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Chilopsis linearis


Figure 1—Desert-willow in Anza Borrego State Park, California. Photo © 2000 Larry Blakely, used with permission.

SUMMARY

This review summarizes the information on the biology, ecology, and effects of fire on desert-willow in that was available in the scientific literature as of 2021.

Desert-willow is native to the southwestern United States and northern Mexico, occurring in the Mojave, Sonoran, Colorado, southern Great Basin, and Chihuahuan deserts. It is an obligate riparian species, and it is most common in wash shrublands and mixed-deciduous riparian woodlands. Desert-willow reproduces sexually and regenerates from seed. It sprouts from the root crown following top-kill. Desert-willow is cross-pollinated by bumble bees. Seed crops are typically large in wet years. Germination occurs on moist substrates, soon after flooding, and growth is typically rapid in moist soils. Desert-willow is most common in open areas. Flooding is the primary disturbance in communities with desert-willow, but desert-willow also grows in early postfire environments. With alterations in hydrology and climate change, fire is becoming the primary disturbance in many riparian ecosystems.

Low- to moderate-severity fire prunes or top-kills desert-willow. It sprouts after top-kill by fire and may also establish from seed. Historically, fuels in plant communities where desert-willow is common were likely mostly sparse and discontinuous, although information on this is limited. Fires were probably infrequent, and they likely varied in severity and extent across southwestern riparian ecosystems. Most fires were likely patchy and of low severity due to patchy and sparse fuels.

Changes in hydrology and vegetation have undoubtedly affected fuels and fire regimes of southwestern riparian communities and surrounding uplands. However, the degree to which fire regimes are changing in southwestern riparian ecosystems—or what the effects might be for riparian ecosystem structure, function, and resilience—are not well known. Severe, intense, and/or frequent fire can interact with altered hydrology to accelerate the replacement of native plants with nonnative and often invasive plants such as tamarisk and Russian-olive. When nonnative species invade the overstory and/or understory and alter riparian vegetation composition and structure, fuel characteristics may also change. In turn, this can alter fire behavior and fire regimes. Fire management in southwestern riparian areas may include fuel reduction and prescribed fire treatments that promote or maintain native vegetation and reduce or eradicate invasives.

The value of southwestern riparian woodlands for both biotic communities and human livelihoods is well established. Communities with desert-willow provide important habitat and food for a wide variety of wildlife species. Riparian areas with desert-willow provide many ecosystem services including recreational opportunities, harboring pollinators, and providing livestock forage. Few naturally functioning riparian areas remain in the Southwest, and those that do are threatened with climate change. Southwestern riparian ecosystems are vulnerable to decline under increasingly hotter and drier conditions. Climate change is expected to alter water flow and lower water tables, and to increase the frequency and severity of extreme weather. This will likely result in more frequent and severe floods as well as more intense droughts. Where natural flow regimes are more intact and tamarisk and other invasive species are suppressed by native vegetation, desert-willow and other native trees are more likely to maintain dominance in postfire landscapes.

Citation:
Fryer, Janet L. 2022. Chilopsis linearis, desert-willow. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Missoula Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/plants/tree/chilin/all.html [].


Table of Contents


INTRODUCTION



FEIS abbreviation:
CHILIN

Common names:
desert-willow
catalpa-willow
desert catalpa
false willow
flowering-willow
mimbre
willowleaf catalpa

TAXONOMY
The scientific name of desert-willow is Chilopsis linearis (Cav.) Sweet (Bignoniaceae) [1,6,55,100,121,129,161,220,224,251]. Chilopsis is a monotypic genus [43,55,85,97]. There are three subspecies:

Chilopsis linearis subsp. arcuate (Fosberg) Henderson
Chilopsis linearis subsp. linearis [6,85,97,119,161,220,224]
Chilopsis linearis subsp. tomenticaulis Henrickson [85]

Common names are used throughout this Species Review. For scientific names of plant and animal species mentioned in this review and links to other FEIS Species Reviews, see the Appendix.

Reviews cited in this Species Review include: [127,134,192,220,244].

SYNONYMS
For Chilopsis linearis subsp. arcuate:
     Chilopsis linearis (Cav.) Sweet var. arcuate Fosberg [68]

For Chilopsis linearis subsp. linearis:
     Chilopsis linearis (Cav.) Sweet var. glutinosa [1,68,100,129]

LIFE FORM
Tree-shrub

DISTRIBUTION AND OCCURRENCE


GENERAL DISTRIBUTION
Figure 2—Distribution of desert-willow. Map from Little (1976) [120] and digitized by Thompson et al. (1999) [213]. Desert-willow also occurs in Kansas, Oklahoma, and Georgia, although precise distributional information is sparse for those states [96,224].

Desert-willow is native to the southwestern United States and northern Mexico [96,120,224] (fig. 2), occurring in the Mojave, Sonoran, Colorado, southern Great Basin, and Chihuahuan deserts [51,159]. It is distributed from southern California eastward through southern Utah to western Texas [63,96,100,120,127,130,226] and southward to Baja California Norte, Durango, and Zacatecas, Mexico [120,130]. Disjunct but apparently native populations occur in Kansas, Oklahoma, and Georgia [96,224]. Desert-willow is planted outside its native range [63].

Chilopsis linearis subsp. arcuate occurs in California and Arizona; Chilopsis linearis subsp. linearis occurs in Kansas, Oklahoma, and Texas. Distributions of these subspecies overlap in Socorro County, New Mexico [224]. Chilopsis linearis subsp. tomenticaulis occurs in northeastern Mexico [85]. PLANTS Database provides distributional maps for the first two subspecies.

States:
United States: AZ, CA, GA, KS, NV, NM, OK, TX, UT [96,120,224]
Mexico: BCN, Chih, Coah, Dgo, Son, Zac [120]

SITE CHARACTERISTICS
Desert-willow is a warm-desert species that grows in lowlands [220,244] with bimodal rainfall. In its northwestern distribution, rain falls mostly in winter; in its eastern distribution, rain falls mostly in summer [9,156,183,220,236].

Arid climate restricts this phreatophyte to moist areas [51,159]; desert-willow is an obligate outwash or riparian species [13,19,39,53,54,102,157]. It is considered an indicator of sites with relatively shallow water tables [138]. Sites with desert-willow include stream- and riverbanks, alluvial fans, washes, arroyos, gullies [10,73,116,220,243,244], and bajadas [51,159,215,251]. It also grows on moist plains, hillsides [51,159,251], mesas, and terraces [51,159,215,226,251]. It is rare on unstable dunes [220].

Desert-willow is flood tolerant [16,73,92,156], and it is frequent on sites that experience intermittent flooding [10,73,116,146,159,220,243,244]. It often occupies the middle and banks of drainage channels, sometimes covering broad expanses in washes [73,92]. On the Gila River Resource Area, Arizona, desert-willow is most frequent in areas that receive severe scouring [137]. It may survive 1 to 3 months of inundation during the growing season, going dormant during prolonged floods [16]. It is somewhat drought intolerant ([210,217]); desert-willow mainly avoids drought by reducing transpiration [149,150] and/or shedding its leaves (see Seasonal Development).

