Open Access
Translator Disclaimer
22 April 2021 Priority Species Lists to Restore Desert Tortoise and Pollinator Habitats in Mojave Desert Shrublands
Todd C. Esque, Lesley A. DeFalco, Gayle L. Tyree, K. Kristina Drake, Kenneth E. Nussear, Joseph S. Wilson
Author Affiliations +

Mojave Desert shrublands are home to unique plants and wildlife and are experiencing rapid habitat change due to unprecedented large-scale disturbances; yet, established practices to effectively restore disturbed landscapes are not well developed. A priority species list of native plant taxa was developed to guide seed collectors, commercial growers, resource managers, and restoration practitioners in support of the Bureau of Land Management's Mojave Desert Native Plant Program. We identify focal plant taxa that are important for habitats of the threatened Mojave desert tortoise (Gopherus agassizii), a widely distributed herbivore in low and middle elevations, and pollinator taxa, including mostly Lepidopterans and Apoidean bees, some of whose populations are in decline. We identified 201 unique plant taxa in the diets of tortoises, and 49 taxa that provide thermal cover for tortoises with some overlapping taxa that provide both diet and cover. We discuss 134 native pollinators associated with plants used for nectaring, larval hosts, or cover and nesting materials. Detailed plant species accounts describing the status-of-knowledge for 57 plant taxonomic groups including detailed information on life history, ecology, and pollinator syndrome relevant to restoration success, methods of seed harvesting, propagation, and historical use in restoration. Our approach for developing a priority plant species list for the Mojave Desert provides a data-guided listing of species for restoration practitioners and identifies knowledge gaps for future investigation.


Landscape-scale disturbances in the Mojave Desert are increasing in frequency and extent (Leu et al. 2008; Carter et al. 2020). As fire and other large surface disturbances increase in area and frequency, biotic communities lose native plant diversity and resilience to future perturbations may decline (Allison 2004; Tilman et al. 2006). The footprint of renewable energy development, such as utility-scale solar and wind farms, is rapidly expanding into areas of low and middle elevation desert shrublands, impacting sensitive habitats (BLM and US-DOE 2012; Vandergast et al. 2013).

Natural restoration of disturbed Mojave Desert vegetation is notoriously slow because of harsh climate conditions (Cody 2000; Miller et al. 2009). It is concerning, therefore, that the native seed reserves representing the regeneration potential of Mojave Desert shrublands are vulnerable to disturbances such as wildfires that incinerate seeds and reduce microsite availability (Esque et al. 2010), or are diminished by surface compaction, excavation, or burial (Scoles-Sciulla and DeFalco 2009). Seed dormancy allows them to survive low annual precipitation and high summer temperatures while awaiting favorable germination and growth conditions (Baskin and Baskin 2014) that coincide with adequate precipitation, when it is available, but are often challenged by seasonal to multi-annual drought (Beatley 1974; Turner 1994). Moreover, future climate for the Desert Southwest is expected to be hotter and drier than current conditions with potential shifts in the magnitude, frequency, and timing of precipitation pulses likely altering plant regeneration and persistence (Dai 2013; IPCC 2013). These challenges add to the difficulty of desert restoration and emphasize the importance of genetically diverse plant materials and appropriate guidelines for restoration practitioners to meet future challenges (Bradford et al. 2018).

Historically, conservation strategies strove to minimize new disturbances with designs to reduce the footprint size and minimize edge-to-area ratios. Such strategies are not always feasible due to the configuration of legacy infrastructure. Alternatively, new mitigation lands may be purchased (Spang et al. 1988); however, availability of suitable habitat has dwindled from high demand. One remaining alternative is active restoration of disturbed lands to support diverse, sustainable, resilient, and connected native biotic communities. This can be challenging and costly because native plant materials are not readily available, and consistently successful restoration methods remain poorly developed (Bainbridge 2007; DeFalco and Esque 2014; Olwell and Riibe 2016).

The availability of local plant materials is pivotal for restoring shrubland habitats (Johnson et al. 2010; Olwell and Riibe 2016); however, availability of commercially provided plant materials is limited for the Mojave Desert. Disturbances in low- and mid-elevation Mojave Desert shrublands are a major concern because these shrublands provide habitat for sensitive plants and wildlife including Mojave desert tortoises (Gopherus agassizii Cooper), which are protected by the Endangered Species Act (USFWS 2011). Restoration of degraded habitat is the highest research priority for the recovery of the tortoise (USFWS 2011; Drake et al. 2015). As an umbrella species (Tracy and Brussard 1994), the desert tortoise enhances habitat for other species through burrowing activities and indirectly through its legal protection (Nussear and Tuberville 2014). Importantly, desert tortoises depend on herbaceous annuals and grasses for food (Esque et al. 2014; Jennings and Berry 2015) and use shrubs for cover from extreme temperatures and predators (Nussear and Tuberville 2014).

In addition to recovering plant communities for desert tortoises, restoration success in the Mojave Desert includes sustaining habitats for diverse pollinator communities—a goal for many natural resource management agencies (Olwell and Riibe 2016). Plant–pollinator relationships are fundamental to the success of plant communities, and are thus critical to long-term, landscape-scale restoration. Environmental heterogeneity in the Southwest has driven speciation and supports a hotspot of arthropod diversity, particularly for scores of native bee plant specialists (Michener 1979; Minckley et al. 2000; Griswold et al. 2006; Gonzalez and Griswold 2013; Carril et al. 2018). Many plants provide pollinators with nectar or pollen, also providing important structure, cover, and nesting materials (Gonzalez and Griswold 2013). Prioritizing restoration species that benefit pollinators supports the National Strategy to Promote the Health of Honeybees and Other Pollinators (Vilsack and McCarthy 2015) and the National Seed Strategy (Olwell and Riibe 2016).

