territories, 

and access to private 

lands is often limited. 

Angela White 

Figure 19—Black oak resprouting shortly after the Rim Fire in summer 2013 on the 

Stanislaus National Forest. 
36 

GENERAL TECHICAL REPORT PSW-GTR-252

species movement with changes in climate. Kauffman and Martin (1990) suggested 

that reproduction from acorns was important for colonizing areas after fires, and 

they reported observing seedlings that were 50 m away from the nearest potential 

parent tree following prescribed burns. 

Although deer browse young black oaks (fig. 20), in many areas browsing may 

thin the oaks and promote the surviving stems to replace the large trees. However, 

studies in some areas have reported a scarcity of recruitment and a population 

skewed toward older trees, including Cuyamaca Rancho State Park in San Diego 

County (Bowyer and Bleich 1980), parts of Sequoia-Kings Canyon National Park 

(Parsons and DeBenedetti 1979), and Yosemite National Park (Kuhn and Johnson 

2008). Researchers have posited several hypotheses for the lack of regeneration, 

including reduced fire, reduced tending by Native Americans, changes in hydrology 

(particularly in the Yosemite Valley black oak grove), and increased deer browsing. 

Many of these possible causes are interrelated, because Native American practices 

would have promoted acorns through fire while keeping deer populations in check Jonathan Long 

Figure 20—Example of browsing by deer on young oaks at the 

Crane Valley site on the Sierra National Forest, July 2013. 
37 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

through hunting. In a recent report on California hardwoods, Merriam (2013) noted 

that in fire-excluded stands, there were mostly mature black oaks with few very 

young trees and almost no saplings. Ripple and Beschta (2008) found support for 

the deer browse hypothesis in reporting greater recruitment in stands that were 

not accessible by deer since the early 1900s because of physical barriers. They 

attributed this to the reduction in mountain lions in the early 20th century, which 

stimulated a trophic cascade of increased deer populations and, in turn, reduced 

numbers of black oak seedlings. They cautioned that prescribed burning in areas 

where ungulate herbivory is high could accelerate decline of black oaks. In discussing the competing explanations for declining recruitment, Kuhn and Johnson (2008) 

agreed that deer browsing seemed the best explanation for trends in the Yosemite 

Valley, although they noted that other factors might also be involved. They also 

pointed out numerous ways in which ecological dynamics in the Yosemite Valley 

may be distinctive as the result of historical management first by Native Americans, 

then by Euro-American settlers, and most recently by the National Park Service. On 

the other hand, concerns over oak regeneration have also been widespread in the 

southeastern United States and have spurred both prescribed burning and attention 

to burgeoning deer populations (Arthur et al. 2012). 

Concerns Regarding Future Climate 

Drought, wildfire, and other stressors— 

Effects of a changing climate, including increased episodes of drought and severe 

wildfire, are expected to favor black oak at the expense of less drought-tolerant, 

non-resprouting conifers (Kershner 2014). However, drought, damage from fire, 

and browsing of resprouts following fire may also impair black oak recruitment 

(Merriam 2013). Furthermore, an increase in severe fires will jeopardize large, 

mature oaks in favor of young sprouts. Large and old black oaks at the limits of the 

range for the species may also die owing to projected increases in moisture stress 

(Lutz et al. 2010). Mature black oaks may be particularly vulnerable to drought 

because they tend to be dominant primarily in sites that are already too harsh for 

competing conifers (Holmes et al. 2008, Vankat and Major 1978). 

These stressors are also expected to interact with diseases, pests, pollutants, 

and other stressors that are affecting black oak in various regions. Sudden oak death 

pathogen (Phytophthora ramorum), which is affecting black oak in coastal regions 

(Sturrock et al. 2011); the gold-spotted oak borer (Agrilus auroguttatus), a pest from 

southern Arizona and Mexico that has killed black oaks in San Diego County and 

could spread widely across the range of black oak (Venette et al., in press); and 

nitrogen deposition and ozone pollution, especially in southern California (Handley 
38 

GENERAL TECHICAL REPORT PSW-GTR-252

Black oaks may 

continue the recent 

trend of increasing 

in some areas, 

particularly in 

woodland types and 

severely burned areas, 

while declining as a 

mature tree within 

mixed-conifer forests. 

The synergistic effects 

of these stressors are 

likely to cause declines 

in acorn production to 

the detriment of Native 

American harvesters 

as well as wildlife. 

and Grulke 2008). As a consequence of these stressors, black oaks may continue 

the recent trend of increasing in some areas, particularly in woodland types and 

severely burned areas, while declining as a mature tree within mixed-conifer forests. The synergistic effects of these stressors are likely to cause declines in acorn 

production to the detriment of Native American harvesters (Ortiz 2008, Voggesser 

et al. 2013), as well as wildlife. 

For many California plant species, including valley oak and blue oak, future 

climates may cause range contractions as well as pressure to move upslope or northward, especially within the California-Oregon border region (Kueppers et al. 2005, 

Loarie et al. 2008). Declines in black oak are expected at more drought-prone, lowelevation, and low-latitude sites, including the Tehachapi Mountains of the southern 

Sierra Nevada (Dingman et al. 2013). Monleon and Lintz (2015) demonstrated that 

black oak seedlings are found at colder temperatures and higher elevations, but 

not higher latitudes, than older, larger black oak trees, which also suggests that its 

distribution may shift upslope with time. Black oaks may be more restricted in their 

ability to move long distances in response to climate change than many other species because they depend on animals to disperse their seeds (Devine et al. 2012). 

Changes in phenology— 

Increasing temperatures and other changes to seasonal weather patterns could influence black oak acorn production, gathering, and tending. McDonald (1990) reported 

that “sound acorns begin dropping from late September to early November and 

cease by November 15 at lower elevation. At higher elevations almost all acorns 

have fallen by early December” (McDonald (1990: 665). However, at the North Fork 

meeting, Ron Goode noted that rains have become less predictable during the fall 

season and that the prime window for gathering in the southern Sierra Nevada has 

shifted toward the two weeks at the end of November. Less predictable and reduced 

precipitation may constrain windows to apply fire. 

Desired Conditions 

Desired conditions are descriptions of goals to be achieved at some time in the 

future. They are most helpful for forest planning when they can be considered 

across scales from individual trees out to the forest landscape (fig. 21). Because 

restoration of black oak ultimately depends on understanding larger social, cultural, 

and ecological forces that affect forests (McShea and Healy 2002), restoration 

strategies ultimately need to consider the full socioecological system (Lake and 

Long 2014). Many Native Americans consider it necessary to restore fire as an 

ecological and cultural process (Norgaard 2014). Many ecologists also emphasize 
39 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

 

Figure 21—A conceptual model of indigenous management of California black oak at different scales 

of biological organization, modified from Anderson (2002). 

the importance of forest restoration based upon reestablishing fire as a fundamental 

process and restoring heterogeneity according to topography, and black oak plays 

an important role in that vision (North et al. 2014). 

Desired conditions will vary with the values and goals that people have for 

forests, and they change over time. For example, in the 20th century, many land 

managers focused on forests in the United States as a source of wood products, and 

therefore desired conditions reflected a concern for producing high-quality lumber 

Many Native Americans 

consider it necessary 

to restore fire as 

an ecological and 

cultural process. 

Many ecologists 

also emphasize the 

importance of forest 

restoration based 

upon reestablishing 

fire as a fundamental 

process and restoring 

heterogeneity 

according to 

topography, and black 

oak plays an important 

role in that vision. 
40 

GENERAL TECHICAL REPORT PSW-GTR-252

Native Americans 

emphasize the need for 

black oaks to produce 

large quantities of 

high-quality acorns 

to support abundant 

harvest on a reliable 

basis. 

primarily from conifer trees, although some scientists have proposed strategies 

to promote black oak and other hardwoods as commercial species (Tappeiner and 

McDonald 1980). This report focuses on desired conditions expressed by many 

Native Americans who gather black oak acorns, but it also considers what conditions might be desirable from a perspective of fire resilience, production of lumber, 

and conservation of sensitive wildlife species. By considering how these different 

perspectives might inform forest practices, people may envision how to design 

an overall strategy that would sustain this range of values. McDonald and Huber 

(1995b) have previously laid the foundation for this type of integrated approach, 

although they did not focus on production for tribal use and conservation of oldforest associated species. 

Conditions Associated with Tribal Values 

Native Americans emphasize the need for black oaks to produce large quantities 

of high-quality acorns to support abundant harvest on a reliable basis. Consistent 

production is unrealistic because many factors besides tree condition influence 

masting (Koenig and Knops 2002). However, gatherers can spread risk by gathering 

from various stands at different elevations, and by storing black oak acorns and 

adding acorns from other oaks. Consequently, desired conditions may be described 

at the scale of the tree and even in terms of individual acorn quality. An idealized 

depiction of desired condition at the tree scale (fig. 22) integrates many qualities 

described in further detail below. 

Mature trees— 

The mere presence of oaks is not sufficient to meet tribal needs because black oaks 

take 80 to 100 years before they become mature enough to consistently produce 

large quantities of acorns (McDonald 1969). However, the time to maturity may be 

greatly accelerated for resprouts of top-killed trees, as the well-developed root systems facilitate regrowth above competing plants and the reach of browsing animals. 

Large oak trees are also particularly important for wildlife species including fisher 

and California spotted owl, and they can be an important source of wood. Anderson 

(2005) noted that the Native American practice of tending and frequent burning 

favored older oaks and may have enhanced their longevity. It would be difficult to 

ensure the survival of large trees under a fire regime that includes large amounts 

of high-intensity fire, which could happen to some extent under a managed fire 

regime. However, extensive loss of mature trees in mixed-conifer forests should be 

expected under fire-suppression-based management without intensive reduction of 

fuels and competing conifers that reduce oak vigor and eventually cause oak mortality, and increase potential for severe fire (Cocking et al. 2012b). 
41 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Figure 22—Idealized tree condition desired by tribal acorn gatherers. Illustration by Steve 

Oerding with guidance from the authors. 

High acorn production— 

A black oak tree may remain an abundant producer until it is at least 200 years old 

and may live for half a millennium (McDonald 1969). However, the oldest trees may 

not necessarily be as productive as the middle-aged ones (Anderson 1993; Garrison 

et al. 2002a, 2002b). McDonald (1969) suggested that a large 150- to 200-year-old 

tree would produce approximately 6,500 acorns annually. In 1989, he communicated to Anderson (1993) that a mature black oak will yield 140 pounds (64 kg) of 

acorns in a good year. A study in southern Placer County of four black oak stands 

with trees between 58 and 356 years of age found that larger diameter trees produced more acorns but the relationship was not strong, in part because 38 percent of 

the sampled trees did not produce acorns, even though they were in the prime age 

to be good producers (Garrison et al. 1998). Moreover, they observed the highest 

acorn production in the youngest stand and the lowest production in the oldest stand 

(Garrison et al. 2002a, 2002b), which does suggest that middle-aged trees might be 

more productive. 
42 

GENERAL TECHICAL REPORT PSW-GTR-252

Gatherers also 

emphasize the 

importance of the 

productivity of black 

oaks for supporting 

the larger ecosystem, 

including the multitude 

of wildlife species that 

utilize acorns. 

Upper values for acorn production and relationships to size might be different in 

tended and burned stands. Tribal practitioners at the 2013 North Fork meeting estimated production at approximately 90 kg (198 lbs) of acorns from typical trees they 

use for gathering, while noting that particularly high-quality trees could produce up 

to twice that amount in a good masting year. Heizer and Elsasser (1980) reported 

yields ranging from 41 to 62 kg (90 to 136 lbs) per tree. These estimates demonstrate that even production is highly variable. To provide a sense of scale, a typical 

traditional Native family would use from 82 to more than 618 kg (180 to more than 

1,360 lbs) of acorns in a year, according to reports summarized by Anderson (1993). 

Gatherers also emphasize the importance of the productivity of black oaks for 

supporting the larger ecosystem, including the multitude of wildlife species that use 

acorns—which are also culturally significant. They note that acorn production is 

a good indicator of tree health, and a mature tree that is not producing a good crop 

may be compromised by the effects of pathogens or fire. However, gatherers do 

recognize the many benefits that oaks provide to humans even when they are not 

producing acorns, including as habitat for wildlife, fuelwood, or lumber. 

High acorn quality— 

An important metric of management practices is whether the desired resources are 

usable (Lake and Long 2014). Useful indicators of acorn quality include the quantity of good acorns per tree or unit area and the ratio of good to bad acorns; these 

metrics are good measures of suitability for gathering. Good quality acorns are not 

infested with weevils or worms (Anderson 2005). A useful visual indicator of acorn 

condition is the color of the “eye,” or the spot on the acorn where it was attached to 

the cap; the eye will usually appear brown or blotchy where an insect has injured 

the acorn, as Frank Lake observed at the North Fork meeting in 2013. Members 

of the North Fork Mono and other tribes comment that exit holes on acorns and 

the weight of acorns in the hand are some of the best indicators of insect damage 

(Anderson 2009). 

It is important to note that scientists have measured acorn production using 

timed, relatively short visual counts of acorns on individual trees (Koenig et al. 

1994), which does not directly evaluate acorn quality. However, measuring sound 

acorns typically involves the arduous task of constructing traps that catch acorns 

while excluding animals. Koenig and Knops (1995) noted that trapping is very 

labor intensive and that birds remove many acorns on the tree before they even 

fall. Research by Garrison et al. (1998) suggested that timed counts of acorns are 

significantly related to acorns counted in traps, with a strong relationship between 
43 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

the visual counts and the total number of sound acorns and an even stronger relationship between visual counts and the weight of sound acorns. Those relationships 

appear strong enough to provide meaningful indicators of black oak condition, 

although they may or may not be sufficient to rigorously evaluate potential effects 

of fire and other tending practices on acorn quality. Where acorn gatherers are 

actively harvesting stands, it may be possible to relate yields of sound acorns to the 

more routine visual counts on the trees.
 

Trees with broad crowns and low branches— 

Trees that are desired for acorn harvest have full crowns (high crown volume). Close
 

proximity of conifer trees tends to degrade the acorn production, crown structure,
 

and vigor of oaks (Harrington and Devine 2006, Holmes et al. 2008). However, as
 

noted by Danny Manning in box 1, tall pine trees in the vicinity of oaks may benefit 

acorn production by providing protection from winds. Gatherers desire low branches 

(fig. 23) that can be reached with knocking sticks to facilitate gathering. At the 2013 

North Fork workshop, Ron Goode pointed out that many trees today have limbs too 

high off the ground to be reachable with a 2.5- to 3.0-m pole. Where the branches 

are too high, the knocking stick may be used on the trunk, taking care to avoid 

Jonathan Long 

Figure 23—Black oak tree with desirable architecture for acorn gatherers, including full crown 

and low branches on Tom Harris ranch, North Fork. 
44 

GENERAL TECHICAL REPORT PSW-GTR-252

 

Neither conifer 

encroachment nor 

relatively intense 

wildfires promote this 

broad-canopied, lowbranched architecture, 

but instead promote 

taller and narrower 

growth forms. 

damaging the bark. Ron Goode and other participants explained that trees with 

low branches were used by children in games of tree tag. Children and young men 

also climbed those low branches to help gather acorns (Anderson 2007, Siegel et al. 

2008). 

Neither conifer encroachment nor relatively intense wildfires promote this 

broad-crowned, low-branched architecture, but instead promote taller and narrower 

growth forms (Cocking et al. 2012b; Crotteau et al. 2015; Harrington and Devine 

2006). This tall tree shape has been common in the Beaver Creek Pinery (fig. 24), 

which has maintained a lightning-dominated frequent fire regime (Taylor 2010). 

The absence of low branches may render the trees less susceptible to tall flames 

from wildfires. This difference in tree architecture might serve as a clue to differences in management history, and it also demonstrates how natural wildfire alone 

would not create conditions found in tended stands. 

Connie Harrington has explained the difference in architecture between fullcrowned trees and tall trees as having a whole “hand” of tree crown rather than just 

“fingernails”—the tops of closely spaced trees.5 She noted that research has shown 

that acorn production is greater when you have the “whole hand” in production 

rather than just the “fingernails,” citing Peter and Harrington (2002). 

Cavities and heart rot fungi— 

Cavities in boles and large limbs are common in both wild and tended oak 

trees. Although cavities are clearly detrimental to producing high-quality wood 

(Tappeiner and McDonald 1980), they are not considered by gatherers to be detrimental to acorn production. Furthermore, they are particularly important for some 

wildlife species, and they may have broader effects on wildlife, including invertebrates, by maintaining pools of water. Although a variety of cavities are used by 

wildlife at different times, trees scorched by fires may produce cavities that are not 

as well insulated—and generally not as useful for wildlife—as those formed from 

heart rot. The principal fungi that cause heart rot in black oak are oak canker rot 

fungi (Inonotus andersoni and Inonotus dryophilus) and sulfur fungus or chickenof-the-woods (Laetiporus gilbertsonii) (McDonald 1969, Swiecki and Bernhardt 

2006). The latter species is a yellow-orange fungus used by various tribes as food 

and medicine (Anderson and Lake 2013). The fruiting bodies (fig. 25) are edible 

when young and after being thoroughly cooked, although caution is warranted 

5 Harrington, C. 2015. Personal communication. Research forester, U.S. Department 

of Agriculture, Forest Service, Pacific Northwest Research Station, 3625 93rd Ave. SW, 

Olympia, WA 98512. 
45 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife Carl Skinner Ted Swiecki 

Figure 24—Black oak in the wildfire-maintained Beaver Creek Pinery in the Ishi Wilderness, 

Tehama County, exhibits full crowns but few low branches. 

Figure 25—Fresh fruiting bodies of the sulfur fungus, also known as chicken-of-the-woods, on 

California black oak.
46 

GENERAL TECHICAL REPORT PSW-GTR-252

We should look at 

whether you can 

survive on the land— 

where is there food for 

survival? 

because some people have allergic reactions after eating these mushrooms (Swiecki 

and Bernhardt 2006). Fruiting bodies on a tree indicate infection, which can hollow 

out the trunk and large branches. Heart-rot-inducing fungi can enter the tree at broken branches and wounds created by fire, especially at the base of the tree (Loarie 

et al. 2008, Plumb 1980). Following a fire, the point of fungal entry may become 

obscured as the tree heals over (Long 1913); this process can result in a tree that has 

well-insulated cavities yet is still productive and resistant to fire. In other cases, the 

scars may continue to enlarge with fires, hollowing out the bole of the tree and leaving it vulnerable to toppling (McDonald 1969). 

