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VOLUME II - SECTION C HABITAT ACCOUNTS VEGETATION ASSOCIATIONS

    Alluvial Scrub

    VEGETATION ASSOCIATION:

    ALLUVIAL SCRUB

    MAPPED SUBASSOCIATIONS

    Alluvial Scrub, Disturbed Alluvial Scrub

    DATA CHARACTERIZATION

    All of the large patches of well-developed alluvial scrub referenced in the published literature are within the MSHCP database. Mapping errors for this vegetation should be low because its distinctive structure is interpretable readily and accurately from aerial photographs. Additional studies regarding the effects of reduced flooding intensity within the stands within larger more urbanized floodplains are necessary to determine whether adaptive management techniques are necessary and sufficient to sustain alluvial scrub in its present or past form.

    BIOGEOGRAPHY

    Large stands of well developed Riversidian alluvial scrub occur from inland Los Angeles to San Bernardino and Riverside counties. Examples of these occur in Big Tujunga Wash, San Gabriel River, San Antonio Creek, Cucamonga Creek, San Sevaine/Etiwanda/Day Creeks, Cajon/Lytle Creeks, and portions of the San Antonio, Santa Ana, and San Jacinto Rivers (Hanes et. al. 1989). Less well developed stands of alluvial scrub supporting only the early successional forms of the vegetation occur along smaller drainages throughout Western Riverside County to northern San Diego County along isolated portions of the Santa Margarita River. A different association of alluvial scrub indicated by broom baccharis (Baccharis sarothroides) is distributed in portions of San Diego County and northern Baja California (Smith 1996).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Alluvial scrub occurs throughout many drainages in the western Riverside Plan Area and comprises approximately 0.6 percent (8,216 acres) of the Plan Area. Large acreages of the vegetation occur on the Santa Ana River near Lake Evans in the City of Riverside, along the San Gorgonio River and tributaries near Banning, on the San Jacinto River from the National Forest to the Soboba Indian Reservation, near Temecula along Temecula Creek, the Aguanga area, Bautista Creek south of Hemet, and near Murrieta and Glen Ivy in the Temescal Valley.

    VEGETATION CHARACTERISTICS

    Alluvial scrub is a Mediterranean shrubland type that occurs in washes and on gently sloping alluvial fans. Alluvial scrub is made up predominantly of drought-deciduous soft-leaved shrubs, but with significant cover of larger perennial species typically found in chaparral (Kirkpatrick and Hutchinson 1977). This vegetation type is distinctive because of the co-occurrence of evergreen shrubs, drought-deciduous shrubs, riparian species, and upland annual species in close proximity to one another (Hanes et al. 1989). Because alluvial scrub is physiognomically intermediate between chaparral and coastal sage scrub, it shares many of the same species. The only dominant species that has a strong fidelity to alluvial scrub is scalebroom (Lepidospartum squamatum) (Smith 1980). Scalebroom generally is regarded as an indicator of Riversidian alluvial scrub (Smith 1980; Hanes et al. 1989).

    Perennial species diversity in alluvial scrub has been measured along different aged stands in the San Gabriel River, revealing higher diversity of perennial species in more mature stands on terraces above the immediate flood channel (Smith 1980). The annual species component of species diversity of scalebroom-dominated alluvial scrub has not been quantitatively studied, but a similar vegetation type in San Diego County (Diegan alluvial scrub vegetation) revealed significantly higher native species richness and diversity than in neighboring disturbed and undisturbed Diegan coastal sage scrub (Smith 1996). Higher diversity in Diegan alluvial scrub was related to a greater diversity of microhabitats that supported greater numbers of evergreen shrub and annual herbaceous species.

    SPECIES COMPOSITION

    In addition to scalebroom, alluvial scrub typically is composed of white sage (Salvia apiana), redberry (Rhamnus crocea), flat-top buckwheat (Eriogonum fasciculatum), our lord’s candle (Yucca whipplei), California croton (Croton californicus), cholla (Opuntia spp.), tarragon (Artemisia dracunculus), yerba santa (Eriodictyon spp.), mule fat (Baccharis salicifolia), and mountain-mahogany (Cercocarpus betuloides) (Hanes et al. 1989; Smith 1980). Annual species composition has not been studied but is probably similar to that found in understories of neighboring shrubland vegetation. Two sensitive annual species are endemic to alluvial scrub vegetation in the Plan Area: slender-horned spineflower (Dodecahema leptocerus), and Santa Ana River woolly-star (Eriastrum densifolium ssp. sanctorum).

    PHYSICAL ENVIRONMENT

    Alluvial scrub is found on recent or actively alluviating surfaces that experience infrequent but severe flood events (Hanes et. al. 1989). Typical settings include drainages and outwash fans on the coastal side of the southern California peninsular range. Alluvial scrub typically is found on coarse-particled Riverwash soils near the flood channels or in areas that are frequently inundated, and upon deeper and more fine textured Soboba soils at higher elevations on floodplain terraces (Smith 1980; Hanes et. al. 1989). Soils supporting alluvial scrub drain rapidly, have slow runoff, and contain low amounts of organic matter. These drainages typically do not support extensive hydrophytic (i.e., wetland) vegetation because of the scarcity of surface water for much of the year.

    ECOSYSTEM PROCESSES

    The spatial distribution and species composition of alluvial scrub largely is determined by the dynamic geomorphic processes within floodplains and upon outwashing alluvial fans. Several different seral stages of alluvial scrub have been identified based on flooding frequency and distance from the floodplain channel (Smith 1980; Hanes et. al. 1989). Floodplain landforms that flood less frequently support a more physiognomically diverse array of plants, while areas flooded more frequently support early successional perennial and annual species (Kirkpatrick and Hutchinson 1980; Smith1980; Hanes et. al. 1989). Smith (1980) estimated that three aged stands were present at roughly 10, 20, 30, and 50 years old along the San Gabriel River. The 10 and 30 year old stands had developed since large-scale flooding in 1969 and 1938 respectively. Since there has not been catastrophic flooding since 1969, the 40-50 aged upper terrace stands presently are approximately 60-70 years old. With increasing urbanization, flood control projects, and associated changes in the frequency and intensity of flooding in many of the watersheds supporting alluvial scrub, it may be necessary to simulate periodic flooding to sustain the age class structure typical of diverse alluvial vegetation.

    The role of fire in the development and maintenance of alluvial scrub vegetation is unknown. Because the species composition of alluvial scrub vegetation largely is made up of fire adapted plants fire may play a role in reducing older aged stands and promoting the growth of younger individuals and annual species similar to upland coastal sage scrub and chaparral systems. Fire is probably less important and less likely in younger-aged stands that are frequently flooded because of the lack of fuel load and open nature of the vegetation.

    More research is needed to determine the flooding frequency and intensity needed to sustain alluvial scrub vegetation in its present or previous undisturbed forms.

    THREATS

    Threats to alluvial scrub vegetation include changes to the flooding regime caused by flood control, irrigation and upstream residential and commercial development projects; urban edge effects; pollution from residential and industrial runoff; and off-highway vehicle use.

    LITERATURE CITED

    Hanes, T. L., R.D. Friesen, and K. Keane. 1989 Alluvial Scrub Vegetation in Coastal Southern California. USDA Forest Service General Technical Report PSW-110. 1989.

    Kirkpatrick J.B. and C.F. Hutchinson 1977. The Community Composition of Californian Coastal Sage Scrub. Vegetation vol. 35, 1:21-33.

    Kirkpatrick J.B. and C.F. Hutchinson 1980. The Environmental Relationships of Californian Coastal Sage Scrub and some of its component communities and species. Journal of Biogeography 7:23-28.

    Smith, D.S. 1996. Composition and Diversity of Diegan Alluvial Scrub. Master’s Thesis. San Diego State University.

    Smith, R.L. 1980. Alluvial Scrub Vegetation of the San Gabriel River Floodplain, California. Madrono vol.27, 3:126-138.

    Big Sagebrush Scrub

    VEGETATION ASSOCIATION:

    BIG SAGEBRUSH SCRUB

    BIOGEOGRAPHY

    Big sagebrush scrub occupies a large portion of the Great Basin region of Utah, Nevada and adjacent surrounding states (Holland and Kiel 1995). In California, big sagebrush scrub is widely distributed east of the Cascade-Sierra crest within the counties of Modoc, Lassen, Mono and Inyo. The most extensive area of sagebrush scrub occurs on the Modoc Plateau. This community extends as far north as the upper drainages of the Cuyama River of cismontane California and as far south as San Diego County. Within southern California, scattered localities of big sagebrush scrub occur along the fringes of the Mojave and Sonoran deserts and in interior cismontane areas.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Big sagebrush scrub occurs mostly in the southeastern portion of the western Riverside Plan Area. One occurrence of big sagebrush scrub is mapped in the southwestern corner of Riverside County. Big sagebrush scrub occupies approximately 0.9 percent (11,852 acres) of the Plan Area. Large acreages of the vegetation occur north and south of SR-371 within Tule Valley, Culp Valley, Wilson Creek, Cahuilla, the Ramona Indian Reservation and the Community of Anza.

    VEGETATION CHARACTERISTICS

    Big sagebrush scrub is mostly a treeless, soft-woody shrub-dominated community, composed of shrubs approximately 0.5 to 2.0 m tall. The understory is characterized by patches of bare ground and scattered herbs. Big sagebrush scrub often is found in close proximity to woodlands and coniferous forests. This vegetation can occur in the understory of tree-dominated, high elevation communities of pinyon-juniper woodlands, montane mixed coniferous forest and alpine forest (Holland and Kiel 1995). At lower elevations, big sagebrush scrub occurs adjacent to saltbush scrub, blackbush scrub, Joshua tree woodland and creosote bush scrub communities. In areas where soil salinity and alkalinity are high, big sagebrush scrub typically is replaced by saltbush scrub.

    SPECIES COMPOSITION

    The dominant shrub within this Habitat type is big sagebrush (Artemisia tridentata), which sometimes forms pure stands but often occurs with other shrubs (Holland and Kiel 1995). Common associates include bitterbrush (Purshia tridentata), rubber rabbit-bush (Chrysothamus nauseosus), yellow rabbitbrush (C. viscidiflorus) black bush (Coleogyne ramosissima), Mormon-tea (Ephedra viridis), horsebrush (Tetradymia canescens), plateau gooseberry (Ribes velutinum) and hopsage (Grayia spinosa). The herbaceous cover generally is dominated by a mixture of perennial bunch grasses such as ricegrass (Achnatherum hymenoides), needle-and-thread (Stipa comata), letterman’s needlegrass (S. lettermanii), needlegrass (S. occidentalis), needlegrass (S. thurberiana), desert needlegrass (S. speciosa), one-sided bluegrass (Poa secunda), bluebunch wheatgrass (Agropyron spicata), and Great Basin wild rye (Leymus cinereus). Annual grasses and forbs may occur also within big sagebrush scrub. Cheatgrass (Bromus tectorum), an introduced annual grass, has become the dominant herbaceous species in many areas.

    The species composition of Big sagebrush scrub in Riverside County was not reviewed.

    PHYSICAL ENVIRONMENT

    Big sagebrush scrub often is found on alluvial washes within interior regions where frost may occur. This community persists at elevations of 1210 to 2730 m (Holland 1986) but may extend to 3600 m in the White Mountains. Big sagebrush scrub is well-suited to a variety of soils and terrain ranging from rocky, well-drained slopes to fine-textured valley soils with high a water table (Holland 1986). This community typically occurs on south or southwest facing slopes of variable steepness (Gordon and White 1994). The Big sagebrush scrub environment is characterized by cold, harsh winters and dry summers (Barbour and Major 1988).

    LITERATURE CITED

    Barbour, Michael and Jack Major. 1988. Terrestrial Vegetation of California. California Native Plant Society. Davis, California.

    Gordon, H.J. and T.C.White 1994. Ecological Guide to southern California chaparral plant series. Technical Publication R5-ECOL-TP-005. USDA, Forest Service, Pacific Southwest Region, San Francisco California.

    Holland, R.F. 1986. Preliminary Descriptions of the Terrestrial Natural Communities of California. State of California Department of Fish and Game, Nongame-Heritage Program, Sacramento, California.

    Holland, V.L. and David J. Kiel. 1995. California Vegetation. Kendall/Hunt Pub. Dubuque, Iowa.

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Black Oak Forest

    VEGETATION ASSOCIATION:

    BLACK OAK FOREST

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Black oak forest is difficult to map from aerial photography and may not be distinguished easily from mixed evergreen forest, broadleaved upland forests or oak woodlands. The limited occurrence of this Habitat in the Plan Area in the Agua Tibia Mountains may be an artifact of the difficulty in mapping. Black oaks occur elsewhere in the Plan Area, especially in the Santa Ana and San Jacinto Mountains, however, many black oak vegetation stands are considered co-dominant or sub-dominant rather than dominants. The distribution of black oak forest may not be well-understood in Riverside County.

    BIOGEOGRAPHY

    The black oak ranges from just north of Eugene, Oregon south through the Cascade and Klamath Mountains to Marin County. Scattered populations occur from the Santa Lucia Mountains south to the Transverse and Peninsular ranges where black oak becomes more common (McDonald 1990). The species extends into Baja California as well (McDonald 1990).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    An occurrence in the Cahuilla Mountains is described in the literature (Sawyer and Keeler-Wolf 1995). The MSHCP database contains only one occurrence in the Agua Tibia mountains, south of Wild Horse peak, in the southeastern region of the county. A total of nine acres of black oak forest are mapped in the Plan Area.

    VEGETATION CHARACTERISTICS

    The black oak forest Habitat type is dominated by large-sized, deciduous black oaks (Quercus kelloggii). The oak canopy may be continuous, intermittent or savanna-like and trees can reach heights of 40 m (Sawyer and Keeler-Wolf 1995). Shrub cover ranges widely and ground cover is sparse on grass (Sawyer and Keeler-Wolf 1995). The understory density of these areas is limited due to shade, heavy litter, tannins and allelopathic substances (Vogl 1976). Small pure stands of black oak at higher elevations may be the result of removal of conifers through intensive logging or crown fires (Thorne 1977).

    Black oak and ponderosa pine (Pinus ponderosa) appear to be associated closely over much of their Oregon and California ranges and may shelter seedling ponderosa pine, big-cone Douglas-fir (Psuedotsuga macrocarpa) or incense-cedar (Calocedrus decurrens) and may occur with Jeffrey pine and white fir as well (McDonald 1990). Black oak groves typically are associated with other hardwoods such as tanoak (Lithocarpus densiflorus), Pacific madrone (Arbutus menziesii), interior live oak (Quercus wislizenii), coast live oak (Q. agrifolia), Engelmann oak (Q. engelmannii) at lower elevations and Pacific dogwood (Cornus nuttallii), bigleaf maple (Acer macrophyllum), California bay (Umbellularia californica) and canyon live oak (Q. chrysolepis) at higher elevations (McDonald 1990; McDonald 1980; Sawyer and Keeler-Wolf 1995). Shrub associates include manzanitas (Arctostaphylos spp.), California-lilac (Ceanothus spp.), buckthorn (Rhamnus spp.)currant, gooseberry (Ribes sp.) and poison-oak (Toxicodendron diversilobum).

    PHYSICAL ENVIRONMENT

    Over the entire California range, the Habitat occurs between 60 and 2500 m elevation although Bakker (1971) states that this species rarely grows below 300 m (Sawyer and Keeler-Wolf 1995). Although the parent soil material may vary widely, black oak occurs on coarse-textured soils which generally are moderately to excessively drained (Sawyer and Keeler-Wolf 1995; McDonald 1990; McDonald 1980). For this reason, black oaks typically are found on thin soils with rocky slopes, however, stands of black oaks also may occur on various aspects and topographic areas (Sawyer and Keeler-Wolf 1995; McDonald 1990). The typical climatic environment of black oaks is hot, dry summers and cool, moist winters (McDonald 1990). Black oaks require an annual rainfall above 65 cm (Bakker 1971).

    In southern California black oak grows well on deep sandy-loam soils derived from granitic rock (McDonald 1990). Within the Santa Ana and San Jacinto Mountain ranges in particular, black oaks occupy gentle slopes with deep soils often on north-facing slopes (Vogl 1976). These typically moist sites offer winter snows and frozen soils (Vogl 1976). Average rainfall in the southern California mountains varies widely, averaging 91 cm (McDonald 1990). In the San Jacinto Mountains, elevations for black oak reach 2,440 m (McDonald 1990).

    ECOSYSTEM PROCESSES

    Most black oaks in black oak forests show signs of past fires. The response of the trees depends on the type of fire and possibly the vegetation association. Mild crown fires cause recovering black oak trees to produce epicormic trunk sprouts, whereas, passing fires do not damage heavily most black oak trees because of their thick bark and therefore no response is necessary (Vogl 1976). More intense crown fires are very damaging to black oaks and ground fires are often fatal, as evidenced by one age-class stands of black oaks (McDonald 1990). However, in observing succession in a Jeffrey pine-black oak dominated setting McDonald (1980) contends that fires of nearly any intensity can be endured and act to encourage black oak re-sprouting and reproduction. A study of prescribed burns in a Jeffrey pine-black oak dominated area showed that the fires had little effect on trees but caused a significant reduction in shrub cover (Lathrop and Martin 1982). Seedling establishment and shrub resprouts following the burn were not significantly changed (Lathrop and Martin 1982). However, McDonald’s (1990) study on black oak stands following a fire noted that shrubs generally do not form dense cover until a fire occurs, at which time they aggressively compete with black oak sprouts. In any case, black oaks seem well-adapted to fire but the frequency of severe fires may be important to this community. Prior to fire suppression activities, Vogl (1976) believes severe crown fires occurred approximately every 100 to 200 years and even surface fires were probably relatively infrequent.

    THREATS

    Because fire was prehistorically a limited occurrence within this community, the suppression of fires in adjacent fuel-types may result in exceptionally hot fires which may invade the otherwise fire-resistant black oak forest (Vogl 1976). Black oak is also susceptible to fungal infections including Inontus dryophilus and Laetiporus sulphureus (McDonald 1990). Air pollution has been shown to be damaging to black oaks in the San Bernardino mountains (McDonald 1990).

    LITERATURE CITED

    Bakker, Elna S. 1971. An Island Called California: An Ecological Introduction to its Natural Communities. University of California, Press. Berkeley, California.

    Lathrop, Earl W. and Bradford D. Martin. 1982. Response of Understory Vegetation to Prescribed Burning in Yellow Pine Forests of Cuyamaca Rancho State Park, California. Alsio 10(2): 329-343.

    McDonald, Philip A. 1980. California Black Oak. In Forest Cover Types of the United States and Canada. ed. F. H. Eyre. Society of American Foresters, Washington, D.C. p. 122.

    McDonald, Philip A. 1990. Quercus kelloggii - California black oak. Pages 661-671 in R.M. Burns and B. H. Honkala, technical coordinators. Silvics of North America, Volume 2. Hardwoods. Agriculture Handbook 654. USDA, Forest Service, Washington, D.C.

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Thorne, Robert F. 1977. Montane and Subalpine Forests of the Transverse and Peninsular Ranges In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 538-557.

    Vogl, Richard J. 1976. An Introduction to the Plant Communities of the Santa Ana and San Jacinto Mountains. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 77-98.

    Broadleaved Upland Forest

    VEGETATION ASSOCIATION:

    BROADLEAVED UPLAND FOREST

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Broadleaved upland forest is a general classification for several different vegetation types that occur within California (Holland 1986). Within the Plan Area, these may include canyon live oak, interior live oak and big-cone Douglas-fir Habitats. This general classification likely was applied to vegetation that was differentiated by patterns on topography from the aerial photography, but that was not field-checked for species composition.

    BIOGEOGRAPHY

    The discussion of biogeography is based on species ranges and distribution known from the Plan Area because of the wide range of Habitats within broadleaved upland forest in California. Canyon live oak ranges from Siskiyou County, in extreme northern California, to San Diego County, in the extreme south (Finch and McClerry 1980). Sawyer and Keeler-Wolf (1995) describe the species in Baja California as well. The species occurs in the Cascade, Sierra Nevada and Coast Ranges and may range from sea level to 2740 m (Finch and McClerry 1980). Interior live oak may occur in monotypic stands or as a dominant canopy species between 500 and 4500 m in the Klamath, Cascade, Coastal, Sierra Nevada, Transverse and Peninsular Ranges of California (Sawyer and Keeler-Wolf 1995). The big-cone Douglas-fir component of this Habitat is limited to the Transverse and Peninsular ranges of southern California only (McDonald 1990).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Keeler-Wolf described stands dominated by interior live oak at Millard Canyon, Hall Canyon, and Fern Canyon (Sawyer and Keeler-Wolf 1995). A location in the Cahuilla Mountain Research Natural Area (RNA) contains vegetation which fits the interior live oak - canyon live oak series (Sawyer and Keeler-Wolf 1995). This Habitat was mapped in the GIS database to include 2,762 acres including populations near Poppet Flats, San Gorgonio and in the Santa Ana Mountains. The Poppet Flats area contains broadleaved upland forest in linear strips, west of the Banning-Idyllwild Highway and northwest of Lake Fulmor. A population is mapped south of Mount San Gorgonio in the northeast corner of the Plan Area, mostly within the San Bernardino National Forest adjacent to the Morongo Indian Reservation. The largest populations are mapped in the Santa Ana Mountains, on the western edge of the Plan Area. This population is mainly on the eastern slope of Santiago Peak but generally occurs between Trabuco Peak and Buds Peak within the Cleveland National Forest.

    VEGETATION CHARACTERISTICS

    Classification of broadleaved upland forest varies widely in descriptions of Habitats within California and the continent. Provided here is a discussion of associations either dominated by canyon live oak (Quercus chrysolepis) or interior live oak (Quercus wislizenii) or a combination of both. Also included in this description is a discussion of big-cone Douglas-fir (Psuedotsuga macrocarpa) and various oaks.

    The canopy in a broadleaved upland forest, where dominated by interior live oak, can reach 15 m and varies from continuous to intermittent and even savanna-like (Sawyer and Keeler-Wolf 1995). Shrubs generally are infrequent to common and ground cover is sparse (Sawyer and Keeler-Wolf 1995). Vegetation composition varies within California including woodlands and forests and areas composed mainly of shrubs (Sawyer and Keeler-Wolf 1995).

    The canopy, where dominated by canyon live oak, can reach 30 m in a continuous cover and may be two-tiered (Sawyer and Keeler-Wolf 1995). The upper tier consists of conifers and the lower tier of canyon live oak. Other species present include black oak (Quercus kelloggii), California bay, (Umbellularia californica), big-cone Douglas-fir, Coulter pine (Pinus coulteri), ponderosa pine (Pinus ponderosa) and/or white fir (Abies concolor) (Sawyer and Keeler-Wolf 1995).

    Cooper (1922) identifies a Quercus chrysolepis-kelloggii association, where the important tree species are canyon live oak and black oak (Quercus kelloggii) with the understory varying widely throughout the state. Other common trees are Pacific madrone (Arbutus menziesii), California bay (Umbellularia californica), bigleaf maple (Acer macrophyllum), and buckeye (Aesculus) (Cooper 1922). Coulter pine may be a component of this Habitat on mesic sites (Holland 1986). These species also may form the understory beneath big-cone Douglas-fir (Cooper 1922). The Habitat has broad transition areas to adjacent Habitat types. The overstory of these woodlands generally has a mix of deciduous and evergreen hardwoods with a typical canopy height of 4 to 21 m tall(Holland 1988).

    In the Santa Ana and San Jacinto Mountains, Vogl (1976) identified an association of evergreen groves of interior live oak or canyon oak. The trees are physically small and crowded, reducing the presence of understory vegetation. The two most commonly found subdominant species are hybrid oaks and California bay (Vogl 1976).

    An alternate form of broadleaved upland forest was described in McDonald and Littrell (1976) where big-cone Douglas-fir is part of oak woodland communities, often on north slopes where canyon live oak predominates. The bulk of the big-cone Douglas-fir distribution occurs in mixed hardwood forest and chaparral (Sawyer et al. 1977). Big-cone Douglas-fir also may occur as the sole or dominant tree species in the canopy of many north-facing slopes (Sawyer and Keeler-Wolf 1995). The canopy height generally is less than 30 m and continuous with infrequent shrubs and a sparse ground layer (Sawyer and Keeler-Wolf 1995). Percent cover of big-cone Douglas-fir also depends on elevation. At lower occurrences (below 700 m) the trees are scattered above a closed canopy of canyon live oak (Barbour and Minnich 2000). Big-cone Douglas-fir cover increases to equal that of canyon live oak at around 1500 m (Barbour and Minnich 2000).

    McDonald (1990) notes that canyon live oak and big-cone Douglas-fir are strongly associated and may be considered a climax community. At higher elevations, tree associates may include ponderosa pine, Jeffrey pine (Pinus jeffreyii), singleleaf pinyon (Pinus monophylla), sugar pine (Pinus lambertiana), incense-cedar (Calocedrus decurrens) and California white fir (Abies concolor) (McDonald 1990). At middle elevations, Coulter pine, California walnut (Juglans californica), California black oak, California ash (Fraxinus dipetala) and California-laurel are common associates (Sawyer and Keeler-Wolf 1995, McDonald 1990). At lower elevations species such as coast live oak (Quercus agrifolia), toyon (Heteromeles arbutifolia), Nuttall’s scrub oak (Quercus dumosa), mountain-mahogany (Cercocarpus minutiflorus), laurel sumac (Malosma laurina), sugar bush (Rhus ovata), big-leaf maple and white alder (Alnus rhombifolia) may co-occur with bigcone Douglas-fir (McDonald 1990). Shrub associates are many and also vary with elevation; typical genera associated with big-cone Douglas-fir include, chamise (Adenostoma spp.), sagebrush (Artemesia), California-lilac (Ceanothus spp., buckwheat (Eriogonum), manzanita (Arctostaphylos spp.), and buckthorn (Rhamnus spp.) (McDonald 1990).

    In the Santa Ana Mountains, Vogl (1976) observed big-cone Douglas-fir growing with chamise, manzanita, coast live oak, canyon oak and interior live oak.

    PHYSICAL ENVIRONMENT

    Broadleaved upland forest typically occurs on steep mesic sites, intermediately between chaparral and Coulter pine (Pinus coulteri) in the Santa Ana and San Jacinto Mountains. In California, the Habitat occurs between 500 and 4500 m.

    Where dominated by canyon live oak, this Habitat can occur on all aspects, on stream sides, canyon bottoms or near streams (Sawyer and Keeler-Wolf 1995). Canyon live oaks prefer shallow, well-drained soils (Sawyer and Keeler-Wolf 1995). The range for this association is between 450 and 2000 m (Sawyer and Keeler-Wolf 1995).

    The interior live oak - canyon live oak series occurs mainly on north-facing slopes, often very steep areas, between 700 and 1900 m on alluvial or bedrock-derived soils, often with rocks (Sawyer and Keeler-Wolf 1995).

    The apparent aspect preferences of big-cone Douglas-fir vary with elevations: at low elevations, as low as 300 m, it occurs mainly on northern aspects and canyon bottoms; at middle elevations it may occur in a variety of aspects; and at high elevations (above 2400 m) on west and south aspects (Sawyer et al.1977). Big-cone Douglas-fir may grow in areas with a precipitation range from 250 to 1270 mm per year, with the majority of it being provided by rain (McDonald 1990). Soils supporting big-cone Douglas-fir vary although inceptisols were frequently found and alfisols occasionally found (McDonald 1990). When associated with canyon live oak, big-cone Douglas-fir occurs on steep north-facing slopes, in ravines, and on bedrock surfaces with sandstone or schist-derived, shallow, well-drained soils (Barbour and Minnich 2000; Sawyer and Keeler-Wolf 1995). The big-cone Douglas-fir component of this community in the Santa Ana mountains is present mainly on steep, unstable and rocky north-facing slopes (Vogl 1976).

    ECOSYSTEM PROCESSES

    Because of the wide variety of Habitats within broadleaved upland forest, no literature specific to the type was available. Information on the effects of fire or regeneration in broadleaved upland forest was not reviewed. Ecosystem processes for oak-dominated Habitats are discussed within the oak woodlands discussion. The effects of fire on big-cone Douglas-fir has been well-studied and a summary is presented below.

    Repeated fires are most often found in lower elevations and are likely to extirpate big-cone Douglas-fir, leaving oaks or chaparral. Less frequent intense fires, on the other hand, result in a scattered distribution of big-cone Douglas-fir trees of several age classes (McDonald 1990). Steep slopes act to protect these trees from fire damage (McDonald 1990). Regeneration of big-cone Douglas-fir will occur only where fire is absent because the trees generally regenerate best in post-burn conditions and with above-average rainfall by producing trunk resprouts (Vogl 1976; McDonald 1990). Others contend that the big-cone Douglas-fir phase is not adapted to fire but also seems to be suffering the effects of fire suppression (Barbour and Minnich 2000). Between 1938 and 1983, fires occurred on the average every 65 years and the overall stand area decreased; i.e., afforestation occurred more frequently that deforestation because of the inability of small patches to recover from fires (Barbour and Minnich 2000). Other studies have hypothesized that big-cone Douglas-fir dominated areas are reduced by fire, invaded by oak woodland or oak chaparral, and that recolonization by big-cone Douglas-fir requires hundreds of years (McDonald and Littrell 1976).

    THREATS

    The Land and Resources Management Plan for the Cleveland National Forest (USDA 1986) identified the gradual reduction of grazing within this community to allow for oak regeneration and the development of shrub and herbaceous layers. Armillaria root disease (Armillaria mellea) infects dead, dying or felled oaks, eventually infecting nearby or planted conifers (Stephenson and Calcarone 1999). Fire is the greatest threat to big-cone Douglas-fir as damage from insects and disease are not well reported (McDonald 1990).

    LITERATURE CITED

    Barbour, Michael G. and Richard A. Minnich. 2000. Californian Upland Forests and Woodlands. In North American Terrestrial Vegetation. Second Edition. Cambridge University Press, Cambridge, UK.

    Cooper, W.S. 1922. The broad-sclerophyll vegetation of California. Publication 319. Carnegie Institution of Washington, Washington, D.C.

    Finch, Sherman J. and Dick McClerry. 1980. Canyon Live Oak. In Forest Cover Types of the United States and Canada. ed. F. H. Eyre. Society of American Foresters, Washington, D.C. 125-126.

    Holland, V. L. 1988. Coastal Oak Woodland. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 78-79.

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company. Dubuque, Iowa.

    McDonald, Philip A. and Edward E. Littrell. 1976. The Bigcone Douglas-Fir-Canyon Live Oak Community in Southern California. Madrono, vol. 23:310-320.

    McDonald, Philip A. 1990. Psuedotsuga macrocarpa - big-cone Douglas-fir. Pages 520-525 in R.M. Burns and B. H. Honkala, technical coordinators. Silvics of North America, Volume 1. Conifers. Agriculture Handbook 654. USDA, Forest Service, Washington, D.C. 1990.

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Stephenson, John R. and Gena M. Calcarone. 1999. Southern California mountains and foothills assessment: habitat and species conservation issues. General Technical Report GTR-PSW-172. Albany, CA: Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture; 402 p.

    USDA, Forest Service. 1986. Land and Resources Management Plan: Cleveland National Forest. USDA, Forest Service, Cleveland Nation Forest, San Diego, CA.

    Vogl, Richard J. 1976. An Introduction to the Plant Communities of the Santa Ana and San Jacinto Mountains. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 77-98.

    Chaparral

    VEGETATION ASSOCIATION:

    CHAPARRAL

    MAPPED SUBASSOCIATIONS

    Chaparral (undifferentiated), Chamise Chaparral, Red Shank Chaparral, and Semi-Desert Chaparral. Vegetation Chaparral

    DATA CHARACTERIZATION

    No objective measurement of the accuracy of the MSCHP vegetation map has been conducted, therefore no definitive statements can be made about the accuracy of chaparral vegetation on a polygon by polygon basis. Because mapping utilized several different sources and was spot-checked in the field, it is probable the general distribution of chaparral within the Plan Area has been accurately depicted. The boundaries between chaparral and most vegetation types (e.g., coastal sage scrub, grasslands, oak woodlands, and coniferous forest) are reasonably interpretable from the 1:24000 aerial photos. Inaccuracy at small scales (e.g., variation in polygon boundaries or mis-classification) due to problems of interpretation are most likely to occur in areas that were recently burned when the photos were taken, areas where the vegetation types are fragmented, and in more arid regions where chaparral shrubs are sparse (e.g, the Badlands and Vail Lake areas). In these areas there is a higher probability for misinterpretation of chaparral and coastal sage scrub. Because much of the mapping was field checked specifically for coastal sage scrub these errors should be infrequent.

    Because of the difficulty in interpreting different types of chaparral from the scale of the aerial photography the acreage of chamise, red shank chaparral, and semi-desert chaparral is likely understated. For the same reason other subassociations of chaparral (montane chaparral, ceanothus chaparral, and manzanita chaparral) known to occur within the Plan Area were not differentiated in the mapping.

    Research concerning the physiology and ecology of chaparral species is extensive; however, important questions are unresolved concerning some of the ecosystem processes important to the development and maintenance of chaparral (e.g., the appropriate fire regimes for long-term management of the vegetation type).

    BIOGEOGRAPHY

    Chaparral vegetation occurs regionally along the Pacific Coast to the mountain foothills (0–2,000 meters) from southern Oregon to the San Pedro Martir Mountains in Baja California (Detling 1961; Axelrod 1973). Most of chaparral’s distribution lies within the state of California where it is one of the most widespread vegetation types, encompassing approximately 29,000 square-kilometers (km) or seven percent of the total land area of the state (Davis et al. 1994). Species composition is varied within California where as many as 50 different subassociations have been recognized (Sawyer and Keeler-Wolf 1995). Additional forms of chaparral are known from Arizona and northeastern Mexico, and the Rocky Mountain Region but these types are isolated by greater than 200 km of desert, and are adapted to summer rainfall and a different fire regime (Keeley 2000). Within the Plan Area many subassociations of this vegetation type occur at both regional and local scales based on variation in rainfall, temperature, elevation, fire regime, slope aspect, soils and disturbance regimes.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Chaparral vegetation is the most abundant and widespread vegetation type in Western Riverside County, covering approximately 35% (435,000 acres) of the Plan Area. Large contiguous stands of chaparral occur along the Santa Ana Mountains in the western portion of the Plan Area, and along the San Bernardino, San Jacinto, and Agua Tibia Mountains in the eastern and southern portions. Although chaparral is less common than other vegetation types in the central lowlands of Riverside County three large chaparral-dominated areas occur on steeper lands near the Gavilan Hills-Gavilan Plateau-Meadowbrook Region, the Lakeview Mountains-Double Butte area, and the Sedco Hills-Hogbacks area.