A focused strategy to understand the genetic variability of native species is paramount to restoring habitats damaged by large disturbances (Shryock et al. 2017). Some native plant species may be suitable for reintroduction across broad environmental gradients; however, others may be at risk of failure under narrower environmental conditions, or their establishment may have negative genetic consequences for local ecotypes (Lesica and Allendorf 1999). In addition, conservation of genetic variation within species is fundamental for adapting to future environmental change. Alternative restoration targets, genetic diversity, genetic structure, and future adaptability need to be considered in restoration programs (Rice and Emery 2003). While there has been progress toward understanding effectiveness of restoration treatments in the Mojave Desert (Bainbridge 2007; Weigand and Rodgers 2009), information is still lacking on appropriate plant materials that are successful in combination with emergent restoration practices. While we emphasize the importance of incorporating native plant species into restoration programs to benefit desert tortoises and native pollinators, the establishment of diverse native plant communities has many important ecosystem consequences (Maron et al. 2014). Diverse plant communities provide energy, essential nutrients, and cover for wildlife while strengthening food chains (Wilson 1987; Tallamy 2004; Burghardt et al. 2009; Burghardt and Tallamy 2013).

Priority species lists have been successfully deployed in other ecoregions to benefit local and regional plant material needs by increasing plant performance and socioeconomic value in Ethiopian drylands (Reubens et al. 2011), identifying phosphorus- and grazing-tolerant species in Australian grasslands (Graff and McIntyre 2014), and evaluating functional diversity of species in Brazilian Amazonian forests (Giannini et al. 2016). Here, our aim is to provide a synthesis of existing information on plants that can be used for restoration of tortoise and pollinator habitats in Mojave Desert shrublands. This priority species list can be used to guide seed collections, seed increase, cultivation of outplantings, and successful deployment of plant materials in disturbed areas in support of Bureau of Land Management's (BLM) Mojave Desert Native Plant Program.


Study Area

This study encompasses the Mojave Desert of the western United States including desert shrublands below ∼1500 m. The Mojave Desert is intermediate between the cold northerly Great Basin and the warmer Sonoran Desert to the south (MacMahon 1980). Block faulting exposes diverse geological parent materials (Keeler-Wolf 2007; Minnich 2007), resulting in a variety of soil textures, compositions, and depths driving the diversity, structure, and function of vegetation communities. The southern Sierra Nevada, Transverse, and San Bernardino mountains to the west cause a rain shadow of increased aridity across the Mojave Desert (MacMahon and Wagner 1985). Precipitation and temperature vary widely in the Mojave Desert on a daily, seasonal, interannual, and decadal basis (Hereford et al. 2006). Seasonal annual precipitation ranges between 65 and 190 mm for most of the Mojave desert with extremes from 47 mm in Death Valley, California, to as high as 253 mm at Pierce Ferry, Arizona (Turner 1994). Average daily minimum temperatures in January can reach 2.9 °C in Bishop, California, and average maximum monthly temperatures in July can soar to 38.3 °C in Death Valley, California (Minnich 2007). The Mojave Desert is classified as a desert shrubland with many woody shrubs and subshrubs generally less than 2 m tall, various herbaceous perennials, and distinct spring and summer annual floras. A diversity of trees, stem and leaf succulents, geophytes, and parasitic epiphytes (Turner 1994; Keeler-Wolf 2007) add regional distinction to shrubland habitat structure.

Plant Taxa Used in Tortoise Diets

We searched the literature for observations of plant taxa eaten by wild Mojave desert tortoises, particularly empirical studies quantifying diets through bite count studies in which every “bite” (opening and closure of mandibles) was counted (Esque et al. 2014). We supplemented that with qualitatively documented studies of diets through fecal analyses and visual behavioral observations (e.g., Hansen et al. 1976 and annotations in Grover and DeFalco 1995, respectively). The data are reported as the number of bites recorded, number of individual observations, or number of scats (fecal pellets) where species were recorded. The frequency of sites where species occurred in diets illustrates the geographic scope of their use across the Mojave Desert. Plant taxonomy follows the Jepson Flora Project (2018) or the Integrated Taxonomic Information System (ITIS 2021).

Figure 1.

Map of Mojave Desert (perimeter in dark gray outline) illustrating where shrub cover species and herbaceous diet species were documented within the Mojave desert tortoise recovery units (thin gray outlines). Site names are as follows: Desert Tortoise Natural Area (1), Hinkley (2), Fort Irwin National Training Center (3), Joshua Tree National Park (4), Stateline Pass (5), Ivanpah Valley (6), Arden Study Area (7), McCullough Pass (8), Piute Valley (9), Hidden Valley (10), Halfway (11), Lower Grand Canyon (12), Littlefield Site (13), Beaver Dam Wash (14), City Creek Site (15).


Plant Taxa Used as Tortoise Cover

Tortoise cover was monitored at nine sites spanning 37 y to identify shrub species used for cover from thermal extremes and protection from predators from the seven past sampling sites (greater Fort Irwin National Training Center and Stateline Pass in California; McCullough Pass, Piute Valley, Hidden Valley, and Halfway, Nevada; Littlefield Study Site, Arizona; and City Creek Study Site, Utah); and Burge (Arden, Nevada; 1978; Figure 1). Cover taxa were quantified by the frequency and percentage of total observations where each species was used by tortoises on first encounter during field work. In addition to species use, we report the frequency of sites where cover plants were used across the Mojave Desert to indicate the breadth of geographic use.

