T
T
T
Fire Regime - Synthesis

Fire regimes of Sonoran desert scrub communities

Written
October, 2023
Contributors
Kris Zouhar - 1st Author, Robin J. Innes - 1st Editor, Ilana L. Abrahamson - 2nd Editor
Fire Regime Type
Synthesis

Zouhar, Kristin. 2023. Fire regimes of Sonoran desert scrub communities. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Missoula Fire Sciences Laboratory (Producer). Available: https://research.fs.usda.gov/feis/fire-regimes/sonoran-desert-scrub

The desert scrub communities covered in this synthesis are represented by three Biophysical Settings—Sonoran paloverde-mixed cacti desert scrub, Sonoran granite outcrop desert scrub, and Sonoran mid-elevation desert scrub. These communities are unique to the Sonoran Desert, have many species in common, are often adjacent to one another, and have similar, clumped and patchy canopies of small trees, tall shrubs, and large cacti. They are also thought to have similar historical fire regimes, characterized by rare or infrequent, high-severity fires.

Estimates of historical fire regime characteristics are based on observations of postfire responses of dominant plants, as well as stand structures and fuel characteristics of contemporary, undisturbed communities. Because many of the dominant, long-lived woody and succulent plants in Sonoran paloverde-mixed cacti desert scrub communities are easily killed or injured by fire, and few are adapted to regenerate in the postfire environment, fires were thought to be historically rare (mean fire interval (MFI) >1,000 years) in those communities. Sonoran mid-elevation desert scrub communities have a larger proportion of resprouting species and are often adjacent to communities with frequent fire historically, so fires are thought to have been more common, but still infrequent (MFI >100–374 years) in those communities. Estimates of fire frequency in Sonoran granite outcrop desert scrub are intermediate to these (MFI >500 years), and no information was found in the literature that was specific to those communities. LANDFIRE classifies historical fires in all three communities as stand-replacing, presumably based on a lack of fire adaptions in dominant species. However, historical fires were probably small and/or patchy, due to patchy stand structure and discontinuity of fuels. Historically, large fires (>50 ha) may have occurred infrequently, but only when extreme fire weather coincided with the presence of adequate fine fuels.

Contemporary fire regimes are likely outside the historical range of variability in Sonoran desert scrub communities. Frequent, large, and severe fires in Sonoran desert scrub are attributed mostly to increased human access and subsequent human-caused ignitions, and to increases in receptive fine fuel loads and continuity caused by nonnative plant invasions. Nonnative plant invasions can increase the likelihood of fire ignition and spread, which may result in larger patches of high-severity effects. Rapid recovery of herbaceous fuels—especially nonnative grasses—may lead to repeated fires on a site. Repeated and/or large patches of high-severity fire can lead to vegetation type conversions that may persist indefinitely.

Sonoran desert scrub communities are not resilient to frequent, large, or severe wildfires, and native, prefire vegetation may not recover for centuries, if at all. Therefore, fire prevention is critical. Fire prevention includes identifying areas where large fires are likely to occur and reducing fire risk by managing nonnative invasive plant fuels or implementing other actions to help prevent fire ignition and spread.

Landscape view of several Sonoran desert scrub communities.
Photo Credit
A) Photo © Burley Packwood, some rights reserved (CC BY-SA). B) Photo © inceptis, some rights reserved (CC BY-NC). C1) Photo © Eric Hough, some rights reserved (CC BY-NC). C2) Photo © Eric Hough, some rights reserved (CC BY-NC).

Figure 1—Examples of Sonoran desert scrub communities covered in this synthesis. A) Sonoran paloverde–mixed cacti desert scrub dominated by saguaro. B) Sonoran granite outcrop desert scrub dominated by elephant tree. C1) Sonoran mid-elevation desert scrub dominated by jojoba. C2) Sonoran mid-elevation desert scrub dominated by crucifixion thorn.

This Fire Regime Synthesis brings together information from two sources: the scientific literature as of 2023, and the Biophysical Settings (BpS) models and associated Fire Regime Data Products developed by LANDFIRE, which are based on literature, local data, and/or expert estimates [71]. The information in this synthesis supplements that provided by FEIS Species Reviews and can assist with revisions or modifications to LANDFIRE models. This synthesis summarizes what is known about historical (i.e., presettlement) and contemporary fire regimes in the following Sonoran desert scrub BpSs: paloverde-mixed cacti, granite outcrop, and mid-elevation desert scrub (e.g., fig.1). “Contemporary” generally refers to the time after Euro-American settlement, although fires were not widely reported in the Sonoran desert until the latter part of the 20th century. In this synthesis, “Sonoran desert scrub” refers only to these three communities, which are mapped only in the Sonoran desert. Other desert scrub communities that occur in the Sonoran and other deserts—such as those dominated by creosotebush, mesquite, blackbrush, or saltbush—are not included here. For information on fire regimes in those communities, see the following Fire Regime publications:

  • Creosotebush–white bursage desert scrub
  • Mesquite scrub and woodland
  • Mojave mid-elevation mixed desert scrub
  • Saltbush desert scrub

Sonoran desert scrub communities were grouped together for this synthesis because they are unique to the Sonoran Desert, have many species in common (although characteristic species differ among them), have similar stand structure, are often adjacent to one another on the landscape, and are thought to have similar historical fire regimes, characterized by rare or infrequent, high-severity fires (see the LANDFIRE Biophysical Settings section). With the exception of paloverde-mixed cacti desert scrub, they are also not typically identified separately in the fire literature.

While the focus of this synthesis is specifically Sonoran desert scrub communities, much of the literature on fire regimes and fire history in warm deserts of North America approaches these topics at a larger scale (e.g., all or part of the Sonoran Desert or the Mojave and Sonoran deserts combined) and does not always distinguish among plant communities or types of “desert scrub” covered. Whenever possible, information specific to Sonoran desert scrub communities was extracted from these studies, which often focused primarily on paloverde-mixed cacti desert scrub, due to the concern for preserving the iconic saguaro cactus and its habitat. References in the literature to "Arizona Upland" vegetation were assumed to refer to areas of paloverde-mixed cacti and/or mid-elevation desert scrub communities, based on the descriptions of the Arizona Upland subdivision by Shreve and Wiggins (1964) [116] and Turner and Brown (1982) [125]. Granite outcrop communities were not specifically identified in any of the fire literature examined. Some publications make inferences about Sonoran desert scrub communities based on observations from similar desert plant communities (e.g., Sonora-Mojave creosotebush–white bursage desert scrub, BpS series 10870; hereafter creosotebush–white bursage desert scrub). The areas and plant communities covered by the studies cited in this synthesis are identified in the text as much as possible, to clarify the potential scope of the results or observations.

Much of the information in this synthesis comes from LANDFIRE (2020) BpS descriptions and models [71], as well as from literature reviews or syntheses in other publications, particularly those shown in table 1. LANDFIRE (2020) BpS descriptions and models [71] are summarized in the LANDFIRE Biophysical Settings section of this review.

Table 1—Publications with literature reviews or syntheses that are frequently cited in this synthesis, organized by the main topics covered.
Main topicsTitleReference
Site characteristics and plant communitiesInternational ecological classification standard: Terrestrial ecological classifications.NatureServe (2009) [90]
Site characteristics and plant communitiesBiotic communities of the American Southwest—United States and Mexico: Sonoran desertscrub.Turner and Brown (1982) [125]
Fire regimesFire in California's ecosystems: Southeastern deserts bioregion.Brooks et al. (2018) [28]
Fire regimesThe effects of fire on Sonoran Desert plant communities.Alford (2001) [7]
Fire regimesMortality of burned Cereus giganteus.Rogers (1985) [107]
Fire regimesEffects of wildfire on a Sonoran Desert plant community.McLaughlin and Bowers (1982) [84]
Invasive plants and fire regimesContrasting geographic patterns of ignition probability and burn severity in the Mojave Desert.Klinger et al. (2021) [67]
Invasive plants and fire regimesFire and nonnative invasive plants in the Interior West bioregion.Rice et al. (2008) [104]
Invasive plants and fire regimesAlien annual grasses and their relationships to fire and biotic change in Sonoran desert scrub.Esque and Schwalbe (2002) [48]
Fuel characteristics and invasive plantsModelling and mapping dynamic variability in large fire probability in the lower Sonoran Desert of south-western Arizona.Gray et al. (2014) [58]
Fuel characteristics and invasive plantsCreating hotter fires in the Sonoran Desert: Buffelgrass produces copious fuels and high fire temperatures.McDonald and McPherson (2013) [83]

Common names are used throughout this synthesis. For a complete list of common and scientific names of species discussed in this synthesis and links to FEIS Species Reviews, see table A1.

General Distribution

The three Sonoran desert scrub BpSs covered by this synthesis occur in the northern and central Sonoran Desert, from southeastern California eastward throughout southern Arizona (fig. 2) and into northwestern Sonora, Mexico [90]. They occur primarily in the Arizona Upland subdivision of the Sonoran Desert [125] along the northern and eastern edge of the Sonoran Desert from the Buckskin Mountains, Arizona, southeastward to about Altar, Sonora, Mexico [125]. Granite outcrop and paloverde-mixed cacti desert scrub communities also occur in scattered areas within the Lower Colorado River Valley subdivision in the northwestern part of the Sonoran Desert [71,90].

Distribution of Sonoran desert scrub communities based on Biophysical Settings and Existing Vegetation Type. A comparison of the images indicates that LANDFIRE Biophysival Settings maps do not accurately reflect the distribution of paloverde-mixed cacti desert scrub (pink color) or granite outcrop desert scrub  (brown color)—under representing the former and over representing the latter.

Figure 2—Distribution of Sonoran desert scrub communities based on A) LANDFIRE (2020) Biophysical Settings (BpS) data layer [69], and B) LANDFIRE Existing Vegetation Type (EVT) data layer [70]. 

LANDFIRE maps of these BpSs (fig. 2A) do not accurately reflect the distribution of paloverde-mixed cacti or granite outcrop desert scrub communities—under representing the former and over representing the latter. LANDFIRE’s existing vegetation type (EVT) maps seem to reflect the distribution of these potential vegetation types more accurately (Malusa, University of Arizona, personal communication [75]), with only very small areas of granite outcrop desert scrub just north of the U.S./Mexico border, and much larger areas of paloverde-mixed cacti desert scrub (fig. 2B). This is also supported by the description of the granite outcrop desert scrub BpS, which says that it occurs primarily in Sonora, Mexico, and extends across the border into southern Arizona [71].

Paloverde-mixed cacti desert scrub communities are a characteristic feature of the Arizona Upland subdivision [71,90] (fig. 3). West of the Colorado River in California, variants of paloverde-mixed cacti desert scrub (e.g., lacking saguaro and yellow paloverde) occur on upper bajadas on the northeastern side of the Chuckwalla and Chocolate mountains, and along the southern sides of the Little San Bernardino, Eagle, McCoy, and Palen mountains [13]. Sonoran mid-elevation desert scrub communities occur mostly in a narrow elevational band along the lower slopes of the Mogollon Rim/Central Highlands region, and in Perris Valley and the Cahuilla Mountains in southern California [91]. Granite outcrop desert scrub communities are widespread in the foothills and mountains of Sonora, reaching their northern extent in extreme southern Arizona (fig. 2) [70,71,90].

Map of Sonoran Desert subdivisions with elevation shown in colors indicating elevation ranging from lowest elevation (blue) to highest elevation (white).
Photo Credit
Map adapted from Shreve (1951) and published by Drake et al. (2016) [45].

Figure 3— Map of Sonoran Desert subdivisions.

Site Characteristics

Climate

The climate of the Sonoran Desert is unique among North American deserts. It ranges from arid to subtropical and is characterized by relatively more precipitation [91,125]. Precipitation is generally bimodal in areas dominated by Sonoran desert scrub communities, with some falling in winter and some during summer monsoons. This bimodal precipitation pattern favors the small trees, large shrubs, large cacti, and other succulents that dominate Sonoran desert scrub communities, while smaller-statured desert scrub communities dominate other North American desert regions [125].

Sonoran desert scrub communities occur throughout the Arizona Upland subdivision of the Sonoran Desert and in relatively mesic sites within the Lower Colorado River Valley subdivision [98,125]. Elevation tends to decrease along an east-west gradient from the Arizona Upland to the Lower Colorado River Valley subdivision coinciding with gradients in temperature (low to high) and precipitation (high to low), which correlate with gradients in soil chemistry (see Landform, Elevation, and Soil, below), all of which affect the occurrence of Sonoran desert scrub communities and species composition and stand structure within them [87]. For example, plant cover and density tend to be higher in the eastern part of the Sonoran Desert where precipitation is greater [92].

In the Arizona Upland subdivision, mean annual temperature ranges from about 17.8 °C to 22.2 °C. Mean annual precipitation is mainly between 200 mm and 425 mm and generally has a bimodal distribution, with 30% to 60% falling in summer (June–August), and 10% to 40% falling in winter. Winter precipitation is generally higher in the Arizona Upland subdivision (averaging 75 mm) than in other subdivisions or in the Mojave Desert (averaging 30–50 mm) [125].

The Lower Colorado River Valley subdivision is the largest and driest of the seven Sonoran Desert subdivisions. Mean annual temperature ranges from about 18.3 °C to 23.3 °C, and mean annual precipitation ranges from about 30 mm to 285 mm. Granite outcrop communities occur throughout this subdivision, whereas paloverde–mixed cacti communities only occur in relatively mesic sites where soil moisture is concentrated (e.g., runnels, washes, and depressions), and mid-elevation communities are absent [71,91,117,125,126].

Episodes of extreme cold limit the extent of Sonoran desert scrub communities [71]. While freezing temperatures occur throughout much of the Sonoran Desert, they are uncommon and typically of short duration in areas where paloverde-mixed cacti and granite outcrop desert scrub communities occur [71,91,125]. Episodes of below-freezing temperatures for long durations (about 24 hours or more) can cause considerable mortality and, in exceptional cases, replacement of dominant cacti, trees, and shrubs [71,126], and thus limit the northern extent, upper elevation, and aspect of these communities [15,91]. Mid-elevation desert scrub communities occur in a transitional zone, at locations and elevations where freezing temperatures are too frequent and prolonged for frost-sensitive species that dominate paloverde-mixed cacti and granite outcrop communities to persist, and the climate is too dry for interior chaparral species to be abundant [90].

To the north and east, where temperatures are colder and elevations higher, Sonoran desert scrub communities transition to interior chaparral, desert and semidesert grasslands, or Chihuahuan desert scrub. To the south and southeast, where summer precipitation is higher and freezing temperatures are rare, paloverde-mixed cacti communities transition to subtropical thornscrub [71,125]. To the south and west, where summer precipitation and winter temperatures are lower, lower-statured desert scrub communities dominate the landscape, and Sonoran desert scrub communities may occur on relatively mesic sites and landscape positions [91,125]. To the west, where the transition zone from the Sonoran Desert to the drier Mojave Desert is long and gradual ("ill-defined and arbitrary" [110]), species characteristic of desert scrub in each desert occur together in some locations [110,125].

Landform, Elevation, Soil

Sonoran desert scrub communities covered in this synthesis occur on similar landforms and sites, and they are often adjacent to one another (but also see Adjacent Communities). They mostly occur on slopes and rock outcrops above alluvial flats, valleys, and basins and below elevations where interior chaparral and lower montane woodlands occur. When adjacent, granite outcrop and mid-elevation communities occur upslope from paloverde-mixed cacti communities [90,98,125].

The Arizona Upland subdivision of the Sonoran Desert—where Sonoran desert scrub communities are most common—is a generally mountainous and topographically complex region characterized by slopes, ridges, broken ground, bajadas, washes, and dissected, sloping plains. Steep gradients in elevation, slope, and aspect can occur over relatively small areas [117,125]. The number and variety of life forms range from few to many in the Arizona Upland and are controlled to a great extent by soil moisture, as mediated by differences in soil texture, slope, elevation, rainfall, and temperature [71,92,125]. The Lower Colorado River Valley subdivision—where Sonoran desert scrub communities are typically restricted to relatively mesic sites or rock outcrops—lies mostly below 600 m, and it is characterized by a series of increasingly lower-elevation plains separated by mountain ridges that get smaller and lower toward the west. Plant communities decrease in stature and complexity as elevation decreases [117,125,126].

Paloverde-mixed cacti desert scrub communities occur mainly from 150 to 1,070 m, with rare occurrences up to 1,400 m [12,71,91,125] on lower slopes of mountains, foothills, hillsides, mesas, and upper bajadas, and less commonly in valleys and on alluvial flats [90,91,98]. Paloverde-mixed cacti desert scrub sites typically have gentle to steep slopes and generally shallow, well-drained, gravelly, coarse-textured soils [91] that permit rapid infiltration and uptake during infrequent rainfall events [98]. In uplands, soils are coarse and may be poorly developed. At lower elevations, soils are more developed [71] and caliche (calcium carbonate) is often present, and paloverde-mixed cacti communities occur mostly in association with xeroriparian features such as arroyos and sandy washes [13,73,123]. Parent materials are usually alluvium and colluvium derived from basalt and other igneous or metamorphic rocks. On slopes, plants are often distributed in patches around rock outcrops [91].

Mid-elevation desert scrub communities occur at elevations ranging from 600 to 1,300 m in Arizona and 580 to 880 m in southern California [71,91] on well-drained upland slopes, alluvial fans, and rock outcrops [91]. Soils are generally coarse textured and rocky (cobbly loam to loamy sand [91]) and derived from limestone, granite, or rhyolite [71].

Granite outcrop desert scrub communities occur on low- to mid-elevation (150–1,070 m) granite outcrops, foothills, and low mountains [90].

Plant Community Composition

The three Sonoran desert scrub communities covered in this synthesis each have characteristic overstory species but also share many dominant and subdominant species—many of which are unique to the Sonoran Desert. Together they comprise the vegetation that defines the Arizona Upland subdivision of the Sonoran Desert as described by Shreve and Wiggins (1964) [116] and Turner and Brown (1982) [125] and the Sonoran paloverde-mixed cacti desert scrub and Sonoran sarcocaulescent desert scrub groups as described by the NatureServe International Vegetation Classification [91]. They correspond with the following NatureServe alliances that comprise those groups as follows:

  • Sonoran paloverde-mixed cacti desert scrub group
    • Saguaro–yellow paloverde–velvet mesquite desert scrub alliance [91], which corresponds with the Sonoran paloverde-mixed cacti desert scrub BpS (series 11090) [71]
    • Jojoba–crucifixion thorn–eastern Mojave buckwheat desert scrub alliance [91], which corresponds with the Sonoran mid-elevation desert scrub BpS (series 10910) [71]
  • Sonoran sarcocaulescent (i.e., fleshy-stemmed) desert scrub group
    • Elephant tree–physicnut–organpipe cactus desert scrub alliance [91], which corresponds with the Sonoran granite outcrop desert scrub BpS (series 10900) [71]

Numerous and unique species contribute to the diverse and complex nature of these communities [12,71,117,125]. For example, over 80 species of perennial vascular plants have been recorded in paloverde-mixed cacti communities, and it is not uncommon to have 30 or more in a 0.1-ha plot [93]. Locally, any or several of these species can dominate [31]. Several cacti species are endemic to or most abundant in these communities. The woody plants are typically spiny, or they possess aromatic terpenes or other chemicals to discourage herbivory [31,125].

