Quercus agrifolia



SPECIES: Quercus agrifolia

photo courtesy of Charles Webber, California Academy of Sciences

 

AUTHORSHIP AND CITATION:
Steinberg, Peter D. 2002. Quercus agrifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].

FEIS ABBREVIATION:
QUEAGR

SYNONYMS:
None

NRCS PLANT CODE [154]:
QUAG

COMMON NAMES:
coast live oak
encina
California live oak

TAXONOMY:
The currently accepted scientific name of coast live oak is Quercus agrifolia Nee. (Fagaceae). Two varieties of coast live oak are recognized [70,76]:

Quercus agrifolia Nee var. agrifolia
Quercus agrifolia Nee var. oxyadenia (Torr.) J.T. Howell

Coast live oak is classified in the red oak subgenus (Erythrobalanus) [7,65,67,92,114]. Several hybrids between coast live oak and other  red oak species have been documented [20,70]. Quercus a. var. oxyadenia California black oak (Q. kelloggii) hybrids, known as Q. ganderi C.B. Wolf, occur in San Diego County [20]. California black oak also hybridizes with the typical variety of coast live oak (Q. a. var. agrifolia). This cross, known as Q. chasei McMinn, is found in Monterey and Santa Cruz counties. Hybrids between interior live oak (Q. wislizenii) and the typical variety of coast live oak are known in many areas in northern California. Coast live oak also hybridizes with Nuttall's scrub oak (Q. dumosa) and Shreve oak (Q. parvula var. shrevii) [44,114]. All these oak species show evidence of introgression with one another [44].

There is some uncertainty about the degree of introgression between interior and coast live oaks. Brophy and Parnell [20] note that limited hybridization has occurred but the 2 species remain relatively distinct. Dodd and others [45] estimate that in the northern part of interior and coast live oaks' ranges, where hybridization is greatest, up to 60% of coast live oak may show signs of introgression.

LIFE FORM:
Tree

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
None

DISTRIBUTION AND OCCURRENCE

SPECIES: Quercus agrifolia
GENERAL DISTRIBUTION:
Coast live oak occurs in California and northern Mexico. It is distributed along the Coast, Transverse, Peninsular, and Sierra de Juarez ranges from Mendocino County, California, south to Canada El Piquillo, Baja California [8,67,106,114,122]. Limited inland populations occur along watercourses in the Central Valley [72,77]. Coast live oak is also found on the Channel Islands of Santa Rosa and Santa Cruz [34,125]. Quercus agrifolia var. oxyadenia occurs in interior cismontane regions of Baja California and in Riverside, Orange and San Diego counties of California [114]. CalFlora provides a distributional map of coast live oak and its varieties.

ECOSYSTEMS [58]:
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES27 Redwood
FRES28 Western hardwoods
FRES34 Chaparral-mountain shrub
FRES42 Annual grasslands

STATES:
CA Mexico

BLM PHYSIOGRAPHIC REGIONS [14]:
3 Southern Pacific Border

KUCHLER [86] PLANT ASSOCIATIONS:
K005 Mixed conifer forest
K006 Redwood forest
K009 Pine-cypress forest
K010 Ponderosa shrub forest
K012 Douglas-fir forest
K029 California mixed evergreen forest
K030 California oakwoods
K033 Chaparral
K035 Coastal sagebrush
K036 Mosaic of K030 and K035
K047 Fescue-oatgrass
K048 California steppe

SAF COVER TYPES [52]:
221 Red alder
232 Redwood
234 Douglas-fir-tanoak-Pacific madrone
244 Pacific ponderosa pine-Douglas-fir
245 Pacific ponderosa pine
246 California black oak
248 Knobcone pine
249 Canyon live oak
250 Blue oak-foothills pine
255 California coast live oak

SRM (RANGELAND) COVER TYPES [147]:
109 Ponderosa pine shrubland
110 Ponderosa pine-grassland
201 Blue oak woodland
202 Coast live oak woodland
203 Riparian woodland
204 North coastal shrub
205 Coastal sage shrub
206 Chamise chaparral
207 Scrub oak mixed chaparral
208 Ceanothus mixed chaparral
214 Coastal prairie
215 Valley grassland
405 Black sagebrush
613 Fescue grassland

HABITAT TYPES AND PLANT COMMUNITIES:
Coast live oak is a conspicuous tree in lower-elevation oak woodlands of California [65]. California oak woodlands occupy about 10 million acres (4 million ha) and may be divided into 3 types: northern, southern, and foothill woodlands. The northern oak woodland type is dominated by blue oak (Q. douglasii), Oregon white oak (Q. garryana), and interior live oak [11]. Southern oak woodlands are dominated by coast live oak with or without Engelmann oak (Q. engelmannii), sometimes in association with bigcone Douglas-fir (Pseudotsuga macrocarpa) [11,33,35,64]. Southern California coast live oak woodland also has a chaparral phase, dominated by shrubby coast live oak. Canyon live oak (Q. chrysolepis), gray pine (Pinus sabiana), California bay (Umbellularia californica), wedgeleaf ceanothus (Ceanothus cuneatus), chamise (Adenostoma fasciculatum), and manzanita (Arctostaphylos spp.) are common associates of coast live oak in chaparral habitats [73,146]. The foothill oak woodland is dominated by blue oak and interior live oak, sometimes with gray pine [11].

Coast live oak frequently codominates in southern California walnut (Juglans californica) woodland [64]. It is also a frequent dominant or codominant in mixed evergreen forests where it grows with valley oak (Q. lobata), tanoak (Lithocarpus densiflorus), Pacific madrone (Arbutus menziesii), blue oak, California black oak, interior live oak, California bay, gray pine, and coast Douglas-fir (Pseudotsuga menziesii var. menziesii) [33,64,142]. In riparian areas, coast live oak occurs with western sycamore (Platanus racemosa), white alder (Alnus rhombifolia), Fremont cottonwood (Populus fremontii), and red alder (A. rubra) [10].

Classification systems describing communities where coast live oak is dominant are available [2,3,33,53,125,142].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Quercus agrifolia

 

GENERAL BOTANICAL CHARACTERISTICS:
Coast live oak is a native, drought-resistant, evergreen tree, ranging in height from 19 to 82 feet (6-25 m) and in diameter from 1 to 4 feet (0.3-1.2 m) [70,129]. The bark of young trees is smooth. With age, it develops deep furrows, ridges, and a thickness of about 8 to 9% of bole or branch diameter [70,129]. The inner bark and cork layers are thick [43,129]. Open-grown crowns are broad and dense, with foliage often reaching the ground [70]. In open areas trunks are usually 4 to 8 feet (1.2-2.8 m) tall; at this height, primary branches originate and grow horizontally. Trees in dense stands generally have irregular crowns and few lower branches. In closed stands trunks may be branchless up to 20 feet (6.2 m) high, where several branches extend diagonally upward [129]. Coast live oak stands are typically from 40 to 110 years old. Individual trees may live over 250 years [43,129].

