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SPECIES: Atriplex lentiformis

• Introductory
• Distribution and occurrence
• Botanical and ecological characteristics
• Fire ecology
• Fire effects
• Management considerations
• References

INTRODUCTORY

AUTHORSHIP AND CITATION FEIS ABBREVIATION SYNONYMS NRCS PLANT CODE COMMON NAMES TAXONOMY LIFE FORM FEDERAL LEGAL STATUS OTHER STATUS
	Michael Charters, Southern California Wildflowers

Big saltbush may not regularly hybridize, even though it occurs with several Atriplex species [46]. However, Hanson [46] reports hybrids of quailbush and beach saltbush (A. leucophylla) and quailbush and Davidson's bractscale (A. serenana var. davidsonii) in the collections of the California Academy of Sciences.

DISTRIBUTION AND OCCURRENCE

SPECIES: Atriplex lentiformis

GENERAL DISTRIBUTION ECOSYSTEMS STATES/PROVINCES BLM PHYSIOGRAPHIC REGIONS KUCHLER PLANT ASSOCIATIONS SAF COVER TYPES SRM (RANGELAND) COVER TYPES HABITAT TYPES AND PLANT COMMUNITIES
	Quailbush. Tony Baker, Natural Landscapes

Riparian zones in which big saltbush occurs include woodlands such as cottonwood-willow (Populus-Salix spp.) and mesquite (Prosopis spp.) cover types, as well as riparian scrub. Big saltbush also grows on the margins of oases and wetlands. In cottonwood-willow woodlands big saltbush typically occurs with low frequency [16,49,93]. According to vegetation classifications based on literature and expert opinion [49,93], more common riparian species such as mule's fat (Baccharis salicifolia), common reed (Phragmites australis), sandbar willow (S. exigua), and Fremont cottonwood (Populus fremontii) occur with big saltbush in riparian areas across big saltbush's range. Goodding willow (S. gooddingii) is another component of these communities, although it is less common in southern California coastal areas and absent from California's central coast [93,109]. Big saltbush is an uncommon species in the riparian areas of California's Central Valley and central and southern coastal regions, which typically contain California sycamore (Platanus racemosa), red willow (S. laevigata), Pacific willow (S. lucida ssp. lasiandra), arroyo willow (S. lasiolepis), Douglas' mugwort (Artemisia douglasiana), and common elderberry (Sambucus nigra ssp. canadensis) [93]. On the Mojave River big saltbush may occur with velvet ash (Fraxinus velutina), white alder (Alnus rhombifolia), and riparian species common throughout its range [93]. Giant reed (Arundo donax) and tamarisks (Tamarix spp.) also co-occur with big saltbush in riparian zones throughout much of big saltbush's range [49,93].

Riparian scrub forms thickets along rivers and streams of the Southwest, which are typically dominated by tamarisks and arrowweed (Pluchea sericea) [15,17,63]. In addition to these dominants, some communities may contain scattered Fremont cottonwood and willows, as well as big saltbush, screwbean mesquite (Prosopis pubescens), honey mesquite (P. glandulosa), saltgrass, Palmer's coldenia (Tiquilia palmeri), mule's fat, and/or spiny chloracantha (Chloracantha spinosa) [15,16,17,49,63,71,106]. Mojave seablite, cattle saltbush, and iodinebush (Allenrolfea occidentalis) can also be components of this vegetation type [71].

Further from the river channel is the mesquite bosque forest. Big saltbush is a typical species of the mesquite bosque understory [16]. Presettlement mesquite bosque forests had relatively open understories with grasses, forbs and saltbushes, such as big saltbush and cattle saltbush (Atriplex polycarpa) forming the ground cover [14,49,71]. These communities can contain honey mesquite, velvet mesquite (P. velutina), and/or screwbean mesquite as well as Mojave seablite (Suaeda moquinii), viney milkweeds (Sarcostemma spp.), gourds (Cucurbita spp.), netleaf hackberry (Celtis reticulata), common elderberry, blue paloverde (Parkinsonia floridum), wolfberries (Lycium spp.), and saltbushes including big saltbush, cattle saltbush, and fourwing saltbush (A. canescens) [49,71,93].

Big saltbush occurs on the margins of oases, wetlands, and in alkali sink communities. Vogl and McHargue [114] report big saltbush as a rare species within the oases along the San Andreas Fault. Typical species within southern Californian oases include California palm (Washingtonia filifera), blue paloverde, mule's fat, desert willow (Chilopsis linearis), common reed, arrowweed, mesquite (Prosopis spp.), screwbean mesquite, sandbar willow, Goodding willow, and desert wild grape (Vitis girdiana). [93,114]. Along the margins of Sonoran wetlands, species from surrounding scrublands including tamarisk (Tamarix spp.), arrowweed, mesquite, and big saltbush mix with species more typical of wetlands, such as common cattail (Typha latifolia), saltgrass (Distichlis spicata), common threesquare (Schoenoplectus pungens), and common reed [71]. Big saltbush may be one of several saltbush species found in alkali sinks of the desert Southwest along with iodinebush and Mojave seablite [15]. According to a literature review by Thorne [106], several other Chenopods occur in this community including goosefoot species (Chenopodium spp.), pickleweeds (Salicornia spp.), seepweeds (Suaeda spp.), and black greasewood (Sarcobatus vermiculatus). Other species found in alkali sinks include fewleaf spiderflower (Cleome sparsifolia), stinkweed (Cleomella spp), spreading alkaliweed (Cressa truxillensis), alkali pepperweed (Lepidium dictyotum), spiny caper (Oxystylis lutea), and spectacle fruit (Wislizenia refracta) [106].

