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The Effects of Irrigation On Revegetation of Semi-Arid Coastal Sage Scrub in Southern California
The Effects of Irrigation On Revegetation of Semi-Arid Coastal Sage Scrub in Southern California
1007/s002670010100
The use of irrigation for the restoration of vegetation In desert ecosystems with precipitation generally less
native to arid and semi-arid regions is controversial that 15 cm, favorable conditions for germination and
(Perry 1987, Luken 1990, Keator 1994). Though not survival of perennial species may occur as infrequently
always feasible or practical, under conditions where as once or twice in 15 years (Bleak and others 1965,
extra water can be supplied, it has the advantage of West and others 1979, Jordan and Noble 1981, Silcock
prolonging the growing season and substituting for 1986). Where rapid restoration of native arid and
unpredictable rainfall during drought years. In semi- semi-arid vegetation is desired, such as following road
arid ecosystems, favorable conditions for shrub recruit- construction, urban development, or wildfire, irriga-
ment are often episodic (Bleak and others 1965, West tion may provide an attractive alternative to depen-
and others 1979, Jordan and Noble 1981, Silcock 1986, dence on erratic precipitation. Irrigation may also
O’Connor 1996). In Mediterranean ecosystems where permit the initiation of revegetation programs outside
the majority of precipitation occurs during the winter of the normal growing season.
months, consistent recruitment following severe distur- Supplying supplemental water may negatively impact
bances is dependent on sufficient winter moisture to establishment of arid-zone plant species, however (Perry
allow seedlings to mature quickly enough to withstand 1987, Luken 1990). Prolonged dry season irrigation of
the 5–6-month summer dry season (Went 1948, Mooney arid-zone plants is known to promote the growth of
and Kummerow 1981, Moreno and Oechel 1992). In pathogenic fungi, resulting in shortened life spans
chaparral ecosystems where precipitation averages 40 to (Perry 1987, Keator 1994). During the early stages of
50 cm, successful recruitment may occur every year seed germination and growth, irrigation may artificially
(O’Leary 1990, Tyler 1995, Moreno and Oechel 1992). enhance the competitive ability of more mesic species
and individuals at the expense of seedlings better
KEY WORDS: Vegetation restoration; Seedling survival; Species even- adapted to surviving under dry conditions (Lauenroth
ness
and Dodd 1978, Gutterman and Evenari 1994). This
1Current address: USDA-Forest Service Fire Research Lab, 4955 Canyon
advantage may not continue unless irrigation is main-
Crest Dr., Riverside, California 92507, USA tained in perpetuity. Particularly when the goal of
*Author to whom correspondence should be addressed. irrigation is for short-term, rapid establishment—but
Environmental Management Vol. 26, No. 4, pp. 427–435 r 2000 Springer-Verlag New York Inc.
428 P. E. Padgett and others
not for long-term maintenance—establishment of spe- mammal, and 26 bird species as threatened or endan-
cies adapted to irrigated conditions may result in a gered (Keeley and Swift 1995). For this reason, restora-
failure in restoration once irrigation is withdrawn (El- tion of CSS has become a serious concern in southern
berse and Breman 1990). California. Yet restoration of Mediterranean ecosystems
Germination of many wildland plants is well regu- present significant management challenges because of
lated to ensure that the timing of emergence is optimal their proximity to human populations, tendency toward
for the survival of the seedling (Went 1948, 1949, fire and flooding, and in some cases lack of aesthetic
Gutterman and Evenari 1990, Bewley and Black 1994). appreciation (Mooney 1982). Because CSS shares a
The known mechanisms regulating germination vary number of features of both chaparral and desert ecosys-
widely. Some, such as the phytochrome-red light/far- tems, information concerning revegetation strategies
red light system and temperature stratification are well may be mutually applicable.