Sonoran desert scrub communities are characterized by a sparse herbaceous layer composed of perennial grasses and forbs with annuals seasonally present and occasionally abundant (see Stand Structure: Temporal Variation). Perennial grasses—such as black grama, low woolygrass, bush muhly, tobosagrass, and large-spike bristlegrass—and perennial forbs and subshrubs—such as trailing windmills, longflower tube tongue, slender janusia, and desert globemallow may be present but are typically sparse. Annual grasses—such as sixweeks grass and sixweeks threeawn and annual forbs such as desert Indianwheat, curvenut combseed, and Menzies' fiddleneck—may be present to abundant. They grow during late winter and the last 2 months of summer [90,91] and may persist as litter in other seasons. In contemporary Sonoran desert scrub communities, the herbaceous layer on some sites is dominated by nonnative species that are often invasive, the most important and widespread of which are red brome and buffelgrass (see Contemporary Fuel Characteristics). Other nonnative grasses such as Mediterranean grass, lovegrass, fountaingrass, and annual forbs such as Sahara mustard and arugula may also be present to abundant.

Sonoran Paloverde–Mixed Cacti Desert Scrub

The Sonoran paloverde-mixed cacti desert scrub BpS is characterized by a sparse, but very conspicuous and diagnostic, emergent tree layer of saguaro (3–16 m tall) and/or a sparse to moderately dense canopy codominated by xeromorphic drought-deciduous and evergreen short and tall shrubs (0.5–5 m tall) [90,91]. Yellow paloverde, creosotebush, desert ironwood, and other tall shrub and small tree species such as blue paloverde, velvet mesquite, ocotillo, and crucifixion thorn are largely responsible for the arborescent form of the vegetation [12,117,125]. White bursage is a dominant understory shrub or subshrub, and it frequently serves as a nurse plant for the dominant overstory plants [81]. Other common shrubs and subshrubs include catclaw acacia, fairyduster, brittle bush, sangre de cristo, littleleaf ratany, desert-thorn, rough menodora, jojoba, whitestem paperflower, Hall's shrubby spurge, and Rocky Mountain zinnia [90,91]. Saguaro is one of several cacti species that are largely confined to, or best represented in, the Arizona Upland subdivision, where this community is most common and widespread [12,117,125]. Many other cacti are typically present to dominant in these desert scrub communities including species of barrel cactus, hedgehog cactus, cholla, pricklypear, foxtail cactus, and globe cactus [90,91]. These mixed cacti are diagnostic even when saguaro is sparse or absent [91].

Plant community composition varies with topography and geography. For example, yellow paloverde, saguaro, and desert ironwood become increasingly prominent with elevation away from valley floors [12,117,125] such that paloverde–mixed cacti desert scrub communities are best developed on upper bajadas and mountain sides [125]. Variants of this succulent-dominated community occur in the western Mojave Desert in California as "cholla gardens", which are characterized by teddybear cholla, ocotillo, desertsenna, and other succulents, but lack saguaro and yellow paloverde [90]. Desert ironwood and blue paloverde may be present instead [13].

Sonoran granite outcrop desert scrub

The Sonoran granite outcrop desert scrub BpS is characterized by the subtropical and frost-sensitive species, elephant tree, physicnut, and organpipe cactus. Other diagnostic species at the northern extent of this type (in Arizona) include Bigelow's nolina, yellow paloverde, Kearney's sumac, and brittle bush [71].

Sonoran mid-elevation desert scrub

The Sonoran mid-elevation desert scrub BpS is a transitional desert scrub system dominated by crucifixion thorn (on limestone or granite soils) or jojoba (on rhyolite soils) over an open shrub layer composed of species such as eastern Mojave buckwheat, triangle bur ragweed, creosotebush, narrowleaf goldenbush, and/or brittle bush, with a variety of subdominant cacti, shrubs, and subshrubs. The canopy layer may also include blue paloverde, ocotillo, and/or acacia. Frost-sensitive species such as saguaro, yellow paloverde, desert ironwood, teddybear cholla, and mesquite are absent or infrequent [91].

Adjacent Communities

Plant communities adjacent to Sonoran desert scrub communities are important to consider with regard to fire regimes at a particular site because they can influence the likelihood that the adjacent site may burn.

At the lower end of their elevational range, Sonoran desert scrub communities could occur adjacent to or on suitable microsites within smaller-statured desert scrub communities such as creosotebush-white bursage desert scrub communities (BpS series 10870) [12,71,91,125] or blackbrush communities (Mojave mid-elevation mixed desert scrub, BpS series 10820). Many Sonoran granite outcrop communities are surrounded by these small-statured, sparsely vegetated desert scrub communities [71].

At the upper end of the elevational range, possible adjacent communities include desert and semidesert grasslands [71,93,125], interior chaparral (Sonora-Mojave Semi-Desert Chaparral, BpS series 11080 or Mogollon chaparral, BpS series 11040) [71,90], and lower montane oak and pine-oak woodlands (Gambel oak, BpS series 11070 or Madrean pine-oak-juniper) [12,71,93].

Fire Adaptations and Postfire Recovery

Few of the dominant, long-lived woody and succulent plants in Sonoran desert scrub communities are adapted to survive fire or regenerate in the postfire environment [28,37,71,104,112], and many are killed or injured by fire, such that even low-intensity fires may have severe effects [76,84,130]. For example, most cacti do not resprout after top-kill or injury [37,84,122]. Many desert trees and shrubs may resprout after top-kill from fire (e.g., [48,50,89,109]), although the degree of postfire sprouting varies both within and among species (e.g., [2,84]) and is likely influenced by several factors including fuel characteristics, fire characteristics, plant condition, degree of damage, and postfire weather [2]. A greater proportion of resprouting species in mid-elevation desert scrub communities suggests that they may be more resilient to occasional fires than paloverde–mixed cacti and granite outcrop desert scrub communities [71]. Resprouting in desert scrub species may have developed as a response to stress damage from freezing or drought, rather than in response to damage from fire [133].

Irregular recruitment from seeds can limit postfire recovery of Sonoran desert scrub species. Postfire regeneration from seeds is slow, variable, and difficult to predict because it not only depends on the presence of seeds in the soil seed bank or the survival of reproductive individuals, but it is also largely dependent on weather and degree of seed and seedling predation (see Stand Structure: Temporal Variation).

Slow postfire recovery of dominant perennials means it may take many decades or even centuries after high-severity, stand-replacing fire for structure and species composition in Sonoran desert scrub communities to resemble prefire conditions [2,14,48,109]. Abiotic factors, such as site characteristics (elevation, topography, mean annual precipitation) and the amount of precipitation received in the first postfire year may have more influence on postfire vegetation structure and composition over the long-term than time-since-fire [117,120], but studies are inconsistent. A chronosequence study of burned sites in the Arizona Upland and Lower Colorado River Valley subdivisions found that recovery of vegetation cover, height, and density showed closer correspondence to gradients in elevation, postfire precipitation, and annual precipitation, respectively, than to time since fire [117]. In contrast, a systematic review and analysis of data from studies of postfire recovery of perennial vegetation in the Sonoran and Mojave deserts found close relationships between time-since-fire and plant cover in four of five studies, and three of these studies found perennial plant cover was within 10% of that in unburned areas within about 40 years. However, postfire species composition differed substantially from unburned species composition in five of six studies, even 47 years after fire [2]. See Plant Community Conversion for more examples of persistent plant community changes after fire.

Stand Structure

Historical stand structure in Sonoran desert scrub communities is inferred from contemporary stand structure on undisturbed, uninvaded sites and based on the assumption that stand-replacing disturbances are rare or infrequent. These communities generally occur as patches or clusters of shrubs, trees, and cacti with a sparse herbaceous layer and high cover of bare ground within and between patches (e.g., see table A2). Reference conditions are difficult to characterize for these communities because cover and density of component species vary both spatially and temporally. Site characteristics that determine moisture availability and exposure to freezing temperatures (e.g., geographic location, landform, parent material, soil characteristics, and topographic position) (e.g., [136]) contribute to spatial variation, while climatic fluctuations that result in erratic recruitment and mortality contribute to temporal variation (e.g., [56,126]). Contemporary stand structure and species composition is increasingly affected by nonnative plant invasions (e.g., [96]) and wildfire (see Fire Management Considerations).

Spatial Patterns and Variation

Sonoran desert scrub communities occur in stands ranging from tens to thousands of hectares. Paloverde-mixed cacti desert scrub commonly occurs at scales of hundreds of hectares and may occur at scales of thousands of hectares [71]. In drier parts of the range (e.g., the Lower Colorado River Valley subdivision), paloverde-mixed cacti communities may form linear stands on flats at low elevations along sites where soil moisture is concentrated (e.g., runnels, washes, and xeroriparian areas) [117,125]. Mid-elevation desert scrub stands range from narrow strips on steep slopes extending for long distances to very broad areas such as in the Verde Valley. Stand size varies with aspect and elevation, ranging from about 40 to >2,000 ha [71]. No information was available on stand size in granite outcrop communities.

Undisturbed, uninvaded Sonoran desert scrub stands are composed of patches of sparse to open scrubland or low woodland, with a typically sparse and discontinuous herbaceous layer and areas of bare soil or rock within and between patches [91,125] (fig. 4). Competition for limited moisture and nutrients helps maintain wide plant spacing within and between small and large patches [71]. Favorable microsite characteristics result in a patchy structure within larger patches. For example, microsites under individual tree and shrub canopies (e.g., nurse plants) tend to accumulate silt, fine sand, litter, and seeds, and thus become favorable microsites for moisture retention and seedling recruitment (reviews by [81,93]), such that plants tend to be clustered in these microsites.

Woody canopy layers can be sparse to moderately dense (1%–50% cover) and are codominated by medium- to tall-statured, drought-deciduous and evergreen shrubs or small trees (mostly about 0.5–5 m tall) over one to several open layers of smaller (<0.5 m tall) shrubs, subshrubs, and succulents [90,125]. Tall, woody cacti are often present [91], and many paloverde-mixed cacti stands are characterized by a sparse, emergent tree layer of saguaro, 3- to 16-m tall [90,125]. Canopy cover and composition varies with site characteristics. For example, saguaro density may be lower and shrub cover higher (reaching 50% or more) on northern aspects [93]. Because disturbances are rare and dominant plants are long-lived, age structures are skewed toward a low density of large, old plants [56].

Various landscapes in  Sonoran desert scrub communities showing structural variability.
Photo Credit
Top left: U.S. Geological Survey photo by Sarah Studd. Top right and bottom left: USDA Forest Service photos by Charlie McDonald. Bottom right: photo © Colin Barrows, some rights reserved (CC BY-NC).

Figure 4—Examples of structural variability in Sonoran desert scrub communities. 

Herbaceous layers are usually sparse and discontinuous on undisturbed sites and consist of perennial and annual grasses and forbs that vary in composition and abundance both within and among years (see Temporal Variation, below). In many contemporary stands, the herbaceous layer is dominated by nonnative invasive plants that can result in uncharacteristically tall, dense, and continuous herbaceous layers (see Contemporary Fuel Characteristics).

Geographically, Sonoran desert scrub communities occur more frequently, continuously, and with greater plant density in the eastern part of their range, and become more scattered and less dense moving west, as vegetation transitions from large areas dominated by medium to tall-statured desert scrub to large areas dominated by lower-statured creosotebush–white bursage communities [56,126]. For example, cover of dominant overstory plants in paloverde-mixed cacti communities may reach 50% on Arizona Upland sites [71,93] but is usually <20% in southeastern California [71]. From east to west, precipitation decreases and temperature, soil calcium content, total organic carbon, particle size, and pH increase. Distribution and abundance of saguaro and several of its nurse plant species (triangle bur ragweed, white bursage, yellow paloverde, creosotebush, mesquite, and desert ironwood) reflect these gradients. For example, the gradient in soil pH likely governs the range of several species (e.g., creosotebush) that are restricted to calcic soils in western areas [87].

Species composition, stand structure, and plant abundance also vary within and among patches [56,126], due to differences in topographic position, soil texture, and associated water availability. For example, the gradient from coarser-textured soils on upper bajadas to finer-textured soils on lower bajadas and valleys was associated with a gradient from higher diversity of plant species and growth forms on upper bajadas to lower diversity on lower bajadas [21,98].

Temporal Variation

Structure and composition of the herbaceous layer of Sonoran desert scrub communities vary in the short-term (seasonally and annually) in response to fluctuations in the timing and amount of precipitation, whereas structure and composition of perennial plants vary over the long-term, in response to climate variability and extremes.

Live annual grasses and forbs are present during two distinct periods when moisture availability is greatest: late winter and during the monsoon season in the last 2 months of summer [90,91]. Moisture is generally unavailable to shallow-rooted plants from April through June and from September to early December, when annual plants senesce [92,117]. Cover and density of native annuals is typically sparse but can be high in years with above-average precipitation [90,91], depending on precipitation timing and amounts and seed bank composition and density. Two or more consecutive years of above-average precipitation can add large numbers of seeds to the soil seed bank, and result in a relatively dense and continuous cover of native annual plants [84,85,104,137]. Contemporary Sonoran desert scrub communities may be dominated by nonnative invasive species that can form a more dense, continuous, and persistent herbaceous layer than those dominated by natives.

Long-term variation in perennial vegetation is generally attributed to the interacting effects of natural attrition and erratic recruitment, which are related more to climate variability and climate extremes than to large-scale, stand-replacing disturbances. However, multi-year, extended drought or occasional catastrophic freezes (>24 hours) can cause widespread mortality of dominant plants and alter stand structure when older individuals die without replacement [21,71,98,117,125]. For example, widespread mortality and limited recruitment of saguaro in Saguaro National Park since 1941 has altered the structure of the population from one dominated by large, old saguaros (> 5.4 m tall) to one dominated by small, young saguaros (<1.8 m tall) since the 1990s [41,97]. Natural attrition more commonly results from old age, pathogens, and root or stem parasites in adults, and from herbivores, seasonal drought, and interspecific interference in seedlings and juveniles [18].

Recruitment of many dominant trees, shrubs, and cacti in Sonoran desert scrub communities is generally episodic and strongly influenced by the timing and amount of precipitation (e.g., [41,97,126]). Low or erratic seedling establishment has been noted for many dominant species including saguaro, yellow paloverde, ocotillo, velvet mesquite, creosotebush, and jojoba [20,36]. Seedling recruitment can be abundant in relatively wet years, but it is scarce to absent in most years (e.g., [20]). Germination and seedling survivorship of saguaro and other tree-layer plants tend to be reduced in hot, dry weather [136]. Studies on Tumamoc Hill near Tucson, Arizona, show large fluctuations in individual species abundance during the 20th century, that were usually, but not always, coincident with fluctuations in rainfall (e.g., [18,20,56]). Over 6 years at an ungrazed site on Tumamoc Hill, only a few species emerged in every year. First-year survival averaged across all 15 species was 3.7%; and only 0.1% of seedlings lived as long as 4 years. First-year seedling survival improved with increased rainfall, although about three times as many seedlings died from predation as from desiccation [20].

Human-caused disturbances (e.g., road building, woodcutting, and livestock grazing) and nonnative species introductions are largely responsible for altering stand structure and species composition in contemporary Sonoran desert scrub communities [56,126]. In addition, wildfires have become increasingly more common in Sonoran desert scrub communities in the late 20th and early 21st centuries (see Contemporary Fire Frequency), coincident with increases in abundance of nonnative plants (mostly grasses) that not only interfere with natural succession, but also have the potential to fuel additional wildfires, establish an invasive plant/fire cycle, and convert areas of desert scrub to nonnative grassland or steppe (see Fire Management Considerations).

Historical Fuel Characteristics

Historical fuel characteristics in Sonoran desert scrub communities are inferred from fuel characteristics on undisturbed, uninvaded contemporary stands. Growth patterns of native vegetation in undisturbed Sonoran desert scrub communities (i.e., patchy distribution of dominant shrubs and trees; sparse, usually discontinuous cover of herbs; and high percentage of bare ground) (see Stand Structure) result in fuels that are inadequate to carry fire in most years [7,62,71,84,99,104,108,135].

Amount and seasonality of precipitation are important large-scale drivers of fire in North American deserts due to their effects on fuels [33,67,104]—fires are generally less frequent in drier deserts because less fuel is produced [62]. Average and below-average precipitation result in sparse and discontinuous fine fuels in most desert scrub ecosystems, and this likely precludes fire spread on many sites [104]. During years of above-average precipitation, especially 2 or more consecutive years, establishment of annual forbs and grasses can result in relatively thick and continuous patches of fine fuels that are sufficient to carry fire (fig. 5) [71,84,85,88,100,104,114,137], and thus increases the likelihood of fire ignition and spread on those sites [7,28,58,71] for one or more fire seasons [114].

Dense patch of desert Indianwheat.
Photo Credit
Photo © David Vander Pluym, some rights reserved (CC BY-NC).

Figure 5—Dense patch of the native forb, desert Indianwheat, a primary fine fuel source for many desert fires in 2005. 

Large fires—with large, contiguous areas burned—were rarely if ever documented in Sonoran desert scrub during the 19th and most of the 20th centuries, making it unclear whether native annuals could reach densities sufficient to carry fire over large areas, and therefore whether large fires could have occurred in these communities historically (e.g., [109]). During the late 20th and early 21st centuries, however, several large fires occurred in the Sonoran and Mojave deserts, and some were fueled largely—although not exclusively—by native annuals (e.g., the 1979 Granite Fire and the 2005 King Valley Fire) (e.g., [24,27,48,50,84,127]). Nonnative annuals were also present on these sites, and the area burned included communities other than Sonoran desert scrub. Nonetheless, these observations led Brooks and Matchett (2006) to suggest that Arizona Upland vegetation occurs in an "ecological zone" (typically on lower mountain slopes at elevations above arid shrublands) that seems "to possess fuels near the threshold of where fire can spread across large areas" [24]. However, observations of the 2020 Bighorn Fire and 2019 Mercer Fire suggest that patchy fires with relatively small burned areas in Sonoran desert scrub communities may have been more likely, historically. Burned areas of Sonoran desert scrub occurred only where native and nonnative annuals had been most abundant, such as northern aspects. Warmer, drier, and/or rockier aspects remained unburned, unless they were invaded by nonnative perennial grasses, such as buffelgrass [133], suggesting that these areas were unlikely to burn historically. See Contemporary Fire History for more details on these fires.