Big saltbush also inhabits comparatively dry habitats such as grasslands and desert scrub, including saltbush vegetation. Other saltbushes are the dominant species in this type, with big saltbush a relatively common associate [43,63,85,106,108,112]. Cattle saltbush and thinleaf fourwing saltbush (A. canescens var. linearis) are frequently dominants of this type [63,107]. Several saltbushes are associated with this community, including big saltbush, silverscale saltbush (A. argentea), fourwing saltbush, shadscale (A. confertifolia), wheelscale saltbush (A. elegans var. fasciculata), Nuttall's saltbush (A. nuttallii), Parry's saltbush (A. parryi), leafcover saltweed (A. phyllostegia), smooth saltbush (A. pusilla), and Torrey's saltbush (A. torreyi) [49]. In addition, fivehorn smotherweed (Bassia hyssopifolia), iodinebush, boraxweed (Nitrophila occidentalis), Parish's pickleweed (Arthrocnemum subterminale), seepweeds, and saltgrass can occur in this community [107]. Other species in this type are shrubby alkaliaster (Machaeranthera carnosa var. intricata), spiny hopsage (Grayia spinosa), white burrobrush (Hymenoclea salsola), rusty molly (Kochia californica), Anderson wolfberry (Lycium andersonii), peach thorn (L. cooperi), western honey mesquite (Prosopis glandulosa var. torreyana), and Pursh seepweed (Suaeda calceoliformis) [49]. Vasek and Barbour [112] report big saltbush as an important species in the salt scrub vegetation of Rabbit Dry Lake in California along with green molly (Kochia americana), Mojave seablite, and shadscale saltbush. Gullion [43] mapped the distribution of a Las Vegas Valley saltbush type that was comprised of saltbushes including big saltbush and Mojave seablite, saltgrass, and globemallow (Sphaeralcea spp.).

Big saltbush also occurs in other desert scrub types. Although rare in the oases studied by Vogl and McHargue [114], big saltbush was one of many species, including catclaw acacia (Acacia greggii) and Schott's pygmycedar (Peucephyllum schottii), that were part of the surrounding desert community. Saltbushes can occur in mixed woody scrub, Joshua tree (Yucca brevifolia) woodland, and creosote bush (Larrea tridentata) scrub [49,105,106]. Confirmation of big saltbush in these communities is not provided by the literature. Big saltbush has been reported as a codominant with coyote bush (Baccharis pilularis) on upland sites [83], and as the dominant species in an area with fourwing saltbush [86].

There are few reports of big saltbush occurring in grasslands. However, Munz [74] includes valley grassland in a list of communities that contain quailbush. In addition, Heady [47] notes the need for more research of the saltbush variations of valley grasslands. He notes the occurrence of cattle saltbush on foothills and other drained areas that border sinks. Other alkali-tolerant associated species include iodinebush, rusty molly, Parish's pickleweed, seablite and grasses including saltgrass, dwarf barley (Hordeum depressum), and alkali sacaton (Sporobolus airoides). It is likely that big saltbush is an uncommon associate in these or similar areas.

Big saltbush, typically quailbush, occurs in coastal sage scrub [4,10,25]. In coastal regions near the mouth of the Ventura River, big saltbush and coyote bush occur amongst other scattered dune vegetation dominated by sand verbena (Abronia spp.) and silver burr ragweed (Ambrosia chamissonis). Other species characteristic of this vegetation type are beach suncup (Camissonia cheiranthifolia ssp. suffruticosa), California croton (Croton californicus), saltgrass, seaside buckwheat (Eriogonum latifolium), and chamisso bush lupine (Lupinus chamissonis) [26]. Pearcy and Harrison [86] studied big saltbush in a coastal area where it occurred with saline saltbush (Atriplex subspicata), California seablite (Suaeda californica), California sagebrush (Artemisia californica), and saltgrass.

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

GENERAL BOTANICAL CHARACTERISTICS RAUNKIAER LIFE FORM REGENERATION PROCESSES SITE CHARACTERISTICS SUCCESSIONAL STATUS SEASONAL DEVELOPMENT
	Quailbush. Ken Gilliland, Thomas Payne Foundation

Big saltbush is a large, perennial, native shrub. It typically grows to between 3.3 and 8.2 feet (1-2.5 m) tall, but can reach 9.8 feet (3 m) [8,57,79,90,119]. Plants are wide spreading [25,113]. Individuals approximately 6.6 feet (2 m) tall were reported to cover areas ranging from 5.6 m² to 7.8 m² [5], and some plants reach coverages of up to 10 m² [79]. Big saltbush is typically evergreen, but can be drought deciduous in some desert environments [25,28,113,119]. The numerous leaves of big saltbush are somewhat thick, about 0.4 to 2 inches (1-5 cm) long, 0.25 to 1.5 inches (0.3-4 cm) wide, and covered in fine scales [8,25,113,119]. Big saltbush branches are numerous and slender. The bark is typically covered in fine scales when young and becomes rough on old trunks [113]. The small, imperfect flowers occur in panicles [25,74,119]. The fruits are utricles with bracts typically 0.1 to 0.15 inch (3-4 mm) long and wide, which contain a seed 0.04 to 0.06 inch (1-1.5 mm) wide [8,69,119].