The root system consists of a deep taproot that is usually nonfunctional in large trees [35,128]. Several deep main roots may tap groundwater if present within approximately 36 feet (11 m) of the soil surface [31,35,128]. Coast live oak develops extensive horizontal root branches and surface-feeding roots [18,35,128]. Tree roots in southwestern California are associated with mycorrhizae that aid in water uptake during the dry season. A network consisting of roots from 3 coast live oak trees and their and associated mycorrhizae covered a 50- 13-foot (15- 4-m) area of the soil profile that reached through weathered granite through to bedrock. Roots in clay soils were not infected with mycorrhizae [18].

RAUNKIAER [132] LIFE FORM:
Phanerophyte

REGENERATION PROCESSES:
Breeding system: Oaks (Quercus spp.) are monoecious with staminate flowers in naked aments and solitary or clustered pistillate flowers [145]. Genetic studies using biochemical and molecular markers show that coast live oak is the least diverse of all California oaks in the subgenus Erythrobalanus. This suggests that coast live oak populations are highly outcrossing, and most genetic diversity is among, rather than within, populations [44].

Pollination: Coast live oaks are wind pollinated [32,44].

Seed production: Coast live oak is the only black oak in California whose acorns develop in 1 year [21,129]. Acorns are 1.0 to 1.5 inches (2.5-3.8 cm) long and 0.4 to 0.6 inch (1.0-1.5 cm) wide [70,129]. Coast live oak California black oak hybrids' acorns mature in 2 years [114]. Information on the age of sexual maturity and age of maximum production of acorns is lacking [32]. 

Acorn production is variable within and among years, but coast live oak is productive relative to other California oak species. Acorn production over a 5-year period was observed for canyon live oak, coast live oak, valley oak, blue oak, and California black oak. Canyon live oak was most productive, and coast live oak was 2nd most productive [32]. Crop failure is frequent, and large crops may occur in consecutive years but no more frequently than expected by chance. In low acorn years, acorns can be absent on up to 90% of trees: this occurred in valley oak, coast live oak, blue oak stands observed in Santa Barbara, San Luis Obispo, and San Benito counties [57]. 

Though crop size is variable, it is often synchronized across oak populations and species. Synchronization is more common among California oak species whose acorns mature in the same year, but it also can occur among species with different acorn maturation times [32,81,83]. Synchronization may be detectable on areas as large as 190 to 390 square miles (500-1000 km2) [81]. Observation of acorn production cycles among canyon live oak, coast live oak, valley oak, blue oak, and California black oak in the Santa Lucia Mountains of central California found no species-specific patterns in acorn production. The 5 oaks were observed over 12 years; at only 2 of 60 date/year combinations (3.3% of sample dates) was there no acorn production from any species. Coast live oak, however, had crop failures (acorns per tree averaging less than 1) 50% of the time [84].

Several nonmutually exclusive hypotheses seek to explain the variability and synchronicity of California oak acorn crop sizes. This may be an adaptation to seed predation as high acorn production years have incomplete seed consumption and allow leftover seed to establish [84]. Synchronicity over large geographic areas may also be because the areas experience similar environmental conditions [83]. One study found that acorn crops for coast live oak and canyon live oak were larger when the previous 1 and 2 years had more rain [82]. Another 5-year study found no correlation between crop size and 14 weather variables [32].

Seed dispersal: Coast live oak retains its acorns longer than other California oaks. Most of its acorns are dropped in fall, but some remain attached to trees until spring. This adaptation reduces seed loss to birds and small mammals such as squirrels, magpies, and scrub jays; in small crop years 100% of the crop may be consumed [32,115]. Though seed predation may be high, it also provides a means of seed dispersal. Seed-caching animals such as the scrub jay and California ground squirrel are important to species survival, as acorns buried by these animals are more likely to develop into seedlings than acorns that have not been cached [28,32]. Scrub jays cache about 5,000 acorns each year over an average home range of about 6.2 acres (2.5 ha), effectively seedling at a rate of 800 acorns per acre (2,000 per ha). Though 95% of cached acorns are found and eaten, remaining acorns have high germination and survival rates because they are typically cached about 0.4 inch (1 cm) below the soil surface [100]. Scrub jays prefer burned sites for acorn caching [115].

Acorn-feeding insects damage coast live oak seed.  Weevil and moth larvae on the Santa Rosa Plateau Ecological Reserve preferred coast live oak acorns to those of Engelmann oak, and researchers concluded that coast live oak acorns provided a more nutritious diet. Although damage inflicted to individual acorns appeared slight, the ability of insect-damaged acorns to germinate and establish is poorly understood. Further studies are required to determine the impact of insect damage to coast live oak establishment [52].

Seed banking: Little information is available regarding coast live oak seed banks, but, because coast live oak acorns have no dormancy requirement, it is reasonable to assume that seed banks are small.

Germination: Coast live oak acorns have no dormancy requirement and germinate 15 to 50 days after falling [25,64,108]. Germination is slower than other California oaks; this limits coast live oak establishment on dry sites [95]. Griffin [64] found 100% viability of acorns collected in the Carmel Valley, while Mirov and Kraebel [108] reported 73% germination of acorns collected in Berkeley. Fifty percent germination after 18 days of 55 to 68 degrees Fahrenheit (13-20 C) controlled climate without stratification has also been reported [96]. Acorns with low moisture content show higher rates of germination and produce taller seedlings than do acorns with high moisture content, but unviable seed is not a serious problem for this species [75,115].

Seedling establishment/growth: Seedling top-growth is slow under natural conditions, with early development concentrated on rapid, early root growth [96]. Lateral root growth of 9.8 inches (25 cm) in 6 months has been observed under moist controlled conditions [27]. In the field, coast live oak taproot elongation of approximately 28 inches (70 cm) in 2 months has been noted [39]. Under ideal nursery conditions, coast live oak seedlings have attained heights of 5 to 8 feet (15.2-24.4 m) in 2 years [144].