studied (Bewley and Black 1994). Others, such as nitrate In this study we evaluate the use of irrigation in the
stimulation (Hilhorst and Karssen 1988), smoke responses establishment of six shrub species native to the CSS of
(Keeley and Frotheringham 1998), or the requirement southern California. Because significant mortality oc-
of some species to pass through a wet/dry/wet cycle curs during the first growing season, particularly in the
prior to germination are less well understood (Bewley presence of weedy annuals (Moreno and Oechel 1992,
and Black 1994). Most mechanisms appear to function Eliason and Allen 1997), temporary irrigation during the
in some way to provide environmental cues. In arid and spring or summer months may provide enough additional
semi-arid environments, the most important environmental resources to enhance survival through the first summer
condition for seedling survival is the availability of water drought. Trial plots were seeded with a mixture of the
during early development (Went 1948, 1949, Elberse and species. The treatments consisted of four different watering
Breman 1990). Temperature and light provide secondary regimes: (1) no additional water, (2) irrigation during the
cues regarding the time of year, thus the expected spring months only, (3) irrigation during the summer
duration of the available moisture (Juhren and others months only, and (4) continual irrigation as needed.
1956, Bewley and Black 1994). Once germinated, the Irrigation was provided during the first year and the
success in establishment is dependent on a successful plots were monitored for 2 years. Monitoring consisted
match of physiological requirements and environmen- of monthly or bimonthly identification and assessment
tal conditions. In semi-arid regions this usually means of numbers and maximum heights of individuals and a
an adequate supply of water and nutrients until the final harvest for determination of biomass.
seedling is mature enough to withstand a dormant or
quiescent period during prolonged dry spells. Materials and Methods
Coastal sage scrub (CSS) is a vegetation type that
occupies a physiographic position between chaparral Seedbed Preparation
and western Mojave shrublands. The distribution of The experiment was conducted at the University of
CSS is limited to the populated coastal foothills and California’s Agricultural Research Station in Riverside
inland valleys of southern California, although similar from November 1995 through June 1997. The site is
ecosystems occur in Mediterranean regions around the located at 33°588 N latitude, 117°178 W longitude, and
world. Rainfall is limited to the winter months in at approximately 300 m elevation. Coastal sage scrub
southern California with a 10-year average for precipita- was the original vegetation type, but the site has been in
tion in CSS of 30 cm. However, total rainfall is quite agricultural production for more than 50 years. The
variable from year to year, ranging between 10 and 45 field was disced and graded smooth in preparation for
cm over the same 10-year period. Summer temperatures planting; no furrows were created. The field was divided
are intermediate to those of chaparral and the Mojave into 1.5-m-by-1.5-m plots with 1-m walkways between
Desert, typically hot and with low relative humidity. plots. After installation of the irrigation system, the
Several plant species common to CSS are also found in plots were raked to roughen the surface and seeds were
chaparral or desert ecosystems (Pavlik and Skinner 1994). hand broadcast on December 5, 1995. Following seed-
Agricultural and urban development, anthropogenic ing, the plots were again raked and lightly watered to
and natural disturbances, and encroachment by non- prevent the seeds from blowing away. A mixture of six
native plant species have resulted in CSS ecosystem species, typical of the CSS community, was used: Artemi-
losses of 70% to 90% of the area originally occupied sia californica, Salvia mellifera, Lotus scoparius, Eriogonum
(Westman 1981, Freudenberger and others 1987, Kee- fasciculatum, Encelia californica, and Baccharis pilularis.
ley and Swift 1995, Minnich and Dezzani 1998). This Species will be referred to by genus in the rest of the
loss of habitat has led to the listing of 10 reptile, 11 text. S&S Seed (Carpinteria, California) donated all of
Restoration of Semi-Arid Vegetation 429
until constant weight was reached. The combined more than 30% of that observed for the no-water and
weight of flowers, leaves, and stems was recorded. summer water treatments. By April 1997, Eriogonum
populations were near zero for all of the irrigated
Statistical Analysis treatments, but remained at about 25 m⫺2 in the
Data were analyzed using Jandel SigmaStat version unirrigated plots.