    Four types of chaparral have been mapped for the Plan Area based on variation in species composition: chamise chaparral, red shank chaparral, semi-desert chaparral, and chaparral (undifferentiated). The undifferentiated chaparral category is an artifact of the amalgamation of several different mapping sources and less detailed mapping effort of chaparral in portions of the Plan Area (PSBS 1995). The areas mapped as undifferentiated chaparral are composed of many different chaparral sub associations including southern mixed chaparral, montane chaparral, scrub oak chaparral, and several forms of manzanita or ceanothus chaparral. Because much of the variation of these vegetation types is continuous with nodes of dominance by one or a few species, these vegetation types will be described within the mixed chaparral category for this Habitat account. It is probable that a significant portion of undifferentiated chaparral includes chamise chaparral and smaller patches of red shank and semi-desert chaparral.

    VEGETATION CHARACTERISTICS

    Chaparral is a shrub-dominated Habitat that is composed largely of evergreen, sclerophyllous species that range from 1 to 4 meters in height (Keeley 2000). Other growth forms including soft-leaved subshrubs, perennial herbs, geophytes (bulbs and corms), and annual herbs are less abundant in mature chaparral but can be present in abundance in early and late successional stands of chaparral (Keeley and Keeley 1988). Sparse stands of trees can occur within chaparral, typically within transition areas with conifers at higher elevations and oaks on north-facing slopes or ravines (Hanes 1977; Keeley and Keeley 1988). Depending on the species composition and underlying topography and soil, the structure of chaparral can range from low, monotonous, smooth-textured vegetation to more heterogeneous stands approaching the vertical structure of woodlands (Keeley 2000 ).

    From inland and high elevations to coastal locations, chaparral occurs in both large continuous stands or within a patchwork of Habitats including coastal sage scrub, grasslands, oak woodlands, coniferous Habitats and several wetland Habitats (Heady 1977; Hanes 1977; Callaway and Davis 1993). Chaparral near the coast tends to occur in disjunct patches occupying more mesic Habitats whereas coastal sage scrub is distributed more extensively in drier Habitats (Kirkpatrick and Hutchinson 1980; Malanson and O’Leary 1994). Mountain foothill and high elevation stands of chaparral are larger and more continuous, with coastal sage scrub occurring in smaller patches generally restricted to steep and south-facing exposures (Keeley 2000; PSBS 1995). Oak woodlands border chaparral in more mesic areas (e.g. ravines, north-facing slopes) that have developed deeper soils (Griffen 1977). Oak woodlands are thought to develop within late successional chaparral in areas with more developed soils (Cooper 1922; Wells 1962). The native grassland-chaparral interface is not well understood; however, research has shown cases of type conversion from chaparral to non-native annual grasslands with frequent fire or mechanical disturbance (Zedler et al. 1983).

    Chaparral generally is thought to be a fire dependent system based on the many adaptations of its characteristic species, and its resilience in form and species composition to periodic burning (Keeley 1986; Keeley 1992). Most of the characteristic shrub species in chaparral can be organized generally into three adaptive strategies related to fire: (1) shrubs that have stems that regenerate following fire from below ground burls (resprouters); (2) shrubs that produce large amounts of dormant seed that persist for long periods of time and germinate by heat or chemical processes initiated by fire (obligate seeders); and (3) plants that apply both strategies (Keeley 1977). Within chaparral vegetation non-shrub plant growth forms may also employ these strategies or fire avoidance to persist within this fire prone system (e.g., geophyte species whose bulbs or corms persist following fire, annual herb species with long seed dormancy and heavy annual seed production, annuals with the ability to disperse seeds over long distances) (Keeley 1986).

    The species composition of a particular chaparral stand is largely influenced by fire. Chaparral generally returns to pre-fire structure and composition within a normal fire regime (Keeley 1986); however, considerable research has documented various effects of fire regime on species mortality (Keeley 2000). Frequency of fire has been shown to affect chaparral species composition, where short fire intervals may eliminate obligate seeding species in favor resprouters (Keeley 1986; 1992). Additional research has shown that fire temperature or intensity also has a strong influence on post-fire species composition (Davis et al. 1989; Rice 1993; Tyler 1995). Stand age following fire is thought to influence the reproduction of species based on reproductive strategies. Research has shown that seedling recruitment is more common for resprouting species in old (> 56 yr.) stands of chaparral whereas seedling recruitment for obligate seeding species was extremely uncommon (Keeley 1986; 1992). This research has led to the conclusion that short intervaled fires may adversely affect the presence of obligate resprouting species in favor of obligate seeders.

    SPECIES COMPOSITION

    The floristic composition of chaparral varies depending on biogeography, local Habitat characteristics and fire history. Of the many growth forms present in chaparral, woody evergreen perennials are the dominant plants and, as such, exert the most influence on the Habitat. The most common and widespread species within chaparral is chamise (Adenostoma fasciculatum) (Hanes 1971 ). This species occurs in most stands of chaparral and is the dominant plant in drier Habitats (Keeley 2000). The ubiquity of this species is likely explained by its many adaptations to drought, fire and disturbance (Hanes 1977). Other common shrub species include representatives from manzanita (Arctostaphylos spp.), wild-lilac (Ceanothus spp.), silk-tassel bush (Garrya spp.), oak (Quercus spp.), redberry (Rhamnus spp.), Rhus spp., laurel sumac (Malosma laurina), mountain-mahogany (Cercocarpus betuloides), toyon (Heteromeles arbutifolia), holly-leaf cherry (Prunus ilicifolia), and mission manzanita (Xylococcus bicolor) (Holland 1986). Soft-leaved subshrubs are less common in chaparral than in coastal sage scrub but occur within canopy gaps of mature stands, and may be more prevalent following fire (Holland 1986; Keeley and Keeley 1988; Sawyer and Keeler-Wolf 1995). Common species include California buckwheat (Eriogonum fasciculatum), sages (Salvia spp.), California sagebrush (Artemisia californica), and monkeyflower (Mimulus spp.). Suffrutescent and perennial herbaceous species commonly include deerweed (Lotus scoparius), nightshade (Solanum spp.), Spanish bayonet (Yucca whipplei), rock-rose (Helianthemum scoparium), golden yarrow (Eriophyllum confertiflorum), Bloomeria spp., Brodiaea spp., onion (Allium spp.), sanicle (Sanicula spp.), Lomatium spp., soap plant (Chlorogalum spp.), and bunch grasses (Nassella spp., and Melica spp.) (Holland 1986; Keeley and Keeley 1988; Sawyer and Keeler-Wolf 1995 ). Vines commonly present in chaparral include wild cucumber (Marah spp.), dodder (Cuscuta spp.), chaparral-pea (Lathyrus spp.), bedstraw (Galium spp.), poison-oak (Toxicodendron diversilobum), and honeysuckle (Lonicera spp.) (Keeley and Keeley 1988). Annual species persisting in mature chaparral or in the post-burn flora vary according to geographic location, but typically include lupine (Lupinus spp.), Lotus spp., California thread-stem (Pterostegia drymarioides), Claytonia spp., Gnaphalium spp., Phacelia spp., Gilia spp., whispering bells (Emmenanthe pendulflora), fiesta-flower (Pholistoma spp.), and many others (Holland 1986; Keeley and Keeley 1988; Sawyer and Keeler-Wolf 1995).

    VARIATION IN SPECIES COMPOSITION, STRUCTURE AND DISTRIBUTION BY MAPPED HABITAT TYPE

    Chamise Chaparral: Chamise chaparral occurs throughout much of the range of chaparral in California from approximately 10 to 1800 meters in elevation (Sawyer and Keeler-Wolf 1995). This vegetation is found on all slope-aspects on shallow soils (Gordon and White 1994; Sawyer and Keeler-Wolf 1995). Seven patches of chamise chaparral totaling 362 acres were delineated within the Plan Area. These patches were identified at Estelle Mountain, Steele Peak, Meadowbrook, southeast of Table Mountain, and in the Lakeview Mountains near the town just west of Hemet. The mapped acreage is an underestimate. Because of the difficulty in differentiating different subassociations of chaparral from aerial photography many stands of chamise chaparral likely were included within the undifferentiated chaparral category.

    Vegetation structure is open to dense between 1 to 4 meters in height, with little litter and few understory species in mature stands (Hanes 1971; Holland 1986; Boyd and Banks 1995). Recent studies describe this association as having greater than 60% cover chamise (Gordon and White 1994; Sawyer Keeler-Wolf 1995). Where another species occurs at greater than 30% cover but chamise remains the dominant cover the stands have been described as a mixed series (e.g., chamise-bigberry, chamise-black sage, chamise-cupleaf ceanothus, chamise-Eastwood’s manzanita, chamise-hoaryleaf ceanothus, chamise-mission manzanita-woollyleaf ceanothus series, chamise-wedgeleaf ceanothus, and chamise-white sage) (Sawyer and Keeler-Wolf 1995). These series all are likely included within the undifferentiated chaparral category in the MSHCP database. Evergreen species that commonly occur at less than 30 % cover within chamise chaparral in the MSHCP Plan Area include bigberry manzanita (Arctostaphylos glauca), Eastwood’s manzanita (Arctostaphylos glandulosa), mission manzanita (Xylococcus bicolor), scrub oak (Quercus berberidifolia), interior live oak (Q. wislizneii), hoary-leaved ceanothus (Ceanothus crassifolius), Spanish bayonet (Yucca whipplei), laurel sumac (Malosma laurina), sugar-bush (Rhus ovata), toyon (Heteromeles arbutifolia), yerba santa (Eriodictyon crassifolium and E. trichocalyx), redberry (Rhamnus crocea), and chaparral beard-tongue (Keckiella antirrhinoides). Chamise chaparral often supports low cover of subshrubs characteristic of sage scrub (e.g., black sage [Salvia mellifera]), California buckwheat [Eriogonum fasciculatum], California sagebrush [Artemisia californica]), and saw-toothed goldenbush [Hazardia squarrosa]). Perennial herbaceous species are few in mature stands of chamise chaparral but cudweed (Gnaphalium spp.), Sanicula spp., southern taushia (Tauschia arguta), California melic (Melica imperfecta), lilac mariposa (Calochortus splendens), Bigelow's spike-moss (Selaginella bigelovii), and other post-burn or gap specialist annuals (e.g., Phacelia spp., whispering bells [Emmenanthe pendulaflora], Cryptantha spp., Plagiobothrys spp., spineflower [Chorizanthe spp.], Camissonia spp., and Pterostegia drymarioides) may be present.

    Undifferentiated Chaparral/Mixed Chaparral: The majority of the chaparral vegetation in the Plan Area is mapped as undifferentiated chaparral. This vegetation covers approximately 363,000 acres, encompassing 29 % of the Plan Area. Several different subassociations may be included within this vegetation type based on variation in slope aspect, elevation and distance from the coast.

    In the western portion of the Plan Area (e.g., the Santa Ana Mountains, Agua Tibia Mountains) undifferentiated chaparral is dominated by chamise in drier Habitats and by a more diverse mixture of species in more mesic areas (analogous to southern mixed chaparral sensu, Holland 1986). Species composition ranges from that described for chamise chaparral (see above) to more diverse shrub associations supporting hoary-leaved ceanothus (Ceanothus crassifolius), birchleaf mountain-mahogany (Cercocarpus betuloides), toyon (Heteromeles arbutifolia), sugar bush (Rhus ovata), holly-leaf redberry (Rhamnus ilicifolia), heart-leaf penstemon (Keckiella cordifolia), southern honeysuckle (Lonicera subspicata), scrub oak (Quercus berberidifolia), black sage (Salvia mellifera), and other species.

    The central portion of the Plan Area (e.g., the Gavilan Hills, Sedco Hills, and Black Hills) is lower in elevation and supports a drier expression of chaparral with abundant stands of chamise along with more arid climate chaparral species [e.g., jojoba (Simmondsia chinensis), chaparral beard-tongue (Keckiella antirrhinoides), and desert thorn (Lycium andersonii)] and Riversidian sage scrub [e.g., brittlebush (Encelia farinosa), California buckwheat (Eriogonum fasciculatum var. polifolium) and white sage (Salvia apiana)]. Mesic areas (e.g., north-facing slopes, narrow ravines) in this region support southern mixed chaparral, or red shank chaparral.

    The eastern portion of the Plan Area including the San Bernardino, San Jacinto, and Agua Tibia Mountains and foothill areas supports the largest continuous stands of undifferentiated chaparral. This area has a large elevational gradient supporting chaparral from about 400 to 2000 meters. This gradient allows for high variation in chaparral species composition and many different species associations. At lower elevations chaparral is composed of chamise, chaparral beard-tongue, toyon, scrub oak, California buckwheat and other species common to coastal southern California. With increasing elevation chamise remains abundant but with Eastwood’s manzanita (Arctostaphylos glandulosa), bigberry manzanita (Arctostaphylos glauca), birch-leaf mountain-mahogany, chaparral whitethorn (Ceanothus leucodermis) and interior live oak (Q. wislizenii) becoming more abundant. At higher elevations where chaparral transitions to coniferous forest, chaparral is largely composed of Eastwood’s manzanita, bigberry manzanita, pink-bract manzanita (A. pringlei), chaparral whitethorn (Ceanothus leucodermis), deer bush (C. integerrimus), and Veatch's silk-tassel (Garrya veatchii). At higher elevations chaparral also may support sparse stands of coniferous and oak species including Jeffrey pine (Pinus jeffreyi), ponderosa pine (P. ponderosa), Coulter pine (P. coulteri), black oak (Quercus kelloggii), canyon live oak (Q. chrysolepis), and interior live oak (Q. wislizenii). Understories are varied along the elevation gradient with species typical of chamise chaparral understories occurring at lower elevations, and species such as Bigelow's spike-moss (Selaginella bigelovii), bedstraw (Galium spp.), bird’s-beak (Cordylanthus sp.), wallflower (Erysimum capitatum), yarrow (Achillea millefolium), rock cress (Arabis perennans), whiskerbrush (Linanthus ciliatus), claytonia (Claytonia parviflora), and Indian paintbrush (Castilleja spp.) occurring within canopy gaps at higher elevations. Presence of fire-following annual species in higher elevation chaparral is not as evident as at lower elevations (Sweeney 1968). Most of the species present after fire are present in open areas prior to burning (Hanes 1977).

    Red Shank Chaparral: Red shank chaparral has a more restricted distribution than other chaparral associations within California with the greatest areal extent occurring within Riverside and northern San Diego counties in the interior valleys and the San Jacinto and Santa Rosa Mountains (Hanes 1977). The only other areas supporting significant stands of red shank chaparral are the Laguna and Palomar Mountains in San Diego County, inland northern Baja California, the Santa Monica Mountains in Los Angeles County, and the Cuyama Valley in Santa Barbara and San Luis Obispo counties (Hanes 1977). This vegetation generally occurs between 300 and 2000 meters and is usually found in areas with colder winters and greater precipitation than chamise chaparral (Holland 1986). Redshank chaparral is the predominate vegetation type in the southeastern portion of the Plan Area. This area supports large blocks of redshank chaparral from the San Diego County border to the Table Mountain area. This vegetation covers approximately 1 percent of the Plan Area totaling 72,000 acres. North of Anza, redshank chaparral occurs in smaller patches interspersed with chamise chaparral, Jeffery pine forest, and oak woodland. Redshank chaparral is structurally taller (2 to 5 meters) than chamise chaparral and is more commonly found on granitic, more mesic and fertile soils, and deep alluvial soils (Beatty 1984; Holland 1986; Sawyer and Keeler-Wolf 1995). Typically red shank chaparral is dominated by red shank (Adenostoma sparsifolium), although many other shrub species may be present including chamise, toyon, holly-leaved cherry, bigberry manzanita, birch-leaf mountain-mahogany, Eastwood manzanita, cupleaf ceanothus (Ceanothus greggii), hoary-leaved ceanothus (Ceanothus crassifolius), scrub oak (Quercus berberidafolia), hollyleaf redberry (Rhamnus ilicifolia), and Veatch silktassel. Because of the more open architecture of Adenostoma sparsifolia, red shank chaparral typically has greater abundances of herbaceous species within the understories of mature stands than chamise chaparral (Hanes 1977; Boyd and Banks 1995). Typical herbaceous species may include southern tauschia (Taushia arguta), mesa saxifrage (Jepsonia parryi), coast figwort (Scrophularia californica), blue dicks (Dichelostemma capitata), miner's lettuce (Claytonia perfoliata), baby blue-eyes (Nemophilia menziesii), dwarf stonecrop (Crassula connata), and other common chaparral understory species (Boyd and Banks 1995).

    Semi-Desert Chaparral: Semi-desert chaparral is a transitional chaparral association that occurs primarily on the interior of the transverse and peninsular ranges on the border of the Mojave and Colorado deserts between 610-1500 meters in elevation (Holland 1986). Semi-desert chaparral was mapped in four separate polygons (19 acres) within a mile of the community of Anza. Because semi-desert chaparral may not appear distinctly from aerial photo interpretation additional acreage of this vegetation type is probably included within undifferentiated chaparral in the desert transition areas. Shrub cover for this vegetation is lower than most other chaparral associations owing to its occurrence within drier regions (Gordon and White 1994). Species composition usually includes chamise and cup-leaved ceanothus as co-dominates with lower cover of sugarbush, scrub oak (Quercus berberidifolia), California buckwheat, Spanish bayonet (Yucca whipplei), birch-leaved mountain mahogany, and desert-apricot (Prunus fremontii ) (Gordon and White 1994; Sawyer and Keeler-Wolf 1995).

    PHYSICAL ENVIRONMENT

    Chaparral generally is limited to the Mediterranean-type climate region of California from sea level to approximately 2000 meters in elevation (Keeley 2000). This region receives from 200 to 1000 mm of precipitation, most of which falls from November to April (Miller and Hajek 1981). Because this region has concentrated rainfall in the winter months and prolonged summer drought much of the current flora has adaptations that allow for rapid growth during the winter and spring and water Conservation during the summer and fall (Harrison et al., 1971; Poole and Miller 1981). Because many of these adaptations have facilitated periodic fires, most chaparral species have evolved mechanisms to succeed within a fire-prone ecosystem as described below.

    Although physical environmental factors typically exert a strong influence over the distribution of vegetation at the landscape-scale, the influence of fire upon chaparral makes these relationships more complex. Few strong relationships have been established between specific physical site factors and the distribution of chaparral on the landscape (Wells 1962; Bradbury 1978; Keeley 2000). Chaparral is known to occur on many different types of soils and geologic substrates (Minnich and Howard 1984). The literature relating physical environmental factors to chaparral species composition showed few strong or repeatable relationships over regions, however, the literature revealed that localized patterns may exist based on topographic position and soils (Gauss 1964; Gordon and White 1994; Keeley 2000). In coastal areas where coastal sage is more widely distributed, chaparral is more likely to occur within more mesic topographic positions (e.g. north-facing slopes, and areas with low slope angles) (Kirkpatrick and Hutchinson 1980). Within more inland and high elevation sites chaparral is more broadly distributed occurring on most slope-aspects and slope angles (Hanes 1971; Gordon and White 1994). Generally chaparral is thought to occur upon thin, porous, and rocky soils that are relatively low in nitrogen, potassium and phosphorous (Vlamis et. al. 1954; Crawford 1962; cited in Hanes 1977). One study has provided evidence that while chaparral and other Mediterranean-type vegetation changes over time depending on fire and disturbance regimes, chaparral is likely to be the edaphic climax vegetation on well drained rocky soils with or without the influence of fire (Callaway and Davis 1993). Most research has asserted that physical environmental factors and fire interact to control the landscape-scale distribution of chaparral versus other vegetation types (e.g., coastal sage scrub, oak woodlands, and grasslands) (Westman 1983; Callaway and Davis 1993; Zedler et. al. 1983).

    ECOSYSTEM PROCESSES

    The dominant driving force in chaparral is fire. The majority of chaparral species are either adapted to occasional fire or are able to persist in fire prone ecological regimes (Hanes 1977; Zedler and Zammit 1989). Chaparral has been described as “autosuccessional,” undergoing a rapid succession from largely herbaceous flora immediately after fire to relatively dense woody vegetation in a short time period with minimal loss of species (Hanes 1971; Zedler and Zammit 1989). Early research suggested that without fire, chaparral would develop into oak woodlands or grasslands (Sampson 1944; Wells 1962). Chaparral succession to oak woodlands may occur in mesic situations adjacent to current stands of oak woodlands (e.g., Callaway and D’Antonio 1991) but most research has provided examples of greater than 100 year-old chaparral stands without evidence of physiognomic succession (Zedler 1981; Keeley 1992). This research has shown that in addition to remaining stable and reproductively viable following long periods without fire, some chaparral species (most resprouting species) sexually reproduce largely within older aged stands (Zedler 1981; Keeley 1992). Additional research has shown that high frequency burning of chaparral in the presence of non-native grasses can cause type-conversion from shrublands to non-native grasslands (Wells 1962; Zedler et al. 1983; Keeley 1990). So that while chaparral appears to be fire-adapted, it can remain healthy for long periods without fire and too-frequent fire may cause conversion to grassland.

    The distribution and species composition of chaparral, at the landscape-scale, is largely influenced by varying interactions between fire regime (frequency, seasonal timing, size, and intensity) and physical environment (Zedler 1981; Zedler et al. 1983; Davis et al. 1988; Moreno and Oechel 1991a; Minnich 1995; Keeley 2000). The primary source of wildfires prior to human alteration of the “natural” fire regime was lightning (Keeley 1982). Although lightning-caused fires remain, fire regimes have changed due to increased anthropogenic ignitions and fire suppression (Keeley 1982; Timbrook et al. 1982; Minnich 1995). Although much research has been conducted concerning the relationship between chaparral and fire, there are varying opinions on the frequency and patterns of the anthropogenic fire regimes. Two basic models have been described for the “natural” fire regime in chaparral: (1) the fuel dynamics model where fire regime is determined primarily by the amount of combustible plant material (fuel dynamics) and where ignition frequency and source are unimportant (Minnich 1989; 1995); and (2) the ignition/allogenic model where fire is dependent on stochastic combinations of events (weather, ignition source, topography, and vegetation flammability) and where fire history and patch structure are not important (Keeley et al. 1989; Minnich 1995). The fuel dynamics model suggests a shorter frequency of fires that preferentially burn where there are suitable fuel loads and avoid recently burned areas (Minnich 1989; 1995). According to the ignition/allogenic model the rates of fire return are presumed to be long-term with large and extensive fires (Keeley et al. 1989). Because questions remain about the fire regimes present prior to human alteration, the implications of the altered fire regimes on the functioning of ecosystems in the long-term is not clearly known (Keeley et al. 1989; Keeley 2000). Nonetheless, both models suggest that equilibrium in species composition of chaparral is likely produced by variable burning regimes.

    Most research has suggested returning to “natural” fire regimes similar to those existing before European settlement in order to improve chaparral Habitat quality (Minnich 1993; Keeley 2000). Because of the high risk involved in allowing fuel loads to develop in urban areas, such fire regimes must be managed in isolated preserves (Tippets et al. 1993). Preserve selection and management strategies that recognize the value of “natural” fire regimes are necessary to maintain Habitat capable of supporting the majority of the existing native plants and animals for long periods of time, and allowing for the evolution of new species.

    Larger preserve areas with less urban edges and more spatially heterogeneous topography and vegetation would be more likely to maintain “natural” fire regimes. Within larger and more topographically diverse areas there are more opportunities for different plant associations, age stands, and potential for unburned patches to occur. Larger areas with smaller urban edges also reduces the public safety risk from fire. Because of the existing pattern of urban development in southern California and the rarity of some Habitats and species near urban areas, smaller Habitat preserves are inevitable. In smaller preserve areas it may be necessary to actively maintain and manage more diverse patch structure to insure both public safety and functioning ecosystems. Because of the presence of annual grasses and higher frequencies of human fire ignitions in urban areas, active management likely will be required to maintain healthy populations of chaparral species. Because of increased fire frequency and the potential for type conversion of chaparral to non-native grasslands in reserves near urban development, management effort should be focused on maintaining a balance of both old (50 to 100+ years) and younger aged stands. Adaptive management programs will need to be tailored to the composition of chaparral, size of the reserve area, and biological goals of the reserve.

    THREATS

    Because chaparral and many of its component species are widely distributed there is no direct threat to chaparral as a vegetation type. Certain stands of chaparral that support sensitive species (e.g., Rainbow manzanita [Arctostaphylos rainbowensis]) or unique species composition may be threatened by urban development or type conversion at local scales. Precise locations of chaparral endemic sensitive species or unique chaparral associations are required to address potential threats. Large-scale changes in climate or pollution may affect the distribution of chaparral species but research on the effects of potential changes is not well developed. Fire suppression has been described as a threat to chaparral but this has not been demonstrated over large areas. Fire suppression that subsequently allows catastrophic fire likely is a threat to chaparral Habitat quality within isolated reserves because widespread catastrophic fire within a single age-class may eliminate species that do not reproduce well following fire.

    LITERATURE CITED

    Axelrod, D.I. 1973. History of Mediterranean ecosystem in California, Pp. 225-277 In F. De Castri and H.A. Mooney (eds.), Mediterranean ecosystems: origin an structure. Springer-Verlag, New York.

    Axelrod, D.I. 1989. Age and Origin of Chaparral, Pp. 7-19 In S.C. Keeley (ed.), The California chaparral: paradigms reexamined. Natural History Museum of Los Angeles County, Los Angeles, Science Series No. 34.

    Beatty, S.W. 1987. Origin and role of soil variability in southern California chaparral. Physical geography 8:1-17.

    Boyd, S. and D.L. Banks. 1995. A botanical assessment of the Agua Tibia Wilderness Area, Cleveland National Forest, California. Rancho Santa Ana Botanic Garden, Claremont, California.

    Boyd, S. 1983. A Floral inventory of The Gavilan Hills. Master’s Thesis. California State University Fullerton.

    Bradbury, D.E. 1978. The evolution and persistence of a local sage/chamise community pattern in southern California. Yearbook of the Association of Pacific Coast Geographers. 40:39-56.

    Callaway, F.M. and C.M. D’Antonio. 1991. Shrub facilitation of coast live oak establishment in central California. Madrono 38:158-169.

    Callaway, F.M. and F.W. Davis 1993. Vegetation dynamics, fire, and the physical environment in coastal central California. Ecology 74:1567-1578.

    Cooper, W.S. 1922. The broad-sclerophyll vegetation of California: an ecological study of the chaparral and its related communities. Carnegie Institution of Washington, Washington D.C.

    Crawford, J.M. 1962. Soils of the San Dimas Experimental Forest. USDA Forest Service, Pacific Southwest Forest and Range Experiment Station, Misc. paper PSW-76. Berkeley, California.

    Davis, F.W., D.E. Hickson, and D.E. Odion. 1988. Composition of maritime chaparral related to fire history and soil, Burton Mesa, Santa Barbara, County, California. Madrono 35:169-195.

    Davis F.W., M.I. Borchert, and D.C. Odion 1989. Establishment of microscale vegetation pattern in maritime chaparral after fire. Vegetatio 84:53-67.

    Davis F.W., P.A. Stine, D.M. Stoms, M.I. Borchert, and A.D. Hollander. 1994. Gap analysis of the actual vegetation of California. 1. The southwestern region. Madrono 42:40-78.

    Detling, L.E. 1961. The chaparral formation of south-eastern Oregon with consideration of its post glacial history. Ecology 42:348-357.

    Evans, R.A. and J.A. Young. 1989. Characterization and analysis of abiotic factors and their influences on vegetation. Pp. 13-28 In L.F. Huenneke and H.A. Mooney, (eds.) Grassland structure and function: California annual grasslands. Kluwer Academic Publishers, Boston, MA.

    Gauss , N.M. 1964. Distribution of selected plant species in a portion of the Santa Monica Mountains, California, on the basis of site. Master’s Thesis, University of California, Los Angeles.

    Gordon, H.J. and T.C.White 1994. Ecological Guide to southern California chaparral plant series. Technical Publication R5-ECOL-TP-005. USDA, Forest Service, Pacific Southwest Region, San Francisco California.

    Griffin, J.R. 1977. Oak Woodland. Pp.383-416. In M.G. Barbour and J. Major (eds.), Terrestrial vegetation of California. John Wiley and Sons, New York.

    Hanes T.L. 1971. Succession after fire in the chaparral of southern California. Ecological monographs 41:27-52.

    Harrison, A.T., E. Small, and H.A. Mooney. 1971. Drought Relationships and distribution of two mediterranean-climate plant communities. Ecology 52:869-875.

    Hanes T.L. 1977. California chaparral. Pp.417-470. In M.G. Barbour and J. Major (eds.), Terrestrial vegetation of California. John Wiley and Sons, New York.

    Heady H.F. 1977. Valley grasslands. Pp.491-514. In M.G. Barbour and J. Major (eds.), Terrestrial vegetation of California. John Wiley and Sons, New York.

    Holland, R.F. 1986. Preliminary descriptions of the terrestrial communities of California. California Department of Fish and Game, Nongame Heritage Program, Sacramento.

    Keeley, J.E. 1982. Distribution of lightning and man-caused wildfires in California, P.p, 431-4437 in C.E. Conrad and W.E. Oechel (eds.), Proceedings of the symposium on dynamics and management of mediterranean-type ecosystems. USDA Forest Service, Pacific Southwest Forest and Range Experiment Station, General Technical Report PSW-58.

    Keeley, J. E. and S.C. Keeley. 1984. Postfire Recovery of California coastal sage scrub. American Midland Naturalist 111:105-117.

    Keeley, J.E. 1977. Seed production, seed populations in soil, and seedling production after fire for two and vegetative sprouts in unburned chaparral. Ecology 73:1194-1208.

    Keeley, J.E. 1986. Resilience of Mediterranean shrub communities to fire. Pp. 95-112 In B. Bell, A.J.M. Hopkins, and B.B. Lamont (eds.), Resilience in mediterranean-type ecosystems. Junk, Dordrecht.

    Keeley, J. E. 1990. The California valley grassland. Pp. 2-23 in A.A. Schoenherr (ed.), Endangered plant communities of southern California. California State University, Fullerton. Southern California Botanists, Special Publication No. 3.

    Keeley, J.E. 1992. Demographic structure in of California chaparral in the long-term absence of fire. Journal of Vegetation Science 3:79-90.

    Keeley, J.E. 1992. Recruitment of seedlings and vegetative sprouts in unburned chaparral. Ecology 73:1194-1208.

    Keeley, J.E. 2000. Chaparral. Pp. 201-253 In M.G. Barbour and W.D. Billings (eds.), North American terrestrial vegetation. Cambridge University Press, New York.

    Keeley, J.E. and S.C. Keeley 1989. Grasslands. Pp. 165-207 In M.G. Barbour and W.D. Billings (eds.), North American terrestrial vegetation. Cambridge University Press, New York.

    Keeley, J.E., P.H. Zedler, C.A. Zammit, and T.J. Stohlgren. 1989. Fire demography, pp. 151-153. In S.C. Keeley (ed.), The California chaparral: paradigms reexamined. Natural History Museum of Los Angeles County, Los Angeles, Science Series No. 34.

    Kirkpatrick J.B. and C.F. Hutchinson 1980. The Environmental Relationships of Californian Coastal Sage Scrub and some of its component communities and species. Journal of Biogeography7:23-28.

    Malanson, G.P. and J.F. O’Leary 1994. The coastal sage scrub-chaparral boundary and response to global change. In J.M. Moreno and W.C. Oechel (eds.), Global change in mediterranean-type ecosystems. Ecological studies 117. Springer-Verlag, New York.

    Miller, P.C. and E. Hajek. 1981. Resource availability and environmental characteristics of Mediterranean-type ecosystems, Pp. 17-41. In P.C. Miller (ed.), Resource use by chaparral and mattoral. Springer-Verlag, New York.

    Minnich, R.A. 1989. Chaparral fire history in San Diego County and adjacent northern Baja California: an evaluation of natural fire regimes and the effects of suppression management, pp. 37-47 In S.C. Keeley (ed.), The California chaparral: paradigms reexamined. Natural History Museum of Los Angeles County, Los Angeles, Science Series No. 34.

    Minnich, R.A. 1995. Fuel-driven fire regimes of the California chaparral, pp. 21-27. In J.E. Keeley and T.Scott (eds.), Brush fires in California wildlands: ecology and resource management. International Association of wildland Fire, Fairfield, Washington.

    Minnich, R.A. and C. Howard. 1984. Biogeography and prehistory of shrublands. Pp. 8-24 In J.J. Devries (ed.), Shrublands in California: literature review and research needed for management. Contribution 191, Water Resources Center, University of California, Davis.

    Minnich, R.A. and R.J. Dezzani 1998. Historical decline of coastal sage scrub in the Riverside-Perris plain, California. Western Birds 29:366-391.

    Moreno, J.M. and W.C. Oechel. 1991. Fire intensity effects on germination of shrubs and herbs in southern California chaparral. Ecology 72:1993-2004.

    O’Leary J.F. and W.E. Westman 1988. Regional disturbance effects on herb succession patterns in coastal sage scrub. Journal of Biogeography. 15:775-786.

    Padgett, P.E., E.B. Allen, A. Bytnerowicz, and R.A. Minnich. 1999. Changes in soil inorganic nitrogenous related to atmospheric nitrogenous pollutants in southern California. Atmospheric Environment.

    Poole, D.K. and P.C. Miller. 1981. The distribution of plant water stress and vegetation characteristics in southern California chaparral. American Midlands Naturalist 105:32-43.

    PSBS. 1995. Western Riverside County Multi-Species Habitat Conservation Plan; Phase1-Information Collection and Evaluation. Prepared for: Western Riverside County Habitat Consortium.

    Rice, S.K. 1993. Vegetation establishment in post-fire Adenostoma chaparral in relation to fine-scale pattern in fire intensity and soil nutrients. Journal of Vegetation Science. 4:115-124.

    Sampson, A.W. 1944. Plant succession on burned chaparral lands in northern California. California Agricultural Experiment Station Bulletin 685:1-144.

    Sawyer, J.O. and T. Keeler-Wolf 1995. A manual of California vegetation. California Native Plant Society, Sacramento.

    Sweeney, J.R. 1968. Ecology of some “fire type” vegetations in northern California, Pp. 111-125. In Proceedings of the tall timbers fire ecology conference. Tall Timbers Research Station. Tallahassee, Florida.

    Timbrook, J., J.R. Johnson, and D.D. Earle. 1982. Vegetation burning by the Chumash. Journal of California and Great Basin Anthropology 4:163-186.

    Tippets, W.E., L.H. Davis, T.A. Stewart, and S.A. Cochrane. 1995. Fire and the Natural Community Conservation Planning (NCCP) Program, pp. 35-40. In J.E. Keeley and T.Scott (eds.), Brush fires in California wildlands: ecology and resource management. International Association of wildland Fire, Fairfield, Washington.

