Weed Science Society of America

Impact, Biology, and Ecology of Saltcedar (Tamarix spp.) in the Southwestern United States
Author(s): Joseph M. Di Tomaso
Source: Weed Technology, Vol. 12, No. 2 (Apr. - Jun., 1998), pp. 326-336
Published by: Weed Science Society of America and Allen Press
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Weed Technology. 1998. Volume 12:326-336

Impact, Biology, and Ecology of Saltcedar (Tamarixspp.) in the

SouthwesternUnited States1

JOSEPHM. DI TOMASO2

Abstract: Eight species of Tamarixwere first broughtto North America in the 1800s from southern
Europe or the eastern Mediterraneanregion. Many of the species escaped cultivation and by the
1920s invaded about 4,000 ha of riparianhabitatin the southwesternUnited States. By 1987, it was
estimated to have increased to at least 600,000 ha. The success of saltcedarin the southwest can be
attributedto several factors related to its growth habit, reproduction,water usage, ability to tolerate
highly saline conditions, and redistributionof salt from deep in the soil profile to the soil surface.
The flowers produce small, numerous, and tufted seeds that can be carried long distances by wind
or water.The seeds, however, have a shortperiod of viability, and must come in contact with suitable
moisture within a few weeks of dispersal. Unlike obligate phreatophytes,such as willows and cot-
tonwoods, saltcedaris a facultativephreatophyteand is often able to survive underconditions where
groundwateris inaccessible. The high evapotranspirationrates of saltcedarcan lower the water table
and alter the floristic composition in heavily infested areas. Mature plants are tolerant to a variety
of stress conditions, including heat, cold, drought, flooding, and high salinity. Saltcedar is not an
obligate halophyte but survives in areas where groundwaterconcentrationsof dissolved solids can
average 8,000 ppm or higher. In addition, the leaves of saltcedarexcrete salts that are deposited on
the soil surface under the plant, inhibiting germinationand growth of competing species.
Nomenclature: Saltcedar,Tamarix./
Additional index words: Phreatophyte,invasive weed, riparianecosystem, Tamarixaphylla, Ta-
marix chinensis, Tamarix gallica, Tamarixparviflora, Tamarix ramossissima, TAAAP, TAACH,
TAAGA, TAAPA, TAARA.

INTRODUCTION shade, stabilize eroding streambanks, or were grown as

Saltcedar was originally thought to have been intro-
duced from southern Europe or the eastern Mediterra-
nean region by the Spaniardsin the 1800s, but now it is
believed that the first introductionto North America was
made by nurserymen on the east coast of the United
States in 1823 (Horton 1964). It was sold at that time
by the Old AmericanNursery operatedby Lawrenceand
Mills and later by Bartram'snursery of Philadelphia.In
1868, the United States Departmentof Agriculturegrew
six different species of saltcedarin its departmentalar-
boretum.At least three species of Tamarixwere brought
from the eastern United States and first appearedfor sale
in western nurseries as early as 1854 (Brotherson and
Winkel 1986). During these early years of introduction,
saltcedars were planted to create wind breaks, provide
I Received for publication June 27, 1997, and in revised form February
12, 1998.
2
Noncrop Weed Ecologist, University of California,Davis, CA 95616.
ornamentalshrubs or trees (Neill 1985). Escapes from
cultivation were first reportedin the 1870s (Brotherson
and Field 1987; Brotherson and Winkel 1986) and oc-
cuffed only slowly until the 1920s (Robinson 1958). By
the 1920s it became clear that some species were becom-
ing a serious problem (Brothersonand Winkel 1986).
Spread and Distribution. Saltcedar infestations in the
1920s covered about 4,000 ha (Neill 1985). In 1960 a
comprehensive survey estimated that saltcedar covered
362,000 ha of riparianhabitatsfrom Oklahomato south-
ern California and from Colorado to Sonora, Mexico
(Friederici1995; Neill 1985). By 1970, infestationswere
estimated to have increased to 540,000 ha (Gay and
Fritschen 1979). Recent estimates indicateinfestationsin
the southwestern United States to exceed 600,000 ha
(Brothersonand Field 1987). This representsat least a 3
to 4% increaseper year.In the Coloradoand GreenRiver

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 WEED TECHNOLOGY
systems, saltcedar spread was calculated to be about 20
km of river length per year (Graf 1978).
Today, saltcedarinfestations are common in most riv-
er systems over much of Utah, Nevada, Arizona, south-
ern California (Cache Creek in northern California),
New Mexico, Colorado, Oklahoma, and Texas (Horton
1977; Robinson 1965). Saltcedaris continuing to spread
northward into Montana and Canada (DeLoach 1989;
Swenson et al. 1982) and southwardinto northwestern
Mexico (DeLoach 1989). Populationshave been reported
at elevations as high as 2,135 m in the southernRocky
Mountains (Everitt 1980) but typically occur below 500
m (Brothersonand Winkel 1986).
Invasions typically occur in bare, moist, exposed ar-
eas. Initially, saltcedaris associated with mesquite (Pro-
sopis spp.), cottonwoods (Populus spp.), and willows
(Salix spp.) but eventually replaces these naturalwoody
riparian species (Frasier and Johnsen 1991). In Afton
Canyon near Barstow, CA, saltcedarreplaced70% of the
original native vegetation since the 1960s (deGouvenain
1996). In the lower Colorado River, saltcedar replaced
up to 90% of the ripariancommunitieshistoricallydom-
inated by cottonwood-willow forests (Crins 1989). Once
established, periodic burning, clearing, and flooding
have allowed saltcedar communities to remain young
and to form nearly impenetrablethickets (Everitt 1980).
FactorsContributingto SaltcedarInfestation.A num-
ber of factors can lead to the establishment and spread
of saltcedar.Perhapsthe most importantis the develop-
ment of water managementprogramsthat severely affect
naturalriver flows (Egan et al. 1993; Kerpez and Smith
1987; Sala and Smith 1996). These alterations include
reservoir and dam construction, river diversions, flow
regulations,and irrigationprojects.Historically,the flow
of these rivers peaked in the late spring and early sum-
mer from snowmelt. However, constructionof diversion
and storage facilities in the 1900s reducedthe magnitude
of these high flows and associated channel movements
and subsequent formation of bare, moist alluvial bars
during the time in which cottonwoods and willow seeds
were viable. These changes in channel geometry and
streamflowcreated conditions unfavorablefor the regen-
eration and survival of native riparian species (Engel-
Wilson and Ohmart 1978; Everitt 1980; Shafroth et al.
1995). As a result, rapid colonization and expansion of
saltcedar occurred throughoutthe western river systems
(Horton 1977; Howe and Knopf 1991; Sala and Smith
1996). In many riparian systems of the southwestern
United States, salinity increases, caused by changes in
water flow, have favored salt-tolerant species such as
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saltcedarand greatly reducedthe recruitmentand growth
of native salt-sensitive riparianspecies (Brothersonand
Field 1987; Sala and Smith 1996).
In addition to altering streamflow,clearing and plow-
ing of floodplains and associated agriculturalactivity
also aided saltcedarcolonization during the 1800s (En-
gel-Wilson and Ohmart 1978; Everitt 1980; Shrader
1977). Saltcedarhas also been reportedto rapidly infest
riparianareas exposed to heavy grazing (Hughes 1993).