Few broken and dead limbs— 

Tribal acorn gatherers noted that trees that are healthy and productive have few broken and dead limbs, which contribute to fuel loads, shade out productive branches, 

and pose hazards to people under oaks. The practice of knocking the trees with 

poles was used along with pruning dead branches with hooked sticks or saws to 

yield firewood and enhance acorn production (Anderson 2009). Conifer encroachment causes dieback that increases the abundance of dead limbs (Harrington and 

Devine 2006). 

Low mistletoe infection— 

Tribal acorn gatherers suggested that heavy infection by oak mistletoe 

(Phoradendron villosum) (fig. 26) is a sign of a declining tree. McDonald (1969) 

reported that mistletoe is often seen in both open-growing black oaks or in the 

crowns of trees growing close together, but that the overall effect on tree growth 

was probably minor. Garrison et al. (2002b) reported that 28 of the 144 trees they 

studied had mistletoe, but they did not report on any relationship between mistletoe 

infestation and acorn production. 

Open ground with low fuel levels— 

To facilitate gathering underneath productive trees, Native Americans prefer to 

have open ground with few logs or branches and low levels of litter. Kauffman and 

Martin (1987) suggested that reducing the duff and litter to less than 18 to 36 Mg/ha 

through a high-consumption burn or several burns of low to moderate consumption 

would help black oak acorns to germinate. 

Stand, Community, and Landscape Scales 

Native Americans may view the health of the landscape in terms of its capacity to 

support people according to their traditional lifeways; as Ron Goode remarked at 

the North Fork meeting, “we should look at whether you can survive on the land— 

where is there food for survival?” However, Native Americans also emphasize the 
47 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Figure 26—Oak tree with heavy oak mistletoe infection at Tom Harris Ranch, North Fork. 

Jonathan Long 

importance of tending to promote the productivity and health of black oaks for 

the non-human elements of the forest ecosystem, including understory grasses, 

shrubs, mushrooms, and the multitude of animals that inhabit oaks and consume 

acorns directly or indirectly. Native Americans prefer sites that offer good access 

to productive black oak trees on gentle topography to facilitate gathering. The 

areas within these desirable gathering groves, and the communities surrounding 

them, should support diverse and abundant understory plants, especially ones with 

cultural uses (Lake and Long 2014), including: 

•	 Foods from understory plants such as woodland strawberry, wild sunflowers (Wyethia spp.), onions (Allium spp.), cluster-lilies (Brodiaea spp.), snake 

lilies (Dichelostemma spp.), mariposa lilies, camas (Camassia spp.), wildrye (Elymus spp.) and other edible native grasses, elderberries (Sambucus 

spp.), gooseberry (Ribes spp.), sourberry (Rhus trilobata), and manzanita 

(Arctostaphylos spp.). 

•	 Fiber and basketry materials, including milkweeds, (Asclepias spp.), soaproot (Chlorogalum spp.), California hazel (Corylus cornuta ssp. californica), 

deergrass (Muhlenbergia rigens), beargrass (Xerophyllum tenax), willows 

(Salix spp.), basket sedge (Carex barbarae), western redbud (Cercis occidentalis), deerbrush, and Pacific dogwood (Cornus nuttallii). 
48 

GENERAL TECHICAL REPORT PSW-GTR-252

Maintaining a variety 

of productive stands 

at different elevations 

across a landscape 

would help to expand 

opportunities for 

gathering across 

space and time. 

•	 Medicinal or ceremonial uses such as wild tobacco (Nicotiana spp.). 

•	 Various fungi, including sulfur fungus (fig. 25) and morels, that yield foods 

and medicines (Anderson and Lake 2013). 

Many of these desirable plants will not necessarily grow well directly underneath oaks, but desired conditions for gathering will intersperse oaks among 

openings that support shrubs and meadows that support plants requiring wetter 

environments. One plant that gatherers do not desire underneath black oaks is 

mountain misery (bearclover), because its intertwined stems make acorn collection 

more difficult (Anderson 2005). However, this plant has important cultural values, 

provides a valued resource for bees and wildlife, has nodules that fix nitrogen 

(Heisey et al. 1980), and provides highly flammable fuels to carry surface fires, so 

it has important roles within the wider landscape (Lake and Long 2014). The fact 

that the desirability of this specific plant varies at a fine scale illustrates the importance of a multiscale view of desired conditions. 

Maintaining a variety of productive stands at different elevations across a 

landscape would help to expand opportunities for gathering across space and time. 

Because Native Americans value the biodiversity of forests with black oak as a 

whole, they do not expect nor desire open forest conditions throughout the landscape. Nevertheless, promoting desired conditions for tribal values has potential 

to affect other desired conditions, including habitat for species of special concern, 

wildfire resilience, and wood production, which all need to be considered as part of 

a holistic restoration strategy. 

Conditions Associated with California Spotted Owls and Fisher 

Promoting large, mature black oaks that produce large quantities of acorns would 

provide habitat and prey to support carnivores of special concern, including the 

California spotted owl and the Pacific fisher (the subspecies of fisher present within 

the range of black oak). For example, the majority of fisher dens in the southern 

Sierra Nevada are located in large black oaks (mean dbh 70.5 cm).6 However, 

fishers and spotted owls have been associated with relatively high levels of conifer 

canopy cover, multi-layered understories, and decadent tree conditions (figs. 27 and 

28). For example, California spotted owls commonly use large black oak trees with 

features such as cavities, broken tops, and greater than 70 percent total canopy for 

nesting and roosting (Keane 2014). Fishers appear to prefer areas with canopy 

6 Purcell, K.L. 2014. Unpublished data for fisher study. On file with: U.S. Department 

of Agriculture, Forest Service, Pacific Southwest Research Station, 24075 Highway 41, 

Coarsegold, CA 93614. 
49 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Figure 27—Idealized tree condition for sensitive wildlife species such as fisher and owl. Illustration by Steve Oerding with guidance 

from the authors. 
50 

GENERAL TECHICAL REPORT PSW-GTR-252

Jonathan Long 

Figure 28—California black oak being overtopped by conifer trees, 

Yosemite National Park. 

cover of at least 50 to 55 percent for resting (Purcell et al. 2012). They also appear 

to prefer riparian areas, canyons, and steep slopes more than ridges (Zielinski 

2014). Although fishers generally use live trees rather than dead snags, dead limbs 

with access to cavities are also important (Zielinski 2014). Dead branches are also 

a particularly valuable and overlooked resource for cavity-nesting birds, and those 

features are more commonly found in large black oaks (Garrison et al. 2002b). 

Older black oaks are particularly important, not only because large trees are 

preferred, but also because the heart rot that forms cavities becomes more prevalent 

in older trees (Tappeiner and McDonald 1980). Indeed, Bolsinger (1980) reported 

a sharp increase in decay indicators (visible rot, hollows, breaks, and fungus) with 

tree diameter, with all trees >73 cm dbh having visible indicators. Consequently, 

it is important not only to preserve decadent trees, but also to recruit and develop 

large black oaks to provide for the distinctive habitat needs of the fisher in the 

future (Zielinski 2014). 
51 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Habitat associations among fishers, oaks, and landscape attributes may differ in 

strength across California. For example, Zielinski et al. (2006) found that average 

hardwood dbh and slope were both important variables in a predictive model of 

suitability for fisher rest sites based upon Forest Inventory and Analysis plot data 

for the southern Sierra Nevada. However, Zielinski et al. (2012) reported that both 

slope and basal area of black oak were not important in a preferred model of fisher 

habitat choice based upon forest inventory plot data from northwestern California. 

These findings suggest possible differences in habitat needs across geographic 

regions, and they may reflect the greater diversity and availability of hardwoods 

in the Klamath Mountains. However, the associations between aspect and other 

attributes to species’ occurrence may have changed as forests have become denser 

and more homogeneous, so relationships in restored forests could be different. Restoration of conditions closer to historical levels of canopy cover, while promoting 

heterogeneity, would provide for many small gaps needed to favor shade-intolerant 

trees such as black oak (Gersonde et al. 2004). 

Impacts of Fires and Tending Practices 

Wildfires 

Severe fires may kill the above ground stems of black oaks (top-kill) or the entire 

tree. Unfortunately, many studies of fire effects simply report mortality in terms 

of whether the stem lives instead of distinguishing between an undamaged tree, a 

damaged stem with resprouting from the crown, a dead stem with basal resprouting, and a completely dead, non-sprouting tree. These distinctions have important 

implications for how quickly the tree can provide important services, such as acorn 

production and a full crown. Regelbrugge and Conard (1993) studied outcomes of 

the August 1987 Stanislaus Complex Fire in the central Sierra Nevada in terms of 

black oak survival. Across a range of burn severities, they found that out of 38 trees, 

11 percent were live at the top with no basal sprouts, 16 percent were live with basal 

sprouts, 66 percent were top-killed but resprouted, and 8 percent were fully killed. 

More recent studies of wildfires (Cocking et al. 2012b, 2014; Safford et al. 2012) 

provide some estimates of both top-kill and complete kill for a number of fires with 

mixed vegetation burn severity (table 3). In an early study from Southern California, 

Plumb (1980) studied 72 trees nearly 5 years after the Vista Fire on the San Bernardino National Forest and found that 13 percent were dead (all less than 46 cm), 

24 percent had been top-killed, and 64 percent still had live crown, although he 

noted that “a few large, old, hollowed-out trees fell over” (Plumb 1980: 211). 

McCreary and Nader (2011) provide guidance on determining the likelihood 

that the trunks of oaks will survive fires, although they do not provide thresholds 
52 

GENERAL TECHICAL REPORT PSW-GTR-252

Table 3—Data on black oak mortality (both top-kill and complete kill) from several fires with variable 

burn severity in California 

Siskiyou Complex Storrie Fire Grass Valley Fire Peterson Fire 

Fire (2008) (2000) (2007) (2008) 

Study reference Cocking et al. 2012 Cocking et al. 2014 Safford et al. 2012 

National forest Klamath Lassen San Bernardino Lassen 

Year of measurements 2010 2010 2010 2010 

Recently treated area? (if No No No Yes No Yes 

yes, then year of treatment) (2006) (2006) 

Number of black oaks 191 227 29 21 12 6 

Percentage of oaks 13 7.5 14 5 42 50 

completely dead 

Percentage of oaks top-killed 58 71 62 10 59 33 

but resprouted 

Percentage of oaks not 28 21 24.1 86 0 17 

top-killed 

Plot size 314 m2 314 m2 Variable Variable 

Number of plots with >1 25 25 7 6 3 2 

black oak 

Percentage of plots with all 0 0 0 0 0 0 

oaks completely dead 

Percentage of plots with all 12 48 57 0 100 50 

top-killed 

Percentage of plots with 88 52 43 0 0 50 

no kill 

Percentage reduction in 65 70 NA NA NA NA 

oak basal area 

Although black oaks 

that are top-killed by 

fire do not disappear 

from a site, the 

ecological services 

that they provide 

may be lost for many 

decades, or even 

longer. 

of burn severity that consistently result in complete kill. According to them, if the 

bark is flattened, then the underlying cambium is likely dead. If this extends around 

the tree, the tree is likely to die. If the bark remains fissured, albeit blackened, there 

will likely be living, greenish-white tissue under the bark. If the stem is alive, then 

the tree can actively shed its leaves in the fall. However, if fire kills the stems, then 

the leaves will remain until they are knocked down by weather. 

Recovery Following Wildfires 

Although black oaks that are top-killed by fire do not disappear from a site, the 

ecological services that they provide may be lost for many decades, or even longer 

if conditions do not facilitate regrowth of mature trees. Researchers working in 

the area of the Storrie Fire in 2000 have observed acorn production on resprouts, 

as well as prolific resprouting in areas that burned a second time in the Chips Fire 
53 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

of 2012 (fig. 29).7 McDonald (1969), who documented that oaks need many decades 

to mature, noted that nearly all trees he had seen were resprouts, which can grow 

much faster than oak seedlings (McCreary and Nader 2011) and competing conifers 

(Skinner et al. 2006). Cocking et al. (2014) reported that resprouted trees reached 8.5 

m tall in 10 years following a wildfire, which is faster than the regrowth observed 

on sprouts in an unburned, productive forest by McDonald and Tappeiner (2002). 

Evidently, where mature oaks are merely top-killed they will reestablish mature 

stems much more rapidly than in areas where acorns must germinate, seedlings 

are planted, or small trees resprout following severe fires. However, Kauffman and 

Martin (1990) reported that reproduction from acorns is common following burns. 

7

 Varner, M. 2015. Personal communication. Assistant professor, Virginia Polytechnic 

Institute and State University, 310 West Campus Drive, Blacksburg, VA 24061. Matt Cocking 

Figure 29—A black oak top-killed by the Storrie Fire in August 

2000 had resprouted and was producing acorns nearly 12 years 

later when it was top-killed again by the Chips Fire in July, 

2012. An abundance of acorn cups are visible on the dead 

branches in this photo. 
54 

GENERAL TECHICAL REPORT PSW-GTR-252

Prescribed Burns
	

Prescribed fires (fig. 30) typically have much less severe effects than wildfires, but 

conditions and fire behavior can be highly variable. Consequently, outcomes are not 

necessarily generalizable outside of the specific conditions and fuel and vegetation 

types included in a particular study. 

Studies at the University of California’s Blodgett Experimental Forest in El 

Dorado County have shown variable responses of black oak to prescribed fire. 

Research in the mid-1980s showed that tree-sized (>3 m tall) black oaks survived fall 

and spring prescribed burns without top-kill (Kauffman and Martin 1990). However, 

some large black oak stems that initially survived the fires had toppled by the second 

year (Kauffman 1986). Subsequent studies confirmed that burning had not only 

killed small stems, but also induced mortality of larger stems after several years 

(Kobziar et al. 2006, Krasnow and Stephens 2012). Another study at Blodgett found 

100 percent top-survival rates for black oak in a unit burned under moderate conditions in October following a recent rain, and more variable results from a higher 

severity burn, also in October, but under drier conditions (Schmidt et al. 2006). 

James Turner 

Figure 30—Members of the Yurok Tribe’s fire crew participated in a training exchange 

(TREX) prescribed fire in a black oak stand near Weitchpec, California. The intent of this burn 

was to promote access to quality shoots of California hazel for basket weaving, reduce hazardous fuels, and restore wildlife habitat. 
55 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Fire Susceptibility of Encroached Oaks 

Oaks underneath overtopping conifers may exhibit greater sensitivity to fire than 

would otherwise be expected because of compromised vigor. Increased fuel loading 

also increases the likelihood that oaks will be top-killed in fires, as shown in a study 

in which mechanical fuel reductions preceding prescribed fire lowered black oak 

mortality as well as risk of damage under modeled wildfires (Stephens and Moghaddas 2005). Although both black oak and pine litter support frequent fire regimes, 

in the absence of fire, accumulating fuels will increase the severity of fires when 

they eventually recur. For many large pine trees, heavy accumulation of litter and 

duff at their base (fig. 31) can induce long-term smoldering that effectively girdles 

and kills the trees (Hood 2010). Oak litter decomposes more rapidly than pine litter, 

so litter accumulations may be more of an issue in stands with higher densities of 

conifers. Sackett et al. (1992) compared litter and soil heating from prescribed fires 

under two black oaks, one that was a large (102 cm dbh) and full-crowned black oak, 

and another that had a small (23 cm dbh), leaning bole and was overtopped by large 

ponderosa pines. The soil under the full-crowned oak was dominated by oak leaf 

litter and experienced less heating during the burn than the ground underneath the 

suppressed oak, which was dominated by ponderosa pine litter. 

Effects of Season of Prescribed Burning 

Because black oaks are deciduous, stands located below snow lines are well suited to 

burning in the winter, when leaves are available as flashy fuels and leafless crowns 

allow sufficient sunlight to dry the forest floor (Knapp et al. 2009). Burning in 

the dormant season when trees do not have their leaves also results in less scorch 

(Knapp et al. 2009). Following budburst, oaks may be more sensitive to heat and the 

reserves needed to sprout may be more depleted at that time of year (Knapp et al. 

2009). Kauffman and Martin (1990) reported greater top-kill of small black oaks (<3 

m tall) in late spring and early fall burns, with the highest survival in late fall burns 

and intermediate survival in the early spring when conditions were moist. Burning 

during the spring may also pose some hazards for wildlife, including nesting birds 

and denning mammals. The primary denning time for fisher is mid-March to midApril, which can coincide with budburst in oaks. Pups in dens may be vulnerable 

to carbon monoxide poisoning, leading to concerns over prescribed burning during 

those weeks in the spring (Long et al. 2014). These natural cycles suggest the need to 

consider potential negative effects when burning during early spring. 
56 

GENERAL TECHICAL REPORT PSW-GTR-252

Jonathan Long 

Figure 31—The bases of pine trees may get charred during fires because of the accumulation 

of duff and litter; such heavy accumulations of conifer litter also pose threats to encroached 

black oaks. 

Effects of Cultural Burns 

Cultural burning of black oak (figs. 32 through 34) may generate a variety of 

cultural and ecological benefits at the scale of individual trees as well as the larger 

plant community (table 4). 

Biological diversity— 

Burning encourages understory growth of many grasses and forbs (Anderson 2003, 

Wayman and North 2007), and it can stimulate fruiting of edible fungi, including 

fire-following sac fungi such as morel and cup mushroom (Peziza sp.), as well as 

mushrooms that are found in burned areas and may benefit from reduced duff, such 

as coccora (Amanita calyptroderma) (Anderson 1993, Anderson and Lake 2013). 

Cultural burning is an important means of enhancing pyrodiversity, or a range of 

ecological effects that promote biodiversity (Anderson and Barbour 2003). 

Under indigenous regimes, burns were so frequent and fuel levels so low that 

they did not pose a hazard to the forest, according to elders’ accounts (Anderson 

1993, 2005). For example, burning from the bottom of a slope was reported as a 

common practice (Anderson 1993), although that practice tends to be hazardous 

in forests with extensive areas of high fuel loads and therefore is often disfavored 
57 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife Ron Goode Ron Goode 

Figure 32—Cultural burn of a sourberry patch under a valley oak near Mariposa, California. 