RAUNKIAER [91] LIFE FORM:
Phanerophyte

REGENERATION PROCESSES:
Big saltbush reproduces by seed [25,79].

Breeding system: Big saltbush can be either monoecious or dioecious [113,119,123]. In plantings done to determine potential of big saltbush as a forage crop the sex ratio was 60% male plants, 10% female plants, and 30% monoecious plants [117]. In their 1984 article, Freeman and others [35] included data from previous research demonstrating change of sex in big saltbush. Plants typically change from dioecious to monoecious, but can also change from female to male. The ability to change sex appeared to enhance survival and may provide a reproductive advantage to the population. The table below shows the number of individuals from a wild population of 70 that exhibited each type of change in sex between 1978 and 1983 [35].

Pollination: According to Meyer [69] saltbush species are wind pollinated, but evidence demonstrating this for big saltbush is lacking. Saltbush sootywing butterflies feed on big saltbush nectar [81]. Whether they or other insects transfer pollen while feeding is not reported.

Seed production: Big saltbush has been reported to produce an abundance of seeds [25], but quantitative data illustrating this are not available.

Seed dispersal: Ohmart and Anderson [79] note that seed dispersal occurs mainly by water and vertebrates. Several species, including ring-necked pheasants and Gambel's quail, are known to eat big saltbush seeds [25,44], but there is no research addressing bird dispersal of big saltbush seeds.

Seed banking: Again there is little information available. However, seeds have been successfully stored in sealed containers for 5 years [8]. Jorgensen [53] reports a maximum storage time of 6 years.

Germination: In laboratory studies, big saltbush germination rates have varied. Watson and others [118] obtained average germination rates of 7% one year and 57% the following from four replicates of 25 seeds incubated on a 68/100 °F (20/40 °C) diurnal temperature regime. Seeds were collected in the fall and winter of both years. In low salinity treatments Jackson and others [51] found 21% to 24% mean germination rates of seeds with utricles removed, and Young and others [125] report a mean germination rate of 39% to 40% over 55 different temperature treatments ranging between 32 °F and 104 °F (0 and 40 °C). Malcolm and others obtained high germination rates, above 80%, with big saltbush seeds that had their utricles removed [62].

Thorough studies on the effect of temperature on big saltbush germination have been performed. Young and others [125] investigated the effects of 55 different alternating and constant temperature regimes on big saltbush germination. They obtained the best germination rates, between 65% and 68%, when the 8-hour warm period temperature was between 68 °F and 86 °F (20 and 30 °C) and the 16-hour cold period temperature was between 41 °F and 59 °F (5 and 15 °C). No germination occurred when warm and/or cold period temperatures were below 41 °F (5 °C). When both were 41 °F (5 °C) the mean germination rate was 5%. The only other temperature regimes that resulted in no germination were at warm-period temperatures of 104 °F (40 °C) when the cold-period temperature was 95 °F (35 °C) or higher. Mean germination was 6% when both warm and cold period temperatures were 95 °F (35 °C). All other temperatures resulted in germination greater than 20%. Results are from 4 replications of 100 seeds each [125]. Young and others [125] did not find a significant effect (p>0.01) of light on big saltbush germination. Mikhiel and others [70] found the highest germination rate with a treatment in which seeds were exposed to 50 °F (10 °C) and 68 °F (20 °C) each for 12 hours per day. The germination rates for the 3 treatments are shown in the table below [70].

In addition to temperature, salinity has a strong influence on germination rates. Jackson and others [51] observed no big saltbush germination in salinity treatments of 18,000 mg/L and higher. At salinity levels of 6,000 mg/L and lower, mean big saltbush germination rates were between 21% and 24%. The table below shows the germination rates of big saltbush subject to 4 salinity treatments [50].

Differences between the control and the 3 salinity treatments were significant (p<0.05), while the differences between salinity treatments were not statistically significant. The germination rate at the ~24 dS/m salinity level was significantly (p<0.05) higher than other saltbushes tested, including cattle saltbush and fourwing saltbush [50]. Mikhiel and others [70] also determined germination rates for big saltbush at 4 salinity levels; their results are shown in the table below. Unlike many other species investigated, big saltbush does not exhibit a significant synergistic effect of temperature and salinity [70].

In the field, estimates for germination rates are from rehabilitation plantings. For example, in a review of several revegetation projects, Biggs and Cornelius [13] report that germination of big saltbush on the Cibola National Wildlife Refuge in Arizona was initially high, with over 1,000 seedlings produced on four 19.7-foot × 29.5-foot (6- × 9-m) plots. Anderson and others' [3] first attempt at establishing big saltbush on the lower Colorado River, by broadcast seeding, was almost entirely unsuccessful. The table below shows big saltbush percent germination per meter of seeds planted in loam in watered areas along the lower Colorado River in their third attempt to establish big saltbush. The number of meters sampled per site is shown in parentheses [3].