Although intensive grazing by domestic livestock can
quickly eliminate the more desirable native seedling or
suckers (Engel-Wilson and Ohmart1978; Horton 1977),
seasonal grazing in regenerationzones has been shown
to provide a competitive advantageto native willows and
cottonwoods (Hughes 1993).
The planting of saltcedar for bank protection or ero-
sion control has also contributedto the rapid spreadand
establishment of saltcedar in stream systems (Everitt
1980). Other forms of disturbance, including off-road
vehicle traffic(Lovich et al. 1994) and harvestingof cot-
tonwood, mesquite, and other trees and larger shrubsfor
fuel and building materialsby early pioneers (Brotherson
and Winkel 1986; Horton 1977) created open sites suit-
able for saltcedarinfestations.
TAXONOMY
Tamarixis one of four genera of Tamaricaceaeand is
representedby 54 species worldwide (Baum 1967). The
genus was named after the TamarisRiver in Spain and
consists of halophytic shrubs and small trees native to
China and Mongolia throughsouthernEurope, the Med-
iterraneancountries, the Middle East, and North Africa
(Baum 1967). Some species also occur in the Canary
Islands and South Africa. Eight species of Tamarixhave
been introducedinto the United States, primarilyas or-
namentalsor for wind breaks and shade. All species ex-
cept T. aphylla (L.) Karst. #3 TAAAP (athel tree) are
generally referredto as saltcedar.The name "saltcedar"
refers to the small scale-like leaves that resemble cedars
and the salty exudate that collects on the foliage (Frasier
and Johnsen 1991).
Of these eight introducedspecies, five are present in
the southwest (DiTomaso 1996). Most species are
weedy, particularlyT. parviflora DC. # TAAPA, previ-
ously known as T. tetrandraPall., T. ramosissimaLed-
eb. # TAARA, previously known as T.pentandraPallas.,
and T. chinensis Lour. # TAACH. Tamarixchinensis is
Letters following this symbol are a WSSA-approved computer code from
Composite List of Weeds, Revised 1989. Available from WSSA, 810 East
10th Street, Lawrence, KS 66044-8897.
327
 DI TOMASO: SALTCEDARIN THE SOUTHWESTERNUS
Table 1. Comparisonof the five most common Tamarixspecies in the southwesternUnited States.
Species Height Leaves
T. aphylla tree < 12 m not overlapping,
strongly clasping
T. chinensis tree < 10 m overlapping, oblong to
narrowlylanceolate
T. gallica shrub or tree < 8 m overlapping, linear to
narrowlylanceolate
T. parviflora shrub or tree 1.5-5 m overlapping, linear
T. ramosissima shrub or tree < 8 m overlapping, ovate
difficult to distinguish from T. ramosissima and is oc-
casionally thought to be the same species. Tamarixgal-
lica is less widely distributedthan other weedy Tamarix
species and is also similar to T. ramosissima. One less
weedy species is the large evergreenathel tree (T. aphyl-
la). Table 1 compares the morphology and distribution
of the saltcedarspecies found in the southwesternUnited
States. Most species of Tamarix, with the exception of
T. aphylla and T. parviflora, are very difficult to distin-
guish from each other.The characteristicstypically used
to separatethese species include featuresassociated with
leaf shape and base, as well as nectar disk shape and
arrangementof the stamens.
IMPACT
Until the twentieth century, saltcedarwas considered
to have a positive effect on riparianecosystems. Its sta-
bilizing effect on the lake edge soils and its ability to
desalinize the lower soil profiles througha salt excretion
mechanism were considered desirable features (Gold-
smith and Smart 1982). In addition, its roots stabilized
streambanksagainst flooding, the foliage provided im-
portant shade and forage for domesticated animals,
woody stems were an importantsource of firewood, and
the showy white or pink flowers were desirable orna-
mentals (Friederici 1995). Beekeepers also utilize salt-
cedar for honey production(Everitt 1980).
However, the negative attributesof saltcedarfar out-
weigh its desirable qualities. Infestations have caused
significant changes in flooding and erosion patterns,fire
frequency, and both plant and animal diversity.
Flooding and Erosion. The extensive root system of
saltcedar species is more stable and resistant to erosion
than most native riparian trees and shrubs. However,
heavy infestations can significantly alter the hydrology
of an area. When stream channels are stabilized they
become more immobile and inflexible (Graf 1978),
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Flowers Distribution
5-parted,nectar disk lobes wider than long, widely cultivated, mainly in southwest,
stamens alternatedisk lobes rarely weedy
5-parted,nectar disk lobes wider than long, common in Arizona, New Mexico, Tex-
stamens alternatedisk lobes as, and Oklahoma,becoming a prob-
lem in Montana
5-parted,nectar disk lobes longer than less common than other species, mostly
wide, stamens together with disk lobes on the Texas Gulf Coast
4-parted,nectar disk lobes longer than widespread,most common in Arizona
wide, stamens together with disk lobes and California
5-parted,nectar disk lobes wider than long, widespread,most common in Arizona
stamens alternatedisk lobes and California
which progressively restricts channel width by increas-
ing sedimentdeposition. Narrowingof the water channel
increases the rate of water flow and the potential and
severity of subsequentfloods (Egan et al. 1993; Frasier
and Johnsen 1991; Friederici 1995; Kerpez and Smith
1987). A saltcedar-infestedarea on the Gila River in Ar-
izona had a 30% increase in water flow velocity and a
13% increase in water depth than a comparable unin-
fested area on the same river (Great Western Research
1989). As the river recedes, saltcedar establishes itself
furtherout into the channel. This process continues until
streamflowis severely reduced. Saltcedarinfestationsin-
creased on the Brazos River in north central Texas be-
ginning in 1941. At this time the mean width of the river
channel along a 121-km stretchwas 155 m. By 1979 the
mean width had been reduced to 66 m. This narrowing
in the channel width increasedthe incidence of flooding,
as well as the area inundatedby the floodwaters(Black-
burn et al. 1982).
Fire Frequency. Saltcedaris a fire-adaptedspecies with
more efficient fire recovery mechanisms than nearly all
other native riparian species (Anderson et al. 1977;
Busch and Smith 1993). It has been suggested that salt-
cedar is able to utilize volatilized nutrients,increasedsoil
concentrationsof mineral elements, increased soil pH,
and reduced available moisturebetterthan native woody
riparianspecies following fire events (Busch and Smith
1993). This adaptationhas likely been a significantfactor
promotingits rapidcolonization of water courses (Busch
and Smith 1992, 1993; Wiesenborn 1996).
In native riparian plant communities dominated by
cottonwood, willows, or mesquite, wildfires appearto be
infrequent(Busch and Smith 1993). In contrast,intervals
between fires are considerably shorter in saltcedar-in-
fested areas. Thus, it has been hypothesized that saltce-
dar, like other plant species that possess resprouting
mechanisms, might have developed adaptive character-
istics that enhance the flammabilityof the communities
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 WEED TECHNOLOGY
where they grow (Busch and Smith 1992). This can lead
to the replacementof nonadaptedfire communitiesdom-
inated by cottonwood and willow (Busch and Smith
1992; Kerpez and Smith 1987). In support of this, An-
derson et al. (1977) demonstratedthat 21 of 25 saltcedar
stands along the lower Colorado River burned within a
15-yr period. Similarly, Busch (1995) noted that fires
burned 35% of saltcedar-dominatedvegetation on the
lower Colorado River floodplain between 1981 and
1992, compared to 2% of communities of honey (Pro-
sopis glandulosa Torr.# PRCJG)or screwbeanmesquite
(Prosopis pubescens Benth.) duringthe same period.The
increased incidence of fire in saltcedar stands has been
attributedto the rapid accumulationof leaf litter,as well
as dead and senesced woody material (Busch 1995;
Busch and Smith 1993; Kerpez and Smith 1987). Lovich
et al. (1994) noted that fuel buildup by saltcedar pro-
motes a fire about every 10 to 20 yr in North American
desert ripariansettings.