Figure 33—Sourberry with desirable shoots, along with lupine and grasses, grew back following 

the cultural burn under a valley oak near Mariposa, California. 
58 

GENERAL TECHICAL REPORT PSW-GTR-252

Carl Skinner 

Figure 34—Prescribed burn in a young stand of black oak on 

Horse Mountain at Shasta Lake. 

Table 4—Potential cultural and ecological benefits of oak cultural burning practices 

Effects of burning 

practices Cultural benefits Ecological benefits 

Reduce incidence of 

diseases and pests 

(especially filbert weevils and filbert worms) 

Possibly reduce 

incidence of mistletoe 

Stimulate production of 

epicormic branches 

Reduce undergrowth 

and fuels 

Formation of charcoal 

Encourage understory 

plant growth (grasses 

and forbs) and mushrooms 

Encourage nitrogen 

fixing legumes 

Enhanced quality of acorns 

(Anderson 2005: 288) 

May increase acorn production 

(Anderson (1993: 45) 

Straight, long, flexible branches 

used for cultural items, including 

baskets Anderson (2005: 288) 

Facilitate acorn collection 

(Anderson 2005: 288) and improve hunting (Anderson 1993, 

Jack 1916) 

Charcoal used for making fires 

Sources of greens, bulbs, edible 

seeds, mushrooms, medicines, 

and basketry materials (Anderson 

1993, Anderson and Barbour 

2003, Anderson and Lake 2013, 

Norgaard 2014) 

Sources of food, thatching and 

other technologies (Anderson 

2009) 

Presumably, enhanced quality 

would also benefit wildlife 

species that consume acorns 

May promote oak health and 

survival (Anderson 1993: 45) 

Epicormic branches create platforms for wildlife in some tree 

species (Ishii et al. 2004) 

Decrease likelihood of more 

severe fires that reach tree 

crowns and promote establishment of oak seedlings and 

saplings 

Charcoal consumed by wildlife, 

sequesters carbon in soils 

Enhance biological diversity; 

encourage nitrogen-fixing 

species 

Compensates for loss of nitrogen 

during burns 
59 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

under contemporary practices. Kauffman and Martin (1987) also contended that 

frequent surface fires and low fuels were typical of presettlement fires in black oak 

forests. 

The typical season of burning reported by tribal practitioners from the central 

and southern Sierra Nevada was autumn or early winter (October through December), typically after or soon before rains (Anderson 1993). Burning during the 

fall prior to gathering of the new acorn crop was beneficial for clearing the forest 

floor and controlling pest incidence, as well as limiting the density of competing 

conifers. However, Anderson (1993: 39) noted an account of late winter burning by 

Hazel Hutchins, Mono, who said that, “The Mono women set the fires in January, 

February, or March before the leaves sprouted and the mushrooms grew.” These 

seasonal relationships may vary from place to place and with time; at the North 

Fork meeting, Ron Goode and Lois Conner Bohna indicated that some oaks in the 

vicinity of North Fork were burned in the early winter (December and January), 

and that mushrooms were out by March. 

Acorn production— 

Reducing insect pests to improve acorn production was one of the most important 

reasons for regular burning under oaks. Two important insect predators on acorns 

are the filbert worm (Cydia latiferreana) and the filbert weevil (Curculio occidentis) (Swiecki and Bernhardt 2006). These two insects can destroy most of a crop of 

black oak acorns; however, acorns that are only partially damaged may still be able 

to grow (Otvos et al. 2012). Anderson (2005) explained that burning treatments can 

provide a very effective biological control on filbert worms and weevils (fig. 35), 

supporting claims written nearly a century ago by Klamath River Jack (1916). The 

combination of knocking to remove acorns from the trees and burning following the 

harvest is likely to be an effective means of controlling a range of pests and diseases, as research in many nut orchard systems has shown that retention of “mummy” 

nuts in the trees and on the ground contributes to such problems (Eilers and Klein 

2009, Siegel et al. 2008). Because black oaks will drop or abort unhealthy acorns 

earlier than healthy acorns to reduce losses from insect predators (Otvos et al. 2012), 

burns conducted prior to gathering can both reduce insect pests and facilitate collection of acorns in the burned area. 

Beginning in the 1970s, Forest Service managers at Shasta Lake sought to 

mimic the historical burning by local Wintu and Pit River people to improve acorn 

production, with a goal of enhancing deer and elk populations (Skinner 1995). They 

found that by burning the stands in January, there would be a large increase in that 

year’s fall acorn crop; that was in contrast to surrounding unburned areas, which 

experienced a poor crop (Skinner 1995). 

Burns conducted prior 

to gathering can both 

reduce insect pests and 

facilitate collection of 

acorns in the burned 

area. 
60 

GENERAL TECHICAL REPORT PSW-GTR-252

Figure 35—Cycles of filbert worms, filbert weevils, and Native American tending practices to manage these two pests of black oak 

acorns. Modified from Anderson (2007). 
61 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Research is currently underway to evaluate effects of cultural burning on 

acorn production (Codding et al., in press). There are several mechanisms by which 

tending practices can enhance production of high-quality acorns for consumption or 

germination: 

•	 Reducing stress on large, acorn-producing trees, which increases production of acorns in terms of total quantity, period within the year, and production across years. 

•	 Reducing the incidence of pests, leading to higher quality acorns. 

•	 Reducing the incidence of tree- or stand-replacing fires, which in turn stabilizes acorn production. 

•	 Facilitating persistence of black oak across a wider elevation range, potentially expanding seasonal availability (Hunt 2004). 

•	 Extending the life and productivity of individual oaks by reducing competition from other trees and shrubs for water, nutrients, and light and shifting 

those resources into fewer, larger, and more productive trees. 

Masting, or years of exceptionally great production, is believed to have developed as a means of contending with species that prey on acorns, by depleting the 

food for predators in mast failure years and satiating them in good crop years 

(Koenig and Knops 2002). To the extent that treatments increase production and 

provide for more continuous food supplies, there is also potential to increase the 

abundance of insects and rodents that prey on acorns, particularly if acorns are not 

being harvested or burned in a manner that controls such pests. This idea suggests 

that it may be important to gather and burn in conjunction with treatments that 

enhance production to avoid creating pest problems, and also that it is important to 

maintain those elements of food webs that keep acorn predators in check. 

Promoting oak tree health— 

Traditional practitioners have suggested many other benefits of tending and burning 

in addition to increasing production of high-quality acorns and reducing insect pests, 

such as enhancing the diversity of desirable plant communities and the health and 

growth of the oak trees. A preliminary experiment found that several fungi, including species causing root disease and heart rot, might be inhibited through smoke from 

forest fires (Parmeter and Uhrenholdt 1974). Moritz and Odion (2005) suggested that 

frequent fires may curb spread of fungal pathogens such as sudden oak death. Several 

mechanisms for this type of effect have been proposed, including releasing calcium 

that could enhance the tree’s defenses, promoting drier microclimates, and producing 

smoke with fungicidal properties (Calder et al. 2010). However, researchers of sudden 

oak death have shown that its relationship to fire is complex (Rizzo et al. 2005). 
62 

GENERAL TECHICAL REPORT PSW-GTR-252

These new growths 

serve various cultural 

uses, may also form 

platforms that are 

desirable for wildlife, 

and may help to 

fill gaps in the tree 

crown, in particular by 

reforming branches 

closer to the ground. 

Participants at the 2013 North Fork meeting noted that burning could reduce 

incidence of mistletoe, which is a claim that had been supported by the pioneering 

fire ecologist Harold Biswell (Anderson 1993). A study in a foothill community 

of blue oak and interior live oak also reported that fire reduced oak mistletoe 

infestation from “abundant” to “completely absent,” although the incidence was 

not quantified nor evaluated over a longer period (Haggerty 1994). In addition, 

Zimmerman and Laven (1987) found that smoke from burning forest fuels for at 

least an hour may inhibit the viability of seeds of dwarf mistletoe (Arceuthobium 

spp.), which infects ponderosa pine. Harrington and Hawksworth (1990) suggested 

that pine stands with light to moderate dwarf mistletoe infection could be sanitized 

using prescribed understory burning. Mathiasen et al. (2008) explained that this 

effect was induced through scorch that kills infected trees, lower infected branches, 

and brooms, as well as smoke and heat that caused “dehiscence” of mistletoe plants. 

At the 2013 North Fork meeting, Carl Skinner noted that following burning treatments at Shasta Lake, a lot of the oak mistletoe fell to ground, still green, where 

it was eaten by deer. His observations suggested that smoke or heat, rather than 

scorch, was responsible. 

Practitioners also contend that ash from the burning can help the trees grow 

better (Anderson 2005). Lois Conner Bohna suggested at the 2013 North Fork 

meeting that tannic acid from the oak leaves might be harmful to the oak trees, 

according to her grandmother, who recommended regularly burning that litter. 

Researchers in France reported a relationship between accumulations of oak leaves 

in litter near the trunk base and increases in soil acidity in the same area, and they 

noted that the trend was stronger in soils that were poorer in clay (Beniamino et al. 

1991). The acidifying potential of oaks is expected to be less than conifers, but more 

than grasses (Chandler and Chappell 2008), and therefore, isolated acidification 

under oaks may be more evident in oak woodlands than in mixed-conifer forests, 

where acidification is presumably more widespread. 

Production of Epicormic Branches or Root Sprouts 

Practices that induce stress to the tree or increase exposure to light, including thinning and burning, can stimulate sprouting of new branches (epicormic branches) 

from the bole or old stems, or sprouts from the roots (Meier et al. 2012). These new 

growths serve various cultural uses, may also form platforms that are desirable 

for wildlife, and may help to fill gaps in the tree crown, in particular by reforming branches closer to the ground (fig. 36). Although opening the canopy through 

treatments can stimulate acorn production, McDonald and Ritchie (1994) cautioned 

that epicormic branches may not produce acorns. However, Devine and Harrington 
63 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife Lenya Quinn-Davidson 

Figure 36—Epicormic sprouts on a black oak five years after a 

release treatment on private land in Laytonville, California, 

May 8, 2015. 

(2006) observed acorn production on 5-year-old epicormic branches on Oregon 

white oak, so production may just require time for the branches to grow. Treatment 

design should consider this potential response, especially when reclaiming overly 

dense stands. Because these physiological responses are complex, they warrant 

further investigation (Meier et al. 2012). 

Effects of Other Native American Tending Practices 

Pruning and knocking can remove dead and dying branches, mummy nuts, 

galls, and mistletoe; stimulate emergence of new sprouts and branches; open the 

crown; and enhance productivity (Anderson 2005). These effects in turn increase 

crown surface area and production of acorns, while reducing incidence of broken 

branches. Weeding around individual oaks and seeding of desired grasses and forbs 

may help promote desired conditions and community composition (Anderson 2005). 
64 

GENERAL TECHICAL REPORT PSW-GTR-252

Approaches for Reclaiming Degraded Stands 

Black oak groves that have been overgrown with conifers because of fire exclusion 

may have changed so dramatically in terms of fuels and structure that restoring 

their condition may not be feasible simply by reintroducing fire. Researchers in the 

southeastern United States have cautioned that because departures from historical 

conditions in their region have increased constraints on burning and made mature 

trees more vulnerable, use of fire needs heightened scrutiny to avoid damaging 

effects (Arthur et al. 2012). One consequence of these changes is that the strategies 

to reclaim degraded forests may initially require practices that differ from those 

typically employed in actively maintained forests. For example, a nontraditional 

practice may involve burning concentrations of fuels (“jackpot burning”) when the 

fire will not spread easily between concentrations. In terms of burns designed to 

spread through the understory, it may be helpful to conceive of various stages of 

burning: 

•	 Initial entry or preparatory burns are intended to reduce continuity and 

amount of surface fuels, but are not likely to modify forest structure. Such 

burns are likely to be conducted well outside the natural wildfire season 

when moisture or other conditions limit intensity. 

•	 Restorative burns reduce fuels and may kill trees of various sizes to realign 

forest structure and composition closer to a reference condition. 

•	 Maintenance burns kill smaller trees and reduce fuels but are not likely to 

kill mature, healthy trees. 

Spring Burns and Multiple Burns 

Reclamation of degraded stands may require multiple burns before the stand is 

prepared for intensive use and frequent dormant-season burning; that principle is 

consistent with the tradition of cultural burning described above. Furthermore, 

burning outside of the historical fall-winter seasons may help to realign the system 

to the point where in-season burning can occur more safely (Knapp et al. 2009). For 

example, burning in the early spring, when moisture levels are higher, may help to 

reduce heavy accumulations of brush and duff and reduce the potential for large 

tree mortality that might occur in the fall (Kauffman and Martin 1987). However, 

out-of-season burning has been a topic of controversy in some areas, because of 

concerns about impacts to wildlife and plant growth in particular (Knapp et al. 

2009). By the time a second or third burn is planned, fuel conditions may be better 

suited for more traditional fall burning. Fall burns may be appropriate for creating 
65 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

conditions on the landscape that help to restore resources, although it is important 

to engage tribal practitioners in designing, monitoring, and evaluating these treatments to determine whether desired outcomes come to fruition. 

Corrective Fires 

The potential use of managed wildfire or more intense prescribed fire as a forest 

restoration strategy becomes very complex when considering black oaks, not only 

because of their inherent susceptibility to fire but also because compromised oaks 

tend to be more sensitive than overtopping conifers to fire. Cocking et al. (2014) 

suggested that to allow black oaks to regain dominance in stands where conifers 

are well established and mechanical thinning is not practical, severe fires would be 

needed to kill the competing conifers, but would also top-kill the oaks. This idea 

recalls the “corrective burning” used by aborigines in Australia and described by 

Lewis (1994) as a way to reclaim forests that had gone too far from their desired 

condition; those more severe burns would effectively reset the system, and then be 

followed up with maintenance burning at intervals that promote desired conditions. 

However, if severe burns were to occur over a wide area, they would reduce for 

decades the flow of ecological services provided by mature black oaks. Corrective 

fires also have potential to generate considerable smoke, which may make them 

challenging to implement. However, once a frequent understory burning regime 

has been reestablished in black oak-dominated forests, maintenance burns should 

generate much less smoke. Therefore, strategies may need to be tailored to reduce 

the potential for unusually large and severe wildfires and to safeguard particularly 

sensitive, high-value trees through multi-staged interventions (Cocking et al. 

2012b). Wildfires, whether managed or not, offer opportunities to reestablish tending practices with a goal of reestablishing mature oak groves (fig. 37). 

Thinning Conifers to Release Oaks 

Forest managers facing atypically dense stands around black oaks commonly 

prescribe thinning treatments to favor oaks. In a 10-year study of Oregon white 

oak, Devine and Harrington (2013) found that removing all conifers around oaks 

within one tree height (“full-release” treatments) significantly increased oak growth 

and acorn production compared with nontargeted thinning in encroached stands. 

If black oak responds similarly, these relatively aggressive treatments may be 

important in restoring desired conditions, including acorn production and crown 

survival. However, McDonald and Ritchie (1994) suggested thinning lightly around 

black oaks at first and retaining shading by conifers to minimize adverse impacts 

Strategies may need to 

be tailored to reduce 

the potential for 

unusually large and 

severe wildfires and to 

safeguard particularly 

sensitive, high-value 

trees through multistaged interventions. 
66 

GENERAL TECHICAL REPORT PSW-GTR-252

Jonathan Long 

Figure 37—The Eldorado National Forest has proposed an area on the Power Fire (2005) for 

special management to promote the oak resource. This stand of mature black oaks survived with 

stems intact. 

on acorn production that might result from stimulating sprouting. They also suggested that integrating thinning with burning to consume some of the lower branch 

sprouts might help ensure that resources are invested more in acorn production than 

branching, as well as to inhibit build-up of insect pests. Yet, epicormic sprouting 

is a means of restoring low branches and filling in the tree crown that has been 

lost. Consequently, experimenting with a variety of treatments may be valuable in 

evaluating potential tradeoffs. 

Current practices include treatments that increase sunlight around oak trees 

and remove conifers under and near the tree’s crown, as well as thinning oaks to 

promote development of fewer, larger trees. Opening up large patches of sun as part 

of variable density thinning (Knapp et al. 2012) would facilitate reproduction of 

both oaks and pines. Some practitioners have considered treatments that focus on 

the south side of oak trees; however, this approach might increase the propensity of 

trees to lean toward the south, which can decrease their structural integrity. These 

growth patterns are already a common occurrence for heavily encroached oaks (fig. 

38). Harrington (2015) observed that a full release treatment benefits oaks by reducing belowground competition and increasing availability of indirect light. Although 

tree form is not generally as important for oaks as conifers that are being grown 
67 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife Lenya Quinn-Davidson 

Figure 38—Black oaks in a dense Douglas-fir stand in the Klamath Mountains. Oaks are leaning toward the south to capitalize on limited sunlight 

availability. Treatments that focus on the south side of trees can exacerbate 

this problem. 

for lumber, having a well-formed tree rather than a leaning tree will facilitate faster 

growth and better acorn production, and should reduce the tendency for toppling, 

all of which would encourage larger, long-lived, and more productive oaks with 

larger cavities. 

Harrington and Devine (2006) outlined strategies for prioritizing oak stands 

for release treatment based upon tree condition, but additional research on black 

oak would help to evaluate which trees might be too decadent to respond. Devine 

and Harrington (2006) found that Oregon white oak trees with less crown dieback 

(measured as pretreatment live-crown ratio) responded better to release treatments, 

and trees rated as healthy yielded more acorns following treatment. However, more 

recent work by the same authors demonstrates the capacity of long-suppressed 

Having a well-formed 

tree rather than a 

leaning tree will 

facilitate faster growth 

and better acorn 

production, and should 

reduce the tendency 

for toppling, all of 

which would encourage 

larger, long-lived, and 

more productive oaks 

with larger cavities. 
68 

GENERAL TECHICAL REPORT PSW-GTR-252

white oaks to recover and produce acorns after release treatments, so it is important 

that even heavily compromised trees be considered for restoration treatments, 

especially if they are valued for their size, location, cultural significance, or other 

attributes (Devine and Harrington 2013). Because it may take 5 to 7 years to evaluate outcomes, studies need to be moderately long; Devine and Harrington (2013) 

offer a good example of a long-term study, as they evaluated restoration outcomes 

10 years after treatment. 

