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Wildlife, Animals, and Plants
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Introductory
SPECIES: Heteromeles arbutifolia | Toyon
ABBREVIATION :
HETARB
SYNONYMS :
Photinia arbutifolia
SCS PLANT CODE :
HEAR5
HEARA2
HEARC2
HEARM
COMMON NAMES :
toyon
Christmasberry
California holly
TAXONOMY :
The currently accepted scientific name of toyon is Heteromeles
arbutifolia (Lindl.) M. Romer. (Rosaceae) [27]. Heteromeles is a
montypic genus [35,45]. Three varieties are recognized based
on differences in fruit size and color [27]:
H. a. var. arbutifolia
H. a. var. cerina (Jepson) E. Murr.
H. a. var. macrocarpa (Munz) Munz
LIFE FORM :
Shrub
FEDERAL LEGAL STATUS :
NO-ENTRY
OTHER STATUS :
NO-ENTRY
COMPILED BY AND DATE :
N. McMurray, February 1990
LAST REVISED BY AND DATE :
NO-ENTRY
AUTHORSHIP AND CITATION :
McMurray, Nancy E. 1990. Heteromeles arbutifolia. In: Remainder of Citation
DISTRIBUTION AND OCCURRENCE
SPECIES: Heteromeles arbutifolia | Toyon
GENERAL DISTRIBUTION :
Toyon occurs in chaparral communities throughout much of California. It
is distributed in the Coast Ranges from Humboldt County southward into
Baja California and in the foothills of the Sierra Nevada from central
California southward into the Transverse Ranges [40,44,45,56].
Heteromeles arbutifolia var. macrocarpa is restricted to the Santa
Catalina and San Clemente islands off the coast of southern California
[35,46]. It is cultivated in Hawaii [71].
ECOSYSTEMS :
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES26 Lodgepole pine
FRES27 Redwood
FRES28 Western hardwoods
FRES34 Chaparral - mountain shrub
STATES :
CA HI MEXICO
ADMINISTRATIVE UNITS :
CABR CHIS FOPO GOGA JOMU MUWO
PINN PORE SAMO WHSH
BLM PHYSIOGRAPHIC REGIONS :
1 Northern Pacific Border
3 Southern Pacific Border
4 Sierra Mountains
7 Lower Basin and Range
KUCHLER PLANT ASSOCIATIONS :
K005 Mixed conifer forest
K006 Redwood forest
K009 Pine - cypress forest
K029 California mixed evergreen
K030 California oakwoods
K033 Chaparral
K034 Montane chaparral
K035 Coastal sagebrush
K036 Mosaic of K030 and K035
SAF COVER TYPES :
232 Redwood
234 Douglas-fir - tanoak - Pacific madrone
243 Sierra Nevada mixed conifer
248 Knobcone pine
255 California coast live oak
SRM (RANGELAND) COVER TYPES :
NO-ENTRY
HABITAT TYPES AND PLANT COMMUNITIES :
Toyon is a shrub component of chaparral, woodland, and forest
communities throughout much of California [4]. It has not been included
as a dominant or indicator in published classification schemes. Hanes
[21] lists it as one of a number of woody dominants in chamise
(Adenostoma fasciculatum) chaparral, Nuttall's scrub oak (Quercus dumosa)
chaparral, and mixed-chaparral communities. In the northern Coast Range
and foothills of the northern Sierra Nevada, toyon and chamise
codominate low, open communities constituting serpentine chaparral
[21,55]. Toyon becomes locally dominant in seral communities which are
transitional between coastal sage scrub and chaparral in southern
California and the southern part of the Coast Range [4,48]. Coast live
oak (Quercus agrifolia) woodlands occupying north slopes in the Coast
Ranges often have toyon as a conspicuous subdominant; toyon sometimes
becomes dominant within these communities in the central portion of the
Coast Range [18].
Toyon is one of a number of tall shrubs constituting scrub oak dominated
chaparral [12,25,60]. Common associates within scrub oak chaparral
include [22,47] Nuttall's scrub oak, California live oak, hollyleaf
cherry (Prunus ilicifolia), birchleaf mountain-mahogany (Cercocarpus
betuloides), redberry (Rhamnus crocea), California coffeeberry (Rhamnus
california), hoaryleaf ceanothus (Ceanothus crassifolius), chaparral
whitethorn (C. leucodermis), chamise, poison-oak (Toxicodendron
diversilobum), and honeysuckle (Lonicera spp.). Toyon is also an
important component of communities which are transitional between
chaparral and coastal sage scrub types [4,21]. California sagebrush
(Artemisia californica), California buckwheat (Eriogonum fasciculatum),
lotus (Lotus scoparius), and sage (Salvia spp.) are understory
associates within transitional communties.