Plant Diversity. Saltcedar stands can be considerably
denser than naturally occurring riparianvegetation his-
torically dominated by cottonwood-willow and screw-
bean mesquite (Egan et al. 1993). In some areas, 70 to
80% of the vegetative cover can consist of saltcedar.
Such infestations lead to the dramaticreductions in na-
tive woody and herbaceousplant composition and abun-
dance (Engel-Wilson and Ohmart 1978; Hughes 1993;
Lovich et al. 1994; Weeks et al. 1987).
Fire frequency, as well as other factors, in saltcedar
communitieshas dramaticallyhinderedthe establishment
of native riparian species. When present in saltcedar-
dominatedcommunities, cottonwood populationsconsist
solely of mature or nearly mature trees. Because these
communities lack a broad age distributiontypical of a
vigorous reproducingcommunity, they are less able to
spreadand reduce present saltcedarstandsthroughshad-
ing (Shrader 1977). Along the Rio Grande,cottonwood
regeneration has not occurred for 30 to 35 yr. Conse-
quently, the ripariancottonwood woodland is typically
35 to 60 yr old and its abundance is expected to dra-
matically decline within the next 50 yr (Howe and Knopf
1991).
Animal Diversity. Saltcedar rarely provides food and
shelter necessary for the survival of wildlife (Shrader
1977). Although some wildlife species successfully sur-
vive in saltcedar-dominatedareas, most species are neg-
atively effected by the displacement of native riparian
plant species and other habitat changes resulting from
encroachment of saltcedar.In addition, the sticky sub-
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stance exuded by saltcedar leaves can also damage the
plumage of birds (Cohan et al. 1978).
Insects. Willow and cottonwood supporta greaterabun-
dance of insect life than does saltcedar(Neill 1985). Per-
haps the only insect species that thrive on saltcedarin
the southwest are cicadas and the Europeanhoney bee
(Apis mellifera) (Horton and Campbell 1974). Saltcedar
can provide not only an early source of pollen for over-
wintering bees (Kerpez and Smith 1987) but can be a
beneficial species for honey production(Shrader 1977).
Aside from bees and cicadas, very few insects are known
to use saltcedar as cover or forage (Egan et al. 1993).
Even when insects are present, their reproductionfluc-
tuates dramatically on saltcedar compared with native
riparianhabitat (Carotherset al. 1976).
Birds. Some obligate riparianbird species can success-
fully utilize saltcedar (Ellis 1995). However, most of
these continue to show preferencefor more diverse, na-
tive plant communities (Shrader 1977). These include
various doves, Gambel'squail (Lophortyxgambelii), and
other granivores(seed feeders) or other types of ground-
feeding birds. Because the seeds of saltcedar are too
small to be eaten by most animals (Neill 1985), bird
populations that roost in saltcedar generally forage in
nearby agriculturalfields. Among the bird species that
utilize saltcedar,doves maintainthe highest density (An-
derson et al. 1977), particularlywhite-winged (Zenaida
asiatica) and mourning doves (Z. macroura). Although
saltcedar provides nesting sites for these doves, their
populations are usually higher in nonsaltcedarcommu-
nities, especially mesquite (Shrader 1977). In addition,
saltcedar provides less food for doves than does mes-
quite (Kerpez and Smith 1987). The endangeredsouth-
western subspecies the willow flycatcher (Empidonax
trailii extimus) will also nest in saltcedarwhen its pref-
erable nesting species, willows, have been displaced
(DeLoach et al. 1996). Some species, such as Gambel's
quail, prefer honey mesquite and cottonwood commu-
nities but will utilize saltcedarstands for shelter,though
not for nesting (Shrader1977).
Although some riparianbird species have continued
to breed as saltcedar has replaced native vegetation,
breeding densities of these populations have declined.
On the Colorado River, migratory and resident spring
and summer breeding species show a trend of declining
use of saltcedar from west to east, and are largely re-
stricted to cottonwood-willow communities (Hunter et
al. 1985, 1988). The Bell vireo (Vireo bellii) on the low-
er Colorado River is nearly excluded as a breeding spe-
329
 DI TOMASO: SALTCEDARIN THE SOUTHWESTERNUS
cies. Summer tanagers (Piranga rubra) and yellow-
billed cuckoos (Coccyzusamericanus) are also in serious
danger of elimination from the lower Colorado River.
The majorityof birds, particularlyriparianspecies, are
more strictly tied to native plant communities (Hunteret
al. 1985, 1988; Ellis 1995). Waterfowl,frugivores (fruit
and seed feeders) and insectivores, usually abundantin
native riparianvegetation, almost completely avoid salt-
cedar (Brotherson and Field 1987; Kerpez and Smith
1987; Shrader 1977). Many frugivores feed on the fruit
of desert mistletoe (Phoradendron californicum Nutt.).
This parasiticplant grows on native woody species (Co-
han et al. 1978), but more uncommonly on saltcedar
(Haigh 1996).
When comparingtotal bird density and species diver-
sity, saltcedar stands consistently had lower values than
communities dominated by cottonwood, willow, and
mesquite (Anderson et al. 1977; Cohan et al. 1978; En-
gel-Wilson and Ohmart 1978; Hunteret al. 1985, 1988;
Kerpez and Smith 1987). In one study, Anderson et al.
(1977) found that cottonwood-willow communities
proved the most valuable to bird populations, followed
by honey mesquite, screwbean mesquite, saltcedar-hon-
ey mesquite, desert wash, saltcedar,orchard, and arro-
wweed [Pluchea sericea (Nutt.) Cov.]. Along the Colo-
rado River, the native riparianareas sustained a density
of 154 birds per 40 ha, whereas the saltcedar-dominated
areas consisted of four birds per 40 ha (Johnson 1987).
Along the Pecos River in New Mexico, more birds were
observed in 39 ha of cottonwood, willow, and mesquite
communities than in 19,600 ha of saltcedar(Engel-Wil-
son and Ohmart 1978).
Restoration projects may have a dramatic effect on
breeding populationsin saltcedarstands. Andersonet al.
(1977) noted that the addition of one or more native tree
species, even in small numbers, greatly enhanced the
overall attractivenessof an area to breeding pairs.
Mammals. With the exception of woodrats (Neotoma
spp.) and the desert cottontail (Sylvilagus audubonii),no
native mammal species are known to feed on mature
saltcedar.Only in some cases is young growth utilized
by mammals. When saltcedarwas cleared from a 20-ha
area along the lower Colorado River and replaced with
native vegetation, the diversity of both birds and rodents
increased significantly (Anderson and Ohmart 1985). In
a study by Engel-Wilson and Ohmart(1978), mammals
such as porcupine (Erethizon dorsatum) and beaver
(Castor canadensis) had a high affinity for the cotton-
wood-willow association, but occurredin very low den-
sities in saltcedar-dominatedcommunities.
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BIOLOGY
Saltcedar species are phreatophytes (deep-rooted to
reach water table) that often depend on groundwaterfor
their water supply. However,undersome conditions salt-
cedar can grow where no groundwater is accessible.