Desert-willow grows at low elevations, occurring between 450 and 1,830 m elevation across its range [98,100,127], usually below 1,600 m [51,159,215] (table 1).

Table 1—Elevational range of desert-willow by area.

Area

Elevational range (m)

Sonoran Desert

≤1,500 [220]

Southwest 1,146–1,433 [208]
Arizona generally ≤1,650 [166], rarely to 1,830 [100]
California <2,100 [6]
New Mexico 1,200–1,700 [34,129]
Nevada, southern 500–1,800 [98]
Texas, Trans-Pecos region 600–1,500 [161]
Utah, southern 1,400 [91]

Desert-willow generally grows in alluvial substrates [73,239], with soils composed of sandy loam [95], sand [6,34,38,141,154,157,239,246], gravel [6,46,129,130,157,226,235,251], and/or cobble [157,159]. Desert-willow is intolerant of saline soils [182].

PLANT COMMUNITIES
Desert-willow is dominant in or a common component of wash shrublands [19,21,112,125,162,166,176,214,215,239]; mixed-deciduous riparian [12,56,84,165], pinyon-oak [115,134,147], and Joshua tree [43] woodlands; and some grasslands [76,238]. Washes and woodlands with desert-willow are typically adjacent to upland desert shrubland, desert grassland, or plains grassland communities [19,159,183,211].

Desert-willow is most common in and may codominate in creosote bush [43,113,184], mesquite [20], and catclaw acacia [21,214,215,242,247] wash shrublands. In southern California and Nevada, desert-willow grows in creosote bush-burrobush [43,113,184] and blackbrush [159,183] communities. In Arizona, it commonly grows in washes with catclaw acacia, desertbroom, eastern Mojave buckwheat, screwbean mesquite, and/or velvet mesquite [21,154,159,242]; and in riparian bosques and scrublands with honey mesquite [21,154,215,247]. In the Colorado, Sonoran, and Chihuahuan deserts, desert-willow sometimes codominates or associates with paloverde [21,215,247]. In Guadalupe Mountains National Park, western Texas [75], and southward to northern Mexico [42,64], it codominates with creosote bush and honey mesquite [42,64,75].

Desert-willow also occurs in other wash shrublands, or other shrublands over shallow water tables. In the Coachella Valley, California, smoketree-desert-willow-burrobrush communities occur near drier yucca/cactus communities [246]. Desert-willow is locally dominant in paloverde-cactus communities of the Mojave and Sonoran deserts [139,159,239]. In Arizona, it may also codominate with spiny hackberry and western soapberry [148], and on the Kofa National Wildlife Refuge, Arizona, desert-willow grows with ironwood, blue paloverde, and smoketree [245]. In the Trans-Pecos region of Texas, desert-willow is a component of Apache plume shrublands along arroyos [211]. In Chihuahua, Mexico, desert-willow grows in tropical-arid riparian shrublands with baccharis, singlewhorl burrobrush, and tree tobacco [156].

Desert-willow is a subcanopy tree in many riparian mixed-deciduous woodlands of the Southwest. These woodlands are codominated by Arizona sycamore, Arizona walnut, Fremont cottonwood, and/or velvet ash [12,84,165]. Willows—including arroyo willow, Goodding’s willow, and narrowleaf willow [12,155]—often codominate as well [56,155]. Shorter-statured deciduous communities dominated by desert-willow mostly occur in lower portions of arroyos, where streambeds widen [159]. Desert-willow riparian woodlands are a minor type in the Santa Ana Mountains of California [233], central New Mexico [56], and western Texas [159]. In southern Utah, desert-willow forms thickets with cottonwoods and nonnative tamarisk [98]. On the Stockton Plateau of west-central Texas, Arizona walnut-desert-willow communities occur on streambanks [86]. Desert-willow/splitleaf brickellbush communities occur on the Edwards Plateau of south-central Texas, particularly on intermittently flooded banks and washes [144]. Little walnut-desert-willow-splitleaf brickellbush communities occur in southern Texas [145]. Along the Rio Conchos in Chihuahua, Mexico, desert-willow is subdominant in black walnut-Lombardy poplar-Texas paloverde communities [42].

Desert-willow occurs in moist areas within pinyon-juniper-oak woodlands [115,134,147]. At the turn of the last century, Leiberg (1900) noted Fremont cottonwood-velvet ash/desert-willow-white alder stands in ravines within singleleaf pinyon-Parry pinyon-canyon live oak woodlands of the San Jacinto Reserve in California (now part of the San Bernardino National Forest) [115]. In the southern Guadalupe Mountains of Texas, desert-willow is a component of wavyleaf oak-alligator juniper-Texas madrone woodlands [147].

Desert-willow occurs adjacent to and may finger into some desert [167,230] and southern plains [76,211,238] grasslands. It grows in upper-elevation (1,700-1,900 m) [238] blue grama-tobosagrass [76,238] desert or plains grasslands. In Mexico, desert-willow codominates or occurs in desertbroom-netleaf hackberry-spiny hackberry riparian scrub [167] that merges into desert grasslands [230] composed of Arizona muhly, Hall's panicgrass, and sideoats grama [167].

See table A3 for a list of representative plant community classifications in which desert-willow occurs.

BOTANICAL AND ECOLOGICAL CHARACTERISTICS


Figure 3—Desert-willow blooming in Springs Preserve Garden in Las Vegas, Nevada. Photo by Stan Shebs, used with permission.
Figure 4—Seed-filled desert-willow capsules from Beaver Dam Wash, Nevada. Photo © 2016 Zoya Akulova, used with permission.

BOTANICAL DESCRIPTION
This description covers characteristics that may be relevant to fire ecology and is not meant for identification. Identification keys are available (e.g., [1,6,43,55,129,251]). Henrickson (1985) provides highly detailed botanical descriptions of desert-willow and its subspecies [85].

Aboveground Description
Desert-willow is a deciduous [85,92,127,130,232] small tree or shrub [1,127,129]. It is typically 3 to 7.5 m tall [127] but sometimes up to 12 m tall [1]. Habit is described as "tall and straggling" [180] (fig. 1). Trunks range from 10 to 40 cm in diameter [1], with one to several stems arising from the root crown [91,129,251]. The bark is "very thin": not more than 0.6 cm thick [91]. The upper crown is spreading [119]. Like its namesake, willow, desert-willow has slender, flexible branchlets [68] and drooping, elongated (linear-lanceolate) leaves [46,68,85,155,237]. Leaves are simple [6,100,130,251], with a thick, waxy cuticle that reduces water loss [220,226]. Some leaves may have extrafloral nectaries [85,157]. In a Sonoran Desert population, presence of extrafloral nectaries was variable both within and among individual trees [157]. The inflorescence is a raceme or panicle with showy, tubular flowers [85,129,161,251] (fig. 3). The fruit is a slender capsule [127,129,251] from 10 to 30 cm long and about 6 mm in diameter [119,127,237]. In part, this genus/species is distinguished by its long, leathery capsules. Its seeds have a fringe of soft white hairs on each end [85,127,130,161,180,232,251] (fig. 4). Seeds are flat, about 8 mm long [127], and have 10 to 15 mm-long hairs [129,161,232,251].