Plant Taxa Used by Pollinators

First, we identified pollinators that used the plant taxa already identified as diet and cover plants by tortoises. Then, we added common and widespread plant species and their pollinators that we found in the literature. From this list, we researched and highlighted the flower characteristics of the taxa thought to influence visitation by pollinators and known as the pollination syndrome (Baldwin et al. 2002). Pollination syndromes represent “suites of convergent floral traits [e.g., color, shape] hypothesized to adapt distantly related angiosperm species to particular types of pollen vectors” (Ollerton et al. 2009). For example, strongly scented white flowers that are tubular and bloom at night are a pollination syndrome particularly attractive to night-feeding moths. The bloom periods of each taxon are compiled (Jepson Flora Project 2018; SEINet 2020) to ensure diversity of species available, and facilitate planning resources for pollinators throughout the growing season. We documented visitation records, pollinator ecology, and general studies on pollinator communities of the Mojave Desert. Specific citations for all such literature are found in association with the plant taxonomic accounts ( Supplement 1 (naar-41-02-09_s01.docx), including independent Literature Cited).

Taxonomic Accounts on Life-History Characteristics and Restoration Potential

We summarized supplemental information on key characteristics of selected plant taxa ( Supplement 1 (naar-41-02-09_s01.docx)). Accounts include common names, plant functional groups, flowering times, elevational range, geographic distribution (relative to the Mojave desert tortoise), habitat types, flower form and shape, pollinator use, tortoise use, propagation and production techniques, and documented outcomes when used in restoration. When available, we included technical details on seed biology, seed collection, storage and handling, and establishment. Taxonomic accounts also include information about the recoverability of taxa based on short- and long-term recovery patterns across multiple wildfire studies (Shryock et al. 2014) and some species' affinities to disturbed habitat such as desert washes and roadsides.


Desert Tortoise Diet Taxa

We documented tortoise diets from fifteen studies at nine sites over a 22 y period across the Mojave and Colorado deserts (Table 1, Figure 1,   Supplement 2 (naar-41-02-09_s02.docx)). Bite-count studies were conducted at four sites (five studies) resulting in 210,095 bites by 98 tortoises across California, Arizona, and Utah ( Supplement 2A and 2B (naar-41-02-09_s02.docx)). Supplementary qualitative observations on foraging tortoises were documented at six sites (seven studies) adding Nevada ( Supplement 2C (naar-41-02-09_s02.docx)), and two fecal studies added an additional site in Arizona ( Supplement 2D (naar-41-02-09_s02.docx)).

We found 167 unique native plant taxa in tortoise diets representing bite counts, observational, and fecal studies ( Supplement 2 (naar-41-02-09_s02.docx)). Seventeen diet taxa accounted for greater than 1% of bites at 1 to 6 sites (Table 2). Over 80% of native taxa in bites were annual or perennial forbs, while shrubs, perennial grasses, and succulents each comprised less than 10% of the plant taxa ( Supplement 2 (naar-41-02-09_s02.docx)). Collectively, native taxa comprised over one-half (59%) of the total bites (including exotic species) in tortoise diets and were moderately distributed among sites. The top five native forbs each comprised between >2% and 7% of total diets ( Supplement 2 (naar-41-02-09_s02.docx)).

In contrast, each of the top three nonnative species comprised >10% of total bites and in combination comprising 42% across geographically diverse sites ( Supplement 2 (naar-41-02-09_s02.docx), Figure 1): the annual forb Erodium cicutarium comprised 14.2% (eight sites), while annual grasses Bromus madritensis L., B. tectorum L., and Schismus P. Beauv spp. collectively comprised 26.9% of bites, and were found in diets at nine, two, and five sites, respectively.

Table 1.

Fifteen studies at nine sites document plant species in the diets of Mojave desert tortoises across the Mojave Desert ecoregion. Methods abbreviated as BC = bite counts, quantified through direct observation, OBS = foraging observations that were not quantified with bites, and FA = fecal analysis of wild tortoises.


Although it was not widely abundant in diets, we include Opuntia basilaris Englemann and J. Bigelow among the priority species ( Supplement 2 (naar-41-02-09_s02.docx)) because it was consumed abundantly during physiologically stressful years when preferred species were unavailable (Turner et al. 1984; Esque et al. 2014). Sphaeralcea ambigua A. Gray was also added to the list because it recovers well after fire, tortoise movements into burned areas were facilitated by S. ambigua cover (Drake et al. 2015), the species is readily consumed by captive and wild desert tortoises when available (TCE pers. obs. and Van Devender et al. 2002, respectively), and the nutrition in Sphaeralcea ambigua is comparable to other diet species (McArthur et al. 1994). Because tortoise diets are mostly herbaceous annual and perennial species and cover plants are usually woody shrubs, our comparison has only one overlapping taxon among the top 1% of diet and cover plants in either list: Krameria erecta.

Table 2.

Native species that comprise ≥1% of total number of bites.


Desert Tortoise Cover Taxa

Forty-nine unique plant taxa were used for cover by tortoises based on 4187 field observations of radio-telemetered tortoises across nine study sites in Nevada, Utah, and California, and observations at one site in southern Nevada (Burge 1978;   Supplement 3 (naar-41-02-09_s03.docx)). Twelve taxa comprised over 93% of tortoise cover sites and the majority were woody species that typically recover poorly following wildfire (Table 3). Larrea tridentata and Ambrosia dumosa alone comprised 64% of use (Table 3). Six taxa had between 100 and 300 observations of use including Ephedra spp., Yucca schidigera, Lycium andersonii, Ambrosia salsola Strother and B.G. Baldwin, Sphaeralcea ambigua, and Y. brevifolia Englemann (Table 3). The genera Atriplex, Encelia, Ephedra, Eriogonum, and Yucca were each represented by three species used by tortoises.