Acorn predation strongly limits establishment in small crop years, but seedling survival rates are also low [115]. At sites in the Central and South Coast ranges, current sapling-to-tree ratios are 1:3 or lower [112]. Common causes of mortality include herbivory and livestock trampling and inadequate moisture [32,115,123]. Mortality results from aboveground or root herbivory by domestic livestock, black-tailed deer, pocket gophers, grasshoppers, cutworms, and feral pigs [32,48,64,123,149,153]. 

Coast live oak appears to be more susceptible to browsing damage than other California oaks while in the seedling stage, but shows better ability to sprout after herbivory damage incurred in the sapling stage [64,111,127]. Coast live oak seedlings are particularly vulnerable to browsing damage because some insect and mammalian browsers prefer coast live oak seedlings over seedlings of associated oaks [48].

Coast live oak is more susceptible to drought than other California oaks. In experimental soils wherein subsurface moisture was controlled and dried while leaving a moist surface horizon, coast live oak did not increase lateral root production when the vertical root tip died in dry soil. In contrast, blue oak and valley oak both responded to the treatment with increased lateral growth; these morphological differences are consistent with the species' different ranges [27]. 

Because of drought and herbivory's deleterious effects on seedling survival, recruitment is best among germinants growing in shade, where herbivory protection and water availability are higher [157]. Recruitment among rock outcrops is relatively common as this offers some protection from large browsers [66]. Common nurse plants are California heathgoldenrod (Ericameria ericoides), chamise, coyote bush (Baccharis pilularis), purple sage (Salvia leucophylla), orange bush monkeyflower (Diplacus aurantiacus ssp. aurantiacus), chamisso bush lupine (Lupinus chamissonis), California sagebrush (Artemisia californica) and poison-oak (Toxicodendron diversilobum) [28,118]. Eighty percent of seedlings found on sites in the Santa Ynez Valley of Santa Barbara County were growing under purple sage or California sagebrush. Another 15% grew under mature coast live oak. Mortality due to herbivory was considerably higher among seedlings under parent trees [28]. Establishment is generally greater in coastal sage scrub than in other types of chaparral [30]. A study of coast live oak seedling establishment, with and without nurse shrubs and cages for protection, found that acorns planted in the open had higher germination but lower 1-year survival. Between 1 and 2 years of age, nurse shrubs had no effect on seedling survival, but cages continued to have a positive effect by reducing black-tailed deer browsing [118]. Another study of coast live oak establishment at 2 sites in central California showed strong association of seedlings with shrubs. Shrub cover was 30%,  but 80% of coast live oak seedlings were under canopies. In 2 years of monitored growth, survivorship was 0% in open areas and 31% under shrubs. Seedlings in the open were grazed more, but drought also increased stress. Mortality related to moisture and temperature stress was 17% under shrubs and 63% in the open. The authors noted that differential dispersal might also increase recruitment under shrubs [28]. 

Asexual regeneration: Coast live oak sprouts from the root crown and/or trunk following cutting or burning [37,43,126,128,130].

SITE CHARACTERISTICS:
Coast live oak occurs in a mediterranean climate characterized by mild, wet winters and hot, dry summers. Climatic extremes are modified near the coast, where trees receive more precipitation than inland populations [43]. Trees generally occur on mesic sites such as north slopes, alluvial terraces, canyon bottoms, or upper streambanks [23,46,101]. Coast live oak's preference for mesic sites is most pronounced in the southern part of its range [106]. Coast live oak may grow where it can access groundwater, but most individuals have extensive shallow root networks [79].

Low-elevation coastal populations of coast live oak generally grow in loam, while higher-elevation coastal populations are associated with shaley clay-loam soil. Inland populations are found on sandy soil, while those in southern California islands grow in clay or clayey loam [34,43]. Coast live oak tolerates serpentine soils [53]. Coast live oak is often associated with depositional environments, deeper soils, and higher organic matter. Coast live oak occurs on soils ranging from silts and clays to weathered granite [18,47].

Elevations of coast live oak populations range from sea level to 3,000 feet (914 m) in central and northern California and from sea level to 5,000 feet (1,524 m) in southern California [70,79,129]. Coast live oak occurs at slightly higher elevations in Baja California. On the coastal side of the Sierra Juarez coast live oak grows below 4,260 (1,300 m), and on western slopes of Sierra San Pedro Martir, elevations range from 3,930 to 5,580 (1,200-1,700 m) [106].

SUCCESSIONAL STATUS:
Ecologists refer to changes in the grassland, chaparral, and oak woodland mosaic of California as "non-directional fluctuations" rather than succession [12]. Coast live oak may be considered seral or climax depending on habitat, but it is tolerant of shade throughout its life [129]. Where their ranges overlap, deciduous trees such as blue oak and California black oak are seral to coast live oak [13]. However, because deer and cattle prefer coast live oak, it is gradually replaced by California bay in some areas of coastal northern California where the 2 species codominate [99]. In mixed evergreen forest that is burned or logged, a coast live oak phase is seral to the climax evergreen deciduous and conifer forest. However, on steep slopes or poor sites within this habitat type, coast live oak represents a topographic or edaphic climax [7]. Succession on coastal dunes at Asilomar State Park on the Monterey Peninsula includes an early stage of beach wormwood (Artemisia pycnocephala) and/or California heathgoldenrod on slopes; the next sere includes dense shrub (coyote bush, seaside woollysunflower (Eriophyllum stoechadifolium), California coffeeberry (Rhamnus californica), poison-oak, and Pacific dewberry (Rubus vitifolius)) development with coast live oak and/or gray pine in the overstory. If fire frequency and deer numbers are low, coast live oak replaces gray pine [9]. In the San Francisco Bay area light browsing allows coyote bush to invade grasslands, and coyote bush subsequently facilitates coast live oak woodland development [139]. In the absence of disturbance, coyote bush brushland almost always gives way to coast live oak and California bay, as coyote bush seedlings do not develop beneath their own canopies [161].

Recruitment of most oaks in California has declined, with some species not regenerating rapidly enough to maintain current density. There is much debate about the causes of change and size of decline, but it appears that recruitment patterns are dependent on both species and locality  [11]. Overall coast live oak and interior live oak are recruiting at higher rates than the deciduous oaks of the region; this may be because of pocket gophers' preference for the roots of deciduous oaks over those of live oaks [66]. Generally, coast live oak woodlands with shrub understories have stable population structures, and more xeric woodlands with herbaceous understories are likely to be recruitment limited [30].