VALUE AND USE
SPECIES: Heteromeles arbutifolia | Toyon
WOOD PRODUCTS VALUE :
NO-ENTRY
IMPORTANCE TO LIVESTOCK AND WILDLIFE :
Toyon browse is generally considered of little importance to livestock
or wildlife [8,56]. Plants often grow beyond the reach of most
livestock [56] and the thick, leathery leaves are largely unpalatable
[38,42,56]. Use typically occurs in the late summer or fall when more
palatable herbaceous plants are cured [2,39]. The current year's growth
is heavily utilized by domestic goats on forested sites dominated by
Monterey pine (Pinus radiata) and red gum (Eucalyptus camaldulensis)
[67]. Mature fruits are extensively utilized by numerous wildlife
species, particularly birds [11,30,37,66]. The California quail,
band-tailed pigeon, and raccoon all readily consume toyon berries.
Toyon is apparently of localized importance as deer browse in portions
of California [56].
PALATABILITY :
Palatability of toyon browse ranges from low to moderate, depending on
plant condition and community associates [2,56]. Mature plants are
rarely utilized by livestock or wildlife because of large concentrations
of tannins and cyanogenic glycosides [38,42,56]. Following fire,
however, toyon produces an abundance of leafy sprouts which are much
preferred by black-tailed deer [2,56,57].
Browse ratings for toyon in California are presented below [56]:
goats good - fair
deer good - fair
sheep poor - useless
cattle useless
horses useless
NUTRITIONAL VALUE :
NO-ENTRY
COVER VALUE :
Toyon is most commonly associated with an array of tall, broad-leaved
shrubs constituting scrub oak chaparral [20]. Tall shrub communities
dominated by scrub oak lend structural and compositional diversity to a
landscape otherwise dominated by shorter statured chamise chaparral and
provide important nesting and hiding cover for numerous birds and small
mammals.
VALUE FOR REHABILITATION OF DISTURBED SITES :
Due to its wide-spreading root system and relatively rapid growth
following disturbance, toyon is useful for erosion control on dry, steep
hillsides [37,58]. On suitable sites, transplants may grow 11 to 18
feet (3.4 to 5.5 m) tall with comparable spreads in approximately 20
years [13]. Toyon is also widely used for wildlife plantings, since the
berries are eaten by a variety of bird species [58].
Toyon seed should be collected in the fall [6]. Seedlings can be
started in nursery beds using unstratified seed in the fall or
stratified seed in the spring. Plants may also be propagated by
grafting and by cuttings [37].
OTHER USES AND VALUES :
Toyon is well known for its large and abundant clusters of bright red
berries [10], and plants are often cultivated for ornamental purposes,
especially var. macrocarpa [13,45,46]. Sprigs of toyon or "Christmas
berry" were once widely used as a commercial substitute for the more
traditional English holly (Ilex aquifolium) throughout much of
California [56,58]. Today, however, California state law prohibits
anyone from collecting the branches of wild toyon [10]. Patches of
toyon become prominent in December when the berries are particularly
conspicuous [56]. It is thought that the community of Hollywood may
have derived its name from the display of toyon on the surrounding
foothills [10].
Toyon berries are sweet and spicy and have been used historically for a
variety of purposes. West Coast Indian tribes gathered the berries for
food and medicinal uses; Spanish settlers concocted a beverage from the
berries [8,10]. Channel Island fishermen apparently used toyon bark to
tan their fishing nets [10].
MANAGEMENT CONSIDERATIONS :
Browsing: Although mature toyon is typically unpalatable, heavy use may
occur on overgrazed rangelands such as those on Santa Catalina Island.
Decades of severe overgrazing by feral animals (pigs, sheep, goats) has
removed more palatable species and has converted chaparral stands into
open, arborescent woodlands. Within these communities, toyon often
exhibits a noticeable browse line and a trend towards increased trunk
diameter, canopy area, and height. Toyon can recover from prolonged
overuse. On sites where feral sheep grazing was excluded, plants
immediately produced basal sprouts and within 3 years lost their pruned
appearance [5].
Herbicides: Toyon is sensitive to such herbicides as 2,4-D and 2,4,5-T
[51,68]. If sprouts are treated following burning, plants are killed by
retreatment [68].