Belowground Description
Desert-willow has a taproot [46] and is deep rooted [51,116,180,226]. Horizontal roots may extend as far as 15 m [220]. On the Jornada Experimental Range, New Mexico, the taproot of a 50-m³ tree was 160 cm deep. Most of the total root biomass was in shallow soil: the distribution of root biomass by soil depth averaged 72% from 0 to 40 cm, 18% from 40 to 80 cm, 8.6% from 80 to 120- cm, and 1.4% from 140 to 160 cm [51].

Stand or Population Structure
Structure of plant communities with desert-willow are variable in structure [175]. Desert-willow is typically a subcanopy tree in mixed-deciduous communities, but it is a canopy species in shrublands or short-statured woodlands. It often forms thickets along washes [91]. Communities in scour zones [154] or that are accessible to off-highway vehicles (OHVs) [69] may lack or have sparse understories.

A few studies quantify stand structure of desert-willow in pure and mixed-deciduous communities. See these references for densities of desert-willow and associated trees in Arizona and New Mexico [208], New Mexico [51], and northwestern Chihuahua, Mexico [208].

Desert-willow may live as long as 100 years [55]. Other information on age class structure was lacking in the literature.

Raunkiaer Life Form
Phanerophyte
Geophyte [164]

SEASONAL DEVELOPMENT
Desert-willow has a long flowering period (table 2). It may produce flowers from April through August throughout its range [127]. Flowering is triggered by winter [220] and summer [138,220] rains. Mass flowering usually occurs early in the growing season. After mass flowering, plants usually produce only one to four flowers at a time, over an extended flowering season. Flowers open and produce nectar from late afternoon through late night, with flowers senescing after 1 to 2 days [172]. Fruits ripen from late summer to late fall and persist on the plant through winter [127]. Seed dispersal coincides with winter and spring floods [101]. Seeds germinate in summer in southern California [246]; other germination times were not available.

Desert-willow may drop its leaves in any season. It is cold-deciduous [46,102,116,220], with leaves falling in late fall or winter [46,116] after temperatures fall below around 5 °C [220]. It is also drought-deciduous [149,155,177,220].

Table 2—Phenology of desert-willow.
Area Event
Intermountain region Flowers May-July [43]
Sonoran Desert Leaves 1st emerge March-May; flowers April-August [172,220]; drought-induced leaf drop June-July; 2nd leaf emergence with August rains [220]
Southwest Flowers May-June; sporadically thereafter following summer rains [232]
Arizona Flowers April-August [100]
California Germinates in summer [246]; flowers May-June [6,28]
New Mexico Flowers April-August [129,237]; leaf drop October-November [102]
Nevada, southern Flowers May-August [98]
Texas, Trans-Pecos region Flowers April-September [161]
Texas, north-central Flowers May-September [55]
Utah, Washington County Flowers May-July [226]
Baja California Flowers August-April [251]

REGENERATION PROCESSES
Desert-willow reproduces sexually and regenerates from seed. It sprouts from the root crown following top-kill. Survival and reproduction of riparian trees is largely determined by favorable precipitation and periodic floods and droughts [192]. Seed crops are typically large in wet years. Germination occurs on moist substrates, soon after flooding. Growth is rapid in moist soils. Droughts can result in high mortality of juveniles and sprouts.

Pollination and Breeding System
The flowers are perfect [127], with a strong, pleasant scent [130,188,226]. Sonoran bumble bees [43,171,172,226,248] are the main pollinators. Hummingbirds [22,80,171,172,220,226], honeybees [171,172,232], lady-beetles, small flies [220], hawkmoths, carpenter bees [43,171,172,226], and other nectar-robber bees [171,172] also visit the flowers; but they are not effective pollinators [171,172].

Desert-willow is mostly cross-pollinated [158,172]. Its flowers close after the stigmata are touched, reopening if pollination was ineffective (e.g., not enough pollen was deposited or self-pollination occurred) [172]. Hand-pollination of flowers near Tucson, Arizona, resulted in a 540% increase in fruit set compared to bumble bee pollination [158].

Seed Production
Desert-willow typically produces good seed crops, although quantitative data are lacking for its seed production. Numerous flowers are produced early in the growing season (see Seasonal Development) [158], when a single tree may produce hundreds of flowers [172]. Subsequently, numerous seeds are contained in each capsule [43,161]. Flower and fruit production are positively associated with precipitation; plants may not produce flowers in a very dry year [51] (see Plant Growth). Fruit set may be limited by insufficient amounts of outcrossed pollen (e.g., inadequate movement of pollinators among individual plants) [158].

Seed Dispersal
The seeds are dispersed by wind [14] and water [101]. The seed hairs (fig. 4) aid dispersal [14].

Seed Banking
Desert-willow has a transient [122,127], soil-stored seed bank [50,127]. Viable desert-willow seed was collected from soil in southern Nevada for the Millennial Seed Bank Project [50], although details of collection were not provided.

Germination and Seedling Establishment
Desert-willow seeds are not dormant [127,128,180,254] and nonrefractory (Fryer 2022, this publication). They probably remain viable only until late in the spring following dispersal [128]. Fresh seeds germinate “readily” [220], although a short period of afterripening enhances germination [127]. Germination rates are generally high, ranging from about 50% [232] to 80% [127]. In a laboratory test, fresh seeds averaged 70% germination in the dark, with germination reduced to 15% for seeds dry-stored for 6 months [122].

A moist substrate is required for germination, and flooding and light shade promote germination and recruitment of desert-willow [141,191,200,243]. During floods, desert-willow seeds often lodge under boulders, which provide shade for emergents [38]. Desert-willow may also establish near its own canopy [141].

Plant Growth
Seedling growth is rapid under favorable conditions [114,126,180]. Desert-willow seedlings may grow as much as 1 m/year if water is not limiting [220]. In the greenhouse, desert-willow seedlings averaged 20.6 cm tall 20 days after emerging, and roots averaged 15.0 cm long [180]. In Lubbock, Texas, desert-willow seedlings planted in gravel averaged 815 g dry weight at initial planting and 1,093 g dry weight at 1 year old. Height gain after 1 year averaged 106 cm [105].