SEASONAL DEVELOPMENT:
Coast live oak defers growth until winter as a drought evasion strategy [60,79]. Approximately 90% of stem elongation occurs in March, but some elongation occurs from December to April, depending on elevation. Prior to stem growth, total non-structural carbohydrate concentration of branches peaks at 5.5% [109]. Leaves emerge from February to April; and old leaves are shed at the same time [109,114,129]. Transpiration and photosynthesis are highest in spring but leaves are functioning year-round [109].

Coast live oak begins flower and fruit production during stem elongation in the spring [109]. Flowering of coast live oak is triggered by warm temperature. This is in contrast to many other black oaks, which flower in response to photoperiod [156]. Catkins emerge in March and April, and acorns ripen in September and October [108,109,114]. Damaged acorns are aborted in August through October [115]. Fall and early winter storms remove acorns from trees [109]. Seed is dispersed between late fall and early winter. Germination is from late November to January in low elevations and in January and February at higher elevations [96]. Shoots are visible in late winter [28].

FIRE ECOLOGY

SPECIES: Quercus agrifolia
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: Coast live oak is exceptionally fire resistant, more so than other California oak species. Adaptations to fire include evergreen leaves, thick bark, and sprouting ability. Evergreen leaves allow coast live oak to allocate greater amounts of energy to recovery from fire than to replacing the entire crown annually. Evergreens are often better able to conserve nutrients than deciduous species, and are favored in fire-prone environments [102]. Coast live oak bark is the thickest among California oaks; it is mainly composed of live inner bark with little dead outer tissue [128,129,130]. Oaks are more likely to be damaged by fall fire than earlier fires [129]. Because of mortality among small-diameter trees, frequent fire limits coast live oak invasion of grasslands [104].

Coast live oaks sprout from the main trunk and upper crown even after severe burning [37,130]. When trees are top-killed, they sprout from the root crown [43]. Vigorous sprouting is supported by food reserves stored in the extensive root system [35]. Sprouting from the root crown often occurs during the first 2 months after top-kill, but some charred trees do not sprout for 2 to 3 years [122,129]. Scorched trees retaining live vascular cambial tissue sprout from the bole and from scaffold and smaller surviving branches. A few scorched trees may produce root crown as well as epicormic sprouts [37]. Prefire crown volume is generally recovered, or nearly so, in about 8 to 10 postfire years [37,129].

Roots are protected from fire by an outer corky layer and soil. Coast live oak roots generally suffer little direct heat damage except to feeder roots near the soil surface [128]. Acorns on the soil surface are killed by low-severity fire, while animal-buried acorns usually survive moderate-severity fire, sometimes allowing high rates of postfire establishment [42,89]. Severe fire may kill even buried acorns, and complete removal of shrubs by fire may reduce coast live oak seedling survival [28,42,118]. By caching acorns in burned areas burned areas, which are preferred caching sites, scrub jays may facilitate postfire establishment of coast live oak seedlings [115]. 

Fire regimes: Fire frequency largely defines the extent of coastal sage scrub, chaparral, and oak woodland; in these habitats decreasing fire frequency tends to favor the development of coast live oak. Conversely, where coast live oak occurs in mixed evergreen forests (or where coast Douglas-fir is invading oak woodlands), frequent fire favors coast live oak and other seral species. One study of vegetation dynamics in coastal sage scrub, chaparral, and oak woodland near Santa Barbara found that without fire or livestock grazing, coastal sage scrub was replaced by coast live oak woodland at a rate of 0.3% annually. Grassland to coastal sage scrub transition occurred at a rate of 0.69% per year, and oak woodland reverted to grassland at a rate of 0.08% per year. On burned areas without livestock grazing or on unburned sites with livestock grazing, rates of transition of grassland to coastal scrub and coastal scrub to oak woodland were lower and the rate of oak woodland reversion to grassland was higher [29].

Oak woodlands: Analysis of oak pollen (primarily that of coast live oak) records showed that oak woodlands remained stable for up to 4 centuries before major European-American settlement (approximately 1470 to 1870). Coinciding with a decrease in fire frequency between 1870 and 1985, percent oak pollen increased to its highest level in 560 years. The 2-fold increase in oak pollen reflected both increase in oak density and in area occupied [104]. Coast live oak woodlands may experience an increase in shrub cover in the absence of frequent fire or heavy grazing, but there are also reports of oak savannas and adjacent grasslands that, without fire for 50 years, have not had a noticeable increase in shrub cover. Sites without shrub increase are generally south facing and/or on shallow soils [66]. Generally, grass is present in open stands, while closed stands have up to 5 inches (12.7 cm) of oak litter [129]. 

Riparian forests: Coast live oak associates in riparian areas include white alder, California sycamore, and Fremont cottonwood, all of which are sprout after fire. Severe fire was apparently historically rare in these habitats. Currently most fire is accidental and of high severity, causing relatively high rates of top-kill and basal sprouting of coast lie oak and associated tree species [10,42].

Mixed evergreen forests: Where coast live oak occurs in mixed evergreen forests of coast Douglas-fir, tanoak, and Pacific madrone, historic fire return intervals were less than 35 years, with fires generally occurring between late August and November after termination of radial growth [5,24]. In many of these stands fire ceased in the early to mid-1900s, and there has been a concomitant increase in coast Douglas-fir density. Bowcutt [19] states that coast Douglas-fir increase in coast live oak woodlands may indicate that such stands, when coast Douglas-fir was not present, represented "a disclimax created by native people through burning." A study of fire frequency and coast Douglas-fir establishment on Point Reyes Peninsula, where coast live oak occurs with California bay, showed coast Douglas-fir establishment greatly accelerated with cessation of fire in the early 1900s. Fire history of 2 such sites is provided below; only 1 fire scar was observed after 1945 [24]: 

  Period analyzed Number of intervals Mean fire interval (years, with standard deviations) Range of intervals (years) Weibull mean (50% exceedance) probability interval (years) 5% to 95% probability interval (years)
Site 1 1820 to 1905 11 7.7 (5.0) 1 to 17 6.9 1.3 to 17.6
Site 2 1825 to 1918 11 8.5 (5.3) 3 to 18 7.8 2.1 to 16.0

Chaparral and coastal sage scrub: Prior to settlement, chaparral communities in which coast live oak grows in a shrubby, shorter form had mean fire intervals of 10 to 30 years [121]. With land use conversion and urbanization, fire frequency has decreased. In the Santa Monica Mountains, chaparral sites dominated by California sagebrush, California brittlebrush (Encelia californica), coastal buckwheat (Eriogonum cinereum), purple sage, and black sage (Salvia mellifera) (with 11 to 19% cover of coast live oak) had fires in 1903, 1945 and 1978. A similar site with 5.2% coast live oak cover had fire in 1903 and 1978 [93]. In most chaparral and coastal scrub areas, fire regimes are entirely anthropogenic, and with urbanization, steep slopes, containment difficultly, and erosion potential, fire's current use is limited [42,68]. The absence of fire, particularly in coastal sage scrub, has allowed coast live oak to increase in density and area. On sites in the Monterey Bay area with fire exclusion for over 70 years, coast live oak has increased significantly (p<0.001) from 2 to 5.5% between 1976 and 2000. Where coast live oak has developed, understory density and diversity have declined dramatically [155].