Thus, it is classified as a facultative ratherthan obligate
phreatophyte(Kerpez and Smith 1987). The typical life
span of an individual saltcedartree has been reportedto
be 75 to 100 yr (Horton 1977).
Shoot Growth. Though the seeds will germinaterapidly,
new seedlings require wet soils for several weeks (Hor-
ton 1977). Under ideal conditions, seedlings can grow 3
to 4 m in a single growing season (Friederici1995; Sis-
neros 1991). Once mature, saltcedar is remarkablytol-
erant to mechanical injury caused by cutting, grazing,
and burning,as well as a variety of environmentalstress
conditions, including, heat, cold, drought, water inun-
dation, and high concentrations of dissolved solids
(Brotherson and Field 1987; Everitt 1980; Frasier and
Johnsen 1991).
After burning or cutting, saltcedar shrubs redevelop
rapidly (Horton 1977). Cutting has been shown to stim-
ulate growth. Goldsmith and Smart (1982) measured
stem growth rate at 75 cm/yr after cutting. This com-
pared with 30 cm/yr for an uncut bush growing in the
same area. In another study, Gary (1960) reportedthat
after cattle removed approximately50% of the saltcedar
foliage, the shrubsrecovered vigorously. By the second
year the stand became so dense that cattle would not
reenterthe area.
Under severe drought, saltcedaris capable of surviv-
ing by dropping its leaves and, thus, reducing evapo-
transpirationrates (Horton and Campbell 1974). As a
result, growth rates are greatly reduced. Despite this,
these populations are capable of withstanding lengthy
periods of drought.In the otherextreme situation,mature
plants are able to survive complete submergencein wa-
ter for as long as 70 d (Brothersonand Field 1987; Ker-
pez and Smith 1987; Warrenand Turner1975).
Reproduction. Seedlings mature rapidly and produce
small white or pinkish flowers, often by the end of the
first year of growth (Neill 1985). Flowers have four or
five sepals and petals, three to five styles, and stamens
borne on a fleshy, lobed, hypogynous disk. The fruit is
a three to five-valved capsule (Kerpez and Smith 1987).
Seeds are quite small and light (0.1 mg) (Sisneros 1991),
and have a tuft of hair on the end to aid in wind dispersal
or can be carried and deposited along sandbarsand riv-
erbanksby water (Brothersonand Field 1987). A single
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 WEED TECHNOLOGY
large saltcedarplant can producehalf a million seeds per
year.
Saltcedarcan bloom almost the entire growing season
from April to October at lower elevations (Everitt 1980)
but appears to have one major and one minor peak of
seed productionduring this period (Shrader 1977). The
ability of saltcedarto producean almost continualsupply
of seeds during the growing season allows it to colonize
areas after summerrains or summerrecession flow (En-
gel-Wilson and Ohmart 1978; Horton 1977). This gives
saltcedar a significant ecological advantage over other
woody riparian species because it can exploit suitable
germinatingconditions over a longer time interval when
seeds of the other species are not present (Howe and
Knopf 1991).
Germination. The tiny seeds of saltcedarhave high ini-
tial viability (Neill 1985). However, seeds remainviable
for only about 5 wks under normal conditions (Everitt
1980). Because of their short-lived viability, saltcedar
seeds must come in contact with suitablemoisturewithin
a few weeks of dispersal. Consequently,for germination
to occur following water dispersal, it is importantthat
the availability of seed coincide with the time of peak
annual discharge, so that seeds will settle and germinate
in a suitable location at highwatermarks (Everitt 1980).
Once wetted, fresh seeds usually germinatewithin 24
h (Kerpez and Smith 1987). Germinationcan occur un-
der a wide variety of conditions in association with
stream banks or receding lake margins (Johnson 1987)
and can even occur while seeds are still afloat. Ideally,
saltcedargerminationis favored in fine-grainedsaturated
sediment (Everitt 1980).
Seedling Establishment. Several factors contribute to
the low seedling survival of saltcedar.Seedlings, partic-
ularly roots, grow more slowly than many other riparian
species (Everitt 1980). For seedlings to establish suc-
cessfully, they requirea combinationof saturatedsoil for
the first 2 to 4 wks of life, open sunny ground, and the
absence of competition (Brotherson and Field 1987;
Brothersonand Winkel 1986; Everitt 1980). These con-
ditions are typically provided by a gently sloping river-
bank, sandbar,or silt bar, where slowly receding water
levels of river, stream,or reservoir create optimum seed-
beds (Shrader 1977). In the initial stages of establish-
ment, roots grow slowly within the first 4 wk and will
not survive more than 1 d if the soil dries (Kerpez and
Smith 1987). Seedling establishment also requires at
least 4 to 6 wks without subsequentinundation(Kerpez
and Smith 1987; Shrader 1977). Mortality is also high
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when soils are scoured by high water flow velocities.
Although seedlings can survive submerged for a few
weeks, they are easily uprootedby even a weak current
within a period of several months subsequentto germi-
nation (Kerpez and Smith 1987).
Occasionally saltcedarcan become establishedin typ-
ically dry locations if these areas experience an unusu-
ally wet spring or early summer, or if rivers or lakes
temporarilyflood their boundaries (Carmanand Broth-
erson 1982). Once established, saltcedarcan survive al-
most indefinitely in the absence of surface saturationof
the soil (Brothersonand Field 1987).
Root Growth. The root system of saltcedaris extensive
and is largely responsible for its competitiveness and
survival under stress. Initially, the primary root grows
steadily downward with little branching from lateral
roots until it reaches the water table (Horton 1977). Al-
though tap roots typically reach depths of about 5 m
(Brotherson and Field 1987; Brotherson and Winkel
1986), they have been reportedto extend as deep at 50
m (Baum 1978). Once the water table is reached, sec-
ondary root branchingbecomes profuse.
Plants can resprout vigorously from roots if the top
growth is damaged or removed (Frasier and Johnsen
1991). In addition, adventitious roots easily develop
from submergedor buried saltcedarstems (Everitt 1980;
Kerpez and Smith 1987). Thus, expansion in saltcedar-
infested areascan be, to some degree, throughvegetative
growth, as well as seed production.
ECOLOGY
Saltcedargrows at elevations no greaterthan 2,100 m
and prefers very saline soils. Typically, saltcedaroccu-
pies sites with silt loams and silt clay loams high in
organic matter,intermediatemoisture,high water tables,
and little erosion (Brotherson and Winkel 1986). They
can resproutvegetatively after fire, severe flood, or treat-
ment with herbicides and are able to accommodatewide
variations in soil and mineral gradients, as well as en-
vironmental stress conditions (Brotherson and Field
1987). Saltcedarhas a slight preferencefor alkaline con-
ditions (pH 7.5) but is also commonly found in more
acidic growing conditions (Brotherson and Winkel
1986).
In dense mature stands of saltcedar with little bare
ground exposed beneath the shrubcanopy, there is little
opportunity for regeneration of any species (Horton
1977). In these heavily infested areas, it would be un-
331
 DI TOMASO: SALTCEDARIN THE SOUTHWESTERNUS
likely that native riparianspecies could reestablishwith-
out some form of human intervention.
The shifts in riparian community structure and the
ability of saltcedarto outcompete native riparianspecies
is primarily due to distinct adaptationsassociated with
water usage, as well as tolerance and interactionswith
salinity (Busch and Smith 1995).