Desert-willow's rate of photosynthesis is relatively high [199], suggesting that it maintains relatively rapid growth rates. A study near Las Cruces, New Mexico, showed desert-willow maintains relatively constant photosynthetic rates and water potentials despite either drought or rapid fluctuations in water availability [46].

Growth rate in a wet year may be double that in a dry year. On an arroyo on the New Mexico State University's Agricultural Experiment Station, the mean number of new desert-willow shoots/plant was 440 over 4 years, with a range from 72 to 960 shoots/plant. The rate and amount of growth was strongly associated with available moisture, based on precipitation. Lowest production of leaf biomass (3 kg/ha) occurred in a year (1971) with a dry spring and relatively low summer precipitation. Almost no fruits were produced that year. The next growing season (1972–1973) had relatively continuous moisture from fall through summer. This resulted in the largest peak of leaf (9 kg/ha) and fruit (2.5 kg/ha) biomass production in the 4 years of study [51].

Dams limit the extent of flooding and germination of riparian trees within the floodplain, and they lower the water table in areas away from the active channel. Thus, seedlings and established trees are more likely to die. Changes to peak discharge timing also limit reproduction of riparian species [192].

Vegetative Reproduction and Regeneration
Desert-willow sprouts from the root crown [18,218,220] after top-kill by fire [18,218], flood damage [18], or cutting [119]. It may also layer [57].

Mortality
Drought causes mortality of riparian species such as desert-willow, which depend on floods and groundwater for survival and growth [192]. Seedling and sprout mortality may be high during hot, dry periods and/or when water tables drop [243]. No further information was available on mortality of desert-willow except in the context of fire.

SUCCESSIONAL STATUS
Desert-willow is most common in open areas [137], although it is somewhat shade tolerant and can establish near parent plants [141].

Flooding is the primary disturbance in mixed-deciduous riparian communities of the Southwest [67,191,200,243]. Due to erosion and deposition during floods, these communities often experience large-scale disturbances that set back succession [30,134,243]. Early-successional stages are common in mixed-deciduous woodlands [30,243], and trees on heavily scoured sites may never reach maturity [243] (i.e., the stem exclusion stage). In the Sonoran Desert in western Chihuahua, Mexico, sporadic floods on the lower banks of major rivers with scrubby desert-willows are described as "torrential" [156]. Trees on upper banks of such riparian areas are most likely mature [243], while trees on lower banks may be dislodged in floods.

Desert-willow also establishes in early postfire environments (see Fire Adaptations and Plant Response to Fire). Fire is becoming the primary disturbance in riparian ecosystems due to changes in hydrology (see Fire Management Considerations), climate, and plant community composition and structure (i.e., fuels) [245] (also see Fire Regimes).

Because it is unpalatable and associated species are preferred, desert-willow cover may increase with overgrazing [64,138]. In Coahuila, Mexico, the importance value of desert-willow was higher on sites with livestock (cattle, domestic goats, donkeys, and horses) than on sites without livestock [64]. However, near Little Ash Creek on the Prescott National Forest, Arizona, desert-willow was present only in livestock exclosures [209], suggesting that trampling and/or browsing of desert-willow had occurred in unprotected areas.

FIRE ECOLOGY AND MANAGEMENT


IMMEDIATE FIRE EFFECTS
Low- to moderate-severity fire prunes or top-kills desert-willow [12,18,218,219,244]. Its "very thin" bark [91] provides little protection from fire. Although desert-willow typically sprouts after top-kill by fire (see below), it may not survive severe or repeated fire [12,244].

Postfire Regeneration Strategy
Tree with a sprouting root crown
Tall shrub with a sprouting root crown [198]

Other Possible Strategies:
Ground residual colonizer (on site, initial community)
Initial off-site colonizer (off site, initial community)
Secondary colonizer (on- or off-site seed sources) [198]

FIRE ADAPTATIONS AND PLANT RESPONSE TO FIRE
Desert-willow sprouts after top-kill by fire, and it may also establish from seed. Most riparian species are adapted to sprouting after damage from flooding; similarly, aboveground damage from fire can also trigger production of new shoots [204]. Desert-willow typically sprouts from the root crown after top-kill from low- to moderate-severity fire [12,18,218,219,244]. Postfire sprout mortality may be high during hot, dry periods and with lowering water tables [243]. Because the seeds are wind- and water-dispersed and have a transient, soil-stored seed bank, desert-willow may establish from seed after fire on moist sites (see Regeneration Processes).

Desert-willow occurs in early postfire environments [12,218,219,245]. Following a July wildfire in sugar sumac-coastal sage scrub oak-hairy yerba santa chaparral in San Diego County, California, more than 90% of desert-willows survived. These survivors sprouted by postfire month 2. They developed a multistemmed growth form and averaged 171 sprouts/plant by postfire month 10 (table 3) [219].

Table 3—Postfire growth of desert-willow after the July 1975 San Ysidro chaparral wildfire in Anza-Borrego Desert State Park, California. Data are means. Adapted from Tratz (1978) [219].
Postfire month Plants/ha Sprouts/plant Sprout length
(cm)
Aboveground mass/plant (g)
2 (Sept.) 30 16 5.5 2
4 (Nov.) 10 21 27.1 40
7 (Feb.) 30 48 47.6 132
10 (June) 18 171 49.3 892

In 2003–2004 and again in 2012–2013, Bock and Bock (2014) measured the condition of riparian trees at the Appleton-Whittell Research Ranch, Arizona, in three canyons above the San Pedro River that had experienced wildfire in 2002, 2009, or both. Fire killed some desert-willows, but most were top-killed and sprouted. For desert-willows on sites that burned once, 10% survived as mature trees, 5% died, and 85% were partially to completely top-killed and sprouted. For desert-willows on sites that burned once or twice, 15% survived as mature trees, 20% died, and 65% were partially to completely top-killed and sprouted [12]. See the Research Project Summary of this study for more information on the response of desert-willow and associated trees (Arizona sycamore, Arizona walnut, Fremont cottonwood, and velvet ash) to these fires.

FUEL CHARACTERISTICS
Historically, fuels in low-elevation deserts of the Southwest were likely mostly sparse and discontinuous, although information on this is limited [60,109,243]. Keane et al. (2000) provide crown bulk density estimates for various structural stages of riparian forests and several other plant communities on the Gila National Forest. See the publication for further information [99].

Desert-willow communities typically have a moderate to deep litter layer, although litter decay rates may be fast. Desert-willow communities codominated by willow typically have deep litter, mostly due to the accumulation of willow leaves [155]. However, on the Jornada Experimental Range, New Mexico, sites dominated by desert-willow had loose litter that averaged 1 cm thick [95]. Over 6 months in Las Cruces, New Mexico, desert-willow leaf litter decayed faster on the soil surface than when buried 10 cm deep. Both surface and buried litter decayed faster in summer than in winter or spring. Schaefer et. al (1985) present a model for predicting rates of litter decay for desert-willow and associated shrub species [179].