Fire intervals: Greenlee and Langenheim [63] described fire regimes of different coast live oak associated communities in the Monterrey Bay area between aboriginal time and present. Their results, presented below, show the remarkable decline in fire frequency in the recent era (1929 was chosen to demarcate the recent fire regime because of restrictions that were put in place against burning). "Probable mean fire interval" refers to estimates of fire intervals that are derived from historical or very limited physical evidence.

Fire regime Vegetation where burning concentrated Vegetation where burning incidental Recorded or calculated mean fire intervals (years) Probable mean fire intervals (years)
Lightning   Prairies   1-15
  Coastal sage   1-15
  Chaparral   10-30
  Oak woodland   10-30
Mixed evergreen     15-30
Redwood forest   135  
Aboriginal (until approximately 1792) Prairies   1-2  
Coastal sage   1-2  
  Chaparral 18-21  
Oak woodland   1-2  
  Mixed evergreen   50-75
  Redwood forest 17-82  
Spanish (1792 to 1848)   Prairies   1-15
  Coastal sage   1-15
Chaparral   19-21  
  Oak woodland   2-30
  Mixed evergreen   50-75
  Redwood forest 82  
European-American (1847 to 1929)   Prairies   20-30
  Coastal sage   20-30
  Chaparral 10-27  
  Oak woodland 50-75  
Mixed evergreen   7-29  
Redwood forest   20-50  
Recent (1929 to present)   Prairies   20-30
  Coastal sage 155  
  Chaparral 155  
  Oak woodland 225  
  Mixed evergreen 215  
  Redwood forest 130  

Fire regimes for plant communities and ecosystems in which coast live oak occurs are presented below. More information regarding fire regimes and fire ecology of these communities can be found in the 'Fire Ecology and Adaptations' section of the FEIS species summary for the plant community or ecosystem dominants below.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
California chaparral Adenostoma and/or Arctostaphylos spp. < 35 to < 100 
coastal sagebrush Artemisia californica < 35 to < 100 
California steppe Festuca-Danthonia spp. < 35 [121]
Pacific ponderosa pine* Pinus ponderosa var. ponderosa 1-47 [159]
coastal Douglas-fir* Pseudotsuga menziesii var. menziesii 40-240 [5,110,135]
California mixed evergreen P. m. var. m.-Lithocarpus densiflorus-Arbutus menziesii < 35 
California oakwoods Quercus spp. < 35 
coast live oak Q. agrifolia 2-75 [63]
canyon live oak Q. chrysolepis <35 to 200
blue oak-foothills pine Q. douglasii-Pinus sabiniana <35 
Oregon white oak Q. garryana < 35 [5]
California black oak Q. kelloggii 5-30 [121
interior live oak Q. wislizenii < 35 [5]
*fire return interval varies widely; trends in variation are noted in the species summary

POSTFIRE REGENERATION STRATEGY [148]:
Tree with adventitious bud/root crown
Crown residual colonizer (on-site, initial community)
Initial off-site colonizer (off-site, initial community)
Secondary colonizer (on-site or off-site seed sources)

FIRE EFFECTS

SPECIES: Quercus agrifolia
IMMEDIATE FIRE EFFECT ON PLANT:
Coast live oak seedlings and saplings less than 3 inches (7.6 cm) in diameter may be top-killed by low- to moderate-severity fire, and severe fire kills trees of this size [46,122,128,129,151]. Because of vascular cambium protection, mature trees have high fire survival rates, even with crown fire. Heavily charred bark has a checkered appearance and frequently exfoliates, but damage typically extends only 0.5 to 0.8 inch (1.3-1.9 cm) into the bark. If cambium death occurs in large trees it is typically in small scars at the base [129]. Trees greater than 6 to 8 inches (15.2 to 20.3 cm) in diameter resist top-kill [122,129]. Saplings 2 to 6 inches (5.1-152 cm) in diameter are top-killed by severe fire. An "extremely hot" crown fire on the San Bernardino National Forest caused only 4% coast live oak mortality. Ninety percent of the oaks less than 3 inches (7.6 cm) in diameter were top-killed, and 2% were killed. Of trees greater than 6 inches (15.2 cm) in diameter, the trunks and crowns of all but 5% survived the fire without top-kill (the 5% includes 3% that were top-killed and 2% killed) [128].

For small-diameter trees, mortality as a result of low- to moderate-severity fire depends on tree height and location with respect to other tree crowns. Fire mortality of Engelmann oak and coast live oak (in a woodland with herbaceous and coastal sage scrub species in the understory) was studied for trees less than 3.9 inches (10 cm) in diameter: of 1,214 small trees surveyed, 531 survived 5 years after fire. Mortality was similar for each species: 56% for coast live oak and 59% for Engelmann oak. Mortality was 50% among trees 11.8 to 15.7 inches (30-40 cm) tall, 30% among those 29.3 to 78.7 inches (1-2 m) tall, and 10% among trees taller than 78.7 inches (2 m). In gaps between tree canopies mortality was significantly lower (p<0.0001). Coast live oak mortality was 40% in gaps and 59% under canopies [89] .Generally mortality of coast live oak is greater when there is a considerable shrub understory or when trees are adjacent to chaparral. Coast live oak in the Santa Monica Mountains was killed in large numbers when a hot chaparral fire burned into the oak woodland [141].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
No entry.