Information about flammability and fuel characteristics of native plant species was lacking in the available literature. A study of desert plant flammability suggests that native plants such as creosotebush, white bursage, and Menzies' fiddleneck spread fire slowly (0.12 cm/s) but burn at relatively high temperatures for a long duration, such that that those growing beneath shrubs can act as “igniters” of dead shrub branches. Flames produced by dead branches can then engulf the entire shrub, resulting in locally intense fire with potentially severe effects on adjacent plants [53]. For example, plants clustered near saguaros may form a potentially lethal fuel bed for the fire-intolerant cactus [135].

The many cacti in Sonoran desert scrub communities typically do not combust, and therefore are not likely to fuel fire spread. However, observations suggest that saguaro may combust and spread fire under some circumstances (e.g., [103]). See the FEIS Species Review on saguaro for details. Common sotol plants rolling in high winds may also spread fire [103].

Historical Fire Regimes

Lacking any means of establishing a fire history for Sonoran desert scrub communities (e.g., witness trees, stand age, historical observations) details of presettlement fire regimes remain uncertain. However, inferences based on observations of fire effects and postfire recovery of dominant plants, stand structures, and fuel characteristics suggest that desert scrub ecosystems, including Sonoran desert scrub communities, probably evolved with little to no fire [7,37,62,84,107,109], and that fire frequency and severity may have differed among the three community types.

Although lightning storms are frequent in the Sonoran Desert during the summer, lightning ignitions were unlikely to spread beyond individual plants or small plant clusters in Sonoran desert scrub communities due to patchy stand structure and discontinuous fuels. Very small or small (<50 ha), patchy fires were more likely than large (>400 ha), contiguous fires.

Observations of wildfires in the late 20th and early 21st centuries indicate that large, severe, or repeated fires can have long lasting effects on species composition and stand structure in Sonoran desert scrub communities, supporting the idea that presettlement fires must have been rare or infrequent, very small, and/or patchy—leaving patches of unburned vegetation to persist and reproduce. If presettlement fires had been more frequent and homogenous or widespread, Sonoran desert scrub communities would be less common on the landscape [48]. Because the dominant and characteristic perennial plants in most Sonoran desert scrub communities are susceptible to fire injury and mortality, any fires that did occur were presumed likely to have high-severity effects [76,84,130].

Historical Fire Ignition and Season

Historically, fires in Sonoran desert scrub communities would have been ignited by dry lightning during summer monsoonal storms (mid-May to early September) when lightning is common [28,48,71,103]. High air temperatures and low relative humidity in the Sonoran Desert are favorable conditions for wildfire ignition during most of the year; however, fuels are inadequate for fire spread in most years (see Historical Fuel Characteristics). Although there are accounts of American Indians using fire in southeastern Arizona (e.g., [11,17]), it is not likely that they intentionally burned desert scrub communities, and no mention of this was found in the literature. However, fires intentionally ignited in adjacent ecosystems could have burned into Sonoran desert scrub communities under some conditions, increasing the likelihood of fire at those locations [109].

Historical Fire Frequency

It is generally thought that fire was rare in paloverde–mixed cacti and granite outcrop desert scrub and infrequent in mid-elevation desert scrub communities, because many of the dominant and characteristic species are poorly adapted to survive fire or thrive in the postfire environment (see Fire Adaptations and Postfire Recovery), and because fuels are usually too sparse and discontinuous to carry fire (see Historical Fuel Characteristics). Fire frequency likely varied within and among communities depending on site characteristics and adjacent plant communities. Fire frequency estimates from LANDFIRE succession modeling (based on expert opinion, literature, and local data) suggest mean fire interval estimates of 103 to 350 years for mid-elevation desert scrub, 500 years for granite outcrop desert scrub, and more than 1,000 years for paloverde-mixed cacti desert scrub (see the LANDFIRE Biophysical Settings section) [71]. The importance of resprouting species in mid-elevation desert scrub and its frequent adjacency to grassland, interior chaparral, and oak and pine-oak woodland communities led reviewers to suggest a greater likelihood of fire (i.e., shorter fire intervals) in those communities than in paloverde-mixed cacti or granite outcrop communities [71].

Estimates of presettlement fire frequency for Sonoran desert scrub communities from the literature are based largely on lack of fire adaptations in many dominant plants. Observations indicate that many of the dominant plants—especially the trees, cacti, and some shrubs—are killed by fire, recruit episodically (on decadal timescales) (e.g., [52]), and grow slowly [121], such that populations recover very slowly after a single fire [2,47,48,84,107,109], and may not recover at all after large, high-severity, or repeated fires (see Plant Community Conversion). For example, Rogers (1985) suggested that fire intervals less than 30 years would eliminate saguaro from paloverde-mixed cacti communities [107], and Esque and Schwalbe (2002) later responded that saguaro persistence would require fire-free intervals "much longer than 30 years", because of its specific requirements for germination and establishment [48]. Thomas (1991) estimated that presettlement fire-free periods in the Sonoran Desert had to be more than 250 years, based on the sensitivity of native succulents to fire [122]. In contrast, Paysen et al. (2000) suggested presettlement fire intervals of 35 to <100 years for "paloverde-cactus shrub" communities [101], although it is unclear what this estimate is based on.

Fire may have been more likely on some sites and at some locations due to microclimatic variability [67], which affects fuel characteristics, and due to frequent fire regimes in adjacent plant communities, which could have increased the likelihood of ignition from adjacent sites. For example, based on observations of contemporary fires fueled largely by native annuals, Sonoran desert scrub communities that occurred in landscape positions with relatively more mesic conditions, such as northern aspects, higher elevations [133], and xeroriparian areas [130], may have been more likely to burn due to higher cover of fine fuels.

Sonoran desert scrub communities on sites adjacent to plant communities with similar fuel structures and rare or infrequent fires—typically at lower elevations—were probably less likely to burn than those adjacent to plant communities with frequent fires (e.g., grassland, interior chaparral, and oak or pine-oak woodland)—typically at higher elevations [62,71,103,104,133]. Estimated presettlement fire intervals for adjacent plant communities at lower elevations, such as creosotebush–white bursage communities (330 years) and blackbrush communities (400–833 years) are long enough that Sonoran desert scrub communities were probably not much more likely to burn due to adjacency. Estimated presettlement fire intervals of adjacent communities at higher elevations are typically much shorter than that of Sonoran desert scrub; for example, interior chaparral fire interval estimates range from about 40 to 80 years, most desert and semidesert grasslands estimates range from about 10 to 100 years, and oak and pine-oak woodlands estimates range from about 4 to 55 years [71]. Of the three communities covered in this synthesis, mid-elevation desert scrub communities occur at the highest elevations and are the most likely to be adjacent to communities where fires were historically more frequent, contributing to shorter fire-interval estimates for those communities [71].

Contemporary Sonoran desert scrub communities that occur at the ecotone with grassland communities may not have been present historically, and instead established when fire was excluded from grassland communities after Euro-American settlement, largely due to the reduction of fine fuels by livestock grazing. In Saguaro National Park, plots at elevations ranging from 990 to 1430 m—the ecotone between Sonoran desert scrub and desert grassland—had burned one or more times since the cessation of livestock grazing 40 years prior and had transitioned from Sonoran desert scrub to dominance by perennial bunchgrasses. Cover and density of trees decreased over time, regardless of time-since-fire, even more than 20 years after fire. The authors speculated that these sites were historically dominated by grassland or steppe vegetation, and that presettlement fire intervals on these sites may have ranged from 20 to 40 years, which would have maintained grass dominance [120]. The timing of widespread saguaro establishment on a slope above the Mount Lemmon Highway at about 1,128 m elevation coincided with the introduction of livestock grazing in the late 19th century. The authors suggest that this area may have been a grassland community, and the few older saguaros that were present prior to that time probably occupied open rocky terrain that lacked fine fuels to carry fires [130]. While early observations indicated that livestock grazing and associated fire exclusion from desert grasslands allowed “desert scrub” dominants to establish and spread into adjacent grasslands (e.g., [11,30,61,74]), evidence of this dynamic is lacking for Sonoran desert scrub communities and is more common in other types of desert scrub—especially that dominated by mesquite (e.g., [30,61,74]).

Although it is unlikely that American Indians intentionally burned desert scrub communities, burning in adjacent communities may have resulted in occasional fires at those locations [109].

Historical Fire Severity

Because many of the dominant and characteristic species in Sonoran desert scrub communities are poorly adapted to survive fire or reproduce in the postfire environment [49,84], it is often assumed that any fires that did occur would have had high-severity effects. LANDFIRE models classify these rare or infrequent historical fires as high-severity, stand-replacement fires (>75% top-kill) [71]. However, observations of the 2020 Bighorn and 2019 Mercer fires [133] and the 2021 Telegraph Fire [72] suggest that fires fueled by native annuals may have left a mosaic of burned and unburned patches within the fire perimeter (i.e., mixed-severity effects), and that overall burned area in Sonoran desert scrub communities would have been small and localized. After the 1979 Granite Fire burned large areas of Sonoran desert scrub, small, unburned patches of vegetation were found throughout burned areas. With the exception of these patches, nearly all trees and shrubs were completely top-killed, and a small percentage resprouted [84].

Historical Fire Size and Pattern

Lack of fire adaptations in dominant plants suggest that fires were not only rare or infrequent in Sonoran desert scrub communities, but also that when fires did occur, they were probably small (<50 ha) and/or patchy, leaving unburned vegetation to repopulate burned areas (e.g., [17]). Patchy and discontinuous distribution of vegetation (i.e., fuels) also suggests that small and patchy fires were more likely than large, contiguous fires.

Large contemporary fires (>400 ha) in the Sonoran and Mojave deserts fueled mostly (but not entirely) by native annuals after periods of above-average precipitation (see Contemporary Fire History) suggest that large fires may have been an infrequent part of the presettlement fire regimes in some desert scrub communities [28,104,106], but these occurred only when adequate fine fuels were present along with weather conditions associated with extreme fire behavior [71]. These large fires may have been linear in shape, as a result of convective storms with high winds [7,28,58,62,67,71,84,104,107,112]. Even so, large fires did not likely carry far into Sonoran desert scrub communities, and then only in locations where conditions resulted in a dense enough cover of annuals to carry it (e.g., [133]).

Contemporary wildfires in Sonoran desert scrub are mostly ignited by humans [7,49,112], tend to be larger and more widespread following periods of above-average precipitation (e.g., [7,44,112]), and are often and increasingly fueled by nonnative invasive plants [26,28,44,48,54,68,104,109,114]. Buffelgrass and red brome are the most common nonnative plant fuels in Sonoran desert scrub communities, although other nonnative plants may be locally abundant and/or widespread in adjacent communities, and thus also responsible for increasing the likelihood of fire ignition and spread on sites where dense and continuous populations of these plants occur.

Although the details of presettlement fire regimes in North American deserts remain uncertain, available evidence suggests that the size and frequency of contemporary wildfires are outside the historical range of variability for Sonoran desert scrub communities. As fires become more common in desert scrub communities, fuel characteristics may be further altered by postfire changes in plant community composition and structure, with a greater dominance of nonnative, fire-adapted grasses, and thus increasing the probability of additional fires (see Fire Management Considerations: Invasive Plant/Fire Cycle).

Role of Nonnative Invasive Plants

Disturbances from human settlements and land use activities coupled with an increase in cool-season precipitation variability over the past half century have facilitated invasion by nonnative plants on many sites throughout the Sonoran and other North American deserts [1,58]. Nonnative plant invasions can have a variety of impacts. Of most concern to fire managers are changes in fuel characteristics and fire regimes. Other impacts might include decreases in total native herb canopy, herb species richness, shrub density and canopy, and insect and bird diversity, as was found on sites invaded by lovegrass in southeastern Arizona (Bock et al. 1986, cited in [54]).

Nonnative invasive plants can add abundant and novel fuels to desert scrub ecosystems, increasing fuel biomass and continuity, and thus increasing the potential for uncharacteristically large and frequent wildfires. By altering fuel characteristics, nonnative plant invasions—especially grasses—can increase the range of conditions favorable for fire ignition and increase the potential for fire spread and high fire intensity [28,55,71,83,104,114,118]. High-intensity fires may be more likely than low-intensity fires to severely damage or kill cacti [83,118] and other dominant native desert scrub perennials and may also limit resprouting (see Contemporary Fire Severity). Nonnative invasive plants may also increase the likelihood of repeated fire by recovering quickly and providing sufficient fuels to carry another fire soon after burning [28,55,71]. The long-term effect of nonnative plant invasions on the seasonality, frequency, and severity of desert wildfires threatens the sustainability of desert ecosystems on many sites [28].

Species of Concern

Photos depicting various invasive annual grasses that cover large areas in the Sonoran Desert.
Photo Credit
A) Photo © Eric Hough, some rights reserved (CC BY-NC). B) U.S. Geological Survey photo by Molly McCormick. C and D) Photo © Steve Jones, some rights reserved (CC BY-NC).

Figure 6—Invasive annual grasses cover large areas in the Sonoran Desert. A) A cluster of red brome plants. B) Dried red brome plants in desert scrub interspaces. C) A cluster of common Mediterranean grass plants. D) A large, dense patch of common Mediterranean grass plants. 

Nonnative species of greatest concern with regard to altered fuel characteristics and increases in fire occurrence in Sonoran desert scrub communities include the annual grasses red brome and Mediterranean grass (fig. 6), the perennial grasses buffelgrass, fountaingrass (especially, crimson fountaingrass) (fig. 7), and lovegrass (Lehmann lovegrass and weeping lovegrass), and the annual forbs Sahara mustard and arugula (fig. 8) [28,54,71,103,104,114], although several other nonnative species, including soft feather pappusgrass, are spreading and of increasing concern (e.g., [16,54,77,130,134]). Presence of red brome, Mediterranean grass, buffelgrass, fountaingrass, and lovegrass has been linked, at least anecdotally, to fire occurrence in Sonoran desert scrub communities [55,103,104] and other Sonoran and Mojave desert ecosystems [44,48,55,71,103,114]. Of these, red brome and buffelgrass are most widespread and problematic in Sonoran desert scrub [28,44,48,49,71,121]. Other species may be abundant in localized areas of Sonoran desert scrub (e.g., arugula [28,44], lovegrass [103], fountaingrass [54,134], and soft feather pappusgrass [16,77,130]) or in adjacent creosotebush communities (e.g., Mediterranean grass [28,48,88] and Sahara mustard [28,63]) and could become more widespread [28,44].

The invasive annual grasses red brome and Mediterranean grass are the most abundant plants below 1,250 m elevation over large areas in the northern Sonoran Desert. Red brome is especially invasive in the Arizona Upland subdivision where Sonoran desert scrub communities are common, while Mediterranean grass is more invasive in creosotebush communities in the Lower Colorado River Valley subdivision. Both can dominate herbaceous layers and have the potential to establish more horizontally continuous, flammable, and persistent fuel beds than native annual plants [28,48,104].

Dense nonnative invasive perennial grasses.
Photo Credit
National Park Service photos.

Figure 7—Nonnative invasive perennial grasses form dense patches of fine fuels. A) Buffelgrass, and B) fountaingrass under Sonoran paloverde–mixed cacti desert scrub in Saguaro National Park.

Nonnative invasive perennial grasses are increasingly prevalent in the monsoonal regions of the Sonoran Desert, and land managers and scientists share a growing concern over their spread and effects on fuel characteristics and fire regimes in invaded communities [47,134]. Buffelgrass, for example, is especially invasive in the Arizona Upland Subdivision, often in Sonoran desert scrub communities [8,44,77,128]. Buffelgrass populations in these communities have rapidly increased in extent since its introduction [49], filling in bare areas between native plants and spreading rapidly to displace native vegetation [83,95,96]. Rapid spread of buffelgrass has been observed in Saguaro National Park [83,95,96], in the Santa Catalina Mountains [95], at the Desert Laboratory in Tucson [19], and at Organ Pipe Cactus National Monument [111]. See the FEIS Species Review on buffelgrass for more details [63]. Crimson fountaingrass, lovegrass, and soft feather pappusgrass also have the potential to invade Sonoran desert scrub communities and alter fuel characteristics [16,130]. Crimson fountaingrass is an ornamental perennial and a close relative of buffelgrass. It has escaped cultivation and is abundant and widespread at Organ Pipe Cactus National Monument, and it has spread into both districts of Saguaro National Park [54,134]. Lovegrass was widely used for postfire reclamation on the Tonto National Forest in the 1960s and 1970s, and subsequently established in saguaro communities on the Tonto National Monument, where they likely helped carry repeated fires [103]. Soft feather pappusgrass was first observed in the Finger Rock Canyon Watershed of the Santa Catalina Mountains, Arizona, in 1990, and has since been increasing rapidly in distribution and abundance [16].

Nonnative invasive annual forbs can form dense patches.
Photo Credit
A) Photo © Kevin W Smith, some rights reserved (CC BY-NC). B) Photo © Eric Hough, some rights reserved (CC BY-NC). C) Photo © kdouglas8787, some rights reserved (CC BY-NC). D) Photo © Mike Plagens, some rights reserved (CC BY-NC).

Figure 8—Nonnative invasive annual forbs can form dense patches of fine fuels in the Sonoran Desert. A) A Sahara mustard plant. B) Dense Sahara mustard plants under a large shrub. C) Scattered, small arugula plants. D) A tall, dense patch of arugula in desert scrub. 

Relationships with Fire Regimes

Sonoran desert scrub communities with an invasive grass component may have greater fine-fuel density and continuity and a higher likelihood of frequent, high-intensity, and/or large fires than uninvaded communities [71]; however, data supporting this relationship are limited to a few species and locations [10] and are lacking for Sonoran desert scrub communities in particular. A 2020 meta-analysis of published literature on the effects of native versus nonnative species on fire frequency, fire intensity, flammability, fuels quantity, and fire spatial extent showed “a strongly significant difference between fire metrics associated with nonnative and native species, with nonnative species linked to enhanced fire effects and risk.” However, this analysis also highlighted the limited scope of our understanding—only a small number of species (22) and ecosystems had been quantitatively examined. Only 30 studies met criteria for inclusion in the analysis, and only two included information relevant to Sonoran desert scrub [10].