Thinning Resprouts 

To encourage faster growth of tall and large-stemmed oaks, researchers and managers have proposed thinning resprouts from root crowns (McCreary and Nader 2011, 

McDonald and Vaughn 2007). Even-aged stands of black oak are common, owing 

to resprouts following severe disturbances, including fires as well as pollution 

from mining (Byers et al. 2001, Skinner 1995). Natural mortality of resprouts in 

unmanaged areas appears to reduce the number of stems growing in clumps below 

10, but clumps of stems often consist of relatively skinny trees, some of which 

curve upward to reach gaps in the canopy (McDonald and Vaughn 2007). In large, 

severely burned areas, the need to accelerate development of large trees may be 

deemed particularly important. McDonald and Vaughn (2007) studied effects of 

thinning resprouts and found that trees in small clumps of two to four trees grew 

at about the same rate as a single tree of the same diameter within relatively open 

stands that had been thinned; however, within denser, unthinned areas, stems in 

such small clumps grew slower than single stems. Tappeiner and McDonald (1980) 

noted that thinning before 4 years promotes more sprouting, and that thinning 

between 4 and 8 years may promote forked trees. Combined, these findings suggest 

there may be opportunity to accelerate recovery of mature oaks through thinning 

clumps with more than four stems. However, treatments should be studied over the 

long term to evaluate whether they are efficient in promoting desired outcomes and 

whether they have any undesirable effects. 

Promoting Black Oak When Replanting Conifers After Wildfires 

As large and severe fires have become a more common occurrence in the Sierra 

Nevada and southern Cascade region, managers have proposed replanting conifers 

to promote desired conditions. Because of their capacity to vigorously resprout, 

black oaks are well adapted to become dominant following wildfires. To conserve 

oaks in areas proposed for planting, the Sierra Nevada Forest Plan Amendment 

(USDA FS 2004) imposed a guideline of buffering 6.1 m from the crown edge, 

which was consistent with recommendations by Tappeiner and McDonald (1980). 
69 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Because that distance is less than half the height of mature oaks, it is less than the 

amount recommended for release treatments of mature oaks proposed by Devine 

and Harrington (2013). Consequently, at least in theory, applying such a relatively 

narrow buffer (compared to the height of mature oaks) might limit the opportunity 

for black oaks to be dominant in the future. On the other hand, using a planting 

buffer also potentially constrains the opportunity to use black oaks as nurse trees 

for planted conifers. Strategies that might promote forest heterogeneity could 

include explicitly reserving stands or groups where conifers were not interplanted 

among black oaks, as well as planting conifers using a more variable buffer. Using 

wide buffers could promote diversity and oak values in places where mature oaks 

are scarce, especially at higher elevations, whereas a smaller buffer could help to 

diversify lower-elevation areas where oaks are more dominant. Wide planting buffers may be more important for California black oak than other oaks such as canyon 

live oak, given black oak’s reduced tolerance of shading. 

Protecting Legacy Black Oaks 

Land managers might consider special actions to conserve especially large and 

productive black oaks during prescribed burns or wildfires because of their importance. The term “legacy tree” has been used to distinguish trees that are larger and 

older than the average trees in a landscape (Mazurek and Zielinski 2004). These 

considerations may heighten the need to develop maps or data layers that locate 

important trees or groves, which could be identified in a wildfire decision support 

system. It is important to promote conditions that will allow fire to enter the stands 

while lowering the risk of top-kill, because many black oaks and other large trees 

do not survive present-day fires. One particular strategy may be to remove down 

branches or boles that may be close to the tree, to avoid causing damage to the trees. 

In addition, many researchers and managers have suggested raking litter, especially 

pine needles, away from the boles of large pines and large oaks to promote their 

survival during burns (Abella and Fule 2008, Truex and Zielinski 2013). Hood 

suggested removing both litter and duff using rakes or leaf blowers in areas with 

low summer moisture (Hood 2010). Researchers have found value in these types of 

raking treatments on yellow pines (Hood 2010, Laudenslayer 2008) and sugar pines 

(Nesmith et al. 2010). 

However, caution is warranted, particularly where organic soil layers harbor 

valuable mycorrhizal fungi; this is recommended to avoid damaging the mycelial 

mats, also called shiro (Anderson and Lake 2013, Pilz and Molina 1996). A study 

from the Pacific Northwest showed that shallow raking (i.e., not raking into the 

mineral soil surface) and/or litter and duff replacement can mitigate impacts to 
70 

GENERAL TECHICAL REPORT PSW-GTR-252

Black oak has a 

particularly important 

role in the evolving 

approach toward 

restoration forestry 

in the Sierra Nevada, 

which emphasizes 

the importance of 

considering multiple 

scales, retaining 

hardwoods, restoring 

heterogeneity, 

and designing 

treatments according 

to productivity and 

topography. 

mushroom production, although those findings might not necessarily extend to 

black oaks and associated fungi (Luoma et al. 2006). Furthermore, reducing litter 

may also favor mushroom production (Luoma et al. 2006, Weigand 1998). Consequently, although raking heavy accumulations of pine litter may be important for 

reducing fire severity under encroached black oaks, studies are needed to test the 

effectiveness and possible risks of that treatment. In addition to studies based upon 

monitoring and experiments, local knowledge of mushroom harvesters may be 

particularly helpful in suggesting appropriate practices (Anderson and Lake 2013). 

Other Treatments 

Although the saw and the flame are likely to be key tools for restoring black oak 

ecosystems, other treatments may have important roles in specific contexts. For 

example, an option for reclaiming large black oaks that are being overtopped is 

to girdle the competing conifers (Engber et al. 2011). In areas of heavy browsing, 

treatments to protect sprouts until they exceed the browse height may be necessary. 

Planting of acorns or seedlings may be necessary where populations have been 

extirpated, as may have occurred on Santa Cruz Island, although any such plantings 

would have to consider a range of issues, including appropriate source materials, 

resilience to future climate, and safeguards to avoid spread of pests and pathogens. 

Furthermore, planting of black oak stock with desirable genetic properties, such 

as resistance to sudden oak death, may be an important treatment option for some 

areas in the future (Rizzo et al. 2005). 

A Landscape Strategy for Intensive Black Oak 

Restoration 

Black oak has a particularly important role in the evolving approach toward restoration forestry in the Sierra Nevada, which emphasizes the importance of considering 

multiple scales, retaining hardwoods, restoring heterogeneity, and designing treatments according to productivity and topography (North et al. 2009). A landscape 

approach considers the condition of individual trees and groups of trees, as well as 

the wider landscape in which they are situated. Furthermore, these strategies need 

to consider interactions with the sociocultural systems that value and depend on the 

organisms in the forest (Long et al. 2014). When brought together, these elements 

suggest a three-part approach for managing landscapes with black oak (table 5). 

Considering the Landscape Context for Acorn Gathering 

A landscape strategy to promote tribal use of black oak needs to consider how, 

when, and where to get the system to a point at which desired levels of gathering 
71 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Table 5—Multiscale restoration strategy 

Context Orchard areas General forest Priority resting/denning areas 

Topography 

Access 

Current oak condition 

General treatment approach 

Expected fire frequency 

Expected fire severity 

Resource benefits 

Flatter areas, near meadows 

Good access via roads, which 

also serve as fire barriers 

Remnant large oaks, often 

suppressed and crowded 

Intensive release treatments 

around productive groves, 

followed by frequent burning and tending 

Frequent 

Low 

Acorn production and supporting wildlife values 

Oaks often thin, snakelike owing to overtopping 

and crowding by conifers 

Highly variable, and often 

less intensive, treatments 

intended to promote oak 

longevity, productivity, 

and recruitment 

Moderate 

Moderate 

Fire resilience 

Canyons, riparian area, steep, 

north-facing slopes 

Poor access via roads 

Dying, decadent trees often 

shaded by high conifer canopy 

Areas not to be intensively 

treated, except perhaps in 

select patches to retain their 

long-term value and promote 

diversity 

Less frequent 

High 

Maintain habitat for old-forest 

associated species in near term 

can reliably occur. Maintaining a more reliable supply of the desired services may 

depend on having a number of areas that have experienced fires in recent years and 

can be managed for acorn production. Treatments need to be rotated through various stands across a broad elevation range to maintain availability of the resource 

over time. The distance that gatherers would need to go to gather acorns and other 

key food resources is an important consideration at the landscape scale (table 5). 

The resources have to be plentiful enough to encourage gatherers to seek them out, 

and desirable gathering areas will feature a diverse mix of resources. Both Tappeiner and McDonald (1980) and Potter and Johnston (1980) suggested guidelines 

for oak basal area and canopy cover to support high quantities of oak production 

within relatively pure groups ranging from 0.2 to 4 ha to facilitate treatments to 

favor oak within conifer areas. 

Gathering areas that are close to areas of flat or gently rolling topography and 

roads will be preferred over steeper, less accessible areas. Sites near meadows (fig. 39) 

may not only afford easier access, but also yield a diversity of resources for gatherers; in particular, desirable locations would be near meadows with sandy soils for 

basket sedge and near stands of shrubs. Locations of historically tended “orchards” 

identified by gatherers or in historical records would provide a useful starting point. 

However, long-term treatments should not be limited to these traditional areas, as 

other locations may be important to support those values in the future. Moreover, 

many of the sites where oaks have better withstood encroachment are likely to be 
72 

GENERAL TECHICAL REPORT PSW-GTR-252

Jonathan Long 

Figure 39—Mature black oak with spreading branches surrounded by conifers near a meadow 

at Grey's Mountain treatment area, Sierra National Forest. 
73 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

poorer, drier sites; to promote production it may be worth targeting more productive 

sites. Prioritizing sites on more level ground and close to meadows, consistent with 

criteria supported by tribal gatherers, could help to restore oak in more productive 

areas. 

In the general forest (i.e., not specific gathering sites, box 2), treatments 

designed to enhance oak growth and acorn production may be undertaken without 

necessarily employing the more intensive tending practices used in gathering areas. 

In addition to considering the landscape-scale context for gathering, the age, condition, and growth form of the oaks will inform which individual trees are best suited 

for intensive treatment, and which are best left alone (fig. 40). To restore the values 

provided by black oaks, release thinning or restoration of fire are needed throughout the general forest to redirect conditions away from the trajectory of conifer 

dominance that has been set by the change in fire regime. 

Avoiding Conflicts with Sensitive Species 

When viewed at a tree- or stand-scale, treatments to increase sunlight to black 

oak trees appear to conflict with tactics to conserve California spotted owls and 

fisher that appear to depend on decadent, dense, and complex cover. Furthermore, 

treatments that remove dead branches, inhibit decadence, and clear the understory 

beneath individual oaks could reduce desirable habitat features for those species. 

However, applying a broader, multiscale perspective can help to address these 

concerns in several ways. First, targeting restoration treatments to promote acorn 

gathering groves in areas with gentle topography, good access by roads, and in 

historically more open stands would avoid conflicts with wildlife that depend 

on steeper, more remote, and historically more closed-canopy forests. At a finer 

scale, creating gaps around mature oaks may encourage both acorn production 

and recruitment of young trees that can take off when mature trees fall (Purcell 

et al. 2012). Meanwhile, treatment strategies could retain tree clusters with high 

canopy cover and features favored by wildlife, including broken branches and other 

resting platforms (fig. 41), for their high present habitat value (North et al. 2009). 

Researchers have suggested that maintaining large oak trees along with areas of 

dense canopy cover is a key element of a larger landscape resilience strategy that 

can sustain both spotted owls and fishers (Keane 2014, Zielinski 2014). Promoting 

high variability at multiple scales could retain decadent conditions to support those 

wildlife species in the near term while also maintaining healthy oaks that can provide food resources and habitat for the long term. Because of the chronic pressure 

of forest densification combined with the risk of severe wildfires, active restoration 

treatments will be important for sustaining and reestablishing large oaks. 

Targeting restoration 

treatments to promote 

acorn gathering groves 

in areas with gentle 

topography, good 

access by roads, and 

in historically more 

open stands would 

avoid conflicts with 

wildlife that depend on 

steeper, more remote, 

and historically more 

closed-canopy forests. 
74 

GENERAL TECHICAL REPORT PSW-GTR-252

Jonathan Long 

Figure 40—A black oak in a small riparian area at Crane Valley, Sierra National Forest, lies on a 

steep slope that is probably unsuitable for acorn gathering. 
75 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Figure 41—A large legacy black oak used by a fisher as a resting site. 

Rebecca Green 

A project at the Sagehen Experimental Forest offers a relevant example of 

this conceptual approach. That project, which was refined through a collaborative 

process (Stine and Conway 2012), was proposed for a high-elevation forest type 

where marten, rather than fisher, was a chief wildlife concern. The plan proposed 

to create variable amounts of Dense Cover Areas (DCAs) and Early Seral Openings 

(ESOs) within areas that prioritized different resources, and to remove smaller trees 

around large “legacy trees” to increase their resiliency to fire, drought, pathogens, 

and disease, except in the DCAs. In lower elevation Sierra Nevada landscapes, that 

treatment strategy may be adapted to favor black oaks as a focal legacy tree. 

Conclusions: Opportunities to Increase 

Socioecological Resilience 

Restoration of California black oak lies at the heart of broader discussions of how 

to promote socioecological resilience in the Sierra Nevada (Long et al. 2014). The 

species has suffered long-standing declines because of fire suppression and other 

historical activities, including widespread efforts to systematically replace it with 

conifers favored for timber production. But today, the many values provided by 

black oak have made it a centerpiece for demonstrating the benefits of fire-centric 

restoration of forests in California (Byers et al. 2001, North et al. 2014). 
76 

GENERAL TECHICAL REPORT PSW-GTR-252

Enhancing forests 

with black oaks to 

support tribal values 

offers a win-win 

opportunity to increase 

social and ecological 

resilience in the face 

of stressors such as 

drought, disease, and 

uncharacteristically 

severe wildfires. 

Promoting Ecological Diversity 

Conditions desired by Native Americans would be likely to support a wide variety 

of wildlife species through increased frequency of fire, enhanced acorn production, and retention of large oaks. These conditions do not necessarily conflict with 

sensitive species when considering the system at larger spatial scales and longer 

temporal scales. Indeed, there are wide-ranging benefits to promoting more heterogeneous and more open stands with fewer conifers, especially shade-tolerant 

ones. At least in the near future, thinning and perhaps girdling to release oaks from 

conifer competition are important tools for conserving legacy oaks (Engber et al. 

2011). Fire also can maintain a diversity of vegetation conditions and edges between 

those areas, and managed fire has been recommended as a way to restore and 

sustain features that were abundant in the pre-fire suppression forest (Taylor 2010). 

Moreover, fire and smoke (fig. 42) promote pyrodiversity, or a range of ecological 

effects that promote biodiversity (Anderson and Barbour 2003) and are not replicated through thinning, including inhibition of pests and pathogens, formation of 

cavities, and stimulation of fire-following fungi and plants. 

Promoting Resilience to Climate-Related Stressors 

Enhancing forests with black oaks to support tribal values offers a win-win 

opportunity to increase social and ecological resilience in the face of stressors 

such as drought, disease, and uncharacteristically severe wildfires, which may 

become more common as the climate changes in the future (Long et al. 2014). 

Reestablishing a more frequent fire regime would promote structural changes that 

might render forests more resilient to sudden oak death (Moritz and Odion 2005). 

Targeted reductions in tree densities, creation of openings around black oaks, and 

greater reliance on low-intensity fires may increase resilience to stressors, as well 

as provide dividends in terms of increased water yield and reductions in the extent 

of severe wildfires (Long et al. 2014). Extreme fires such as the Rim and King 

Fires (fig. 43) kill overtopping conifers and may allow resprouting oaks to reclaim 

dominance and extend into higher elevation areas, but not without losing mature 

oaks and the valuable benefits they provide for many decades. Moreover, repeated 

high-severity fires, which have become increasingly common, are likely to inhibit 

development of mature oaks, and drying conditions are likely to inhibit reestablishment at lower elevations and latitudes. Intentional caretaking through judicious use 

of fire and removal of conifers can safeguard and promote legacy trees and their 

associated values in ways that reduce vulnerability of Native Americans, other 

people living in rural areas, and wildlife (Charnley et al. 2007, Devine et al. 2012). 
77 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife Lenya Quinn-Davidson Jonathan Long 

Figure 42—Prescribed burning implemented through a low-cost training program (TREX) on 

45 ha of private lands with black oak, white oak, and ponderosa pine near Hayfork, California, 

October 29, 2013. 

Figure 43—The Rim Fire of 2013 featured many extensive high-severity patches that were 

marked by bare soils, white ash, holes around tree boles, charred organic matter, dead conifer 

trees, and loss of mature black oaks. 
78 

GENERAL TECHICAL REPORT PSW-GTR-252

Promoting Adaptive Management 

Useful indicators for 

evaluating trends in 

black oak conditions 

include the abundance 

of trees with large, 

full canopies, and the 

availability of highquality acorns for 

gathering. 

To promote an integrated approach to managing forests with hardwood, McDonald 

and Huber (1995b) had previously emphasized the need for teamwork involving 

agency researchers, planners, workers, and various landowners. Their framework 

can be expanded to recognize the importance of tribal governments, Native 

Americans, oak enthusiasts, and other stakeholders who value these important 

resources and have a vision for restoring this important component of our forests. 

In particular, continuing research partnerships among scientists, land managers, 

and tribal gatherers (fig. 44) can guide adaptive management efforts in forests with 

black oak. Useful indicators for evaluating trends in black oak conditions include 

the abundance of trees with large, full crowns, and the availability of high-quality 

acorns for gathering. Research has suggested that trees need to be 80 years old to be 

significant producers of acorns, which translates to a dbh of at least 33 to 50 cm and 

crown diameters of at least 8 m, depending upon site productivity (McDonald and 

Tappeiner 2002, Potter and Johnston 1980). Forest management plans can use these 

indicators to track progress in retaining and restoring legacy black oaks and their 

benefits. Research at the scale of large landscape demonstration areas and extensive 

monitoring, as well as smaller experimental plot studies, will help to understand the 

benefits of more intensive caretaking of mature black oaks. 