PLANT RESPONSE TO FIRE:
Coast live oak sprouts from the bole, branches, and /or root crown after fire damage. Mature trees with live branches generally sprout from the branches. Trees with damaged branches sprout from the bole, which is better protected from fire by its thicker bark [37,130]. Seedlings and saplings are generally top-killed, and sprout from the root crown after fire [43,151]. Coast live oak recovers rapidly from moderate-severity fire. Severely burned crowns, trunks, and root crowns may require several years to sprout. If sprouting occurs within several postfire months, basal sprouts can be 2 to 3 feet (0.6-0.91 m) tall in 2 years, and crown density can be 80 to 100% of prefire levels within 10 years [88,130]. The most common fire damage to the trunk is a basal wound resulting in potential cambium death. Wounds less than a few inches in size may eventually heal with no accompanying heart rot, but larger wounds are susceptible to fungal and bacterial pathogens and insect infestation [138].

Light-severity surface fire has little effect on mature coast live oak. An October prescribed fire in northern San Luis Obispo-southern Monterey counties varied from light to moderate severity. The fire had no effect on postfire coast live oak canopy coverage, measured at postfire year 1. Six percent of the canopy was singed. No mature trees died, and the number of coast live oak snags was unchanged [151,158].

Saplings and seedlings generally recover quickly from light- to moderate-severity fire. For the fall prescribed fire in southern Monterey and northern San Luis Obispo counties, 22 percent of coast live oak saplings were killed. Survival was best for saplings with light fuel loads within 3.3 feet (1 m) of their stems. The remaining 78% of saplings were top-killed and had root crown sprouts by the next fall. Average number of sprouts was 6.4 (+1.2 SE) per sapling. Average length of the longest sprout was 24.1 inches (61.2 cm) (+4.1 inches (10 cm) SE) [151].

Recovery of coast live oak following severe fire on the San Bernardino National Forest showed patterns typical of the species. Mortality and top-kill rates were highest among the smallest size classes. Basal sprouting was common in smaller size classes and crown sprouting (or crown sprouting in addition to basal sprouting) was more common among larger size classes. Sprouting and mortality rates were observed after 5 years on 0.64-acre (0.25 ha) transects; rates are summarized below by size class [128]:

Tree diameter (inches) Number of charred trees Dead (%) Basal sprouts only (%) Basal and Crown sprouts (%) Crown sprouts only (%)
0-3 90 2 88 1 9
3-6 54 11 26 28 35
6-12 65 1 5 52 42
12-18 24 0 0 62 38
18+ 22 4 5 50 41

Coast live oak experiences severe fire in riparian habitats. After the high-severity Wheeler Fire near Ojai, California, coast live oak, white alder, and California sycamore all sprouted. Most burnt tree trunks remained after fire until storms the following winter, when some were uprooted. Six of 19 coast live oak boles blew down in winter. Sprouting rates were 7% for white alder, 83% for California sycamore, and 70% for coast live oak. Basal sprouting of coast live oak was positively correlated (p<0.001) with tree size [42].

Regeneration from acorns following fire is variable, depending on fire severity and site characteristics. Acorn survival is high in low- to moderate-severity fire, particularly when acorns have been cached by animals. Acorn survival in severe fire is low, and postfire seedling recruitment is slower. Following a severe fire in Ventura County, burned sites supported no coast live oak germinants the following spring, while adjacent unburned areas produced new seedlings [42]. Fire in an Engelmann oak/coast live oak stand in southern California had positive effects on coast live oak seedling establishment. In the 2 years preceding fire there was no establishment; in 5 postfire years 1,118 oak seedlings established, of which 1,025 were coast live oak. Establishment was greatest under the outer edges of canopies, particularly for coast live oak [89].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
The 1993 Topanga Fire, driven by November Santa Ana Winds, damaged hundreds of coast live oaks in Los Angeles County. Recovery of 90 trees was evaluated through postfire year 8. Thirty trees were randomly selected from a valley riparian site, 30 from a ridgeline site, and 30 from an open riparian area. All 3 sites were surrounded by mixed chaparral and annual grassland. Immediate fire effects were [37]:

Valley riparian site: The site last burned in 1943. The riparian coast live oak-California sycamore community had a closed canopy prior to the wildfire. Eleven coast live oaks were burned down to trunks or scaffold branches; the remaining 29 trees were scorched and lost all their leaves.

Ridge site: The ridgeline had a history of frequent fire. Prescribed burned in 1988, the coast live oak woodland had experienced previous wildfires in 1970, 1942, and 1938. Prefire fuel loads were not as abundant as the other 2 sites. Four mature trees and 1 sapling were top-killed by the fire. The remaining 25 study trees  received some scorch damage but retained some green leaves.

Open riparian site: This site last burned in 1943, from the same fire affecting the valley riparian site. A heavy fuel load of dead and live chaparral vegetation surrounded the site. Within the coast live oak community, the canopy was closed and understory vegetation was sparse. Five trees were top-killed; the remaining 25 trees were scorched but retained some green leaves.

Recovery: Overall recovery of coast live oak was excellent, with 96% survivorship at postfire year 8. Among the 21 most severely burned trees, only 4 trees died. Those 4 were all close to the fire front. Surviving trees showed rapid growth in trunk diameter. Trees from all but the valley riparian site, which had colder temperatures and less shade than the other 2 sites due to steep canyon walls, recovered 80% of their prefire canopy by postfire year 8. Few trees sprouted from the root crown; nearly all postfire growth was initially from epicormic sprouts. At postfire year 8, growth was mostly from terminal branches; epicormic growth was minimal and there were no root crown sprouts. Size distribution and vigor of surviving coast live oaks are shown below [37].

Number of trees in DBH1 class (cm)

Site <25 25-50 51-75  >75 25-50 51-75 76-100 101-125 >125
1994
valley riparian 2 11 3 1 1 2 4 3 3
ridgeline 6 11 2 0 0 3 2 2 3
open riparian 3 1 8 2 2 1 3 2 8
Total# trees 11 23 13 3 3 6 9 7 14
2001
valley riparian 3 7 5 1 1 1 2 4 4
ridgeline 3 11 2 4 2 2 3 2 2
open riparian 2 3 3 5 1 0 1 4 9
Total# trees 8 21 10 10 4 3 6 10 15

Vigor of trees in DBH2 class (cm)

Site <25 25-50 51-75  >75 25-50 51-75 76-100 101-125 >125
1996
valley riparian 3.5 2.5 3 4 3 3 3 3 3
ridgeline 3.5 3 4 0 4 3.5 3 3 3
open riparian 3 3 4 3 3 2 3 3 3.5
2001
valley riparian 4 4 3.5 4.5 5 3 3.5 3.5 4.5
ridgeline 4.5 4 4.5 0 4.5 4 4 4 4
open riparian 3 4 4 4 4 0 3 4.5 4.5
1Some trees had multiple stems. Values are sums for all stems.
2Values based on a rating scale of 1-5, where 1= dead and 5=excellent

The Research Project Summary Response of vegetation to prescribed burning in a Jeffrey pine-California black oak woodland and a deergrass meadow at Cuyamaca State Park, California, provides information on prescribed fire and postfire responses of many plant community species including coast live oak.