While anecdotal evidence suggests a strong potential for negative impacts that may not yet be detectable, an analysis of the effects of 12 invasive grass species on fire regime characteristics in U.S. ecoregions showed significant impacts for some but not all invasive species in the Sonoran Desert. In the Sonoran Basin and Range and Sonoran Desert ecoregions, fire was significantly more likely to occur in areas with invasive populations of buffelgrass than in uninvaded areas, and fire frequency in buffelgrass-invaded areas was more than twice that in uninvaded areas. Also in those ecoregions, fire was significantly more likely to occur in areas invaded by common Mediterranean grass than in uninvaded areas [55]; however, that most likely reflects fires in creosotebush-white bursage desert scrub, where Mediterranean grass is most invasive [28,48,88]. Across five ecoregions, including the Sonoran Basin and Range (but excluding the Sonoran Desert ecoregion, where red brome abundance was limited) no significant differences in fire occurrence or fire size was detected between areas invaded by red brome and uninvaded areas. Insufficient spatial data were available to examine effects of Lehmann lovegrass, weeping lovegrass, and crimson fountaingrass on fire regime characteristics [55].

Although data confirming effects on fire regime characteristics are lacking for red brome, lovegrass, and crimson fountaingrass [55], observations suggest that these grasses have helped fuel wildfires that burned areas of Sonoran desert scrub. Anecdotally, reported impacts of red brome—primarily from the Mojave Desert—include increased fuel load and persistent flammability; and increased frequency, occurrence, and spread of low-intensity fire (plants are 0.1–0.7 m tall) (e.g., [24,25]). Observations also indicate that several wildfires in Sonoran desert scrub since 1979 were fueled by red brome, including the 1994 Mother’s Day Fire in Saguaro National Park, which burned 138 ha of Arizona Upland desert scrub [47,49]; and a 6,500-ha fire in the Harcuvar Mountains in 1999 [48]. Lehmann lovegrass, weeping lovegrass, and crimson fountaingrass are reported to be fire promoters in some ecosystems. For example, repeated wildfires in upper Cave Creek on the Tonto National Monument were likely fueled by lovegrass that spread onto the monument from the adjacent Tonto National Forest, where it had been planted to stabilize soils after the 1964 Schultz Fire [103].

Buffelgrass-fueled fires can be exceptionally detrimental to Sonoran desert scrub communities because they burn at high temperatures and intensities (see Contemporary Fire Severity), spread quickly, and can occur any time of year [49,83]. In addition, buffelgrass recovers quickly after fire, so it has the potential to carry very frequent fires (see Fire Management Considerations: Invasive Plant/Fire Cycle).

Contemporary Fuel Characteristics

Fine surface fuel layers in contemporary desert scrub communities are increasingly dominated by nonnative invasive grasses and forbs, which provide novel fuels and contribute to fine fuel load, height, continuity, and persistence on many sites in the Sonoran and Mojave deserts [2,28,47,49,83,88,112,114]. Across 2,530 plots in the Sonoran Desert in southwestern Arizona, plots with nonnative plants had higher total fine fuel biomass estimates (about 33 kg/ha) than plots with only native plants (about 9 kg/ha). Biomass estimates were extremely low across the study area for both native and nonnative plants because field work took place during years with below-average precipitation (2011 and 2012) [114]. Sonoran desert scrub sites invaded by perennial buffelgrass are likely to have considerably higher fine fuel biomass than uninvaded sites or sites dominated by native and nonnative annuals (table 2), although data are limited. Nonnative herbaceous fuels may burn during different seasons and at greater intensity and spread fires faster and farther than native fuels, although fuel characteristics differ among nonnative species, and information on this topic is lacking.

Surface fine fuel beds in uninvaded Sonoran desert scrub communities are typically dominated by annual plants, therefore fine fuel loads and continuity can change dramatically from year to year, depending on precipitation patterns [114]. Nonnative annual plant invasions may amplify this fuel–climate relationship in the Sonoran Desert [44,48,58]. Like native annual plants in desert scrub communities, nonnative annual plants can establish dense populations in understories and interspaces following periods of above-average precipitation [7,48,104,112,114,121] and form nearly continuous beds of fine surface fuels that dry out in late spring and persist during the summer lightning season or longer [48,104,133]. For example, annuals were so dense in spring 2005 in some areas that large amounts of fine fuel persisted into the middle of 2007 [50,58,114,127]. These fine fuels can be sufficient to sustain a wildfire and thereby increase the likelihood of fire ignition and spread [48,104,133].

Unlike native annual plants, which are mostly forbs, nonnative annual plants are mostly grasses. Grasses have fine leaves with large surface area to volume ratios that can dry out quickly and provide fuels that carry fire under a broader range of conditions than do woody fuels and surface leaf litter [7,25,43]. Nonnative annual grass litter and standing dead material can accumulate rapidly and decompose very slowly in desert climates [25,28,48,104], persisting for 2 years or longer [28,48].

Nonnative annual forbs (e.g., Sahara mustard and arugula) are often large and can also produce abundant, persistent litter on sites where they invade [28,29]. In comparison, native annual fuels tend to be small and decompose rapidly after they die [28,48,62,104].

Nonnative perennial grass invasions can form large, continuous fine fuel loads [48,49,114] that persist with less interannual variation than sites dominated by annual plants [7,29,83]. For example, fine-fuel biomass in buffelgrass-dominated desert scrub sites can be orders of magnitude greater than similar sites dominated by red brome [47,49], and it may approach or exceed that reported for nearby native and nonnative desert grassland sites (table 2). In the absence of intense grazing, buffelgrass can fill in bare interspaces between native perennial plants [22,51,83] and provide continuous fine fuel—particularly, but not exclusively, after high rainfall years [34,42,79,104]. Fine fuel loads on buffelgrass-invaded sites fluctuate much less than fine fuel loads on sites dominated by native and/or nonnative annuals in the Sonoran Desert [47,77,83,94,119], which can increase the probability of fire ignition and spread in any season of any given year [7,29,47,83]. Ungrazed buffelgrass accumulates flammable material every growing season, and after several years of growth it has been described as an “almost-woody subshrub” [46]. See table S1 in Wilder (2021) for information on fuel models used in desert scrub ecosystems, including adjustments made for buffelgrass fuels [133].

Although contemporary fires are often fueled by nonnative plant invasions, native plants also helped fuel large wildfires in some years (e.g., [50,58,84,114]). The King Valley Fire, for example, was carried largely by desert Indianwheat and other annual fine fuels and by traveling along xeroriparian desert wash ecosystems with "substantial tree cover" composed of species such as desert ironwood, blue paloverde, yellow paloverde, desert willow, and/or smoketree [127,130]. However, it is unlikely that fuels were exclusively native plants in most contemporary fires, given that nonnative plants were already present in Arizona and often locally abundant.

Table 2—Examples of herbaceous layer fuel loads reported from different fuel types in desert ecosystems.
Ecosystem, dominant fuelsLocation, yearBiomass (kg/ha)Reference
Sonoran desert scrub, nonnative perennial (buffelgrass)Javelina Picnic Area, 20032,828 ± 335 (SE)[47]
Sonoran desert scrub, nonnative perennial (buffelgrass)Panther Peak, 20032,480 ± 227 (SE)[47]
Sonoran desert scrub, nonnative perennial (buffelgrass)Saguaro National Park, several locations, July 20082,600–6,850[83]
Sonoran desert scrub, nonnative annual (red brome)Tonto National Forest, 1981–82203–799[37]
Sonoran desert scrub, nonnative annual (Mediterranean grass)Tonto National Forest, 1981–823–19[37]
Sonoran desert scrub, mix of native and nonnative annualsTonto National Forest, 1981–82203–799[37]
General desert, mix of native and nonnative annualsNorth American desert biomes0 to >700Review by [47]
General desert, mix of native and nonnative annualsSonoran Desert90–950Patten 1978, cited in [83]
Desert grassland, native perennial (big sacaton)Arizona desert grasslands3,900–5,150[83]
Desert grassland, nonnative perennial (Lehmann lovegrass)Southwest desert grasslands1,100–6,000[83]

Contemporary Fire History

Fires were rarely if ever documented in Sonoran desert scrub communities until several large wildfires occurred near Tucson and Phoenix during the late 1970s and early 1980s that included areas of Sonoran desert scrub (e.g., [44,80,89]). These fires prompted several studies of contemporary fire regimes and their effects on native plants and animals (table 3). Seemingly unprecedented "intense and dramatic" fires continued to occur throughout the northern Sonoran Desert in the late 20th and early 21st centuries [48,114]. From the early 1970s through 2021, years with high fire activity in the Sonoran Desert typically followed periods of above-average precipitation that resulted in periodic abundance of both native and nonnative annual plants [32,120]. The number of fires and total area burned in and around the Sonoran Desert in Arizona were especially high in 2004, 2005, 2019, and 2020 (based on data from Short (2022) [115]). Descriptions of a few of these fires are available in the literature (see Contemporary Fire Frequency), although data and analyses are lacking, and it is unclear how much of the total area burned in these fires was in Sonoran desert scrub communities.

Several sources indicate large areas of Arizona Upland vegetation or saguaro-dominated communities burned in contemporary fires. On and near the Tonto National Forest, for example, "a considerable portion" of mature saguaro-shrub habitat burned in the late 20th century [89], including an estimated 30% of saguaro habitat on the Mesa Ranger District between about 1970 and 1995 [7,89,107,135]. Esque (2004) describes several "major" fires in the Arizona Uplands including fires in the Santa Catalina Mountains in 1986 and the Mother's Day Fire in Saguaro National Park in 1994 [49]. One of the largest fires in 1979, the Granite Fire, south of Florence, Arizona, was said to have burned >10,000 ha of Arizona Upland, including paloverde-mixed cacti communities [49,84,127]. The 2021 Telegraph Fire started in desert scrub, burned 3,326 ha of desert scrub on the Tonto National Forest, spread into other plant communities, and grew to 73,150 ha [72].

Table 3—Contemporary fire histories that include areas dominated by Sonoran desert scrub communities.
Area studiedStudy area size (ha)YearsFire variables examinedReference
Sonoran Desert portion of the Tonto National Forest391,0001955–1983Frequency, season, annual area burned, size, and ignition source[106,108,112]
Sonoran Desert portion of the Tonto National Forest391,0001955–2000Frequency, season, annual area burned, size, and ignition source[6,7]
Tonto National Monument4501942–1991Season, ignition source, and postfire photo points[103]
Sonoran Desert in Arizona6,654,2001989–2010Frequency, season, annual area burned, size, and fuels[44]
BLM and adjacent lands in the Sonoran Desert of southwestern Arizona1,160,0001970–2005Fuels[114]

Frequent, large, and severe fires in contemporary Sonoran desert scrub are attributed mostly to increased human access (i.e., increased road and trail density in and adjacent to wildlands) and subsequent human-caused ignitions (see Contemporary Fire Ignition and Season, below), in addition to increases in receptive fine fuel loads and continuity brought about by nonnative species invasions. These fuels can increase the likelihood of fire ignition and spread (see Contemporary Fuel Characteristics) that may result in larger patches of high-severity effects (see Contemporary Fire Size and Pattern). Rapid recovery of herbaceous fuels—especially nonnative grasses—may lead to repeated fires on a site (see Fire Management Considerations: Invasive Plant/Fire Cycle). Repeated and/or large patches of high-severity fire can lead to vegetation type conversions that may persist indefinitely (see Fire Management Considerations: Plant Community Conversion).

Contemporary Fire Ignition and Season

Contemporary fires in the Sonoran Desert are mostly ignited by humans and can occur in any month of the year. Fires are most frequent from late spring to mid-summer, when both human- and lightning-caused ignitions are most common. From 1955 to 2000, nearly 70% of fires on the Tonto National Forest were human-caused. Most fires occurred in May (26.9%) and June (25.2%), largely before the summer monsoon and lightning season, followed by July (21.6%), August (8.8%), and September (5.3%). About 6% occurred in fall and winter (October–March), when lightning is uncommon [6]. From 1955 to 1983, all fires on the Tonto National Forest during the cooler months of November through March were human-caused, and only during July and August were lightning-caused fires more frequent than human-caused fires. Fires were numerous from May through August, but the area burned was greater during June than all other months combined [112]. The number of fires on the Tonto National Forest from 1955 to 2000 increased with the population of Maricopa County and with traffic along major highways in the forest. However, there was no relationship between these factors and area burned, presumably because human-caused fires were mostly smaller than lightning-caused fires [6].

In the Sonoran and Mojave deserts, lightning-ignited fires are most common when lightning is most frequent—during the summer monsoon. From 1985 to 2001, 78% of lightning strikes in the desert bioregion of California occurred from July to September [28]. Lightning-ignited fires generally occur from mid-May to September in the Sonoran Desert and are most frequent and burn the greatest area from June through August [103]. From 1942 to 1991 all fires in Sonoran desert scrub communities on the Tonto National Monument were started by lightning between mid-May and mid-August—most frequently in July. Most area was burned by fires that started in late June or July, including four large fires (1947, 1964, 1970, and 1980) that came onto the monument from the national forest to the south during conditions of extreme heat and, in some cases, high winds [103]. The 2019 Mercer Fire was ignited in August when lightning struck a dense saguaro-paloverde stand on a south-facing slope heavily infested with buffelgrass. It burned 10 ha before running out of fuel (buffelgrass) and dying out [133].

Contemporary fires in Sonoran desert scrub communities are often ignited in adjacent communities (e.g., grassland, interior chaparral, or pine-oak woodland) and spread into desert scrub through ecotone areas with high densities of fine fuels. For example, the 2020 Bighorn Fire was ignited in June by lightning in semidesert grassland, on a steep and rugged, north-facing slope at about 1,200 m in the Santa Catalina Mountains. Fueled primarily by dead winter annuals (about 50% cover on unburned plots), as well as native bunchgrasses and nonnative lovegrass, the fire backed downslope from its ignition point into transitional, Sonoran mid-elevation desert scrub communities dominated by sotol, ocotillo, velvet mesquite, pricklypear, and hopbush. The fire spread into areas of paloverde-saguaro desert scrub on northern aspects, where fuel loads from dried winter annuals were higher than on other aspects, and in areas where fuels consisted of patches of nonnative perennial grasses. Other paloverde-saguaro communities were mostly unburned in the Bighorn Fire [133]. In some areas and years, dense patches of fine fuels along roadsides can extend into native plant communities and potentially carry fire into wildlands [7].

Nonnative invasive plants may increase the potential for fire in any season by providing receptive fuels nearly year-round [49,78,83]. For example, although nonnative perennial buffelgrass may be particularly dry and flammable during the monsoon season when lightning is most frequent [47], it can burn in any season [49], even when green [46]. See the buffelgrass Species Review for more information.

Contemporary Fire Frequency

Fires have become much more common across large parts of North American warm deserts, and they can pose significant land management challenges in localized areas [28]. Limited analyses from a small number of fire history studies (table 3) suggest that fire frequency in Sonoran desert scrub communities generally increased during the late 20th century. Most fires were small, human-ignited fires along roads (e.g., [6,49,112,114]). Arizona Bureau of Land Management records show that 210 fires burned 36,621 ha in the Arizona Upland and Lower Colorado River Valley subdivisions between 1973 and 1979, suggesting fire cycles shorter than the life span of long-lived dominant species [109]. In contrast, fire frequency was still sufficiently low (fire cycle of about 274–294 years) in the approximately 391,000-ha Sonoran Desert portion of the Tonto National Forest (dominated by Arizona Upland desert scrub) from 1955 to 1983, to support the idea that fires in the Sonoran Desert had been historically rare or infrequent, despite a trend of increasing fire frequency and annual area burned during that time [112]. Increased fire frequency was attributed to increased human-caused ignitions, changes in fine fuels—which would have been more abundant during the wetter than normal winters in the late 1970s—and/or improvements in fire detection and reporting [112]. Annual number of fires in that area remained high through 2000, especially during relatively wet periods [6,7]. These early studies (e.g., [112]) are often cited to support the contention that fire frequency and extent are “increasing” in the Sonoran Desert. An updated analysis including fires from 2000 to present and identifying area burned in specific plant communities within the Sonoran Desert is needed to validate and clarify trends [24].

Because fine fuel loads and continuity vary dramatically from year to year (e.g., [44,58,114]), contemporary wildfires in the Sonoran Desert are most common (and larger, see below) after abundant precipitation produces dense patches of native and/or nonnative herbaceous vegetation that cures during dry, hot conditions in late spring and forms relatively continuous fine fuel beds [7,28,48,58]. For example, on the Tonto National Forest, both number of fires and area burned from 1955 to 2000 were greater after two and three consecutive winters with above-average precipitation [6,7]. In dry shrub communities in southern Arizona, area burned by wildfires from 1980 to 2000 was greatest 1 to 2 years after periods of above-average precipitation [132]. While lacking analyses, additional observations from the early 1970s through 2021 support this pattern. For example:

  • During late spring and summer of 1974, "numerous" fires occurred in the Arizona Upland after above-average precipitation in the winter of 1973–1974 resulted it a high density of annual plants—mostly the nonnative annuals redstem stork’s bill and red brome—on many sites [130].
  • During spring and summer of 1979, several fires occurred in the desert regions of Arizona [49,84,127] and in creosotebush desert scrub along the western margins of the Sonoran and Mojave deserts of southern California [32], following two consecutive winters of above-average precipitation.
  • In 2005, at least 25 fires >400 ha occurred in the Sonoran Desert [44,88,115,130] after 3 years of above-average precipitation, including exceptionally high rainfall in the winter of 2004–2005 (nearly 4 times the mean in some areas) [127]. In spring 2005, large areas of the Sonoran Desert were dominated by dense stands of the native forb, desert Indianwheat, and lesser amounts of nonnative Mediterranean grass [44,50,88,130].
  • In 2019, 2020, and 2021, large fires in Arizona and California that included areas of desert scrub (fig. 9) were preceded by periods of above-average precipitation. Heavy rains in October 2018 and 2 to 3 inches of snow in January 2019 led to dense patches of red brome, which fueled the 50,100-ha Woodbury Fire in June 2019 [72,115]. The following wet winter and early spring (2019–2020) resulted in abundant winter annuals in the Sonoran Desert—about half of which were native species and half red brome. This was followed by a delayed and dry monsoon season, periodic strong winds, and the hottest summer in Tucson’s 125-year weather record [133], as well as the 48,600-ha Bighorn Fire [133] and the 78,300-ha Bush Fire [72,115] in Arizona, and the 17,500-ha Dome Fire in southeastern California [133]. In June 2021, the Telegraph Fire was ignited in desert scrub and eventually burned a total of 73,150 ha, including >3,200 ha of desert scrub on the Tonto National Forest [72].
Burned Sonoran desert scrub.
Photo Credit
U.S. Geological Survey photos by Molly McCormick.