    Tyler, C.M. 1995. Factors contributing to postfire seedling establishment in chaparral: direct and indirect effects of fire. Ecology. 83:1009-1020.

    Vlamis, J.A., E.C. Stone, and C.L. Young. 1954. Nutrient Status of brushland soils in southern California. Soil Science 78:51-55.

    Wells, P.V. 1962. Vegetation in relation to geological substratum and fire in the San Luis Obispo Quadrangle, California. Ecological Monographs 32:79-103.

    Westman 1983. Xeric mediterranean-type shrubland association of Alto and Baja California and the community/continuum debate. Vegetation 52:3-19.

    Zedler, P.H. Vegetation change in chaparral and desert communities in San Diego County, California, pp. 406-430 in D.C. West, H.H. Shugart, and D. Botkin (eds.), Forest succession: Concepts and applications. Springer-Verlag, New York.

    Zedler, P.H., C.H. Gautier, and G.S. McMaster. 1983. Vegetation change in response to extreme events. The effect of a short return interval between fires in California chaparral and coastal scrub. Ecology 64:809-818.

    Zedler, P.H. and C.A. Zammit. 1989. A population based critique of concepts of change in the chaparral, pp. 73-78 In S.C. Keeley (ed.), The California chaparral: paradigms reexamined. Natural History Museum of Los Angeles County, Los Angeles, Science Series No. 34.

    Coastal Sage Scrub

    VEGETATION ASSOCIATION:

    COASTAL SAGE SCRUB

    MAPPED SUBASSOCIATIONS

    Diegan Coastal Sage Scrub

    Riversidean Sage Scrub

    Undifferentiated Coastal Scrub

    OTHER CLASSIFICATIONS:

    Riversidean Upland Sage Scrub (Holland)

    Coastal Sage-Chaparral Scrub (Holland)

    Black Sage Series (Sawyer & Keeler-Wolf)

    California Buckwheat Series (Sawyer & Keeler-Wolf)

    California Buckwheat-White Sage Series (Sawyer & Keeler-Wolf)

    California Sagebrush Series (Sawyer & Keeler-Wolf)

    California Sagebrush-Black Sage Series (Sawyer & Keeler-Wolf)

    Mixed Sage Series (Sawyer & Keeler-Wolf)

    Purple Sage Series (Sawyer & Keeler-Wolf)

    White Sage Series (Sawyer & Keeler-Wolf)

    DATA CHARACTERIZATION

    The data set for coastal sage scrub within the Plan Area is fairly complete.

    BIOGEOGRAPHY

    Coastal sage scrub is represented by several major associations that occur discontinuously from the San Francisco Bay area south to El Rosario in Baja California, Mexico. Some classification systems are based on dominant species (e.g., Holland 1986; Sawyer and Keeler-Wolf; White and Padley 1997), while others are based on geographic location (e.g., Axelrod 1978; Westman 1982). The most commonly cited geographic-based associations include those of Axelrod (Franscican, Diablan, Lucian, Venturan, Diegan, and Riversidian) and Westman (Diablan, Venturan, Riversidean, Diegan, Martirian, and Vizcainan). Coastal sage scrub is found most extensively at lower elevations of coastal southern California, but occurs up to 1300 m in elevation in the Coast Ranges. It transitions into to Mojave desert vegetation to the east and to Sonoran vegetation in Baja California, Mexico ( Axelrod 1978; Westman 1981).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Riversidean sage scrub is distributed throughout Western Riverside County, occupying approximately 159,000 acres (12 percent) of the Plan Area (PSBS 1995) where it is represented by three subassociations: Diegan coastal sage, Riversidian sage scrub and coastal scrub. It occurs from the eastern slopes of the Santa Ana Mountains to elevations in the San Jacinto Mountains less than 5,000 feet. Mapped boundaries between the subassociations are complicated and can be the result of different map source data or differences in dominant species composition. Intergradation between the subassociations near mapped boundaries exists and is dependent on distance inland (east), slope-aspect, elevation, and fire/ disturbance history.

    VEGETATION CHARACTERISTICS

    Coastal sage scrub is dominated by a characteristic suite of low-statured, aromatic, drought-deciduous shrubs and subshrub species. Composition varies substantially depending on physical circumstances and the successional status of the Habitat, however, characteristic species include California sagebrush (Artemisia californica), California buckwheat (Eriogonum fasciculatum), laurel sumac (Malosma laurina), California encelia (Encelia californica), and several species of sage (e.g., Salvia mellifera, S. apiana) (Holland 1986; Sawyer-Wolf 1995). Other common species include brittlebush (E. farinosa), lemonadeberry (Rhus integrifolia), sugarbush (Rhus ovata), yellow bush penstemon (Keckiella antirrhinoides), Mexican elderberry (Sambucus mexicana), sweetbush (Bebbia juncea), boxthorn (Lycium spp.), shore cactus (Opuntia littoralis)coastal cholla (O. prolifera), tall prickly-pear (Opuntia oricola), and species of Dudleya. Sage scrub often is patchily distributed throughout its range (O’Leary 1992). Over a scale of several miles, it can be found in diverse Habitat mosaics with other plant communities, particularly grassland and chaparral, and oak/riparian woodland in more mesic areas. Coastal sage scrub may convert to chaparral or grassland, depending on slope, aspect, climate, fire history, and other physical factors and biological phenomena; conversely, chaparral or grassland areas may convert to coastal sage scrub (Axelrod 1978; White 1995; O’Leary 1995; Allen et al. 1998).

    According to Westman (1982), all coastal sage scrub in Riverside County is contained in the Riversidean subassociation, however, a recent classification of sage scrub in Western Riverside County (White and Padley 1997) has identified seven subassociations (or series) based upon dominant shrub cover. These series include California sagebrush, California buckwheat, California sagebrush-California buckwheat, California sagebrush-white sage, brittlebush, black sage, and deerweed. A number of researchers now are stressing the need for consideration of such variation in Conservation planning and in order to make sound management decisions (DeSimone and Burk 1992; Read 1994; Weaver 1998; White and Padley 1997)

    PHYSICAL ENVIRONMENT

    Coastal sage scrub typically is found on xeric sites, notably steep, south-facing slopes with thin and/or rocky soils. It also is found on exposed sea bluffs, coastal and river terraces composed of coarse alluvial outwash, and coastal dunes (Axelrod 1978). The more open nature of the canopy permits persistence of a diverse herbaceous component of forbs, grasses, and succulents in mature stands than usually is associated with chaparral. It often is interdigitated with chaparral and grassland communities and the distinct boundaries between each can sometimes be difficult to delineate.

    ECOSYSTEM PROCESSES

    As one of the Habitats that has evolved in a Mediterranean climate, it generally is assumed, based upon studies conducted in chaparral, that coastal sage scrub is adapted to periodic wildfire disturbance. These inferences should not be generalized to all coastal sage scrub as there is a number of characteristics in coastal sage scrub that differ from chaparral which could affect fire ecology. Sage scrub’s resilience to periodic wildfire is not completely understood but seems to be a product of the reproductive strategies of the constituent species and the nature of the fire regime. Coastal sage scrub has lower shrub cover, higher volatile oil content, greater cover by herbaceous (or understory) species, shorter duration of nitrogen-fixing species, and more marked variation in post-fire sprouting patterns (Westman et al. 1981). Typically, coastal sage scrub has much less standing biomass and litter accumulation and constituent shrub species also are capable of continual reproduction by seed, unlike chaparral species. There appears to be a difference in recovery strategy dependent upon the geographic locations of the coastal sage scrub and, perhaps, fire regimes (White 1995). In coastal areas, most sage scrub species resprout from below ground root crowns, although there can be substantial seedling germination. This is not the case in inland areas where there is little or no regeneration from sprouting and virtually all recovery is dependent upon seed germination. Habitat recovery in these areas is low. This may be due to an adaptation to a fire interval that was longer than occurs today or that these species once were more effective in recolonizing from seed. Coastal sage scrub assemblages that regenerate primarily by seeding may be inherently more vulnerable to the effects of non-natives than stands that regenerate by sprouting (O’Leary 1990, White 1995) which has management implications which should be considered in Conservation planning.

    Wildfires and controlled burns occur with increasing frequency in southern California (Zedler et. al. 1983). High fire frequency (i.e., short intervals between fires) may alter permanently the floristic composition and structure of a site, including the extirpation of weak resprouting species such as California sagebrush (Malanson and O’Leary 1982). Fires at five to ten year intervals may result in type conversion from chaparral to coastal sage scrub (Keeley 1987; O’Leary, Murphy, Brussard 1992). Type conversion from coastal sage scrub or chaparral to grassland may be accomplished by repeated burning in successive or alternate years (Zedler et. al. 1983).

    AREA PLAN SUBREGIONS

    Riversidean sage scrub is the dominant sage scrub subassociation in the Plan Area, occupying approximately 10.3 percent (136,278 acres) of the Plan Area; it occurs in all of the Area Plans except Eastvale and on March Air Force Base. It is common in the Gavilan Hills, Steele Peak, above Canyon Lake along the San Jacinto River, Wildomar, Lake Skinner, north of Temecula Creek at Vail Lake, Lake Perris, and Lakeview Mountains areas, and along the Peninsular Mountain foothills from the Badlands southeast to the Soboba Indian Reservation. Smaller but significant stands of Riversidian sage scrub also are scattered throughout the Plan Area .

    Diegan coastal sage scrub is distributed in the westernmost portion of the Plan Area within an area generally bounded by the Santa Ana River (south of SR 91), Interstate 15, Temecula Creek (near State Route 79), and the Santa Ana Mountains. The majority occurs southwest of Temecula and also in theTemescal, Greater Elsinore, and Southwest Area Plans. It occupies 1.2 percent (15,805 acres) of the Plan Area.

    Undifferentiated coastal scrub occupies less than one percent (0.46 percent; 7,159 acres) of the Plan Area. This subassociation occurs at the higher elevations of the Plan Area, and was mapped primarily south of Banning-Beaumont Pass, northeast of Soboba Hot Springs, as well as in scattered patches between Bautista Creek and Mountain Center.

    THREATS

    O’Leary (1995) cites several threats to coastal sage scrub rangewide: fragmentation (including edge effects), invasion of non-native species, altered fire cycle, and air pollution. In addition to these, Minnich and Dezzani (1998) include, for Western Riverside County, land clearing, grazing, and competitive exclusion (in conjunction with grazing and altered fire cycles).

    LITERATURE CITED

    Allen, E.B. et al. 1998. What are the Limits to Restoration of Coastal Sage Scrub in Southern California? Interface Between Ecology and Land Development in California, Second Symposium. J.E. Keeley, M.B Keeley, and C.J. Fotheringham, eds. International Associate of Wildland Fire, Fairfield, WA.

    Axelrod. D.I. 1978. The Origin of Coastal Sage Vegetation, Alta and Baja California. American Journal of Botany 65(10): 1117-1131.

    DeSimone, S. 1995. California’s Coastal Sage Scrub. Fremontia 23(4): 3-8.

    Keeley, J.E. 1987. Role of fire in seed germination of woody taxa in California chaparral. Ecology 68: 434-442.

    Malanson, G.P. and J.F. O’Leary. 1982. Post-fire regeneration strategies in California coastal sage shrubs. Oecologia 53:355-358.

    Minnich, R. A. and R.J. Dezzani. 1998. Historical Decline of Coastal Sage Scrub in the Riverside-Perris Plain, California. Western Birds 29(4): 366-391.

    O’Leary, J.F. 1990. Post-fire diversity patterns in two subassociations of California coastal sage scrub. Journal of Vegetation Science 1: 173-180.

    O’Leary, J.F. 1990. Californian Coastal Sage Sscrub: General Characteristics and Considerations for Biological Conservation. Pages 24-41 in Endangered Plant Communities of Southern California, Alan Schoenherr, ed. Southern California Botanists, Special Publications No. 3

    O’Leary, J.F., Murphy, D., And Brussand, P. 1992. The coastal sage scrub community Conservation planning region: An NCCP special report. Natural Community Conservation Planning/Coastal Sage Scrub Special Report 2.

    O’Leary, J.F. 1995. Coastal Sage Scrub: Threats and Current Status. Fremontia 23(4): 27-31Kirkpatrick J.B and C.F. Hutchinson. 1977. The community composition of Californian coastal sage scrub. Vegetation 35:21-33

    Pacific Southwest Biological Services (PSBS). 1995. Draft MSHCP Strategy Report Vegetation Mapping. Prepared for the Western Riverside County Habitat Consortium. February 20.

    Read, E.A. 1994. The Importance of Community Classification to Mitigation and Restoration of Coastal Sage Scrub. Restoration Ecology 2: 80-86.

    Weaver, K. L. 1998. Coastal Sage Scrub Variations of San Diego County and Their Influence on the Distribution of the California Gnatcatcher. Western Birds 29: 392-405.

    Westman, W.E. 1981. Diversity Relations and Succession California Coastal Sage Scrub. Ecology 62(1): 170-184.

    Westman, W.E., J.F. O’Leary, and G.P. Malanson. 1981. The effects of fire intensity, aspect, and substrate on post-fire growth of Californian coastal sage scrub. P: 151-179 in N.S. Margaris and H.A. Mooney, editors. Components of Productivity of Mediterranean-climate Regions: Basic and Applied Aspects. Dr. W. Junk, The Hague, Netherlands.

    Westman, W.E. 1982. Coastal Sage Scrub Succession. Pages 91-99 in: Proceedings of the Symposium on Dynamics and Management of Mediterranean-Type Ecosystems. C.E. Conrad and W.C. Oechel, technical coordinators.

    Westman, W.E. 1983. Xeric Mediterranean-type Shrubland Associations of Alta and Baja California and the Community/Continuum Debate. Vegetation 53: 3-19.

    White, S.D. 1995. Disturbance and Dynamics in Coastal Sage Scrub. Fremontia 23(4): 9-16.

    White, S.D. and W. D. Padley. 1997. Coastal Sage Scrub Series of Western Riverside County, California. Madrono 44(1): 95-105

    Zedler, P.H., C.R. Gautier, and G.S. McMaster. 1983. Vegetation change in response to extreme events: the effects of a short interval between fires in California chaparral and coastal sage scrub. Ecology 64: 809-818.

    Dairy and Livestock Feedyards

    VEGETATION ASSOCIATION:

    DAIRY AND LIVESTOCK FEEDYARDS

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Because dairy and livestock farms are registered with the County and State, and information regarding land use is recorded with the County by parcel, the mapping of this Vegetation Community is considered to be very accurate. There may be areas of open space which are under grazing pressures but still support native or slightly-altered vegetation communities and are not mapped under this category.

    BIOGEOGRAPHY

    Dairy and livestock feedyards occur in nearly all human-inhabited areas of the world.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Milk is Riverside County’s largest agricultural commodity, yielding $335 million in 1998 (CDFA 1999). In 1999, 116,131 cows on 106 dairies were present in Riverside County (CDFA 2000). According to the vegetation classification of this planning process, dairy and livestock feedyards occur in 123 separate locations covering 5853 acres of the Plan Area. The largest areas of dairy and livestock feedyards are located north of San Jacinto and north of Juniper Flats in the communities of Lakeview, Mystic Lake, Nuevo, southeast Perris, Eastvale, and Lake Norconian off of Bellegrave Avenue. Other occurrences include: along Tenaja Road south of the Santa Rosa Plateau, along Wilson Creek, in the Cahuilla Valley, south of Black Mountain off of De Portola Road, in the area of Canyon Lake off of Newport Road, in the City of Menifee, in and around the Domenigoni Valley off SR-79, in the community of Winchester off of Simpson Road, in Diamond Valley west of the Santa Rosa Hills, in Moreno Valley, in Cherry Valley, between Lake Matthews and Gavilan Plateau south of Cajalco Road, in the community of Woodcrest, in Norco, and in Glen Avon. The SR-91 corridor through the City of Riverside also contains a number of small dairy and livestock farms.

    VEGETATION CHARACTERISTICS

    The type of vegetation occurring in dairy and livestock feedyards varies widely due to location and farming practices. Pasture lands typically consist of a mix of perennial grasses and legumes that provide 100 percent cover (Zeiner 1988). The height of the vegetation varies by season and livestock type (Zeiner 1988). Wetter areas often contain more weeds which can grow several feet tall (Zeiner 1988). In southern California, Bermuda grass (Cynodon dactylon) often dominates pasture lands (Zeiner 1988). The dominant weedy vegetation which persists in a feedyard, or pastoral community, are often either low-growing, distasteful, and/or bear spines (Holland and Keil 1995). Often weedy plants which establish in these areas reproduce vegetatively through rhizomes (Holland and Keil 1995). Feedyards may also support isolated patches of annual or native grasslands (Holland and Keil 1995).

    PHYSICAL ENVIRONMENT

    Although dairy and livestock feedyards have been established under a variety of physical conditions, some general statements can be made. Pastures generally are limited to flat ground or gently sloping hills and often are placed on soils not suitable for other crops (Zeiner 1988). Many farmers also irrigate their pasture lands and rotate pasture use with crops (Zeiner 1988).

    ECOSYSTEM PROCESSES

    It is thought that historically there were few native herbivore species in California and for that reason native vegetation is not well-adapted to grazing (Holland and Keil 1995). Grazing was important in creating an environment that allowed exotic grasses to become established in California, even though in many areas non-native grasses now perpetuate themselves and exclude natives without grazing pressures (Holland and Keil 1995).

    Other ecosystem processes, such as fire and regeneration of native plant communities, were not reviewed for this account.

    THREATS

    No threats to dairy and livestock production have been identified in the literature review for this account. It can be stated that the main threat to dairy and livestock production is economic competition for land.

    LITERATURE CITED

    California Department of Food and Agriculture. 1999. California Agriculture Resource Directory State of California, Sacramento, California.

    California Department of Food and Agriculture. 2000. California Dairy Statistics. State of California, Sacramento, California.

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company. Dubuque, Iowa.

    Zeiner, David C. 1988. Pasture. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 128-129.

    Field Croplands

    VEGETATION ASSOCIATION:

    FIELD CROPLANDS

    MAPPED SUBASSOCIATIONS

    None

    BIOGEOGRAPHY

    Field croplands occur in nearly all human-inhabited areas in the world today, although, historically, the indigenous peoples of Western Riverside County were not known to grow crops.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Field croplands are mapped extensively throughout the Plan Area. The largest areas are: around SR-371 in the vicinity of Anza; in an east-west strip from Murrieta Hot Springs; through French Valley; Antelope Valley; Paloma Valley; Menifee Valley; Winchester; Domenigoni Valley; to West Hemet; and the Diamond Valley area; and in Eastvale. More medium-sized tracts of croplands are mapped around SR-79 in the Radee Valley east of Vail Lake, in Wolf Valley and Pauba Valley, around Murrieta north of Temecula, in Tres Cerritos in the San Jacinto Valley, around the communities of Lakeview, Nuevo, Romoland and Perris, in Mockingbird Canyon and Mead Valley, and in the area of Beaumont and Banning Bench. Small and scattered occurrences are in the Terwilliger and Cahuilla Valleys, along Wilson Creek in the Lewis Valley from the community of Sage up to Black Mountain, on the Santa Rosa Plateau, east of the Gavilan Hills, along I-15 through Alberhill and Corona in the foothills of the Santa Ana mountains, south of SR-91 in the City of Riverside, along the Santa Ana River in the communities of Mira Loma, Glen Avon and Jurupa, and along San Timoteo Canyon.

    VEGETATION CHARACTERISTICS

    Crop vegetation varies widely from ten-foot tall corn to two-inch tall strawberries (Zeiner 1988). Some crops are planted in rows, whereas others form dense stands (Zeiner 1988). Some croplands support annual plants which can be rotated, whereas others are long-term monocultures (Zeiner 1988). The leading crops in Riverside County are dates, hay, and bell peppers (CDFA 1999). According to 1992 statistics, 6200 acres of barley, 600 acres of dried beans, 1000 acres of corn, 12100 acres of cotton, 7400 acres of oats and 29,000 acres of wheat were planted in Riverside County (CASS 1993a). The Perris-Hemet area is noted for its production of cantaloupes, other melons, carrots, onions and potatoes (CDFA 1988). In 1987 12,300 acres of lettuce, 700 acres of onions, 4600 acres of potatoes, 5060 acres of asparagus, 1700 acres of broccoli, 5300 acres of cantaloupe, 2,000 acres of carrots, 800 acres of cauliflower, 100 acres of celery and 4,800 acres of sweet corn were planted in Riverside County (CDFA 1988).

    PHYSICAL ENVIRONMENT

    Crop rotations of a five to seven-year period are common and switch between annual and perennial species (Zeiner 1988). This method serves to enhance soil productivity by retaining nutrients (Zeiner 1988). Common cultivation techniques used on croplands act to repeatedly disturb the soil (Holland and Keil 1995).

    ECOSYSTEM PROCESSES

    Many annual crop species are self-fertile or set seeds apomictically (Holland and Keil 1995). Seeds are also disseminated by machinery and some species may have seeds which can lay dormant in a seed bank (Holland and Keil 1995). Some weedy species common to croplands exclude growth of nearby plants; e.g., wild oat (Avena fatua) produces allelopathic chemicals and sow-thistles (Sonchus spp.) develop a basal rosette which shades out other seedling plants (Holland and Keil 1995). For these reasons, within a few years of converting a natural Habitat area to cropland, native plants are effectively eliminated (Holland and Keil 1995). Native perennials are especially susceptible to disturbance (Holland and Keil 1995). Abandoned crop lands seldom re-establish dense native plant communities and often native species are absent (Holland and Keil 1995).

    THREATS

    No threats to crop production have been identified in the literature review for this account. It can be stated that the main threat to crop production is economic competition for land.

    LITERATURE CITED

    California Agricultural Statistics Service. 1993a. California Field Crop Statistics 1983-1992. County Data 1991-92. Sacramento, California.

    California Department of Food and Agriculture. 1988. California Vegetable Crop Statistics: County Data 1986-87. Sacramento, California.

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company. Dubuque, Iowa.

    Zeiner, David C. 1988. Cropland. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 138-139.

    Freshwater Wetlands

    VEGETATION ASSOCIATION:

    FRESHWATER WETLANDS

    MAPPED SUBASSOCIATIONS

    Coastal and Valley Freshwater Marsh, Cismontane Alkali Marsh, Marsh (undifferentiated), Wet Montane Meadow and Open Water.

    STATUS:

    State: California Fish and Game Wetlands

    Federal: U.S. Army Corps of Engineers Jurisdictional Waters of the United States

    DATA CHARACTERIZATION

    Much of the information provided in this section was obtained from published papers written by Holland and Keil (1995), Barbour and Major (1977), Sawyer and Keeler-Wolf (1995), Kramer (1988) and Grenfell (1988). Distribution data for the vegetation association and subassociations was obtained primarily from the University of California, Riverside GIS database.

    Because of the regional scale of the mapping and the small size of most marsh and wet meadow Habitats, many areas supporting marsh and wet meadow Habitats probably were overlooked. Most of the potential wet montane meadow Habitat is located within National Forest Service lands that have not been extensively surveyed and mapped; therefore, the distribution of wet montane meadow may be greater than indicated by the current data. In addition, marsh Habitat develops quickly in freshwater areas and the occurrence of many small marsh areas associated with urban and agricultural land uses, as well as small patches within native riparian Habitats, may be more frequent than indicated in this study.

    BIOGEOGRAPHY

    Marsh Habitats typically occur at elevations ranging from sea level to 2,000 meters in association with stream channels, ponds and reservoirs (Sawyer and Keeler-Wolf 1995; Holland and Keil 1995). Montane meadows typically occur at elevations above 1,000 meters (Stephenson and Calcarone 1999) within the montane North Coast Ranges; montane Central Coast Ranges; montane Klamath Ranges; montane Cascade Range; montane Sierra Nevada; montane Transverse Ranges; montane Peninsular Ranges; montane Warner Range; and montane White, Inyo and Sweetwater Ranges (Sawyer and Keeler-Wolf 1995).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Marsh (coastal and Valley, cismontane alkali and undifferentiated ) Habitat is scattered sparsely over the western Riverside region, occupying approximately 0.1 percent (1,732 acres) of the Plan Area. Cismontane alkali marsh was mapped in two general localities along Cahuilla Creek south of Anza and upstream from Lake Mathews along the Colorado River aqueduct. The mapped locations of coastal and Valley freshwater marsh are more widely distributed, with the largest acreage occurring in the Prado Basin in the Santa Ana River Valley. Smaller patches are located upstream on the Santa Ana River near Pedley, north of Lake Elsinore in Walker Canyon, near San Jacinto, along the shores of Lake Skinner and Vail Lake, and adjacent to the cismontane alkali marsh on Cahuilla Creek. Undifferentiated marsh was mapped in three locations including the shore of Lake Mathews, near Mystic Lake, and upstream from Vail Lake along Temecula Creek in the Aguanga Valley. Wet montane meadows were mapped only in the San Jacinto Mountains in the San Bernardino National Forest, primarily within the vicinity of Hemet Lake. Open water was mapped at Vail Lake, Lake Skinner, Eastside Reservoir, Lake Perris, Mystic Lake, Canyon Lake, Lake Elsinore, Lee Lake, Lake Mathews, Hemet Lake, portions of the San Jacinto River, and portions of the Santa Ana River as well as various small ponds, private reservoirs and portions of stream channels.

    VEGETATION CHARACTERISTICS

    Because all freshwater wetlands experience periodic flooding, the vegetation is adapted to an anaerobic environment (Kramer 1988). Many freshwater marsh and wet meadow species are able to reproduce through rhizomes (Barbour and Major 1977; Weller 1981). Vegetation heights can vary from a few millimeters to two meters (Kramer 1988).

    SPECIES COMPOSITION

    Coastal and Valley Freshwater Marsh (including undifferentiated Marsh). Coastal and Valley freshwater marsh typically is dominated by perennial monocots up to two meters in height (Kramer 1988). This Habitat type includes cattails (Typha spp.), bulrush (Scirpus spp.), sedges (Carex spp.), spike rushes (Eleocharis spp.), flatsedges (Cyperus spp.), smartweed (Polygonum spp.), watercress (Rorippa spp.) and yerba mansa (Anemopsis californica) (Barbour and Major 1977; Holland and Keil 1995; Sawyer and Keeler-Wolf 1995). Rooted aquatic plant species with floating stems and leaves also may be present, such as pennywort (Hydrocotyle spp.), water smartweed (Polygonum amphibium), pondweeds (Potamogeton spp.) and water-parsley (Oenanthe sarmentosa) (Holland and Keil 1995).

    Cismontane Alkali Marsh. Typical cismontane alkali marsh species include yerba mansa, saltgrass (Distichlis spicata), alkali-heath (Frankenia salina), cattails (Typha spp.), common pickleweed (Salicornia virginica), rushes (Juncus spp.), marsh flea-bane (Pluchea odorata) and sedges (Carex spp.) (Holland 1986).

    Wet Montane Meadow. Montane meadows are dominated by sedges (Carex spp.), spike rushes (Eleocharis spp.), and bulrushes (Scirpus spp.), but also contain perennial and biennial herbs (e.g.Oenothera spp., Polygonum spp., Lupinus spp., Potentilla spp., and Sidalcea spp.) and grasses (e.g.Agrostis spp., Deschampsia spp. and Muhlenbergia spp.) (Barbour and Major 1977; Sawyer and Keeler-Wolf 1995; Stephenson and Calcarone 1999). Wet montane meadows that dry out by mid-summer have a higher percentage of perennial grasses than meadows that remain moist during the entire growing season (Holland and Keil 1995).

    Open Water. Open water Habitat typically is unvegetated due to a lack of light penetration. However, open water may contain suspended organisms such as filamentous green algae, phytoplankton (including diatoms) and desmids (Grenfell 1988). Floating plants such as duckweed (Lemna spp.), water buttercup (Ranunculus aquatilis) and mosquito fern (Azolla filiculoides) also may be present (Holland and Keil 1995).

    PHYSICAL ENVIRONMENT

    Herbaceous freshwater wetlands, including freshwater marsh and wet montane meadow, occur in both flowing and still water. The bodies of water range from lakes and reservoirs to small pools. Periods of hydrology can vary from perennial to seasonal. Variables that can affect herbaceous wetlands include rate of water flow, fluctuations in water level, water depth, water and air temperatures, pH and dissolved salts, depth and nature of bottom sediments, organic content of the water and past history of the body of water (Holland and Keil 1995).

    Where freshwater marsh and wet montane meadow Habitats occur together, the water depth demarcates the boundary of the two Habitats (Ratliff 1988). Freshwater marsh typically has a water table at or above the soil surface and occurs along the margins of lakes, ponds, reservoirs and slow-flowing stream channels (Barbour and Major 1977). Following spring runoff, wet montane meadows typically have a water table at or near the soil surface during the growing season (Ratliff 1988).

    Wet montane meadows generally occur at elevations above 5,900 feet (Ratliff 1988). Wet montane meadow Habitat can be characterized by two physical conditions: fine-textured (i.e., clay) and richly organic surface soil; and a shallow water table (usually within two feet of the soil surface) during mid-summer. Meadows generally occur on gentle gradients and relatively impervious bedrock in combination with adequately-sized drainage areas (Stephenson and Calcarone 1999) as well as along the margins of stream channels (Barbour and Major 1977). The soils of wet montane meadows are typically more aerated than marsh soils (Holland and Keil 1995).

    Wet montane meadows include three hydrologic types: lotic, sunken concave and hanging sites. Lotic sites have an input flow and a downstream flow and consist of topographic basins with a slight slope. Percolation is low to zero at lotic sites due to the saturated or impermeable substrate. Sunken concave sites receive an input flow but the primary outflow is limited to evapotranspiration. The hydrologic source for hanging sites (seeps and springs) consists of underground hydrostatic flows. Hanging sites frequently occur on steep slopes and the main outflow is surface runoff (Ratliff 1988). Many montane meadows are wet only during the spring and early summer, becoming dried-out by mid-summer (Holland and Keil 1995).

    The boundary between open water Habitat and emergent wetlands is generally at a depth of two meters (6.6 feet) (Kramer 1988). Open water Habitat includes inland depressions and stream channels containing standing water. Depth may vary from hundreds of meters to a few centimeters. This Habitat includes ponds, lakes and reservoirs. Within the relatively calm waters of ponds and lakes, temperatures vary by depth and by season, and light penetration depends on water turbidity. The oxygen content of ponded water is relatively low due to the small proportion of water that is in contact with the air and because decomposition of organic materials is occurring on the substrate below (Grenfell 1988).

    ECOSYSTEM PROCESSES

    Bottom sediments and runoff are the major sources of inorganic and organic nutrients. The decomposition of organic sediments may be the most important source of nutrients in still or slowly-flowing water; however, the decomposition of organic materials can be inhibited by acidic pH conditions or low temperatures, resulting in low nutrient availability (Holland and Keil 1995).

    The presence of organic materials promotes the growth of microorganisms, including bacteria. The respiration of these decomposers substantially reduces the oxygen supply. Thus, the decomposition of organic matter can deplete the supply of oxygen and the bottom sediments of a nutrient-rich lake or pond may be completely anaerobic. As oxygen is a requirement for root growth, many plant species are unable to grow in anaerobic conditions (Holland and Keil 1995).

    COMMUNITY RELATIONSHIPS

    Freshwater herbaceous wetlands and open water Habitats often occur in conjunction with riparian Habitats and upland Habitats. Freshwater herbaceous wetlands and open water areas provide food, water, cover and reproduction Habitat for mammals, birds, reptiles and amphibians (Weller 1981; Grenfell 1988; Kramer 1988). Within open water, zooplankton or animal organisms (including copepods, cladocerans and rotifers) may graze upon the minute plants (Grenfell 1988). (The reader should refer to the species accounts for specific information about sensitive wildlife and plant species and riparian Habitats.)

    THREATS

    Threats to herbaceous freshwater wetlands and open water include pollution, grazing, and conversion to other land uses, including agriculture. Pollution in the form of fertilizer, pesticides and untreated sewage can increase the amount of organics entering an aquatic ecosystem. This can increase the turbidity of the water and reduce the oxygen levels in the water, which can kill the vegetation (Holland and Keil 1995).

    Grazing of freshwater marshes by domestic animals can result in trampling of the soil, invasion by exotic plant species and erosion. Grazing of wet montane meadows by domestic animals (particularly sheep) can severely disturb the soils, resulting in bare patches that are invaded by exotic plant species. Moreover, grazing can result in population shifts of the palatable and unpalatable plant species (Holland and Keil 1995).

    LITERATURE CITED

    Barbour, M.J. and J. Major. 1977. Terrestrial Vegetation of California. Wiley Press. New York, New York. 1002 pp.

    Grenfell, W.E., 1988 Lacustrine in A Guide to Wildlife Habitats of California. California Department of Forestry and Fire Protection. 166 pp.

    Holland R.F. 1986. Preliminary Descriptions of the Terrestrial Natural Communities of California. State of California Department of Fish and Game, Nongame-Heritage Program, Sacramento, California.

    Holland, V.L. and D.J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company, Dubuque, Iowa. 516 pp.

    Kramer, G. 1988. Fresh Emergent Wetland in A Guide to Wildlife Habitats of California. California Department of Forestry and Fire Protection. 166 pp.

    Ratliff, R.D. 1988. Wet Meadow in A Guide to Wildlife Habitats of California. California Department of Forestry and Fire Protection. 166 pp.

    Sawyer, J.O. and T. Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, California. 471 pp.

    Stephenson, J.R. and G.M. Calcarone. 1999. Southern California Mountains and Foothills Assessment: Habitat and Species Conservation Issues. General Technical Report GTR-PSW-172. Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture, Albany, California. 402 pp.

    Weller, M.W. 1981. Freshwater Marshes: Ecology and Wildlife Management. University of Minnesota Press, Minneapolis, Minnesota. 146 pp.

    Grasslands

    VEGETATION ASSOCIATION:

    GRASSLANDS

    VEGETATION SUBASSOCIATIONS:

    Valley and Foothill Grasslands, Non-native grasslands

    DATA CHARACTERIZATION

    The distribution of Valley and foothill grasslands is larger than that shown in the MSCHP mapping. Because native grasslands are typically small, difficult to differentiate from non-native grasslands using aerial photographs, and require field verification to identify, the true extent of native grasslands in the Plan Area remains unknown. A cost-effective method to estimate the location of existing native grasslands within the Plan Area would be to digitize maps of clay soils from the Riverside County soil survey (Knecht 1971) and combine these with the mapped extent of existing grasslands. This mapping also would be helpful to determine locations that may support Valley and foothill grasslands given future changes in the landscape. Without additional data it would be difficult to evaluate preserve designs in regard to the Conservation of native grasslands.