Table 3.

Plant taxa comprising ≥1% of use for cover by Mojave desert tortoises across eight sites in the Mojave Desert (Figure 1; see  Supplement 3 (naar-41-02-09_s03.docx) for complete list). Species occurrences include these sites and add Burge (1978) for a total of nine sites for frequency (Figure 1). Plant Functional Type (PFT) is adapted from Shryock et al. (2014) and denotes growth forms that have high (“+”) or low (“–”) recovery following fire: perennial forbs (PF+) and woody species (W+ or W–) based on life-history traits (i.e., lifespan, seed mass, dispersal, height class, and leaf longevity). Sites refers to the number of the total nine sites where cover was documented based on numeric codes in Figure 1.


Pollinator Host Taxa

We compiled 57 plant taxonomic groups including 130 species to identify their use as pollinator hosts (Table 4). Pollinators using these plants include 78.4% of the Apocrita (hereafter bees), 9.7% of Hymenoptera (wasps and others except Apocrita), 5.2% of Diptera (flies), 3.0% of Coleoptera (beetles), 63.4% of Papilionidae (butterflies), 81.3% of Lepidoptera (moths) except Papilionidae, and 5.2% of Apodiformes (hummingbirds) (Table 4). We documented 13 plant taxa used exclusively by moths, 3 only by butterflies, and 1 by bees. Fly, wasp and others, or beetle pollinator groups were not documented using plant taxa exclusively. At least 11 of the 13 plants identified as having exclusive relationships with moths are wind-pollinated, and thus likely function as larval host plants. These included six grasses, three Ephedra spp., and two Ambrosia spp. The exclusive butterfly relationships also are associated with wind-pollinated plants, including Plantago spp. and the perennial grass Pappostipa sp. (Table 4).

Fifty-three (93%) of the plant taxa on our list are larval host plants for at least one butterfly or moth taxon. Of these, 37.1% of larval host taxa are herbaceous annuals, 21.4% are herbaceous perennials, and 30.0% are woody shrubs or trees (Table 4). There was only one annual grass, four succulents, and three perennial grasses used by larval pollinators. Nine of 11 moth-pollinated species we listed have white or very light colored corollas (Table 4), and 10 of 11 are tubular (e.g., Mirabilis spp.) or have deep hypanthia (e.g., Chylismia spp., Oenothera spp.); this shape and color combination is the classic syndrome of moth-pollinated plants, particularly hawkmoths (Sphingidae; Raguso and Willis 2003). Eighty-one percent (N = 76) of the listed taxa (excluding genera, as well as wind-, self-, and hummingbird-pollinated taxa) have corollas that are blue (N = 3), purple (N = 23), white (N = 38), or yellow (N = 20), which are also all attractive to bees (Leleji 1973; Real 1981). Only four plant species (Echinocereus mojavensis [= E. triglochidiatus] Engelmann and J.M. Bigelow, Euphorbia micromera Boissier, E. parryi Engelmann, Penstemon pseudospectabilis M.E. Jones, and Sphaeralcea ambigua) have red to orange corollas, and of these, only the cactus and penstemon have tubular-shaped flowers expected to attract hummingbirds (Table 4,  Supplement 1 (naar-41-02-09_s01.docx)).

There were six plant taxa that are in tortoise diets, but for which no pollinator relationships were found in the literature: Cryptantha micrantha (Torr.) I.M. Johnst., C. nevadensis A. Nelson & P.B. Kenn., Eriogonum maculatum A. Heller, Malacothrix coulteri Harv. & A. Gray, Plantago patagonica Jacq, and Stipa hymenoides Roem. & Shult. While Plantago spp. are self-pollinating, P. ovata Forsk is a larval host to at least one moth, and many Eriogonum spp. are larval hosts to several moths or butterflies. Many Cryptantha spp. also have close relationships with native bee pollinators ( Supplement 2A (naar-41-02-09_s02.docx)), and close relatives of Malacothrix coulteri have relationships with several bee species and host moth larvae.

Taxonomic Accounts

The taxonomic accounts provide detailed information in support of key species for 57 taxonomic groups (  Supplement 1 (naar-41-02-09_s01.docx)). Many of the accounts combine several species within a genus because of their morphological and ecological similarities, and rarely some genera were combined into functional groups for similar reasons. We found a wide variety of online resources designed to assist gardeners, restorationists, hobbyists, and professionals on each of the topics in the species accounts, and the availability of these sites is growing rapidly. Their individual development status and usefulness for technical work varies widely.

Table 4.

Mojave Desert priority plant taxa (134 spp. among 57 taxonomic groups) for use in restoration and pollinated by animals, by selfing, or by wind. Higher order pollinator names include Hymenoptera (bees and wasps); Diptera (flies); Coleoptera (beetles); Lepidoptera (butterflies) and (moths). Detailed explanations for codes found in this table are at the bottom of the table. Habit: annual forb (AF), annual grass (AG), biennial (Bien), leaf succulent (LS), perennial forb (PF), perennial grass (PG), shrub (Shr), subshrub (sShr), geophyte (Geo), tree (Tr), stem succulent (SS).









Priority plant species were identified using multiple selection criteria (Giannini et al. 2016), an approach that in other ecoregions recognizes the benefits of native plants on wildlife and pollinators, local biodiversity, ecosystem services and function, and socioeconomics (Tallis et al. 2008). Because the Mojave desert tortoise is an umbrella species with a broad distribution, the Mojave Desert priority species list includes diverse taxa providing food and cover for this generalist herbivore while recognizing the value of host plants to support an array of native pollinators. By identifying all known plant taxa consumed by tortoises at sites across the Mojave Desert, we present a broad spectrum to guide seed collection and plant propagation to restore diet species in disturbed tortoise habitats.