Seedling establishment/growth: Big saltbush can successfully establish on many types of sites [3,13,30,41,60,80]. After 1 growing season in the northern portions of the Crescent Bypass riparian revegetation area, about 34 miles (56 km) south of Fresno, California, most big saltbush had grown quickly and survival was 88% [80]. In a review of revegetation projects, Briggs and Cornelius [13] report that some big saltbush grew to over 3 feet (1 m) and were producing seed after 3 growing seasons at Mittry Lake in Arizona. In rehabilitation plantings in western Australia, big saltbush had relatively high establishment compared to several other saltbush species and reached 14.6 inches (37 cm) tall and 40.6 inches (103 cm) diameter after 2 years' growth [60]. Three years following planting of quailbush in San Onofre State Beach, California, coverages reached 11.9% on a site that had been hydroseeded and 13.8% on a site that had been broadcast seeded and raked [48].

Despite the ability of big saltbush to tolerate drought, establishment and survival are probably improved with greater availability of water. Young big saltbush plants have been reported to be more susceptible to drought (Anderson and Ohmart, personal communication in [58]). Goodin and McKell [41] noted a lack of regeneration when subject to only 3.9 inches (100 mm) mean precipitation and speculated that the dry conditions were interfering with seedling establishment. Big saltbush growth to 6 feet (1.8 m) in height and diameter within a year was documented in plantings done near the Fresno Slough from 1959 to 1960 [30]. Given the short distance to the slough, the water table was likely shallow and plants were also irrigated during the summer months. In addition, Watson and others [118] found a general trend toward larger numbers of big saltbush establishing closer to the irrigation source. In 1992, they found 5 big saltbush established per linear meter when 1.5 meters from the irrigation source; this decreased with distance from the irrigation source. In 1993, a drier year, less than 1 big saltbush established per linear meter when 1.5 meters from the irrigation source. Again establishment decreased as distance from the irrigation source increased, although at a lower rate than observed in 1992 [118]. For more general information, see the Water section below.

High levels of salinity appear to affect seedling development. Jackson and others [51] reported no effect of salinity on the growth of big saltbush seedlings at salinity levels of 18,000 mg/L and lower. However at 36,000 mg/L and 60,000 mg/L,  shoot biomass was significantly (p<0.05) less than the 6,000 mg/L and 18,000 mg/L treatments. The total stem growth over 120 days was also significantly less (p<0.05) at 36,000 mg/L and 60,000 mg/L salinity levels compared to less saline treatments. The effects of the higher salt concentrations became larger over time. However, none of these salinity treatments resulted in seedling mortality [51]. In addition, Malcolm and others [62] found increased cotyledon width and more rapid formation of true leaves in delayed salinity treatments, where big saltbush seeds were subject to a period of low salinity (160 mS/m) before salinities were increased (1900 mS/m).

Asexual regeneration: There have been no reports of big saltbush reproducing asexually in the wild.

Elevations where big saltbush occurs range from sea level [90] up to 4,000 feet (1,219 m) in Arizona and Mexico [57,90]. Elevational ranges by state are shown below.

Temperature: The optimal temperatures for big saltbush depend on the location of the populations sampled. It has been demonstrated that for big saltbush in coastal areas, temperatures below 97 °F (36 °C) result in higher CO₂ uptake than big saltbush in desert areas. However, above 97 °F (36 °C) big saltbush in desert areas have more efficient CO₂ uptake. Photosynthesis is most efficient at 111 °F (44 °C) [86]. Big saltbush in desert areas have been shown to have a greater capacity to acclimate to high temperatures [87]. For effects of temperature on germination see Germination.

Soil: Big saltbush is typically found in moist to dry alkaline or saline soils [37,57,90,111], and has low tolerance for acidity [90]. The range of pH and electrical conductivity found on 3 sites, one in southwestern Utah and two in Nevada, containing big saltbush are shown in the table below [111].

Values of pH and electrical conductivity measured on sites near Safford, Arizona, where big saltbush was grown to test its use as a forage crop, are shown in the table below [117].

Big saltbush occurs in a variety of soil textures from quite coarse soils, especially in the case of the quailbush, which can grow in pure sand [96] to silty loams and silty clay loams [63]. Big saltbush was established in an area of revegetation with sandy loams [80]. Turner and Brown [108] note the occurrence of the saltbush series, a community type containing big saltbush, in areas with soil textures that are generally finer than sandy loams. Marks [63] reports saltbush communities typically growing in soils with intermediate textures, such as silty loams to silty clay loams. Soils with poor aeration can result in severe growth limitations in big saltbush [6]. Soil texture has not been shown to have a significant (p>0.05) effect on germination [125]. See the Germination and Seedling establishment/growth sections for more information on factors affecting big saltbush in these stages.

Water: Big saltbush may grow in areas with or without an accessible water table [37,41,57,66]. In California, Pearcy and Harrison [86] found moist soil at depths between 7.9 to 15.8 inches (20-40 cm) on a desert and a coastal site containing big saltbush. In an experiment investigating water use of several species, big saltbush was grown in tanks with an experimentally set water table depth of between 3.3 and 5.5 feet (1-1.7 m) [66]. According to McDonald and Hughes [66] big saltbush has been found on sites with deep water tables. These plants are likely to survive on surface sources of soil moisture and be less vigorous than specimens that occur in areas with a shallow water table.