FIRE MANAGEMENT CONSIDERATIONS:
Use of fire is recommended for managing coast live oak woodlands [46,94,128,129]. The outstanding fire resistance of this species ensures that few large trees are killed by low to moderate-severity prescribed fire. Fire reduces coast and big-cone Douglas-fir where they invade or increase in density in oak woodland [73]. Oak regeneration is favored because scrub jays prefer burns as acorn-caching sites [115]. Large trees do not always sprout during the 1st postfire growing season, and Plumb and Gomez [129] recommend waiting at least 1 growing season, possibly up to 3, if there is a desire to remove large trees with uncertain potential for crown survival. Dougherty and Riggan [46] report that in San Diego County, prescribed fire in coast live oak woodland (with desert ceanothus (Ceanothus greggii) and Nuttall's scrub oak in the understory) achieved litter reduction goals only temporarily. Within 1 to 3 years, they estimate, litter falling from lightly burned parts of the lower canopy would increase surface litter to its former coverage.

Fire exclusion in coastal sage scrub and mesic chaparral communities allows coast live oak to increase in density and reduce understory diversity and abundance [104,155]. Van Dyke and Holl [155] recommend prescribed burning in coastal sage scrub to maintain scrub species and associated herbaceous species and to slow coast live oak expansion. Fire managers in southern California have made fire control lines in coast live oak riparian woodlands when planning broadcast burning of adjacent chaparral and coastal scrub. Control lines are burned to create a fuelbreak [46]. Green [61] offers much information on prescribed fire planning in chaparral.

Flammability of coast live oak and chaparral communities with a coast live oak component is of particular concern because of their high fuel loading and proximity to urban areas. Some fire-excluded chaparral habitats have fuel accumulations of 30 to 40 tons per acre (73 to 97 t/ha) [68]. Fuel modification is of greatest value in the immediate area around development. Recommendations for property protection include: planting trees away from structures, trimming low branches, selecting less flammable landscaping species, using nonflammable construction materials, and, where trees already grow near homes, removing overhanging branches [49,54]. See the Firewise website for specific recommendations. Domestic goat grazing (at a rate of 240 goats/acre (600 goats/ha) for 1 day), in conjunction with prescribed fire, has been used to reduce fuel loading and continuity in dense coast live oak chaparral near housing developments [152].

MANAGEMENT CONSIDERATIONS

SPECIES: Quercus agrifolia
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Coast live oak woodlands are some of the most important habitats to wildlife in California [57]. These communities are preferred habitat for black bear and black-tailed deer. In chaparral areas where shrubs predominate, coast live oak stands with an understory of mixed shrubs and forbs are critical to black-tailed deer [131]. Coast live oak provides browse for black-tailed deer and various rodents and lagomorphs [140]. Feral pigs and pocket gophers eat the roots, and black bear, feral pig, black-tailed deer, rodents, and various upland game and nongame birds consume the acorns heavily [123,140]. Coast live oak litter provides excellent hunting opportunities for insectivorous vagrant and ornate shrews [131]. Cattle eat coast live oak sprouts [124]. Most livestock do not readily browse mature foliage, although domestic goats eat it year-round [62]. Livestock readily consume the acorns [140].

Coast live oak communities support a number of bird species including the federally endangered least Bell's vireo and least tern [87,117]. Acorns of coast live oak are of particular value to acorn feeders, as they are retained on tree for up to 8 months [32]. Acorn-dependent birds include the acorn woodpecker, yellow-billed magpie, and scrub jay [64,88]. Acorns comprise over 50% of diets of the acorn woodpecker and scrub jay in fall and winter [88].

Palatability/nutritional value: Nutritional information on coast live oak foliage is sparse. One study found the protein content of leaves was 4.6% in December. Springtime protein levels were not determined; however, protein content of interior live oak, a closely related species, was 17.6% in May [15]. Coast live oak acorns are 6.26% protein, 16.7% fat, and 54.7% carbohydrate [6]. The palatability of coast live oak foliage has been rated useless for cattle and horses, poor to useless for domestic sheep, and fair to poor for black-tailed deer [140].

Cover value: Western sycamore/coast live oak communities in the South Coast Ranges provide wintering grounds for 32 species of birds, at a density of approximately 251 birds per acre (620/ha) [51]. An estimated 41% of owl territories in southern California are interior and coast live oak/bigcone Douglas-fir forests. Owls are benefited by fire in adjacent chaparral, which increases prey populations [160]. Red-shouldered hawks rely heavily on coast live oak woodlands in southern California but adapt to urban woodlands as well [16].

Several cavity-nesting birds use coast live oak for nesting. In a Santa Barbara County study, western bluebird and ash-throated flycatcher were the most common nest occupants. Bewick's wren, oak titmouse, tree swallow, and violet-green sparrow also use coast live oak cavities [113].

VALUE FOR REHABILITATION OF DISTURBED SITES:
Coast live oak is favored for use in rehabilitation projects throughout its range. It is used in watershed improvement, restoration, and wildlife habitat rehabilitation projects [59,74,124]. 

Artificial regeneration of coast live oak is typically from acorn plantings or transplanting seedlings and saplings.  The Oak Habitat Restoration Project in Walnut Creek has had good success with direct plantings of coast live oak acorns. About 1 in 3 acorns established as seedlings, and survivorship of 1st-year seedlings has been about 60%. Kraetsch [85] provides techniques for artificial regeneration of coast live oak from acorns. In the Sepulveda Wildlife Reserve of Los Angeles County, nursery seedlings were planted for slope stabilization and wildlife habitat improvement. Survival of these seedlings was between 40 and 75% over 3 years [119]. Transplants along the Santa Margarita River in San Diego County have also shown good survival [134]. In eastern Ventura County, coast live oak was planted to help restore the ecotone between saltbush (Atriplex spp.), black sage, and annual grassland communities [120]. Recommendations for nursery propagation of coast live oak are 36 to 41 degrees Fahrenheit (2.5-5 C) acorn storage, germination in wet vermiculite, and planting in tar paper tubes filled with 25% topsoil, 25% organic matter, and 50% sand. Burger and others [26] also recommend transferring seedlings to glass houses after 0.8 inch (2 cm) of aboveground growth. Root fragments seldom host both ecto- and endomycorrhizae; inoculating with both types is generally detrimental [50].