Diet and cover use vary among individual tortoises, populations, and years (Esque et al. 2014), and the available studies on tortoise diet are limited in duration, location, or seasonality; thus, every project should be tailored to local diversity and conditions whenever possible, and more work on local diets would be useful for restoration. The studies that quantify tortoise plant use in the Mojave Desert are strongly biased toward winter/spring flora. Similarly, most plant taxa we identified as pollinator resources are spring and early-summer flowering, and restoration programs in the Mojave Desert could benefit from more information on the summer flora and their pollinators.

Despite their prevalence in some tortoise diets, nonnative annual species are excluded from our priority list because they reduce native plant diversity (Brown and Minnich 1986; Brooks 1999), cause bodily harm to young tortoises (Medica and Eckert 2007; Drake et al. 2016), and reduce growth and survival of Mojave desert tortoises, possibly influencing negative population growth trends (Drake et al. 2015).

Native forbs and grasses are required in seed mixes to replenish seedbanks that are depleted, such as following wildfire (Esque et al. 2010) and surface compaction or excavation activities (DeFalco et al. 2009). With sufficient winter precipitation, native annuals can be competitively released when perennial species are lost to disturbance. However, recolonization success is hindered by the invasion and rapid dominance by competitive Mediterranean annual grasses like Bromus spp. and Schismus spp. (Brooks 1999). Wildfires can severely deplete the seed bank, particularly species beneath shrubs (Cave and Patten 1984; Esque et al. 2010), yet annual species with affinities for shrub interspaces may persist following disturbance. Annual species that are known colonizers (Acmispon/Lotus spp., Stephanomeria exigua) may establish well in disturbed areas if seeded in concert with herbicide suppression of invasive annual grasses to reduce competition (DeFalco and Esque 2014), although details of herbicide effects on native communities must be worked out, and potential effects of herbicide treatments on tortoise health have not been documented.

As an alternative to seed dispersal, species that resprout from above- or below-ground structures have the potential to persist after disturbance (Clarke et al. 2013). Resprouting success depends on how and if the regenerating buds are protected from damage, and the location and amount of resources available for resprouting (Clarke et al. 2013). Many of the woody “resprouters” we identified are facultative (e.g., Lycium spp.) or obligate inhabitants of riparian areas that experience frequent natural disturbance (e.g., Chilopsis linearis, Prunus sp.). Long-lived wash species that evolved to resprout after damaging flood events may have an advantage when recovering after other disturbances as well (Bock and Bock 2014). Among perennials we identified as “resprouters,” one is a geophyte (Androstephium breviflorum), two are perennial grasses with rhizomatous root systems (Muhlenbergia porteri and Hilaria rigida), and one reproduces from nodal offshoots (Euphorbia albomarginata Torr. & A. Gray). Others, like Mirabilis sp., Delphinium sp., Allium sp., and some Asclepias sp., also have root systems or tubers capable of resprouting. Although resprouting can be a successful means to persist after damage to aboveground tissues, some desert shrubs and trees resprout poorly after disturbance (Abella 2009) and may only survive low-intensity injury when the root crown remains intact and post-disturbance conditions favor growth (Gibson et al. 2004; DeFalco et al. 2010; Steers and Allen 2011; Esque et al. 2013).

While propagating “colonizers” from seed, and their inclusion in seed mixes may be an economical means to restore large disturbances, poor colonizers such as many native annuals may require nursery propagation from seed or cuttings before outplanting. Fortunately, the cacti included in this group all grow readily from cuttings and may be planted directly into restoration sites without growing to size in a nursery setting. Positioning outplanted seedlings and cuttings in groups to form habitat “islands” may enhance shrubland establishment and eventually restore ecological processes such as facilitation by nurse plants and fertile island development for annual species (Badano et al. 2016) and is being formally tested in the Mojave Desert (LAD, pers. comm.).

We used pollination syndrome as an indicator of pollinator relationships because empirical studies for many of the taxa are lacking. Many of the priority Mojave Desert taxa support classic pollination syndromes found in the literature (Fenster et al. 2004), including a large proportion of the species with purple/blue, yellow, and white corollas, which are bee pollinated (Leleji 1973; Real 1981). Other taxa have shape and color combinations consistent with moth pollination (Raguso and Willis 2003). In contrast, few vertebrate-pollinated floral syndromes were among the taxa we identified: Echinocereus triglochidiatus and Penstemon pseudospectabilis have red-purple tubular flowers with little or no scent and are considered attractive to hummingbirds. While classical pollination syndromes are useful for indicating visitation by certain groups, they are not always indicative of the whole pollinator guild. We acknowledge that accounts of pollinator visitations must be validated whenever possible because visitation does not always equate to pollination and is a poor proxy for successful plant outcrossing (King et al. 2013).