Jonathan Long 

Figure 44—A group of tribal members, scientists, and managers at the North Fork meeting 

considered how the context of black oak sites should influence treatments. 
79 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Supporting Community Engagement in Restoration 

Reintroducing fire using their own traditional knowledge allows Native peoples to 

reengage as caretakers of their ancestral lands (Eriksen and Hankins 2014). Tribal 

participation in efforts to promote resilience, especially by enhancing the quality 

of black oaks, can strengthen the adaptive capacity of both tribal communities and 

government resource managers (Lynn et al. 2013). Specifically, tribal practitioners 

note that actively tending gathering areas cultivates an important sense of place 

and teaches respect for those places and organisms within the community, especially among tribal youth. Various models exist for cultivating tribal engagement, 

including the use of tribal fire crews with traditional knowledge and training in the 

use of fire for cultural resources, as well as co-management agreements (Eriksen 

and Hankins 2014, Lake and Long 2014). Fire-use training could be developed to 

emphasize how to use fire from both cultural and ecological perspectives. Having 

cultural expertise available would help institutions manage both prescribed fires 

and wildfires to sustain black oaks and other forest values. 

Efforts to engage communities in restoration can extend broadly beyond Native 

American tribes. Examples of black oak restoration projects include the Forest 

Service program of prescribed burning at Shasta Lake inspired by traditional 

knowledge of local tribes (Skinner 1995), several demonstration sites on the Sierra 

National Forest pictured in this report and designed with guidance from Mono 

tribal members, and partnership-based efforts led by the Lomakatsi Restoration 

project, U.S. Fish and Wildlife Service, Natural Resources Conservation Service, 

Klamath Bird Observatory, and several other organizations in southern Oregon 

(Cocking et al. 2012a). The Eldorado National Forest has also proposed special 

management for oaks in parts of the Power Fire (fig. 37). 

The majestic, widely distributed black oak trees serve as an appropriate centerpiece for illustrating the benefits of active, fire-centric restoration in the forests of 

California and southern Oregon. Promoting forest conditions that increase Native 

American access to high-quality acorns are likely to enhance habitat for a broad 

array of wildlife species, as well as yield other societal benefits. Rather than designing strategies to promote these values individually, an integrated approach would 

seize the opportunity to link conservation of cultural diversity, biological diversity, 

and other ecological services (Charnley et al. 2007). Researchers studying forest 

ecosystems recognized for having high biocultural diversity have concluded that 

strategies that reinforce traditional cultural values, including use of wild resources, 

are likely to be more effective in conserving forests because they foster supportive 

attitudes and behaviors (Byers et al. 2001). As more people understand the complex 

The majestic, widely 

distributed black oak 

trees serve as an 

appropriate centerpiece 

for illustrating the 

benefits of active, firecentric restoration in 

the forests of California 

and southern Oregon. 
80 

GENERAL TECHICAL REPORT PSW-GTR-252

web of relationships that bind a variety of species through acorns and oak dwellings, they can better appreciate the value of caretaking to sustain the many benefits 

provided by these trees of life. 

Acknowledgments 

The authors express thanks to the many individuals who have contributed information, photos, or data to this report, including other individuals who participated in 

and supported the initial California black oak meeting from July 23–25, 2013, in 

North Fork, California: 

•	 Jessica Einhorn, Australia National University 

•	 Dyann Eckstein, Chukchansi Rancheria 

•	 Ruby Pomona, North Fork Mono Rancheria 

•	 Lois Conner Bohna and Nancy Tex, North Fork Mono Tribe 

•	 Jared Aldern, Prescott College 

•	 Eric Knapp, Kathryn Purcell, and Craig Thompson, USDA Forest Service, 

Pacific Southwest Research Station 

•	 Dirk Charley, David Martin, Mark Smith, and Burt Stalter, USDA Forest 

Service, Sierra National Forest 

•	 Chathurika Goonawardena, USDA Forest Service volunteer 

•	 Kat Anderson, USDA Natural Resources Conservation Service 

In addition, we greatly appreciate contributions by Danny Manning, Ted 

Swiecki, Bill Zielinski, John Duggett, Matthew Cocking, Steve Oerding, Angela 

White, Brandon Collins, Kevin Krasnow, John Keane, Scott Stephens, Malcolm 

North, and Morgan Varner. We also thank the many individuals who reviewed 

drafts of this report, including Joanna Clines, Matthew Cocking, Phil McDonald, 

and Connie Harrington. 
81 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

List of organisms with common, scientifi c, and western Mono (Nium) names and usesa

Common name Scientific name Mono/Nium name Mono/Nium cultural use 

Birds: 

Acorn woodpeckerBand-tailed pigeonBrown creeperCalifornia spotted owlDowny woodpeckerMountain chickadeeMountain quailNorthern pygmy-owlRed-breasted nuthatchSteller’s jay Valley quail Western scrub jay White-breasted nuthatchWild turkey 

Mammals: 

Black bearDouglas’ squirrel Dusky-footed woodratFisher (Pacific fisher)Hog, wildMule deerWestern gray squirrel Filbert weevil 

Insects: 

Filbert wormGold-spotted oak borerCanker rot fungi 

Melanerpes formicivorusPatagioenas fasciataCerthia americanaStrix occidentalis ssp. lucida Picoides pubescensPoecile gambeliOreortyx pictusGlaucidum gnomaSitta canadensisCyanocitta stelleriCallipepla californicaAphelocoma californicaSitta carolinensis Meleagris gallopavoUrsus americanusTamiasciurus douglasii Neotoma fuscipesPekania pennanti (ssp. pacifica) Sus scrofa Odocoileus hemionusSciurus griseusCurculio occidentisCydia latiferreanaAgrilus auroguttatusInonotus andersoni and Inonotus dryophilus 

Pa-na-ta-daSoo-weCeet-ne-sahtMuhuMuhu-ceeChsg-o-nohChot-nu-key-nehKa-be-ka-be-nahPahubichKah-wahTah-huit Mau-we 

RegaliaFoodSpiritualitySpiritualitySpiritualityResourceMeatFur/hide for pouchesFood, resourceFood, resource 
82 

GENERAL TECHICAL REPORT PSW-GTR-252 Common name Scientific name Mono/Nium name Mono/Nium cultural use Fungi: Charcoal-loving elf cup Geopyxis carbonaria Chicken-of-the-woods (also known Laetiporus gilbertsonii Food as sulfur fungus) Coccora Amanita calyptroderma Toap’ Food Cup mushroom Peziza spp. Nuc-ca Food Morel Morchella sp. Chu-yu/Tsu-yu Food Sudden oak death pathogen Phytophthora ramorum Ancestral black oak (fossil species) Quercus pseudolyrata Trees: Blue oak Quercus douglasii Pa-wi-yap’ Acorns, dyes 

California black oak Quercus kelloggii Wi-yap’ Acorns, wood, walking/

digging sticks, dyes, othersin text 

California buckeye Aesculus californica Wo-yo-nobi Nuts California hazelnut Corylus cornuta ssp. californicaCalifornia nutmeg Torreya californicaCanyon live oak Quercus chrysolepisChase oak Quercus × chasei Giant chinquapin Chrysolepis chrysophyllaCoast live oak Quercus agrifolia Coast live oak, interior variety Quercus agrifolia var. oxyadenia Douglas-fir Pseudotsuga menziesii var. menziesii Sa-qwa’abee Wood Gander oak Quercus × ganderi Incense cedar Calocedrus decurrens Wau-up Wood, food, nuts, pitch, 

medicine, food, drink

Interior live oak Quercus wislizeni Tsigi-nobe Acorns Jeffrey pine Pinus jeffreyi Kusi-wo-qobi Nuts Oracle oak Quercus × morehus Oregon white oak Quercus garryanaPacific madrone Arbutus menziesiiPonderosa pine Pinus ponderosa Woh-koob’ Wood, basketry, nutsRed maple Acer rubrum 
83 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Common name Scientific name Mono/Nium name Mono/Nium cultural use Santa Cruz island oakShreve oakSugar pineTanoak Walnut Western redbud White firYellow pine Dwarf mistletoe

Mistletoes: 

Oak mistletoeBasket sedge 

Shrubs and

herbs: 

Beargrass Big deervetchBush chinquapinCamasCluster-lilies DeerbrushDeergrassElderberry (blue)Elderberry (Mexican)GooseberriesLupinesManzanita (green-leaf)

Quercus parvula var. parvula Quercus parvula var. shrevei Pinus lambertianaNotholithocarpus densiflorusJuglans hindsiiCercis occidentalisAbies concolorPinus jeffreyi or Pinus ponderosaArceuthobium spp.Phoradendron villosumCarex barbarae Xerophyllum tenax Hosackia crassifoliaChrysolepis sempervirensCamassia spp. Brodiaea spp.Ceanothus integerrimusMuhlenbergia rigensSambucus nigra ssp. ceruleaSambucus mexicanaRibes spp.Lupinus spp.Arctostaphylos patula 

Pa-ne-wi-nip’Ta-qwa-qwa-abee Sa-qwa-abeeWoh-koob’ Pa-zo-o-biTe-de-nap’ Tsi-ni-pin Po-no-o-wi/Qo-no-o-wiTsi-ni-pab’ Mo-nop”Hu-bu-he-yab’Hu-bu-he-ya/Ku-nu-geb’Ta-bap’/Ta-ba’ya Pa-na-qo-zabi 

Wood, basketry, nuts, medicineBasketryWood, food, nuts, pitch, medicineMedicine, poisonbindingNutsFoodBasketryBasketry, cordageFood, musical instrumentsFood, musical instrumentsFoodFood, medicine, walking/digging sticks 
84 

GENERAL TECHICAL REPORT PSW-GTR-252 Common name Scientific name Mono/Nium name Mono/Nium cultural use Manzanita (Mariposa) Arctostaphylos viscida ssp. mariposa Aposo-wabi Food, medicine, walking/ digging sticks 

Mariposa lilies Calochortus spp. Food Milkweeds Asclepias spp. Wi-si-bi/We-tsib’ Cordage, medicineMountain misery or bearclover Chamaebatia foliosa Sa-a-tu-wib/ Medicine 

Sat-tu-wib 

Onions Allium spp. Ped-is Food Pacific dogwood Cornus nuttallii Medicine Pinemat Ceanothus diversifoliusSnake lilies Dichelostemma spp. Food Soaproots Chlorogalum spp. So-o-sibe/So-sib’ Brush, medicineSourberry Rhus trilobata Ta-ka-te/ Food, medicine

Kwa-nu-nup’ 

Whitethorn Ceanothus cordulatusWild sunflowers Wyethia spp. Aki-bi Food, medicineWild tobacco Nicotiana spp. Pa-mo Medicine Wildrye Elymus spp. Food Willows Salix spp. Se-he-be/See-hibe? Arrows and bows, basketry, 

medicine, musical instruments 

Woodland strawberry Fragaria vesca Food 

a Scientific names are based upon records in the Integrated Taxonomic Information System (http://www.ITIS.gov). 
85 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

 

English Equivalents 

When you know: Multiply by: To find: 

Centimeters (cm) .394 Inches 

Meters (m) 3.28 Feet 

Hectares (ha) 2.47 Acres 

Square meters (m2) 10.76 Square feet 

Kilograms (kg) 2.205 Pounds 

Square meters per hectare (m2/ha) 4.37 Square feet per acre 

Trees per hectare .405 Trees per acre 

References 

Abella, S.R.; Fule, P.Z. 2008. Fire effects on gambel oak in southwestern ponderosa 

pine-oak forests. Res. Note RMRS-RN-34. Fort Collins, CO: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station. 6 p. 

Abrams, M.D.; Nowacki, G.J. 2008. Native Americans as active and passive 

promoters of mast and fruit trees in the eastern USA. The Holocene. 18(7): 

1123–1137. 

Agee, J.K. 1993. Fire ecology of Pacific Northwest forests. Covelo, CA: Island 

Press. 493 p. 

Aginsky, B.W. 1943. Culture element distributions XXIV. In: Kroeber, A.L.; 

Gifford, E.W.; Lowie, R.H.; Olson, R.L., eds. Anthropological records. Berkeley, 

CA: University of California Press: 393–468. 

Anderson, M.K. 1993. Indian fire-based management in the sequoia-mixed conifer 

forests of the central and southern Sierra Nevada. Final report submitted to 

Yosemite Research Center, Yosemite National Park Cooperative agreement order 

number 8027-2-002. 126 p. 

Anderson, M.K. 2002. An ecological critique. In Stewart, O.C.; Lewis, H.T.; 

Anderson, M.K., eds. Forgotten fires: Native Americans and the transient 

wilderness. Norman, OK: University of Oklahoma Press: 37–64. 

Anderson, M.K. 2005. Tending the wild: Native American knowledge and the 

management of California’s natural resources. Berkeley, CA: University of 

California Press. 555 p. 

Anderson, M.K. 2007. Indigenous uses management, and restoration of oaks of the 

far Western United States. Tech. Note No. 2. Washington, DC: U.S. Department 

of Agriculture, Natural Resources Conservation Service, National Plant Data 

Center. 20 p. 
86 

GENERAL TECHICAL REPORT PSW-GTR-252

Anderson, M.K. 2009. Historical Native American resource use, harvesting, and 

management of California’s oak communities. Final report to the University of 

California Integrated Hardwood Range Management Program. Berkeley, CA: 

University of California–Berkeley. 387 p. 

Anderson, M.K.; Barbour, M.G. 2003. Simulated indigenous management: a new 

model for ecological restoration in national parks. Ecological Restoration. 21(4): 

269–277. 

Anderson, M.K.; Lake, F.K. 2013. California Indian ethnomycology and 

associated forest management. Journal of Ethnobiology. 33 (1): 33–85. 

Anderson, M.K.; Moratto, M.J. 1996. Native American land-use practices and 

ecological impacts. In: SNEP Science Team and Special Consultants, ed. Status 

of the Sierra Nevada, Sierra Nevada Ecosystem Project Final Report to Congress. 

Vol. 2, Chapter 9. Davis, CA: University of California–Davis: 187–206. 

Anderson, R.S.; Carpenter, S.L. 1991. Vegetation change in Yosemite Valley, 

Yosemite National Park, California, during the protohistoric period. Madrono. 

38(1): 1–13. 

Arthur, M.A.; Alexander, H.D.; Dey, D.C.; Schweitzer, C.J.; Loftis, D.L. 2012. 

Refining the oak-fire hypothesis for management of oak-dominated forests of the 

eastern United States. Journal of Forestry. 110(5): 257–266. 

Ascending the Giants. 2015. Oregon champion tree registry. [Updated 1 May 2013]. 

http://ascendingthegiants.com/oregon-big-tree-registry.html. (22 June 2015). 

Barnett, H.G. 1937. Culture element distributions: VII, Oregon Coast. In: Kroeber, 

A.L.; Gifford, E.W.; Lowie, R.H.; Olson, R.L., eds. Anthropological records. 

Berkeley, CA: University of California Press: 155–204. 

Barr, P.M. 1946. The research program of Blodgett Forest of the University of 

California. Journal of Forestry. 44: 738–741. 

Barrett, R.H. 1980. Mammals of California oak habitats—management 

implications. In: Plumb, T.R., tech. coord. Proceedings of the symposium on 

the ecology, management, and utilization of California oaks. Gen. Tech. Rep. 

PSW-44. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific 

Southwest Forest and Range Experiment Station. 275–291. 

Baumhoff, M. 1978. Environmental background. In: Heizer, R.F., ed. California. 

Washington, DC: Smithsonian Institution: 16–24. 
87 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Baumhoff, M.A. 1963. Ecological determinants of aboriginal California 

populations. In: University of California Publications in American Archaeology 

and Ethnology, vol. 49(2). Berkeley, CA: University of California Press: 155–236. 

Beniamino, F.; Ponge, J.F.; Arpin, P. 1991. Soil acidification under the crown 

of oak trees. I. Spatial distribution. Forest Ecology and Management. 40(3–4): 

221–232. 

Bettinger, R.L.; Malhi, R.; McCarthy, H. 1997. Central place models of acorn 

and mussel processing. Journal of Archaeological Science. 24(10): 887–899. 

Bigelow, S.W.; North, M.P.; Salk, C.F. 2011. Using light to predict fuels-reduction 

and group-selection effects on succession in Sierran mixed-conifer forest. 

Canadian Journal of Forest Research. 41(10): 2051–2063. 

Bingham, M.A.; Simard, S. 2012. Ectomycorrhizal networks of Pseudotsuga 

menziesii var. glauca trees facilitate establishment of conspecific seedlings under 

drought. Ecosystems. 15(2): 188–199. 

Blicharska, M.; Mikusiński, G. 2014. Incorporating social and cultural 

significance of large old trees in conservation policy. Conservation Biology. 

28(6): 1558–1567. 

Bolsinger, C.L. 1980. Oaks in California’s commercial forests—volume, stand 

structure, and defect characteristics. In: Plumb, T.R., ed. Proceedings of the 

symposium on the ecology, management, and utilization of California oaks. Gen. 

Tech. Rep. PSW-GTR-44. Berkeley, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Forest and Range Experiment Station: 101–106. 

Bonner, F.T.; Karrfalt, R.P. 2008. The woody plant seed manual. Washington, 

DC: U.S. Department of Agriculture. 1223 p. 

Bottorff, J. 2007. Managing Pacific Northwest forests for band-tailed pigeons. 

Woodland fish and wildlife. Extension Bulletin MISC0564. Pullman, WA: 

University of Washington Cooperative Extension. 8 p. 

Bowyer, R.T.; Bleich, V.C. 1980. Preliminary recommendations for managing 

California black oak in the Sierra Nevada. In: Plumb, T.R., ed. Proceedings of 

the symposium on the ecology, management, and utilization of California oaks. 

Gen. Tech. Rep. PSW-44. Berkeley, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Forest and Range Experiment Station: 107–111. 

Brand, J.C.; Snow, B.J.; Nabhan, G.P.; Truswell, A.S. 1990. Plasma glucose and 

insulin responses to traditional Pima Indian meals. American Journal of Clinical 

Nutrition. 51(3): 416–420. 
88 

GENERAL TECHICAL REPORT PSW-GTR-252

Byers, B.A.; Cunliffe, R.N.; Hudak, A. 2001. Linking the conservation of culture 

and nature: A case study of sacred forests in Zimbabwe. Human Ecology. 29(2): 

187–218. 

Calder, W.J.; Lifferth, G.; Moritz, M.A.; Clair, S.B.S. 2010. Physiological 

effects of smoke exposure on deciduous and conifer tree species. International 

Journal of Forestry Research. 2010: 7. 