2.0. Seedling emergence data were analyzed by re- Salvia demonstrated very poor survival in all treat-
peated measures ANOVA for the total data set and t test ments (Figure 2C) even though initial emergence was only
for selected dates. The biomass data were analyzed by slightly less than that observed for Eriogonum. Similar to
ANOVA. The data were checked by a normality test Eriogonum seedlings, Salvia seedlings in spring water
function prior to the application of statistical test, and plots had significantly lower densities than in the other
no transformations were needed. treatments. The number of surviving seedlings began to
decline in early March of the first growing season.
Neither spring nor summer irrigation appeared to
Results
rescue the declining populations. At the end of the first
Emergence and Survival summer the numbers of seedlings were essentially zero
in the sampled subplots and remained that way through
Artemisia seedlings in the continuous water plots
the onset of the rainy season until the final harvest.
emerged earlier than those dependent on natural rainfall
Encelia, in contrast with the other species, demon-
(Figure 2A). However, seedling numbers in all of the
strated very little difference in the emergence and
treatments increased rapidly with precipitation so that
survival under different irrigation regimes (Figure 2D).
maximum densities occurred in all treatments within a
The observed low densities recorded in late summer of
week of each other. Seedling numbers in the plots not
the first year is most likely due to mistaken notation of
receiving irrigation (none, spring, summer) were very
dormant, leafless seedlings as dead. These dead seed-
similar from January until March. Once the spring water
lings releafed with the winter rains. As with Eriogonum
treatment was begun in March, germination slowed and
and Artemisia, emergence occurred earliest in the con-
densities became significantly lower than in the other
tinuous water plots. By early February 1996, the num-
treatments. The summer-watered plots contained signifi-
bers of seedlings growing in the continuous water plots
cantly more seedlings in February than either the spring
were equivalent to the numbers of seedlings in all the
or no-watered plots, but by June, seedling numbers in all
other plots. Other than a slight increase in survival of
but the continually watered plots were statistically similar.
plants in the continuous water treatment during the
In these plots the densities declined during the first
first summer, the lack of statistical difference (P ⬍ 0.1)
growing season to less than 300 m⫺2, with a small degree
among treatments continued until the end of the
of continued mortality until the conclusion of the
experiment. Once the rainy season began in October
experiment. Neither the spring water nor summer
1996, all treatments contained similar numbers of
water treatment appeared to reverse this trend, nor did
Encelia plants.
the return of natural rainfall in October 1996. The
Lotus seedling emergence occurred at much lower
seedling numbers in the continuous water plots re-
rates relative to the other species, averaging only 10 to 30
mained significantly higher from mid-July 1996 until
seedlings per m2 (Figure 2E). By summer of the second
the seedling count at the conclusion of the experiment.
year, survival of these seedlings was significantly higher
Beginning in February 1997, the average number of
in the no-water plots. Irrigation appeared to be deleteri-
seedlings in the continuous water plots dropped from
ous to this species as spring water resulted in suppressed
about 750 m⫺2 to about 200 m⫺2. By June 1997, there
germination. The greatest survival was obtained in the
was no statistically significant difference among the four
summer water plots prior to initiation of the treatment,
irrigation treatments for Artemisia seedlings.
but once irrigation began, survival plummeted.
Eriogonum seedlings in the continuous water plots
Baccharis was not well suited to this experimental site.
emerged rapidly, but the numbers of seedlings declined
No seedlings survived in the sampling subplots after the
quickly to 20% of the initial emergence within 3 months
first summer. Continual irrigation did, however, extend
(Figure 2B). Emergence of Eriogonum in the no-water
survival nearly a month longer than any of the other
and summer water plots occurred later and maintained
treatments.
significantly higher seedling numbers than the continu-
ous water treatments until early summer when all of the
treatments were statistically similar. Initiation of spring Survival of Seedlings
irrigation suppressed germination so that maximum The maximum germination rate for Eriogonum was
seedling numbers in the spring water plots were never poorly correlated with the calculated number of live
Restoration of Semi-Arid Vegetation 431
Figure 2. Seedling densities of the six species planted and the early germination and growth phase and monthly as the
grown under four irrigation regimes. A 50-cm ⫻ 25-cm subplot seedlings became established. Data shown are the means of six
was established in the center of each plot, and the number of replicate plots with a pooled standard error bar (LSD) shown
seedlings occurring in the subplot was recorded weekly during in the upper left corner of each graph.