Information on the flammability of desert-willow was lacking in the literature, but a few publications provide measures of live desert-willow fuels. Leaf area index is used in fuel models to help predict fire behavior and severity [212]. Studies in the Chiricahua Desert (location not provided) found desert-willow had lower mean specific leaf area (4.30 m²/kg) than surrounding desert shrub species [231], suggesting that it has less canopy fuel than some associated species. In southern New Mexico, surface area of desert-willow leaves averaged 3.1 cm², and specific leaf mass averaged 12.5 mg/cm² [102].

Climbing ladder fuels are present on some sites with desert-willow. For example, canyon grape often occurs in communities where desert-willow is dominant or important [7,140,207], and it can grow into upper canopies. At the Fort Bowie National Historic Site, Arizona, canyon grape formed continuous fuels from surface vegetation to the upper canopy of an Arizona walnut-netleaf hackberry-gum bully riparian woodland in which desert-willow was an associate [239].

Nonnative species can alter fuel distributions and loads. When riparian vegetation composition and structure change as a result of nonnative species in the overstory and/or understory, fuel characteristics may also change. In turn, this can alter fire intensity, severity, and/or rates of spread [243] (see Fire Regimes and Management Under a Changing Climate). Near Yuma, Arizona, fine fuels helped carry and spread the 2005 King Valley Fire, which burned a desert ironwood-blue paloverde-yellow paloverde-desert-willow community. The fine fuels were mostly Mediterranean grass (nonnative annual grasses) and desert Indianwheat (a native annual forb). These herbs had grown at high densities in response to the extremely wet winter of 2004–2005 [221].

FIRE REGIMES
In shrub and mixed-deciduous riparian communities of the Southwest and northern Mexico, wildfires are ignited by lightning in June and July [9]. Contemporary fires often spread into riparian areas from adjacent uplands [12,40], but fires can also start in riparian zones [243]. It is uncertain how frequent fires were within riparian areas of these deserts historically. Fires in small drainages likely had different frequencies and behaviors than fires in larger drainages where riparian zones were wider and vegetation more developed [12,160]. Fire intervals may be long and severity low in washes and riparian mixed-deciduous woodlands where periodic flooding scours the understory, limiting buildup of fine fuels and litter [243,244].

Information on historical fire regimes in riparian areas of the Southwest is limited due to lack of data [72,191,244], and the variability of historical fire regimes within and among desert riparian areas is not well known. The dynamic nature of riparian ecosystems makes their fire regimes hard to characterize [243,244]. Historically, sparse surface fuels and the variable patchiness of desert plant communities probably limited fire size, severity, and intensity, particularly in riparian zones [60,109,243]. Webb (2017, 2019) suggests that fires varied in severity and extent across southwestern riparian ecosystems [243,244]. Most fires were likely patchy and of low severity due to the patchiness of vegetation in desert landscapes; however, mixed- and high-severity fires likely occurred as well [111,243,244]. Fire might have been historically uncommon in some riparian areas [60,72,192,194,243,244]. In many cases, fires that started in the uplands might have slowed or stopped in riparian zones because temperatures were cooler and live fuel moistures higher [18,60,72,243]. Brooks et al. (2018) report that prior to the 1940s, ranchers who tried to burn in desert shrublands and Joshua tree woodlands were mostly unsuccessful due to fuel discontinuity [18].

In many areas, stand structure and function of mixed-deciduous riparian woodlands have been significantly altered since European-American exploration and settlement began [243]. Before that time, American Indians likely burned riparian zones, although the extent of such burning is unknown. Archeological studies along the Salt-Gila Aqueduct in southern Arizona revealed desert-willow charcoal deposits near prehistorical Hohokam settlements, and the researchers suggested these deposits were remnants of prehistorical agricultural burning [132,133]. Riparian areas experienced rapid ecological change [47,243,244] during European-American exploration (1540-1821) and settlement (1822-1912) periods [47]. In Arizona, for example, Spanish explorers imported livestock to the San Pedro Valley as early as the 1500s; European-Americans cut timber throughout the settlement period; miners diverted water from the San Pedro River to process ore in the 1800s; and fur trappers hunted American beavers until the animals were extirpated from the valley [243,244].

Changes in hydrology and vegetation have undoubtedly affected fire regimes of southwestern riparian communities and the surrounding uplands. However, the degree to which fire regimes are changing—or what the effects might be for riparian ecosystem structure, function, and resilience—are not well known [243,244]. The greatest ecological change in riparian areas since large-scale livestock ranching was introduced is probably a shift in fire regimes from mostly patchy surface fires to larger crown fires [5,243]. Some research suggests that fire frequency and severity are increasing in some southwestern riparian ecosystems [26,27,59,62,72,90,204,206,244], although the extent of these increases are largely unknown. Wildfire became a significant disturbance agent in southwestern riparian areas at the end of the 20th century. With the reduction in frequency and magnitude of floods, litter and debris accumulated in the forest understory. This accumulation, along with increased density of native and nonnative vegetation, resulted in greater fuel loads, fire sizes, and fire intensities than existed prior to riparian modification (review by [192]).

Factors that can increase fire severity include drought, altered flood disturbance, anthropogenic ignitions in high-use areas [18,26,59,62,206,243,250], flood and fire exclusion and associated high fuel loads, and more continuous fuels [18,59,62,72,77,243,250] resulting, in part, from spread of nonnative plant species [18,26,59,206,243]. Severe or intense fires can induce uniform changes in structure and composition across riparian zones by causing widespread mortality of native trees [8,243], increasing the likelihood of postfire establishment of nonnative plants [72,243,244]. Severe fires may be less common in areas with natural flood regimes [62,243,244].

Drought and/or human engineering can cause changes in hydrology (e.g., surface flows, groundwater levels, and channel and floodplain morphology) that favor nonnative plant species over native ones [27,59,72,90,169,244,257]. Nonnative annual grasses—particularly red brome, Mediterranean grass, and cheatgrass [17,18]—and nonnative trees Russian-olive [256] and tamarisk (saltcedar and five-stamen tamarisk [17,255], and potentially Athel tamarisk [234] and French tamarisk [29]) can increase fuel continuity and fire frequency in riparian communities of the Southwest [17,18] (see Fire Management Considerations). Fires may be larger, more frequent, and more severe in invaded communities. For example, fire-induced mortality of native riparian trees can increase dramatically when tamarisk occurs in the prefire plant community [244]. On 30 sites across the Southwest, high fire consumption and postfire mortality of Fremont cottonwoods and willows were positively associated with high prefire tamarisk cover. Fire was more severe where tamarisk was present than where it was not, resulting in more tissue damage to native trees. When prefire tamarisk cover was >50%, fire completely consumed the fine tissues of all native trees present [59].