BOTANICAL AND ECOLOGICAL CHARACTERISTICS
SPECIES: Heteromeles arbutifolia | Toyon
GENERAL BOTANICAL CHARACTERISTICS :
Toyon is a native, broad-leaved, sclerophyllous, arborescent shrub which
typically grows from 6 to 10 feet (1.8 to 3 m) tall [9,24,45,56,58]. On
more favorable sites toyon may occasionally attain tree stature,
reaching heights of approximately 33 feet (10 m); however, in these
instances it typically retains a shrublike form [9]. Plants are erect,
freely branched, and unarmed [45]. Older branches have gray bark [45].
The dense foliage is composed of simple, evergreen leaves which are from
2 to 4 inches (5 to 10 cm) long and arranged alternately on the stem;
leaf margins are sharply toothed and bristly pointed [56,58]. The
inconspicuous, bisexual flowers are white in color and occur in loose,
somewhat flat, terminal clusters [56]. The bright red, berrylike fruits
are approximately 0.25 inch (5 to 6 mm) in diameter and contain one or
two, small brown seeds [37,45,46]. Toyon has a strong and much-branched
root system that is deeply penetrating and wide spreading [24]. Feeder
roots are abundant in the surface humus around the plant base as well as
elsewhere throughout the extensive root system. In response to repeated
postburn sprouting, toyon sometimes develops an enlarged rootcrown which
is irregularly shaped and platformlike [30]; this structure, however, is
not a lignotuber [30,31]. Longevity of toyon is estimated to be from
100 to 200 years [30].
Morphological distinctions between varieties are presented below
[45,46]:
H. a. var. arbutifolia - typical variety; fruit red
H. a. var. cerina - fruit yellow
H. a. var. macrocarpa - large fruited form; fruit red; leaves subentire
RAUNKIAER LIFE FORM :
Phanerophyte
REGENERATION PROCESSES :
Toyon regenerates by both sexual and vegetative means. Within the
fire-prone chaparral environment, it maintains itself primarily through
vigorous sprouting [29,31,32]. Little or no seedling establishment
occurs immediately following fire [31,63]. Extended fire-free periods
are required for successful seedling establishment and population
expansion [32,33].
Vegetative regeneration: During extended fire-free intervals, toyon
rejuvenates its crown by continually producing new sprouts from
established rootcrowns [31,33]. Following disturbances such as fire or
cutting, toyon sprouts from surviving adventitious buds on the rootcrown
[55,58].
Seedling reproduction: Onset of seed production occurs early in toyon;
2-year-old transplants can produce seed during the second season
following outplanting [13]. Abundant seed is produced annually after
the first flowering [37,54], and seed production apparently does not
decrease with age [29]. Seeds are dispersed in the late fall and winter
[32]. Significant, widespread dispersal of the persistant, fleshy
fruits occurs through animals, particularly birds [11,30,43,63,64,66];
if not dispersed by animals, seeds fall directly beneath the parent
plant. Seeds are short lived and retain their viability for not more
than 9 months when dried at room temperature [32]. Germination occurs
readily under favorable moisture and temperature conditions, often
within 10 to 40 days [37,41]. Seeds germinate without the stimulus of
heat or charred wood; in fact, heat treatments are generally lethal to
toyon seed [32]. Germinative capacity under greenhouse conditions
ranges from 73 to 99 percent [32,37,41]. Although fresh seeds do not
need stratification, stored seeds require stratification for 3 months at
temperatures of 35 to 41 degrees F (1.6 to 5 degrees C) to enhance
germination [37,41].
Toyon seedlings are never very abundant within chaparral communities
[63]. Limited observations suggest that seedling establishment is
episodic and coincides with periods of above-normal rainfall [31,63]. A
series of wet years apparently produces an abundant seed crop and also
creates a moisture regime conducive to seedling establishment [31].
Successful establishment occurs within mature chaparral in gaps
resulting from the death of senescing, shorter lived species [64].
Although abundant initial establishment may occur in burned or unburned
stands during years of above-normal precipitation [64], continued
survival is favored beneath mature stands on sites that are relatively
mesic (north slopes) and which possess a well-developed litter layer
[31]. Seedlings are frequently stunted and susceptible to browsing from
small mammals [26]. Consequently, long-term survival beneath mature
chaparral is rare [31].