Insect diversity is high in the southwestern US (Allred 1969; Moldenke and Neff 1974; Michener 1979), and insect pollinators far outnumber vertebrate pollinators (Simpson and Neff 1987). Yet published studies of desert pollinators are biased toward bats, hummingbirds, and specialist relationships. The ubiquity of hymenopteran pollination, particularly the bees (Apoidea), illustrates their importance to desert plants. However, the most comprehensive invertebrate studies are limited to a few plant taxa, including Larrea tridentata (Hurd and Linsley 1975a, 1975b; Minckley et al. 2000), Oenothera (Raven 1979; Thorp and LaBerge 2005), Krameria (Simpson and Neff 1987; Simpson 1989), and Opuntia (Grant and Grant 1979), or encompass small geographic areas with high levels of bee endemism (e.g., Griswold et al. 2006). Pollinators may be afforded better protection from predators by woody vegetation, which provides sustainable, reliable nutrition and protection for hosted larvae compared with the short lifespans and the spatiotemporal unpredictability of annuals. Our survey indicates that almost all woody plant taxa are host plants for Lepidopteran larvae, compared to <50% of herbaceous plants. Pollinator studies are needed for the Mojave Desert ecoregion, particularly ecological studies that describe the many oligolectic native bees (i.e., bees that specialize in collecting pollen from a limited number of genera or species of flowering plants), the influences that disturbance has on pollinator diversity and function, how pollinator abundance and diversity respond to restoration of degraded habitats, and how climate change may influence pollinator/host relationships.

While we focused on many generalist species, many native plant species co-evolved with native arthropod consumers such that their life history stages are wholly dependent on plants and can only be replaced by few if any other species (Tallamy 2004; Burghardt and Tallamy 2013). Therefore, the loss of plant species potentially reduces overall species diversity further along the food web. For example, many terrestrial birds are dependent on insect protein, especially for feeding their young during growth periods (Burghardt et al. 2009). While such work has not been quantified for Mojave Desert communities, we predict that failure to restore diverse shrub communities over large areas may have negative consequences for ecoregional biodiversity.

Our priority species list can be used to guide resource managers and practitioners in collecting and storing seeds for landscape-scale restoration projects (e.g., Seeds of Success; Haidet and Olwell 2015), establishing production fields for seed increase (e.g., USDA, Tucson Plant Materials Center, commercial growers), cultivating nursery stock for outplanting (National Park Service, Joshua Tree National Park; Lake Mead National Recreation Area, Mojave Desert Land Trust, Nevada Division of Forestry), and prioritizing research topics. By identifying Mojave Desert priority species for restoration a priori, opportunities may grow for entrepreneurs to develop a diversity of species for socioeconomic benefits, thereby increasing business opportunities while enhancing restoration and conservation. The priority taxa and taxonomic accounts presented here for the Mojave Desert, in combination with seed transfer zones derived from climate and genetic information (Shryock et al. 2017, 2018, 2020), will assist practitioners in creating customized species menus for use on restoration projects across this desert ecoregion. While this priority species listing presents a start, it would be useful to create an online, living repository of Mojave Desert native plant cultivation methods and data that are regularly curated to increase the incentive for community participation.


We are grateful for funding provided by C. Lund (BLM, California State Office), F. Edwards (BLM, Nevada State Office), J. Fox (NPS, Grand Canyon-Parashant National Monument), and K. Harcksen (BLM, Arizona Strip District Office, retired). California BLM Plant Conservation and Restoration Program and BLM Mojave Desert Native Plant Program provided funding support. At the time of publication, data are not available from the Bureau of Land Management, U.S. Fish and Wildlife Service, or Fort Irwin National Training Center. Please contact the lead author for information. We thank J. Perkins (BLM, California) for her insightful comments on this manuscript. Any use of trade, product, or firm names is for descriptive purposes only and does not imply endorsement by the U.S. government.



Abella, S.R. 2009. Post-fire plant recovery in the Mojave and Sonoran Deserts of western North America. Journal of Arid Environments 73:699–707. Google Scholar


Allison, G. 2004. The influence of species diversity and stress intensity on community resistance and resilience. Ecological Monographs 74:117–134. Google Scholar


Allred, D.M. 1969. Bees of the Nevada test site. Great Basin Naturalist 29:20–24. Google Scholar


Avery, H.W. 1998. Nutritional ecology of the desert tortoise (Gopherus agassizii) in relation to cattle grazing in the Mojave Desert. Dissertation, University of California, Los Angeles. Google Scholar


Badano, E.I., O.R. Samour-Nieva, J. Flores, J.L. Flores-Flores, J.A. Flores-Cano, and J.P. Rodas-Ortíz. 2016. Facilitation by nurse plants contributes to vegetation recovery in human-disturbed desert ecosystems. Journal of Plant Ecology 9:485–497. Google Scholar


Bainbridge, D.A. 2007. A Guide for Desert and Dryland Restoration: New Hope for Arid Lands. Society for Ecological Restoration International. Island Press, Washington, DC. Google Scholar


Baldwin, B.G., S. Boyd, B.J. Ertter, R.W. Patterson, T.J. Rosatti, D.H. Wilken, and M. Wetherwax. 2002. The Jepson Desert Manual: Vascular Plants of Southeastern California. University of California Press, Berkeley and Los Angeles. Google Scholar


Barrow, J.T. 1979. Aspects of ecology of the desert tortoise, Gopherus agassizii, inJoshua Tree National Monument, Pinto Basin. Proceedings of the Desert Tortoise Council 1979:105–131. Google Scholar


Baskin, C.C., and J.M. Baskin. 2014. Seeds: Ecology, Biogeography, and Evolution of Dormancy and Germination. 2nd ed. Academic Press, San Diego, CA. Google Scholar


Beatley, J.C. 1974. Phenological events and their environmental triggers in Mojave Desert ecosystems. Ecology 55:856–863. Google Scholar


Bickett, J.E. 1980. Aspects of the natural history of the desert tortoise, Gopherus agassizii in southeastern California. Thesis, California State University, Sacramento. Google Scholar


BLM and US DOE [Bureau of Land Management and U.S. Department of Energy]. 2012. Final Programmatic Environmental Impact Statement (PEIS) for Solar Energy Development in Six Southwestern States. Bureau of Land Management, Washington, DC. Google Scholar