Carrillo, H.M.R. [N.d.]. Quercus kelloggii Newb. in the State of Baja California, 

Mexico. Manuscript in preparation. Centro de Investigaciones Forestales y 

Agropecuarias. On file with: J. Long, Pacific Southwest Research Station, 

1731 Research Park Dr., Davis, CA 95618. 

Chandler, K.R.; Chappell, N.A. 2008. Influence of individual oak (Quercus 

robur) trees on saturated hydraulic conductivity. Forest Ecology and 

Management. 256(5): 1222–1229. 

Chaney, R.W. 1918. The ecological significance of the Eagle Creek flora of the 

Columbia River Gorge. Journal of Geology. 26(7): 577–592. 

Charnley, S.; Fischer, A.P.; Jones, E.T. 2007. Integrating traditional and local 

ecological knowledge into forest biodiversity conservation in the Pacific 

Northwest. Forest Ecology and Management. 246(1): 14–28. 

Christensen, G.A.; Campbell, S.J.; Fried, J.S. 2008. California’s forest 

resources, 2001–2005: five-year Forest Inventory and Analysis report. Gen. 

Tech. Rep. PNW-GTR-763. Portland, OR: U.S. Department of Agriculture, 

Forest Service, Pacific Northwest Research Station. 183 p. 

Cocking, M.I.; Bey, M.; Schrodt, L. 2012a. The central Umpqua-mid-Klamath 

oak habitat conservation project: a cooperative conservation partnership 

initiative. Ashland, OR: Lomakatsi Restoration Project. 14 p. 

Cocking, M.I.; Varner, J.M.; Sherriff, R.L. 2012b. California black oak 

responses to fire severity and native conifer encroachment in the Klamath 

Mountains. Forest Ecology and Management. 270: 25–34. 

Cocking, M.I.; Varner, J.M.; Knapp, E.E. 2014. Long-term effects of fire 

severity on oak-conifer dynamics in the southern Cascades. Ecological 

Applications. 24(1): 94–107. 
89 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Codding, B.F.; Bohna, L.C.; Bird, R.B.; Aldern, J.D.; Bird, D.W.; Goode, R.W. 

[In press]. Co-evolutionary dynamics between indigenous fire regimes & acorn 

(Quercus kelloggii) use in the Sierra Nevada Range of California. Ethnobiology. 

Collins, B.M.; Everett, R.G.; Stephens, S.L. 2011. Impacts of fire exclusion and 

managed fire on forest structure in an old growth Sierra Nevada mixed-conifer 

forest. Ecosphere. 2(4): art 51. 

Collins, B.M.; Lydersen, J.M.; Everett, R.G.; Fry, D.L.; Stephens, S.L. 2015. 

Novel characterization of landscape-level variability in historical vegetation 

structure. Ecological Applications. 25(5): 1167–1174. 

Cristancho, S.; Vining, J. 2004. Culturally defined keystone species. Human 

Ecology Review. 11(2): 153–164. 

Crotteau, J.S.; Ritchie, M.W.; Varner, J.M.; Berrill, J.-P. 2015. Quercus 

kelloggii (Newb.) sprout response to fire severity in Northern California. In: 

Standiford, R.B., ed. Proceedings of the 7th California Oak Symposium. Gen. 

Tech. Rep. PSW-GTR-251. Albany, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Research Station: 377–386. 

De Leo, G.A.; Levin, S. 1997. The multifaceted aspects of ecosystem integrity. 

Conservation Ecology. 1(3): art3. 

de Magalhães, R.M.Q.; Schwilk, D.W. 2012. Leaf traits and litter flammability: 

evidence for non-additive mixture effects in a temperate forest. Journal of 

Ecology. 100(5): 1153–1163.
 

Devine, W.; Aubry, C.; Bower, A.; Miller, J.; Maggiulli Ahr, N. 2012. Climate 

change and forest trees in the Pacific Northwest: a vulnerability assessment and 

recommended actions for national forests. Olympia, WA: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Region. 102 p. 

Devine, W.D.; Harrington, C.A. 2006. Changes in Oregon white oak (Quercus 

garryana Dougl. ex Hook.) following release from overtopping conifers. Trees— 

Structure and Function. 20(6): 747–756. 

Devine, W.D.; Harrington, C.A. 2013. Restoration release of overtopped Oregon 

white oak increases 10-year growth and acorn production. Forest Ecology and 

Management. 291: 87–95. 

Dixon, R.B. 1907. The Shasta. Bulletin of the American Museum of Natural 

History. 17(5): 381–498. 
90 

GENERAL TECHICAL REPORT PSW-GTR-252

Dodd, R.S.; Kashani, N.; Afzal-Rafii, Z. 2002. Population diversity and evidence 

of introgression among the black oaks of California. In: Standiford, R.B.; 

McCreary, D.; Purcell, K.L., eds. Proceedings of the fifth symposium on oak 

woodlands: oaks in California’s changing landscape. Gen. Tech. Rep. PSWGTR-184. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific 

Southwest Research Station. 775–785. 

Dunning, D. 1925. Methods of cutting study, Sierra National Forest, California. 

Unpublished report on file. Davis, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Research Station. 34 p. 

Egerton-Warburton, L.M.; Querejeta, J.I.; Allen, M.F. 2007. Common 

mycorrhizal networks provide a potential pathway for the transfer of 

hydraulically lifted water between plants. Journal of Experimental Botany. 

58(6): 1473–1483. 

Eilers, E.J.; Klein, A.-M. 2009. Landscape context and management effects on 

an important insect pest and its natural enemies in almond. Biological Control. 

51(3): 388–394. 

Engber, E.A.; Varner, J.M.; Arguello, L.A.; Sugihara, N.G. 2011. The effects 

of conifer encroachment and overstory structure on fuels and fire in an oak 

woodland landscape. Fire Ecology. 7(2): 32–50. 

Engber, E.A.; Varner, J.M. 2012. Patterns of flammability of the California oaks: 

the role of leaf traits. Canadian Journal of Forest Research. 42(11): 1965–1975. 

Eriksen, C.; Hankins, D.L. 2014. The retention, revival, and subjugation of 

indigenous fire knowledge through agency fire fighting in Eastern Australia and 

California. Society & Natural Resources. 27(12): 1288–1303. 

Everett, R.G. 2008. Dendrochronology-based fire history of mixed-conifer forests 

in the San Jacinto Mountains, California. Forest Ecology and Management. 

256(11): 1805–1814. 

Folke, C. 2006. Resilience: The emergence of a perspective for social-ecological 

systems analyses. Global Environmental Change-Human and Policy Dimensions. 

16(3): 253–267. 

Fralish, J.S. 2004. The keystone role of oak and hickory in the Central hardwood 

forest. Gen. Tech. Rep. GTR-SRS-73. Asheville, NC: U.S. Department of 

Agriculture, Forest Service, Southern Research Station. 78–87. 
91 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Fry, D.L.; Stephens, S.L. 2006. Influence of humans and climate on the fire 

history of a ponderosa pine-mixed conifer forest in the southeastern Klamath 

Mountains, California. Forest Ecology and Management. 223(1–3): 428–438. 

Fryer, J.L. 2007. Quercus kelloggii. [Updated]. http://www.fs.fed.us/database/feis/ 

plants/tree/quekel/all.html. (22 June 2015). 

Fujimura, K.E.; Smith, J.E.; Horton, T.R.; Weber, N.S.; Spatafora, J.W. 2005. 

Pezizalean mycorrhizas and sporocarps in ponderosa pine (Pinus ponderosa) 

after prescribed fires in eastern Oregon, USA. Mycorrhiza. 15(2): 79–86. 

Garibaldi, A.; Turner, N. 2004. Cultural keystone species: implications for 

ecological conservation and restoration. Ecology and Society. 9(3): 1. 

Garibaldi, A. 2009. Moving from model to application: cultural keystone 

species and reclamation in Fort McKay, Alberta. Journal of Ethnobiology. 

29(2): 323–338. 

Garrison, B.A.; Otahal, C.D.; Triggs, M.L. 2002a. Age structure and growth 

of California black oak (Quercus kelloggii) in the Central Sierra Nevada, 

California. In: Standiford, R.B., McCreary, D.; Purcell, K.L., tech. eds. 

Proceedings of the fifth symposium on oak woodlands: oaks in California’s 

challenging landscape. Gen. Tech. Rep. PSW-GTR-184. Albany, CA: U.S. 

Department of Agriculture, Forest Service Pacific Southwest Research Station. 

665–679. 

Garrison, B.A.; Wachs, R.L.; Jones, J.S.; Triggs, M.L. 1998. Visual counts 

of acorns of California black oak (Quercus kelloggii) as an indicator of mast 

production. Western Journal of Applied Forestry. 13(1): 27–31. 

Garrison, B.A.; Wachs, R.L.; Giles, T.A.; Triggs, M.L. 2002b. Dead branches 

and other wildlife resources on California black oak (Quercus kelloggii). In: 

Laudenslayer, W.F., Jr.; Valentine, B.; Weatherspoon, C.P.; Lisle, T.E., eds. 

Proceedings of the symposium on the ecology and management of dead wood 

in western forests. Reno, NV: U.S. Department of Agriculture, Forest Service, 

Pacific Southwest Research Station: 593–604. 

Gersonde, R.; Battles, J.J.; O’Hara, K.L. 2004. Characterizing the light 

environment in Sierra Nevada mixed-conifer forests using a spatially explicit 

light model. Canadian Journal of Forest Research. 34(6): 1332–1342. 

Gifford, E.W. 1923. Western Mono myths. The Journal of American Folklore. 

36(142): 301–367. 
92 

GENERAL TECHICAL REPORT PSW-GTR-252

González-Cabán, A.; Loomis, J.B.; Griffin, D.; Wu, E.; McCollum, D.; 

McKeever, J.; Freeman, D. 2003. Economic value of big game habitat 

production from natural and prescribed fire. Res. Paper PSW-RP-249. Albany, 

CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research 

Station. 38 p. 

Goode, R.W. 1992. Cultural traditions endangered. Clovis, CA: Eagle Eye 

Enterprises, Inc. 52 p. 

Goode, R.W. 2014. Cultural burn. Clovis, CA: Eagle Eye Enterprises, Inc. 12 p. 

Greene, D.F.; Hesketh, M.; Pounden, E. 2010. Emergence of morel (Morchella) 

and pixie cup (Geopyxis carbonaria) ascocarps in response to the intensity of 

forest floor combustion during a wildfire. Mycologia. 102(4): 766–773. 

Greene, E.L. 1889. Illustrations of West American oaks from drawings by the 

late Albert Kellogg, M.D. San Francisco, CA: Sosqui Engraving and Printing Co. 

83 p. 

Griffin, J.R.; Critchfield, W.B. 1976. The distribution of forest trees in California, 

with supplement. Research Paper PSW-82. Berkeley, CA: U.S. Department of 

Agriculture, Forest Service, Pacific Southwest Research Station. 114 p. 

Haggerty, P.K. 1994. Damage and recovery in southern Sierra Nevada foothill 

oak woodland after a severe ground fire. Madroño. 41: 185–198. 

Handley, T.; Grulke, N.E. 2008. Interactive effects of O3 exposure on California 

black oak (Quercus kelloggii Newb.) seedlings with and without N amendment. 

Environmental Pollution. 156(1): 53–60. 

Harrington, C.A.; Devine, W.D. 2006. A practical guide to oak release. Gen. 

Tech. Rep. PNW-GTR-666. Portland, OR: U.S. Department of Agriculture, 

Forest Service Pacific Northwest Research Station. 24 p. 

Harrington, J.P. 1932. Karuk Indian myths. In: Smithsonian Institution Bureau of 

American Ethnology, ed. Bulletin. Vol. 107. Washington DC: U.S. Government 

Printing Office. 34 p. 

Harrington, M.G.; Hawksworth, F.G. 1990. Interactions of fire and dwarf 

mistletoe on mortality of southwestern ponderosa pine. In: Krammes, J.S., ed. 

Effects of fire management of Southwestern natural resources. Gen. Tech. Rep. 

RMRS-GTR-191. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Forest and Range Experiment Station: 234–240.
 
93 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Heffner, K. 1984. Following the smoke: contemporary plant procurement by the 

Indians of northwest California. Administrative report. On file with: Six Rivers 

National Forest, 1330 Bayshore Way, Eureka, CA 95501. 

Heisey, R.M.; Delwiche, C.C.; Virginia, R.A.; Wrona, A.F.; Bryan, B.A. 

1980. A new nitrogen-fixing non-legume—Chamaebatia foliolosa (Rosaceae). 

American Journal of Botany. 67(3): 429–431. 

Heizer, R.F.; Elsasser, A.B. 1980. The natural world of the California Indians. 

Berkeley, CA: University of California Press. 271 p. 

Holmes, K.A.; Veblen, K.E.; Young, T.P.; Berry, A.M. 2008. California oaks and 

fire: a review and case study. In: Merenlender, A.; McCreary, D.; Purcell, K.L., 

eds. Proceedings of the sixth California oak symposium: today’s challenges, 

tomorrow’s opportunities. Gen. Tech. Rep. PSW-GTR-217. Albany, CA: U.S. 

Department of Agriculture, Forest Service, Pacific Southwest Research Station: 

551–565. 

Holt, C. 1946. Shasta ethnography. University of California Anthropological 

Records. 3(4): 299–349. 

Hood, S.M. 2010. Mitigating old tree mortality in long-unburned, firedependent forests: a synthesis. Gen. Tech. Rep. RMRS-GTR-238. Fort Collins, 

CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research 

Station. 71 p. 

Huber, D.W.; McDonald, P.M. 1992. California’s hardwood resource: history 

and reasons for lack of a sustained hardwood industry. Gen. Tech. Rep. PSWGTR-135. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific 

Southwest Research Station. 14 p. 

Hunt, D. 2004. The power of the acorn: late Holocene settlement and resource 

distribution in the Central Sierra. Proceedings of the Society for California 

Archaeology. 14: 143–150. 

Innes, R.J.; Van Vuren, D.H.; Kelt, D.A.; Johnson, M.L.; Wilson, J.A.; Stine, 

P.A. 2007. Habitat associations of dusky-footed woodrats (Neotonia fuscipes) 

in mixed-conifer forest of the northern Sierra Nevada. Journal of Mammalogy. 

88(6): 1523–1531. 

Ishii, H.T.; Van Pelt, R.; Parker, G.G.; Nadkarni, N.M. 2004. Age-related 

development of canopy structure and its ecological functions. In: Lowman, M.D.; 

Rinker, H.B., eds. Forest canopies. San Diego, CA: Academic Press: 102–117. 

Chapter 5. 
94 

GENERAL TECHICAL REPORT PSW-GTR-252

Jack, K.R. 1916. An Indian’s view of burning, and a reply. California Fish and 

Game Journal. 2(4): 194–196. 

Jepson, W.L. 1910. The silva of California. Berkeley, CA: University of California 

Press. 480 p. 

Jimerson, T.M.; Carothers, S.K. 2002. Northwest California oak woodlands: 

environment, species composition, and ecological status. In: Standiford, R.B.; 

McCreary, D.; Purcell, K.L., tech. eds. Proceedings of the fifth symposium on 

oak woodlands: oaks in California’s changing landscape. Gen. Tech. Rep. PSWGTR-184. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific 

Southwest Research Station: 705–717. 

Johnson, E.M.; Rosenberg, D.K. 2006. Granary-site selection by acorn 

woodpeckers in the Willamette Valley, Oregon. Northwest Science. 80(3): 

177–183. 

Kauffman, J.B. 1986. The ecological response of the shrub component to 

prescribed burning in mixed conifer ecosystems. Berkeley, CA: University of 

California–Berkeley. 235 p. Ph.D. dissertation. 

Kauffman, J.B.; Martin, R.E. 1987. Effects of fire and fire suppression on 

mortality and mode of reproduction of California black oak (Quercus kelloggii 

Newb.). In: Plumb, T.R.; Pillsbury, M.N., eds. Proceedings of the symposium on 

multiple-use management of California hardwood resources. Gen. Tech. Rep. 

PSW-100. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific 

Southwest Forest and Range Experiment Station: 122–126. 

Kauffman, J.; Martin, R. 1990. Sprouting shrub response to different seasons 

and fuel consumption levels of prescribed fire in Sierra Nevada mixed conifer 

ecosystems. Forest Science. 36: 748–764. 

Keane, J.J. 2014. California spotted owl: scientific considerations for forest 

planning. In: Long, J.W.; Quinn-Davidson, L.N.; Skinner, C.N., eds. Science 

synthesis to support socioecological resilience in the Sierra Nevada and southern 

Cascade Range. Gen. Tech. Rep. PSW-GTR-247 Albany, CA: U.S. Department 

of Agriculture, Forest Service, Pacific Southwest Research Station: 437–467. 

Kerns, S.J. 1980. Observations on wildlife abundance in several California black 

oak habitats in Northern California. In: Plumb, T.R., ed. Proceedings of the 

symposium on the ecology, management, and utilization of California oaks. 

Gen. Tech. Rep. PSW-44. Berkeley, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Forest and Range Experiment Station: 354–359. 
95 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Kershner, J. 2014. Climate change vulnerability assessment for focal resources of 

the Sierra Nevada. Version 1.0. Report to the California Landscape Conservation 

Cooperative and U.S. Forest Service Region 5. Bainbridge Island, WA: 

EcoAdapt. 418 p. 

Kilgore, B.M. 1973. The ecological role of fire in Sierran conifer forests: its 

application to national park management. Quaternary Research. 3(3): 496–513. 

Knapp, E.E.; Estes, B.L.; Skinner, C.N. 2009. Ecological effects of prescribed 

fires season: a literature review and synthesis for managers. Gen. Tech. Rep. 

PSW-GTR-224. Albany, CA: U.S. Department of Agriculture, Forest Service, 

Pacific Southwest Research Station. 80 p. 

Knapp, E.; North, M.; Benech, M.; Estes, B. 2012. The variable-density thinning 

study at Stanislaus-Tuolumne Experimental Forest. In: North, M., ed. Managing 

Sierra Nevada forests. Gen. Tech. Rep. PSW-GTR-237. Albany, CA: U.S. 

Department of Agriculture, Forest Service, Pacific Southwest Research Station: 

127–139. Chapter 12. 

Kobziar, L.; Moghaddas, J.; Stephens, S.L. 2006. Tree mortality patterns 

following prescribed fires in a mixed conifer forest. Canadian Journal of Forest 

Research. 36(12): 3222–3238. 

Koenig, W.D.; Knops, J. 1995. Why do oaks produce boom-and-bust seed crops? 

California Agriculture. 49(5): 7–12. 