Established seedlings are most common in very old stands (60 to 100+
years) [31,32]. Toyon seedling densities (plants established in recent
decades) within a 117-year-old stand of scrub oak chaparral equalled
approximately 50 per hectare [33]. Long fire-free intervals apparently
allow for the buildup of seedling populations resulting from episodes of
establishment. Zedler [64] speculates that gap size is crucial to a
seedlings ability to survive to the stage where it can resprout
following fire. The average size of unoccupied gaps tends to be larger
in infrequently burned stands where the size of senescing individuals is
also larger [63,64]. Obligate sprouters such as toyon apparently
require fire-free intervals of 100 years or more for significant
recruitment of new individuals [31].
Newly emerged toyon seedlings have been observed as early as
mid-February on undisturbed mixed-evergreen sites in central California
[49]. Mortality was high within these communities with nearly all
seedlings dying from damping-off fungus or leaf spot disease [49].
SITE CHARACTERISTICS :
Toyon is a characteristic species of chaparral and foothill woodland
communities throughout its distribution [20,45]. It usually grows in
scattered stands [9,56] on semidry, rocky slopes within foothills,
mountains, and canyon bottoms at elevations below 4,000 feet (1,220 m)
[37,45]. Soils supporting toyon are typically dry and well drained and
may occasionally be somewhat saline [58]. Although occurring on a
variety of aspects, toyon is most often associated with relatively mesic
chaparral communities, occupying cool, north exposures, erosion
channels, arroyos, depressions, and toeslopes [20,60].
SUCCESSIONAL STATUS :
Toyon is a long-lived and relatively shade-tolerant species which is
highly persistent within chaparral and woodland chaparral communities
[20,22,30]. Although widespread, toyon is not usually abundant [9]. It
becomes more common within mesic types of chaparral, particularly stands
dominated by scrub oak. Scrub oak chaparral typically experiences
reduced fire frequencies relative to chamise chaparral [30]. During
extended fire free intervals, toyon is able to outlive, overtop, and
shade out many shorter lived species [64]; seedling establishment then
occurs in newly created gaps beneath the mature canopy [47,63].
Successional studies in scrub oak chaparral indicate that limited toyon
seedling establishment may occur in stands which remain unburned for 10
to 20 years [23,25]. As a stand matures, toyon gradually increases in
prominence until it is an important codominant of 65-year-old stands
[47]. In stands beyond 65 years of age, scrub oak and hollyleaf cherry
increase in dominance while toyon decreases in size and prevalence
[19,23]. However, toyon typically persists within mature chaparral
until the next fire occurs, at which time resprouted individuals become
part of the initial postburn vegetation [7,33,58]. Toyon is also
capable of pioneering eroded sites [58].
SEASONAL DEVELOPMENT :
Studies of seasonal cambial growth indicate that toyon is active from
December through mid-June, with peak activity occurring between February
and April; stem growth ceases from late June to December during the
prolonged summer drought [1,43]. Leaves are initiated in the early
spring, and maximum size is reached soon afterwards; secondary leaf
growth occurs during the remainder of the growing season, resulting in
increased tissue density [11]. Flowering occurs from June through July
[45,56]. Fruit development is prolonged; berries first appear in August
or September and persist until December when they reach maturity
[11,58].
Although toyon is able to photosynthesize year-round, carbon is
partitioned differentially among plant functions and varies according to
season [11]. During the stem growth period in the spring, most fixed
carbon goes to the development of the canopy (new leaf and stem tissue)
which gives toyon a competitive advantage. During periods of little
stem growth, carbon is allocated to root growth, fruit production, and
into compounds which apparently provide a degree of predator protection
[43]. Although leaves are characterized by high levels of both tannins
and cyanogenic glucosides from the time of initiation, tannin levels
reach a maximum in September and October when herbivore pressures are
particularly high. The pulp of immature fruits contains a toxic
cyanogenic glucoside that protects developing fruits from bird
predation. During the long maturation process, however, bird dispersal
of the seeds is encouraged as cyanogenic glucosides are gradually
shifted from pulp to seed, pulp carbohydrate levels increase, and fruits
turn from green to bright red. Subsequent seed predation is prevented
by the localization of cyanogenic glycosides in the seeds [11].
Compared with other chaparral shrubs, toyon apparently has moderate
water use requirements [53]. Maximum water stress generally declines
well before before the onset of fall rains, apparently in response to a
decrease in ambient air temperatures [50].
FIRE ECOLOGY
SPECIES: Heteromeles arbutifolia | Toyon
FIRE ECOLOGY OR ADAPTATIONS :
Following fires which kill aerial stems, toyon sprouts vigorously from
dormant buds located on a rootcrown [55,56,58]. The rootcrown serves as
a source of numerous perennating buds and stored carbohydrates, enabling
toyon to rapidly reoccupy the initial postburn environment [31,38].