Water Use. Evapotranspirationrates of saltcedar are
among the highest of any phreatophyte evaluated in
southwestern North America (Brotherson et al. 1984;
van Hylckama 1974), including native riparian trees
(Busch and Smith 1995; Neill 1985). Its consumptionof
water can be so extensive that in heavily infested areas
saltcedarcan desiccate springs,drainpools, and even dry
up perennial streams (Johnson 1987). One large tree can
absorb 760 L of water a day. Saltcedarin all the heavily
infested areas of the southwest are estimatedto consume
almost twice as much water per year as the major cities
of southernCalifomia (Friederici 1995; Johnson 1987).
Numerous studies have been conducted on the total
water use in saltcedar stands (Davenport et al. 1982a,
1992b; Gay and Hartman1982; Kerpez and Smith 1987;
Weeks et al. 1987). The values obtained in these studies
show a wide range in total water usage and evapotrans-
piration rates. For example, Robinson (1965) reported
that saltcedar in Arizona used between 1.2 and 1.5 m3/
yr water, and Davenport et al. (1982a, 1982b) showed
yearly transpirationrates of 1.2 to 4.0 m3. In a Bureau
of Reclamation (1992) study, water uptake estimates for
saltcedar varied from 0.4 to 3.2 m3/yr.Davenport et al.
(1982a) reported saltcedar transpiredfrom 1.2 to 4 m3/
yr and from 5 to 10 mm/d. Estimates of about 1.8 m3/
yr evapotranspirationwas noted for a dense saltcedar
infestation along the lower Colorado River (Gay 1985).
When saltcedar vegetation was thinned to 50% of the
original density, the reductionin evapotranspirationwas
only 10% (van Hylckama 1974). Along the Colorado
River, water loss up to 433,000 m3/hais lost per year to
channel vegetation, of which saltcedar is a major com-
ponent (Brothersonand Field 1987).
Many studies have also compared the water use of
saltcedarwith other riparianor agriculturalspecies. Ball
et al. (1994) calculated that the evapotranspirationrate
of monotypic stands of saltcedar adjacent to the lower
Colorado River near Blythe, CA, was between 0.7 and
0.8 m3/yr.By comparison,the evapotranspirationrate of
honey mesquite and quailbush or big saltbush [Atriplex
lentiformis (Torr.)S. Watson] was estimated to be 0.5
and 0.7 m3/yr,respectively. The midsummerevapotrans-
piration rates from saltcedarcan be as high as those re-
332
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ported for irrigatedcropland (Gay and Hartman 1982).
Wateruse of saltcedaralong the Colorado River was in
general agreement with water use by irrigated alfalfa
(Medicago satira L.). At moderate density, evapotrans-
piration was about the same as that for grass, but at
heavy density, it was about 2.4 times that of a grassland
ecosystem (Davenportet al. 1982a).
Evapotranspirationrates also vary considerably de-
pending on the time of day or year. Gay and Hartman
(1982) reported daily evapotranspirationtotals ranging
from 2.9 mm/d in early April, up to 11 mm/d in late
June, and again down to 1.8 mm/d in late October.Sim-
ilar results were also reportedby Gay (1985). In a study
conducted by Williams and Anderson (1977), transpira-
tion rates of saltcedar were high until about noon and
then began a steady, gradual decrease that continued
throughoutthe afternoon.
Although total water usage was considerably higher
in saltcedar stands than in areas with native woody ri-
parian species, transpirationrates of individual plants
have been shown to be similar to those of several her-
baceous plants and cooccurringphreatophytes,including
willows, cottonwoods, and mesquite (Anderson 1982;
Sala and Smith 1996). The discrepancy between total
and individual evapotranspiration rates in saltcedar
stands and native riparianhabitatwas attributedto great-
er leaf area index in saltcedarcommunities comparedto
any other riparianpopulations (Sala and Smith 1996).
Thus, the total transpiringarea was the primary reason
for loss of high quantitiesof water in saltcedarinfested
areas. It has also been suggested that salt glands on the
leaf surfaces of saltcedarmay contributemore to water
loss than stomates by acting as a significant "pulling
force" that furtherenhances continued soil water extrac-
tion (Sala and Smith 1996). This is supportedby evi-
dence showing maximum evapotranspirationby saltce-
dar in the early morning when salt gland exudate was
most visible (Wiesenborn1996).
Under dry or extremely hot conditions, saltcedardoes
not always transpireat potential rates (Davenportet al.
1982a). Water conservation under these situations is of
ecological significance because it enables Tamarixspe-
cies that grow in hot desert environmentsto open their
stomata just at daybreak during the coolest and most
humid hours of the day. This allows plants to acquire
adequate CO2 without losing much water. The stomata
close during the hotter afternoonhours, furtherreducing
water loss (Hagemeyer and Waisel 1990). Summer
evapotranspirationrates can also vary considerablywith
stand density and other stress conditions (Davenport et
Volume 12, Issue 2 (April-June) 1998
 WEED TECHNOLOGY
Table 2. Characteristicsof floodplain zones at varying groundwaterdepths (from Horton and Campbell 1974).
Depth to
groundwater
Zone m Saltcedargrowth
1 dwarfed and multi-
stemmed
2 1.5-2.5 major stands
3 2.5-6 major stands
4 > 6 scatteredindividuals
al. 1982a, 1982b). In addition to stomatal regulation,
saltcedar also has a greater ability to tolerate drought
stress through osmotic adjustment compared to native
species (Busch and Smith 1995).
As a facultative phreatophyte, saltcedar may draw
moisture from the saturatedzones below the water table
and is also capable of extracting soil moisture from less
saturatedsoils in areas with deeper water tables (Everitt
1980). Native woody taxa are obligate phreatophytesand
obtain much of their water from the saturated zone
(Busch et al. 1992; Horton and Campbell 1974). This
may partially explain the competitive exclusion of these
native shrubsby saltcedarin southwesternriparianareas.
Thus, the longer a community has been invaded by salt-
cedar, the greaterwill be the capacity to lower the water
table (Brothersonet al. 1984). With this overall drying
out of the habitat, more xeric plant species will occupy
the understoryin established saltcedarstands (Table 2).
If the water table is shallow, saltcedar does not de-
velop densely and the intershrubspaces are usually dom-
inated by saltgrass [Distichlis spicata (L.) Greene #
DISSP], bermudagrass[Cynodon dactylon (L.) Pers. #
CYNDA], or alkali sakaton (Sporobolusairoides Torr.#
SPZAI). Dropping water tables in some areas will allow
saltcedar to outcompete other coexisting grass species
(Horton 1977). Dense stands of saltcedartypically occur
where water tables are between 1.5 and 6 m from the
surface (Horton and Campbell 1974; Kerpez and Smith
1987; Sala and Smith 1996; Table 2).
Restoration of saltcedar-infestedareas has the poten-
tial to dramatically increase moisture availability.
Hughes (1970) estimated that removal of 16,000 ha of
saltcedarfrom Las Vegas to Carlsbad,NM, could prob-
ably yield between 46,000 and 53,000 m/ha of additional
water each year. Weeks et al. (1987) reportedthat annual
water use by saltcedar in the Pecos River is probably
about 0.3 m3 greaterthan that of replacementvegetation.
With control of saltcedar,they predicteda net water gain
of 3,000 m3/hain the infested area.