Transplanting older trees has been successful. Planting saplings, rather than seedlings, may be indicated if herbivory pressure is expected to be high. Studies show that while coast live oak mortality from herbivory can be great, saplings are resistant to browsing damage [48,111,127]. Large coast live oak trees slated for removal because of roadway or other construction have been successfully transplanted onto favorable sites using heavy equipment [36]. Dagit and Downer [38] provide information on transplanting mature trees.

Coast live oak is also established by acorn plantings. In the Berkeley Hills, seedlings grown from locally collected acorns had 75% survival in their 1st year. Seedlings were hand-watered twice during summer because of drought [133]. Seedling survival is enhanced by weeding competitors, providing protection against herbivores and acorn predators with above- and belowground wire caging, and providing microsite shading. Acorns are collected from local oaks in late fall. Planting density recommendations and other cultivation methods have been detailed [115,144]. 

OTHER USES:
Coast live oak is used as an urban ornamental tree [26]. Native Americans planted coast live oaks to harvest the acorns as food. Coast live oak may have expanded its range into interior live oak habitats of northern California when coast live oak acorns were planted inland [137].

Wood Products: Coast live oak wood is primarily used for fuel [129] and can be managed for firewood production. Coast live oak is a good candidate for coppice management, as it sprouts vigorously from cut stumps. Few studies have been conducted on coast live oak response to thinning. Coast live oak stands on 4 sites on the central and southern California coastline responded to thinning with significantly greater basal area compared to unthinned stands [126].

Some commercial charcoal is made from coast live oak wood. The wood is hard, heavy, and fine-grained, but it is unsuitable for lumber because it cracks easily and warps badly [129]. 

OTHER MANAGEMENT CONSIDERATIONS:
Mesic coast live oak woodlands with shrub understories have adequate regeneration, while xeric woodlands are likely to have limited recruitment [30]. Thus, northern coastal populations of this tree are generally stable, but the species is in decline in southern California [112,143]. Coast live oak regeneration in xeric sites can be improved by reducing grazing intensity or protecting individual seedlings from livestock [32,64,123,124]. Other threats to this oak include urban and rural development, increased recreational use of oak woodlands, and cutting trees for firewood [129,143]. Some coast live oak woodlands were cleared to increase forage for domestic livestock, but since the early 1970s, urban and residential development have had a much larger impact [1]. Coast live oak populations in Baja California were not as heavily logged as in California, as natural gas and propane were available when human populations in the area expanded. Here heavy grazing around population centers has been the primary anthropogenic limitation on recruitment [107]. Presence of honey-fungus (Armillaria mellea) in the soils of sites now bereft of coast live or other oaks is a good indicator of oak dominance in the past [22].

Diseases: The most serious threat to coast live oak, other red oaks, and related non-oak species in the beech family is sudden oak death disease.  The primary pathogen response for sudden oak death is the fungus-like water mold Phytophora ramorum. Previously described only as a greenhouse pathogen from northern Europe, its origins are uncertain [55,103,136]. Another pathogen, Hypoxylon thouarsianum, is associated with the disease as a secondary fungus. Mature coast live oak are highly susceptible to sudden oak death, which can kill apparently healthy trees within a few weeks to several years. Sudden oak death disease has reached epidemic proportions in California and southwestern Oregon and is particularly virulent on California's central coast, where coast live oak is the dominant red oak species [103,136]. For example, coast live oak on 2 sites in Marin County showed infection rates of 35% in 2000 and 38% in 2001, and 16% in 2002 and 19% in 2001, respectively. Coast live oak mortality at the 2 sites rose from 8 to 15% and 6 to 8% during that time. About 1/3rd of coast live oak on the 2 sites showed evidence of infection.

Etiology of sudden oak death is unclear as of this writing (2002). Bark and ambrosia beetles (Scolytidae) are associated with infected trees and may be vectors for the fungal pathogens [103] Coast live oak on moist sites may be more vulnerable to Phytophora ramorum infection than trees on drier sites [150]. Ability of infected trees to overcome the disease is unknown [103]. Besides species in the oak family, a variety native woody plant species serve as alternate hosts for Phytophora ramorum (i.e., Rhododenron spp., huckleberries (Vaccinium spp.), and manzanitas). Although currently isolated to 2 west coast states, seedling inoculum trials suggest that some eastern red oak species may be even more susceptible to sudden oak death than coast live and related California red oaks [136]. Standard fungicide treatments (e.g., metalaxyl, cupper sulfate, and phosphoric acid) have shown positive control of Phytophora ramorum in preliminary tests. Fungicides may provide Phytophora ramorum control in urban settings and protect small groups of wildland trees, but are not likely to be practical in large wildland settings [78]. Information on how sudden oak death is transmitted [40], diagnosed [136], and can be monitored [78]  is available.

Coast and interior live oaks are susceptible to oak drippy-nut disease, which develops after wasps, acorn weevils, or other acorn feeders puncture acorns and allow the bacteria Erwinia quercina to develop within [17]. Diplodia quercina, a fungal pathogen that grows hyphae through wounds, may cause dieback of large branches of coast live oak, valley oak, and California black oak. "Twigblight" in coast and interior live oaks is caused by the fungal pathogens Cryptocline cinerescens and Discula quercina; dieback may range from a few twigs to the entire crown [69]. Epidemics of oak wilt disease, caused by the pathogen Ceratocystis fagacearum, have occurred among Texas oaks in urban forests and in live oak (Q. virginiana) savannas; coast and interior live oaks are more resistant [4]. Watering during summer may cause root rot as a result of Armillaria mellea infection or crown rot as a result of Phytophthora spp. infection [69]. 

Insects: The California oakworm (Phryganidia californica) feeds on older leaves in early summer and leaves of all ages later in summer [98]. This moth occurs at low density in most years but periodically, in approximately 5- to 7-year intervals, increases to outbreak levels that defoliate coast live oak [71]. The California oakworm may feed on coast live oak year-round, as it has summer and winter generations that feed without diapause. Outbreak frequency may be related to climate: cold winters cause mortality [105]. Acorn feeders include the acorn weevil (Curculio occidentalis) and to a lesser extent, the filbertworm (Melissopus latiferreanus). One survey in northern California found that 38% of coast live oak acorns had some boring and larval infestation, 70% of which were acorn weevils and 30% were filbertworms. Infestation was greater on cooler, more shaded aspects of trees [90].

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