seed sown (Table 1, Figure 2). An estimated 50% more sia, Encelia, and Salvia were about 50% and for Baccharis
seedlings emerged than were theoretically sown. The 17%. Of the total number of seedlings that emerged,
calculated densities of Lotus indicated very poor germi- nearly 25% of the Artemisia in the continuous water and
nation, at less than 3% relative to the theoretical live spring water treatments survived the first two growing
seed planted. The average germination rates for Artemi- seasons (Figure 2). The higher survival rate in the
432 P. E. Padgett and others
timing of seed sowing may have been particularly poorly under all irrigated conditions and the continu-
advantageous, however, in that the seeds were sown just ous water treatment appeared to be deleterious to both
a month before significant accumulation of rain, and Lotus and Salvia.
germination is known to be related to precipitation Coastal sage scrub contains elements of both chapar-
levels in arid lands (Gutterman and Evenari 1994). It is ral and desert vegetation. Like chaparral, the shrubs
unknown whether irrigation would have been more native to CSS established well with seeding but required
advantageous under either earlier or later sowing dates. no supplemental water, under less than average precipi-
These data suggest that increasing water availability and tation. Like desert vegetation, irrigation had a pro-
concomitant stimulation of earlier germination does nounced effect on the species composition and even-
not promote a more rapid long-term restoration as ness of the vegetation.
compared to nonirrigated conditions.
Although the advantage of earlier germination did
not appear to enhance survival, it did increase the
Conclusion
growth rate of Artemisia and speed closure of the Irrigation of a mixture of six species native to CSS
canopy. Closed canopies have an advantage in the resulted in a near monoculture of A. californica. This
reduction of annual and herbaceous weeds. Although seemed to be due to the very high seeding rate for
the removal of weedy species was not quantified during Artemisia and the ability of this species to respond
this study, it can be a significant variable in the success- positively to increased water availability under continu-
ful restoration of native vegetation (Silcock 1986). ous irrigation. Without supplemental water, five of the
Furthermore, earlier establishment and rapid growth six species seeded exhibited emergence and survival
may aid in slope stabilization and reduce erosion of rates sufficient to close the canopy by the end of the first
bare ground. However, the advantage of rapid growth growing season even in years with lower than normal
was also relatively short-lived as the canopies of all rainfall. These data suggest that where diversity of plant
treatments were closed by the end of the first growing species is an objective, irrigation was neither necessary
season, eliminating the need for hand weeding in the nor beneficial. However, there may be advantages of
second year of the experiment. earlier emergence with irrigation where slope stabiliza-
Irrigation during the spring or summer only was of tion and erosion control are important.
little benefit to seedling recruitment or to establish-
ment of surviving individuals and even seemed to be
deleterious in some cases. The seeding density and Acknowledgments
emergence rates were sufficiently high to provide Funding for this project was provided in part by the
enough survivors for stand establishment even in the California Department of Transportation. We would
absence of supplemental water. Poor survival during the like to thank the University of California, Riverside
first summer is typical for Mediterranean species (Elia- Agricultural Operations personnel, particularly Lisa
son and Allen 1997), but summer irrigation did not Warren, for the assistance with the irrigation system and
affect summer survival. The initiation of spring irriga- field preparation. The assembly of the irrigation system
tion suppressed germination in all six species exam- was made more efficient by the help of Scott Eliason. We
ined, but a greater percentage of the individuals that are very appreciative of S&S Seed for their generous
had emerged prior to the irrigation treatment survived. supply of seed. The field crew consisted of Elise Focht,
Summer irrigation had little effect over all; germination Stu Hemstreet, Lisa Jones, Nancy Storms, Jane Wahr-
and survival were not significantly different from plots man, and Tho Vo, without whom this project would
receiving no supplemental water. have never been completed.
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