Volume 12, Issue 2 (April-June) 1998
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Water salvage
Other vegetation prospects Other uses
vigorous saltgrass and bermu- little good grazing, flood pas-
dagrass sage, minimal wildlife
use
excellent saltgrass large savings wildlife utilization
(doves), some grazing,
bees
xeric types great water savings wildlife utilization
(doves), bees
xeric types none expected limited use
Salinity. Saltcedar is a facultative halophyte that is ca-
pable of toleratingsoluble salt concentrationsin the soils
ranging from 650 to 36,000 ppm and averagingbetween
6,000 and 8,000 ppm (Brothersonand Winkel 1986; Car-
man and Brotherson 1982; Gatewood et al. 1950; Jack-
son et al. 1990). Jacksonet al. (1990) reportedinhibition
in cottonwood and willow growth by salinity greater
than 1,500 ppm. Anderson (1996) also reportedreduced
stem growth in willows, cottonwoods, and mesquite
growing undersaline conditions. Consequently,saltcedar
species have a distinct advantage over other native
woody riparian species in environments with high salt
concentrations(Shafrothet al. 1995).
The ability of saltcedar to tolerate high soluble salt
concentrationsin the soil partially contributes to their
rapidrecovery and competitive advantagefollowing fire.
Ash deposits subsequentto fire are high in salinity and
have elevated concentrationsof phytotoxic boron. Cot-
tonwood and willow seedlings are not only sensitive to
high saline conditions, but also exhibit low tolerance to
boron (Busch and Smith 1993).
The salt glands of Tamarixspecies are on the adaxial
and abaxial leaf surfaces as well as young stem surfaces.
They consist of eight radially arranged,epidermal cells
(Bosabalidis and Thomson 1984, 1985). Six outer cells
are densely cytoplasmic secretory cells and two inner
cells are highly vacuolated and often called collecting
cells. These collecting cells accumulatehigh concentra-
tions of salts prior to secretion. Studies conducted with
T. aphylla (Bosabalidis and Thomson 1984, 1985) noted
that, unlike typical glandulartrichomes of other species,
the salt glands of saltcedar are sunken slightly in the
epidermis and only occasionally protrudeabove the epi-
dermal surface.
Salt glands of T. aphylla, and presumablyother spe-
cies of Tamarix,are not permanentand continue to ini-
tiate and differentiatethroughoutleaf expansion and sub-
sequent maturation(Bosabalidis 1992). In contrast, the
leaf glands of most species are not replaced by new
333
 DI TOMASO: SALTCEDARIN THE SOUTHWESTERNUS
glands once they have senesced. This differenceprovides
saltcedar species with a very efficient ecological adap-
tation to high salinity conditions (Bosabalidis 1992).
The movement of salts from the soil to the foliar
glands has been demonstratedto be by apoplastictrans-
port in the xylem (Campbell et al. 1974; Campbell and
Thomson 1975). The composition of the secreted salts
has been shown by several investigatorsto be nonselec-
tive and dependent on the salt composition of the root
environment (Berry 1970; Story and Thomson 1994;
Thomson et al. 1969). Numeroussalts and minerals,both
macro- and micronutrients, are excreted by saltcedar
glands (Berry 1970; Bosabalidis and Thomson 1984;
Dreesen and Wangen 1981; Kleinkopf and Wallace 1974;
Story and Thomson 1994; Thomson et al. 1969), includ-
ing sodium, potassium,nitrate,calcium, magnesium,sul-
fur, sulfate, phosphorus, bicarbonate, chloride, molyb-
denum, boron, copper,manganese, aluminum,silica, and
zinc. Even trace elements, such as lithium, barium, ti-
tanium, and strontiumwere found in the glands of salt-
cedar. Many of these salts were found in the glands at
concentrations50-fold higher than the external solution,
even with plants grown hydroponicallyunder laboratory
conditions (Berry 1970). Once excreted, these salts form
small white residues on the epidermal surface (Bosabal-
idis and Thomson 1984). The broad range of excreted
salts clearly indicate that the glands of saltcedarhave a
low level of selectivity for salts. A high tolerance for
such a variety of different ions has undoubtedlyallowed
saltcedar species to adapt to many different saline soil
types.
Increased external salt concentrations have demon-
stratedsome fitness cost on saltcedargrowth. Kleinkopf
and Wallace (1974) reportedthat a reduction in growth
rate with increasing salt treatmentdid not correlatewith
a concomitant change in the rate of photosyntheticcar-
bon fixation and transpirationalwater loss. They con-
cluded that the observed growth decreases under in-
creased saline conditions were due to energy losses re-
quired to transportsalt and maintainnormal water rela-
tions.
Salt gland exudates have been reported to contain
41,000 ppm dissolved solids in the guttation sap (Gate-
wood et al. 1950). These salts are eventually deposited
on the soil surface under the plant, sometimes forming
a hard crust (Kerpez and Smith 1987). Consequently,
salts are redistributedover time from deep within the
soil profile to become concentratedon the soil surface
of floodplains (Shafroth et al. 1995). Excessive surface
deposits of salt can inhibit the germinationof other spe-
334
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cies (Egan et al. 1993), thus restrictingcompetitionwith
other under-or overstory vegetation for space and water
(Brothersonand Field 1987). In some communities,salt-
cedar is the dominant overstory species, whereas salt-
tolerant grasses, such as saltgrass, dominate the under-
story (Brothersonand Winkel 1986).
LITERATURE
CITED
Anderson,B. W. 1996. Salt cedar,revegetationand riparianecosystems in the
southwest.In J. E. Lovich, J. Randall,and M. Kelly, eds. Proc. California
Exotic Pest Plant Council Symposium 1995. pp. 32-41.
Anderson,B. W., A. Higgins, and R. D. Ohmart.1977. Avian Use of Saltcedar
Communities in the Lower Colorado River Valley. Fort Collins, CO:
USDA, US Rocky Mt. For. Range Exp. Stn. Gen. Tech. Rep. 43:128-
136.
Anderson, B. W. and R. D. Ohmart. 1985. Riparianrevegetation as a miti-
gating process in stream and river restoration.In J. A. Gore, ed. The
Restoration of Rivers and Streams: Theories and Experience. Boston,
MA: Butterworth.pp. 41-80.
Anderson, J. E. 1982. Factors controlling transpirationand photosynthesisin
Tamarixchinensis Lour. Ecology 63:48-56.
Ball, J. T., J. B. Picone, and P. D. Ross. 1994. Evapotranspirationby Riparian
Vegetation along the Lower Colorado River. Boulder City, NV: Bureau
of Reclamation, Lower Colorado Region, Biol. Sci., Cent., Desert Res.
Inst., Reno, NV. Final Rep. I-CP-30-08910.
Baum, B. R. 1967. Introducedand naturalizedtamarisksin the United States
and Canada (Tamaricaceae).Baileya 15:19-25.
Baum, B. R. 1978. The Genus Tamarix.Jerusalem:Israel Academy of Sci-
ences and Humanities.209 p.
Berry, W. L. 1970. Characteristicsof salts secretedby Tamarixaphylla. Amer.
J. Bot. 57:1226-1230.
Blackburn,W. H., R. W. Knight, and J. L. Schuster. 1982. Saltcedarinfluence
on sedimentation in the Brazos River. J. Soil Water Conserv. 37:298-
301.
Bosabalidis, A. M. 1992. A morphological approachto the question of salt
gland lifetime in leaves of Tamarixaphylla L. Israel J. Bot. 41:115-121.
Bosabalidis, A. M. and W. W. Thomson. 1984. Light microscopical studies
on salt gland development in Tamarixaphylla L. Ann. Bot. 54:169-174.