Figure 9—In 2019 and 2020, fires carried by the invasive red brome burned hundreds of thousands of acres of Sonoran desert scrub and resulted in some of the biggest fires in Arizona history. Burned areas are highly vulnerable to vegetation type conversion and further degradation. The Restoration Assessment and Monitoring Program for the Southwest (RAMPS) mapped the spread of red brome to help land managers make decisions on how to prevent large fires in the Sonoran Desert. 

Contemporary Fire Severity

Clear accounts of fire severity in Sonoran desert scrub communities are not common in the available literature. While some accounts generally indicate high levels of mortality and high-severity effects (see Fire Adaptations and Postfire Recovery), others suggest more of a mosaic of unburned patches and burned patches with varied fire severity within fire perimeters (i.e., mixed-severity fires) (e.g., the 2021 Telegraph Fire [72] and 2020 Bighorn Fire [133]). Nonetheless, high-severity fire effects are very likely more widespread in contemporary Sonoran desert scrub communities than they were historically, in part because nonnative invasive grass fuels are generally taller and/or more continuous than native herbaceous fuels. Therefore, fires in nonnative fuels are likely to burn at higher intensity and spread farther than those in native fuels, resulting in larger burned patches with more severe effects [83,133]. Greater frequency of extreme fire weather can also contribute to greater frequency and patch size of high-severity fire in desert scrub.

Few postfire data are available, and those may be biased toward areas with high-severity effects or repeated fires because these are the areas that fire scientists have focused on. Fires on and around the Tonto National Forest in the late 20th century caused considerable mortality to saguaro and other dominant plants (e.g., [7,89,107]) and were likely considered high-severity fires, although severity data were not reported or were not comparable. The 1979 Granite Fire was considered a high-severity fire because nearly all trees and shrubs were top-killed in burned areas, and overall mortality was high (50%–92%) for five dominant trees and shrubs 19 months after the fire. Mortality was especially high among cacti, and very few individuals sprouted; 91% of hedgehog cactus, 59% of barrel cactus, 98% of globe cactus, and 98% of cholla were dead after the fire. Although many plants survived in small, unburned patches within burned areas, and some individuals of each species sprouted after fire (25% of yellow paloverde, 32% of velvet mesquite, 1% of triangle bur ragweed, 9% of Nevada jointfir, and 37% of creosotebush), no seedlings of woody perennials were observed on burned transects either 7 or 19 months after fire [84], and mortality was delayed for some species. While initial mortality of saguaro was estimated at 11%, remeasurements [107] and repeat photographs [130] indicate delayed mortality for saguaro and codominant trees, eventually reaching 100% in some areas. Within 6 years after the fire, "an open plant community of widely scattered low shrubs is all that remains of the former arborescent community". Native dominants were still mostly absent 20 years later [130]. Observations of delayed mortality in paloverdes were similar after the 2005 King Valley Fire, and resprouting of woody dominants seemed less common than after the Granite Fire. Rainfall was well below average during the 2 years after the King Valley Fire, which may have inhibited resprouting [130].

While historical fires in Sonoran desert scrub communities are classified by LANDFIRE as stand-replacing (i.e., high-severity), observations of many contemporary fires suggest that mixed-severity fires may have been more likely. For example, data from the 2021 Telegraph Fire suggest a mixed-severity fire in the 3,326 ha of desert scrub within the fire perimeter on the Tonto National Forest, with most area (65%) burned at high and moderate severity. Fire severity maps based on postfire changes in vegetation show 1,140 ha burned at high severity, 1,016 ha at moderate severity, 839 ha at low severity, and 331 ha remained unburned within the fire perimeter [72]. The 1994 Mother’s Day Fire in Saguaro National Park may also have been classified as mixed severity, based on 24% mortality of saguaro and 73% of paloverde in 2000 [47,49]. Small, unburned patches of vegetation were found throughout severely burned areas after the 1979 Granite Fire [84]. The King Valley Fire was carried, in part, by woody plants in desert wash ecosystems, which were more severely burned than surrounding creosotebush desert scrub [130].

Fires fueled by nonnative annual grasses may burn at greater intensity and spread faster and farther than those fueled by native plants—and therefore have different effects—although fuel characteristics differ among nonnative species, and information on this topic is lacking for most species. Fires in Mediterranean grass tend to burn with low intensity and negligible soil heating, and they leave most woody shrubs unburned. In contrast, nonnative annual brome grasses produce greater fuel loads and depths than do Mediterranean grasses, and accordingly produce longer flame lengths and more intense fires [28] and are likely to have higher-severity effects. Laboratory experiments suggest that Mediterranean grass is a “spreader” of fire, carrying fire quickly (1.32 cm/second) and burning for short duration (0.5 minute) at low temperatures. In contrast, native plants such as creosotebush, white bursage, and Menzies' fiddleneck spread fire slowly (0.12 cm/second) and burn up to eight times longer (4 minutes) and at higher temperatures [53], suggesting higher severity effects from these native fuels.

Fires fueled by nonnative perennial grasses like buffelgrass may burn at higher intensity and have higher severity effects (i.e., greater postfire mortality of dominant plants and less postfire resprouting) than those fueled by native or nonnative annual plants. For example, in paloverde-mixed cacti communities burned in the 2020 Bighorn Fire, paloverde, brittlebush, and saguaro were "burned more severely" (i.e., scorched and/or killed) in areas where the fire was fueled by nonnative perennial grasses (mostly buffelgrass, although fountaingrass was also common) than in areas where the fire was fueled by winter annuals [133]. Based on fuel characteristics, buffelgrass-fueled fires likely have longer flame lengths, more rapid rates of spread, higher temperatures, and higher-severity effects (i.e., cause greater mortality of native flora and fauna) than fires fueled by native and nonnative annuals [47,83]. Mean maximum temperatures generated by buffelgrass-fueled fires in the Avra Valley, Arizona—where buffelgrass was planted in the late 1970s, and its biomass ranged from 6,710 to 12,100 kg/ha—ranged from 568 °C at the soil surface to 799 °C at 30 cm above ground, with a maximum recorded temperature of 871 °C (height not given). The authors noted that even thin but continuous patches of buffelgrass could carry fire and produce temperatures lethal to native plants [83]. In comparison, using similar but not identical methods, mean maximum temperatures during a June prescribed fire in an uninvaded paloverde-saguaro community on the Tonto National Forest—where fuel loads ranged from 700 kg/ha to 3,200 kg/ha—were 76 °C in interspaces, 167 °C under yellow paloverdes, and 210 °C within triangle bur ragweeds at 30 cm above ground. Maximum temperatures were 88 °C in interspaces, 299 °C under yellow paloverdes, and 405 °C within triangle bur ragweeds—all recorded at 1 cm above ground [99].

Fires that occur during periods with higher fuel moisture may have lower severity effects than those occurring during more extreme fire weather (i.e., hotter, drier, and/or windier), regardless of fuel type. For example, the 2019 Mercer Fire burned 10 ha of a dense saguaro-paloverde community and was fueled by buffelgrass; however, it occurred during a period of high humidity, which "raised dead fuel moisture and dampened fire behavior". The authors suggest that this may explain the strong spatial variation in saguaro mortality within the fire perimeter, and why most other perennial species "exhibited at least some regrowth four months after fire". Although many species (e.g., yellow paloverde, hopbush, saguaro, and most other cacti) had “high rates” of fire injury and postfire mortality during the first postfire year, and the authors anticipate additional, delayed mortality [133], data for saguaro alone suggest a mixed-severity rather than a stand-replacement fire: Of 95 saguaros sampled in the burned area, 89 (94%) had some fire injury; 39 (41%) had >50% scorching; and 28 (29%) had >80% scorching; 17 individuals (18%) died in the first year, most of which were young plants, and the authors estimated that 28 (36%) of the 78 living saguaros had a 50% or greater chance of dying in the future [133].

Contemporary Fire Size and Pattern

Most contemporary fires in Sonoran desert scrub communities are small (<20 ha), and fire size is generally smaller in the Sonoran and Mojave deserts—where 97% of total area burned from 1992¬ to 2016 was by fires <50 ha—compared to other North American desert regions—where 97% of total area burned from 1992 to 2016 was by fires <300 ha (Short 2017, cited in [59]). Nonetheless, several large fires occurred in and around the Sonoran Desert during the late 20th and early 21st centuries, and many exceeded 400 ha in total size (based on data from Short (2022) [115]). Large fires are more likely following periods of above-average precipitation that result in continuous patches of fine fuels. Many contemporary fires in the Sonoran Desert burned areas of desert scrub, including Sonoran desert scrub; however, data on area burned by plant community or BpS were not reported in the literature reviewed. Information on fire pattern is similarly lacking.

Fine surface fuel density and connectivity can limit or enhance fire spread and thus, fire size and patchiness in desert scrub communities. Fires in general, and especially large fires and large area burned in the Sonoran Desert, typically coincide with periods of above-average precipitation that result in increased continuity of fine surface fuels. In and around the Sonoran Desert in Arizona, maximum fire size, total area burned, and number of large fires (>400 ha) were high in 2004, 2005, 2019 and 2020 (based on data from Short (2022) [115]), coinciding with periods of above-average precipitation (see Contemporary Fire Frequency). From 1989 to 2010, 316 small (<20 ha) and 79 large (≥20 ha) fires occurred in the Sonoran Desert in Arizona [114]. Most large fires occurred (n = 36) and most area burned (51,700 ha) in 2005 [44]. These fires were fueled by dense patches of native and nonnative annual plants that had established in typically bare interspaces and connected patches of native perennial plants [44,88,114]. Median size of large fires in the Sonoran Desert in 2005 was 95 ha, compared to a median size of 60 ha over the 22-year study period [44,114]. Across a 45,100-km2 study area in the Sonoran Desert of Arizona (mostly within the Lower Colorado River subdivision), the probability of large fires (≥20 ha) from 1989 to 2010 was greatest in years with high maximum annual normalized difference vegetation index (NDVI, which represents patterns of vegetation greenness), and in areas at low elevations and with low road density [44,58]. NDVI is used as a proxy for annual fine fuel loads to estimate fire risk over large, contiguous extents (see Fire Management Considerations: Fire Prediction and Fuels Management). Twenty hectares was used as a cut off for large fires because it “represents a low-end estimate for large fire size in desert fuels" (W. Reaves, BLM, personal communication cited in [58]).

Published observations of contemporary fires suggest that because fire spread (and by extension, fire size) is limited by patchy fuels, the largest burned areas in Sonoran desert scrub communities tend to occur on relatively mesic sites and/or sites with dense populations of invasive grasses. For example, burned areas within the 2020 Bighorn Fire had dense patches of winter annuals and/or perennial grasses, and these patches varied with elevation and slope characteristics. Patches of buffelgrass, fountaingrass, and Lehmann’s lovegrass were most dense and common at the semidesert grassland/desert scrub ecotone, and patches were generally small and discontinuous along rocky hillsides. Fire spread was limited along those hillsides, and by areas of bare ground, hiking or game trails, and rock outcrops, which helped prevent fire spread into areas dominated by yellow paloverde, brittlebush, and saguaro that occurred downslope. Remnant winter annuals were much less dense (<5% total cover) [133], and nonnative perennial grasses were presumably absent from these unburned Sonoran desert scrub sites. Continued spread and coalescence of buffelgrass patches would increase fuel continuity and fire spread on these sites [133].

Large fires are most often ignited by lightning and driven by extreme fire weather. Lightning-caused fires tended to be larger on average, and they burned approximately twice the area burned by human-caused fires on the Tonto National Forest from 1955 to 1983 [112]. The larger size of lightning-caused fires might be attributed to the timing of dry thunderstorms at the end of the spring drought when vegetation is most flammable, and to the strong, gusty winds that accompany them. Lightning-caused fires are also more likely to occur in remote, inaccessible regions where they are not discovered or suppressed before growing large [112].

In contrast, human-caused fires typically occur in accessible places such as along roads and near campsites, where there is greater access for both human-caused ignitions and faster detection and suppression [112]. As of this writing (2023), these analyses are over 30 years old, and none has been published since, so it is unclear if this relationship has persisted. However, of 78 fires larger than 400 ha reported on the Tonto National Forest (in all vegetation types) from 1993 through 2020, 52 were lightning-caused, 25 were human-caused, and 1 had an undetermined cause (data from Short (2022) [115]), suggesting that this relationship persists.

While observations are limited, patchy fuels suggest that fires in desert scrub burn in a mosaic of varied severities, including unburned patches within the fire perimeter. One of the largest fires in 1979, the Granite Fire, south of Florence, Arizona, was said to have burned >10,000 ha of Arizona Upland, including paloverde-mixed cacti communities [49,84,127]. One observation suggests this was a patchy fire, with small, unburned patches of vegetation in the fire perimeter [84]. The 2021 Telegraph Fire started in desert scrub and grew to 73,150 ha. The fire burned through 3,326 ha of desert scrub (including but not limited to Sonoran desert scrub) on the Tonto National Forest, which resulted in a mosaic of high-severity (1,140 ha), moderate-severity (1,016 ha), low-severity (839 ha), and unburned patches (331 ha) [72].

Sonoran desert scrub communities, especially those invaded by nonnative grasses and forbs, are not resilient to frequent, large, and/or severe wildfires. Because postfire recovery of predisturbance vegetation may not occur for centuries [48,114], if at all, the primary fire management consideration is fire prevention, part of which requires fuels management in the form of reducing cover of nonnative invasive plants.

An aggressive program of fire suppression in the Sonoran Desert might be viewed as "inconsistent by a ranching community that is increasingly accepting fire as an ecological process capable of holding shrub invasion of grasslands, bordering the Sonoran Desert, at bay" [121]. However, widespread occurrence of nonnative invasive plants coupled with a changing climate increases the likelihood of large and widespread, high-intensity fires with large patches of high-severity effects in Sonoran desert scrub [71], which may lead to profound, undesirable changes that are difficult to predict. Changes may include persistent losses of native vegetation cover and diversity, losses to native seed banks, changes in demographic processes, increased erosion, changes in nutrient availability, increased dominance of invasive species, and reduced habitat for vulnerable wildlife that rely on mature desert scrub communities—including numerous threatened and endangered species [9,48,58,114]. The potential for an invasive plant/fire cycle and conversion from native desert scrub to nonnative grassland or other alternative state is of increasing concern [117]. Plant community conversion has already occurred—to a relatively limited extent—on many sites at middle and low elevations of the Sonoran Desert [28,48,58].

Many aspects of postfire vegetation recovery remain poorly understood in this and similar desert ecosystems, and more work is needed to improve understanding of how plant recovery differs among fires, sites, species, and climates [2]. For example, the degree to which topographic and climatic gradients influence vegetation recovery patterns is poorly understood. Understanding the relative influences of time-since-fire and abiotic conditions is important for managing burned desert landscapes, particularly in terms of prioritizing restoration activities or evaluating effects on habitat for threatened species such as the Sonoran desert tortoise [117].

A simulation study that maps the degree of resilience, risk, and vulnerability to change by fire across the landscape of the Sonoran Desert suggests that sites in the Arizona Upland subdivision have some of the highest fire risk and lowest ecological resilience. The simulation incorporates a combination of social factors (e.g., availability of fire management resources) and ecological factors (e.g., fuel abundance, topographic diversity, geophysical diversity, vegetation diversity, habitat connectivity, species richness, human modification, and water availability) [9].

Invasive Plant/Fire Cycle

In many ecosystems globally, including areas of the Great Basin and Mojave Desert, invasion by fire-adapted nonnative grasses has been identified as a major driver of increased fire frequency that can eventually lead to an invasive plant/fire cycle and plant community type conversion [26,43,48,55,104]. Regional alteration of fire regimes has not been quantitatively assessed for most species and ecosystems, including Sonoran desert scrub. However, mechanisms by which invasive grasses promote fire are likely applicable across large spatial scales and in a range of ecosystems [55]. Because nonnative grasses such as buffelgrass, red brome, crimson fountaingrass, and nonnative forbs such as Sahara mustard can establish and increase in cover soon after fire, dominate many previously burned sites in the Sonoran and Mojave deserts, and thrive under a regime of frequent fires [28,47,104,112], a feedback cycle may establish that promotes further spread of nonnative species and diminishes occurrence of natives [10,55].

While an invasive plant/fire cycle has not been identified in Sonoran desert scrub, substantial anecdotal information, and data from ecosystems with similar fuel structures indicate a strong potential on many sites. An invasive grass/fire cycle has been reported to occur in the Mojave Desert for more than 20 years, but it has only been substantiated in specific elevation zones and regions [67]. Although fire regimes over most of the low-elevation zone are still within the historical range of variation [28], repeated burning has converted large areas of creosotebush–white bursage to annual grasslands in parts of the western Sonoran (i.e., in the Lower Colorado River Valley subdivision) and Mojave deserts [48].

For desert scrub communities in the Sonoran and Mojave deserts, susceptibility to invasive grass/fire cycles tends to increase from high to low elevations and is greatest where native vegetation lacks fire-tolerant traits and seasonal precipitation is sufficient for invasive grasses to establish, spread, and eventually dominate landscapes [38]. Large, high-severity burns may predispose areas to further establishment and spread of invasive plants, and these areas could act as large-scale nascent foci from which invasive plants could spread into adjacent unburned areas and increase the likelihood and frequency of fire on those sites [66,67]. Over a 30-year projected period, state-and-transition models suggested that large fires and greater area burned were more likely in areas of the Tucson Basin where buffelgrass was present than where it was absent, especially below 1,200 m elevation. A noticeable increase in fires after 20 years was associated with more continuous areas of buffelgrass cover, which was projected to spread from 440 ha of invaded area to 4,920 ha of invaded area over the 30 years [133]. Frequent and/or severe buffelgrass-fueled fires in Sonoran desert scrub are likely to lead to a decline of trees (e.g., desert ironwood and bursera), shrubs (e.g., yellow paloverde and creosotebush), and cacti (e.g., saguaro and organ pipe cactus) that are not adapted to fire [39,82,105].

Plant Community Conversion

Resilience in vegetation structure following fire in many arid communities may be limited to single, small, low-severity fires in areas that have not burned in many decades [66]. Esque et al. (2006) estimate that it may take 200 years to replace mature saguaro stands killed by fire, assuming the sites do not burn again during recovery [47]. Repeated fires, and possibly a single, high-severity fire, could lead to long-term or relatively permanent changes in Sonoran desert scrub [2,3,66,104,112], including local extirpations of long-lived species such as saguaro and paloverde, and could convert saguaro-paloverde desert scrub to more fire-tolerant vegetation such as mesquite and acacia savannas [47,54] or nonnative grasslands [50]. Sonoran mid-elevation desert scrub may be more resilient to fire than paloverde-mixed cacti and granite outcrop communities—because more of the dominant species resprout after top-kill—however, information is lacking.