Bock, C.E., and J.H. Bock. 2014. Effects of wildfire on riparian trees in southeastern Arizona. Southwestern Naturalist 59:570–576. Google Scholar


Bradford, J.B., J. Betancourt, B.J. Butterfield, S.M. Munson, and T.E. Wood. 2018. Anticipatory natural resource science and management for a changing future. Frontiers in Ecology and the Environment 16:295–303. Google Scholar


Brooks, M.L. 1999. Habitat invasibility and dominance by alien annual plants in the western Mojave Desert. Biological Invasions 1:325–337. Google Scholar


Brown, D.E., and R.A. Minnich. 1986. Fire and changes in creosotebush scrub of the Western Sonoran Desert, California. American Midland Naturalist 116:411–422. Google Scholar


Burge, B.L. 1978. Physical characteristics and patterns of utilization of cover sites used by Gopherus agassizii in southern Nevada. Proceedings of the Desert Tortoise Council Symposium 1978:80–111. Google Scholar


Burge, B.L., and W.G. Bradley. 1976. Population density, structure and feeding habits of the desert tortoise, Gopherus agassizii, in a low desert study area in southern Nevada. Pp. 51–74 in N.J. Engberg, S. Allan, and R.L. Young, eds., The Desert Tortoise Council Proceedings of 1976 Symposium. Desert Tortoise Council, Las Vegas, NV. Google Scholar


Burghardt, K.T., and D.W. Tallamy. 2013. Plant origin asymmetrically impacts feeding guilds and life stages driving community structure of herbivorous arthropods. Biodiversity and Distributions 19:1553–1565. Google Scholar


Burghardt, K.T., D.W. Tallamy, and W.G. Shriver. 2009. Impact of native plants on bird and butterfly biodiversity in suburban landscapes. Conservation Biology 23:219–224. Google Scholar


Carril, O.M., T. Griswold, J. Haefner, and J.S. Wilson. 2018. Wild bees of Grand Staircase-Escalante National Monument: Richness, abundance, and spatio-temporal beta-diversity. PeerJ 6:e5867. < Scholar


Carter, S.K., K.E. Nussear, T.C. Esque, I.I.F. Leinwand, E. Masters, R.D. Inman, N.B. Carr, and L.J. Allison. 2020. Quantifying development to inform management of Mojave and Sonoran desert tortoise habitat in the American Southwest. Endangered Species Research. < Scholar


Cave, G.H., and D.T. Patten. 1984. Short-term vegetation responses to fire in the upper Sonoran Desert. Journal of Range Management 37:491–496. Google Scholar


Clarke, P.J., M.J. Lawes, J.J. Midgley, B.B. Lamont, F. Ojeda, G.E. Burrows, N.J. Enright, and K.J.E. Knox. 2013. Resprouting as a key functional trait: How buds, protection and resources drive persistence after fire. New Phytologist 197:19–35. Google Scholar


Cody, M.L. 2000. Slow motion population dynamics in Mojave Desert perennial plants. Journal of Vegetation Science 11:351–358. Google Scholar


Dai, A. 2013. Increasing drought under global warming in observations and models. Nature Climate Change 3:52–58. Google Scholar


DeFalco, L.A., and T.C. Esque. 2014. Soil seed banks: Preserving native biodiversity and repairing damaged desert shrublands. Fremontia 42:20–23. Google Scholar


DeFalco, L.A., T.C. Esque, J.M. Kane, and M.B. Nicklas. 2009. Seed banks in a degraded desert shrubland: Influence of soil surface condition and harvester ant activity on seed abundance. Journal of Arid Environments 73:885–893. Google Scholar


DeFalco, L.A., T.C. Esque, S.J. Scoles, and J. Rodgers. 2010. Desert wildfire and severe drought diminish survivorship of the long-lived Joshua tree (Yucca brevifolia; Agavaceae). American Journal of Botany 97:243–350. Google Scholar


Drake, K.K., L. Bowen, K.E. Nussear, T.C. Esque, A.J. Berger, N.A. Custer, S.C. Waters, A.K. Miles, and R.L. Lewison. 2016. Negative effects of invasive plants on conservation of sensitive desert wildlife. Ecosphere 7(10):e01531. < Scholar


Drake, K.K., T.C. Esque, K.E. Nussear, L.A. DeFalco, S.J. Scoles-Sciulla, A.T. Modlin, and P.A. Medica. 2015. Desert tortoise use of burned habitat in the eastern Mojave Desert. Journal of Wildlife Management 79:618–629. Google Scholar


Esque, T.C. 1994. Diet and diet selection of the desert tortoise (Gopherus agassizii) in the northeast Mojave Desert. Master's thesis. Colorado State University, Fort Collins. Google Scholar


Esque, T.C., K.K. Drake, and K.E. Nussear. 2014. Water and food acquisition and their consequences on life history and metabolism of North American tortoises. Pp. 85–95 in D. Rostal, E.D. McCoy, and H. Mushinsky, eds., Biology and Conservation of North American Tortoises. Johns Hopkins University Press, Baltimore, MD. Google Scholar


Esque, T.C., R.H. Webb, C.S.A. Wallace, C. van Riper III , C. McGreedy, and L. Smythe. 2013. Desert fires fueled by native annual forbs: The 2005 King Valley fire and its impacts on plant and bird communities in the Lower Sonoran Desert of Arizona. Southwestern Naturalist 58:223–233. Google Scholar


Esque, T.C., J.A. Young, and C.R. Tracy. 2010. Short-term effects of experimental fires on a Mojave Desert seed bank. Journal of Arid Environments 74:1302–1308. Google Scholar