Bosabalidis, A. M. and W. W. Thomson. 1985. Ultrastructuraldevelopment
and secretion in the salt glands of Tamarixaphylla L. J. Ultrastructure
Res. 92:55-62.
Brotherson,J. D., J. G. Carman,and L. A. Szyska. 1984. Stem-diameterage
relationshipsof Tamarixramosissimain centralUtah. J. Range Manage.
37:362-364.
Brotherson,J. D. and D. Field. 1987. Tamarix:impacts of a successful weed.
Rangelands9:110-112.
Brotherson,J. D. and V. Winkel. 1986. Habitatrelationshipsof saltcedar(Ta-
marix ramosissima) in central Utah. Great Basin Nat. 46:535-541.
Bureau of Reclamation. 1992. Vegetation Management Study, Lower Colo-
rado River.BoulderCity, NV: Lower ColoradoRegion Phase I Rep. 103 p.
Busch, D. E. 1995. Effects of fire on southwesternriparianplant community
structure.Southwest. Nat. 40:259-267.
Busch, D. E., N. L. Ingraham,and S. D. Smith. 1992. Wateruptake in woody
riparianphreatophytesof the southwesternUnited States: a stable isotope
study. Ecol. Appl. 2:450-459.
Busch, D. E. and S. D. Smith. 1992. Fire in a RiparianShrub Community:
Postburn Water Relations in the Tamarix-Salix Association along the
Lower Colorado River. Ogden, UT: USDA, For. Serv. Intermt.Res. Stn.
Gen. Tech. Rep. 289. pp. 52-59.
Busch, D. E. and S. D. Smith. 1993. Effects of fire on water and salinity
relations of riparianwoody taxa. Oecologia 94:186-194.
Busch, D. E. and S. D. Smith. 1995. Mechanisms associated with decline of
woody species in riparianecosystems of the southwestern U.S. Ecol.
Monogr. 65:347-370.
Campbell, N. and W. W. Thomson. 1975. Chloride localization in the leaf of
Tamarix.Protoplasm83:1-14.
Campbell, N., W. W. Thomson, and K. Platt. 1974. The apoplastic pathway
of transportto salt glands. J. Exp. Bot. 25:61-69.
Volume 12, Issue 2 (April-June) 1998
 WEED TECHNOLOGY
Carman,J. G. and J. D. Brotherson. 1982. Comparisonsof sites infested and
not infested with saltcedar (Tamarix pentandra) and Russian olive
(Elaeagnus angustifolia). Weed Sci. 30:360-364.
Carothers,S. W., S. E. Aitchison, M. M. Karpiscak,G. A. Ruffner,and J. J.
Sharber.1976. An ecological survey of the riparianzone of the Colorado
River between Lee's Ferry and Grand Wash Cliffs, Arizona. U.S. Na-
tional Park Service, Colorado River Res. Serv. Tech. Rep. 10. 251 p.
Cohan, D. R., B. W. Anderson, and R. D. Ohmart. 1978. Avian population
responses to saltcedaralong the lower Colorado River. In R. R. Johnson
and J. F McCormick, eds. Strategies for Protectionand Managementof
Floodplain Wetlands and Other RiparianEcosystems. Washington,DC:
U.S. Forest Service Gen. Tech. Rep. WO-12. pp. 371-381.
Crins, W. J. 1989. The Tamaricaceaein the southwestern United States. J.
Arnold Arboretum70:403-425.
Davenport, D. C., P E. Martin, and R. M. Hagan. 1982a. Evapotranspiration
from riparianvegetation:water relationsand irrecoverablelosses for salt-
cedar. J. Soil WaterConserv. 37:233-236.
Davenport, D. C., P. E. Martin,and R. M. Hagan. 1982b. Evapotranspiration
from riparianvegetation: conserving water by reducing saltcedar tran-
spiration.J. Soil WaterConserv. 37:237-239.
deGouvenain, R. C. 1996. Origin, history and currentrange of salt cedar in
the U.S. In Proc. SaltcedarManagementWorkshop.Rancho Mirage, CA.
CaliforniaExotic Pest Plant Council. pp. 1-3.
DeLoach, C. J. 1989. Prospects for biological control of saltcedar (Tamarix
spp.) in riparian habitats of the southwestern United States. In E. S.
Delfosse, ed. Proc. VII InternationalSymposium of the Biological Con-
trol of Weeds, 1988, Rome, Italy. 1st Sper. Patol. Veg. (MAF). pp. 307-
314.
DeLoach, C. J., M. J. Pitcairn, and D. Woods. 1996. Biological control of
saltcedar in southern California. In Proc. SaltcedarManagementWork-
shop. Rancho Mirage, CA. California Exotic Pest Plants Council. pp.
30-31.
DiTomaso, J. M. 1996. Identification,biology and ecology of saltcedar.In
Proc. SaltcedarManagementWorkshop.Rancho Mirage, CA. pp. 4-8.
Dressen, D. R. and L. E. Wangen. 1981. Elemental composition of saltcedar
(Tamarixchinensis) impacted by effluents from a coal-fired power plant.
J. Environ. Qual. 10:410-416.
Egan, T B., R. A. Chavez, and B. R. West. 1993. Afton Canyon saltcedar
removal first year status report.In L. Smith and J. Stephenson,eds. Proc.
Symposium of Vegetation Management,Hot Desert RangelandEcosys.,
Phoenix, AZ. p. 18.
Ellis, L. M. 1995. Bird use of saltcedar and cottonwood vegetation in the
Middle Rio GrandeValley of New Mexico, U.S.A. J. Arid Environ. 30:
339-349.
Engel-Wilson, R. W. and R. D. Ohmart. 1978. Floral and attendantfaunal
changes on the lower Rio Grande between Fort Quitman and Presidio,
Texas. In Proc. National Symposium Strategies for Protectionand Man-
agement of Floodplain Wetlands and Other Riparian Ecosystems. pp.
139-147.
Everitt, B. L. 1980. Ecology of saltcedar-a plea for research.Environ.Geol.
3:77-84.
Frasier,G. W. and T N. Johnsen, Jr. 1991. Saltcedar(tamarisk):classification,
distribution,ecology, and control. In L. F James, J. 0. Evans, M. H.
Ralphs, and R. D. Child, eds. Noxious Range Weeds. Boulder,CO: West-
view Press. pp. 377-386.
Friederici, P 1995. The alien saltcedar.Am. For. 101:45-47.
Gary, H. L. 1960. Utilization of Five-StamenTamariskby Cattle.FortCollins,
CO: U.S. Department of Agriculture Forest Service, Rocky Mt. For.
Range Exp. Stn. Res. Note 51. 4 p.
Gatewood, J. S., T. W. Robinson, R. B. Colby, J. D. Hem, and L. C. Halpenny.
1950. Use of Waterby BottomlandVegetation in Lower Safford Valley,
Arizona. U.S. Geol. Survey WaterSupply Paper 1103.
Gay, L. W. 1985. Evapotranspirationfrom saltcedaralong the lower Colorado
River. First North. Am. In Conf. Proc., RiparianEcosystems and their
Management: Reconciling Conflicting Uses. Tucson, AZ. Fort Collins,
CO: U.S. Forest Service. pp. 171-174.
Gay, L. W. and L. J. Fritschen. 1979. An energy budget analysis of water use
by saltcedar.WaterResour. Res. 15:1589-1592.