Woody perennials and cacti in Sonoran desert scrub communities may be reduced substantially by a single fire, and repeated fires can eliminate them from a site (see Fire Adaptations and Postfire Recovery). Published postfire information from these communities is limited to a handful of fires and sites and is largely anecdotal. After fires in the 1970s and 1980s, Rogers (1985) suggested that two fires within a few years could result in the local extirpation of many desert species, including saguaro. He reported that repeated fires ignited by motorists eliminated saguaro and many other desert perennials from large areas near major roads, such as along Interstate Highway 17, north of Phoenix [107]. Similar changes have been observed along Highway 87 northeast of Scottsdale (T. Esque 1995, personal observation cited in [48]), and along the Bush Highway, where an area of desert scrub that burned 4 times over about 25 years had largely been converted to native and nonnative grasses [7]. Similarly, repeat photography at two locations on the Tonto National Monument showed sites once dominated by saguaro and paloverde that had no trees or large shrubs, few saguaros, and abundant grasses (e.g., lovegrass) after burning 4 or 5 times between 1942 and 1984 [103]. Five “major” wildfires that occurred in upper Cave Creek on the Tonto National Monument from 1964 to 1980 were likely fueled by lovegrass. A 1981 photo shows scattered saguaro plants that are “very badly damaged”, and a 1988 repeat photo shows only one saguaro on the site and no trees remaining [103]. Repeat photography from several sites that had burned in one or more fires shows declines and losses of many common and dominant native desert scrub perennials—especially saguaro and yellow paloverde, but also buckhorn cholla, barrel cactus, and teddybear cholla—sometimes accompanied by increases in nonnative grasses. The authors suggest that it could take decades for native perennials to reestablish [130].

Observations of the Granite Fire and Dead Man Wash Fire suggest that Sonoran desert scrub communities in areas of smooth, level topography are more vulnerable to damage and extirpation by fire than those on rocky slopes. These areas are subject to both greater likelihood of ignition (due to greater human access) and fewer natural impediments to fire spread [7,107,130].

Sonoran desert scrub communities adjacent to frequent-fire communities at upper elevations (e.g., grassland, interior chaparral, or pine-oak woodland) may also be more susceptible to conversion because fire is more likely to spread from these communities and postfire dominance of grasses may be more likely. For example, a 1975 grassland-adjacent fire in a saguaro community at its upper elevational limit (around 1,500 m) killed the few saguaros that survived a freeze in 1962. In February 2006 the burned area was dominated by grasses, including a large population of nonnative perennial soft feather pappusgrass [130].

While evidence of plant community conversion is relatively limited for Sonoran desert scrub communities, a chronosequence study from desert scrub in the Mojave Desert revealed long-term (30–40 years) conversion of native, woody-dominated communities to nonnative, herbaceous-dominated communities and raises similar concerns for the Sonoran Desert. Communities were converted after single, moderate- to high-severity fires, or after periods of high fire frequency, independent of burn severity. Across elevation zones, woody cover was 1.5 to 3 times greater in unburned plots than high-severity burned plots 40 years after fire, and herbaceous cover was greater in moderate- to high-severity burned plots than unburned plots 20 to 40 years after fire. Herbaceous cover remained higher than woody cover for several decades, especially on plots burned more than once than those burned once. The authors noted "virtually no resilience in shrub cover when woody-dominated areas burn more than twice in a half-century" [66].

Fire Prediction and Fuels Management

Fire prevention includes identifying where large fires are likely to occur [58,114] and reducing fire risk by managing nonnative invasive plant fuels. For example, information about herbaceous plant location and phenology, landscape structure, and predicted fire behavior can help managers prioritize high risk areas for treatments to control invasive grasses or implement other actions, such as placement of new hiking trails, in ways that help prevent fire spread. This information might also be used to help prioritize strategic areas to apply flame retardant during wildfires [133].

Mapping fine fuel loads annually, including nonnative plant distribution and spread potential, is important for identifying where large fires are likely to occur in a given year and can help fire managers with decision making (e.g., [44,59,114]). Tools are available for mapping and tracking the annual buildup of fine fuel, which is a strong predictor of large fire probability [44,58,59]. For example, the Rangeland Productivity Monitoring Service and associated Fuelcast.net allows users to see past and projected fuel loads by map zone. Normalized difference vegetation index (NDVI) was the strongest predictor of large fire probability in the Sonoran Desert from 2005 to 2018, followed by fire weather, topography, and climate [59]. Maximum NDVI the year before the fire was less influential than that the year of the fire, but it was still a strong predictor of large fires [44]. NDVI is used as a proxy for the annual buildup of fine fuel [58,114]. Maximum NDVI reflects the total accumulation of herbaceous fuels (it does not distinguish between native and nonnative fuels) [58].

Because of their disproportional contribution to fine fuel loads and the risk of invasive plant/fire cycles and plant community conversion, nonnative invasive plant distributions—current and projected—are a focus for fuel management in the Sonoran Desert (e.g., [44,114,129]). Combining bioclimatic envelope modeling with mechanistic modeling targeted to a given species can help land managers identify locations and species that pose the highest risk for conversion [1]. Biomass maps created using field sampling and remote sensing data were used in conjunction with landscape features (e.g., elevation and road density) to model invasion risk (i.e., habitat suitability modeling) and fire likelihood associated with five nonnative invasive plant species (red brome, Mediterranean grass, buffelgrass, Sahara mustard, and arugula) on two military installations, two wildlife refuges, and one national monument in the Sonoran Desert of Arizona. Based on total area with “high fire likelihood”, the Kofa (36,500 ha) and Cabeza Prieta (78,700 ha) wilderness areas were most likely to burn, and with high or very high negative effects likely. These areas overlapped with two of the largest wildfires that occurred in 2005: the King Valley Fire (13,000 ha) in the Kofa Wilderness and the Growler Peak Fire (11,000 ha) in the Cabeza Prieta Wilderness. These areas fall mostly within the Lower Colorado River Valley subdivision. Large portions of the North Maricopa Mountains (18,000 ha) and Woolsey Peak (17,700 ha) wilderness areas also had high fire likelihood, and these areas largely fall within the Arizona Upland subdivision and had not experienced a large fire since 1984 [44].

Fine fuels management should be closely tied to invasive plant management from the perspective of both managing invasive plant fuels that are currently present, preventing their spread, and preventing establishment and spread of additional species that may change fuel structure and potentially cause even greater fire management challenges in the future. Even when nonnative annual grasses represent <1% vegetative cover and may be overlooked, they can still be a dominant component of the soil seed bank, making it difficult to detect and evaluate their population trends. Although populations of nonnative annual grasses can be depleted by drought, they can recover to ecologically significant numbers relatively quickly [28]. Dispersal abilities of invasive plants can be grossly underestimated, and nonnative invasive plants should be considered a threat not only where they are currently found, but also in adjacent or similar communities [48]. Land managers have observed damage caused by fires fueled by red brome in southern Arizona and are even more concerned about the potential effects of buffelgrass-fueled fires (S. Rutman, Organ Pipe Cactus National Monument, personal communication cited in [28]). Larger fires, more area burned, and loss of native species can be expected if buffelgrass spreads and increases in abundance [133].

A well-funded, systematic approach over an extended period might control the spread of nonnative invasive plants in susceptible areas (e.g., [4,83]). Areas most susceptible to nonnative grass invasion are those along road corridors, near urban areas with high propagule pressure, areas where current or historical livestock grazing has reduced native plant cover, and areas burned by repeated or large, high-severity fires [38,66,133]. Atmospheric nitrogen deposition downwind of urban or agricultural areas can increase soil nitrogen availability, which may increase biomass of invasive annual grasses in those areas [38].

Management in a Changing Climate

As the global climate warms, conditions favorable for large fire occurrence are likely to increase in the Sonoran Desert, where steadily increasing temperatures and increasingly sporadic and intense rainfall are expected to increase the length of the fire season and spur production of fine fuels—especially nonnative grasses—and result in elevated risk of large wildfires [1,58]. Frequency of days with extreme fire danger in the Sonoran Desert nearly doubled from 2000 to 2010 due to increased temperature and reduced humidity in spring and summer months [1]. Unusual precipitation and drought patterns associated with climate change may be as likely to result in increased biomass production and subsequent drying of native plant species as nonnative plant species [10]. Areas susceptible to an invasive plant/fire cycle and extensive, long-term plant community conversion might remain localized for many decades and then spread to other areas following an extreme event such as an "abnormally high magnitude" wildfire [67]. Lengthening of the freeze-free season, increased frequency of wet winters, and an earlier onset and lengthening of the fire season are predicted to further the invasive plant/fire feedback loop [1,133].

Several publications are available that discuss implications and potential effects of climate change in the Sonoran Desert. Examples are shown in table 4.

Table 4—Examples of literature addressing aspects of the effects of climate change in the Sonoran Desert.
TopicTitleReference
Impacts on cacti; reviewGlobal change impacts on cacti (Cactaceae): Current threats, challenges and conservation solutions.Hultine et al. (2023) [60]
Impacts on succulents; reviewTissue succulence in plants: Carrying water for climate change.Pérez-López et al. (2023) [102]
Impacts on invasive plants; reviewThe biogeography of invasive plants – projecting range shifts with climate change.Bradley (2023) [23]
Impacts on winter annualsWhat common-garden experiments tell us about climate responses in plants.Schwinning et al. (2022) [113]
Impacts on dominant plantsDominant Sonoran Desert plant species have divergent phenological responses to climate change.Zachmann et al. (2021) [138]
Desert vegetation response to droughtAssessing vegetation response to multi-scalar drought across the Mojave, Sonoran, Chihuahuan Deserts and Apache Highlands in the Southwest United States.Khatri-Chhetri (2021) [65]
Impacts on desert plantsLife history traits predict colonization and extinction lags of desert plant species since the Last Glacial Maximum.Butterfield et al. (2019) [35]
Impacts on succulent plantsSucculent plant diversity as natural capital.Grace (2019) [57]
Methods for estimating impactsClimate change vulnerability and adaptation strategies for natural communities: Piloting methods in the Mojave and Sonoran Deserts.Comer et al. (2012) [40]
Older reviewDisturbance and climate change in United States/Mexico borderland plant communities: A state-of-the-knowledge review.McPherson and Weltzin (2000) [86]

As climate conditions change, temperatures increase, frequency of freezing temperatures decreases, and precipitation patterns shift, specific sites and total area susceptible to nonnative plant invasion are likely to change, and dominant nonnative species are likely to change. Seasonal changes in precipitation may be more influential than total rainfall on the persistence of many plant species [44], and thus impact which nonnative species dominate areas of invaded Sonoran desert scrub. For example, while wetter winters since 1976 encouraged the spread of red brome in the upper Sonoran Desert of central and southern Arizona, a shift to drier winters and wetter summers might favor buffelgrass dominance [121]. Warming trends, in general, are likely to favor buffelgrass range expansion in the Sonoran Desert [5,64,131].

2020 LANDFIRE Biophysical Settings — Historical Fire Regime Characteristics
Biophysical SettingMean Fire Interval (years)Fire Severity Percent (%)
CodeFire Regime GroupLowMixedReplacementAllLowMixedReplacement
Series 11090 - Sonoran Paloverde-Mixed Cacti Desert Scrub
11090_15_25V-B1049104900100
11090_14V-B1056105600100
11090_4_13V-B1284128400100
Series 10910 - Sonoran Mid-Elevation Desert Scrub
10910_25V-A37437400100
10910_15V-A35535500100
10910_13_14IV-B10410400100
Series 10900 - Sonoran Granite Outcrop Desert Scrub
10900_14V-B51351300100
Summary
Minimum10410400100
Maximum1284128400100
Mean67667600100
Median51351300100
Percentage of fires in 3 fire severity classes, derived from LANDFIRE succession modeling. Replacement-severity fires cause >75% kill or top-kill of the upper canopy layer; mixed-severity fires cause 26%-75%; low-severity fires cause <26%.
LANDFIRE. 2020. Biophysical settings models and descriptions, [Online]. Washington, DC: U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior; U.S. Geological Survey; Arlington, VA: The Nature Conservancy, (Producers). Available: https://www.landfirereview.org/search.php [96496]

Table A1

List of plant species mentioned in this synthesis. Asterisks indicate nonnative species.
Life formCommon nameScientific name
Cactusbarrel cactusFerocactus spp.
Cactusbuckhorn chollaCylindropuntia acanthocarpa
CactuschollaCylindropuntia spp.
Cactusfoxtail cactusEscobaria spp.
Cactusglobe cactusMammillaria spp.
Cactushedgehog cactusEchinocereus spp.
Cactusorganpipe cactusStenocereus thurberi
CactuspricklypearOpuntia spp.
CactussaguaroCarnegiea gigantea
Cactusteddybear chollaCylindropuntia bigelovii
Cactuspinkflower hedgehog cactusEchinocereus fendleri
Annual and perennial graminoidbromeBromus spp.
Annual graminoidArabian schismusSchismus arabicus*
Annual graminoidcommon Mediterranean grassSchismus barbatus*
Annual graminoidMediterranean grassSchismus spp.*
Annual graminoidred bromeBromus rubens*
Annual graminoidsixweeks grassVulpia octoflora
Annual graminoidsixweeks threeawnAristida adscensionis
Perennial graminoidbig sacatonSporobolus wrightii
Perennial graminoidblack gramaBouteloua eriopoda
Perennial graminoidbuffelgrassPennisetum ciliare*
Perennial graminoidbush muhlyMuhlenbergia porteri
Perennial graminoidcrimson fountaingrassPennisetum setaceum*
Perennial graminoidfountaingrassPennisetum spp.
Perennial graminoidlarge-spike bristlegrassSetaria macrostachya
Perennial graminoidLehmann lovegrassEragrostis lehmanniana*
Perennial graminoidlovegrassEragrostis spp. *
Perennial graminoidlow woolygrassDasyochloa pulchella
Perennial graminoidsoft feather pappusgrassEnneapogon cenchroides*
Perennial graminoidtobosagrassPleuraphis mutica
Perennial graminoidweeping lovegrassEragrostis curvula*
Annual forbarugulaEruca vesicaria*
Annual forbcurvenut combseedPectocarya recurvata
Annual forbdesert IndianwheatPlantago ovata
Annual forbMenzies' fiddleneckAmsinckia menziesii
Annual forbredstem stork's billErodium cicutarium*
Annual forbSahara mustardBrassica tournefortii*
Perennial forb/vineslender janusiaJanusia gracilis
Perennial forbtrailing windmillsAllionia incarnata
Forb/subshrubdesert globemallowSphaeralcea ambigua
Forb/subshrubdesert zinniaZinnia acerosa
Forb/subshrublongflower tube tongueJusticia longii
Forb/subshrubRocky Mountain zinniaZinnia grandiflora
Forb/subshrubrough menodoraMenodora scabra
Forb/subshrubwhitestem paperflowerPsilostrophe cooperi
Shrub/subshrubbrittle bushEncelia farinosa
Shrub/subshrubcommon sotolDasylirion wheeleri
Shrub/subshrubeastern Mojave buckwheatEriogonum fasciculatum
Shrub/subshrubfairydusterCalliandra eriophylla
Shrub/subshrubHall's shrubby spurgeTetracoccus hallii
Shrub/subshrublittleleaf ratanyKrameria erecta
Shrub/subshrubNevada jointfirEphedra nevadensis
Shrub/subshrubsotolDasylirion spp.
Shrub/subshrubtriangle bur ragweedAmbrosia deltoidea
Shrub/subshrubwhite bursageAmbrosia dumosa
Shrub/subshrubwhite ratanyKrameria grayi
ShrubBerlandier's wolfberryLycium berlandieri
ShrubblackbrushColeogyne ramosissima
ShrubcreosotebushLarrea tridentata
ShrubdesertsennaSenna armata
Shrubdesert-thornLycium spp.
ShrubjojobaSimmondsia chinensis
Shrubnarrowleaf goldenbushEricameria linearifolia
ShrubocotilloFouquieria splendens
ShrubphysicnutJatropha cuneata
ShrubsaltbushAtriplex spp.
Shrubsangre de cristoJatropha cardiophylla
Shrub/treeacaciaAcacia spp., Senegalia spp., Vachelia spp.
Shrub/treeBigelow's nolinaNolina bigelovii
Shrub/treeburseraBursera spp.
Shrub/treecatclaw acaciaSenegalia greggii
Shrub/treecrucifixion thornCanotia holacantha
Shrub/treedesert ironwoodOlneya tesota
Shrub/treehopbushDodonaea spp.
Shrub/treeKearney's sumacRhus kearneyi
Shrub/treepaloverdeParkinsonia spp.
Shrub/treeyellow paloverdeParkinsonia microphylla
Treeblue paloverdeParkinsonia florida
TreejuniperJuniperus spp.
TreeoakQuercus spp.
TreepinePinus spp.
Tree/shrubdesert willowChilopsis linearis
Tree/shrubelephant treeBursera microphylla
Tree/shrubGambel oakQuercus gambelii
Tree/shrubmesquiteProsopis spp.
Tree/shrubsmoketreePsorothamnus spinosus
Tree/shrubvelvet mesquiteProsopis velutina