Presettlement fire regimes of mixed-deciduous riparian communities of the Southwest likely varied among sites [243,244]. LANDFIRE (2008) models classify fire regimes of Biophysical Settings dominated by cottonwood-willow communities in the Southwest into two groups: Fire Regime Group I (<35-year return intervals, mixed severity) and Fire Regime Group V (>200-year fire return intervals, replacmente severity). Communities in the frequent-fire group are generally those along larger rivers, and those in the infrequent fire group are generally smaller desert riparian areas (i.e., stringers) [111].

See these FEIS publications for information on historical fire regimes in plant communities in which desert-willow is common to dominant: See table A3 for a list of additional plant communities in which desert-willow is common to dominant in wash zones, and search FEIS for fire regime information on those communities.

FIRE MANAGEMENT CONSIDERATIONS
There is limited research on the impacts of fire on southwestern riparian communities [244]. Dams, waterway diversions, and climate change have greatly altered the hydrology of many streams and rivers. In the absence of natural flooding and with lowering water tables, fire is becoming the dominant disturbance in some riparian ecosystems of the Southwest. Groundwater depth and streamflow greatly influence vegetation type and extent in riparian ecosystems [26,77,90,243], and fire regimes seem to be changing in some lowland riparian communities with altered hydrology. Changing fire regimes are likely to have drastic, potentially irreversible effects on biodiversity and function of affected riparian communities. Alterations in plant community structure and plant and animal community composition may occur, including declines in wildlife species of concern [244]. Desert-willow may not survive severe, intense, or very frequent fire [12].

More severe, intense, and/or frequent fire can interact with altered hydrology to accelerate the replacement of native plants with nonnative plants [118,243]. In many riparian areas in the Southwest, there have been increases in tamarisk, Russian-olive, Mediterranean grass, and other nonnative plants that have different fuel characteristics, fire tolerances, and postfire fire responses than native species. Establishment and spread of tamarisk [170,255] and other nonnative species have changed fuel and fire regime characteristics in many riparian areas, and increased fire risk has been associated with high cover of tamarisk and Russian-olive [26,59,142,169,174,243,244]. Where groundwater levels have dropped, tamarisk has a competitive advantage over cottonwoods [118,243]. Its higher tolerance to drying soils [196] and frequent fire are driving changes in riparian stand structure and fire regimes, resulting in decreasing cover of formerly dominant, native trees such as desert-willow, cottonwoods, and willows [244,255]. Fire intervals in mixed-deciduous woodlands may be as frequent as 15 years in areas where tamarisk cover is high [59,110,163,243,244]. Where natural flow regimes are more intact and tamarisk is suppressed by native vegetation, desert-willow and other native trees are more likely to dominate postfire landscapes [244].

Fire management in southwestern riparian areas may include fuel reduction and prescribed fire treatments that promote or maintain native vegetation [244] and reduce or eradicate invasives [187,243]. Although simulated fuel reduction treatments showed potential to mitigate fire severity in some mixed-deciduous woodlands and forests under projected climate change in the Huachuca Mountains, projected basal area and mortality rates in riparian zones were unchanged by simulated fuel treatments, and a reduction in basal area and spatial extent of riparian species occurred with or without fire and regardless of treatments [151]. Prescribed fire may be most effective when used in combination with other restoration, such as restoring natural waterway flows [244]. However, little information was available on using prescribed fire in riparian plant communities of the Southwest [9]. The effects of fire, including prescribed burning, have been better studied in southwestern grasslands and savannas than in adjacent riparian woodlands and forests [12,60,72,191], so more information is needed on the use of prescribed fire and postfire restoration in riparian areas. Fire may be severe, and lethal to overstory deciduous trees, in riparian woodlands that have not experienced fire for many decades.

Bock and Bock (1990) do not recommend prescribed fire in late-successional riparian woodlands of the Southwest because "fire is difficult to manage and potentially very destructive" in those habitats [11]. Refined fuel and fire behavior models are needed for riparian ecosystems of the Southwest, including models for areas in which tamarisk has become dominant. Webb et al. (2019) review the information available for using prescribed fire in southwestern riparian areas, with and without invasive species [244].

OTHER MANAGEMENT CONSIDERATIONS


FEDERAL LEGAL STATUS
None [225]

OTHER STATUS
Desert-willow shrublands are ranked Vulnerable (G3, S3) [144,146,159]. Other information on state- and province-level protection status of plants in the United States and Canada is available at NatureServe.

IMPORTANCE TO WILDLIFE AND LIVESTOCK
A myriad of wildlife species use riparian habitats [48,49,81,124,134,237,241,244]. Riparian areas comprise a small part of total land area in the southwestern United States and northern Mexico, but they are extremely important to biotic communities [244]. Riparian areas are among the most diverse and productive ecosystems in desert bioregions, supporting floras and faunas that are biologically richer than adjacent uplands [21,32,35,244]. Riparian areas and washes provide water, cover, and breeding habitat for many species of wildlife, including obligate riparian species [70,134]. For wildlife that also use uplands, riparian areas provide refuge from the harsher, surrounding environments [134]. Riparian areas and washes also provide water, cover, and breeding habitat for livestock [24,70,134].

Wildlife that use riparian ecosystems for breeding, migration, and wintering habitat include large ungulate [48,244], small mammal, bird [2,48,81,168,205,241], reptile, amphibian, and arthropod species [22,25,48,168,244]. For example, On the Stockton Plateau of Texas, Arizona walnut-desert-willow communities provide habitat for white-tailed deer, ringtails, eastern fox squirrels, and Merriam's pocket mice [86]. Western white-throated woodrats along the Rio Grande use desert-willow wash habitats [253]. On the New Mexico State University's Agricultural Experiment Station, most (54%) of the litter and soil fauna collected under mature desert-willows were mites. Springtails were also common [95].

Riparian ecosystems provide habitat for many threatened or endangered species. Of 40 Federally listed Threatened or Endangered wildlife species in New Mexico—which includes several endemic species—at least 70% require aquatic and/or riparian habitat to feed, reproduce, and/or carry out their life cycles [222,244]. See these sources for checklists of birds that may use riparian communities with desert-willow for breeding and nesting habitat: [2,81,168,197,205]. Reynolds (1964) provides checklists of mammals and fish that may use riparian/aquatic habitats with desert-willow [168].

Communities with desert-willow provide important breeding, nesting, and migrant habitat for a wide variety of bird species [48,104,197,205]. For example, cactus wrens use desert-willow habitats for nesting and the fluffy desert-willow seeds for nesting material [135]. On the Barry M. Goldwater Air Force Range, southeastern Arizona, species richness of breeding and passing neotropical migrants is positively associated with high cover of mature (>2.5 m) desert-willow, paloverde, mesquite, and catclaw acacia. Migrant hummingbirds use desert-willow for cover and its nectar as a food source [83].