Gay, L. W. and R. K. Hartman.1982. ET measurementsover ripariansaltcedar
on the Colorado River. Hydrol. WaterResour. Arizona Southwest 12:9-
15.
Goldsmith, F. B. and N. Smart. 1982. Age, spacing and growthrateof Tamarix
Volume 12, Issue 2 (April-June) 1998
This content downloaded from 170.140.26.180 on Wed, 21 Aug 2013 18:35:31 PM
All use subject to JSTOR Terms and Conditions
as an indicationof lake boundaryfluctuationsat SebkhetKelbia, Tunisia.
J. Arid Environ. 5:43-51.
Graf, W. L. 1978. Fluvial adjustmentsto the spread of tamariskin the Col-
orado Plateau region. Bull. Geol. Soc. Am. 89:1491-1501.
Great Western Research, Inc. 1989. Economic analysis of harmful and ben-
eficial aspects of saltcedar.In Final Report. Boulder City, NV: Bureau
of Reclamation,Lower Colorado Region. p. 259.
Hagemeyer,J. and Y Waisel. 1990. Phase-shift and memorizationof the cir-
cadian rhythm of transpirationof Tamarixaphylla. Experientia46:876-
877.
Haigh, S. L. 1996. Saltcedar(Tamarix ramosissima), an uncommon host for
desert mistletoe (Phoradendroncalifornicum).GreatBasin Nat. 56:186-
187.
Horton, J. S. 1964. Notes on the Introductionof Deciduous Tamarix. Fort
Collins, CO: U.S. For. Serv. Res. Note R-16. 7 p.
Horton,J. S. 1977. The Development and Perpetuationof the PermanentTam-
arisk Type in the PhreatophyteZone of the Southwest. Fort Collins, CO:
USDA, U.S. Rocky Mt. For. Range Exp. Stn. Gen. Tech. Rep. 43.
Horton, J. S. and C. J. Campbell. 1974. Management of Phreatophyteand
Riparian Vegetation for Maximum Multiple Use Values. Fort Collins,
CO: U.S. Forest Service, Res. Pap. RM-117. 23 p.
Howe, W. H. and F L. Knopf. 1991. On the imminentdecline of Rio Grande
cottonwoods in central New Mexico. The Southwest. Nat. 36:218-224.
Hughes, L. E. 1993. "The devil's own"-tamarisk. Rangelands 15:151-155.
Hughes, W C. 1970. Economic Feasibility of Increasing Pecos Basin Water
Supplies throughReduction of Evaporationand Evapotranspiration. Wa-
ter Resources Research InstituteRep. 9.
Hunter, W C., B. W. Anderson, and R. D. Ohmart. 1985. Summer avian
communitycomposition of Tamarixhabitatsin three southwesterndesert
riparian systems. In Conf. Proc., Riparian Ecosystems and their Man-
agement: Reconciling Conflicting Uses, Tucson, AZ. Fort Collins, CO:
U.S. Dept. Agr., Forest Service. pp. 128-134.
Hunter,W. C., R. D. Ohmart,and B. W. Anderson. 1988. Use of exotic salt-
cedar (Tamarixchinensis) by birds in arid ripariansystems. Condor90:
113-123.
Jackson, J., J. T Ball, and M. R. Rose. 1990. Assessment of the salinity
tolerance of eight Sonoran Desert riparian trees and shrubs. In Final
Report. Yuma, AR: Bureauof Reclamation,YumaProjectOffice. p. 102.
Johnson, S. 1987. Can tamariskbe controlled?Fremontia 15(2):19-20.
Kerpez, T. A. and N. S. Smith. 1987. SaltcedarControl for Wildlife Habitat
Improvementin the SouthwesternUnited States. Washington,DC: USDI.
Fish and Wildlife Serv. Resource Publ. 169. 16 p.
Kleinkopf, G. E. and A. Wallace. 1974. Physiological basis for salt tolerance
in Tamarixramosissima. Plant Sci. Lett. 3:157-163.
Lovich, J. E., T B. Egan, R. C. de Gouvenain. 1994. Tamariskcontrol on
public lands in the desert of SouthernCalifornia:two case studies. Proc.
CaliforniaWeed Conf. 46:166-177.
Neill, W. M. 1985. Tamarisk.Fremontia 12:22-23.
Robinson, T. W. 1958. Phreatophytes.Washington, DC: U.S. Geol. Survey
WaterSupply Paper. 1423. 85 p.
Robinson, T. W 1965. Introduction,Spread, and Aerial Extent of Saltcedar
(Tamarix) in the Western States. Washington, DC: U.S. Geol. Survey
Prof. Paper 491-A. 12 p.
Sala, A. and S. D. Smith. 1996. Water use by Tamarix ramosissima and
associated phreatophytesin a Mojave Desert floodplain. Ecol. Appl. 6:
888-898.
Shafroth,P. B., J. M. Friedman,and L. S. Ischinger. 1995. Effects of salinity
on establishment of Populus fremontii (cottonwood) and Tamarix ra-
mosissima (saltcedar) in southwestern United States. Great Basin Nat.
55:58-65.
Shrader,T H. 1977. Selective management of phreatophytesfor improved
utilization of natural food-plain resources. Water management for irri-
gation and drainage.Proc. Soc. Civil Eng. 2:16-44.
Sisneros, D. 1991. Herbicide Analysis: Lower Colorado River SaltcedarVeg-
etation Management Study. Denver, CO: Bureau of Reclamation, U.S.
Dept. Int. R-91-06. 167 p.
Story, R. and W. W. Thomson. 1994. An x-ray microanalysisstudy of the salt
glands and intracellularcalcium crystals of Tamarix.Ann. Bot. 73:307-
313.
Swenson, J. E., P. Hendricks,and A. Farjon. 1982. Arrival and occurrenceof
Tamarix chinensis (tamarisk) along the Yellowstone River in Treasure
and Rosebud Counties, Montana. Proc. Mont. Acad. Sci. 41:67-70.
Thomson, W. W., W. L. Berry, and L. L. Liu. 1969. Localizationand secretion
335
 DI TOMASO: SALTCEDARIN THE SOUTHWESTERNUS

of salt by the salt glands of Tamarixaphylla. Proc. Natl. Acad. Sci. USA
63:310-317.
van Hylckama,TE.A. 1974. WaterUse by Saltcedaras Measuredby the Water
Budget Method. Washington,DC: USGS ProfessionalPaper491-E.
Warren,D. K. and R. M. Turner.1975. Saltcedarseed production,seedlingestab-
lishment,and responseto inundation.ArizonaAcad. Sci. J. 10:131-144.
Weeks, E. P., H. L. Weaver,G. S. Campbell, and B. D. Tanner.1987. Water
Use by Saltcedar and by Replacement Vegetation in the Pecos River
Floodplain between Acme and Artesia, New Mexico. Washington,DC:
U.S. Geol. Survey Prof. Pap., 491-G. 33 p.
Wiesenborn,W. D. 1996. Saltcedarimpacts on salinity, water,fire frequency,
and flooding. In Proc. SaltcedarManagementWorkshop.RanchoMirage,
CA, 1996. CaliforniaExotic Pest Plant Council. pp. 9-12.
Williams, M. E. and J. E. Anderson. 1977. Diurnal trends in water status,
transpiration,and photosynthesisof saltcedar.Hydrol. WaterResour.Ar-
izona Southwest 12:119-124.

Herbicide Handbook-7th Edition, 1994

Editedby W.H. Ahrens

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