Big saltbush can tolerate drought and flooding. Precipitation in areas with big saltbush can be very low. For instance, Goodin and McKell [41] noted their observation of big saltbush occurring in conditions of 3.9 inches (100 mm) mean annual precipitation. A catalog of native seeds recommends planting in areas which receive at least 4 inches (101.6 mm) of rain per year [90]. Although these areas may not typically receive much rain, big saltbush occurs in areas, such floodplains and valley bottoms that are subject to flooding [108]. Big saltbush has been classified as flood tolerant. It can survive flooding for most of a growing season, with some root growth likely during this period [115].

Big saltbush can also tolerate irrigation with saline water. Electrical conductivity and pH of water used to irrigate big saltbush at a site near Safford, Arizona were measured twice during a study investigating its use as a forage crop. In May of 1984 the water had an electrical conductivity (EC) of 10.3 dS/m and a pH of 8.6. In September 1984 the electrical conductivity was 9.3 dS/m and the pH was 8.8 [117]. The feasibility of irrigating big saltbush with saltwater has also been investigated. Wiley and others [124] used big saltbush from a site irrigated with seawater discharge from a shrimp aquiculture facility in Sonora, Mexico. They reported total dissolved salts of 40 parts per trillion (ppt); typical levels for seawater are between 30 to 35 ppt. In another investigation in the same area using the same water source (hypersaline, 39%-41%), the growth rate of big saltbush was less than that for sites irrigated with freshwater in Bodega Head and Death Valley in California. However, mean annual productivity was 1,794 grams of dry weight/m² (mean = 149), which is similar to the more northern sites due to the longer growing season in Sonora [38]. See Palatability/nutritional value for more information on the forage production of big saltbush.

In riparian areas, disturbances, typically floods, occur regularly. Johnson and others [52] note that desert riparian communities that establish after a disturbance typically have the same species assemblages as before the disturbance. Species composition is influenced more by site characteristics, such as depth to water table, than time since last disturbance. The frequency and intensity of disturbances does affect which species can establish. For instance, consistent flooding can allow for establishment of species that do not tolerate high levels of salt [13]. This interaction between disturbance and site characteristics results in a dynamic mosaic of vegetation types [18]. Whether flooding was recent or occurred some time ago, big saltbush could occur on a site with appropriate conditions. Whether this trend would be observed after other types of disturbances, such as fire, has not been reported. However, Busch [16] was unable to detect a postfire successional trend in burned cottonwood-willow woodland along the lower Colorado River.

Although healthy riparian forests of the Southwest may not exhibit Clementsian succession, changes to disturbance regimes have resulted in changes in species composition. In altered habitats, such as those with decreased water tables, decreased frequency of flooding, and a resulting increase in fire, tamarisks (Tamarix spp.) can replace native species over time [17,24], resulting in older sites being dominated by dense stands of tamarisk [24]. Due to big saltbush being a "vigorous competitor" on sites where it is already established, these areas may be less likely to follow this pattern [28].However, the strong response of tamarisks after fire [16] could negatively affect big saltbush exposed to increased fire frequencies. For a comprehensive review of tamarisks see the FEIS Tamarisk review and Glenn [37].

Few studies have addressed succession in saltbush scrub. Karpiscak [54] summarizes succession in saltbush and creosotebush shrublands. Species typical of saltbush or creosotebush vegetation typically recolonize abandoned agricultural areas after a series of mostly invasive annuals. Russian-thistle (Salsola kali) gives way to several mustard species (Brassicaceae) after two or three years, which is followed by annual grasses. Goldenbush (Isocoma spp.) and desertbroom (Baccharis sarothroides) follow, and typically establish just before saltbush or creosotebush species [54].

FIRE ECOLOGY

• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

Fire regimes: In desert shrublands fire is rare due to lack of continuous fuels [67,82,100,120,121]. The expansion of invasive annuals such as cheatgrass (Bromus tectorum) and red brome (B. madritensis ssp. rubens) can increase the frequency of fire in these ecosystems [121]. Fires in saltbush vegetation are likely to be more severe and spread faster with increasing fuel porosity, decreasing levels of moisture, and increasing amounts of fine fuels and dead vegetation [76].

Little is known of the role of fire in riparian habitats of the desert Southwest. The flammability of riparian habitats would likely vary temporally with drought and spatially due to fire frequency of surrounding landscapes [110]. There is evidence that the fire frequency in riparian areas is longer and more variable than that of the surrounding landscape [98,99,110]. Riparian areas can slow or impede the spread of fire [29,98,99,104]. However, in drought conditions higher fuel loads compared to other saltbush containing communities (250 to 1000 lb/acre for mesquite bosque forest compared to 40 to 100 lb/acre for saltbush/greasewood types [84]) can make riparian areas more susceptible to fire [29]. Human ignitions sources [29,110] and the invasion of tamarisk [16] will likely increase the frequency of fire in these habitats.

The following table provides fire return intervals for plant communities and ecosystems where big saltbush occurs. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".