    With the exception of differentiating potential native grasslands, the distribution of non-native grasslands is adequately represented. Grasslands are relatively simple to differentiate from other vegetation types using aerial photography. Some errors may be present due to recent type conversion or misinterpretation of agricultural fields.

    BIOGEOGRAPHY

    The historic distribution of Valley and foothill grasslands has been described from central California to Baja California Norte, from the coast into the Sierra foothills (Keeley 1990). Valley and Foothill grasslands have an elevational range from sea level to about 1200-1700 m (Heady 1977; Sawyer and Keeler-Wolf 1995). These native grasslands are widely distributed but are considered uncommon due to the long history of grazing and agricultural usage in lands previously supporting native grasslands. Non-native grasslands occur widely throughout the Mediterranean-type climate portion of California and Baja California, Mexico.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Grasslands occur throughout most of Western Riverside County cover approximately 11.8% (154,421 acres) of the Plan Area. The only Valley and foothill grasslands mapped within the Plan Area are distributed over approximately 2,736 acres (0.2 %) of the Plan Area on the Santa Rosa Plateau. Non-native grasslands occur throughout the majority of the Plan Area (11.6%), usually within close proximity to urbanized or agricultural land uses. Large patches of non-native grasslands occur in the Riverside east area near March Air Reserve base, Lake Mathews, Lake Perris, Lake Elsinore, near Banning, Cahuilla, and in the Terwilliger Valley south of Anza. Non-native grasslands are not common or extensive in high elevation areas that support large blocks of native vegetation (e.g., the Santa Ana and San Jacinto Mountains). Because much of the Plan Area was mapped with the use of aerial photo interpretation, some Valley and foothill grasslands were included within the non-native grassland category. Additional Valley and Foothill grasslands are known from the Santa Ana Mountains (MARKEL and Associates 1982 ), the northwestern portion of the Agua Tibia Wilderness (Boyd and Banks 1995), and the Gavilan Hills (Boyd 1983). Additional ground truthing of areas supporting grasslands and clay or deep, well developed soils is necessary to determine the location of other important stands of Valley and foothill grasslands.

    VEGETATION CHARACTERISTICS

    The nature and extent of grasslands in California have long been subject to varying opinions ranging from Clements’ (1920) view that much of the extent of contemporary non-native grasslands formerly was occupied by native perennial grasses, to Cooper’s (1922) more restrictive view where most of the area currently supporting non-native grasslands was formerly shrublands or oak woodlands (cited in Hamilton 1997). Lacking definitive evidence to support either assertion most reviews of California grasslands have described two grassland associations: (1) a non-native dominated, primarily annual grassland; and (2) a native dominated, perennial grassland (Heady 1977; Keeley 1989; Sims and Risser 2000). These and other authors have suggested that the extant perennial grasslands represent relictual stands of “pristine” native grasslands. Hamilton (1997) has provided a critical review of past research in native grasslands in California, positing that most of the current distribution of non-native grasslands in central and southern California historically was not extensively perennial grasslands but shrublands, woodlands or desert scrub vegetation. Although there is debate about the distribution and pristine nature of native grasslands it is known that areas supporting native grasses in southern California are currently uncommon and support high diversity of both native and sensitive plant species.

    Within the Western Riverside MSHCP Plan Area perennial and annual grasslands were mapped as Valley and foothill grasslands or non-native grasslands (PSBS 1995). Currently used synonyms for these types include Valley needlegrass grassland (Holland 1986), purple needlegrass and foothill needlegrass series for the native grasslands, and California annual grassland series for the non-native grasslands (Sawyer and Keeler-Wolf 1995).

    SPECIES COMPOSITION

    Valley and Foothill Grasslands. Valley and foothill grasslands occur in a variety of forms ranging from scattered perennial bunch grasses (typically Nassella pulchraor N. lepida) with high abundance of non-native grasses and forbs to stands dominated by native perennial grasses in an assemblage of geophytes (plants with underground bulbs or corms), and herbaceous annual species (Holland 1986; Keeley 1989; Sims and Risser 2000). Melica spp., Leymus spp, beard grass (Bothriochloa barbinodis), Muhlenbergia spp., and other native perennial grasses may also occur in Valley and foothill grasslands but typically not in high abundance (Holland 1986; Keeley 1989; Sawyer and Keeler-Wolf 1995; pers. obs.). The percent cover of perennial grasses in undisturbed Valley and foothill grasslands has been estimated at 50 to 75% cover (Keeley 1989). Some researchers have defined current stands of native grasslands as having as little as 10% percent cover of native grasses (California Department of Fish and Game; Keeley 1993). Valley and foothill grasslands usually support substantial cover of non-native grasses (see examples below) and exotic forbs but at lower abundance than in non-native grasslands. Valley and foothill grasslands also support native geophytes, including representatives from the following species or genera: onion (Allium spp.), wild- celery (Apiastrum angustifolium)common golden star (Bloomeria crocea), Brodiaea spp., Calochortus spp., blue dicks (Dichelostemma capitata), Muilla spp., blue-eyed grass (Sisyrinchium bellum), and Dudleya spp. (Holland 1986; Keeley 1989; pers. obs.). Native herbaceous plants commonly found within Valley and foothill grasslands include yellow fiddleneck (Amsinckia menziesii), Calandrinia spp., common calyptridium (Calyptridium monardum)suncup (Camissonia spp.), owl’s-clover (Castilleja spp.), Chinese houses (Collinsia heterophylla), Cryptantha spp. Delphinium spp. California poppy (Eschcholzia californica), Gilia spp., tarweed (Hemizonia spp.)coast goldfields (Lasthenia californica), common tidy-tips (Layia platyglossa), Linanthus spp., Lomatium spp., Lotus spp., Lupinus spp., Microseris spp., Plagiobothrys spp., Sanicula spp., checker mallow (Sidalcea malvaeflora), and clover (Trifolium spp.) (Holland 1986; Keeley 1989; pers. obs.). Because this vegetation is primarily herbaceous it is relatively simple structurally, rarely exceeding one meter in height (Sawyer and Keeler-Wolf 1995). Mapped native grasslands support early successional sub-shrub and suffrutescent species (e.g., Astragalus spp., Ericameria spp., Eriogonum spp., gum plant (Grindelia camporum), Hazardia spp., golden bush (Isocoma menziesii), virgate cudweed aster (Lessingia filaginifolia), and deerweed (Lotus scoparius) but these species occur at low abundances.

    Few descriptions of the distribution and species composition of perennial grasslands in Western Riverside County appear in peer-reviewed publications. Many technical documents from Conservation and development projects exist with descriptions of native grasslands in the Plan Area (e.g., Lathrop and Thorne (1985); MARKEL and Associates 1982; Boyd and Banks 1995; Boyd 1983). Although these and other technical documents have not been peer-reviewed and may be uneven in quality, they are the best information available for specific places within the Plan Area.

    The Valley and foothill needlegrass grassland within the Agua Tibia Wilderness area occur on cobbly clay soils and reportedly supports dense stands of foothill stipa (Nassella lepida) and nodding needlegrass (N. cernua) with lower abundance of other native grasses, such as California melic (Melica imperfecta), junegrass (Koeleria macrantha)Malpais bluegrass (Poa secunda), Pacific fescue (Vulpia microstachys), and six-weeks fescue (V. octoflora) (Boyd and Banks 1995). These grasslands are relatively rich in native geophyte and annual species, and support “well established” stands of non-native grasses and herbs (Boyd and Banks 1995). Additional unmapped Valley and foothill grasslands are reported within the southwestern portion of the Santa Ana Mountains including Elsinore Peak, Bluewater Flats, and Oak Flats area (MARKEL and Associates 1982). These grasslands were not surveyed for species composition but purple needlegrass was generally reported as the dominant native grass with occasional presence of geophyte species (MARKEL and Associates 1982). In a floral inventory of the Gavilan Hills, Boyd (1983) described a native grassland as dominated by foothill stipa and California melic with lower cover of California buckwheat, saw-toothed goldenbush (Hazardia squarrosa), virgate cudweed aster, and several herbs found on clay soils (e.g., Munz’s onion [Allium munzii], Cleveland’s shooting-star [Dodecatheon clevelandii], and Palmer’s grappling-hook [Harpagonella palmeri]).

    Two additional native grassland types are known to occur within the western Riverside MSHCP Plan Area but were not specifically mapped: alkali playa and cismontane alkali marsh. Alkali playas support a native annual grassland type dominated by little barley (Hordeum intercedens) and annual hairgrass (Deschampsia danthonioides). Cismontane alkali marsh and disturbed alkali areas occasionally support small stands of saltgrass (Distichlis spicata) or beardless wild ryegrass (Leymus tritichoides). These grasslands were not specifically identified as part of the regional mapping effort but occur in small patches within their more broadly defined Habitats. These stands are addressed within the vernal pool and alkali playa, and cismontane alkali marsh descriptions.

    Non-Native Grasslands. Non-native grasslands primarily are composed of annual grass species introduced from the Mediterranean basin and other mediterranean-climate regions with variable presence of non-native and native herbaceous species (Baker 1989; Mack 1989). Species composition of non-native grasslands may vary over time and place based on grazing or fire regimes, soil disturbance, and annual precipitation patterns (McNaughton 1968; Heady 1977; Keeley 1989). Non-native grasslands typically produce deep layers of organic matter which is inversely related to the abundance of non-native and native forbs (Heady 1956a). Non-native grasslands are likely to be dominated by several species of grasses: slender oat (Avena barbata)wild oat (A. fatua), fox tail chess (Bromus madritensis), soft chess (B. hordeaceus), ripgut grass (B. diandrus), barley (Hordeum spp.), rye grass (Lolium multiflorum)English ryegrass (L. perrene), rat-tail fescue (Vulpia myuros), Mediterranean schismus (Schismus barbatus) that have evolved to persist in concert with human agricultural practices (Jackson 1985 in Sims and Risser 2000). Non-native grasslands also typically support an array of annual forbs from the Mediterranean-climate regions (e.g., red-stemmed filaree [Erodium cicutarium]), broad-loabed filaree [E. botrys], mustard [Brassica spp.], short-podded mustard [Hirschfeldia incana]wild radish [Raphanus sativus], Centaurea spp., Italian thistle [Carduus pycnocephalus]artichoke thistle [Cynara cardunculus]common catchfly [Silene gallica]), Medicago spp., and Hypochaeris spp) (Keeley 1989; pers. obs.). Low abundances of native species are sometimes present within non-native grasslands. These species usually include disturbance specialists with several different growth forms: subshrubs (e.g., Lotus spp., Eriogonum spp., Lessingia spp, Isocoma, spp., Ericameria spp.); succulents (Opuntia spp.); perennial geophytes (e.g., Dichelostemma capitata); and herbaceous annuals (e.g., doveweed (Eremocarpus setigerus)vinegar weed (Trichostemma lanceolatum), and tarweed (Hemizonia spp). (Holland 1986; Sawyer and Keeler-Wolf 1995; Keeley 1989).

    PHYSICAL ENVIRONMENT

    Because of a lack of agreement on the original Habitat and distribution of perennial (i.e., native) grasslands in central and southern California, and because annual grasslands probably have developed from some form of disturbance (e.g., grazing, discing, fire), the existing descriptions of the physical environment associated with native and non-native grasslands necessarily are vague. Discussion of the site preferences for this review consequently are limited to where perennial grasslands are currently known to occur rather than where they should occur or have occurred in the past.

    Valley and Foothill Grasslands. Perennial grasslands have been described as occurring in many topographic locations within California (Sawyer and Keeler-Wolf 1995), with affinities toward more mesic north and east slope-aspects within a limited region (Keeley 1991; 1993). It is more likely, however, that perennial grasslands are closely associated with soil characteristics particular to a local area. Statewide, perennial grasslands occur on a large variety of soil series; however, most of these support oak woodlands and other vegetation types (Barry 1972; Heady 1977). Most research has provided descriptive accounts of the soil conditions supporting perennial grasslands as deep, brown, fertile and having high clay content (Adams 1964; Heady 1977; Keeley 1989; Sims and Risser 2000). Soil affinities for Valley and foothill grasslands have been established within southeastern Ventura County where soil depth and percentage clay particles were positively related, and percentage rock was negatively related to percentage cover of native perennial grasses (Keeley 1993). Few soil chemical studies have been conducted within Valley and foothill grasslands and no strong relationship has been established between native grasses and soil nutrients (nitrogen, potassium or phosphate) (Keeley 1993). As part of this research, Keeley (1993) found a high fidelity of native grasslands to three soil series: Diablo, Santa Lucia, and San Andreas. None of these soil series occur within the MSHCP Plan Area; however, other series with clayey structure are known from Western Riverside County (e.g., Altamont, Bosanko, Las Posas) (Knecht 1971). Another consistent theme in the physical environment supporting perennial grassland is that they occur upon soils that remain saturated during the winter and become completely dry during summer months (Keeley 1989; Holland 1986).

    Non-Native Grasslands. Most non-native grasslands likely have developed as a result of past agricultural or urban development related activities including discing, brushing, grading, or overgrazing of native Habitats. Because non-native grasslands generally are associated with these disturbances, abiotic factors (excluding fire) probably play a diminished role in determining their distribution. Some large-scale physical environmental factors (e.g., climates with summer drought) may facilitate development of annual grassland within native Habitats (Sims 1988; Keeley 1990). However, it is doubtful that non-native grasslands would develop in most Habitats in the absence of fire, grazing, or other form of disturbance. Species composition varies from one site to another but several annual grass species appear to show site preferences based on annual rainfall (Janes 1969). This research described grassland species along a rainfall gradient with soft chess and broad-lobed filaree on the mesic end (>20cm rainfall) with fox-tail chess and red-stemmed filaree in more xeric conditions(<19 cm) (Janes 1969). Abiotic factors are thought to influence the species composition of annual grasslands on a local scale. Seasonal variation in temperature, rainfall, and physical microsite differences have been shown to influence annual grassland species composition (Evans and Young 1989; [in S&K-W]).

    ECOSYSTEM PROCESSES

    Valley and foothill and non-native grasslands within the MSHCP Plan Area occur within a landscape mosaic consisting of native shrublands and woodlands (primarily chaparral and coastal sage scrub, and oak woodlands), and wetland vegetation (e.g., riparian scrub, forest, and freshwater marsh) and urban and agricultural lands (PSBS 1995). The ecosystem processes regarding Valley and foothill grasslands and non-native grasslands are not well understood. The interacting roles of fire, grazing, cultivation, non-native species invasions, pollution, and urban development combined with only speculative understanding of the pre-European state of grasslands, complicates understanding the underlying formative and sustaining processes of grasslands. In light of these complications, discussion of ecosystem processes must address what is known currently about the known extent and nature of grasslands. Primary driving forces within grasslands in terms of changes in distribution, species composition, and ecosystem function probably include fire, grazing, and human settlement. Other ecological processes, including competition between species, nutrient cycling, and variation in precipitation and temperature probably are also important factors but many of these are difficult to analyze because of the changes introduced by humans.

    Valley and Foothill Grasslands. The current distribution of Valley and foothill grasslands within southern California is limited to areas supporting deep clayey soils that have not been heavily disturbed by mechanical disturbance (Keeley 1993). Research concerning the ecosystem processes with Valley and foothill grasslands should focus not only on extant native grasslands, but also should consider other vegetation types that are currently present on or have acted to form heavy or clay soils similar to those that currently support native grasslands.

    No conclusive evidence has emerged concerning the relationship between Valley and foothill grasslands and other shrubland or woodland Habitats within the same landscape. Research concerning the role of fire in the distribution and maintenance of Valley and foothill grasslands has offered few conclusive facts. Some research has suggested the distribution of native grasslands was related to a long history of burning by native Americans (Sampson 1944; Bean and Lawton 1973; Timbrook et al. 1982). Others dismiss native American burning as not playing a significant role in the distribution of native grasslands suggesting that lightning-caused fires were more likely in the process maintaining grassland ecology (Heady 1977). Evidence supporting this assertion includes the finding that more common perennial grassland dominants (Nassella pulchra, N. lepida) are adapted to fire; resprouting and producing greater volumes of seed following fire (Ahmed 1983; Keeley and Keeley 1984). Several field studies have reported an increasing cover Nassella spp. based on burning treatments (Hatch et al. 1991; Dyer et al. 1996; Wills pers. comm. 1995), while other studies have shown mixed effects on species abundance from burning (Hatch et al. 1999). Though preliminary research has pointed to increasing abundance of perennial grasses following fire, there is little research describing the role of fire on maintaining the role of fire on other native species within Valley and foothill grassland Habitat.

    The effects of grazing on Valley and foothill grasslands also remains unclear. In spite of the fact that a long history of intensive grazing in California has been cited as one of the primary reasons for the demise of perennial grasslands (Burcham 1957; Dasmann 1966 in Keeley 1990; Bartolome and Gemmill 1981) most research has found evidence suggesting that some intensity of grazing is beneficial to or does not negatively affect perennial grasses (Huntsinger et al. 1996). Several researchers have documented cases where native grasses have not increased in abundance on sites that have been excluded from grazing over relatively long (e.g., 20 to 40 year) periods (White 1967; Bartolome and Gemmill 1981; Goode 1981; Heady et al. 1991). Heady (1968 1977) has suggested that large native herbivores present prior to European colonization may have been an important factor in grassland formation and ecology. This assertion supports studies that have found that some form of grazing may be necessary to maintain native perennial grasses (Heady et. al. 1991). Two related field studies have produced mixed results with respect to grazing. Hatch et al. (1991) found perennial grasses to increase in abundance with the exclusion of grazing, while more recent study, in a more coastal location, found no relationship between grazing exclusion and perennial grass cover. Much of the research that has been conducted concerning grazing and native grasslands has focused on the cover of perennial grass species. Additional research is needed to determine the effects of grazing on the abundance and diversity of the other native plant and animal species within perennial grasslands. Based on these studies, it appears that the interactive effect of grazing and native grassland reflects local conditions, and patterns and intensity of grazing. A generic grazing or burn management program may not work for all sites, and proper management treatments would have to be determined on a site-by-site basis.

    Non-Native Grasslands. Without question the distribution of non-native grasslands has increased to the former distribution of native grasslands (sensu. Clements 1920) or beyond, occupying lands formerly consisting of coastal sage scrub (O’Leary and Westman 1988; Minnich and Dezzani 1999), chaparral (Zedler et al., 1983), and oak woodlands (Callaway and Davis 1993). The literature describing this type conversion of native systems generally has shown that altered fire frequencies, grazing pressure or other physical disturbance combined with competitive exclusion by non-native species have caused the expansion of annual grasslands into native Habitats previously occupied by perennial species. Recent work by Minnich and Dezzani (1999) has documented changes in the distribution of coastal sage scrub and annual grassland within a portion of Western Riverside County. This research has shown that non-native grasslands currently are increasing in distribution in areas formerly supporting coastal sage scrub. Some authors have noted that these grasslands have remained stable to an extent over time that it has been proposed to accept many non-native grassland species as “new natives” and manage non-native grasslands as though they were natural systems (Heady 1977). Acceptance of the current distribution of annual grasslands may be shortsighted because recent research in the coastal sage scrub/annual grassland interface has shown that the stability of non-native grasslands may be related to permanent changes in soil nutrient and moisture regimes caused by the presence of exotic species (Heunneke et al. 1990) and air pollution (Allen et al. 1996; Padgett et al. 1997) (Minnich and Dezzani 1998). Because non-native grasslands appear to be invading other Habitat-types and are themselves stable over long periods of time, management is necessary to stabilize or reduce their extent. Preserve design and management also must consider the fact that non-native grasslands have been present long enough to develop as Habitat, forage, and Habitat linkage for native animal species including listed species such as the Stephens’ kangaroo rat. Management strategies should focus on reduction in the abundances of exotics that displace native species in other Habitats, increasing the abundances of native herbaceous species in grasslands, and minimizing the effects that cause non-native grasslands to be self-perpetuating.

    THREATS

    Valley and Foothill Grasslands. Threats to Valley and foothill grasslands may include disturbance of clay soils by agricultural activities, invasion of exotic species, grazing, fire, and urban development. Although Habitat management may be necessary for the preservation of native grasslands, it is also a potential threat to Valley and foothill grasslands because the character of Valley and foothill grasslands, and the processes that control them have not been established. Inappropriate grazing and fire treatments may not be productive and could result in Habitat degradation. Further study is needed to determine whether fire or grazing are necessary to maintain or develop native grasslands. If grazing and/or fire are necessary to maintain existing or to develop new perennial grasslands, studies must also determine the proper regimes under which native grasslands will persist. Because these regimes are not likely present under current and future conditions, management likely will be required to maintain or develop native grasslands. Quantitative descriptions of the extant native grasslands should be used to develop goals for what types of management would be implemented (i.e., species composition, cover, etc.).

    Non-Native Grasslands. Threats to non-native grasslands largely include direct loses due to urban development, increased agricultural activity, and invasion by noxious weedy species (e.g., artichoke-thistle) that reduce the Habitat qualities necessary for native species.

    LITERATURE CITED

    Adams, M.S. 1964. Ecology of Stipa pulchra, with special reference to certain soil characteristics. Master’s Thesis. University of California Davis.

    Ahmed, E.O. 1983. Fire Ecology of Stipa pulchra in California annual grassland. PhD. dissertation, University of California, Davis.

    Allen, E.B., P.E. Padgett, A. Bytnerowicz, and R.A. Minnich. 1998. Nitrogen-deposition effects on coastal sage vegetation of southern California, In Proceedings of the International Symposium on Air Pollution and climate change effects on forest ecosystems, 5-9 February, Riverside (A. Bytnerowicz, M. Arbauge, and S, Schilling technical coordinators) U.S. Forest Service Rep. GTR-164.

    Baker, H.G. 1989. Sources of the naturalized grasses and herbs. In California grasslands, Pp 29-38. In L.F. Huenneke and H.A. Mooney, (eds.) Grassland structure and function: California annual grasslands. Kluwer Academic Publishers, Boston, MA.

    Barry, W.J. 1972. The Central Valley prairie. California Department of Parks and Recreation. 82pp.

    Bartolome, J.W., and B. Gemmill. 1981. The ecological status of Stipa pulchra (Poaceae) in California. Madrono 28:172-184.

    Bean, L.J. and H.W. Lawton. 1973. Some explanations for the rise of cultural complexity in native California with comments on proto-agriculture and agriculture, Pp. V-xlvii In H.T. Lewis, (ed.) Patterns of Indian burning in California: ecology and ethnohistory. Ballena Press, Ramona, California.

    Boyd S. And D.L. Banks. 1995. A botanical assessment of the Agua Tibia Wilderness Area, Cleveland National Forest, California. Rancho Santa Ana Botanic Garden, Claremont, California.

    Boyd 1983. A Floral inventory of The Gavilan Hills. Master’s Thesis. California State University Fullerton.

    Burcham, L.T. 1957. California range land: an historic-ecological study of the range resource of California. Division of Forestry, Department of Forestry, Department of Natural Resources, State of California, Sacramento.

    Callaway, F.M. and F.W. Davis 1993. Vegetation dynamics, fire, and the physical environment in coastal central California. Ecology 74:1567-1578.

    Clements, F.E. 1920. Plant Indicators. Carnegie Institution of Washington, Washington D.C.

    Cooper, W.S. 1922. The broad-sclerophyll vegetation of California: an ecological study of the chaparral and its related communities. Carnegie Institution of Washington, Washington D.C.

    Dasmann, R.F. 1965. The destruction of California. MacMillan Company, New York.Dyer, A.R., H.C. Fossum, and J.W. Menke. 1996. Emergence and survival of Nassella pulchra in a California grassland. Madrono 43:316-333.

    Evans, R.A. and J.A. Young. 1989. Characterization and analysis of abiotic factors and their influences on vegetation. Pp. 13-28 In L.F. Huenneke and H.A. Mooney, (eds.) Grassland structure and function: California annual grasslands. Kluwer Academic Publishers, Boston, MA.

    Goode, S. 1981.The vegetation of La Jolla Valley. Masters Thesis, California State University, Los Angeles.

    Hamilton, J.G. 1997. Changing perceptions of pre-European grasslands in California. Madrono 44:311-333.

    Hatch, D.A., J.W. Bartolome, and D.S. Hillyard. 1991. Testing a management strategy for restoration of California’s native grasslands. Pp. 343-349 In Yosemite Centennial Symposium Proceedings: Natural Areas Conference with the Yosemite Centennial Celebration, October 13-20, 1990, Davis, California. National Park Service, Branch of Publications and Graphic Design, Denver Service Center, Denver, Colorado.

    Hatch, D.A., J.W. Bartolome, J.S. Fehmi, and D.S. Hillyard 1999. Effects of burning and grazing on a coastal California grassland. Restoration Ecology 7:376-381.

    Heady H.F. 1956. Changes in a California annual plant community induced by manipulation of natural mulch. Ecology 37:798-812.

    Heady H.F. 1968. Grassland response to changing animal species. Journal of Soil Water Conservation 23:173-176.

    Heady H.F. 1977. Valley grasslands. Pp.491-514. In M.G. Barbour and J. Major (eds.), Terrestrial vegetation of California. John Wiley, New York.

    Holland, R.F. 1986. Preliminary descriptions of the terrestrial communities of California. California Department of Fish and Game, Nongame Heritage Program, Sacramento.

    Huenneke, L.F., and Mooney, H.A. (eds.) 1989. Grassland structure and function: California annual grassland. Tasks for Vegetation science, volume 20. Dordrecht, Boston.

    Huntsinger, L.H., M.P. McClaran, A. Dennis, and J.W. Bartolome. 1996. Defoliation response and growth of Nassella pulchra (A. Hitchc.) Barkworth from serpentine and non-serpentine populations. Madrono 43:83-103.

    Jackson, L.E. 1985. Ecological origins of California’s Mediterranean grasses. Journal of Biogeography 12:349-361.

    Janes, E.B. 1969. Botanical composition and productivity in the California annual grassland in relation to rainfall. Master’s Thesis. University of California, Berkeley.

    Keeley, J. E. 1990. The California valley grassland. Pp. 2-23 in A.A. Schoenherr (ed.), Endangered plant communities of southern California. California State University, Fullerton. Southern California Botanists, Special Publication No. 3.

    Keeley, J. E. 1993. Native grassland restoration: the initial stage-assessing suitable sites. Pp. 277-281 in J.E. Keeley (ed.), Interface between ecology and land development in California. Southern California Academy of Sciences, Los Angeles.

    Keeley, J. E. and S.C. Keeley. 1984. Postfire Recovery of California coastal sage scrub. American Midland Naturalist 111:105-117.

    Knecht , A.A. 1971. Soils survey of western Riverside area, California. United States Department of Agriculture, Soil Conservation Service, United States Department of the Interior (BIA), and University of California Agricultural Experiment Station. U.S. Government Printing Office, Washington D.C.

    Lathrop, E.W. and R.F. Thorne. 1968. Flora the Santa Rosa Plateau of the Santa Monica Mountians, California. Aliso 6:17-40.

    Lathrop, E.W. and R.F. Thorne. 1985. A new preserve on the Santa Rosa Plateau. Fremontia 13:15-19.

    Markel and Associates 1982. Steppe grass ecosystem review Trabuco District, Cleveland National Forest. Submitted to Southern California Edison Company Environmental and Regulatory Affairs Division.

    Mack, R.N. 1989. Temperate grasslands vulnerable to plant invasions: characteristics and consequences. Pp. 155-179 In J.A. Drake, H.A. Mooney, F. di Castri, R. H. Groves, F.J. Kruger, M. Rejmanek, and M. Williamson (eds.), Biological invasions: a global perspective. John Wiley and Sons, Chichester, N.Y.

    Mc Naughton S.J. 1968. Structure and function in California grasslands. Ecology 49:962-972.

    Minnich, R.A. and R.J. Dezzani 1998. Historical decline of coastal sage scrub in the Riverside-Perris plain, California. Western Birds 29:366-391.

    O’Leary J.F. and W.E. Westman 1988. Regional disturbance effects on herb succession patterns in coastal sage scrub. Journal of Biogeography. 15:775-786.

    Padgett, P.E., E.B. Allen, A. Bytnerowicz, and R.A. Minnich. 1999. Changes in soil inorganic nitrogenous related to atmospheric nitrogenous pollutants in southern California. Atmospheric Environment.

    PSBS. 1995. Western Riverside County Multi-Species Habitat Conservation Plan; Phase1-Information Collection and Evaluation. Prepared for: Western Riverside County Habitat Consortium.

    Sampson, A.W. 1944. Plant succession on burned chaparral lands in northern California. California Agricultural Experiment Station Bulletin 685:1-144.

    Sawyer, J.O. and T. Keeler-Wolf 1995. A manual of California vegetation. California Native Plant Society, Sacramento.

    Sims, P.L. Grasslands. Pp. 265-286 In M.G. Barbour and W.D. Billings (eds.), North American terrestrial vegetation. Cambridge University Press, New York.

    Sims, P.L. and P.G. Risser 2000. Grasslands. Pp. 325-356 In M.G. Barbour and W.D. Billings (eds.), North American terrestrial vegetation. Cambridge University Press, New York.

    Timbrook, J., J.R. Johnson, and D.D. Earle. 1982. Vegetation burning by the Chumash. Journal of California and and Great Basin Anthropology 4:163-186.

    White, K.L. 1967. Native bunchgrass Stipa pulchra on Hastings Reservation, California. Ecology 48:949-955.

    Zedler, P.H., C.H. Gautier, and G.S. McMaster. 1983. Vegetation change in response to extreme events. The effect of a short return interval between fires in California chaparral and coastal scrub. Ecology 64:809-818.

    Grove/Orchard

    VEGETATION ASSOCIATION:

    GROVE/ORCHARD

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Because grove and orchard farms are registered with the County and State, and information regarding land use is stored with the County by parcel, the mapping of this Vegetation Community is considered to be very accurate.

    BIOGEOGRAPHY

    Groves and orchards are planted in human-inhabited areas throughout the world, where climatic conditions are appropriate.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    The largest area of grove/orchard is in Santa Rosa East between Gavilan Mountain and Mesa de Colorado. Smaller tracts of grove/orchard are mapped in the Pauba Valley east of Temecula (Long Valley, Buck Mesa, Glen Oak Valley), east of Hemet, in Temescal Valley, on the Gavilan Plateau, southwest of Lake Matthews, in south Corona and El Cerrito, in Eagle Valley, Arlington Mountains, Woodcrest, and Highgrove. Small and scattered mappings include areas around Sage, in the Lewis and Reed Valleys between Anza and Cahuilla Mountain, in the Elsinore Mountains west of Wildomar, in the Antelope Valley and El Cariso, in the Lakeview Mountains and in the community of Nuevo, and in Moreno Valley, Banning, and Beaumont.

    VEGETATION CHARACTERISTICS

    Agricultural groves are generally open and of single species with often low bushy trees which result from pruning (Schultze 1988). The undergrowth usually contains low-growing grasses and other herbs but is mostly composed of bare ground (Schultze 1988). Aside from agricultural purposes, groves can also be planted as windbreaks, for aesthetic purposes or as firewood or lumber (Holland and Keil 1995). Eucalyptus (Eucalyptus spp.) has been planted widely in California for windbreaks, aesthetic purposes, and for firewood and lumber (Holland and Keil 1995). These stands shade the ground and litter the soil surface (Holland and Keil 1995).

    Plantations of orchard crops may include walnuts (Juglans spp.), plums (Prunus domestica), almonds (Prunus dulcis), peaches (Prunus persica), and apples (Malus sylvestris) (Holland and Keil 1995). Grapes (Vitis vinifera) are commonly grown in Riverside County; it is the second largest agricultural commodity (CDFA 1999). Other leading grove/orchard producing commodities are grapefruit, avocados (Persea spp.), and lemons, in that order (CDFA 1999). According to 1992 statistics, 21 acres of almonds, 199 acres of apples, 42 acres of apricots, 8,462 acres of avocados, 117 acres of cherries, 13 acres of figs, 11,708 acres of grapefruit, 136 acres of kiwi fruit, 5,478 acres of lemons, 54 acres of limes, 125 acres of nectarines, 125 acres of olives, 19 acres of pecans, 12,813 acres of oranges, 189 acres of peaches, 2 acres of pears, 39 acres of pistachios, 51 acres of plums, 598 acres of tangelos, 2,079 acres of tangerines, 43 acres of walnuts, and 18,253 acres of grapes were planted in Riverside County (California Agricultural Statistics Service 1993b).

    PHYSICAL ENVIRONMENT

    Orchards and groves may be found on flat alluvial soils on valley floors, in rolling hill areas, or relatively steep sloped areas (Schultze 1988). Most orchards are irrigated and occur in non-mountainous areas to avoid frost damage (Schultze 1988).

    ECOSYSTEM PROCESSES

    Agricultural cultivation practices are the biggest influences to plant growth. Typically, the desired species are planted as seedlings and are managed until they are grown small to medium size (Schultze 1988). Old and/or damaged trees are usually individually replaced although sometimes the entire plot is replaced or abandoned (Schultze 1988).

    Other processes which influence these areas are a result of the physiographic structure. Typically, grove trees shade the ground and draw nutrients and water from the soil thus influencing plant communities and wildlife interactions (Holland and Keil 1995). Allelopathic chemicals in the leaves of eucalyptus trees, for example, inhibit understory growth (Holland and Keil 1995). The chemicals typically are transferred to the soil via fog-drip or rainwater (Holland and Keil 1995).

    THREATS

    No threats to grove and orchard production have been identified in the literature review conducted for this account. It can be stated that the main threat to grove and orchard production is economic competition for land.

    LITERATURE CITED

    California Agricultural Statistics Service. 1993b. California Fruit and Nut Crop Statistics 1983-1992. County Data 1991-92. Sacramento, California.

    California Department of Food and Agriculture. 1988. California Vegetable Crop Statistics: County Data 1986-87. Sacramento, California.

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company. Dubuque, Iowa.

    Schultze, Ronald F. 1988. Orchard-Vineyard. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 140-141.

    Jeffrey Pine

    VEGETATION ASSOCIATION:

    JEFFREY PINE

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Because of its distinctive character, as well as signature on aerial photographs, and the lack of suitable climate for the dominant species throughout much of the Plan Area, it is likely that the majority of Jeffrey pine forest within the Plan Area was mapped accurately in the San Jacinto mountains. It is possible that areas mapped as Jeffrey pine forest contains areas dominated by ponderosa pine.