Many wildlife species browse desert-willow or consume its nectar or litter [22,218]. Large ungulates browse it, although their use may be limited. On the Elephant Mountain Wildlife Management Area in the Chihuahuan Desert, Texas, female bighorn sheep only browsed desert-willow in winter, although males browsed it year-round [15]. Desert-willow leaves and fruit provide minor forage for desert mule deer from spring through fall [25,103,106,186]. Various birds, including Gambel's quail [79,80], eat the seeds [232]. The sucrose in desert-willow nectar is a good energy source [22]. Hummingbirds and various insects, particularly bumble bees, consume the nectar [22]. Sonoran desert tortoises eat the fallen flowers [25]. Fly species feed from the extrafloral nectaries on the leaves [157]. In the Mojave Desert in southern Nevada, Cooper thrips live in and feed on decaying desert-willow litter [250].

Palatability and Nutritional Value
Desert-willow is not highly palatable to wildlife and livestock [45,100,232] because the foliage and bark contain glycosides and phenolic acids [138,252]. Substantial use typically occurs only when more palatable forage is not available [100,232], although wildlife may browse new sprouts after fire [219] or other top-killing disturbances.

See these sources for information on the nutritional value of desert-willow twigs: [33], leaves: [102] and seeds: [94,228].

Cover Value
Plant communities with desert-willow provide cover for many wildlife species [108,127]. They provide birds with hiding, resting, and nesting sites and foraging opportunities, some of which are absent or rare in adjacent plant communities [192]. Bird species using these communities include ferruginous pygmy-owls [173], northern bobwhites [82,88], Gambel’s quail [79,93], road runners [93], ladder-backed woodpeckers [93,95], hummingbirds [83], and many passerines [93] (e.g., gray vireos [249] and summer tanagers [205]).

High-quality fish habitat depends on suitable water temperatures for spawning and survival [48,78]. Desert-willow provides shade cover for fish, including the Federally Endangered [225] Rio Grande silvery minnow [41], and the Apache and Gila trout [48].

Burned areas create habitat for some wildlife guilds. They tend to increase hunting opportunities for predators that prefer hunting in open areas such as coyotes [36], common black hawks [178], and peregrine falcons [192]. Mixed-severity and severe fires create wildlife snags for cavity-nesting species [192].

VALUE FOR REHABILITATION OR RESTORATION
Desert-willow is used for erosion control [91,98,100,119,127], stream stabilization, and revegetation [89,127]. Because of the relatively fast growth rates of desert-willow and other riparian species, riparian communities generally recover faster than upland plant communities after restoration is implemented [134].

Desert-willow is propagated from seed, cuttings [58,119,127,220], and mound layering [57]. Cultivars are available [105,189,216,220]. See these sources for information on propagating desert-willow: [37,105,127].

OTHER USES
Riparian areas with desert-willow provide many ecosystem services including recreational opportunities, harboring pollinators, providing livestock forage [190,244], and enhancing water quality [244]. Riparian vegetation reduces nonpoint source pollution in waterways by filtering sediments, chemicals, and nutrients from overflow. The roots of streamside plants help prevent excessive erosion during floods [223] and promote groundwater infiltration [66], and vegetation surrounding active channels helps slow water velocity during floods [223]. Riparian trees and shrubs shade surface water, reducing water temperatures [223] and lost to evaporation [244].

Several products or potential products involve desert-willows. Desert-willow is planted as an ornamental [43,63,100,127,194,195,220] and used in shelterbelts [82]. The flowers are a bee food from which good honey is made [232]. Desert-willow can uptake gold from gold-enriched media, so it has potential as a gold mining plant [175]. Desert-willow tissues are being investigated for use as medicines. The foliage and bark have antibacterial and antifungal properties [138]. Fatty acids in the seeds may have anticarcinogenic and antidiabetic properties, as well as properties that may promote immune system and heart health [228].

Desert-willow wood provides timber, fenceposts, and firewood [91,98,119,148,161,220,232]. The wood is durable but not strong [91], and it has no commercial value [237]. See these sources for information on the properties of desert-willow wood: [152,153].

Native peoples have many traditional uses for desert-willow. They use the wood for house frames, granaries, and bows; the fibrous bark for weaving nets, shirts, and breechcloth [127]; and the branches for making baskets [220,232] and bows [188]. The flowers and seed capsules are eaten [127].

ADDITIONAL MANAGMENT CONSIDERATIONS
The value of southwestern riparian woodlands for both biotic communities and human livelihoods is well established [48,49,87,143,192,201,229], and maintaining and/or restoring riparian areas of the Southwest is a critical management issue [48,49,131,192]. Management goals for southwestern riparian areas may include floodplain restoration, safeguarding high-quality riparian areas, fuel reduction treatments, preventing spread of fire, and preventing establishment and spread of nonnative invasive plants [187,243].

Many riparian communities of the Southwest are highly imperiled due to anthropogenic disturbances [190]. Few naturally functioning riparian areas remain in the region, and those that do are threatened with climate change [134,244]. Within the last 100 to 150 years, humans have modified many riparian areas extensively by irrigating, pumping groundwater, building dams and diversions, clearing floodplains, cutting wood, OHV use [69], grazing livestock [23], and spreading nonnative invasive plants. These modifications have resulted in significant changes in the physical environment and biota [64,134,136,202,244]. For example, livestock generally use riparian areas more than adjacent, less productive upland communities, compacting soils and reducing recruitment of palatable deciduous seedlings and saplings into the overstory [44,134]. Use of the relatively unpalatable desert-willow by livestock generally indicates overbrowsing or overstocking [227,240].

Altered hydrological regimes are often associated with establishment and spread of nonnative plants. Altered hydrology and sediment deposition resulting from river impoundment may reduce or eliminate germination and recruitment of native deciduous trees while promoting germination and recruitment of tamarisk [244]. Consequently, tamarisk species have established in many riparian areas of the Southwest [4,134,170,203,204,255], including wash and woodland communities in which desert-willow was formerly dominant or important [20,123,125] (see Fire Management Considerations). Maintaining an ecologically healthy riparian community can help stop establishment and spread of tamarisk and other invasive plant species [3]. Dense shade in closed-canopy communities with desert-willow may inhibit establishment of tamarisk [52].

Flow regimes that fall within the natural range of variability for undammed waterways can have multiple benefits, including:

MANAGEMENT UNDER A CHANGING CLIMATE
Southwestern riparian ecosystems are vulnerable to decline under hotter and drier conditions [31,201,243,244]. Climate change and variation—and associated impacts on hydrology, channel morphology, and riparian vegetation—may alter fire regimes by increasing the likelihood of more frequent or severe wildfires [12,60,193,244]. For example, prolonged drought increases fire risk by decreasing water availability and reducing fuel moisture [160,243,244].