FIRE EFFECTS

SPECIES: Atriplex lentiformis

• IMMEDIATE FIRE EFFECT ON PLANT
• DISCUSSION AND QUALIFICATION OF FIRE EFFECT
• PLANT RESPONSE TO FIRE
• DISCUSSION AND QUALIFICATION OF PLANT RESPONSE
• FIRE MANAGEMENT CONSIDERATIONS

IMMEDIATE FIRE EFFECT ON PLANT:
In laboratory experiments, big saltbush has been shown to exhibit reduced flammability compared to highly flammable chaparral shrubs [72] and plants with lower silica-free mineral content than big saltbush [88]. The effects of wildfire or prescribed burns on big saltbush have not been investigated.

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
Big saltbush has been shown to have reduced flammability due to high moisture and ash contents. Montgomery and Cheo [72] reported reduced flammability of 3.2 to 4 inch (8-10 cm) cuttings of big saltbush terminal growth due to its naturally high moisture content. The table below shows the ignition times and type of combustion (ignition times for char combustion are approximations) for big saltbush of varying moisture content at 1202 °F (650 ?C). Heat-dried samples were exposed to 212 °F (100 ?C) for 45 minutes then placed in a desiccator until just before burning. They showed no reduction in flammability compared to heat-dried samples of species considered to be highly flammable [72].

rainy season	dry season	heat-dry
char (18-24 seconds)	char (12-20 seconds)	flame (2.2 ? 0.28 seconds)

Field collected big saltbush moisture contents were high, 62.3% fresh weight in the dry season and 71.5% fresh weight in the rainy season. Big saltbush also lost moisture at a slower rate than the other species investigated. Ash content was not found to have an effect on big saltbush flammability. However, burning was at temperatures much higher than those where effects of ash content are likely to be observed (Philpot, personal communication in [72]). Philpot [88] demonstrated species with high ash content, including big saltbush, exhibited reduced flammability compared to species with low ash content when burned at temperatures between 225 °F and 720 °F (125? and 400 °C).

Despite reports of big saltbush producing large amounts of seed [25], even less is known about the colonization of burned areas by big saltbush. High ambient temperatures have been shown to decrease germination rates [125]. However, the effects of a more intense heat source for a shorter duration have not been investigated. Although the same moisture and soil factors important on unburned sites are probably important, factors affecting the ability or inability of big saltbush to establish on recently burned sites are unknown.

MANAGEMENT CONSIDERATIONS

• IMPORTANCE TO LIVESTOCK AND WILDLIFE
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER USES
• OTHER MANAGEMENT CONSIDERATIONS

Palatability/nutritional value: Leaves and seeds of big saltbush are eaten by many species. Mule deer, pronghorn, and livestock browse the leaves [6,25,57,84,94,113]. In literature reviews, small mammals such as rabbits and rodents have been reported to eat big saltbush [44,84]. Briggs and Cornelius [13] report rabbit damage to big saltbush in a revegetation plot and Everett and others [32] report that, although big saltbush was one of the least preferred species in their laboratory trials, deer mice ate its seeds. Reviews have included big saltbush as a component of the diet of several game birds [30,44]. In a literature review, Gullion [44] noted use of big saltbush by ring-necked pheasants and Gambel?s quail. Big saltbush is also important to some insects. The saltbush sootywing uses big saltbush as one of its hosts as a caterpillar and feeds on the nectar of big saltbush flowers as an adult [81].

Several studies have investigated the nutritional content of big saltbush. Cibils and others [23] found the mean and range of several nutritional measurements available in the literature. These are shown in the table below and represent values for a number of saltbush species.

Although the values from several studies specifically investigating big saltbush fall within these ranges [22,59,124], there are exceptions. For example, gross energy reported by Wiley and others [124] was less than that reported in the Cibils and others's literature review [23]. Wiley and others found 3.1 Mcal/kg in untreated big saltbush and 3.9 Mcal/kg gross energy in washed big saltbush. This data was obtained from big saltbush irrigated with highly saline water (40 ppt) [124]. Khalil and others [59] included digestible energy, which was 3.215 Mcal/ Kg, and metabolizable energy, which was 2.636 Mcal/ Kg. They also found a lower fiber content, 8.0% dry weight [59], than the minimum reported by Cibils and others [23].

Wiley and others [124] and Khalil and others [59] included other big saltbush measurements. Wiley and others [124] reported mean ash content measurements of 37.4% dry matter for untreated big saltbush and 21.7% dry matter for washed big saltbush, while Khalil and others [59] found an ash content of 22.0% dry weight. In addition, both of these studies measured acid detergent fiber (ADF). Wiley and others [124] found an ADF of 23.2% dry matter in untreated big saltbush and 27.5% dry matter in washed big saltbush, while Khalil and others [59] measured ADF as 18.5% dry weight. Wiley and others [124] measured neutral detergent fiber as 41.6% dry matter for untreated big saltbush and 53.6% dry matter for washed big saltbush. Khalil and others [59] included the results from their measurement of digestible dry matter, which was 74.5% dry weight. They also measured several important minerals. The results are included in the table below [59].

Other studies investigating nutritional values of big saltbush included results over time or variations due to site conditions, such as high salinity. Two studies have addressed the effect of time on nutritional content of big saltbush. The table below shows the results of Goodin and McKell [41] from two harvests in percent dry weight.