    BIOGEOGRAPHY

    Jeffrey pine is nearly endemic to California, with stands occurring from the Klamath Mountains of northern California and southwestern Oregon through the Sierra Nevadas and south in the Transverse and Peninsular ranges of southern California and northern Baja California (Jenkinson 1990). Over its range, Jeffrey pine occurs from 150 to 2900 m. Areas where Jeffrey pine shows dominance are in the Klamath Mountains, North Coast Ranges, Cascade Range, Modoc Plateau, Sierra Nevada, and Transverse and Peninsular ranges (Jenkinson 1980).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Vogl (1976) describes the Jeffrey pine forest within the San Jacinto Mountains, noting that Round Valley, Long Valley, and Tahquitz Valley are exceptional sites in terms of species composition and maturity. In the MSHCP database, this Habitat covers 15,424 acres, comprising 1.2% of the Plan Area. Jeffrey pine forest is mapped throughout the San Jacinto Mountains from north of Pine Meadow (on both sides of SR-74) in Garner Valley, on Thomas Mountain south of Lake Hemet, around the community of Mountain Center, around Pine Cove, and Indian Mountain (west of Tahquitz Peak, in the vicinity of Fern Valley), north to Lake Fulmor within May Valley and on Baldy Mountain. Most areas are within U.S. Forest Service land, except around the Pine Cove and Pine Meadow.

    VEGETATION CHARACTERISTICS

    Barbour and Minnich (2000) describe a “midmontane forest” which probably includes areas mapped as Jeffrey pine forest and Ponderosa pine forest. Jeffrey pine typically is situated at higher elevations above ponderosa pine (Pinus ponderosa) (McBride 1988). Jeffrey pine forest typically is composed of four layers, although in most xeric sites the herbaceous layer may not be significant and in moderately dry sites Jeffrey pine occurs as a single canopy layer (McBride 1988). The canopy layer may be composed of only Jeffrey pine (Pinus jeffreyi), or also may include ponderosa pine, Coulter pine (Pinus coulteri), sugar pine (Pinus lambertiana), lodgepole pine (Pinus contorta ssp. murrayana ssp. murrayana), limber pine (Pinus flexilis), white fir (Abies concolor), incense-cedar (Calocedrus decurrens) and black cottonwood (Populus balsamifera ssp. trichocarpa) (McBride 1988). Conifer tree canopy varies from 30 to 60 m forming a fairly continuous canopy of 50-80% cover (Barbour and Minnich 2000, Sawyer and Keeler-Wolf 1995). A subdominant tree canopy occurs patchily and varies in species composition. Heights for these trees range from 5 to 15 m and include winter-deciduous species such as bigleaf maple (Acer macrophyllum), Pacific dogwood (Cornus nuttallii), hazelnut (Corylus cornuta) and black oak (Quercus kelloggii), and broad-leaved evergreens, canyon live oak (Quercus chrysolepis), and curl-leaf mountain-mahogany (Cercocarpus ledifolius). Shrub cover generally is infrequent to common, composing 10-30% of the overall cover and including deciduous and evergreen species and many sclerophylls of the following genera: manzanita (Arctostaphylos)California-lilac (Ceanothus)mountain-misery (Chamaebatia)chinquapin (Chrysolepis)tan oak (Lithocarpus)snowberry (Symphoricarpos), cherry (Prunus)oaks (Quercus)currant (Ribes) and blueberry (Vaccinium) (Barbour and Minnich 2000, Sawyer and Keeler-Wolf 1995, McBride 1988). Herbaceous cover is usually between 5 and 10% but may reach 20% and is dominated by perennial forbs or grasses (Barbour and Minnich 2000, Sawyer and Keeler-Wolf 1995). The following genera occur commonly: trail plant (Adenocaulon), Clintonia, Disporum, bedstraw (Galium)iris (Iris)lupine (Lupinus), Osmorhiza, bracken fern (Pteridium)wintergreen (Pyrola), false Solomon’s seal (Smilacina), rockcress (Arabis), bird’s-beak (Cordylanthus), buckwheat (Erigonum), fritillary (Fritillaria), ivesia (Ivesia), melic (Melica), beardtongue (Penstemon) and needlegrass (Nassella) and violet (Viola) (Barbour and Minnich 2000, McBride 1988).

    In the San Jacinto Mountains, Jeffrey pine occurs in a park-like Habitat with very old trees (Jenkinson 1980). Old-aged Jeffery pines, often over 50 to 100 years old but also reaching 350 to 800 years, dominate this Habitat with few sapling-sized and smaller trees. The understory is mostly non-continuous, although circular clumps of manzanita (Arctostaphylos parrryana var. pinetorum)spiny snowbush (Ceanothus cordulatus) and bush chinquapin (Chrysolepis sempervirens) do occur occasionally (Vogl 1976). Other occasional understory plants include herbs which grow close to the soil surface usually with woolly or fleshy leaves (Vogl 1976). A few level areas contain ideal conditions for a well-developed canopy of pines (Pinus spp.) and understory of bracken fern (See discussion of level areas in Physical Environment section below). Generally, this Habitat within Riverside County does not exhibit high litter accumulation.

    Black oak becomes an important component of Jeffrey pine forest between 1525 and 2135 m in the San Jacinto Mountains (Thorne 1977). Herbaceous cover on dry, rocky slopes and flat areas include rock cress (Arabis repanda), bird’s-beak (Cordylanthus spp.), buckwheat (Eriogonum parishii), fritillary (Fritillaria pinetorum), Gayophytum diffusum ssp. parviflorum, mousetail ivesia (Ivesia santolinoides), Peirson’s lupine (Lupinus peirsonii), melic (Melica stricta), beardtongues (Penstemon rostriflorus and P. caesius), and needlegrass (Stipa parishii) (Thorne 1977).

    PHYSICAL ENVIRONMENT

    Under Barbour and Minnich’s (1995) midmontane forest classification, Jeffrey pine forest would range from 800 to 2500 m. Areas supporting Jeffrey pine have a mean annual temperature of 11oC and annual precipitation is about 100 cm (Barbour and Minnich 1995). Over its California range, Jeffrey pine-dominated areas occur from 60 to 2900 m elevation (Sawyer and Keeler-Wolf 1995). In the San Jacinto Mountains, the typical, park-like, old-aged groves of Jeffrey pine occur from 2125 m to the mountain peak (Vogl 1976).

    Jeffrey pines occur where many other conifer species cannot. Due to greater tolerance of serpentine soils, drought, smog and warm temperatures, Jeffrey pine outcompetes ponderosa pine in areas with these elements (Thorne 1977). The Jeffrey pine is also more competitive on cold, xeric and infertile sites (Jenkinson 1990). A large portion of Jeffrey pine sites are on ultramafic soils; otherwise they are typically found on well-drained soils of a variety of types (Jenkinson 1990; Sawyer and Keeler-Wolf 1995). At lower elevations and moderate slopes, soils which support this Habitat typically are loamy alfisols or ultisols (Barbour and Minnich 2000). Where slopes are steeper, inceptisols generally are found (Barbour and Minnich 2000). Jeffrey pine is known to have a limited need for soil nutrients and moisture probably due to the tree’s short growing season and early dormancy (Jenkinson 1980).

    In the San Jacinto Mountains the trees are most common on moderate slopes and flat areas (Vogl 1976). Certain level areas contain the unique combination of deep soils and flat topography allowing for heavy snow accumulations and retained moisture from summer thunderstorms. Much of this Habitat in Riverside County, however, occurs on steep slopes. This fact, added with the presence of dry summers and high winds prevent the accumulation of litter (Vogl 1976).

    ECOSYSTEM PROCESSES

    Barbour and Minnich (2000) describe a change in the Jeffrey pine forest physiognomy of the past 100 years due to fire suppression activities where shrub and tree cover have shifted. Pre-suppression studies of fire frequency estimate an interval between four and 20 years (Barbour and Minnich 2000). Estimates are that fires were of low-intensity, small (between 1 and 800 ha), patchy, and irregularly shaped (Barbour and Minnich 2000). In the Sierra San Pedro Martir, Baja California, Mexico, which still maintains a relatively natural wildlife regime, the average fire return interval is 26 years, with most fires being less than 16 ha in size and less than 3% resulting in crown replacement (Barbour and Minnich 2000). It appears that most constituent species of this Habitat are adapted to this fire regime. Bracken fern recovers quickly following fires (Vogl 1976). Pines are well-served by fires which burn local accumulations of litter creating optimum conditions for seedbed growth (Vogl 1976). Fire is thought to help reduce stress and mortality caused by competition by eliminating “over-mature, sick and insect- or disease-infected” Jeffery pines (Barbour and Minnich 2000; Vogl 1976). This is evidenced by the large die-offs in the San Jacinto Mountains of ponderosa pine (42% mortality), Jeffrey pine (50%) and Coulter pine (49%) during the drought period in the 1980's and 1990's when compared to the stability of pine populations in Baja California (Barbour and Minnich 2000).

    Jeffrey pines are intolerant of shade and seedlings do not grow well in the presence of old-growth roots (Jenkinson 1990). These species are especially adapted for their seasonal environment. For example, reproduction occurs by vegetative layering, stiff stems help prevent snow crushing, evergreen leaves are adapted to the short growing periods, and the dome-shaped growth of these plants regulates snow deposition (Vogl 1976).

    THREATS

    According to the USDA (1999), fire suppression is thought to result in three major problems for this forest type: “(1) a large increase in the number of understory trees (particularly shade-tolerant white fir and incense-cedar), (2) an increased risk of stand-replacing crown fires due to fuel buildup, and (3) increased mortality and reduced recruitment of large trees (due to increased understory competition).” These hypotheses were supported by a study comparing tree diameters and species composition from a population in 1930 and the same population in 1997. Fewer Jeffrey pines and more white fir and incense-cedar were found in 1997; more small-diameter trees and fewer large trees also were recorded in 1997 (Stephenson and Calcarone 1999).

    Jeffrey pine is especially susceptible to infection from a strain of Heterobasidion annosum, a fungal root disease which colonizes the roots of a freshly cut conifer and eventually kills other trees which come in contact with the infected roots (Stephenson and Calcarone 1999). Also, the western dwarf mistletoe (Arceuthobium campylopodum) infects Jeffrey pine in this Habitat. The mistletoe is managed mainly through the pruning and removal of infected trees (Stephenson and Calcarone 1999). In general, sick and diseased Jeffery pines may suffer their final demise due to the Jeffrey pine bark beetle (Dendroctonus jeffreyi)Ips emarginatus is another bark beetle known to attack Jeffrey pine (Stephenson and Calcarone 1999). Elytroderma disease (Elytroderma deformans) has reached epidemic proportions in some stands (Jenkinson 1990). Other diseases include Medusa needle blight (Davisomycella medusa), Cenangium limb canker (Cenangium ferruginosum), stalactiform rust (Peridermium stalactiforme), filamentosum rust (Peridermium filamentosum), sweetfern rust (Cronartium comptoniae), tarweed rust (Coleosporium madiae), and western gall rust (Peridermium harknessii) (Jenkinson 1990).

    Although timber harvests largely have been discontinued except to supply local demand, high intensity harvests from the 1950's to the mid-1970's contributed to the reduction in the number of large trees and possibly to the increase of white fir in this Habitat (Stephenson and Calcarone 1999).

    Jeffrey pines have been shown to suffer foliage damage from ozone depletion (Stephenson and Calcarone 1999). The major phytotoxins are ozone, nitrogen oxides, hydrocarbons and sulfur dioxide, of which ozone seems to be the most important (Barbour and Minnich 2000). Symptoms of excess ozone are evident on ponderosa and Jeffrey pines through “mottling on the needles, premature needle drop, reduction in net photosynthesis and reduction in needle size (Barbour and Minnich 2000).” The highest rates of mortality occur when ozone-damaged trees enter a drought period and cannot withstand attacks by bark beetles (Stephenson and Calcarone 1999). Forest areas exposed to heavy air pollution are most susceptible to this injury.

    Other threats to Jeffrey pine include a susceptibility to extremely low temperatures, the creation of anaerobic conditions, either by flooding or filling, and highway de-icing salts (Jenkinson 1990).

    LITERATURE CITED

    Barbour, Michael G. and Richard A. Minnich. 2000. Californian Upland Forests and Woodlands. In North American Terrestrial Vegetation. Second Edition. Cambridge University Press, Cambridge, UK.

    Jenkinson, James L. 1990. Pinus jeffreyi - Jeffrey pine. Pages 359-367 in R.M. Burns and B. H. Honkala, technical coordinators. Silvics of North America, Volume 1. Confiers. Agriculture Handbook 654. USDA, Forest Service, Washington, D.C.

    Jenkinson, James. L. 1980. Jeffrey Pine. In Forest Cover Types of the United States and Canada. ed. F. H. Eyre. Society of American Foresters, Washington, D.C. pp. 123.

    Lathrop, Earl W. and Bradford D. Martin. 1982. Response of Understory Vegetation to Prescribed Burning in Yellow Pine Forests of Cuyamaca Rancho State Park, California. Alsio 10(2): 329-343.

    McBride, Joe R. 1988. Jeffrey Pine. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 54-55.

    McDonald, Philip A. 1990. Psuedotsuga macrocarpa - bigcone Douglas-fir. Pages 520-525 in R.M. Burns and B. H. Honkala, technical coordinators. Silvics of North America, Volume 1. Conifers. Agriculture Handbook 654. USDA, Forest Service, Washington, D.C. 1990.

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Stephenson, John R. and Gena M. Calcarone. 1999. Southern California mountains and foothills assessment: Habitat and species Conservation issues. General Tehnical Report GTR-PSW-172. Albany, CA: Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture; 402 p.

    Thorne, Robert F. 1977. Montane and Subalpine Forests of the Transverse and Peninsular Ranges In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 538-557.

    Vogl, Richard J. 1976. An Introduction to the Plant Communities of the Santa Ana and San Jacinto Mountains. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 77-98.

    Lodgepole Pine Forest

    VEGETATION ASSOCIATION:

    LODGEPOLE PINE FOREST

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Because lodgepole pine forest is confined to a specific climatic condition and an elevation which is present only in the San Jacinto Mountains, the mapping of the Habitat is most likely to be accurate. Additionally, the indicator species, lodgepole pine forest, is fairly conspicuous and discernable from other pines.

    BIOGEOGRAPHY

    Munz and Keck (1949) only described this Habitat, in California, as reaching from northernmost California to the central Sierra Nevada between 2530 and 2900 m. However, lodgepole pine forest occurs on high summits in the Transverse and Peninsular ranges as well as the Cascades and Sierra Nevada ranges (Thorne 1976). As a species, lodgepole pine forest can be found from sea level, along the Pacific Coast from Alaska to Baja California, to elevations up to 3600 m as far east as the Rocky Mountains and the Black Hills of South Dakota (Holland and Keil 1995). Lodgepole pine forest has a wide latitudinal range from 63o to 36o north (Pfister and McDonald 1980).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    In the Plan Area, lodgepole pine forest is known from the upper slopes of the San Jacinto Mountains (Sawyer and Keeler-Wolf 1995). Within the MSHCP database, Lodgepole pine forest is mapped in only one general area, west of San Jacinto Mountain, in the vicinity of Little Round Valley, Marion Mountain, and Marion Ridge. The total area of this Habitat is 1,654 acres.

    VEGETATION CHARACTERISTICS

    Lodgepole pine forest occur at slightly higher elevations than mixed coniferous forests in southern California (Holland and Keil 1995). Lodgepole pine forest (Pinus contorta ssp. murrayana ssp. murrayana) may grow in pure stands or as a component of other Habitat types (Lotan and Critchfield 1990). Typically, Lodgepole pine forest forms open stands with similar sized, shorter individuals and a sparse understory (Bartolome 1988). Other conifers found in these forests include western white pine (Pinus monticola) and mountain hemlock (Tsuga mertensiana). When located near wet meadows or ponds, quaking aspen (Populus tremuloides) and willows (Salix spp.), along with a variety of shrubs and showy perennial herbs, are occasionally present (Thorne 1976).

    Where lodgepole pine forest is the dominant tree species, the canopy can range from continuous to open (Sawyer and Keeler-Wolf 1995). Typical heights range from 15 to 20 m and typical crown cover is 50 to 60 percent, but can reach 80 percent (Bartolome 1988, Barbour and Minnich 2000). Canopies may reach slightly greater heights at higher elevations where lodgepole pine forest often forms dense forest with limber pine (Pinus flexilis).

    Shrub density varies from scarce to about 15% and may include manzanita (Arctostaphylos spp.), California-lilac (Ceanothus spp.)chinquapin (Chrysolepis spp.)currant (Ribes spp.), mountain-mahogany (Cercocarpus minutiflorus), spotted coralroot (Corallorhiza maculata), pinedrops (Pterospora andromedea), snowplant (Sarcodes sanguinea), and wintergreen (Pyrola ssp.) (Barbour and Minnich 2000, Holland and Kiel 1995, Thorne 1977). Understory plants typically are sparse to abundant and include species such as sandwort (Arenaria spp.), mariposa lily (Calochortus invenustus), fleabane daisy (Erigeron breweri ssp. jacinteus), gray monardella (Monardella cinerea), and campion (Silene verecunda ssp. platyota) (Sawyer and Keeler-Wolf 1995). Munz and Keck (1949) list these additional species: rothrock sagebrush (Artemisia rothrockii), cinquefoil (Potentilla breweri), owl’s-clover (Castilleja lemmonii), little elephant’s head (Pedicularis attolens), and groundsel (Senecio lugens). In southern California huckleberry (Vaccinium spp.) and mountain-heather (Phyllodoce spp.) are common (Bartolome 1988). In drier areas, Jeffrey pine and lodgepole pine forest may be associated with montane chaparral species or mycotrophic plants such as wintergreensnowplant (Sarcodes sanguinea), pinedrops and spotted coralroot. Only where lodgepole pine forest occurs near a meadow is herb cover significant and comprised of hydrophytic species (Holland and Keil 1995, Barbour and Minnich 2000). Other areas may be composed of dry, glacial scoured areas where herbs are sparse (Holland and Keil 1995).

    In the San Jacinto Mountains lodgepole pine forest occurs in clusters on upper slopes. The large trees have many small and open cones and usually show conspicuous spiral sapwood splits caused by lightning (Vogl 1976).

    PHYSICAL ENVIRONMENT

    In southern California, lodgepole pine forest occurs from 2,450 to 3,600 m, but also may be present at lower elevations in stands around saturated soil conditions; i.e., bogs and meadows (Holland and Keil 1995). Average precipitation is between 75 and 150 cm, most (70-90%) being received as snow in the winter (Holland and Keil 1995, Barbour and Minnich 2000). The mean annual temperature in lodgepole pine forest Habitat is about 5oC (Barbour and Minnich 2000).

    Lodgepole pine forest stands may occur in wet, flat areas or dry, steep slopes. The soils within this Habitat may be seasonally flooded and become saturated or are well-drained and on slopes (Sawyer and Keeler-Wolf 1995).

    As part of Barbour and Minnich’s (1995) “upper montane forests” classification, this Habitat generally occurs on shallow, rocky inceptisols or entisol soils (Barbour and Minnich 2000). Within California, lodgepole pine forest can dominate areas that are arid and windy on shallow soils or flat, wet sites adjacent to meadows or lakes (Barbour and Minnich 2000).

    Lodgepole pine forest in the San Jacinto Mountains occurs from 2600 m to the peak at 3234 m, although it may found as low as 2440 m on moist, shaded north-facing slopes (Thorne 1977). Within southern California, the Habitat most often occurs in upland areas interspersed with subalpine meadows (Sawyer and Keeler-Wolf 1995).

    Lodgepole pine forest, as a species, grows in a variety of environmental conditions, with extreme recorded temperatures between from -57oC to 38oC and precipitation ranging from 25 to 500 cm (Lotan and Critchfield 1990). Lodgepole pine forest is intolerant of the shade, growing best in full sunlight, and most often in moist soils (Lotan and Critchfield 1990). On the other hand, soils with underlying hardpan also support lodgepole pine forest and exclude other common trees (Lotan and Critchfield 1990).

    ECOSYSTEM PROCESSES

    There are varying opinions expressed in the literature as to the effects of fire on lodgepole pine forest. A study of fire within the San Jacinto Mountains showed that lightning-caused fires occurred every few years but were small and of low intensity (Stephenson and Calcarone 1999). This presumably matches historical trends, however, the cycle of large-scale fires in not known. Some stands have been fire free for 350 years whereas others appear to have cycle of 60 to 80 years (Barbour and Minnich 2000).

    According to Holland and Keil (1995), fire is important in the regeneration of lodgepole pine forest and result in even-aged stands. But fire is not essential for cones to open and seeds to be released. Lodgepole pine forest cones quickly mature, open and fall in the absence of fire (Sawyer and Keeler-Wolf 1995). According to Sawyer and Keeler-Wolf (1995), the stands are self-replacing and do not appear to be closely associated with or dependent on fire.

    Where fire has not occurred and soils are favorable, e.g., well-drained, red fir (Abies magnifica) may succeed this community (Holland and Keil 1995). Conversely, lodgepole pine forest may be invading montane meadows where, historically, fire kept seedling trees from reaching maturity (Holland and Keil 1995). Fire may allow lodgepole pine forest to overwhelm a site because seeds may be stored in a serotinous cone (Lotan and Critchfield 1990). Without fire, lodgepole locations may give way to common surrounding Habitats (Lotan and Critchfield 1990). Drought-tolerant species may also outcompete lodgepole pine forest in some settings (Holland and Keil 1995).

    There is debate as to whether lodgepole pine forest occurs as a climax state, evidenced by age structure, or whether it is regulated by fire, precipitation, or even episodic infections of the needle miner (Coleotechnites milleri) (Barbour and Minnich 2000). Either by fire, insect infestation, or logging, gaps in the lodgepole pine forest forest occur and undergo succession. Initially, herbaceous species quickly colonize the gaps, but lodgepole pine forest, with its large seed output, soon closes the gap (Bartolome 1988). Stands may become overcrowded, leaving them susceptible to insect infestation which may increase fuel availability for wildlife (Bartolome 1988). According to Pfister and McDonald (1980) the majority of extant stands of lodgepole pine forest are the result of catastrophic fires.

    THREATS

    Several fungal-infecting species threaten lodgepole pine forest growth and survivability. Lodgepole pine forest may be susceptible to pitch canker fungus (Fusarium subglutinans ssp. pini), a relatively new fungal pathogen in the southwestern region of the United States (Stephenson and Calcarone 1999). Pine engraver (Ips emarginatus) is another bark beetle known to attack lodgepole pine forest (Stephenson and Calcarone 1999). Lodgepole pine forest needle miner (Coleotechnites milleri) has been known to occur in outbreaks and result in stand replacement (Sawyer and Keeler-Wolf 1995). Lodgepole pine forest needle miner has always been a part of lodgepole pine forest Habitat; however, the insect population was regulated by natural predators and infestation caused forest openings which quickly regenerated (Holland and Keil 1995). DDT spraying allowed for outbreaks of needle miner which continue to be a problem (Holland and Keil 1995); today many natural predators have been eliminated resulting in larger infestations which threaten the regeneration of these forests. The most severe pest of lodgepole pine forest is the mountain pine beetle (Dendroctonus ponderosae) which often results in the thinning of the infected tree (Lotan and Critchfield 1990). The most serious parasite of lodgepole pine forest is dwarf mistletoe (Arceuthobium campylopodum), which severely damages its host plant (Lotan and Critchfield 1990).

    Historically, sheep grazing threatened lodgepole pine forest reproduction, however the removal of this pressure in areas such as Yosemite National Park has resulted in re-colonization of lodgepole pine forest (Holland and Keil 1995).

    LITERATURE CITED

    Barbour, Michael G. and Richard A. Minnich. 2000. Californian Upland Forests and Woodlands. In North American Terrestrial Vegetation. Second Edition. Cambridge University Press, Cambridge, UK.

    Bartolome, James W. 1988. Lodgepole pine forest. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 44-45.

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company. Dubuque, Iowa.

    Lotan, James E. and William B. Critchfield. 1990. Pinus contorta ssp. murrayana - lodgepole pine forest. Pages 302-313 in R.M. Burns and B. H. Honkala, technical coordinators. Silvics of North America, Volume 1. Conifers. Agriculture Handbook 654. USDA, Forest Service, Washington, D.C.

    Munz, Philip A. and David D. Keck. 1949. California Plant Communities. El Aliso, Vol. 2, No. 1: 87-105.

    Pfister, Robert D. and Philip M. McDonald. 1980. Lodgepole pine forest In Forest Cover Types of the United States and Canada. ed. F. H. Eyre. Society of American Foresters, Washington, D.C. pp. 97-98

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Sheppard, Paul R. and James P. Lassoie. 1998. Fire Regime of the Lodgepole pine forest Forest of Mt. San Jacinto, California. Madrono, Vol 45, No. 1:47-56.

    Stephenson, John R. and Gena M. Calcarone. 1999. Southern California mountains and foothills assessment: habitat and species Conservation issues. General Technical Report GTR-PSW-172. Albany, CA: Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture; 402 p.

    Thorne, Robert F. 1976. The Vascular Plant Communities of California. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 1-31.

    Thorne, Robert F. 1977. Montane and Subalpine Forests of the Transverse and Peninsular Ranges In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 538-557.

    Vogl, Richard J. 1976. An Introduction to the Plant Communities of the Santa Ana and San Jacinto Mountains. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 77-98.

    Lower Montane Coniferous Forest

    VEGETATION ASSOCIATION:

    LOWER MONTANE CONIFEROUS FOREST

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Because of the diversity of subassociations which may be included in this Habitat type, precise species composition for many of the mapped vegetation stands remains unknown. It is not likely that oak woodlands, grasslands, or shrub-dominated Habitat would be missed-mapped as lower montane coniferous forest. However, other Habitats dominated by conifers; i.e., Jeffrey pine forest and lodgepole pine forest, may be included in this Habitat category.

    BIOGEOGRAPHY

    The range described by Munz and Keck (1949) probably includes: areas mapped as Jeffrey pine forest; however, they describe this Habitat to occur, in California in the North Coast ranges between 915 and 1825 m, in northern California between 365 and 1675 m, in the Sierra Nevada between 600 and 2290 m, and in southern California between 1525 and 2450 m. Coulter pine forests range from 230 m in the Bay Area to 2290 on San Jacinto Mountain (Thorne 1977). The Coulter pine phase of this Habitat does not occur in the Sierra Nevada range, but rather along the coast ranges from San Francisco to Baja California, Mexico, including the Transverse and Peninsular ranges (Thorne 1977). Information regarding the biogeography of lodgepole pine forest was not reviewed.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Sawyer and Keeler-Wolf (1995) identify this Habitat at Hall Canyon and Millard Canyon Research Natural Area (RNA). A Coulter pine-black oak woodland is described at Cahuilla Moutain (Sawyer and Keeler-Wolf 1995). Ponderosa pine forest surrounds Lake Fulmor in the San Bernardino Mountains of Riverside County (Thorne 1977).

    The MSHCP vegetation map shows three areas supporting lower montane coniferous forest in the Plan Area: San Jacinto Mountains and Agua Tibia, totaling 9,097 acres. The San Jacinto Mountains stands occur both east and west of SR-243 from north of Lake Fulmor through Pine Cove, with large populations around Idyllwild, Mountain Center, and Lake Hemet. Lower montane coniferous forest has smaller occurrences in the Thomas Mountains east of Pine Meadow. This vegetation is located mainly within the Cleveland National Forest with the exception of a few large blocks around Idyllwild and Mountain Center. The Agua Tibia occurrence reaches slightly into the southern edge of the Plan Area, within National Forest land.

    VEGETATION CHARACTERISTICS

    As an ecotone between lower manzanita chaparral and higher conifer forests, lower montane coniferous forest represents a variety of subassociations. Due to wide discrepancies in the categorization of montane forests, several classification systems are presented here. However, the varying Habitat categories generally list similar constituent species within this Habitat type.

    Vogl (1976) identifies a mixed coniferous forest Habitat type occurring in the Santa Ana and San Jacinto Mountains. This Habitat type probably includes areas categorized in the Pacific Southwest Biological Services (PSBS 1995) mapping as lower montane coniferous forest, Jeffrey pine forest, and southern California white fir forest. Dominant tree species in this Habitat occur in a variety of densities and include ponderosa pine, black oak, interior live oak, canyon oak, Coulter pine, incense cedar, white fir, sugar pine and Jeffrey pine (Vogl 1976). Bracken fern (Pteridium aquilinum) is a common understory plant often occurring with perennial bunch grasses and meadow plants (Vogl 1976).

    Vogl’s (1976) description of lower montane coniferous forests is similar to the mixed conifer series of Sawyer and Keeler-Wolf (1995), where three or more coniferous trees are co-dominant in the canopy. The list of co-dominant trees, which occur in Western Riverside County, are black oak, big-cone Douglas-fir, incense-cedar, Jeffrey pine, ponderosa pine and/or white fir. The canopy is generally less than 70 m in height and intermittent, with shrubs occurring infrequently to commonly and ground cover varying from sparse to abundant (Sawyer and Keeler-Wolf 1995).

    Thorne (1976) identifies three subassociations of lower montane coniferous forests: Coulter pine forest, yellow pine forest, and mixed conifer forest. The last subassociation has not been described in Western Riverside County and a type of yellow pine forest is described in a separate Habitat account as Jeffrey pine forest.

    The Coulter pine subassociation of lower montane coniferous forest, over its California range, can include bigcone Douglas-fir (Pseudotsuga macrocarpa), black oak (Quercus kelloggii), canyon live oak (Q. chrysolepis), coast live oak (Q. agrifolia), interior live oak (Q. wislizenii) and/or ponderosa pine (Pinus ponderosa) (Sawyer and Keeler-Wolf 1995). Tree heights reach up to 30 m and generally form a continuous canopy (Sawyer and Keeler-Wolf 1995). The shrub layer underneath Coulter pines varies from frequent to infrequent with a sparse ground layer (Sawyer and Keeler-Wolf 1995). Coulter pine (Pinus coulteri) may also occur as a co-dominant with canyon live oak in a two-tiered canopy with infrequent shrubs and a sparse ground layer (Sawyer and Keeler-Wolf 1995).

    Coulter pine in southern California usually forms open stands with an understory of chaparral shrubs and pineland annual and perennial herbs (Thorne 1977). The lowest zone supports Coulter pine stands often along with black oaks. Manzanita chaparral species are also commonly found in this area, with chaparral whitethorn (Ceanothus leucodermis) as the most common shrub species (Thorne 1977, Vogl 1976). Most of the chaparral whitethorn present occurs in mature stands, awaiting fire to stimulate seed growth (Vogl 1976). This lower zone may “give-way” to dense brush or oak-Ceanothus chaparral (Thorne 1977). The upper portion of the Coulter pine subassociation may contain a variety of other pine species (Pinus spp.) and white fir (Abies concolor). In the Santa Ana mountains, Coulter pine stands intergrade with manzanita chaparral, stands of oaks (Quercus spp.), or big-cone Douglas-fir (Vogl 1976).

    The yellow pine forest identified by Thorne (1976) is dominated by either ponderosa or Jeffrey pine (Pinus jeffreyii) depending on the elevation and exposure, although the two species may occur together and may hybridize in some situations (Thorne 1977). Areas dominated by Jeffrey pines are described in a separate Habitat account. Ponderosa pine associates at lower elevations include Coulter pine and black oak (Thorne 1977). On north- and east-facing slopes, i.e., more mesic conditions, ponderosa pine is associated with big-cone Douglas-fir, incense-cedar, canyon live oak and Pacific dogwood. At higher elevations, under similar conditions, white fir and sugar pine (Pinus lambertiana) may co-occur (Thorne 1977). Scattered shrubs include manzanitas, deer brush (Ceanothus integerrimus), yerba santa (Eriodictyon trichocalyx), chinquapin (Chrysolepis sempervirens), thimbleberry (Rubus parviflorus), silk tassel bush (Garrya flavescens), grape soda lupine (Lupinus excubitus), lupine (L. formosus), cherry (Prunus spp.)California coffeeberry (Rhamnus californica), Sierra gooseberry (Ribes roezlii) and nightshade (Solanum xanti). A diverse herbaceous layer is comprised of many of the following species, which are absent from Jeffrey pine forests: morning-glory (Calystegia occidentalis ssp. fulcrata); sedge (Carex multicaulis); clarkia (Clarkia rhomboidea); collinsia (Collinsia childii); bird’s-beak (Cordylanthus rigidus); eriastrum (Eriastrum densifolium); splendid gilia (Gilia splenden); iris (Iris hartwegii); whisker brush (Linanthus ciliatus); phacelia (Phacelia imbricata); campion (Silene lemmoneii); Laguna Mountains jewelflower (Streptanthus bernardinus) and violet (Viola purpurea). Other species include native perennial grasses like California brome (Bromus carinatus var. carinatus), brome (B. orcuttianus), melic (Melica imperfecta), and bluegrass (Poa scabrella). Other common understory species include: western dwarf mistletoe (Arceuthobium campylopodum); Indian milkweed (Asclepias eriocarpa); desert Indian paintbrush (Castilleja angustifolia); pincushion (Chaenactis santolinoides); little prince’s-pine (Chimaphila menziesii); Johnston’s bedstraw (Galium johnstonii); monkeyflower (Mimulus johnstonii); beardtongue (Penstemon grinnellii); and P. labrosus, snowplant (Sarcodes sanguinea); and squirreltail (Elymus elymoides ssp. elymoides) (Thorne 1976; Thorne 1977).

    Two unique species occurrences may fall within lower montane coniferous forest, knobcone pine (Pinus attenuata) and Tecate cypress (Cypressus forbesii). Knobcone pine is found in the Santa Ana mountains in the vicinity of Pleasants Peak and from Sugarloaf Peak (Thorne 1977, Vogl 1976). The closed-cone knobcone pine also is known from the south face of the San Bernardino Mountains (Thorne 1977) and along Hwy 84 below Idyllwild. Trees are short-lived and occur in even-aged stands (Vogl 1976). Another unique species occurrence is that of Tecate cypress which occurs on the northwest slopes of Sierra Peak (Vogl 1976).

    PHYSICAL ENVIRONMENT

    Average precipitation in the lower montane zone is between 65 and 200 cm and the growing season is between four to seven months (Munz and Keck 1949). Yearly maximum temperatures are between 27oC and 34oC and minima are between –5oC and 1oC (Munz and Keck 1949). According to Sawyer and Keeler-Wolf (1995), this general Habitat occurs most commonly in shallow soils on well-drained slopes of any aspect, between 900 and 2200 m. Lower montane coniferous forest typically occurs between 1525 and 2500 m in the San Jacinto Mountains (Vogl 1976). This Habitat occurs on ridgetopes, knolls, and southern exposures above chaparral but below higher coniferous forest types (Vogl 1976).

    Elevations for the Coulter pine association stretch from 700 to 2000 m over the state (Sawyer and Keeler-Wolf 1995). Coulter pine stands generally occur on shallow, well-drained soils on all slope aspects (Sawyer and Keeler-Wolf 1995). In the San Jacinto Mountains, this lower zone may occur as low as 600 m. Dense brush and oak-Ceanothus chaparral, typical of the lowest part of this area, occur on dry, steep slopes on southern exposures. The upper zones, often dominated by ponderosa pine, as described by Thorne (1976), occur on mesic slopes, between 1,375 and 2,135 m (Thorne 1977).