Table A2

Examples of studies that measured stand structure and species composition in undisturbed Sonoran desert scrub communities or those disturbed by fire or nonnative plant invasions.
Site type(s)LocationData collected and site detailsReference
UndisturbedOrgan Pipe Cactus National Monument, ArizonaExamined vegetation patterns relative to site characteristics using cover and density data from 99 sites and analyzed via ordination analysisParker (1991) [98]
UndisturbedSaguaro National Park, ArizonaCompared frequency, cover, density, and diversity of all perennial plant species in permanent plots (with varied fire history) in 1990 and 2000, 30–40 years after cessation of livestock grazingFunicelli et al. (2001) [54]
UndisturbedSaguaro National Park, ArizonaAnnual census data spanning 75 years (1942–2016) documenting mortality and regeneration in a population of saguaroOrum et al. (2016) [97]
UndisturbedSanta Catalina Mountains, ArizonaDensity by size classes for shrubs and trees and by height classes for saguaro in 6 plant community types along an elevation gradient; age class composition; sites had history of livestock grazing from around 1880–1847Niering and Lowe (1984) [93]
UndisturbedSilverbell Mountains, ArizonaCover of dominant species at 10 stations across an elevation gradientTurner and Brown (1982) [125]
UndisturbedTumamoc Hill, ArizonaCoverage and density of all woody and succulent plants, including seedlings; data collected by mapping stems and crowns of all plants in study plots at irregular intervals from 1906 to 1978; age class compositionGoldberg and Turner (1986) [56]
UndisturbedMacDougal Crater, Sonora, MexicoCoverage and density of dominant perennial plants, 1907-1986; age class composition in 1959Turner (1990) [124]
Burned and UnburnedArizona Upland subdivision of the Sonoran Desert, 13 sites in ArizonaCover and density of perennial plant species on paired burned and unburned plots with time-since-fire ranging from 8 to 33 yearsShryock et al. (2015) [117]
Burned and UnburnedGranite Burn (1979) near Florence, ArizonaCompared cover and density of dominant perennials on burned and unburned sites, 7 to 19 months after fireMcLaughlin and Bowers (1982) [84]
Burned and UnburnedTonto National Monument, ArizonaRelative frequency, density, cover, and importance values given for perennial plants in burned and unburned plots; decades of repeat photography at several locationsPhillips (1997) [103]
Burned and UnburnedTonto National Forest (Arizona Upland subdivision), ArizonaCompared height, cover, and density between paired burned and unburned sites at 5 locations with time-since-fire ranging from <5 to 21 yearsAlford et al. (2001, 2005) [6,7]
Burned and UnburnedTonto National Forest, Mesa Ranger District, ArizonaHeight and diameters of plants (grouped by life forms) on 4 unburned, 1-ha sites; focused on fuel propertiesWilson et al. (1998) [135]
Burned and UnburnedTonto National Forest, Vista View Fire area, ArizonaPlant cover and status values (live, resprout, or dead) on paired burned and unburned plots over a 10-year postfire period; data grouped by growth forms similar to those used by Wilson et al. (1998) (tree—woody plant with primary trunk (3.0-3.5 m), shrub—large woody multi-stemmed plant (1.5-2.5 m), bush—low growing multi-stemmed woody plant (0.4-0.8 m), cactus—succulent (0.14-0.19 m), and yucca—fibrous woody (1.4 m))Narog and Wilson (2005) [89]
Invaded and UninvadedSanta Catalina Mountains, ArizonaPlant species cover and density, and saguaro size structure at 10 unburned and ungrazed buffelgrass patches with varied buffelgrass cover; regression models quantified differences in diversity, cover, and density with respect to buffelgrass coverOlsson et al. (2012) [96]

1. Abatzoglou, John T.; Kolden, Crystal A. 2011. Climate change in western US deserts: Potential for increased wildfire and invasive annual grasses. Rangeland Ecology & Management. 64(5): 471-478. [92501]

2. Abella, S. R. 2009. Post-fire plant recovery in the Mojave and Sonoran Deserts of western North America. Journal of Arid Environments. 73(8): 699-707. [81859]

3. Abella, Scott R. 2010. Disturbance and plant succession in the Mojave and Sonoran Deserts of the American Southwest. International Journal of Environmental Research and Public Health. 7(4): 1248-1284. [95419]

4. Abella, Scott R.; Chiquoine, Lindsay P.; Backer, Dana M. 2013. Soil, vegetation, and seed bank of a Sonoran Desert ecosystem along an exotic plant (Pennisetum ciliare) treatment gradient. Environmental Management. 52(4): 946-957. [95873]

5. Albuquerque, Fabio Suzart; Macias-Rodriguez, Miguel Angel; Burquez, Alberto; Astudillo-Scalia, Yaiyr. 2019. Climate change and the potential expansion of buffelgrass (Cenchrus ciliaris L., Poaceae) in biotic communities of Southwest United States and northern Mexico. Biological Invasions. 21(11): 3335-3347. [95639]

6. Alford, Eddie J.; Brock, John H.; Gottfried, Gerald J. 2005. Effects of fire on Sonoran Desert plant communities. In: Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; Edminster, Carleton B., comps. Connecting mountain islands and desert seas: Biodiversity and management of the Madrean Archipelago II; 5th conference on research and resource management in the southwestern deserts; 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 451-454. [61785]

7. Alford, Eddie Jim. 2001. The effects of fire on Sonoran Desert plant communities. Phoenix, Arizona: Arizona State University 122 p. Dissertation. [95759]

8. Arriaga, Laura; Castellanos, Alejandro E.; Moreno, Elizabeth; Alarcon, Jesus. 2004. Potential ecological distribution of alien invasive species and risk assessment: A case study of buffel grass in arid regions of Mexico. Conservation Biology. 18(6): 1504-1514. [95670]

9. Aslan, Clare E. 2021. Land management objectives and activities in the face of projected fire regime change in the Sonoran Desert. Journal of Environmental Management. 280: 111644. [95429]

10. Aslan, Clare E.; Dickson, Brett G. 2020. Non-native plants exert strong but under-studied influence on fire dynamics. NeoBiota. 61: 47-64. [95774]

11. Bahre, Conrad J. 1985. Wildfire in southeastern Arizona between 1859 and 1890. Desert Plants. 7(4): 190-194. [37739]

12. Barbour, Michael G.; Billings, William Dwight, eds. 1988. North American terrestrial vegetation. Cambridge, New York: Cambridge University Press. 434 p. [13876]

13. Barbour, Michael G.; Keeler-Wolf, Todd; Schoenherr, Allan A., eds. 2007. Terrestrial vegetation of California, 3rd ed. Berkeley, CA: University of California Press. 712 p. [82605]

14. Barron, Kara Lynn. 2018. Fire and reseeding effects on Arizona upland plant community composition and a preliminary floristic inventory of Cave Creek Regional Park. Phoenix, AZ: Arizona State University: 75 p. Thesis. [96491]

15. Benson, Lyman. 1982. The cacti of the United States and Canada. Stanford, CA: Stanford University Press. 1044 p. [1513]

16. Bertelsen, David C. 2018. Thirty-seven years on a mountain trail: Vascular flora and flowering phenology of the Finger Rock Canyon Watershed, Santa Catalina Mountains, Arizona. Desert Plants. 34(1-2): 1-247. [95896]

17. Bohrer, Vorsila L. 1992. New life from ashes II: A tale of burnt brush. Desert Plants. 10(3): 122-125. [18805]

18. Bowers, Janice E. 2005. Effects of drought on shrub survival and longevity in the northern Sonoran Desert. Journal of the Torrey Botanical Society. 132(3): 421-431. [96659]

19. Bowers, Janice E.; Bean, Travis M.; Turner, Raymond M. 2006. Two decades of change in the distribution of exotic plants at the Desert Laboratory, Tucson, Arizona. Madrono. 53(3): 252-263. [95675]

20. Bowers, Janice E.; Turner, Raymond M.; Burgess, Tony L. 2004. Temporal and spatial patterns in emergence and early survival of perennial plants in the Sonoran Desert. Plant Ecology. 172(1): 107-119. [96658]

21. Bowers, Michael A.; Lowe, Charles H. 1986. Plant-form gradients on Sonoran Desert bajadas. Oikos. 46(3): 284-291. [10864]

22. Bracamonte, Jesus Arturo; Tinoco-Ojanguren, Clara; Coronado, Maria Esther Sanchez; Molina-Freaner, Francisco. 2017. Germination requirements and the influence of buffelgrass invasion on a population of Mammillaria grahamii in the Sonoran Desert. Journal of Arid Environments. 137: 50-59. [91502]

23. Bradley, Bethany A. 2023. The biogeography of invasive plants - projecting range shifts with climate change. In: Ziska, Lewis H., ed. Invasive species and global climate change. 2nd ed. CAB International: 248-259. [98367]

24. Brooks, M. L.; Matchett, J. R. 2006. Spatial and temporal patterns of wildfires in the Mojave Desert, 1980-2004. Journal of Arid Environments. 67(Supplement): 148-164. [65283]

25. Brooks, Matthew L. 1999. Alien annual grasses and fire in the Mojave Desert. Madrono. 46(1): 13-19. [34386]

26. Brooks, Matthew L.; Esque, Todd C. 2002. Alien plants and fire in desert tortoise (Gopherus agassizii) habitat of the Mojave and Colorado deserts. Chelonian Conservation Biology. 4(2): 330-340. [44468]

27. Brooks, Matthew L.; Minnich, Richard A. 2006. Southeastern deserts bioregion. In: Sugihara, Neil G.; van Wagtendonk, Jan W.; Shaffer, Kevin E.; Fites-Kaufman, Joann; Thode, Andrea E., eds. Fire in California's ecosystems. Berkeley, CA: University of California Press: 391-414. [65559]

28. Brooks, Matthew L.; Minnich, Richard A.; Matchett, John R. 2018. Southeastern deserts bioregion. In: van Wagtendonk, Jan W.; Sugihara, Neil G.; Stephens, Scott L.; Thode, Andrea E.; Shaffer, Kevin E.; Fites-Kaufman, Jo Ann, eds. Fire in California's ecosystems. 2nd ed. Oakland, CA: University of California Press: 353-378. [93914]

29. Brooks, Matthew L.; Pyke, David A. 2001. Invasive plants and fire in the deserts of North America. In: Galley, Krista E. M.; Wilson, Tyrone P., eds. Proceedings of the invasive species workshop: The role of fire in the control and spread of invasive species. Fire conference 2000: 1st national congress on fire ecology, prevention, and management; 2000 November 27 - December 1; San Diego, CA. Misc. Publ. No. 11. Tallahassee, FL: Tall Timbers Research Station.: 1-14. [40491]

30. Brown, Albert L. 1950. Shrub invasion of southern Arizona desert grassland. Journal of Range Management. 3(3): 172-177. [4452]

31. Brown, David E., ed. 1982. Biotic communities of the American Southwest--United States and Mexico. Desert Plants: Special Issue. 4(1-4): 1-342. [62041]

32. Brown, David E.; Minnich, Richard A. 1986. Fire and changes in creosote bush scrub of the western Sonoran Desert, California. The American Midland Naturalist. 116(2): 411-422. [537]

33. Brunelle, A.; Minckley, T. A.; Blissett, S.; Cobabe, S. K.; Guzman, B. L. 2010. A 8000 year fire history from an Arizona/Sonora borderland cienega. Journal of Arid Environments. 74(4): 475-481. [81933]

34. Burquez-Montijo, Alberto; Miller, Mark E.; Martinez-Yrizar, Angelina. 2002. Mexican grasslands, thornscrub, and the transformation of the Sonoran Desert by invasive exotic buffelgrass (Pennisetum ciliare). In: Tellman, Barbara, ed. Invasive exotic species in the Sonoran region. Arizona-Sonora Desert Museum Studies in Natural History. Tucson, AZ: The University of Arizona Press; The Arizona-Sonora Desert Museum: 126-146. [48657]

35. Butterfield, Bradley J.; Holmgren, Camille A.; Anderson, R. Scott; Betancourt, Julio L. 2019. Life history traits predict colonization and extinction lags of desert plant species since the Last Glacial Maximum. Ecology. 100(10): e02817. [98368]

36. Castellanos, A. E.; Molina, F. E. 1990. Differential survivorship and establishment in Simmondsia chinensis (jojoba). Journal of Arid Environments. 19(1): 65-76. [14982]

37. Cave, George H.; Patten, Duncan T. 1984. Short-term vegetation responses to fire in the upper Sonoran Desert. Journal of Range Management. 37(6): 491-496. [610]

38. Chambers, J. C.; Brooks, M. L.; Germino, M. J.; Maestas, J. D.; Board, D. I.; Jones, M. O.; Allred, B. W. 2019. Operationalizing resilience and resistance concepts to address invasive grass-fire cycles. Frontiers in Ecology and Evolution. 7: 185. [95797]

39. Cohn, Jeffrey P. 2005. Tiff over Tamarisk: Can a nuisance be nice, too? Bioscience. 55(8): 648-654. [55523]

40. Comer, P. J.; Young, B.; Schulz, K.; Kittel, G.; Unnasch, D.; Hammerson, G.; Smart, L.; Hamilton, H.; Auer, S.; Smyth, R.; Hak, J. 2012. Climate change vulnerability and adaptation strategies for natural communities: Piloting methods in the Mojave and Sonoran Deserts. Report to the U.S. Fish and Wildlife Service. Arlington, VA: NatureServe. 77 p. [97233]

41. Conver, Joshua L.; Foley, Theresa; Winkler, Daniel E.; Swann, Don E. 2017. Demographic changes over >70 yr in a population of saguaro cacti (Carnegiea gigantea) in the northern Sonoran Desert. Journal of Arid Environments. 139: 41-48. [96660]

42. Cox, Jerry R.; Martin-R., Martha; Ibarra-F., Fernando. 1995. Climatic effects on buffelgrass productivity in the Sonoran Desert. In: Wester, David B.; Britton, Carlton M., eds. Research highlights: Noxious brush and weed control: Range, wildlife and fisheries management. Lubbock, TX: Texas Tech University, College of Agricultural Sciences and Natural Resources. Vol. 26: 25-26. [26623]

43. D'Antonio, Carla M.; Vitousek, Peter M. 1992. Biological invasions by exotic grasses, the grass/fire cycle, and global change. Annual Review of Ecology and Systematics. 23: 63-87. [20148]

44. Dickson, Brett G.; Sisk, Thomas D.; Sesnie, Steven E.; Bradley, Bethany A. 2015. Integrated spatial models of non-native plant invasion, fire risk, and wildlife habitat to support conservation of military and adjacent lands in the arid Southwest. SERDP Project RC-1722. Flagstaff, AZ: Northern Arizona University Landscape Conservation Initiative, School of Earth Sciences and Environmental Sustainability. 106 p. [95722]

45. Drake, Joseph; Griffis-Kyle, Kerry; McIntyre, Nancy. 2016. Landscape connectivity of isolated waters for wildlife in the Sonoran Desert. Report submitted to the Desert Landscape Conservation Cooperative and the Bureau of Reclamation. Lubbock, TX: Texas Tech University. 173 p. [95872]

46. Esque, Todd C.; Burquez, Alberto; Schwalbe, Cecil R.; Van Devender, Thomas R.; Anning, Pamela J.; Nijhuis, Michelle J. 2002. Fire ecology of the Sonoran Desert tortoise. In: Van Devender, Thomas R., ed. The Sonoran Desert tortoise: Natural history, biology, and conservation. Arizona-Sonora Desert Museum Studies in Natural History. Tucson, AZ: University of Arizona Press; Arizona-Sonora Desert Museum: 312-333. [67598]

47. Esque, Todd C.; Schwalbe, Cecil; Lissow, Jessica A.; Haines, Dustin F.; Foster, Danielle; Garnett, Megan C. 2006. Buffelgrass fuel loads in Saguaro National Park, Arizona, increase fire danger and threaten native species. Park Science. 24(2): 33-37. [69872]

48. Esque, Todd C.; Schwalbe, Cecil R. 2002. Alien annual grasses and their relationships to fire and biotic change in Sonoran Desert scrub. In: Tellman, Barbara, ed. Invasive exotic species in the Sonoran region. Arizona-Sonora Desert Museum Studies in Natural History. Tucson, AZ: University of Arizona Press; Arizona-Sonora Desert Museum: 165-194. [48660]

49. Esque, Todd C.; Schwalbe, Cecil R.; Haines, Dustin F.; Halvorson, William L. 2004. Saguaros under siege: Invasive species and fire. Desert Plants. 20(1): 49-55. [91075]

50. Esque, Todd C.; Webb, Robert H.; Wallace, Cynthia S. A.; van Riper, Charles, III; McCreedy, Chris; Smythe, Lindsay. 2013. Desert fires fueled by native annual forbs: Effects of fire on communities of plants and birds in the lower Sonoran Desert of Arizona. The Southwestern Naturalist. 58(2): 223-233. [87384]

51. Falk, Donald A. 2013. Are Madrean ecosystems approaching tipping points? Anticipating interactions of landscape disturbance and climate change. In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane G.; Collins, Loa C. Merging science and management in a rapidly changing world: Biodiversity and management of the Madrean Archipelago III and 7th conference on research and resource management in the southwestern deserts; 2012 May 1-5. RMRS-P-67. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 40-47. [90340]

52. Felix-Burruel, Ricardo E.; Larios, Eugenio; Gonzalez, Edgar J.; Burquez, Alberto. 2021. Episodic recruitment in the saguaro cactus is driven by multidecadal periodicities. Ecology. 102(10): e03458. [96643]

53. Fuentes-Ramirez, Andres; Veldman, Joseph W.; Holzapfel, Claus; Moloney, Kirk A. 2016. Spreaders, igniters, and burning shrubs: Plant flammability explains novel fire dynamics in grass-invaded deserts. Ecological Applications. 26(7): 2311-2322. [91269]

54. Funicelli, Carianne S.; Anning, Pamela J.; Turner, Dale S. 2001. Long-term vegetation monitoring at Saguaro National Park: A decade of change. Tech. Rep. No. 70. Tucson, AZ: USGS Sonoran Desert Field Station, University of Arizona. 76 p. [95824]

55. Fusco, Emily J.; Finn, John T.; Balch, Jennifer K.; Nagy, R. Chelsea; Bradley, Bethany A. 2019. Invasive grasses increase fire occurrence and frequency across US ecoregions. PNAS. 116(47): 23594-23599. [95452]

56. Goldberg, Deborah E.; Turner, Raymond M. 1986. Vegetation change and plant demography in permanent plots in the Sonoran Desert. Ecology. 67(3): 695-712. [4410]

57. Grace, Olwen M. 2019. Succulent plant diversity as natural capital. Plants People Planet. 1(4): 336-345. [98369]

58. Gray, Miranda E.; Dickson, Brett G.; Zachmann, Luke J. 2014. Modelling and mapping dynamic variability in large fire probability in the lower Sonoran Desert of south-western Arizona. International Journal of Wildland Fire. 23(8): 1108-1118. [88832]

59. Gray, Miranda E.; Zachmann, Luke J.; Dickson, Brett G.; Gage, Josh. 2018. An early-warning mapping tool for forecasting fire risk on DoD lands in the arid west. Project 17-834. Washington, D.C: Department of Defense Legacy Resource Management Program. 15 p. [96479]

60. Hultine, Kevin R.; Hernandez-Hernandez, Tania; Williams, David G.; Albeke, Shannon E.; Tran, Newton; Puente, Raul; Larios, Eugenio. 2023. Global change impacts on cacti (Cactaceae): Current threats, challenges and conservation solutions. Annals of Botany. [In press]: 1-13. [98365]

61. Humphrey, Robert R. 1958. The desert grassland: A history of vegetational change and an analysis of causes. The Botanical Review. 24(4): 193-252. [5270]

62. Humphrey, Robert R. 1974. Fire in the deserts and desert grassland of North America. In: Kozlowski, T. T.; Ahlgren, C. E., eds. Fire and ecosystems. New York: Academic Press: 365-400. [14064]

63. Innes, Robin J. 2023. Brassica tournefortii, Sahara mustard. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Missoula Fire Sciences Laboratory (Producer). Available: https://www.fs.usda.gov/database/feis/plants/forb/bratou/all.html. [98218]

64. Jarnevich, Catherine S.; Young, Nicholas E.; Talbert, Marian; Talbert, Colin. 2018. Forecasting an invasive species' distribution with global distribution data, local data, and physiological information. Ecosphere. 9(5): e02279. [95749]