Koenig, W.D.; Knops, J.M.H. 2002. The behavior ecology of masting in oaks. 

In: McShea, W.J.; Healy, W.M., eds. Oak forest ecosystems: ecology and 

management for wildlife. Baltimore, MD: Johns Hopkins University Press: 

129–148. Chapter 9. 

Koenig, W.D.; Knops, J.M.H.; Carmen, W.J.; Stanback, M.T.; Mumme, R.L. 

1994. Estimating acorn crops using visual surveys. Canadian Journal of Forest 

Research. 24(10): 2105–2112. 

Kotliar, N.B. 2000. Application of the new keystone-species concept to prairie 

dogs: how well does it work? Conservation Biology. 14(6): 1715–1721. 

Krasnow, K.D.; Stephens, S.L. 2012. The burning QuKe question. In: York, R., 

ed. Proceedings of the Blodgett Forest Research Station workshop. Berkeley, CA: 

University of California–Berkeley: 32–33. 

Kroeber, A.L. 1925. Handbook of the Indians of California. Bulletin 78. 

Smithsonian Institute Bureau of American Ethnology. Washington, DC: U.S. 

Government Printing Office. 995 p. 
96 

GENERAL TECHICAL REPORT PSW-GTR-252

Kueppers, L.M.; Snyder, M.A.; Sloan, L.C.; Zavaleta, E.S.; Fulfrost, B. 

2005. Modeled regional climate change and California endemic oak ranges. 

Proceedings of the National Academy of Sciences of the United States of 

America. 102(45): 16281–16286. 

Kuhn, B.; Johnson, B. 2008. Status and trends of black oak (Quercus kelloggii) 

populations and recruitment in Yosemite Valley (a.k.a. preserving Yosemite’s 

oaks). Yosemite, CA: National Park Service. 50 p. 

Kuo, M.; Dewsbury, D.R.; O’Donnell, K.; Carter, M.C.; Rehner, S.A.; Moore, 

J.D.; Moncalvo, J.-M.; Canfield, S.A.; Stephenson, S.L.; Methven, A.S.; 

Volk, T.J. 2012. Taxonomic revision of true morels (Morchella) in Canada and 

the United States. Mycologia. 104(5): 1159–1177. 

Lake, F.K. 2013. Trails, fires and tribulations: tribal resource management and 

research issues in Northern California. Occasion: Interdisciplinary Studies in the 

Humanities. 5(March): 22 p. 

Lake, F.K.; Long, J.W. 2014. Fire and tribal cultural resources. In: Long, J.W.; 

Quinn-Davidson, L.N.; Skinner, C.N., eds. Science synthesis to support 

socioecological resilience in the Sierra Nevada and southern Cascade Range. 

Gen. Tech. Rep. PSW-GTR-247 Albany, CA: U.S. Department of Agriculture, 

Forest Service, Pacific Southwest Research Station: 173–185. 

Laudenslayer, W.F. 2008. Survivorship of raked and unraked trees through 

prescribed fires in conifer forests in northeastern California. In: Narog, M.G., 

tech. coord. Proceedings of the 2002 fire conference: managing fire and fuels 

in the remaining wildlands and open spaces of the Southwestern United States. 

Gen. Tech. Rep. PSW-GTR-189. Albany, CA: U.S. Department of Agriculture, 

Forest Service, Pacific Southwest Research Station: 73–82. 

Leach, H.R.; Hiehle, J.L. 1957. Food habits of the Tehama deer herd. California 

Fish and Game. 43(3): 161–178. 

Lee, G. 1998. Walking where we lived: memoirs of a Mono Indian family. Norman, 

OK: University of Oklahoma Press. 224 p. 

Lewis, H.T. 1994. Management fires vs corrective fires in Northern Australia—an 

analog for environmental-change. Chemosphere. 29(5): 949–963. 

Loarie, S.R.; Carter, B.E.; Hayhoe, K.; McMahon, S.; Moe, R.; Knight, C.A.; 

Ackerly, D.D. 2008. Climate change and the future of California’s endemic flora. 

PLOS ONE. 3(6): e2502. 
97 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Long, J.W.; Quinn-Davidson, L.N.; Goode, R.W.; Lake, F.K.; Skinner, C.N. 

2015. Restoring California black oak to support tribal values and wildlife. In: 

Standiford, R.B., ed. Proceedings of the 7th California Oak Symposium. Gen. 

Tech. Rep. PSW-GTR-251. Albany, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Research Station: 113–122. 

Long, J.W.; Quinn-Davidson, L.N.; Skinner, C.N. 2014. Science synthesis to 

support socioecological resilience in the Sierra Nevada and southern Cascade 

Range. Gen. Tech. Rep. PSW-GTR-247. Albany, CA: U.S. Department of 

Agriculture, Forest Service, Pacific Southwest Research Station. 723 p. 

Long, W.H. 1913. Three undescribed heart rots of hardwood trees especially of 

oak. Journal of Agricultural Research. 1(1): 109–128. 

Loomis, J.B.; Loft, E.R.; Garrison, B.A. 1995. An economic assessment of 

hardwoods as habitat components for black-tailed deer in Northern California 

conifer forests. Journal of Forestry. 93(8): 41–45. 

Luoma, D.L.; Eberhart, J.L.; Abbott, R.; Moore, A.; Amaranthus, M.P.; 

Pilz, D. 2006. Effects of mushroom harvest technique on subsequent American 

matsutake production. Forest Ecology and Management. 236(1): 65–75. 

Lutz, J.A.; van Wagtendonk, J.W.; Franklin, J.F. 2009. Twentieth-century 

decline of large-diameter trees in Yosemite National Park, California, USA. 

Forest Ecology and Management. 257(11): 2296–2307. 

Lutz, J.A.; van Wagtendonk, J.W.; Franklin, J.F. 2010. Climatic water deficit, 

tree species ranges, and climate change in Yosemite National Park. Journal of 

Biogeography. 37(5): 936–950. 

Lynn, K.; Daigle, J.; Hoffman, J.; Lake, F.; Michelle, N.; Ranco, D.; Viles, 

C.; Voggesser, G.; Williams, P. 2013. The impacts of climate change on tribal 

traditional foods. Climatic Change. 120(3): 545–556. 

MacRoberts, M.H.; MacRoberts, B.R. 1976. Social organization and behavior of 

the acorn woodpecker in central coastal California. Ornithological Monographs. 

21: 115. 

Mathiasen, R.L.; Nickrent, D.L.; Shaw, D.C.; Watson, D.M. 2008. Mistletoes: 

pathology, systematics, ecology, and management. Plant Disease. 92(7): 

988–1006. 
98 

GENERAL TECHICAL REPORT PSW-GTR-252

Mazur, R.; Klimley, A.P.; Folger, K. 2013. Implications of the variable 

availability of seasonal foods on the home ranges of black bears, Ursus 

americanus, in the Sierra Nevada of California. Animal Biotelemetry. 1(1): 16. 

Mazurek, M.J.; Zielinski, W.J. 2004. Individual legacy trees influence vertebrate 

wildlife diversity in commercial forests. Forest Ecology and Management. 

193(3): 321–334.
 

McCreary, D.; Nader, G. 2011. Burned oaks: Which ones will survive? 

Publication 8445. Davis, CA: University of California, Agriculture and Natural 

Resources. 5 p. 

McDonald, P.M. 1969. Silvical characteristics of California black oak (Quercus 

kelloggii Newb.). Res. Pap. PSW-53. Berkeley, CA: U.S. Department of 

Agriculture, Forest Service, Pacific Southwest Research Station. 26 p. 

McDonald, P.M. 1990. Quercus kelloggii Newb. California black oak. In: Burns, 

R.M.; Honkala, B.H., eds. Silvics of North America. Volume 2, hardwoods. 

Washington, DC: U.S. Department of Agriculture: 661–671. 

McDonald, P.M.; Huber, D.W. 1995a. California’s hardwood resource: status of 

the industry and an ecosystem management perspective. Gen. Tech. Rep. PSWGTR-153. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific 

Southwest Research Station. 24 p. 

McDonald, P.M.; Huber, D.W. 1995b. California’s hardwood resource: managing 

for wildlife, water, pleasing scenery, and wood products. Gen. Tech. Rep. PSWGTR-154. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific 

Southwest Research Station. 23 p. 

McDonald, P.M.; Ritchie, M.W. 1994. Epicormic branching of California black 

oak—effect of stand and tree characteristics. Northwest Science. 68(1): 6–10. 

McDonald, P.M.; Tappeiner, J.C. 2002. California’s hardwood resource: seeds, 

seedlings, and sprouts of three important forest-zone species. Gen. Tech. Rep. 

PSW-GTR-185. Albany, CA: U.S. Department of Agriculture, Forest Service, 

Pacific Southwest Research Station. 39 p. 

McDonald, P.M.; Vaughn, N.R. 2007. Growth of thinned and unthinned hardwood 

stands on a good site in northern California. Gen. Tech. Rep. PSW-GTR-204. 

Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest 

Research Station. 23 p. 
99 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

McElwain, J.C. 2004. Climate-independent paleoaltimetry using stomatal density 

in fossil leaves as a proxy for CO2 partial pressure. Geology. 32(12): 1017–1020. 

McMinn, H.E.; Babcock, E.B.; Righter, F.I. 1949. The Chase oak, a new giant 

hybrid oak from Santa Clara County, California. Madrono. 10: 51–55. 

McShea, W.J.; Healy, W.M. 2002. Oaks and acorns as a foundation for ecosystem 

management. In McShea, W.J.; Healy, W.M., eds. Oak forest ecosystems: 

ecology and management for wildlife. Baltimore, MD: Johns Hopkins University 

Press: 1–9. 

Meier, A.R.; Saunders, M.R.; Michler, C.H. 2012. Epicormic buds in trees: 

a review of bud establishment, development and dormancy release. Tree 

Physiology. 32(5): 565–584. 

Menge, B.A.; Freidenburg, T.L. 2001. Keystone species. In: Levin, S.A., ed. 

Encyclopedia of biodiversity. Volume 3. San Diego, CA: Academic Press: 

613–631. 

Merriam, K. 2013. Natural range of variation in hardwood vegetation in the Sierra 

Nevada, California over the Holocene epoch. [Updated]. http://www.fs.usda.gov/ 

Internet/FSE_DOCUMENTS/stelprdb5434343.pdf. (26 February 2015). 

Messner, T.C. 2011. Acorns and bitter roots: starch grain research in the prehistoric 

Eastern woodlands. Tuscaloosa, AL: University of Alabama Press. 216 p. 

Meyer, M.D.; Schroer, G.L.; Stalter, B. 2013. “Methods of cutting” ecological 

inventory final report. Unpublished report. On file with: Marc Meyer, Southern 

Sierra Nevada Province ecologist, 1600 Tollhouse Road, Clovis, CA 93611. 25 p. 

Mills, L.S.; Soule, M.E.; Doak, D.F. 1993. The keystone-species concept in 

ecology and conservation. BioScience. 43(4): 219–224. 

Monleon, V.J.; Lintz, H.E. 2015. Evidence of tree species’ range shifts in a 

complex landscape. PLOS ONE. 10(1): e0118069. 

Morgan, C. 2008. Reconstructing prehistoric hunter-gatherer foraging radii: a 

case study from California’s Southern Sierra Nevada. Journal of Archaeological 

Science. 35(2): 247–258. 

Moritz, M.A.; Odion, D.C. 2005. Examining the strength and possible causes of 

the relationship between fire history and sudden oak death. Oecologia. 144(1): 

106–114. 
100 

GENERAL TECHICAL REPORT PSW-GTR-252

Nesmith, J.C.B.; O’Hara, K.L.; van Mantgem, P.J.; de Valpine, P. 2010. The 

effects of raking on sugar pine mortality following prescribed fire in Sequoia 

and Kings Canyon National Parks, California, USA. Fire Ecology. 6(3): 97–116. 

Norgaard, K.M. 2014. The politics of fire and the social impacts of fire exclusion 

on the Klamath. Humboldt Journal of Social Relations. 36: 77–101. 

North, M.; Stine, P.A.; O’Hara, K.L.; Zielinski, W.J.; Stephens, S.L. 2009. 

An ecosystems management strategy for Sierra mixed-conifer forests, with 

addendum. Gen. Tech. Rep. PSW-GTR-220. Albany, CA: U.S. Department 

of Agriculture, Forest Service, Pacific Southwest Research Station. 49 p. 

North, M.; Trappe, J.; Franklin, J. 1997. Standing crop and animal consumption 

of fungal sporocarps in Pacific Northwest forests. Ecology. 78(5): 1543–1554. 

North, M.P.; Collins, B.M.; Keane, J.; Long, J.W.; Skinner, C.N.; Zielinski, 

W.J. 2014. Synopsis of emergent approaches. In: Long, J.W.; Quinn-Davidson, 

L.; Skinner, C.N., eds. Science synthesis to support socioecological resilience in 

the Sierra Nevada and southern Cascade Range. Gen. Tech. Rep. PSW-GTR-247. 

Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest 

Research Station: 55–69. 

Ortiz, B.R. 1991. It will live forever: traditional Yosemite Indian acorn making. 

Berkeley, CA: Heyday Books. 148 p. 

Ortiz, B.R. 2008. Contemporary California Indians, oaks, and sudden oak death 

(Phytophthora ramorum). In: Merenlender, A.; McCreary, D.; Purcell, K.L., 

eds. Proceedings of the sixth California oak symposium: today’s challenges, 

tomorrow’s opportunities. Gen. Tech. Rep. GTR-PSW-217. Albany, CA: U.S. 

Department of Agriculture, Forest Service Pacific Southwest Research Station: 

39–56. 

Otvos, I.S.; Mills, D.A.; Conder, N. 2012. Within-crown distribution, attack, 

and germination of Curculio occidentis-damaged and Cydia latiferreanadamaged Garry oak acorns in Victoria, British Columbia, Canada. Canadian 

Entomologist. 144(3): 419–434. 

Paine, R.T. 1995. A conversation on refining the concept of keystone species. 

Conservation Biology. 9(4): 962–964. 

Parmeter, J.R.J.; Uhrenholdt, B. 1974. Some effects of pine-needle or grass 

smoke on fungi. Phytopathology. 65: 28–31. 
101 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Parsons, D.J.; DeBenedetti, S.H. 1979. Impact of fire suppression on a mixedconifer forest. Forest Ecology and Management. 2: 21–33. 

Pavlik, B.M.; Muick, P.C.; Johnson, S.G.; Popper, M. 1995. Oaks of California. 

Los Olivos, CA: Cachuma Press and the California Oak Foundation. 184 p. 

Peter, D.; Harrington, C. 2002. Site and tree factors in Oregon white oak 

acorn production in western Washington and Oregon. Northwest Science. 

76(3): 189–201. 

Pilz, D.; McLain, R.; Alexander, S.; Villareal-Ruiz, L.; Berch, S.; Wurtz, T.L.; 

Parks, C.G.; McFarlane, E.; Baker, B.; Molina, R.; Smith, J.E. 2007. Ecology 

and management of commercially harvested morels in Western North America. 

Gen. Tech. Rep. PNW-GTR-710. Portland, OR: U.S. Department of Agriculture, 

Forest Service, Pacific Northwest Research Station. 161 p. 

Pilz, D.; Molina, R. 1996. Managing forest ecosystems to conserve fungus 

diversity and sustain wild mushroom harvest. Gen. Tech. Rep. PNW-GTR-371. 

Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest 

Research Station. 104 p. 

Pilz, D.; Weber, N.S.; Carter, M.C.; Parks, C.G.; Molina, R. 2004. Productivity 

and diversity of morel mushrooms in healthy, burned, and insect-damaged 

forests of northeastern Oregon. Forest Ecology and Management. 198(1–3): 

367–386. 

Plumb, T.R. 1980. Response of oaks to fire. In: Plumb, T.R., ed. Proceedings of 

the symposium on the ecology, management, and utilization of California oaks. 

Gen. Tech. Rep. PSW-44. Berkeley, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Forest and Range Experiment Station: 202–215. 

Potter, D.A.; Johnston, B. 1980. An approach to managing California black oak 

and hardwoods on a deer winter range in Central California. In: Plumb, T.R., ed. 

Proceedings of the symposium on the ecology, management, and utilization of 

California oaks. Gen. Tech. Rep. PSW-GTR-44. Berkeley, CA: U.S. Department 

of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment 

Station: 112–116. 

Power, M.E.; Tilman, D.; Estes, J.A.; Menge, B.A.; Bond, W.J.; Mills, L.S.; 

Daily, G.; Castilla, J.C.; Lubchenco, J.; Paine, R.T. 1996. Challenges in the 

quest for keystones. BioScience. 46(8): 609–620. 
102 

GENERAL TECHICAL REPORT PSW-GTR-252

Purcell, K.L.; Drynan, D.A. 2008. Use of hardwood tree species by birds nesting 

in ponderosa pine forests. In: Merenlender, A.; McCreary, D.; Purcell, K.L., 

eds. Proceedings of the sixth California oak symposium: today’s challenges, 

tomorrow’s opportunities. Gen. Tech. Rep. PSW-GTR-217. Albany, CA: U.S. 

Department of Agriculture, Forest Service, Pacific Southwest Research Station: 

417–431. 

Purcell, K.L.; Mazzoni, A.K.; Mori, S.R.; Boroski, B.B. 2009. Resting 

structures and resting habitat of fishers in the southern Sierra Nevada, California. 

Forest Ecology and Management. 258(12): 2696–2706. 

Purcell, K.L.; Thompson, C.M.; Zielinski, W.J. 2012. Fishers and american 

martens. In: North, M., ed. Managing Sierra Nevada forests. Gen. Tech. Rep. 

PSW-GTR-237. Albany, CA: U.S. Department of Agriculture, Forest Service, 

Pacific Southwest Research Station: 47–59. 

Regelbrugge, J.C.; Conard, S.G. 1993. Modeling tree mortality following 

wildfire in Pinus ponderosa forests in the central Sierra-Nevada of California. 

International Journal of Wildland Fire. 3(3): 139–148. 

Ripple, W.J.; Beschta, R.L. 2008. Trophic cascades involving cougar, mule deer, 

and black oaks in Yosemite National Park. Biological Conservation. 141(5): 

1249–1256. 

Rizzo, D.M.; Garbelotto, M.; Hansen, E.M. 2005. Phytophthora ramorum: 

integrative research and management of an emerging pathogen in California and 

Oregon forests. Annual Review of Phytopathology. 43(1): 309–335. 