Changes in water flow are expected with climate change, including lower water tables and increased frequency and severity of extreme weather that will likely result in more frequent and severe floods as well as more intense droughts [61,72,181,191,192,243]. Warming in montane areas may increase the number of severe floods downstream by increasing the rate of snowmelt into downstream waterways such as the Middle Rio Grande [243]. Monsoons may occur later in the year and with greater severity [192]. Droughts are likely to be increasingly severe and prolonged, resulting in further reductions in discharge volume [192].

Distribution and composition of upland communities may change with climate warming [70,243,244]. In turn, this may affect composition of riparian vegetation in ways that increase fire risk (e.g., facilitating spread of nonnative species [191]). Climate change could alter competitive relationships between desert-willow and other native species, and also between desert-willow and nonnative species such as tamarisk and Russian-olive [142]. For example, if flood timing does not coincide with spring dispersal of desert-willow, cottonwood, and willow seeds, reproductive success of these native trees is likely to decrease while that of nonnative trees is likely to increase [71,191].


APPENDIX


Table A1—Common and scientific names of plants mentioned in this review. Links go to FEIS Species Reviews.
Common name Scientific name
Cacti
cactus Cactaceae
Forbs
desert Indianwheat Plantago ovata
Graminoids
Arizona muhly Muhlenbergia arizonica
blue grama Bouteloua gracilis
cheatgrassa Bromus tectorum
Mediterranean grassa Schismus spp.
Hall's panicgrass Panicum hallii
red bromea Bromus rubens
sideoats grama Bouteloua curtipendula
tobosagrass Pleuraphis mutica
Lianas
canyon grape Vitis arizonica
eastern poison-ivy Toxicodendron radicans
Shrubs
Apache plume Fallagia paradoxa
baccharis Baccharis spp.
blackbrush Coleogyne ramosissima
burrobush Ambrosia dumosa
catclaw acacia Senegalia greggii
coastal sage scrub oak Quercus dumosa
creosote bush Larrea tridentata
desertbroom Baccharis sarothroides
desert ironwood Olneya tesota
eastern Mojave buckwheat Eriogonum fasciculatum
gum bully Sideroxylon lanuginosa
hairy yerba santa Eridictyon trichocalyx
honey mesquite Prosopis glandulosa
netleaf hackberry Celtis laevigata var. reticulata
mesquite Prosopis spp.
paloverde Parkinsonia spp.
screwbean mesquite Prosopis pubescens
singlewhorl burrobrush Hymenoclea monogyra
splitleaf brickellbush Brickellia laciniata
smoketree Psorothamnus spinosus
spiny hackberry Celtis ehrenbergiana
sugar sumac Rhus ovata
Texas paloverde Parkinsonia texana
tree tobacco Nicotiana glauca
velvet mesquite Prosopis velutina
yellow paloverde Parkinsonia microphylla
yucca Yucca spp.
Trees
alligator juniper Juniperus deppeana
American sycamore Platanus occidentalis
Arizona sycamore Platanus wrightii
Arizona walnut Juglans major
arroyo willow Salix lasiolepis
Athel tamarisk Tamarix aphylla
black willow Salix nigra
blue paloverde Parkinsonia florida
canyon live oak Quercus chrysolepis
cottonwood Populus spp.
desert-willow Chilopsis linearis
five-stamen tamariska Tamarix chinensis
Fremont cottonwood Populus fremontii
French tamariska Tamarix gallica
Goodding’s willow Salix gooddingii
green ash Fraxinus pennsylvanica
Joshua tree Yucca brevifolia
juniper Juniperus spp.
little walnut Juglans microcarpa
Lombardy poplara Populus nigra
narrowleaf willow Salix exigua
oak Quercus spp.
Parry pinyon Pinus quadrifolia
Russian-olivea Elaeagnus angustifolia
saltcedara Tamarix ramosissima
singleleaf pinyon Pinus monophylla
tamariska Tamarix spp.
Texas madrone Arbutus xalapensis
velvet ash Fraxinus velutina
wavyleaf oak Quercus × pauciloba
western soapberry Sapindus saponaria var. drummondii
white alder Alnus rhombifolia
willow Salix spp.
aNonnative species.

Table A2—Common and scientific names of wildlife species mentioned in this review. Links go to FEIS Species Reviews.
Common name Scientific name
Arthropods
bees Apoidea
bumble bee Bombus spp.
carpenter bees Xylocopinae
Cooper thrip Hoplothrips cooperi
flies Diptera
hawkmoth Hyles spp.
honeybee Apis mellifera
lady-beetles Coccinellidae
mites Trombidiformes
Sonoran bumble bee Bombus sonorous
springtails Collembola
Fish
Apache trout Oncorhynchus apache
Gila trout Oncorhynchus gilae
Rio Grande silvery minnow Hybognathus amarus
Reptiles
Sonoran desert tortoise Gopherus morafkai
Birds
cactus wren Campylorhynchus brunneicapillus
common black hawk Buteogallus anthracinus
ferruginous pygmy-owl Glaucidium brasilianum
Gambel’s quail Callipepla gambelii
gray vireo Vireo vicinior
ladder-backed woodpecker Dryobates scalaris
road runner Geococcyx spp.
hummingbirds Trochilidae
northern bobwhite Colinus virginianus
passerines Passeriformes
summer tanager Piranga rubra
Mammals
American beaver Castor canadensis
bighorn sheep Ovis canadensis
desert mule deer Odocoileus hemionus eremicus
eastern fox squirrel Sciurus niger
Merriam's pocket mouse Perognathus merriami
ringtail Bassariscus astutus
western white-throated woodrat Neotoma albigula
white-tailed deer Odocoileus virginianus

Table A3—Representative plant community classifications in which desert-willow occurs.
FRES Ecosystems
FRES30 Desert shrub
FRES32 Texas savanna
FRES33 Southwestern shrubsteppe
FRES 35 Pinyon-juniper
FRES38 Plains grasslands
FRES40 Desert grasslands [74]
Kuchler Plant Associations
K023 Juniper-pinyon woodland
K027 Mesquite bosque
K031 Oak-juniper woodlands
K041 Creosotebush
K042 Creosotebush -bursage
K043 Paloverde-cactus shrub
K044 Creosotebush-tarbush
K054 Grama-tobosa prairie
K058 Grama-tobosa shrubsteppe
K059 Trans-Pecos shrub savanna
K061 Mesquite-acacia savanna [107]
SAF Cover Types
222 Black cottonwood-willow
235 Cottonwood-willow
239 Pinyon-juniper
242 Mesquite [65]
SRM (Rangeland) Cover Types
203 Riparian woodland
211 Creosote bush scrub
212 Blackbush
412 Juniper-pinyon
422 Riparian
502 Grama-galleta
504 Juniper-pinyon pine
506 Creosotebush-bursage
507 Paloverde-cactus
508 Creosotebush-tarbush
707 Blue grama-sideoats grama-black grama
729 Mesquite
735 Sideoats grama-sumac-juniper [185]

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