    The knobcone pine occurrence in the Santa Ana Mountains occurs on hydrothermally-modified serpentinite (Vogl 1976). It is thought that the characteristics of the tree, including “scattered growth, multiple-trunked trees, spreading crowns, and medium-length needles, contribute to the maintenance of the pines by enhancing their ability to intercept marine air and produce considerable fog drip which is readily held by the soil (Vogl 1976).” The knobcone pines have a closed-cone habit which requires periodic fires for shedding of the seed (Vogl 1976). Similarly, Tecate cypress cones generally remain closed until a fire (Vogl 1976). Tecate cypress occurs on eroded clay, shales, sandstone, and conglomerate soils (Vogl 1976).

    ECOSYSTEM PROCESSES

    Ponderosa pine groves show signs of surviving lightning and past fires, although knowledge concerning the effects of fire or other ecosystem processes is not well developed. Coulter pine forests are adapted for rapid regeneration following fires (Stephenson and Calcarone 1999). However, in the Santa Ana mountains, Vogl (1976) observed that Coulter pines do not compete well with manzanitas after a burn. This is despite several adaptations to fire including semi-serotinous cones and relatively short life-span, which allow it to readily reseed burn sites and grow quickly after crown fires (Vogl 1976, Stephenson and Calcarone 1999). Vogl (1976) hypothesizes that the trees may require a specific natural fire frequency and intensity and may suffer from fire suppression activity, as evidenced by Vale (1979).

    THREATS

    Vogl (1976) warns that fire suppression activities, which cause the excessive accumulation of dead chaparral whitethorn, also may result in burns which may be “abnormally hot and destroy the pines and reduce the oaks, thereby raising the lower limits of the treeline” (Vogl 1976). The USDA (1999) identifies the largest threat to Coulter pine as multiple fires in short succession (e.g., less than twenty-five years). If trees are killed before an adequate seed crop has developed, the Habitat may revert to chaparral.

    Insect and disease also threaten the Coulter pine (Stephenson and Calcarone 1999). An example of insect threats to Coulter pine was the loss of nearly 70 percent of overstory Coulter pine at Palomar Mountain in the late 1980's due to a bark beetle epidemic (Stephenson and Calcarone 1999). Coulter and ponderosa pines may be susceptible to pitch canker fungus (Fusarium subglutinans ssp. pini), a relative new fungal pathogen in the southwestern region of the United States (Stephenson and Calcarone 1999). The western dwarf mistletoe (Arceuthobium campylopodum) infects both ponderosa and Coulter pines in this Habitat. Mistletoe is managed mainly through the pruning and removal of infected trees (Stephenson and Calcarone 1999). Bark beetles in southern California are generally of the genera Dedroctonus and Ips, with the western pine beetle (Dendroctonus brevicomis) causing widespread Coulter pine mortality during the early 1990's (Stephenson and Calcarone 1999). Ponderosa pines are attacked by ponderosa pine beetle (Dendroctonus ponderosae). When the populations of these beetles are high, often during or following a drought, they may kill even healthy trees (Stephenson and Calcarone 1999).

    Although timber harvests largely have been discontinued except to supply local demand, high intensity harvests from the 1950's to the mid-1970's contributed to the reduction in the number of large trees and possibly to the increase of white fir in this Habitat (Stephenson and Calcarone 1999). In (1976), Vogl observed that a considerable portion of the Coulter pines in the Santa Ana mountains had been cut for sanitation, fire control, and reforestation.

    Ponderosa pines have been shown to suffer foliage damage from ozone (Stephenson and Calcarone 1999). The highest rates of mortality occur when ozone damage occurs during a drought and trees cannot withstand attacks by bark beetles (Stephenson and Calcarone 1999). Forest areas exposed to heavy air pollution are most susceptible to this injury. Nitrogen deposition is thought to occur in those areas of high air pollution as well. Although it has some positive effects, e.g., increased soil fertility and surface litter decomposition rates, negative effects of nitrogen deposits include nutrient deficiencies, soil acidification, altered species composition (i.e., increase in white fir understory), decreases in mycorrhizal root symbiosis, and elevated nitrate in the soil (Stephenson and Calcarone 1999). Ponderosa pine is also susceptible to smog damage which resulted in clear-felling of dying stands in the San Bernardino Mountains (Thorne 1977).

    LITERATURE CITED

    Munz, Philip A. and David D. Keck. 1949. California Plant Communities. El Aliso, Vol. 2, No. 1: 87-105.

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Stephenson, John R. and Gena M. Calcarone. 1999. Southern California mountains and foothills assessment: habitat and species Conservation issues. General Technical Report GTR-PSW-172. Albany, CA: Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture; 402 p.

    Thorne, Robert F. 1976. The Vascular Plant Communities of California. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 1-31.

    Thorne, Robert F. 1977. Montane and Subalpine Forests of the Transverse and Peninsular Ranges In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 538-557.

    Vale, Thomas R. 1979. Pinus coulteri and Wildfire on Mount Diablo, California. Madrono, Vol.26, No. 3:135-140.

    Vogl, Richard J. 1976. An Introduction to the Plant Communities of the Santa Ana and San Jacinto Mountains. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 77-98.

    Mixed Evergreen Forest

    VEGETATION ASSOCIATION:

    MIXED EVERGREEN FOREST

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Because of the ambiguity in the definition for mixed evergreen forest, precise descriptions of mixed evergreen forest do not occur in the literature. For that reason, it is difficult to determine the specific vegetative characteristics of the areas mapped as mixed evergreen forest in the San Jacinto Mountains. It is likely that these areas generally contain an odd mixture of broadleaved species with various conifer species, described in other conifer Habitats.

    BIOGEOGRAPHY

    Again, the literature differs in characterization of mixed evergreen forest range. Some authors separate northern and southern California Habitats while others describe the Habitat to cover much of California and portions of Oregon. Thorne (1976) describes this Habitat as two varieties, a northern and southern. Northern California mixed evergreen forest occurs inland from redwood forest to the Klamath and Coast ranges on more xeric sites (Thorne 1976). Southern mixed evergreen forest occurs in areas where there is a transition between riparian forests and yellow pine (i.e., lower montane coniferous forest). As defined by Barbour and Minnich (2000) the community occurs around the Central Valley between 600 and 1,200 m on the Sierra side and between 300 and 1,500 m on the Coast range side. In the north it reaches through the Klamath mountains into Oregon; in the south it occurs between 1,275 and 1,425 m in the Transverse, Peninsular and Baja ranges (Barbour and Minnich 2000). Holland and Keil (1995) describe the range of mixed evergreen communities to stretch from southern Oregon through the Klamath-Siskiyou Mountains, on the coastal mountains, and into San Diego County.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Occurrences of mixed evergreen forest were mapped west of Lake Hemet in the vicinity of Rouse Ridge, north of Marion Ridge, east of Dark Canyon and south of Black Mountain all within the San Jacinto Mountains. Mixed evergreen forest occupies 7,965 acres of the total Plan Area.

    VEGETATION CHARACTERISTICS

    Due to the general nature of the Habitat description, the literature concerning the constituent species of mixed evergreen forest is similarly vague. In general, there is overlap with the descriptions of broadleaved upland forest and lower montane coniferous forest. Mixed evergreen forest generally has been used to map vegetation occurring between oak woodlands and montane conifer forests, including the following associations: Coulter pine, big-cone Douglas fir, mixed hardwood, Douglas fir-hardwood, tan oak-madrone, and Santa Lucia fir forests (Barbour and Minnich 2000). Presented below are different interpretations of the Habitat occurrence.

    Thorne (1976) and Brown (1982) describe mixed evergreen forest as a Habitat containing a canopy of bigcone Douglas-fir (Pseudotsuga macrocarpa), Pacific madrone (Arbutus menziesii), incense-cedar (Calocedrus decurrens), Coulter pine (Pinus coulteri), bigleaf maple (Acer macrophyllum), coast live oak (Quercus agrifolia), canyon live oak (Q. chrysolepis), and California bay (Umbellularia californica). As Munz and Keck (1949) describe this Habitat, it does not occur in southern California. Anderson (1988) describes a montane hardwood-conifer series composed of at least one-third broadleaf trees and one-third conifers. This Habitat probably corresponds to areas mapped as mixed evergreen forest in the Plan Area. In southern California, Anderson (1988) adds black oak (Quercus kelloggii) and ponderosa pine (Pinus ponderosa) to the species listed above. Coulter pine can become a dominant element in this Habitat type in southern California, sometimes occurring co-dominant with big-cone Douglas-fir (Barbour and Minnich. 2000). This description most likely corresponds to the lower phase of lower montane coniferous forest.

    The occurrence of big-cone Douglas-fir is likely to be scattered and similar to its occurrence in broadleaved upland forest. Because an extensive discussion of big-cone Douglas-fir ecology is presented in the broadleaved upland forest Habitat account, it will not be repeated here. Please refer to the discussion in that section as it likely applies to portions of this Habitat as well.

    Typically, the conifers are generally between 30 to 65 m in height and the lower broadleaved canopy 10 m to 30 m (Anderson 1988). The conifer overstory may not always be present. The broadleaf understory is generally a closed canopy with evergreen and some deciduous trees (Barbour and Minnich 2000).

    Usually, this Habitat contains little understory although shrubs may be important in ecotone or post-burn situations (Anderson 1988). The understory typically is comprised of: elk clover (Aralia californica); hillside gooseberry (Ribes californicum); and poison-oak (Toxicodendron diversilobum), and a set of herbs and herbs including round-leaved boykinia (Boykinia rotundifolia), miner’s lettuce (Calytonia perfoliata), Darisca glomerata, common chickweed (Stellaria media), and wood fern (Dryopteris arguta). Understory shrubs, herbs and mosses generally comprised less than 25% of the overall percent cover (Barbour and Minnich 2000). The annual grasses common in oak woodlands do not extend into the mixed evergreen forests.

    PHYSICAL ENVIRONMENT

    Because descriptions of the Habitat vary widely, descriptions of the physical environment also vary. In general, the Habitat occurs between 500 and 1800 m in southern California in moist, mesic areas with the exception of the Coulter pine phase which occurs on xeric sites.

    Thorne (1976) places the elevation limits of this community between 500 to 1675 m within southern California. The Habitat generally occurs in moister canyons associated with riparian woodlands but also may intergrade with chaparral or yellow pine forest in drier areas. In southern California, Anderson (1988) identifies mixed evergreen forest in coarse, well-drained mesic sites, often on steep terrain within narrow valleys, from 605 to 1760 m (Anderson 1988). Average rainfall is between 60 and 170 mm and air temperatures are between -2oC and 36oC (Anderson 1988). When compared with oak woodlands, this Habitat occurs in significantly cooler areas (mean annual temperature is 14C) which receive greater precipitation (Barbour and Minnich 2000).

    The Coulter pine phase, which more likely belongs in the lower montane coniferous forest Habitat, occurs between 1200 and 1800 m in the Transverse and Peninsular ranges (Barbour and Minnich 2000). The upper portions of this Habitat can be considered to have three dominants: canyon live oak, Coulter pine and big-cone Douglas-fir, where the first two occur on more xeric and frequently disturbed sites and the first and last occur on more mesic, fire-protected sites (Barbour and Minnich 2000).

    ECOSYSTEM PROCESSES

    Following a fire or other disturbance, recovery in mixed evergreen forest will occur at different rates among conifers and hardwoods (Anderson 1988). Typically, mesic sites support more rapid conifer regrowth whereas hardwoods are dominant for longer on xeric sites (Anderson 1988). The conifer component is typically fully grown after 30 to 50 years to develop while the broadleaved phase usually requires 60-90 years (Anderson 1988). The Coulter pine phase of this Habitat type seems to be an obligate fire type due to the presence of serotinous cones, although some contend that this only is true where the trees are adjacent to chaparral.

    THREATS

    Information regarding specific pest infections of this Habitat are not readily available. However, different pests associated with specific trees are likely to affect trees in this Habitat. Specific information regarding the effects of fire suppression also is lacking in the literature. It can be stated, however, that fire suppression activity has led to denser growth hindering the establishment of those seedlings requiring bare ground (Holland and Keil 1995).

    LITERATURE CITED

    Anderson, Richard. 1988. Montane Hardwood-Conifer. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 70-71.

    Barbour, Michael G. and Richard A. Minnich. 2000. Californian Upland Forests and Woodlands. In North American Terrestrial Vegetation. Second Edition. Cambridge University Press, Cambridge, UK.

    Brown, D.E. 1982. Californian evergreen forest and woodland. Desert Plants 4:66-69.

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company. Dubuque, Iowa.

    Munz, Philip A. and David D. Keck. 1949. California Plant Communities. El Aliso, Vol. 2, No. 1: 87-105.

    Sawyer, John O., Dale A. Thornburgh and James R. Griffin. 1977. Mixed Evergreen Forest In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 359-381.

    Thorne, Robert F. 1976. The Vascular Plant Communities of California. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 1-31.

    Oak Woodland

    VEGETATION ASSOCIATION:

    OAK WOODLAND

    MAPPED SUBASSOCIATIONS

    Dense Engelmann Oak; Coast Live Oak

    DATA CHARACTERIZATION

    Because of the use of an undifferentiated oak woodland community type and the similarity between oak woodland and broadleaved upland forest, it is difficult to determine the species composition of mapped areas without field verification. It is probable that areas containing oaks were mapped within a category which includes oaks as a dominant species; i.e., oak woodland, dense Engelmann oak woodland, coast live oak woodland, broadleaved upland forest.

    Oak woodlands are generally easily interpreted from other vegetation types. Mapping errors may occur near areas of mesic chaparral.

    BIOGEOGRAPHY

    Holland and Keil (1995) describe the oak woodlands as being nearly restricted to the state of California. Most authors distinguish southern oak woodlands from other oak woodlands in other parts of the state. There are discrepancies in the description of the northern limit of southern oak woodlands, varying from Mendocino County to northern Los Angeles County. Coast live oaks range through a large portion of the state, extending outside the range of the typically-defined southern oak woodlands.

    Thorne (1976) distinguishes between northern, foothill, southern, and island oak communities in California. Southern and coastal woodlands, including coast live oak woodland, extend from eastern Mendocino County at 40N latitude through the North Coast, Central Coast, and Transverse ranges on north-facing and coast-facing slopes and in canyons below 1200 m (Barbour and Minnich 2000). The range continues through the interior valleys and foothill slopes of the Peninsular ranges, mainly between 150 and 1,400 m, and south to the Sierra San Pedro Martir at 30N latitude (Barbour and Minnich 2000). According to Munz and Keck (1949), the southern oak woodlands are found in the valleys of southern California between Los Angeles and San Diego counties to about 1,525 m in the San Jacinto Mountains. According to Holland and Keil (1995), coast live oak woodlands range from Sonoma County to Baja California, generally in mesic areas including canyon bottoms and north-facing slopes, whereas southern oak woodlands extend from Ventura County southward. This roughly corresponds with Griffin (1977) who distinguishes oak woodlands from the Santa Ynez Mountains of Santa Barbara County southward as southern oak woodland.

    The Englemann oak, in the U.S., occurs only in San Diego, Riverside, Orange and Los Angeles counties (Scott 1991). Engelmann oak woodlands, which are confined within the distribution of the more general southern oak woodlands, occur from the San Gabriel Mountains to Baja California but are most common in the foothills of San Diego and southwestern Riverside counties (Stephenson and Calcarone 1999). Major occurrences of Engelmann oak woodland are generally between the Palomar and Cuyamaca Mountains and on the Santa Rosa Plateau (Stephenson and Calcarone 1999).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Oak woodlands occupy approximately 2.7 percent of the Plan Area (35,330 acres). These woodlands generally are distributed near drainages in the steeply sloped portions of the western Riverside Plan Area. Large acreage of live oak woodlands occur in three large clusters including the Cleveland National Forest from the Santa Ana Mountains near Glen Ivy south toward San Mateo Canyon; between Hemet and Sage from north of Lake Skinner to Bautista Creek; and from Calimesa to the Cherry Valley and the Morongo Indian Reservation. Large individual patches occur south of Banning east of Mount Edna, north of Estelle Mountain, and in the Santa Rosa East area west of Sandia Canyon. A few scattered stands of live oak woodlands occur south of the urbanized highway 91 corridor, and on the Gavilan Plateau.

    From the literature, Engelmann oak woodlands are described from the following areas: Organ Valley (Cleveland National Forest) on Black Mountain is in an area which was nominated as a Research Natural Area (RNA) (USDA 1986); the Santa Rosa Plateau contains one of the largest, undeveloped Engelmann oak savannas; scattered populations of Engelmann oak occur on the southern and western edges of the Perris Plain (including Corona and Hemet); and on the ridge between the Coachella Valley and mountains south of Santa Rosa peak (Scott 1990).

    Dense Engelmann oak woodlands were mapped in the southwestern portion of the Plan Area from San Mateo Canyon to Sandia Canyon on the Santa Rosa Plateau. Two stands of dense Engelmann oak woodland occur south of Lake Elsinore near Elsinore Peak, and along a drainage on the Gavilan Plateau.

    Undifferentiated oak woodlands which may include live oak, Engelmann oak or possibly canyon live oak and scrub oak stands, occurred in three general regions. The largest area is the eastern portion of the Plan Area in the San Jacinto Mountains from the San Gorgonio River south to Table Mountain. Substantial acreage of oak woodland also occurs in a ring around the urbanized portions of Rancho California with the largest areas in the Santa Rosa East, Pechanga Indian Reservation, and north of the Black Hills area. The Santa Ana mountains also support several patches of oak woodland.

    VEGETATION CHARACTERISTICS

    The two variants of oak woodlands within the Plan Area are either dominated by Engelmann oak (Quercus engelmannii), often in drier areas, or coast live oak (Q. agrifolia), often on moister slopes (Thorne 1976).

    Coast live oak woodland. Many understory plants in a coast live oak woodland are shade tolerant and include wild blackberry (Rubus ursinus), snowberry (Symphoricarpos mollis), California walnut (Juglans californica), California-lilac (Ceanothus spp.), Rhus spp.currant (Ribes spp.), toyon (Heteromeles arbutifolia), California bay (Umbellularia californica), Engelmann oak, manzanita (Arctostaphylos spp.), laurel sumac (Malosma laurina), poison-oak (Toxicodendron diversilobum) and herbaceous plants including bracken fern (Pteridium aquilinum), polypody fern (Polypodium californicum), fiesta flower (Pholistorma auritum) and miner’s lettuce (Claytonia perfoliata) (Holland and Keil 1995, Sawyer and Keeler-Wolf 1995, Thorne 1976, Brown 1982). Munz and Keck (1949) identify similar species for this Habitat and include that a variety of grasses and soft shrubs also are commonly found.

    The physiognomy of coast live oak woodlands varies from savanna-like, with few to no woody associates, to dense woodlands, typically with canyon live oak and/or Engelmann oak (Brown 1982). Coast live oak trees can reach a canopy height of 30 m, but usually vary from nine to 22 m (Sawyer and Keeler-Wolf 1995; Barbour and Minnich 2000). Canopy coverage varies between continuous to open (Sawyer and Keeler-Wolf 1995). Shrub cover is occasional or common with the ground layer varying from grassy to absent (Sawyer and Keeler-Wolf 1995). Woodlands may intergrade with grasslands in which cases shrub cover becomes diminished and herbaceous cover can reach 80% (Holland and Keil 1995; Barbour and Minnich 2000). The shrub component can be quite developed in areas where coast live oak intergrades with chaparral or coastal sage scrub.

    Dense Engelmann oak woodland. Engelmann oak occurs infrequently as pure stands and more commonly as a subdominant to coast live oak (Stephenson and Calcarone 1999). As a subdominant, it may be found in typical southern oak woodlands, with a canopy cover varying from 10 to 50%. Engelmann oak may also be found in dense riparian / oak woodlands with a closed-canopy in which it is usually associated with a water-course or canyon bottom (Scott 1990). The riparian / oak woodlands very rarely support dense stands of Engelmann oaks. Scott (1991) sampled over 30,000 ha of Engelmann oak occurrences and found only 1.6% to occur as pure stands compared with 52% occurrence of Engelmann oak as a subdominant to coast live oak.

    Although accurate historical data regarding Engelmann oak woodland composition is lacking, Wieslander’s vegetation type map (VTM) surveys indicate that some stands were composed of 10-50% tree cover with 90% of the trees being Engelmann oaks while other stands contained 100% tree cover with a mixture of Engelmann and coast live oak or Engelmann oak, coast live oak, and black oak (Barbour and Minnich 2000). Sawyer and Keeler-Wolf (1995) describe portions of this Habitat which are dominated by Engelmann oak or where it is an important component with coast live oak as having a closed or open canopy up to 18 m tall. Engelmann oaks, when mature, typically have a canopy height between 10 and 20 m (Scott 1990). Shrubs may occur commonly or infrequently and ground cover is generally grassy or sparse (Sawyer and Keeler-Wolf 1995). The grassy understory of Engelmann oak woodlands usually is dominated by exotic species: i.e., bromes (Bromus spp.)oats (Avena spp.)barley (Hordeum spp.)filaree (Erodium spp.), but may also contain native grasses; i.e., needlegrass (Nassella spp.) (Scott 1990). Engelmann oak hybridizes with scrub oak (Quercus berberidifolia) and hybrids generally occur on the margins between scrub oak stands and Engelmann oak stands (Sawyer and Keeler-Wolf 1995).

    PHYSICAL ENVIRONMENT

    In general, oak woodlands occur from 10 to 1500 m (Holland and Keil 1995). Average annual rainfall is between 38 and 64 cm with rapid runoff, and the growing season is seven to 10 months (Munz and Keck 1949). Yearly maximum temperatures are between 290C and 33C and minima are between 00C and 70 C (Munz and Keck 1949). Engelmann oak appears limited to areas below 1300 m that receive more than 37 cm annual precipitation (Scott 1990).

    Generally, oak woodlands are open where moisture is limited, in drier more exposed aspects, and densest in moist areas (Holland and Keil 1995). North-facing slope occurrences are also denser than south-facing slope occurrences (Holland and Keil 1995). Coast live oaks occur more frequently on cooler, steeper slopes and Engelmann oaks on warm slopes (Scott 1990). Engelmann oak woodland can be found in relatively flat valleys, on foothill slopes and adjacent to, but above, stream channels (Stephenson and Calcarone 1999). However, in a canyon containing a stream bottom, Engelmann oaks are more likely to occur on the upper margins of the canyon whereas coast live oak will most often occur on the terraces adjacent to the stream channel forming the interior of the woodland canopy (Scott 1990). Scott (1991) found that Engelmann oaks were most concentrated where slopes were southwest-facing, between 0o and 10o and elevation between 700 and 1250 m. Snow (1973) found that coast live oaks are associated with rock outcrops whereas Engelmann oaks are not.

    Common soils which support coast live oak include sandstone and shale-derived soils (Sawyer and Keeler-Wolf 1995). Coast live oak typically occupies slopes with deep soils, alluvial terraces, and the recent alluvium of canyon bottoms (Griffin 1977; Brown 1982). Open woodlands form when soils are shallow (Holland and Keil 1995). Engelmann oak tends to occur on deep clay soils from a gabbro or basalt substrate (Stephenson and Calcarone 1999).

    ECOSYSTEM PROCESSES

    In comparing burned and non-burned oak woodlands on the Santa Rosa Plateau, Lathrop and Osborne (1991) found that although more seedling and sapling oaks were top-killed in burned areas, they resprouted more vigorously and total overall survival was slightly higher in the burned sites. In comparing coast live oak and Engelmann oak survival rates after a burn, coast live oak had a lower long-term survival rate, although mature individuals of both species were generally unaffected (Lathrop and Osborne 1991). Lathrop and Osborne (1991) suggest that prescribed burns occur in the late summer or fall so that resprouts may avoid the summer drought conditions. Following a fire or during a drought, scrub oak and hybrid oaks were found to outcompete Engelmann oak, which may account for the limited distribution of Engelmann oak (Scott 1990).

    Oak trees, in general, require 60 to 80 years to mature (Holland 1988). Engelmann oaks typically live between 50 and 80 years, but can reach 350 years of age (Scott 1990). The two species have distinct reproductive characteristics. Natural germination of Engelmann oak acorns occurs in the early winter on the Santa Rosa Plateau compared with later winter and spring germination of coast live oak acorns (Lathrop and Osborne 1990). As an adaptation to the seasonal timing of germination, coast live oak acorns appear much more sensitive to available moisture when compared with Engelmann oak acorns which germinate more easily due to a higher initial moisture content (Snow 1991). Engelmann oak also appears to delay shoot development, compared with the more rapid development of coast live oak, which may allow Engelmann oaks to absorb more moisture on more exposed Habitats (Snow 1991).

    Regeneration of Engelmann oaks is more dependent on the site conditions where individual acorns fall than percent of viable acorns produced (Lathrop and Osborne 1990). Engelmann oak seedlings are more abundant in the shade. However, within a general limit, seedling survival increases as shade decreases (Lathrop and Osborne 1990). Seedlings also show an affinity to be under the dripline of the parent tree, an area which presumably retains more soil moisture (Lathrop and Osborne 1990). This area, under the canopy, is also the most active area for pocket gophers (Thomomys bottae), which have a detrimental effect on regeneration because they predate acorns; they also aerate the soil which is a beneficial effect on regeneration (Lathrop and Osborne 1990). Seedling survival is enhanced when acorns are buried in litter or soil, protecting them from predation and enhancing their ability to root in the ground (Lathrop and Osborne 1990). Because of the need for fairly specific site conditions, regeneration of Engelmann oaks is dependent on undisturbed conditions. Studies have shown that although cattle grazing does significantly decrease seedling survivorship, drought conditions cause the most substantial seedling mortality (Lathrop and Osborne 1990). By comparison, coast live oak is more well adapted to drought conditions (Snow 1972).

    THREATS

    Holland and Keil (1995) state that in the vast majority of California oak woodland sites, oak reproduction ceased around 1900. The loss of acorn viability can be attributed to cattle and sheep in rangelands and an overabundance of deer in many northern California areas (Holland and Keil 1995). The oak woodland Habitat also has been altered by the replacement of native bunch grasses with exotic annual grasses which produce many more seeds. Man’s reduction in the number of predators of seed-eating animals which predate on oak acorns, also has been found to be a threat (Holland and Keil 1995). Introduced annual grasses, due to their rapid growth and uptake of available surface water, also contribute to the loss of native grasses historically present in oak woodlands and savannas as well as diminishing water supplies for oak seedlings (Stephenson and Calcarone 1999). In some areas, it appears that California bay is replacing coast live oak, possibly due to grazing (Holland 1988). In general, regeneration problems in coast live oak are not considered to be as serious as that of blue oak and valley oak (Holland 1988). Wood cutting, although not as prevalent in the southern portion of the state, has left areas of stumps because oaks were not able to reestablish (Holland 1988). Root rot, caused by overwatering during the summer in urban oaks, also has been known to cause mortality (Holland and Keil 1995).

    In a study of size class on the Santa Rosa Plateau, Lathrop et al. (1991) found that Engelmann oak reproduction in sites where grazing has ceased for the past four years is not sufficient to “maintain the present occurrence and abundance of older age classes in the future.” In the one site where grazing has been excluded for the past 20 years, young trees were more readily observed but regeneration rates still would not maintain current stand characteristics (Lathrop et al. 1991).

    Management techniques to improve oak regeneration include rotational grazing practices and retention of oaks on rangelands. The benefits of oaks in rangelands include enriched soils and reduced erosion (Holland and Keil 1995). Prescribed fires in broadleaf woodlands were seldom used in the Cleveland National Forest as of 1986 (USDA 1986). However, in the 1986 Forest Service’s Land and Resources Management Plan for the Cleveland National Forest, prescribed fires were identified as beneficial and were therefore increased in frequency in order to strengthen oak regeneration and help manage chaparral. The use of dead and down oak wood as fuel wood was identified as an opportunity in the Forest Service Management Plan, whereas the cutting of healthy oaks for timber purposes was not identified and therefore is not thought to be a threat on public land (USDA 1986).

    Declines in oak woodland Habitat type are due mainly to Habitat loss on private lands (Stephenson and Calcarone 1999). The majority of Engelmann oaks (57 percent) are held in private lands, with the next largest portion (24 percent) controlled by the Cleveland National Forest (Scott 1991). Because of the scattered distribution of Engelmann oaks within its range, Scott (1991) suggests that this oak is vulnerable to adjacent conditions and human activities. The Land and Resource Management Plan for the Cleveland National Forest (USDA 1986) identifies Engelmann oak woodland as a Habitat suffering from a limited distribution and deteriorating state due to overgrazing and development. The Plan also identifies these woodlands as important for recreation, recognizing that many campgrounds are in oak groves and dispersed recreation use mainly occurs in oak groves. These uses, thought to be increasing, along with grazing usage, contribute to reduction of shrub and herbaceous layers and disruption of natural regeneration (USDA 1986).

    LITERATURE CITED

    Barbour, Michael G. and Richard A. Minnich. 2000. Californian Upland Forests and Woodlands. In North American Terrestrial Vegetation. Second Edition. Cambridge University Press, Cambridge, UK.

    Brown, D.E. 1982. Californian evergreen forest and woodland. Desert Plants 4:66-69.

    George, Melvin R. 1991. Grazing and Land Management Strategies for Hardwood Rangelands. In Proceedings of the Symposium on Oak Woodlands and Hardwood Rangeland Management. Richard B. Standiford, Technical Coordinator. USDA Forest Service General Technical Report, PSW-126. Davis, California: 315-319.

    Griffin, James R. 1977. Oak Woodlands In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 383-415.

    Holland, V. L. 1988. Coastal Oak Woodland. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 78-79.

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company.. Dubuque, Iowa.

    Lathrop, Earl. W. and Chris D. Osborne. 1990. From Acorn to Tree: Ecology of the Engelmann Oak. Fremontia. Vol.18, No.3: 30-35.

    Lathrop, Earl. W. and Chris D. Osborne 1991. Influence of Fire on Oak Seedlings and Saplings in Southern Oak Woodland on the Santa Rosa Plateau Preserve, Riverside County, California In Proceedings of the Symposium on Oak Woodlands and Hardwood Rangeland Management. Richard B. Standiford, Technical Coordinator. USDA Forest Service General Technical Report, PSW-126. Davis, California: 366-370.

    Lathrop, Earl. W., C. Osborne, A. Rochester, K. Yeung, S. Soret and R. Hopper. 1991 Size Class Distribution of Quercus engelmannii on the Santa Rosa Plateau, Riverside County California. In Proceedings of the Symposium on Oak Woodlands and Hardwood Rangeland Management. Richard B. Standiford, Technical Coordinator. USDA Forest Service General Technical Report, PSW-126. Davis, California: 371-376

    McDonald, Philip A. and Edward E. Littrell. 1976. The Big-cone Douglas-Fir-Canyon Live Oak Community in Southern California. Madrono, vol. 23:310-320.

    Munz, Philip A. and David D. Keck. 1949. California Plant Communities. El Aliso, Vol. 2, No. 1: 87-105.

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Scott, Thomas A. 1990. Conserving California’s Rarest White Oak: The Engelmann Oak. Fremontia. Vol 18, No. 3: 26-29.

    Scott, Thomas A. 1991. The Distribution of Engelmann Oak in California. In Proceedings of the Symposium on Oak Woodlands and Hardwood Rangeland Management. Richard B. Standiford, Technical Coordinator. USDA Forest Service General Technical Report, PSW-126. Davis, California: 351-359.

    Snow, Gerald E. 1991. Germination Characteristics of Engelmann Oak and Coast Live Oak from the Santa Rosa Plateau, Riverside County, California. In Proceedings of the Symposium on Oak Woodlands and Hardwood Rangeland Management. Richard B. Standiford, Technical Coordinator. USDA Forest Service General Technical Report, PSW-126. Davis, California: 360-365.

    Snow, Gerald E. 1973. Some Factors Controlling the Establishment and Distribution of Quercus agrifolia and Q. engelmannii in Certain Southern California Oak Woodlands. Doctor of Philosophy Thesis submitted to Oregon State University, Oregon.

    Stephenson, John R. and Gena M. Calcarone. 1999. Southern California mountains and foothills assessment: Habitat and species Conservation issues. General Technical Report GTR-PSW-172. Albany, CA: Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture; 402 p.

    Thorne, Robert F. 1976. The Vascular Plant Communities of California. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 1-31.

    USDA, Forest Service. 1986. Land and Resources Management Plan: Cleveland National Forest. USDA, Forest Service, Cleveland Nation Forest, San Diego, CA.

    Peninsular Juniper Woodland and Scrub

    VEGETATION ASSOCIATION:

    PENINSULAR JUNIPER WOODLAND AND SCRUB

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Peninsular juniper woodland and scrub is perhaps the least studied Habitat within the Plan Area. Species composition and ecological factors affecting this community appear to be distinct from typical transmontane juniper-pinyon communities. The mapping of this community is likely based on occurrence of California juniper, an easily identifiable, unique component of the County’s flora. For these reasons, it is likely that the mapping of this community, although it probably accurately represents the distribution of California juniper, it may include distinct communities where species composition and ecological processes more closely match other shrub communities (e.g., coastal sage scrub and chaparral).

    BIOGEOGRAPHY

    California juniper (Juniperus californicus) ranges from San Francisco to Los Cedros Island off the coast of Baja California, Mexico with occurrences mainly on the eastern slopes of the Central Coast, Transverse and Peninsular Ranges but also a few isolated coastal occurrences and extensions into the foothills of the southern Sierra Nevada (Harvey 1951). Peninsular juniper woodland and scrub is most often found on ridges, slopes, and valleys between 1000 and 2450 m (Sawyer and Keeler-Wolf 1995). In San Diego County, California juniper is mainly isolated to abrupt eastern escarpments and desert foothills with an elevational range from 140 to 550 m (Harvey 1951).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Harvey (1951) mentions a few isolated occurrences of California juniper near Aguanga, that are 15 miles west of more dense juniper patches in the Terwilliger Valley.

    Several populations were mapped in the Plan Area totaling 1,106 acres. A small population occurs in the vicinity of Table Mountain and Tule Spring. A single occurrence is mapped west of Pine Cove, south of Indian Mountain. Other single occurrences are northeast of Squaw Mountain, west of the peak of the Lakeview Mountains, and west of Monument Peak. Scattered populations are mapped in the Gavilan Hills, Gavilan Plateau, north of Estelle Mountain and on the south and east sides of Lake Mathews. Boyd (1983) hypothesizes that this desert-like community may have historically been more prevalent in the Gavilan Plateau area.