Compared with other chaparral shrubs, toyon is relatively nonflammable
[58].
POSTFIRE REGENERATION STRATEGY :
Tall shrub, adventitious-bud root crown
FIRE EFFECTS
SPECIES: Heteromeles arbutifolia | Toyon
IMMEDIATE FIRE EFFECT ON PLANT :
Toyon is quite resistant to fire mortality [64]. Although aerial
portions may be killed, most plants survive fire [23,51,59]. In fact,
toyon appears to suffer very little fire mortality even when subjected
to short-interval fires. On chaparral sites in southern California,
toyon sprouted following a 1979 wildfire and resprouted in 1980 when a
grassfire reburned the site [65].
DISCUSSION AND QUALIFICATION OF FIRE EFFECT :
NO-ENTRY
PLANT RESPONSE TO FIRE :
Toyon is an obligate sprouter following fires on chaparral sites
[30,31,64]. Vigorous sprouting is the primary means by which toyon
restablishes in the postburn community [29,32]. Seedlings rarely occur
immediately following fire [30,63].
Vegetative regeneration: Toyon sprouts vigorously following fires which
kill the aerial stems [24,51,55]. Following a hot, July wildfire in
southern California chaparral, toyon plants occupying relatively moist
sites produced sprouts within 10 days [51]. Elsewhere on the burn, the
majority of toyon individuals had sprouted by December at which time
plants usually exhibited at least 12 sprouts per plant and sprout
heights of over 6 inches (15 cm) [51]. Although toyon cover is
initially reduced following burning, most plants rapidly regain their
preburn size and biomass [19,47,61]. Following a wildfire on scrub oak
chaparral sites in southern California, toyon produced sprouts 4 to 5
feet (1.2 to 1.5 m) tall within 4.5 years [23].
Seedling reproduction: Unlike many chaparral species, toyon seeds are
not well adapted to resist fire or for long-term survival in the soil
[32,63]. However, since these short-lived seeds germinate readily under
favorable temperature and moisture conditions, some postburn
establishment may occur through bird dispersal of off-site seed [63].
Generalized information on obligate sprouters indicates resprouted
plants begin to produce seed crops within 1 to 2 years of burning and
that postburn fruit crops are often substantial [31].
While seedlings are rarely observed during the first postburn season
[51,62,65], exploitation of fire-created gaps can occur during periods
of above-normal precipitation [31,63]. Following two wet winters, toyon
established seedlings on a 3-year-old burn in Tecate cypress (Cupressus
forbesii) chaparral [63]. Unusually large numbers of toyon seedlings
occurred on both burned and adjacent unburned sites, but few survived
[63]. Although periods of above-normal precipitation are apparently
adequate for the initial establishment of toyon seedlings, successful
seedling establishment seems restricted to mesic sites beneath mature
chaparral where litter layers are well developed [31]. Keeley [31]
speculates that recruitment of new toyon individuals is never very
abundant and occurs primarily between fires rather than after fire [63].
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE :
NO-ENTRY
FIRE MANAGEMENT CONSIDERATIONS :
Prescribed burns: Toyon is a characteristic component of scrub oak
chaparral. These communities generally lack an herbaceous understory
and do not carry fire as readily as chamise or coastal sage scrub
communities [17].
Burn frequency: Although toyon is a characteristic species of
relatively infrequently burned stands of chaparral [22,31], it appears
adapted to a wide range of fire frequencies [31]. Few individuals die
as a direct result of fire, and seedlings are not established in the
immediate postburn environment [62]. Keeley [31] generalizes that
obligate sprouting species such as toyon are resilient to a burning
regime with a recurrence interval of 10 to 100 years. Stand age at the
time of burning can have a significant impact on the recruitment of new
toyon individuals, however [23]. Microsites necessary for significant
population expansion are largely unavialable in stands with fire-free
intervals of less than 100 years [31,64].
Fuels reduction: Domestic goats can be used as a method of fire hazard
reduction on forested sites where toyon is a conspicuous component of
the understory. In communities dominated by Monterey pine and red gum,
cover of toyon at heights of 1.6 to 4.9 feet (0.5-1.5 m) was reduced 60
percent in 1 day with stocking rates of 600 Spanish goats per hectare
[67].
Wildlife management: Burning initially increases the palatability of
toyon browse; sprouts are generally utilized for up to 2 postburn
growing seasons [56,57].
REFERENCES
SPECIES: Heteromeles arbutifolia | Toyon
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Index
Related categories for Species: Heteromeles arbutifolia
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