Watson and others [117] found an effect of harvest treatment and phenological stage on the nutritional content of big saltbush. Like Goodin and McKell [41] they observed a decrease in ash and protein contents over time. The table below shows nutritional information determined by Watson and others [117] from harvests of aboveground plant material when plants were harvested near 60 cm tall. All measurements are shown in percentages of dry weight, except fluoride, which is in ppm.

Salinity has also been shown to have an effect on nutritional aspects of big saltbush. Ibrahim [50] measured the contents of several minerals in four salinity treatments. The results of this investigation, in percent dry weight, are shown in the table below.

The following table shows the ion contents in mol/g of dry weight for several durations of salinity treatments found by Malcolm and others [62]. The 1st number in each column is the result for the low salinity treatment of 160 mS/m, while the 2nd number is from the high salinity treatment, 1,900 mS/m.

Toxicity of big saltbush may be a problem in some areas. Concern over selenium and oxalate levels have been addressed. In a literature review Guillion [44] notes that saltbush species can reach toxic levels of selenium. In a laboratory study soil was injected with 3 mg sodium selenate per kilogram of soil. This was repeated every ten days until the soil had been injected with 18 mg of Se/kg of soil. The harvested big saltbush contained selenium at a concentration of 7 ppm, while the combined, root free soil had a concentration of 2 ppm. These selenium levels in big saltbush could be toxic if it comprised the majority of the diet [27]. Glenn and O'Leary [38] measured oxalate levels and found mean of 3.57% dry weight. The highest oxalate values Watson and others [117] obtained was 4.57% dry weight. This was considered nontoxic. Watson and others [117] also determined that fluoride levels were sufficiently low.

In addition to the possibility of toxicity, there are other disadvantages of use of big saltbush as a forage species that should be addressed when considering its use. It has been found to lack phosphorus [117] and carbohydrates [41]. Other authors have noted the high levels of ash [38] and salts, especially sodium [38,50,59]. Acceptability of big saltbush is also an issue. Domestic goats found diets with 25% big saltbush (leaves and smaller stems) palatable, but big saltbush had been shown in previous investigations to have low acceptance when comprising larger proportions of the goat diet [124]. The Cibilis and others's [23] literature review also addressed the issue of adaptation to the diet. They suggest that if a period of adjustment is necessary for animals to uptake the nitrogen in big saltbush, as some evidence suggests, saltbush species may be much less useful in filling gaps in food availability. Despite these concerns most agree that big saltbush is valuable forage for livestock, when not the sole food source [41,59,117,124].

In addition to the nutrition value, productivity of big saltbush is another attribute which increases its forage potential. Ibrahim [50] notes that big saltbush can produce over five kilograms of dry matter every 3 months. Goodin and McKell [41] reported yields from big saltbush planted as a forage crop. Fresh weight yields (kg/ha) for various harvesting strategies are included in the table below.

They found no statistically significant (p > 0.05) advantage of harvesting twice when plants reached 17.7 inches (45 cm) compared to harvesting once at 23.6 or 29.5 inches (60 or 75 cm) height. Watson and others [117] also determined there was no advantage to harvesting more than once during the establishment year. The highest estimated yield of big saltbush they obtained was 14.7 tonnes/hectare. In addition, transplant survival of big saltbush was high [117]. In a later investigation, Watson and O'Leary [116] found yield rates of 6.3 t/ha, 9.3 t/ha, and 3.5 t/ha in the subsequent harvests. Only one treatment with saline water (EC 18 dS/m) occurred before the first harvest, but thereafter the irrigation water had a mean electrical conductivity of 18 dS/m. The drop in the yield of the third harvest reflected loss of individuals from previous harvests [116]. Glenn and O'Leary [38] measured the productivity of big saltbush when irrigated with hypersaline (39%-41%) seawater and obtained an estimate of 794 grams of dry weight/ m²/ year (s=149) over a year, which was similar to productivities of big saltbush irrigated with freshwater on sites with shorter growing seasons. Despite encouraging results with forage crops, sustainable browsing of livestock on natural stands of big saltbush is complex. According to the Cibils and others [23] literature review, the processes of recruitment and mortality in saltbush species are more sensitive to changes than other shrubs. Care must be taken to incorporate both the effects of herbivory and its interactions with intra- and interspecific "competition" of the plant species when managing these communities.

Cover value: Dense stands of big saltbush provide excellent cover for several species. Most work has investigated avian use of big saltbush. Anderson and others [3] found that plots with big saltbush had higher avian densities (p<0.05) than vegetatively similar sites without big saltbush in 3 out of the 4 seasons investigated. Species including blue grosbeak, blue-gray gnatcatcher, Crissal thrasher, ruby-crowed kinglet, Gambel's quail, and verdin had higher densities on sites with big saltbush [3]. Similar studies have found comparable results [2,28]. Disano and others [28] found that densities of several avian species and guilds, such as passerine granivores, Gambel quail, and to a lesser extent permanent residents and visiting insectivores, were higher in areas where big saltbush and Mohave seablite coverage was approximately 1,500 plants per hectare compared to densities averaged over all riparian vegetation types along the lower Colorado River. Gullion [42] lists big saltbush as one of several species in habitats that provide excellent Gambel quail cover. In a literature review he notes big saltbush's importance to Gambel's quail in southern Nevada [44]. Ohmart and Anderson [79] note the possibility of high densities of foliage arthropods contributing to the high avian habitat quality of big saltbush, although the mechanisms behind the interaction between big saltbush and avian populations are uncertain. In addition to providing anecdotal evidence suggesting that big saltbush revegetation efforts improved habitat quality to the point of allowing for successful establishment of Gambel's quail, Ermacoff [30] reported an increase in ring-necked pheasant activity and populations as well as increased cottontail populations. Little else has been published regarding small mammal use of big saltbush. However, information on rodent use of saltbush vegetation in general is available [1].