65. Khatri-Chhetri, Pratima; Hendryx, Sean M.; Hartfield, Kyle A.; Crimmins, Michael A.; van Leeuwen, Willem J. D.; Kane, Van R. 2021. Assessing vegetation response to multi-scalar drought across the Mojave, Sonoran, Chihuahuan Deserts and Apache Highlands in the Southwest United States. Remote Sensing. 13(6): 1103. [98373]

66. Klinger, Rob; Brooks, Matt. 2017. Alternative pathways to landscape transformation: Invasive grasses, burn severity and fire frequency in arid ecosystems. Journal of Ecology. 105(6): 1521-1533. [92303]

67. Klinger, Rob; Underwood, Emma C.; McKinley, Randy; Brooks, Matthew L. 2021. Contrasting geographic patterns of ignition probability and burn severity in the Mojave Desert. Frontiers in Ecology and Evolution. 9: 593167. [95796]

68. Klinger, Robert C.; Brooks, Matthew L.; Randall, John M. 2018. Fire and invasive plants. In: van Wagtendonk, Jan W.; Sugihara, Neil G.; Stephens, Scott L.; Thode, Andrea E.; Shaffer, Kevin E.; Fites-Kaufman, Jo Ann, eds. Fire in California's ecosystems. 2nd ed. Oakland, CA: University of California Press: 459-476. [92960]

69. LANDFIRE. 2020. Biophysical settings layer, CONUS (LANDFIRE 2.2.0). In: LANDFIRE download data mosaic products, [Online]. U.S. Department of the Interior, Geological Survey; U.S. Department of Agriculture (Producer). Available: https://www.landfire.gov/version_download.php [2023, June 20]. [98050]

70. LANDFIRE. 2022. Existing Vegetation Type Layer, LANDFIRE 2.3.0. In: LANDFIRE download data mosaic products, [Online]. U.S. Department of the Interior, Geological Survey; U.S. Department of Agriculture (Producer). Available: https://www.landfire.gov/version_download.php [2024, March 13]. [99091]

71. LANDFIRE. 2020. Biophysical settings models and descriptions, [Online]. Washington, DC: U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior; U.S. Geological Survey; Arlington, VA: The Nature Conservancy, (Producers). Available: http://www.landfirereview.org/search.php [2022, February 2]. [96496]

72. Lata, Mary. 2022. After the fire? Setting the stage for fire, [webinar]. Miami, AZ: University of Arizona Cooperative Extension, (Producer). Available: https://www.youtube.com/watch?v=a08sh_ma1BA. [98013]

73. Latting, June, ed. 1976. Symposium proceedings: Plant communities of southern California. Special Publication No. 2. Berkeley, CA: California Native Plant Society. 164 p. [1414]

74. Leopold, Aldo. 1924. Grass, brush, timber, and fire in southern Arizona. Journal of Forestry. 22(6): 1-10. [5056]

75. Malusa, James R. 2024. Personal communication [Email to Kris Zouhar]. February-April. Information regarding distribution of Sonoran desert scrub Biophysical Settings and Existing Vegetation Types. Tucson, AZ: School of Natural Resources and the Environment, The University of Arizona. Documents on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT; FEIS files. [99090]

76. Margolis, Ellis Q.; Malevich, Steven B. 2016. Historical dominance of low-severity fire in dry and wet mixed-conifer forest habitats of the endangered terrestrial Jemez Mountains salamander (Plethodon neomexicanus). Forest Ecology and Management. 375: 12-26. [91078]

77. Marshall, V. M.; Lewis, M. M.; Ostendorf, B. 2012. Buffel grass (Cenchrus ciliaris) as an invader and threat to biodiversity in arid environments: A review. Journal of Arid Environments. 78: 1-12. [95607]

78. Martin-R., Martha; Cox, Jerry R.; Ibarra-F., F.; Alston, Diana G.; Banner, Roger E.; Malecheck, John C. 1999. Spittlebug and buffelgrass responses to summer fires in Mexico. Journal of Range Management. 52(6): 621-625. [69189]

79. Martin-R., Martha H.; Cox, Jerry R.; Ibarra-Flores, Fernando. 1995. Climatic effects on buffelgrass productivity in the Sonoran Desert. Journal of Range Management. 48(1): 60-63. [24432]

80. McAuliffe, J. R. 1995. The aftermath of wildfire in the Sonoran Desert. The Sonoran Quarterly. 49: 4-8. [46026]

81. McAuliffe, Joseph R. 1988. Markovian dynamics of simple and complex desert plant communities. The American Naturalist. 131(4): 459-490. [6744]

82. McDonald, C. J.; McPherson, G. R. 2011. Fire behavior characteristics of buffelgrass-fueled fires and native plant community composition in invaded patches. Journal of Arid Environments. 75(11): 1147-1154. [83896]

83. McDonald, Christopher J.; McPherson, Guy R. 2013. Creating hotter fires in the Sonoran Desert: Buffelgrass produces copious fuels and high fire temperatures. Fire Ecology. 9(2): 26-39. [87751]

84. McLaughlin, Steven P.; Bowers, Janice E. 1982. Effects of wildfire on a Sonoran Desert plant community. Ecology. 63(1): 246-248. [1619]

85. McPherson, Guy R. 1995. The role of fire in the desert grasslands. In: McClaran, Mitchel P.; Van Devender, Thomas R., eds. The desert grassland. Tucson, AZ: The University of Arizona Press: 130-151. [26576]

86. McPherson, Guy R.; Weltzin, Jake F. 2000. Disturbance and climate change in United States/Mexico borderland plant communities: A state-of-the-knowledge review. Gen. Tech. Rep. RMRS-GTR-50. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 24 p. [38752]

87. Medeiros, Scott; Drezner, Taly Dawn. 2012. Vegetation, climate, and soil relationships across the Sonoran Desert. Ecoscience. 19(2): 148-160. [96666]

88. Moloney, Kirk A.; Mudrak, Erika L.; Fuentes-Ramirez, Andres; Parag, Hadas; Schat, Marjolein; Holzapeel, Claus. 2019. Increased fire risk in Mojave and Sonoran shrublands due to exotic species and extreme rainfall events. Ecosphere. 10(2): e02592. [96102]

89. Narog, Marica; Wilson, Ruth. 2005. Post-fire saguaro community: Impacts on associated vegetation still apparent 10 years later. In: Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; Edminster, Carleton B., comps. Connecting mountain islands and desert seas: Biodiversity and management of the Madrean Archipelago II; 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 421-425. [61774]

90. NatureServe. 2009. International ecological classification standard: Terrestrial ecological classifications. In: NatureServe Central Databases. Arlington, VA: NatureServe (Producer). 1172 p. Available: http://downloads.natureserve.org/get_data/data_sets/veg_data/nsDescriptions.pdf [2023, August 3]. [94380]

91. NatureServe. 2023. NatureServe Explorer, [Online]. Arlington, VA: NatureServe (Producer). Available: https://explorer.natureserve.org/. [94379]

92. Nichol, A. A. The natural vegetation of Arizona. Tech. Bull. 68. Tucson, AZ: University of Arizona, Agricultural Experiment Station: 189-230. [Revisions by W. S. Phillips]. [3928]

93. Niering, William A.; Lowe, Charles H. 1984. Vegetation of the Santa Catalina Mountains: Community types and dynamics. Vegetatio. 58(1): 3-28. [12037]

94. O'Connor, Christopher D.; Garfin, Gregg M.; Falk, Donald A.; Swetnam, Thomas W. 2011. Human pyrogeography: A new synergy of fire, climate and people is reshaping ecosystems across the globe. Geography Compass. 5/6: 329-350. [85677]

95. Olsson, Aaryn D.; Betancourt, Julio L.; Crimmins, Michael A.; Marsh, Stuart E. 2012. Constancy of local spread rates for buffelgrass (Pennisetum ciliare L.) in the Arizona upland of the Sonoran Desert. Journal of Arid Environments. 87: 136-143. [95751]

96. Olsson, Aaryn D.; Betancourt, Julio; McClaran, Mitchel P.; Marsh, Stuart E. 2012. Sonoran Desert ecosystem transformation by a C4 grass without the grass/fire cycle. Diversity and Distributions. 18(1): 10-21. [95472]

97. Orum, Thomas V.; Ferguson, Nancy; Mihail, Jeanne D. 2016. Saguaro (Carnegiea gigantea) mortality and population regeneration in the cactus forest of Saguaro National Park: Seventy-five years and counting. PLoS One. 11(8): e0160899. [95473]

98. Parker, Kathleen C. 1991. Topography, substrate, and vegetation patterns in the northern Sonoran Desert. Journal of Biogeography. 18(2): 151-163. [14979]

99. Patten, Duncan T.; Cave, George H. 1984. Fire temperatures and physical characteristics of a controlled burn in the upper Sonoran Desert. Journal of Range Management. 37(3): 277-280. [181]

100. Patten, Duncan. 1978. Productivity and production efficiency of an upper Sonoran Desert ephemeral community. American Journal of Botany. 65(8): 891-895. [95817]

101. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; Gottfried, Gerald J.; Haase, Sally M.; Harrington, Michael G.; Narog, Marcia G.; Sackett, Stephen S.; Wilson, Ruth C. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. [36978]

102. Perez-Lopez, Arely V.; Lim, Sung Don; Cushman, John C. 2023. Tissue succulence in plants: Carrying water for climate change. Journal of Plant Physiology. 289: 154081. [98366]

103. Phillips, Barbara G. 1997. History of fire and fire impacts at Tonto National Monument, Arizona. Technical Report No. 59. Tucson, AZ: University of Arizona. 79 p. [95476]

104. Rice, Peter M.; McPherson, Guy R.; Rew, Lisa J. 2008. Fire and nonnative invasive plants in the Interior West bioregion. In: Zouhar, Kristin; Smith, Jane Kapler; Sutherland, Steve; Brooks, Matthew L., eds. Wildland fire in ecosystems: Fire and nonnative invasive plants. Gen. Tech. Rep. RMRS-GTR-42-vol. 6. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 141-173. [70332]

105. Rodriguez-Rodriguez, Lucero; Stafford, Erin; Williams, Anna; Wright, Brian; Kribs, Christopher; Rios-Soto, Karin. 2017. A stage structured model of the impact of buffelgrass on saguaro cacti and their nurse trees. Tucson, AZ: Arizona State University. Tech. Rep. 2017: 29 p. [96069]

106. Rogers, G. F.; Vint, M. K. 1987. Winter precipitation and fire in the Sonoran Desert. Journal of Arid Environments. 13(1): 47-52. [5547]

107. Rogers, Garry F. 1985. Mortality of burned Cereus giganteus. Ecology. 66(2): 630-631. [2020]

108. Rogers, Garry F. 1986. Comparison of fire occurrence in desert and nondesert vegetation in Tonto National Forest, Arizona. Madrono. 33(4): 278-283. [5225]

109. Rogers, Garry F.; Steele, Jeff. 1980. Sonoran Desert fire ecology. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 15-19. [16036]

110. Rundel, Philip W.; Gibson, Arthur C. 1996. Ecological communities and processes in a Mojave Desert ecosystem: Rock Valley, Nevada. Cambridge; New York: Cambridge University Press. 369 p. [61799]

111. Rutman, Sue; Dickson, Lara. 2002. Management of buffelgrass on Organ Pipe Cactus National Monument, Arizona. In: Tellman, Barbara, ed. Invasive exotic species in the Sonoran region. Arizona-Sonora Desert Museum Studies in Natural History. Tucson, AZ: University of Arizona Press; Arizona-Sonora Desert Museum: 311-318. [48674]

112. Schmid, Mary K.; Rogers, Garry F. 1988. Trends in fire occurrence in the Arizona upland subdivision of the Sonoran Desert, 1955 to 1983. The Southwestern Naturalist. 33(4): 437-444. [6103]

113. Schwinning, Susanne; Lortie, Christopher J.; Esque, Todd; C.; DeFalco, Lesley A. 2022. What common-garden experiments tell us about climate responses in plants. Journal of Ecology. 110(2): 986-996. [98372]

114. Sesnie, Steven E.; Dickson, Brett G. 2015. A new time series remote sensing approach to mapping fine fuels in Sonoran Desert ecosystems. Final Report, Project ID: 10-1-04-7. Boise, ID: Joint Fire Science Program. 39 p. [96092]

115. Short, Karen C. 2022. Spatial wildfire occurrence data for the United States, 1992-2020 [FPA_FOD_20221014], [Online]. 6th Edition. Fort Collins, CO: Forest Service Research Data Archive. Available: https://doi.org/10.2737/RDS-2013-0009.6. [97936]

116. Shreve, Forrest; Wiggins, Ira L. 1964. Vegetation and flora of the Sonoran Desert. Stanford, CA: Stanford University Press. Vol. 1-2. 1740 p. [21016]

117. Shryock, Daniel F.; Esque, Todd C.; Chen, Felicia C. 2015. Topography and climate are more important drivers of long-term, post-fire vegetation assembly than time-since-fire in the Sonoran Desert, US. Journal of Vegetation Science. 26(6): 1134-1147. [92517]

118. Stephan, Kirsten; Miller, Melanie; Dickinson, Matthew B. 2010. First-order fire effects on herbs and shrubs: Present knowledge and process modeling needs. Fire Ecology. 6(1): 95-114. [81913]

119. Stevens, Jason; Falk, Donald A. 2009. Can buffelgrass invasions be controlled in the American Southwest? Using invasion ecology theory to understand buffelgrass success and develop comprehensive restoration and management. Ecological Restoration. 27(4): 417-427. [80810]

120. Summers, Ryan; Hovland, Matthew; Gibson, Yvette; Mata-Gonzalez, Ricardo. 2021. Climate-mediated shifts toward bunchgrass dominance 40 years after grazing cessation in Saguaro National Park, Arizona, USA. Applied Vegetation Science. 24(4): e12623. [97294]

121. Swetnam, Thomas W.; Betancourt, Julio L. 1998. Mesoscale disturbance and ecological response to decadal climatic variability in the American Southwest. Journal of Climate. 11(12): 3128-3147. [37431]

122. Thomas, P. A. 1991. Response of succulents to fire: A review. International Journal of Wildland Fire. 1(1): 11-22. [14991]

123. Thorne, Robert F. 1982. The desert and other transmontane plant communities of southern California. Aliso. 10(2): 219-257. [3768]

124. Turner, Raymond M. 1990. Long-term vegetation change at a fully protected Sonoran Desert site. Ecology. 7(2): 464-477. [10866]

125. Turner, Raymond M.; Brown, David E. 1982. Sonoran desertscrub. In: Brown, David E., ed. Biotic communities of the American Southwest—United States and Mexico. Desert Plants. 4(1-4): 181-221. [2375]

126. Turner, Raymond M.; Webb, Robert H.; Bowers, Janice E.; Hastings, James Rodney. 2003. The changing mile revisited: An ecological study of vegetation change with time in the lower mile of an arid and semiarid region. 1st ed.: Tucson, AZ: University of Arizona Press. 334 p. [95976]

127. Turner, Raymond M.; Webb, Robert H.; Esque, Todd C.; Rogers, Garry F. 2010. Repeat photography and low-elevation fire responses in the southwestern United States. In: Webb, Robert H.; Boyer, Diane E.; Turner, Raymond M. Repeat photography: Methods and applications in the natural sciences. Washington, DC: Island Press: 224-235. [95737]

128. Van Devender, Thomas R.; Reina, Ana Lilia. 2005. The Forgotten Flora of la Frontera. In: Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; Edminster, Carleton B., comps. Connecting mountain islands and desert seas: Biodiversity and management of the Madrean Archipelago II; 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 158-161. [95664]

129. Wang, Ophelia; Zachmann, Luke J.; Sesnie, Steven E.; Olsson, Aaryn; Dickson, Brett G. 2014. An iterative and targeted sampling design informed by habitat suitability models for detecting focal plant species over extensive areas. PLoS ONE. 9(7): e101196. [97437]

130. Webb, Robert H.; Boyer, Diane E.; Turner, Raymond M. 2010. Fire in the Sonoran Desert. In: Webb, Robert H.; Boyer, Diane E.; Turner, Raymond M. Repeat photography: Methods and applications in the natural sciences. 2010. Washington, D. C.: Island Press Publishing: 224-235. [96419]

131. Weiss, Jeremy L.; Overpeck, Jonathan T. 2005. Is the Sonoran Desert losing its cool? Global Change Biology. 11(12): 2065-2077. [95504]

132. Westerling, A. L.; Gershunov, A.; Brown, T. J.; R., Cayan. D.; Dettinger, M. D. 2003. Climate and wildfire in the western United States. American Meteorological Society. 84(5): 595-604. [92187]

133. Wilder, Benjamin T.; Jarnevich, Catherine S.; Baldwin, Elizabeth; Black, Joseph S.; Franklin, Kim A.; Grissom, Perry; Hovanes, Katherine A.; Olsson, Aaryn; Maulsa, Jim; Kibria, Abu S. M. G.; Li, Yue M.; Lien, Aaron M.; Ponce, Alejandro; Rowe, Julia A.; Soto, Jose R.; Stahl, Maya R.; Young, Nicholas E.; Betancourt, Julio L. 2021. Grassification and fast-evolving fire connectivity and risk in the Sonoran Desert, United States. Frontiers in Ecology and Evolution. 9: e955561 [+Supplement]. [96620]

134. Williams, David G.; Baruch, Zdravko. 2000. African grass invasion in the Americas: Ecosystem consequences and the role of ecophysiology. Biological Invasions. 2(2): 123-140. [70478]

135. Wilson, Ruth C.; Narog, Marcia G.; Corcoran, Bonni M. 1998. Unburned fuels in an Arizona upland saguaro-shrub community. In: Reynolds, J., comp. Finding faults in the Mojave: Abstracts of proceedings, 1998 desert research symposium; 1998 April 24; Redlands, CA. SBCMA Quarterly; 45(1-2). 109. [95501]

136. Winkler, Daniel D.; Conver, Joshua L.; Huxman, Travis E.; Dwann, Don E. 2018. The interaction of drought and habitat explain space-time patterns of establishment in saguaro (Carnegiea gigantea). Ecology. 99(3): 621–631. [93138]

137. Wright, Henry A.; Bailey, Arthur W. 1982. Fire ecology: United States and southern Canada. New York: John Wiley & Sons. 501 p. [2620]

138. Zachmann, Luke J.; Wiens, John F.; Franklin, Kim; Crausbay, Shelley D.; Landau, Vincent A.; Munson, Seth M. 2021. Dominant Sonoran Desert plant species have divergent phenological responses to climate change. Madrono. 68(4): 473-486. [96645]

Last updated June 4, 2025