Rossi, R.S. 1980. History and cultural influences on the distribution and 

reproduction of oaks in California. In: Plumb, T.R., tech. coord. Proceedings of 

the symposium on the ecology, management, and utilization of California oaks. 

Gen. Tech. Rep. PSW-44. Berkeley, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Forest and Range Experiment Station: 7–18. 

Royer, D.L. 2012. Leaf shape responds to temperature but not CO2

 in Acer rubrum. 

PLOS ONE. 7(11): e49559. 

Royer, D.L. 2015. RE: phenotypic plasticity and genetic determination in Quercus 

kelloggii. to Long, J.W. 12 May 2015. 

Royer, D.L.; McElwain, J.C.; Adams, J.M.; Wilf, P. 2008. Sensitivity of leaf 

size and shape to climate within Acer rubrum and Quercus kelloggii. New 

Phytologist. 179(3): 808–817. 
103 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Royer, D.L.; Meyerson, L.A.; Robertson, K.M.; Adams, J.M. 2009. Phenotypic 

plasticity of leaf shape along a temperature gradient in Acer rubrum. PLOS 

ONE. 4(10): e7653. 

Sackett, S.S.; Haase, S.M.; Burke, G.M. 1992. Soil temperatures related to 

prescribed burning in ponderosa pine stands in Yosemite Valley that include 

California black oak. In: Ffolliott, P.F., ed. Ecology and management of oak 

and associated woodlands: perspectives in the southwestern United States and 

northern Mexico. Gen. Tech. Rep. RMRS-GTR-218. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Forest and Range 

Experiment Station: 144–146. 

Safford, H.D.; Stevens, J.T.; Merriam, K.; Meyer, M.D.; Latimer, A.M. 2012. 

Fuel treatment effectiveness in California yellow pine and mixed conifer forests. 

Forest Ecology and Management. 274: 17–28. 

Sapir, E. 1907. Notes on the Takelma Indians of Southwestern Oregon. American 

Anthropologist. 9(2): 251–275. 

Schenck, S.M.; Gifford, E.W. 1952. Karok ethnobotany. University of California 

Anthropological Records. 13(6): 377–392. 

Siegel, J.; Kuenen, L.; Higbee, B.; Noble, P.; Gill, R.; Yokota, G.; Krugner, 

R.; Daane, K. 2008. Postharvest survival of navel orangeworm assessed in 

pistachios. California Agriculture. 62(1): 30–35. 

Skinner, C.N. 1995. Using prescribed fire to improve wildlife habitat near Shasta 

Lake. Unpublished report. Albany, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Research Station. 26 p. 

Skinner, C.N.; Chang, C. 1996. Fire regimes, past and present. In: SNEP Science 

Team and Special Consultants, ed. Status of the Sierra Nevada, Sierra Nevada 

Ecosystem Project Final Report to Congress. Vol. II. Davis, CA: University of 

California–Davis: 1041–1069. 

Skinner, C.N.; Taylor, A.H.; Agee, J.K. 2006. Klamath Mountains bioregion. In 

Sugihara, N.G.; van Wagtendonk, J.W.; Fites-Kaufman, J.; Shaffer, K.E.; Thode, 

A.E., eds. Fire in California’s ecosystems. Berkeley, CA: University of California 

Press: 170–194. 

Smith, H.V. 1941. A Miocene flora from Thorn Creek, Idaho. American Midland 

Naturalist. 25(3): 473–522. 
104 

GENERAL TECHICAL REPORT PSW-GTR-252

Southworth, D.; Donohue, J.; Frank, J.L.; Gibson, J. 2011. Mechanical 

mastication and prescribed fire in conifer–hardwood chaparral: differing 

responses of ectomycorrhizae and truffles. International Journal of Wildland 

Fire. 20(7): 888–896. 

St. John, T.; Rundel, P. 1976. The role of fire as a mineralizing agent in a Sierran 

coniferous forest. Oecologia. 25(1): 35–45. 

Stephens, S.L. 1997. Fire history of a mixed oak-pine forest in the foothills of the 

Sierra Nevada, El Dorado County, California. In: Pillsbury, N.H.; Verner, J.; 

Tietje, W.D., eds. Proceedings of the symposium on oak woodlands: ecology, 

management, and urban interface issues. Gen. Tech. Rep. PSW-GTR-160. 

Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest 

Research Station: 191–198. 

Stephens, S.L. 2000. Mixed conifer and upper montane forest structure and uses in 

1899 from the central and northern Sierra Nevada, CA. Madroño. 47(1): 43–52. 

Stephens, S.L.; Lydersen, J.M.; Collins, B.M.; Fry, D.L.; Meyer, M.D. 2015. 

Historical and current landscape-scale ponderosa pine and mixed conifer forest 

structure in the Southern Sierra Nevada. Ecosphere. 6(5): 79. 

Stephens, S.L.; Moghaddas, J.J. 2005. Experimental fuel treatment impacts 

on forest structure, potential fire behavior, and predicted tree mortality in a 

California mixed conifer forest. Forest Ecology and Management. 215(1–3): 

21–36. 

Stevenot, K.R. 2008. Ti-Le-Li: Northern Mewuk acorn. In Dubin, M.; Tolley, 

S.-L., eds. Seaweed, salmon and manzanita cider: A California Indian feast. 

Berkeley, CA: Heyday Books: 99–103. 

Stevens, N.E. 2002. Spatial and temporal patterning of bedrock mortar sites in the 

southern Sierra Nevada: a regional exploration. Proceedings of the Society of 

California Archaeology. 16: 175–182. 

Stine, P.A.; Conway, S. 2012. Applying GTR 220 concepts on the Sagehen 

Experimental Forest. In: North, M., ed. Managing Sierra Nevada forests. Gen. 

Tech. Rep. PSW-GTR-237. Albany, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Research Station: 141–147. 

Sturrock, R.N.; Frankel, S.J.; Brown, A.V.; Hennon, P.E.; Kliejunas, J.T.; 

Lewis, K.J.; Worrall, J.J.; Woods, A.J. 2011. Climate change and forest 

diseases. Plant Pathology. 60(1): 133–149. 
105 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Swetnam, T.W.; Baisan, C.H. 2003. Tree-ring reconstructions of fire and climate 

history in the Sierra Nevada and Southwestern United States. In: Veblen, T.T.; 

Baker, W.L.; Montenegro, G.; Swetnam, T.W., eds. Fire and climatic change in 

temperate ecosystems of the western Americas. Vol. 160. New York: Springer: 

158–195. Chapter 6. 

Swiecki, T.J.; Bernhardt, E.A. 2006. A field guide to insects and diseases of 

California oaks. Gen. Tech. Rep. PSW-GTR-197. Albany, CA: U.S. Department 

of Agriculture, Forest Service, Pacific Southwest Research Station. 156 p. 

Tappeiner, J.; McDonald, P.M. 1980. Preliminary recommendations for managing 

California black oak in the Sierra Nevada. In: Plumb, T.R., ed. Proceedings of 

the symposium on the ecology, management, and utilization of California oaks. 

Gen. Tech. Rep. PSW-GTR-44. Berkeley, CA: U.S. Department of Agriculture, 

Forest Service, Pacific Southwest Forest and Range Experiment Station: 107–111. 

Taylor, A.H. 2010. Fire disturbance and forest structure in an old-growth Pinus 

ponderosa forest, southern Cascades, USA. Journal of Vegetation Science. 21(3): 

561–572. 

Taylor, A.H.; Beaty, R.M. 2005. Climatic influences on fire regimes in the 

northern Sierra Nevada mountains, Lake Tahoe Basin, Nevada, USA. Journal of 

Biogeography. 32(3): 425–438. 

Taylor, A.H.; Skinner, C.N. 2003. Spatial patterns and controls on historical fire 

regimes and forest structure in the Klamath Mountains. Ecological Applications. 

13(3): 704–719. 

Tevis, L.J. 1952. Autumn foods of chipmunks and golden-mantled ground squirrels 

in the northern Sierra Nevada. Journal of Mammalogy. 33(2): 198–205. 

Thorne, J.; Bjorkman, J.; Thrasher, S.; Kelsey, R.; Morgan, B.J. 2007. 

1930s extent of oak species in the central Sierra Nevada. In: Merenlender, 

A.; McCreary, D.; Purcell, K.L., eds. Proceedings of the sixth California oak 

symposium: today’s challenges, tomorrow’s opportunities. GTR-PSW-217. 

Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest 

Research Station: 569–587. 

Thorne, J.H.; Morgan, B.J.; Kennedy, J.A. 2008. Vegetation change over sixty 

years in the Central Sierra Nevada, California, USA. Madrono. 55(3): 223–237. 

Thysell, D.R.; Carey, A.B. 2001. Quercus garryana communities in the Puget 

Trough, Washington. Northwest Science. 75(3): 219–235. 
106 

GENERAL TECHICAL REPORT PSW-GTR-252

Truex, R.L.; Zielinski, W.J. 2013. Short-term effects of fuel treatments on 

fisher habitat in the Sierra Nevada, California. Forest Ecology and Management. 

293: 85–91. 

Urban Forest Ecosystems Institute [UFEI]. 2015. California registry of big trees. 

[Updated 1/10/2012]. http://californiabigtrees.calpoly.edu/. (16 January 2015). 

U.S. Geological Survey. 1999. Digital representations of tree species range 

maps from “Atlas of United States Trees” by Elbert L. Little, Jr. (and other 

publications). [Updated]. Reston, VA: Geosciences and Environmental Change 

Science Center. http://esp.cr.usgs.gov/data/little/. (26 February 2015). 

U.S. Department of Agriculture, Forest Service [USDA FS]. 2004. Sierra 

Nevada forest plan amendment, final supplemental enviromental impact 

statement. Volume 4, Appendix E—Adaptive Management Strategy. Vallejo, 

CA: Pacific Southwest Region. 134 p. 

U.S. Department of Agriculture, Forest Service [USDA FS]. 2014. The Pacific 

Northwest Forest Inventory and Analysis database (PNW-FIADB), 2001–2012. 

[Updated May 2014]. http://www.fs.fed.us/pnw/fia/publications/data/. 

(21 November 2014). 

Van de Water, K.M.; Safford, H.D. 2011. A summary of fire frequency estimates 

for California vegetation before Euro-American settlement. Fire Ecology. 7(3): 

26–58. 

van Wagtendonk, J.W.; Fites-Kaufman, J. 2006. Sierra Nevada bioregion. In: 

Sugihara, N.G.; van Wagtendonk, J.W.; Fites-Kaufman, J.; Shaffer, K.E.; Thode, 

A.E., eds. Fire in California’s ecosystems. Berkeley, CA: University of California 

Press: 264–294. 

Vankat, J.L.; Major, J. 1978. Vegetation changes in Sequoia National Park, 

California. Journal of Biogeography. 5(4): 377–402. 

Venette, R.C.; Coleman, T.W.; Seybold, S.J. 2015. Assessing the risks posed 

by goldspotted oak borer to California and beyond. In: Standiford, R.B., ed. 

Proceedings of the 7th California Oak Symposium. Gen. Tech. Rep. PSWGTR-251. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific 

Southwest Research Station: 317–329. 

Voggesser, G.; Lynn, K.; Daigle, J.; Lake, F.; Ranco, D. 2013. Cultural impacts 

to tribes from climate change influences on forests. Climatic Change. 120(3): 

615–626. 
107 

Restoring California Black Oak Ecosystems to Promote Tribal Values and Wildlife

Waddell, K.L.; Barrett, T.M. 2005. Oak woodlands and other hardwood forests 

of California, 1990s. Res. Bull. PNW-RB-245. Portland, OR: U.S. Department 

of Agriculture, Forest Service, Pacific Northwest Research Station. 94 p. 

Waring, K.M.; O’Hara, K.L. 2008. Redwood/tanoak stand development and 

response to tanoak mortality caused by Phytophthora ramorum. Forest Ecology 

and Management. 255(7): 2650–2658. 

Wayman, R.B.; North, M. 2007. Initial response of a mixed-conifer understory 

plant community to burning and thinning restoration treatments. Forest Ecology 

and Management. 239(1–3): 32–44. 

Webster, K.M.; Halpern, C.B. 2010. Long-term vegetation responses to 

reintroduction and repeated use of fire in mixed-conifer forests of the Sierra 

Nevada. Ecosphere. 1(5): art9. 

Weigand, J.F. 1998. Management experiments for high-elevation agroforestry 

systems jointly producing matsutake mushrooms and high-quality timber in the 

Cascade Range of southern Oregon. Gen. Tech. Rep. PNW-GTR-424. Portland, 

OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research 

Station. 42 p. 

Wolf, C.B. 1944. The gander oak, a new hybrid oak from San Diego County, 

California. Proceedings of the California Academy of Sciences. 25(5): 177–188. 

Yaeger, J.S. 2005. Habitat at fisher resting sites in the Klamath Province of 

Northern California. Arcata, CA: Humboldt State University. 64 p. M.S. thesis. 

Zielinski, W.J.; Truex, R.L.; Dunk, J.R.; Gaman, T. 2006. Using forest 

inventory data to assess fisher resting habitat suitability in California. Ecological 

Applications. 16(3): 1010–1025. 

Zielinski, W.J.; Dunk, J.R.; Gray, A.N. 2012. Estimating habitat value using 

forest inventory data: the fisher (Martes pennanti) in northwestern California. 

Forest Ecology and Management. 275: 35–42. 

Zielinski, W.J. 2014. The forest carnivores: marten and fisher. In: Long, J.W.; 

Quinn-Davidson, L.N.; Skinner, C.N., eds. Science synthesis to support 

socioecological resilience in the Sierra Nevada and southern Cascade Range. 

Gen. Tech. Rep. PSW-GTR-247 Albany, CA: U.S. Department of Agriculture, 

Forest Service, Pacific Southwest Research Station: 393–435. 

Zimmerman, G.T.; Laven, R.D. 1987. Effects of forest fuel smoke on dwarf 

mistletoe seed-germination. Great Basin Naturalist. 47(4): 652–659. 
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Glossary
 

Corrective fire—Fires set in relatively dense stands of vegetation to reduce accumulated litter and understory growth in areas that have missed expected recurrence 

of fires (Lewis 1994). 

Cultural burn—Use of fire to stimulate desired conditions for traditional cultural 

purposes. 

Cultural keystone—Species that significantly shape the cultural identity of a 

people, as reflected in diet, materials, medicine, and/or spiritual practice. 

Deciduous—The propensity of a tree to shed its leaves (in winter in the case of 

black oak). 

Desired condition—Descriptions of goals to be achieved at some time in the 

future, from the landscape-scale down to fine-scales. 

Diameter at breast height (dbh)—A measure of the diameter of a tree trunk at 

1.37 m (4.5 ft) above the ground. 

Duff—A component of the forest floor, below the litter and above the mineral soil, 

consisting of partly decomposed organic material. 

Ecological keystone—Species that have disproportionately large ecosystem 

impacts relative to their abundance. 

Ecosystem services—Goods and services provided by natural ecosystems that are 

important for human well-being. These may be tangible goods like acorns, intangible benefits such as spiritual renewal and sense of place, and ecological functions 

that regulate processes such as fire and water flows. 

Epicormic branches—Shoots that arise from adventitious or dormant buds on the 

stem or branch of a woody plant, often after fire or increases in sunlight. 

Interception—Precipitation that does not reach the soil, but is instead retained on 

leaves and branches of plants and the forest floor. 

Food web—A system of interconnected and interdependent food chains. 

Goldilocks principle—Conditions need to fall within certain margins (“not too 

hot, not too cold”) to support a desired condition (“just right”). 

Granary tree—A tree, often dead, in which woodpeckers drill holes to store 

acorns. 

Heart rot—Rotting of the bole and large limbs of living trees by fungal pathogens. 
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Hybrid—An offspring of two animals or plants of different breeds, varieties, species, or genera. 

Initial entry burn—A preparatory burn set to reduce continuity and amount of 

surface fuels, but not likely to modify forest structure. Such burns are likely to be 

conducted well outside the natural wildfire season when moisture limits intensity. 

Jackpot burning—A prescribed burn that consumes “jackpots” or concentrations of fuels that are discontinuous, so the fire will not spread easily between 

concentrations. 

Knocking—A traditional practice for gathering acorns by hitting the trees with 

long poles. 

Legacy tree—A tree that is especially large and old. 

Live-crown ratio—The vertical distance from the top of the tree to the base of the 

lowest live branch (excluding epicormic branches) divided by the tree’s total height. 

Maintenance burn—A fire that kills smaller trees and reduces fuels but is not 

likely to kill mature, healthy trees due to already low fuel levels. 

Mast—Fruit produced by trees or shrubs that is eaten by animals. Hard mast refers 

to nuts in a hard shell, in contrast to soft mast such as berries. 

Mummy nut—An unharvested nut left in the tree or on the ground, which can be a 

reservoir for pests and diseases. 

Mutualism—A relationship in which two organisms of different species benefit 

from the activity of each other. 

Mycelial mats (also called shiro)—The vegetative part of a fungus, consisting of a 

mass of branching, thread-like structures called hyphae. 

Mycorrhizal association or fungi—Colonization of a plant’s roots by a fungus, 

often in a way that benefits the host plant. 

Out-of-season burn—Use of fire outside of the historical fire season in order to 

reduce fuels and alter stand structure sufficiently to prepare the site to more safely 

accept fire during the historical fire season. 

Phenology—The timing of life cycle events (such as acorn ripening) and how these 

are influenced by climate, elevation, and other factors. 

Release—Removing competing overstory trees to increase light, water, and nutrients to benefit trees that are currently suppressed. 
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GENERAL TECHICAL REPORT PSW-GTR-252

Restorative burn—A fire that reduces fuels and may kill trees of various sizes 

to realign forest structure and composition closer to a reference condition. Some 

wildfires could meet this description. 

Socioecological resilience—The capacity of systems to cope with, adapt to, and 

influence change; to persist and develop in the face of change; and to innovate and 

transform into new, more desirable configurations in response to disturbance. 

Top-kill—Aboveground portion of plant is killed while the belowground portion 

survives and is able to sprout. Alternative postfire outcomes for black oaks include 

no damage, resprouting from their crowns, or being entirely killed. 
Pacific Southwest Research Station

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This publication is available online at www.fs.fed.us/psw/.