    VEGETATION CHARACTERISTICS

    Peninsular juniper woodland and scrub most commonly occurs on the eastern slopes of the Peninsular Ranges. However in the case of this discussion, the Habitat occurs in several distinct low-lying areas of Western Riverside County. The Habitat account presented below reveals some information regarding this community as it more commonly occurs but also attempts to focus on describing the Habitat as it occurs within the Plan Area.

    Four-needle pinyon (Pinus quadrifolia), single-leaf pinyon pine (Pinus monophylla) and California juniper (Juniperus californica) are the canopy species which most commonly occur in southern California, forming a scattered canopy from three to 15 m tall (Sawyer and Keller-Wolf 1995; Holland and Keil 1995). Southern California occurrences of juniper woodlands often grade into Joshua tree woodlands or desert scrub communities (Holland and Keil 1995). Shrub cover within this Habitat type is generally intermittent or open and the ground layer is sparse to grassy (Sawyer and Keller-Wolf 1995). Common species on the lower elevational slopes include Mojave yucca (Yucca schidigera) (Y. baccata) (Holland and Keil 1995). Higher elevation species include chamise (Adenostoma fasciculatum) and desert ceanothus (Ceanothus greggi var. vestitus) (Holland and Keil 1995). Some other dominant vegetation species include phlox (Leptodactylon pungens), desert needlegrass (Achnatherum speciosum), matchweed (Gutierrezia sarothrae), beargrass (Nolina parryi), and shrub live oak (Quercus turbinella) (Vasek and Thorne 1977).

    Only one study of Peninsular juniper woodland and scrub in Western Riverside County was reviewed for this account. The vegetative information found in that study is presented below. The community structure presented below is thought to generally fit the areas where this Habitat occurs in the Plan Area, however, vegetative composition may vary widely.

    On the Gavilan Plateau, California juniper occurs in open stands which represent remnants of more extensive historical stands (Boyd 1983). Where juniper woodland is most prevalent on the plateau, topographic relief is moderate. California juniper is the dominant shrub with other scattered shrubs including California buckwheat (Eriogonum fasciculatum), spiny redberry (Rhamnus crocea), snake cholla (Opuntia parryi), chamise (Adenostoma fasciculatum), and Nuttall’s scrub oak (Quercus dumosa) (Boyd 1983). In open areas, often between junipers and other shrubs, grows a diverse herbaceous understory of native annuals and perennials as well as non-native annual grasses (Boyd 1983). Common native wildflowers include goldfields (Lasthenia californica), baby blue eyes (Nemophila menziesii), angel gilia (Gilia angelensis), clarkia (Clarkia purpurea), common owl’s clover (Castilleja exerta), California poppy (Eschscholzia calfornica), cream cups (Platystemon californicus), Johnny jump-up (Viola pedunculata) and common wild onion (Allium praecox). Vegetation composition is somewhat altered in other parts of the Plateau where the juniper woodland community occurs and topographic relief is low. Here few understory shrubs occur, mainly cholla (Opuntia spp.) and redberry (Rhamnus crocea), and the dominant annual is everlasting nest-straw (Stylocline gnaphalioides). Other low, desert-like herbaceous species also may be found in these areas and include everlasting nest-straw (Stylocline filaginea), Palmer’s grappling-hook (Harpagonella palmeri)slender pectocarya (Pectocarya linearis), strigose deerweed (Lotus strigosus), evening- primrose (Camissonia spp.), ground-pink (Linanthus dianthiflorus), and spine flower (Chorizanthe spp.) (Boyd 1983). Under the canopy of junipers, herbaceous growth includes an entirely separate suite of species, including: minor’s lettuce (Claytonia perfoliata), bedstraw (Galium aparine), and nettle (Parietaria floridana). On the margins of junipers occur the following annual species: fiddleneck (Amsinkia intermedia), common forget me not (Cryptantha intermedia), caterpillar phacelia (Phacelia cicutaria), and blue phacelia (Phacelia distans).

    PHYSICAL ENVIRONMENT

    Peninsular juniper woodland and scrub generally occur on bedrock or alluvium-derived soils (Sawyer and Keller-Wolf 1995). California juniper in San Diego County is seldom found on fine-textured lower Bajada and playa soils (Harvey 1951). California juniper appears restricted to sandy and gravelly soils; it is completely absent from clay soils (Harvey 1951).

    Trees in desert woodland Habitats are limited by available moisture and temperature extremes (Holland and Keil 1995). California juniper has been recorded as tolerating temperature extremes from –9oC to 38oC with mean annual temperatures in San Diego County between 10oC and 21oC (Harvey 1951). California juniper occurrences in San Diego County tend to correspond with areas with a steppe climate; i.e., dry climates which are more humid than deserts (Harvey 1951). The average annual precipitation in these areas is between five and 50 cm (Harvey 1951). California junipers have been observed using lateral roots and are thought to also utilize deep penetrating roots allowing the trees to absorb surface and ground water (Harvey 1951). In more easterly, low-elevation occurrences, nearer to the desert, precipitation is the limiting factor in California juniper growth (Harvey 1951). This is evidenced by the observations of California juniper in this region located on mountain tops and lower down on north-facing slopes where available moisture is greater (Harvey 1951).

    ECOSYSTEM PROCESSES

    Ecosystem processes which have been studied in Peninsular juniper woodland and scrub communities are mainly limited to the ecology of California juniper and its responses to fire. The species’ reproductive and growth characteristics help to define its occurrence between chaparral and desert environments.

    The reproductive cycle of California juniper extends over four years, from fertilization to germination (Harvey 1951). Birds, such as jays, as well as rodents are responsible for transporting juniper seeds which can survive the digestive tract of most animals (Harvey 1951). California juniper is not known to stump-sprout following a burn or cut (Harvey 1951). The long period necessary for reproduction, combined with the inability to stump-sprout and an intolerance of shade lessens the opportunity for California juniper invasion of typically highly fire-influenced chaparral communities. Where fires occur in existing juniper stands, it is thought that there is a high likelihood of extirpation if shrubs become dense before new junipers can grow. As a result, intense fires may extirpate California juniper stands or leave isolated patches or individuals to remain (Harvey 1951). An example of this is the Monument Peak area of San Diego County, where California juniper was collected in 1924 but has not been subsequently relocated, presumably, due to a 1944 fire (Harvey 1951). Re-colonization of California juniper may require large tracts of connected Habitat where seeds may be transported via animals.

    THREATS

    There is evidence which indicates that California juniper is susceptible to extirpation following fires (See Physical Environment discussion above). This is likely to be true if placed under intensive human disturbance as well. In the Mojave Desert, stands of juniper woodland are severely restricted due to systematic removal for agricultural development (Sawyer and Keller-Wolf 1995).

    Although true mistletoe (Phoradendron densum) occurs commonly with California juniper, it does not appear to be seriously injurious (Harvey 1951). A list of insect pests to California juniper includes: scale (Aonidia shastae), flattened mealybug (Pseduococcus sp.)dark aphid (Cinara juniperensis), Callimomid wasps (Coleophora cinerella), beetle (Paracotalpa puncticollis)(Mitoura loki), and (Coleophora cinerella). Of these, only the wasps were found to cause significant damage to trees (Harvey 1951).

    LITERATURE CITED

    Boyd, S. D. 1983. A flora of the Gavilan Hills, Western Riverside County, California. Master’s thesis, University of California, Riverside, CA.

    Harvey, H. T. 1951. Distribution of Juniperus californica in San Diego County with interpretations based on ecological factors. M. A. Thesis, San Diego State College, San Diego, CA.

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company. Dubuque, Iowa.

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Thorne, Robert F. 1977. Montane and Subalpine Forests of the Transverse and Peninsular Ranges In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 538-557.

    Vasek, Frank C. and Robert F. Thorne. 1977. Transmontane Coniferous Vegetation In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 797-832.

    Residential/Urban/Exotic

    VEGETATION ASSOCIATION:

    RESIDENTIAL/URBAN/EXOTIC

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Because the County maintains land-use information per parcel, the mapping of urban and residential areas is probably very accurate. Abandoned lots are also most likely well mapped because they usually occur in urban settings rather than isolated patches.

    BIOGEOGRAPHY

    The first urban areas were situated along the coastlines or major rivers (McBride and Reid 1988). Currently, urban areas are most often on what were historically grassland or scrub environments on generally flat slopes (McBride and Reid 1988).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    The largest areas of developed land are in the cities of Temecula, Murrieta/Murrieta Hot Springs, Lake Elsinore (Canyon Lake, Meadowbrook, Alberhill, El Cariso and Lakeland Village), Hemet, East Hemet and Valle Vista, in Moreno Valley, and along the SR-91 corridor from Riverside through Corona and Norco. More medium-sized tracts of developed land are located in the communities of Canyon Lake, Quail Valley, Sun City, Homeland, in Perris along I-215, Mead Valley, Gavilan Hills, Woodcrest, Beaumont, Banning, Cherry Valley, and Calimesa. Small and scattered occurrences include in the southeast portion of the Plan Area along SR-371 in Terwilliger Valley, Sage, Aguanga and Anza, east of Temecula, and in the Sedco Hills.

    VEGETATION CHARACTERISTICS

    There is a variety of ways to classify vegetation within urbanized areas. One method is presented here along with a discussion of weed communities.

    McBride and Reid (1988) divide vegetation within developed areas into four categories: tree grove, street strip, shade tree/lawn, and shrub cover. Tree groves are in parks, green-belts, and cemeteries where a continuous or intermittent canopy is formed and ground coverage varies (McBride and Reid 1988). Street strips and shade trees and lawns generally do not have a continuous cover and vary widely in species and structure (McBride and Reid 1988). These two categories are distinguished by their location. Shrub cover is the most limited vegetation type also occurring as a variety of species and structures (McBride and Reid 1988). A result of these largely ornamental plantings is the establishment of escaped exotics, defined as species originally planted for ornamental or agricultural purposes which have invaded historically natural plant communities. Some commonly escaped exotic species include: acacias (Acacia spp.), pepper-trees (Schinus spp.)pampas grass (Cortaderia spp.), brooms (Cytisus spp.), and English ivy (Hedera helix) (Holland and Keil 1995). In addition to the community types listed above, many areas do not support any vegetation (McBride and Reid 1988).

    Weed communities are also common in urban areas, often occurring on roadsides and abandoned areas. In larger areas these weed populations may represent the early stages of natural succession (Holland and Keil 1995). Some of these areas are known as ruderal communities. A ruderal community occupies waste areas, roadsides often on heavily compacted soils with little available oxygen (Holland and Keil 1995). Typical species include pineapple-weed (Chamomilla suaveloens), common knotweed (Polygonum arenastrum), sow-thistle (Sonchus oleraceus), horseweed (Conyza canadensis), and goosefoot (Chenopodium spp.) (Holland and Keil 1995). Escaped ornamentals also may proliferate in ruderal communities (Holland and Keil 1995). The weed flora in California represents one-sixth of all plant species (Holland and Keil 1995).

    PHYSICAL ENVIRONMENT

    Heat zones form in the most densely populated areas where conditions are 3o to 5oC higher than normal (McBride and Reid 1988). Wind velocities often are lessened in urban areas compared with areas supporting native plant communities (McBride and Reid 1988). A major part of the physical environment in urban areas is human-created such as repeated disturbances from vehicles, trampling, oil, dust, etc. (Holland and Keil 1995).

    ECOSYSTEM PROCESSES

    Most sites maintain the same set of weedy species and herbaceous structure due to human disturbance activities (McBride and Reid 1988). Weeds often become established in disturbed areas due to the large quantities of seeds produced by these plants and lack of natural pests (Holland and Keil 1995). Weedy exotic plants have several reproductive advantages over most natives. The seeds of weedy plants may have effective dispersal mechanisms, may remain dormant in the soil, or the plants themselves may be self-fertile or reproduce apomictically, increasing their ability to compete (Holland and Keil 1995). When human disturbance is suspended or ceased, invasive exotic and some native species often invade (McBride and Reid 1988).

    LITERATURE CITED

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company. Dubuque, Iowa.

    McBride, Joe R. and Chris Reid. 1988. Pasture. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 142-143.

    Riparian Forest/Woodland/Scrub

    VEGETATION ASSOCIATION:

    RIPARIAN FOREST/WOODLAND/SCRUB

    MAPPED SUBASSOCIATIONS

    Riparian Forest, Arundo/Riparian Forest, Southern Willow Scrub, Mulefat Scrub, Riparian Scrub, Tamarisk Scrub, Southern Cottonwood/ Willow Riparian Forest, Southern Sycamore/Alder Riparian Woodland, Montane Riparian Forest, and Montane Riparian Scrub.

    DATA CHARACTERIZATION

    Much of the information provided in this section was obtained from published papers, i.e., Holland and Keil (1995), Faber and Keller (1985), Sawyer and Keeler-Wolf (1995) and Grenfell (1988). Distribution data for the vegetation association and subassociations was obtained primarily from the University of California, Riverside GIS database.

    BIOGEOGRAPHY

    Riparian Habitats generally occur among mid- to large-order streams below 4,000 feet, primarily within the foothills and valleys (Stephenson and Calcarone 1999). The present distributional ranges of riparian Habitat apparently have been influenced more by long-term climatic history than the surrounding upland Habitats. Several million years ago California experienced considerable rain and a warm, humid climate. The flora and fauna that evolved during this period adapted to these conditions. Over the next few millennia, the climate gradually became cooler and drier with summer drought and winter rains. As a result, many plant species that require summer moisture were forced to contract their ranges to riparian zones. Those species not able to persist in the riparian refugia had to adapt to the progressively drier uplands. For these reasons, the dominant riparian tree species in California are not confined to any single floristic region or land form province, or to a single hydrologic or climatic regime (Warner and Hendrix 1984; Faber and Keller 1985).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Riparian vegetation including forest, woodland, and scrub subtypes spatially is distributed in drainages throughout much of Western Riverside County covering approximately 1.1 percent (14,545 acres) of the Plan Area. Southern cottonwood/willow forest makes up the largest proportion of the riparian vegetation in the Plan Area comprising nearly one-half (6,610 acres) of the acreage. Most of the southern cottonwood/willow forest Habitat occurs along the Santa Ana River drainage from Lake Evans to beyond the Prado basin, along the San Gorgonio River north of Banning and along Temecula Creek east of Vail Lake. Other large stands of southern cottonwood/willow forest are scattered evenly throughout the western portion of the Plan Area co-occurring in drainages with other riparian types.

    Large complexes containing several of the riparian forest, woodland and scrub types are located in several portions in the Plan Area. The Santa Ana River basin supports largely southern cottonwood/willow riparian scrub interspersed with smaller patches of riparian scrub, southern willow scrub and mulefat scrub. Drainages flowing toward Highway 91 or Interstate 15, between El Cerrito and eastern Riverside largely support southern willow scrub and riparian scrub with smaller patches of mulefat scrub and southern cottonwood/willow vegetation. Riparian drainages in this area largely are separated by urban and agricultural land uses. Temescal Canyon Wash and its tributaries largely support riparian scrub and mulefat scrub. The stream channels within San Mateo Canyon watershed, within the Cleveland National Forest, generally support riparian forest, southern sycamore/alder riparian woodland and riparian scrub in connected stands. The Temecula area supports a diversity of riparian vegetation types among urban and agricultural land uses along Temecula Creek, Sandia Canyon and portions of Wolf Valley. The riparian system above the Vail Lake area, including Wilson Creek, supports each of the mapped riparian vegetation types.

    Montane riparian forest is distributed at higher elevations in the San Bernardino and San Jacinto Mountains. Most of the montane riparian forest is located along the San Gorgonio River north of Banning, within the Indian Creek watershed on the Soboba Indian Reservation, and along the San Jacinto River west of Hemet Lake. This system also supports significant stands of lower elevation riparian Habitats. Montane riparian scrub is mapped at only one location: the hills south of Highway 60 between Jack Rabbit Trail and Gilman Springs Road.

    Disturbed riparian vegetation types were mapped in only two locations, i.e., arundo/riparian forest was mapped along the urbanized portions of the Santa Ana River from Riverside to Norco and near Corona and tamarisk scrub was mapped only at Vail Lake.

    VEGETATION CHARACTERISTICS

    Riparian communities typically consist of one or more deciduous tree species with an assorted understory of shrubs and herbs (Holland and Keil 1995). The transition between riparian Habitats and adjacent non-riparian Habitats often is abrupt, especially in montane areas where the topography is steep (Grenfell 1988). Vegetation height can vary from one to three meters in scrub Habitats to 30 meters in riparian forest Habitats (Grenfell 1988).

    Riparian Habitats are successional in nature and undergo a predictable sequence of revegetation following flood events. Succession from exposed alluvial soil to mature riparian forest or woodland may take 50 to 75 years or more (Faber and Keller 1985).

    SPECIES COMPOSITION

    Riparian Forest. Riparian forest can include any combination of the following species along perennial stream channel banks: box elder (Acer negundo), big-leaf maple (A. macrophyllum), Valley oak (Quercus lobata), coast live oak (Q. agrifolia), white alder (Alnus rhombifolia), Oregon ash (Fraxinus latifolia), California dogwood (Cornus californica), California bay (Umbellularia californica), sycamore (Platanus racemosa), Fremont’s cottonwood (Populus fremontii), California walnut (Juglans californica), and several species of willow (Salix lasiandraS. lasiolepisS. laevigataS. gooddingii, S. exigua), Mexican elderberry (Sambucus mexicana), wild grape (Vitis girdiana) and poison-oak (Toxicodendron diversilobum). Where the stream channel receives perennial flows in some years but intermittent flows in other years, alder species drop out of the vegetation. Where the stream channel receives only intermittent flow, the willow and cottonwood species become less common and the sycamore, coast live oak and California bay tend to move down into the channel. Along ephemeral stream channels, coast live oak and California walnut can grow within the channel as a continuum or ecotone from uplands on north-facing slopes (Faber and Keller 1985).

    Arundo/Riparian Forest. Arundo/riparian forest is dominated by non-native giant reed (Arundo donax) but also may contain emergent shrubs and trees (Sawyer and Keeler-Wolf 1995) such as arroyo willow (Salix lasiolepis) and mulefat (Baccharis salicifolia).

    Riparian Scrub. This Habitat type has the same potential species composition as riparian forest, but at a younger successional stage, either because of a more recent disturbance or more frequent flooding (Faber and Keller 1985). In addition to the species listed in the description of riparian forest, riparian scrub also may include mulefat.

    Southern Willow Scrub. Southern willow scrub is dominated by willow trees and shrubs (Salix spp.) and also may contain gooseberry (Ribes spp.) and elderberry. When disturbance is high within this Habitat type, the dominant species typically is sandbar willow (Salix exigua). When disturbance is less, the dominant species typically is Goodding’s black willow (Salix gooddingii). Willows are fast-growing and can reproduce vegetatively from root sprouts. Red willow (Salix laevigata) occupies fast-flowing perennial streams at elevations up to 1,200 m and often occurs with yellow willow. Yellow willow (Salix lasiandra) grows along stream channels and in perennially wet places at elevations of 2,500 m. Sandbar willow occurs along sandbars and riverbeds at elevations below 900 m. Arroyo willow occupies Habitat within perennial and intermittent stream channels at elevations up to 750 m. Goodding’s black willow occurs along streambanks and in wet places within drier Habitats at elevations below 450 m (Faber and Keller 1985).

    Mulefat Scrub. Mulefat scrub is dominated by mulefat, but also may include willows (Salix spp.), sedges (Carex spp.) and stinging nettle (Urtica dioica) (Holland 1986; Sawyer and Keeler-Wolf 1995).

    Tamarisk Scrub. Tamarisk scrub is dominated by tamarisk (Tamarix spp.) but also may contain willows (Salix spp.), salt bushes (Atriplex spp.), catclaw acacia (Acacia greggii), and salt grass (Distichlis spicata) (Holland 1986; Sawyer and Keeler-Wolf 1995).

    Southern Cottonwood/Willow Riparian. Southern cottonwood and willow riparian Habitat is dominated by cottonwood (Populus spp.) and willow (Salix spp.) trees and shrubs (Faber and Keller 1985). Understory species may include mugwort (Artemisia douglasiana), stinging nettle and wild cucumber (Marah macrocarpus) (Holland 1986). This riparian Habitat is considered to be an early successional stage as both species are known to germinate almost exclusively on recently deposited or exposed alluvial soils. Like the willow, the cottonwood can reproduce vegetatively from roots. In the absence of disturbance, this Habitat type will transition to include oaks (Quercus spp.) and sycamores or, at higher elevations, will include white alder (Faber and Keller 1985).

    Southern Sycamore/Alder Riparian Woodland. Below 2,000 m sycamore and alder often occur along seasonally-flooded banks; cottonwoods and willows also are often present (Faber and Keller 1985). Poison-oak, mugwort, elderberry and wild raspberry (Rubus spp.) may be present in the understory (Holland 1986). Sycamore and alder are both able to withstand long periods of flooding. The distribution of white alder is restricted to permanent streams and consistent saturation of the root zone by well-aerated, cool water (Faber and Keller 1985).

    Montane Riparian Forest. Montane riparian forest consists of cottonwood (Populus spp.), sycamore, willows (Salix spp.) and alders (Alnus spp.) (Faber and Keller 1985; Grenfell 1988; Sawyer and Keeler-Wolf 1995).

    Montane Riparian Scrub. Montane riparian scrub consists primarily of shrubby species of willows (Salix spp.), dogwood (Cornus spp.) and/or alders (Alnus spp.) (Holland 1986; Faber and Keller 1985; Sawyer and Keeler-Wolf 1995). Jeffrey pine (Pinus jeffreyi) and incense cedar (Calocedrus decurrens) often are found near the edges of these Habitats (Faber and Keller 1985).

    PHYSICAL ENVIRONMENT

    Riparian communities are not restricted to specific climates or soil types, but they are primarily dependent on a permanent supply of water. Variables that affect the community structure and composition include the nature of the water supply (i.e., the amount of water carried by a stream or present within a lake, and the lateral extent and depth of subterranean aquifers), altitudinal gradients, north-south and east-west axes, historical land uses, and the nature and size of the stream banks and flood plains (Bowler 1988; Holland and Keil 1995). The amount of water carried by a stream or present within a lake is determined by the climate (precipitation patterns) and the size of the watershed (Warner and Hendrix 1984).

    Riparian Habitats develop between adjacent mountains and hills and along fault lines. In areas where the geologic forces have upthrust the land into mountain masses, the water flows swiftly and the streams are incised into the underlying rocks. The riparian Habitat is often limited to narrow strips due to the limited soil over the rocks. In areas where the geologic forces have resulted in valleys and gentle slopes, the riparian Habitat is often wider, due to the deposition of eroded soils. Alluvial silt, gravel and sand deposited within the floodplain create a raised terrace which is ideal for bottomland riparian vegetation (Warner and Hendrix 1984).

    Because most of the canopy species are deciduous, there is a seasonal light fluctuation within riparian understories. During winter dormancy, direct sunlight reaches the ground in most parts of the community. It is at this time that some herbaceous and shrub species actively grow and flower. As the canopy leafs-out in the spring, conditions at ground-level change sharply, particularly in broad-leaved deciduous forests. The resulting shade reduces the quantity of light energy beneath the canopy and also reduces daytime temperatures by several degrees. Wind velocity also is decreased by the tree canopy. Moisture evaporation from the soils and transpiration from the vegetation result in increased humidity. These characteristics result in a generally more mesic environment than the surrounding environment (Bowler 1988; Warner and Hendrix 1984; Holland and Keil 1995).

    Within drier riparian zones bordering small, intermittent watercourses, the riparian communities often consist of scattered trees such as sycamore (Platanus racemosa) growing along the stream banks with open areas between the trees. These physical characteristics result in alternating groups of sun-tolerant and shade-tolerant species. The herbaceous species may temporarily die back as the surface water dries up, but the scattered trees and/or shrubs are able to persist due to ground water (Holland and Keil 1995).

    Soils within riparian corridors usually consist of interbedded layers of fine and coarse sediments ranging from clay, silt, sand and gravel to rounded river-rocks and large boulders. The fine-grained particles generally collect in areas where the water movement is slight and the coarse particles generally accumulate where the water flows more quickly. Meandering stream channels within broad floodplains will deposit and redistribute sediments over time, creating a horizontal patchwork and vertical layers. Soils closest to the stream channel are usually relatively young while the seldom-flooded areas within the floodplain are often deep and well-developed (Holland and Keil 1995). Organic materials (e.g., decomposing plant litter) are often present within the soils and nutrient levels are comparatively high. These organic materials are the primary food source for the vegetation within shady headwater situations (Bowler 1988; Holland and Keil 1995).

    Riparian communities can be divided into two general groups: Valley and foothill riparian communities and montane riparian communities. The valley and foothill riparian communities include riparian forest, arundo/riparian forest, southern willow scrub, mulefat scrub, riparian scrub, tamarisk scrub, southern cottonwood/willow riparian, southern sycamore/ alder riparian woodland. Montane riparian communities include montane riparian forest, and montane riparian scrub (Holland and Keil 1995).

    Valley and foothill riparian communities occur at elevations from near sea level to the lower margins of the montane coniferous forest areas within cismontane California. These communities range from the broad valley flood plains to narrow steep canyons. Within valley and foothill riparian communities, the climate is comparatively warm during the winter with precipitation falling primarily as rain and the summers are dry and long (Holland and Keil 1995).

    Montane riparian communities occur in high mountain areas and typically experience lower temperatures than the Valley and foothill riparian communities. Mountainous valleys often serve as drainages for water and cold air. In addition, mountain streams are usually very cold and swiftly flowing. The combined cold air and water temperature limit the types of dominant trees in these communities. Species such as sycamore occur in lower foothill areas but fail to grow at higher elevations. Concurrently, the cold temperatures permit Jeffrey pine and ponderosa pine (P. ponderosa) to colonize riparian areas at lower elevations than they can occupy on exposed slopes. This results in a localized inversion of the usual vegetation zones (Holland and Keil 1995).

    The understory is generally sparse within riparian communities, while the dense broad-leaved deciduous trees can reach heights up to 30 meters (30 m). At higher elevations, the riparian communities are usually less than 15 meters (15 m) in height with a greater concentration of shrubs. At very high elevations (2,000 m or more), the riparian community may only occur in the shrub/scrub stage (Faber and Keller 1985; Grenfell 1988).

    ECOSYSTEM PROCESSES

    Riparian communities are dynamic systems. The stream channels may be swept clean of vegetation during floods as sediments are shifted during erosion flood events. Flood waters may undercut stream banks and, over time, can cut through exposed bedrock, grinding organic debris into small fragments. As a result of channel-cutting and sediment deposition, streams may shift their banks, particularly in areas with gentle topography. Streams in steep areas gradually cut downward, forming canyons or ravines (Holland and Keil 1995).

    In southern California, most streams have very low flow during the summer and in many cases surface flow may dry up (Stephenson and Calcarone 1999). Seasonably variable water flows allow herbaceous annuals and perennials to colonize newly exposed and denuded sites such as an exposed stream channel or a sand or gravel bar. These plants may be swept away during the next winter’s storm events (Holland and Keil 1995).

    Annually variable water flows result in a gradation of environments as well. Some areas of the floodplain may be flooded annually while other areas may be flooded only during years of extremely high waters. Areas that regularly are flooded are often in a condition of perpetual succession. That portion of the riparian community nearest to the stream channel may not advance beyond the pioneer stage, while areas farther from the stream channel may advance through several stages of succession before being swept clean by periodic floods (Holland and Keil 1995).

    Many riparian woody and herbaceous species are adapted to periodic flooding. Some have deep root systems that anchor them against the flood waters and some have flexible stems that bend with the flood waters. Many have rhizomes that are protected by layers of sediments and others have no adaptations but are able to persist if they become established at sites that protect them from the full force of the flood waters (e.g., among large rocks) (Holland and Keil 1995). Cottonwood and willow species germinate almost exclusively on recently deposited or exposed alluvial soils (Faber and Keller 1985).

    COMMUNITY RELATIONSHIPS

    Riparian communities are very important wildlife Habitats. The multiple strata (e.g., canopy, shrubs, herbaceous species) provide a diversity of feeding and nesting sites for mammals and birds. Fish and other aquatic species benefit from important shading and other attributes. (The reader should refer to the species accounts for specific information about sensitive wildlife and plant species and riparian Habitats.) Riparian areas are of particular importance because the moisture of the stream channels is important as a water source in the dry California landscape and are productive during the summer months at times when other plant communities are dormant (Warner and Hendrix 1984; Grenfell 1988; Holland and Keil 1995).

    Linear riparian Habitats, by providing shade, cover, water and food, can function as important corridors for faunal species’ migration and dispersal (Warner and Hendrix 1984).

    THREATS

    Riparian Habitats are threatened by cattle grazing, conversion to other land uses and flood control activities. These activities degrade the water quality, instream ecology, reduce the floodplain and/or encourage the colonization of non-native floral species (Bowler 1988).

    LITERATURE CITED

    Bowler, P.A. 1988. Riparian Woodland: An Endangered Habitat in Southern California.

    Faber, P.M. and E. Keller. 1985. The Ecology of Riparian Habitats of the Southern California Coastal Region: A Community Profile. USDI Fish and Wildlife Service Biological Report. 152 pp.

    Grenfell, W.E. Jr. 1988. Montane Riparian in A guide to Wildlife Habitats of California. California Department of Forestry and Fire Protection. 166 pp.

    Holland R.F. 1986. Preliminary Descriptions of the Terrestrial Natural Communities of California. Sate of California Department of Fish and Game, Nongame-Heritage Program, Sacramento, California.

    Holland, V.L. and D.J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company, Dubuque, Iowa. 516 pp.

    Sawyer, J.O. and T. Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, California. 471 pp.

    Stephenson, J.R. and G.M. Calcarone. 1999. Southern California Mountains and Foothills Assessment: Habitat and Species Conservation Issues. General Technical Report GTR-PSW-172. Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture, Albany, California. 402 pp.

    Warner, R.E. and K.M. Hendrix. 1984. California Riparian Systems: Ecology, Conservation and Productive Management. University of California Press, Berkeley, California. 1035 pp.

    San Jacinto Valley Vernal Pools (Sawyer & Keeler-wolf)

    VEGETATION ASSOCIATION:

    SAN JACINTO VALLEY VERNAL POOLS (Sawyer & Keeler-Wolf)

    MAPPED SUBASSOCIATIONS

    San Jacinto Valley Vernal Pools

    OTHER CLASSIFICATIONS:

    Southern/Perris Basin Vernal Pools (CDFG)

    Interior Cismontane Vernal Pools (Cheatham & Haller)

    Vernal Pool Ephemeral (Thorne)

    Palustrine, non-persistent, seasonally flooded wetland (Cowardin)

    DATA CHARACTERIZATION

    BIOGEOGRAPHY

    The San Jacinto Valley is situated between two arms of the Peninsular Ranges: the Santa Ana Mountains to the west and the San Jacinto/Santa Rosa Mountains to the east. Seismic activity along faults at the valley edges has resulted in the valley floor sinking relative to the adjacent mountains. Alluvial materials have been deposited on the valley floor, and these have developed into saline-alkali clay soils. The vernal pools in this valley are a remnant of a once extensive seasonal wetland landscape that occurred in the valley bottomlands, the majority of which are now in agricultural production. The remaining pools occur in a changing mosaic with a series of alkali-flat plant communities, including alkali (vernal) playas and alkali grassland.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    The San Jacinto Valley/Perris Basin.

    VEGETATION CHARACTERISTICS

    The flora of vernal pools includes species that are restricted to vernal pools, that occur in vernal pools and other types of seasonal wetlands, and/or that are typical of upland Habitats but establish around pool margins or colonize pool basins during the summer or fall (Jones & Stokes 1990). Vegetation cover varies in response to the amount and distribution of rainfall and air temperatures and, consequently, the presence or absence of an individual species and its relative abundance varies from year to year. During drought years, a vernal pool flora may include abundance of upland species and cover not characteristic of vernal pools under normal climatic conditions.

    SPECIES COMPOSITION

    Vernal pools are well-known for their high level of endemism (Stone 1989) and abundance of rare, threatened, or endangered species (Sawyer and Keeler-Wolf 1995). Many vernal pools are characterized by concentric rings of plants that flower sequentially as the pools dry. Vernal pools are dominated by native annual plants, with low to moderate levels of perennial herbaceous cover. Common vernal pool plant species in the pools in the San Jacinto Valley include woolly marbles (Psilocarphus brevissimus), toad rush (Juncus bufonius), spike rush (Eleocharis sp.), wire-stem popcorn flower (Plagiobothrys leptocladus), Mexican speedwell (Veronica peregrina ssp. xalapensis), annual hairgrass (Deschampsia danthonioides), alkali pepper-grass (Lepidium dictyotum), and water pygmy weed (Crassula aquatica). Sensitive or listed plant species that occur in these vernal pools/alkali playas include: California Orcutt grass (Orcuttia californica), little mousetail (Myosurus minimus ssp. apus), thread-leaved brodiaea (Brodiaea filifolia), Coulter’s goldfields (Lasthenia glabrata ssp. coulteri), little mousetail (Myosurus minimus ssp. apus), spreading navarretia (Navarretia fossalis), Orcutt’s brodiaea (Brodiaea orcuttii), thread-leaved brodiaea (Brodiaea filifolia), Parish brittlescale (Atriplex parishii), San Jacinto Valley crownscale (Atriplex coronatavar. notatior), and smooth tarplant (Hemizonia pungens ssp. laevis).

    PHYSICAL ENVIRONMENT

    Vernal pools are ephemeral wetlands that form in shallow depressions underlain by a substrate near the surface that restricts the downward percolation of water. Depressions in the landscape fill with rainwater and runoff from adjacent areas during the winter and may remain inundated until spring or early summer, sometimes drying more than once during the wet season. Smaller pools can fill, and dry, and larger pools can hold water longer and may in the deeper portions, support species that are more representative of freshwater marshes.

    The San Jacinto Valley vernal pools occupy the prehistoric bed of San Jacinto Lake (and its remnant, Mystic Lake) and intergrade with alkali playa and alkali grassland Habitats. Vernal pools and alkali playa occupy approximately 42 acres and 6,081 acres, respectively within the Plan Area (PSBS 1995). Vernal pool basins are few, shallow, and relatively small in size, occurring in both the San Jacinto and Salt Creek drainages within the Perris Basin. Vernal pools, alkali playa, and alkali grasslands also are found southwest of Hemet and scattered along the margins of the San Jacinto River, extending from Mystic Lake to Perris, including portions of the San Jacinto Wildlife Area owned and managed by the California Department of Fish and Game. These pools occupy plains on soils mapped as Domino-Travers-Willows association. This soil association is alkaline and has a subsoil layer of impervious caliche (White 1994). The Willows series is also found in the Sacramento Valley and underlies Northern Claypan pools there, underscoring their similarity to the Riverside pools (CDFG 1998). Hydrology of the Perris Basin pools is variable; some are filled when the San Jacinto River overflows its banks.