Although pretreatment of seeds is not necessary for germination, some pretreatments may enhance germination rates. For example, a month-long afterripening period is suggested by Jorgensen and Stevens [53], and improved germination occurs if seeds are rinsed or soaked and then wrung [8,125]. Young and others [125] reported increases in germination rate of 20% or more in seeds that were rinsed and wrung before planting when incubated at temperatures between 50 °F and 77 °F (10?-25 °C). The maximum germination rate, just over 80%, was observed when rinsed and wrung seeds were germinated at 68 °F (20 °C).

Conditions that improve germination include covering seeds with a small amount of soil and germinating at optimal temperatures and salinities. Depth of planting appears to be important factor for germination success. Goodin [40] reports that big saltbush should not be planted at depths of more than 0.2 to 0.4 inches (5-10 mm). Also germination is significantly (p < 0.05) enhanced when a thin layer of soil covers the seeds. The table below shows germination rates for different soil types when seeds were planted at either a depth of 0.2 inches (0.5 cm) or on the soil surface. Differences between soil types were not statistically significant (p > 0.05) [125].

The ease of propagating saltbush species, including big saltbush from cuttings [64,77,122] provides another method for production. Wieland and others [122] report that cuttings of many of the saltbush species tested rooted without the application of rooting hormone, although rooting success in big saltbush was best when rooting hormone was used. Richardson and others [92] investigated factors affecting rooting success of fourwing saltbush, shadscale, and valley saltbush. They found that 4.7 inch (12 cm) cuttings had much higher rooting success than 2.4 inch (6 cm) cuttings, that concentrations of rooting hormone between 0.3% and 2.0% had the best results depending on the species, and that propagating cuttings in spring and summer resulted in the highest success while propagating cuttings in fall had the lowest success. In a study of 54 Nevada shrubs, big saltbush was one of the easiest species to propagate from cuttings and had higher rooting success when cuttings were semi-hardwood and/or collected during the stage of twig growth [31].

Several methods and levels of effort toward subsequent maintenance have allowed successful establishment of big saltbush. Hydroseeding [10,48], use of a niche seeder [60], broadcast seeding then raking [48], and transplanting individuals older than 9 months [95] have all resulted in successful establishment of big saltbush. Briggs and Cornelius [13] provide a review of several revegetation efforts; some of these plant big saltbush and planting methods used differ. Given the susceptibility of young saltbush to drought, irrigation during the first summer is likely to improve establishment (Anderson and Ohmart, personal communication in [58]). Reductions in salinities, even for short periods after planting, may also assist in establishing big saltbush [62]. For general information on effects of water and salinity in big saltbush establishment see Seedling establishment/growth. Fencing may be included to reduce browsing [95], as several revegetation projects have reported losses from livestock and small mammals such as rabbits and gophers [13,39,95]. Seed predation by deer mice has also been investigated. Although not a preferred species, big saltbush seeds may suffer from high predation rates when other food sources are lacking [32]. Collection of seed from on-site sources is another option. This is recommended for riparian revegetation efforts [58] and has produced good results [10]. Seeds can be stored for a maximum of 3 to 6 years [8,53,89]. Winter has been suggested as the best time for planting seeds [58,90], while transplants 2 to 3 years old have been successfully planted throughout the summer when irrigated [95].

Large rates of big saltbush mortality and low growth rates have been reported, but the cause of these outcomes is unknown. For instance, on the third attempt Anderson and others [3] successfully established big saltbush on three watered sites along the lower Colorado River. Mean growth rates from the time of planting to 6 months later were between 0.8 and 1.2 inches (2-3 cm). Big saltbush planted at a mine reclamation site exhibited 100% mortality. Whether this was due to the level of soluble salts, which ranged between 335 and 3182 ppm, or some other factor or factors is not discussed [78].

Traditional Uses: Castetter's [21] literature review of studies on tribes of the American southwest included information regarding the Pima Indians' practice of pit curing and drying big saltbush seeds before using them to make a thick gruel. Bean and Saubel [7] report a similar practice among the Chauilla as well as use of the flour to make small cakes, use of leaves as a soap, and use of flowers, stems and leaves as a treatment for nasal congestion. Conrad [25] suggests that seeds were likely used in a similar way to fourwing saltbush. Seeds of fourwing saltbush were also reportedly ground into flour. Other uses for fourwing saltbush that may have been similar for big saltbush are the use of the ground meal as an emetic, use of ground flowers or roots moistened with saliva in treating ant bites, and addition of ashes to water for dyeing meal greenish-blue [25].

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