    Alkali grasslands, alkali playas, and vernal pools share much of their floristic composition and are often difficult to differentiate (White 1994). Vernal pools are considered to be those areas which pond reliably year-to-year as evidenced by the presence of standing water durations adequate to support a vegetation characteristic of vernal pool Habitat in most years. In contrast, alkaline vernal playas are physical situations (large, shallow, lakes, some of which are the result of man-made topographic features which impede the natural flow of water) which are subject to seasonal flooding and ponding on a less reliable basis, but which possess characteristic soils and vegetation developed in response to periodic flooding and low soil permeabilities. In the wettest years, large areas of vernal playa Habitat may support plant taxa normally observed primarily in vernal pools. In drier years, these playas may support alkali grassland Habitat (RECON 1995).

    The isolated pool at Skunk Hollow near Temecula is relatively large (up to 10 acres), resembling vernal lake. It may remain wet through much of the year and only contain vernal pool species on the drying margins. The Skunk Hollow pool is located on Willows silty clay and is likely to have some similarities with pools in the old Salt Creek drainage west of Hemet. This 33-acre pool provides Habitat for the Riverside fairy shrimp (Streptocephalus woottoni) and other sensitive species (e.g., Ambrosia pumila in surrounding watershed).

    ECOSYSTEM PROCESSES

    Vernal pools are seasonally flooded landscape depressions that support a distinctive biota adapted to periodic or continuous inundation during the wet season and the absence of either ponded water or wet soils during the dry season. Vernal pools that meet this definition are circumscribed by a framework of topographic, edaphic, hydrological, and biotic parameters. These parameters also distinguish vernal pools from other types of seasonal and perennial wetlands (Jones & Stokes 1990).

    Vernal pools have a unique hydrology primarily because of soil/substrate properties and the Mediterranean climate of lowland California. Pool basins are periodically or continuously flooded during late fall, winter, and spring and are dry and desiccated through summer and early fall, undergoing a sequence of hydrologic phases reflected by the phenology and composition of the dominant biota that characterize each phase. As described in Zedler (1987), vernal pool development during a year can be broken down into four phases: wetting, aquatic, drying, and drought. Fall rains initiate the “wetting” stage and stimulate the germination of dormant seeds and the resprouting of perennials that eventually form an herbaceous turf before and during the transition to the “aquatic” phase. This phase begins after soils reach field capacity (i.e., the soil profile is saturated) and water is either exposed where pool depressions intercept a seasonally perched water table or accumulates from surface runoff. The subsequent “drying” phase begins as water levels recede during spring and is characterized by partially flooded pool basins and/or wet or saturated soil. The final “drought” phase occurs when the pool basin and soils are dry and nearly all of the biota that previously was actively growing dies or becomes dormant. Vernal pools derive water from direct precipitation, overland runoff, and groundwater in seasonal perched water tables. Vernal pool hydrology for pools in the northwestern region seems to differ from other pools because they are filled when the San Jacinto River overflows its banks. Historically, the San Jacinto River drained into Mystic Lake, an expansive playa in the northern region of the valley, and vernal pool hydrology likely was linked to the lake itself (White 1994).

    THREATS

    The most heavily threatened and most severely reduced vernal pool types in Riverside County are the Perris Basin pools (CDFG 1998). San Jacinto Valley vernal pools are threatened by a variety of factors, including: Habitat destruction and fragmentation from agricultural and urban development; pipeline construction; alterations in wetland hydrology by draining or channelization; clay mining; off-road vehicle activity; cattle and sheep grazing; weed abatement; fire suppression practices; and competition from alien plant species.

    The San Jacinto Wildlife Area managed by the CDFG protects a small portion of the San Jacinto Valley vernal pools but the majority of those remaining occur outside its boundaries. Additional protection and study are necessary for these pools and unclassified pools such as Skunk Hollow. Skunk Hollow is conserved as part of a mitigation bank for general wetland impacts which occur within the Plan Area (Western Riverside County). The bank formally is permitted by the Army Corps of Engineers, U.S. Fish and Wildlife Service, and Department of Fish and Game.

    LITERATURE CITED

    Eriksen, C.H. and D. Belk. 1999. Fairy Shrimps of California’s Puddles, Pools, and Playas. Mad River Press, Inc. 196 pp.

    Jones & Stokes. 1990. Sacramento County Vernal Pools: their Distribution, Classification, Ecology, and Management. Unpublished report prepared for the County of Sacramento, Planning and Community Development Department. June 15.

    Keeler-Wolf, T. et al. 1998. California Vernal Pool Assessment Preliminary Report, Western Riverside County Region. California Department of Fish and Game. May

    RECON. 1995. The Distribution, Status, and Conservation of Vernal Pool and Alkali Playa Wetlands of the Upper Salt Creek Drainage, Hemet, California. Unpublished report prepared for the City of Hemet and the Riverside Habitat Conservation Agency. June.

    Sawyer, J.O. and T. Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society

    Stone, R.D. 1990. California’s Endemic Vernal Pool Plants: Some Factors Influencing their Rarity and Endangerment. Pages 89-107 in: Vernal Pool Plants: Their Habitat and Biology, Ikeda & Schlising, eds. Studies from the Herbarium, California State University, Chico. Number 8. June.

    White, Scott D. 1994. Vernal Pools in the San Jacinto Valley. Fremontia 22(3): 17-19.

    Zedler, P.H. 1987. The Ecology of Southern California Vernal Pools: A Community Profile. U.S. Fish and Wildlife Service Biological Report 85(7.11). 136 pp.

    Sonoran Desert Scrub

    VEGETATION ASSOCIATION:

    SONORAN DESERT SCRUB

    BIOGEOGRAPHY

    Sonoran desert scrub is distributed widely within the southwestern desert biomes from the Colorado Desert region to the Little San Bernardino Mountains (Holland 1986). This vegetation dominates arid landscapes at elevations below 760 or 910 m (Holland 1986). Sonoran desert scrub vegetation ranges from the northeastern edge of the Plan Area south and east into southern Arizona, Baja California, and Sonora, Mexico (Holland 1986). In southeastern San Bernardino County and eastern Riverside County, Sonoran desert scrub tends to intergrade with Mojave creosote bush scrub.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Sonoran desert scrub Habitat is sparsely scattered over the southeastern portion of Western Riverside County, occupying approximately 0.02 percent (282 acres) of the Plan Area. The mapped locations of Sonoran desert scrub were limited specifically to the southeast corner of the Plan Area, located east of the City of Aguanga and Lake Riverside.

    VEGETATION CHARACTERISTICS

    The Sonoran desert scrub Habitat type typically is associated with a predominant floristic character comprised of microphyllous shrubs such as burro weed (Ambrosia dumosa), brittlebush (Encelia farinosa), crucifixion-thorn (Canotia holacantha) and Creosote scrub (Larrea tridentata) (Holland 1986; California Gap Analysis Report 1998). Stem and leaf succulents such as ocotillo (Fouquieria splendens) also are associated with Sonoran desert scrub (Holland 1986). Shrubs within this Habitat type prefer to grow in well-drained secondary soils of slopes, and fan and valleys where the soils are rich with biotic components. Sonoran desert scrub is very similar in appearance to the Mojave creosote bush scrub and the two often are mistaken for one another.

    PHYSICAL ENVIRONMENT

    Sonoran desert scrub encompasses great species and life form diversity, often displaying ephemeral, colorful annual flowers, microphyllous shrubs, and succulents following particularly wet winters (California Gap Analysis 1998). Flowers tend to bloom in late February and early March if seasonal rainfall is sufficient, which is earlier than in Mojave creosote bush scrub, whose flowers typically bloom from late March to early April (Holland 1986). Growth typically occurs from winter to early spring and is dependent on rainfall. If the seasonal rainfall is sparse, shrubs from this Habitat type may be dormant for long periods of time until proper resources are sufficient (Holland 1986). Under adequate growing conditions, Sonoran desert scrub vegetation can range in height from 0.5 to 3 m (Holland 1986).

    In southern California, Sonoran desert scrub grows well on well-drained secondary soils of slopes, fans, and valleys rather than on upland sites with thin residual soils or sites with high soil salinity (Holland 1986). Plants within this community also inhabit areas with high alkalinity and rocky, unweathered substrate associated with steep, rocky environments (California Gap Analysis Report 1998).

    THREATS

    Threats to Sonoran desert scrub vegetation includes runoff from agricultural irrigation and upstream residential and commercial development projects; urban edge effects; pollution from residential and industrial runoff; and off-highway vehicle use.

    LITERATURE CITED

    Gray, Violet, David Stoms, and Frank Davis. 1998. California Gap Analysis Report - The Sonoran Desert Region. [http://www.biogeog.ucsb.edu/projects/gap/ report/son_rep.html] Accessed 26 September 2000.

    Holland, R.F. 1986. Preliminary Descriptions of the Terrestrial Natural Communities of California. State of California Department of Fish and Game, Nongame-Heritage Program, Sacramento, California.

    Southern California White Fir Forest

    VEGETATION ASSOCIATION:

    SOUTHERN CALIFORNIA WHITE FIR FOREST

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Because of the limited area within the Plan Area that contains suitable climatic conditions for white fir (Abies concolor) stands and the easily discernable differences (i.e., on aerial photographs) between pine forests and white fir-dominated forest, it is likely that the mapping for this Habitat is accurate.

    BIOGEOGRAPHY

    Found in forested mountains throughout California, white fir generally occurs at elevations between red fir (Abies magnifica) and yellow pine (Pinus ponderosa) forests on moister slopes between 1,675 to 2,300 m (Thorne 1976). White fir, including both varieties, occurs from the Pacific coast to central Colorado and from central Oregon to northeastern Mexico (Laacke 1990).

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Southern California white fir forest was mapped on the western slopes of Tahquitz Peak east of Idyllwild, and west of Pine Cove near Pigeon Spring. Other occurrences are south of Indian Mountain and a large population in the Black Mountains and on Marion Ridge. A total of 6,899 acres of southern California white fir occurs in the Plan Area.

    VEGETATION CHARACTERISTICS

    It should be noted that two varieties of white fir have been defined: Rocky Mountain white fir (Abies concolor var. concolor) and Sierran white fir (A. c. var. lowiana); the Rocky Mountain white fir is the type occurring in the Transverse and Peninsular ranges of southern California (Sawyer and Keeler-Wolf 1995).

    Over its California range, the Habitat may occur in monotypic, even-aged stands or with other conifers (Shimamoto 1995). Thorne (1977) characterized this Habitat as white fir-sugar pine forest, indicating the importance of sugar pine (Pinus lambertiana), and includes incense-cedar (Calocedrus decurrens) as a major component. White fir also may occur with lodgepole pine forest (Pinus contorta ssp. murrayana) in more xeric conditions (Barbour and Minnich 2000). Shrub cover varies considerably but typically includes mountain pink currant (Ribes nevadense), Sierra gooseberry (R. roezlii), thimbleberry (Rubus parviflorus), Sitka willow (Salix sitchensis), and blue elderberry (Sambucus caerulea) (Sawyer and Keeler-Wolf 1995; Thorne 1977). Herbaceous cover usually is sparse but may approach 100 percent in moist swales and drainage bottoms (Shimamoto 1988). Understory growth is limited by shade and fallen woody material (Shimamoto 1988).

    Tree canopy cover has been measured up to 75% and individual trees, in southern California, reach an average of 30 m (Shimamoto 1988; Barbour and Minnich 2000). In the southern ranges of California, white fir often intergrades with mixed conifer at low elevations and lodgepole pine forest at higher elevations (Shimamoto 1988).

    In Riverside County, Southern California white fir is dominated by white fir but also supports sugar pineincense-cedarand a variety of understory plantsOccurring in fairly dense stands, white fir in the San Jacinto mountains also often have “witches’ brooms” caused by dwarf mistletoe (Arceuthobium campylopodum) (Vogl 1976).

    PHYSICAL ENVIRONMENT

    Over its California range, white fir stands generally occur between 1400 and 2700 m, usually on slopes of any aspect or along raised stream benches or terraces (Sawyer and Keeler-Wolf 1995). Although white fir may be found on a variety of slope types, it is most well suited for gentle slopes and level ground (Laacke 1990). In the Peninsular ranges, white fir occurs between 1800 m and 2135 m (Shimamoto 1988). Mean temperatures are between 0oC and 10oC and precipitation is between 76 cm and 178 cm (Shimamoto 1988). White fir trees do not appear to be limited by soil type, growing in a variety of soil types and parent materials (Laacke 1990). Growth of the trees appears to be best suited to moderately deep and well-drained, coarse textured sandy loam to clay loam soils (Laacke 1990, Shimamoto 1988).

    White fir occurring in Riverside County typically is found on more mesic sites between 1830 and 2590 m (Vogl 1976; Thorne 1976).

    ECOSYSTEM PROCESSES

    Historically, white fir did not penetrate low elevations due to fire (Laacke 1990). However, recent fire suppression activity has resulted in greater numbers of white fir at lower elevations (Laacke 1990). This likely is due to the white fir’s ability to tolerate shade and its fire resistance when compared with other high-elevation trees (Laacke 1990).

    Separate from the discussion of white fir invading lower elevation areas where fire has been suppressed, white fir communities also undergo a long period of succession following fires. In white fir-dominated areas in the northern Sierra Nevada, post-fire conditions led to a shift in dominance from montane chaparral shrubs to white fir trees (Conrad and Radosevich 1982). Following a burn, shrub seedlings including manzanita (Arctostaphylos spp.), California-lilac (Ceanothus spp.), willows (Salix spp.), bitter cherry (Prunus emarginata), huckleberry oak (Quercus vaccinifolia), currant (Ribes spp.), gooseberry (Ribes spp.), and chinquapin (Chrysolepis spp.), will establish up to a 75 percent coverage (Shimamoto 1988). These shrubs along with white fir saplings may persist for 30-50 years before white fir becomes fully established (Shimamoto 1988). This reveals a pattern of succession where white fir-dominated sites with a primarily herbaceous and sub-shrub understory is the climax community and white fir in association with montane chaparral species is a successional community (Conrad and Radosevich 1982).

    Other ecosystem processes include the establishment of white fir in montane meadows where lodgepole pine forest is established. It is thought that white fir uses the relatively dry ground created by the pine roots to tolerate the high soil moisture, although white fir’s ability to maintain itself in these conditions is questionable because spring and fall frosts can kill or damage young or old white fir (Laacke 1990). The presence of white fir in some areas may be due to selective logging of ponderosa and sugar pine (Gordon 1980).

    THREATS

    Several parasites affect white fir communities. White fir is susceptible to true mistletoes (Phoradendron spp.) which are less destructive than the dwarf mistletoes (Arceuthobium campylopodum) but still may cause the death of some trees (Stephenson and Calcarone 1999). One study found that one-third of white fir stands is infected by mistletoes (Laacke 1990). Annosus root rot (Heterobasidion annosum) damages white fir trees by causing moisture stress and loss of vigor (Laacke 1990). Other pests include yellow cap fungus (Phliota limonella), Indian paint fungus (Echindontium tinctorium), and white pocket rot (Phellinus pini) (Laacke 1990).

    Many insects attack white fir including the fir engraver beetle (Scolytus ventralis), which is most common in white fir, the roundheaded fir borer (Tetropium abietes), seed maggots (Earomyia spp.), and fir cone looper (Eupithecia spermaphaga) (Laacke 1990); some of these are constant attackers and others occur as episodic epidemics (Laacke 1990). Other similar pests and parasites infect sugar pine and incense-cedar within white fir communities (Thorne 1977). Relative to species like ponderosa pine, white fir is less susceptible to high ozone levels (Laacke 1990).

    White fir invasions of other Habitats may be only controlled by prescribed burns, the method by which is not described here (Laacke 1990).

    LITERATURE CITED

    Conrad, S. G. and S. R. Radosevich. 1982. Post-fire succession in white fir vegetation of the northern Sierra Nevada. Madrono, Vol. 29, No. 1:42-56.

    Gordon, Donald T. 1980. White Fir. In Forest Cover Types of the United States and Canada. ed. F. H. Eyre. Society of American Foresters, Washington, D.C. pp. 92-93.

    Laacke, Robert J. 1990. Abies concolor - white fir. Pages 36-46 in R.M. Burns and B. H. Honkala, technical coordinators. Silvics of North America, Volume 1. Conifers. Agriculture Handbook 654. USDA, Forest Service, Washington, D.C.

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Shimamoto, Karen. 1988. White Fir. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 48-49.

    Stephenson, John R. and Gena M. Calcarone. 1999. Southern California mountains and foothills assessment: Habitat and species Conservation issues. General Technical Report GTR-PSW-172. Albany, CA: Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture; 402 p.

    Thorne, Robert F. 1976. The Vascular Plant Communities of California. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 1-31.

    Thorne, Robert F. 1977. Montane and Subalpine Forests of the Transverse and Peninsular Ranges In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 538-557.

    Vasek, Frank C. 1985. Southern California White Fir. Madrono, Vol. 32, No. 2:65-77.

    Vogl, Richard J. 1976. An Introduction to the Plant Communities of the Santa Ana and San Jacinto Mountains. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 77-98.

    Southern Interior Basalt Vernal Pools

    VEGETATION ASSOCIATION:

    SOUTHERN INTERIOR BASALT VERNAL POOLS

    Mapped Subassociations:

    Southern Interior Basalt Vernal Pools

    OTHER CLASSIFICATIONS:

    Southern Interior Basalt Flow Vernal Pool (Holland)

    Santa Rosa Plateau Vernal Pools (Sawyer & Keeler-Wolf)

    Interior Cismontane Vernal Pools (Cheatham & Haller)

    Vernal Pool Ephemeral (Thorne)

    Palustrine, non-persistent, seasonally flooded wetland (Cowardin)

    DATA CHARACTERIZATION

    The data set for southern interior basalt vernal pools within the Plan Area is fairly complete.

    BIOGEOGRAPHY

    The Santa Rosa Plateau vernal pools represent the only known locality of basalt flow vernal pools in southern California. Other basalt vernal pools can be found in northeastern California as part of a huge system found on the Modoc Plateau. The Modoc pools are classified as northern basalt flow vernal pools (Smith and Keeler-Wolf 1995) and also are underlain by basalt bedrock and basalt-derived clay soils.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Southern interior basalt vernal pools are found only on the Santa Rosa Plateau in Western Riverside County. Approximately 31 acres of this Habitat type have been mapped within the Plan Area (PSBS 1995).

    VEGETATION CHARACTERISTICS

    The flora of vernal pools includes species that are restricted to vernal pools and other types of seasonal wetlands, and/or that are typical of upland Habitats but establish around pool margins or colonize pool basins during the summer or fall (Jones & Stokes 1990). Vegetation cover varies in response to the amount and distribution of rainfall and air temperatures and, consequently, the presence or absence of an individual species and its relative abundance varies from year to year. During drought years, a vernal pool flora may support an abundance of upland species and cover that is not characteristic of vernal pools under normal climatic conditions.

    SPECIES COMPOSITION

    Vernal pools are well-known for their high level of endemism (Stone 1989) and abundance of rare, threatened, or endangered species (Sawyer and Keeler-Wolf 1995). Many vernal pools are characterized by concentric rings of plants that flower sequentially as the pools dry. Vernal pools are dominated by native annual plants, with low to moderate levels of perennial herbaceous cover. Common vernal pool plant species in the pools on the Santa Rosa Plateau include annual hairgrass (Deschampsia danthonioides), waterfern (Marsilea vestita), pillwort (Pilularia americana), woolly marbles (Psilocarphus brevissimus, P. tenellus), water starwort (Callitriche spp.), downingia (Downingia cuspidata, D. bella), Blennosperma nanum, water crow-foot (Ranunuculus aquatilis), spike-rush (Eleocharis acicularis), and toad rush (Juncus bufonius) (Rosario and Lathrop 1984). In addition, the following sensitive or listed plant species are found in one or more of these pools: California Orcutt grass (Orcuttia californica), Coulter’s goldfields (Lasthenia glabrata ssp. coulteri), little mousetail (Myosurus minimus ssp. apus), spreading navarretia (Navarretia fossalis), low navarretia (N. prostrata), Orcutt’s brodiaea (Brodiaea orcuttii), thread-leaved brodiaea (Brodiaea filifolia), Parish brittlescale (Atriplex parishii), Parish meadowfoam (Limnanthes gracilis ssp. parishii), San Diego button-celery (Eryngium aristulatum var. parishii), and Wright’s trichocoronis (Trichocoronis wrightii var. wrightii) (Sawyer and Keeler-Wolf 1995). The Santa Rosa Plateau fairy shrimp (Linderiella santarosae) occurs only here as well as the southernmost record for the vernal pool fairy shrimp (Branchinecta lynchi) (Erikson and Belk 1999).

    PHYSICAL ENVIRONMENT

    Vernal pools are ephemeral wetlands that form in shallow depressions underlain by a substrate near the surface that restricts the downward percolation of water. Depressions in the landscape fill with rainwater and runoff from adjacent areas during the winter and may remain inundated until spring or early summer, sometimes drying more than once during the wet season. Smaller pools can fill, dry, and larger pools can hold water longer and may, in the deeper portions, support species which are more representative of those found in freshwater marshes.

    The Santa Rosa Plateau lies at the southern end of the Santa Ana Mountains, topographically positioned at an elevation of approximately 730 m. The plateau is comprised of several mesas upon which 13 vernal pools exist in a landscape of native grassland (Holing 1988). The pools that are found on these mesas are unique in southern California as, instead of a clay or hardpan, the subsurface layer beneath is olivine basalt base rock from past volcanic flows. The basalt layer ranges in depth from 30-50 centimeters (cm), except under some vernal pools where it may be as shallow as 10 cm. The basalt breaks down to create an extremely expansive clay soil that allows very little water loss through seepage. Rainwater fills the low-lying basin areas creating pools which occur as a sequence of playa-like temporary lakes. Four vernal pools are found on Mesa de Colorado, one on Mesa de la Punta, and eight on Mesa de Burro. Basin areas range from 0.25 hectare (ha) for some of the smaller pools to approximately 10 ha for the largest one on Mesa de Colorado. The size of the pools is dependent upon the interaction of the amount of rainfall and air temperatures of any given year.

    ECOSYSTEM PROCESSES

    Vernal pools are seasonally flooded landscape depressions that support a distinctive biota adapted to periodic or continuous inundation during the wet season and the absence of either ponded water or wet soils during the dry season. Vernal pools that meet this definition are circumscribed by a framework of topographic, edaphic, hydrological, and biotic parameters. These parameters also distinguish vernal pools from other types of seasonal and perennial wetlands (Jones & Stokes 1990).

    Vernal pools have a unique hydrology primarily because of soil/substrate properties and the Mediterranean climate of lowland California. In the case of the Santa Rosa Plateau vernal pools, pool substrate is either basalt or the highly impermeable clay soils which develop from this parent material. Pool basins are periodically or continuously flooded during late fall, winter, and spring and are dry and desiccated through summer and early fall, undergoing a sequence of hydrologic phases reflected by the phenology and composition of the dominant biota that characterize each phase. As described in Zedler (1987), vernal pool development during a year can be broken down into four phases: wetting, aquatic, drying, and drought. Fall rains initiate the “wetting” stage and stimulate the germination of dormant seeds and the resprouting of perennials that eventually form an herbaceous turf before and during the transition to the “aquatic” phase. This phase begins after soils reach field capacity (i.e., the soil profile is saturated) and water either is exposed where pool depressions intercept a seasonally perched water table or accumulates from surface runoff. The subsequent “drying” phase begins as water levels recede during spring and is characterized by partially flooded pool basins and/or wet or saturated soil. The final “drought” phase occurs when the pool basin and soils are dry and nearly all of the biota that previously was actively growing dies or becomes dormant. Vernal pools derive water from direct precipitation, overland runoff, and groundwater in seasonal perched water tables. Although direct precipitation appears to be the primary water source for the pools on the Santa Rosa Plateau, the contribution from other sources may be important or crucial to maintaining their natural hydrology.

    The alteration of very wet and very dry conditions creates an unusual ecological situation that supports a unique biota. To survive in a vernal pool, species must be able to either tolerate a wide range of conditions, or to grow and reproduce in the short time that there is a favorable environment.

    THREATS

    The Santa Rosa Plateau pools are largely protected in the Santa Rosa Plateau Ecological Reserve managed by The Nature Conservancy (TNC). Over 7,300 acres of the Santa Rosa Plateau have been made available as a wilderness ecological preserve through a cooperative management project of the Nature Conservancy, the Riverside County Regional Park and Open Space District, the California Department of Fish and Game, the U.S. Fish and Wildlife Service, and the Metropolitan Water District of Southern California.

    LITERATURE CITED

    Eriksen, C.H. and D. Belk. 1999. Fairy Shrimps of California’s Puddles, Pools, and Playas. Mad River Press, Inc. 196 pp.

    Holing, D. 1988. California Wildlands: A Guide to the Nature Conservancy Preserves. Chapter 23--The Santa Rosa Plateau. Chronicle Books, San Francisco.

    Jones & Stokes. 1990. Sacramento County Vernal Pools: their Distribution, Classification, Ecology, and Management. Unpublished report prepared for the County of Sacramento, Planning and Community Development Department. June 15.

    Keeler-Wolf, T. et al. 1998. California Vernal Pool Assessment Preliminary Report, Western Riverside County Region. California Department of Fish and Game. May.

    Lathrop, E.W. and R.F. Thorne. 1983. A Flora of the Vernal Pools on the Santa Rosa Plateau, Riverside County, California. Aliso 10(3):449-469.

    Lathrop, E.W. and R.F. Thorne. 1976. Vernal Pools of the Santa Rosa Plateau. Fremontia 4(3): 9-11.

    Pacific Southwest Biological Servics (PSBS). 1995. Draft MSHCP Strategy Report Vegetation Mapping. Prepared for the Western Riverside County Habitat Consortium. February 20.

    Rosario, J.A. and E.W. Lathrop. 1984. Distributional Ecology of Vegetation in the Vernal Pools of the Santa Rosa Plateau. Pages 210-217 in: Vernal Pools and Intermittent Streams, S. Jain and P. Moyle, eds. Institute of Ecology Publication Number 28.

    Rosario, J. A. 1979. Distributional Ecology of Vegetation in the Vernal Pools of the Santa Rosa Plateau, California. Dissertation in partial fulfillment of the requirements for Doctor of Philosophy in the Field of Biology, Loma Linda University. 89 pp.

    Sawyer, J.O. and T. Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society.

    Stone, R.D. 1990. California’s Endemic Vernal Pool Plants: Some Factors Influencing their Rarity and Endangerment. Pages 89-107 in: Vernal Pool Plants: Their Habitat and Biology, Ikeda & Schlising, eds. Studies from the Herbarium, California State University, Chico. Number 8. June.

    Zedler, P.H. 1987. The Ecology of Southern California Vernal Pools: A Community Profile. U.S. Fish and Wildlife Service Biological Report 85(7.11). 136 pp.

    Subalpine Coniferous Forest

    VEGETATION ASSOCIATION:

    SUBALPINE CONIFEROUS FOREST

    MAPPED SUBASSOCIATIONS

    None

    DATA CHARACTERIZATION

    Because of the very limited range where climatic conditions are appropriate, the single occurrence of this Habitat within the Plan Area seems to be correct. It should be noted that other occurrences were noted in the literature, however these areas likely were mapped as southern California white fir or lodgepole pine forest.

    BIOGEOGRAPHY

    Subalpine coniferous forests occur in the Klamath, Cascades, Sierra Nevada, Transverse and Peninsular ranges (Holland and Keil 1995). In the southern ranges, south of Mount Whitney, it occurs mainly on the western slopes (Holland and Keil 1995).

    Elevation range estimates for this Habitat vary widely, probably due to differences in the community classification, and are presented below. Holland and Keil (1995) place the California lower elevation range between 2000 m, in the Klamath mountains, and 2900 m in the southern mountains (Holland and Keil 1995). Northern limits range from 2500 m in the Klamath mountains to 3500 m in southern California (Holland and Keil 1995). Sawyer and Keeler-Wolf (1995) describe the California range slightly more narrowly: between 2200 and 3350 m. Grouped to include lodgepole pine forest forest, whitebark pine-mountain hemlock forest, and limber pine forest, Thorne (1976) states that limber pine, as a species, occurs below the treeline between 1675 and 2900 m in the northern part of the state and 2400 to 3500 m in the southern part.

    RANGE AND DISTRIBUTION WITHIN WESTERN RIVERSIDE COUNTY

    Limber pine is the dominant tree species on the ridges and high slopes of Marion and San Jacinto peaks and is common on Tahquitz Peak (Thorne 1977). It also is known from between 2440 m and the summit of Toro Peak in the Santa Rosa Mountains. This Habitat was mapped in a small area immediately west of the San Jacinto peak on the eastern edge of the Plan Area, totaling 541 acres.

    VEGETATION CHARACTERISTICS

    There is general agreement that limber pine (Pinus flexilis) dominates this Habitat. The extent and diversity of other conifers occurring in this community variably are described in the literature. Several interpretations are presented below.

    Thorne (1976) refers to this Habitat as limber pine forest. Although pure stands are common, other pines, namely Jeffrey pine (Pinus jeffreyi) and lodgepole pine forest (Pinus contorta ssp. murrayana), may occur. Munz and Keck (1949) list the following species: whitebark pine (Pinus albicaulis), foxtail pine (Pinus balfouriana), western hemlock (Tsuga mertensiana), and willow (Salix petrophila) in addition to the list aboveSawyer and Keeler-Wolf (1995) describe limber pine occurring with lodgepole pine forest and white fir (Abies concolor) in the subalpine region of the Transverse and Peninsular ranges. According to Holland and Keil (1995) lodgepole pine forest and red fir (Abies magnifica) occur often in this community, but do not form stands, whereas Jeffrey pine and white fir occur only occasionally. Limber pine dominates an open canopy which is generally less than 18 m in height (Sawyer and Keeler-Wolf 1995).

    Shrub cover is infrequent to common and ground cover usually is sparse (Sawyer and Keeler-Wolf 1995). Many timberline shrubs occur in this Habitat including southern alpine buckwheat (Eriogonum kennedyi var. alpigenum) (Thorne 1976). Other shrub and herbaceous species include buckwheat (Eriogonum incanum), wax currant (Ribes cereum), willows (Salix spp.), western blueberry (Vaccinium uliginosum ssp. occidentale), dwarf bilberry (V. caespitosum), mountain-laurel (Kalmia polifolia), mountain gooseberry (Ribes montigenum), columbine (Aqilegia pubescens), stonecrop (Sedum obtusatum), shrubby cinquefoil (Potentilla fruitcosa), white heather (Cassiope mertensiana), mountain-heather (Phyllodoce breweri), rock-spiraea (Holodiscus microphyllus), Great Basin sagebrush (Artemisia tridentata) and beardtongue (Penstemon heterodoxus) (Holland and Keil 1995; Munz and Keck 1949). Herbaceous vegetation varies from very sparse in dry areas to lush and meadow-like in wet areas (Holland and Keil 1995).

    PHYSICAL ENVIRONMENT

    These trees generally form a relatively narrow band from about 2600 m to the timberline. Munz and Keck (1949) cite the following environmental conditions: average precipitation from 76 to 130 cm mostly as snow, growing season from seven to nine weeks, and mean summer temperatures probably less than 650 F.

    Soils at these high elevations generally are shallow, contain little organic matter, and do not retain water well (Holland and Keil 1995). Soils generally consist of coarse sand, gravel, volcanic debris, and rock materials (Verner and Purcell 1988). Wind, often carrying ice particles, are a common environmental factor in this region and limit tree growth near the timberline (Holland and Keil 1995). In this harsh environment, killing frost can occur at any time (Parsons 1980).

    In a general description, Parsons (1980) describes the distribution of subalpine conifers as “scattered among the sparsely vegetated ridges, barren rocky outcrops and draws inhabited by dense, low, riparian vegetation.” Tree density decreases as elevation increases within this Habitat type (Verner and Purcell 1988).

    ECOSYSTEM PROCESSES

    Fire is an infrequent occurrence at these high elevations. When fires occur, they usually are severe and replace the stand (Stephenson and Calcarone 1999). This may explain why limber pine outcompetes other conifers, due to its ability to regenerate without fire, its longevity and its ability to tolerate extreme cold (Verner and Purcell 1988). This Habitat occurs most frequently on low-quality soils although the limber pine grows best on calcareous soils (Barney 1980).

    THREATS

    Subapline areas are relatively undisturbed with the exception of some heavy recreation use on mountaintop trails (Stephenson and Calcarone 1999). The western dwarf mistletoe (Arceuthobium campylopodum) infects limber pine in this Habitat. Mistletoe is managed mainly through the pruning and removal of infected trees (Stephenson and Calcarone 1999).

    LITERATURE CITED

    Holland, V. L. and David J. Keil. 1995. California Vegetation. Kendall/Hunt Publishing Company.. Dubuque, Iowa.

    Munz, Philip A. and David D. Keck. 1949. California Plant Communities. El Aliso, Vol. 2, No. 1: 87-105.

    Parsons, D. J. 1980. California mixed subalpine. Pages 90-91 In F. H. Eyre, editor. Forest cover types of the United States and Canada. Society of American Foresters, Washington, D.C.

    Sawyer, John O. and Todd Keeler-Wolf. 1995. A Manual of California Vegetation. California Native Plant Society, Sacramento, CA.

    Steele, Robert. 1990. Pinus flexilis - limber pine. Pages 348-353 in R.M. Burns and B. H. Honkala, technical coordinators. Silvics of North America, Volume 1. Confiers. Agriculture Handbook 654. USDA, Forest Service, Washington, D.C.

    Stephenson, John R. and Gena M. Calcarone. 1999. Southern California mountains and foothills assessment: Habitat and species Conservation issues. General Tehnical Report GTR-PSW-172. Albany, CA: Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture; 402 p.

    Thorne, Robert F. 1976. The Vascular Plant Communities of California. In Symposium Proceedings: Plant Communities of Southern California. ed. June Latting. California Native Plant Society, Special Publication No. 2. Berkeley, California. 1-31.

    Thorne, Robert F. 1977. Montane and Subalpine Forests of the Transverse and Peninsular Ranges In Terrestrial Vegetation of California. ed. Michael G. Barbour and Jack Major. John Wiley and Sons, New York: 538-557.

    Verner, Jared and Kathryn L. Purcell. 1988. Subalpine Conifer. In A Guide to Wildlife Habitats of California. ed. Kenneth E. Mayer and William F. Laudenslayer, Jr. California Department of Forestry and Fire Protection, Sacramento, California. 40-41.