Fenster, C.B., W.S. Armbruster, P. Wilson, M.R. Dudash, and J.D. Thomson. 2004. Pollination syndromes and floral specialization. Annual Review of Ecology, Evolution, and Systematics 35:375–403. Google Scholar


Giannini, T.C., A.M. Giulietti, R.M. Harley, P.L. Viana, R. Jaffe, R. Alves, C.E. Pinto, N.F.O. Mota, C.F. Caldeira Jr., V.L. Imperatriz-Fonseca, et al. 2016. Selecting plant species for practical restoration of degraded lands using a multiple-trait approach. Austral Ecology 42:510–521. Google Scholar


Gibson, A.C., M.R. Sharifi, and P.W. Rundel. 2004. Resprout characteristcs of creosotebush (Larrea tridentata) when subjected to repeated vehicle damage. Journal of Arid Environments 57:411–429. Google Scholar


Gonzalez V.H., and T.L. Griswold. 2013. Wool carder bees of the genus Anthidium in the Western Hemisphere (Hymenoptera: Megachilidae): Diversity, host-plant associations, phylogeny, and biogeography. Zoological Journal of the Linnean Society 168:221–425. Google Scholar


Graff, P., and S. McIntyre. 2014. Using ecological attributes as criteria for the selection of plant species under three restoration scenarios. Austral Ecology 39:907–917. Google Scholar


Grant, V., and K.A. Grant. 1979. Pollination of Opunta basilaris and O. littoralis. Plant Systematics and Evolution 132:321–325. Google Scholar


Griswold, T., S. Higbee, and O. Messinger. 2006. Pollination ecology final report for biennium 2003, Clark County, Nevada (2004-2005). USDA-ARS Bee Biology and Systematics Laboratory, Utah State University, Logan, UT. Google Scholar


Grover, M.C., and L.A. DeFalco. 1995. Desert tortoise (Gopherus agassizii): Status-of-knowledge outline with references. INT-GTR-316. USDA Forest Service, Intermountain Research Station, Ogden, UT. Google Scholar


Haidet, M., and P. Olwell. 2015. Seeds of success: A national seed banking program working to achieve long-term conservation goals. Natural Areas Journal 35:165–173. Google Scholar


Hansen, R.M., M.K. Johnson, and T.R. Van Devender. 1976. Foods of the desert tortoise, Gopherus agassizii, in Arizona and Utah. Herpetologica 32:247–251. Google Scholar


Hereford, R., R.H. Webb, and C.I. Longpré. 2006. Precipitation history and ecosystem responses to multidecadal precipitation variability in the Mojave Desert region, 1893 to 2001. Journal of Arid Environments 67:13–34. Google Scholar


Hohman, J.P., and R.D. Ohmart. 1980. Ecology of the desert tortoise on the Beaver Dam slopes, Arizona. Report or Contract No. YA-510∼ph7-54. USDI - Bureau of Land Management, St. George, UT. Google Scholar


Hurd, P.D., and E.G. Linsley. 1975a. The principal Larrea bees of the southwestern United States (Hymenoptera: Apoidea). Smithsonian Contributions to Zoology No. 193. Smithsonian Institution Press, Washington, DC. < Scholar


Hurd, P.D., and E.G. Linsley. 1975b. Some insects other than bees associated with Larrea tridentata in the southwestern United States. Proceedings of the Entomological Society of Washington 77:100–120. Google Scholar


IPCC [Intergovernmental Panel on Climate Change]. 2013. Climate Change 2013: The physical science basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. T.F. Stocker, D. Qin, G.-K. Plattner, M. Tignor, S.K. Allen, J. Boschung, A. Nauels, Y. Xia, V. Bex, and P.M. Midgley, eds. Cambridge University Press, Cambridge, UK and New York, NY. Google Scholar


ITIS [Integrated Taxonomic Information System]. 2021. Accessed 5 February 2021 from < >. Google Scholar


Jennings, W.B. 1993. Food preferences and feeding behavior of desert tortoises (Xerobates agassizii) at the Desert Tortoise Natural Area, Kern County, California. Report to USDI-Bureau of Land Management. Google Scholar


Jennings, W.B., and K.H. Berry. 2015. Desert tortoises (Gopherus agassizii) are selective herbivores that track the flowering phenology of their preferred food plants. PLoS ONE 10(1):e0116716. < Scholar


Jepson Flora Project. 2018. B.G. Baldwin, D.J. Keil, S. Markos, B.D. Mishler, R. Patterson, T.J. Rosatti, and D.H. Wilken, eds. Accessed 30 March 2020 from <>. Google Scholar


Johnson, R., L. Stritch, P. Olwell, S. Lambert, M.E. Horning, and R. Cronn. 2010. What are the best seed sources for ecosystem restoration on BLM and USFS lands? Native Plants 11:117–131. Google Scholar


Keeler-Wolf, T. 2007. Mojave desert scrub vegetation. Pp. 609–656 in M.G. Barbour, T. Keeler-Wolf, and A.A. Schoenherr, eds., Terrestrial Vegetation of California. 3rd ed. University of California Press, Berkeley. Google Scholar


King, C., G. Ballantyne, and P.G. Willmer. 2013. Why flower visitation is a poor proxy for pollination: Measuring single-visit pollen deposition, with implications for pollination networks and conservation. Methods in Ecology and Evolution 4:811–818. Google Scholar


Leleji, O.I. 1973. Apparent preference by bees for different flower colours in cowpeas (Vigna sinensis (L.) Savi ex Hassk.). Euphytica 22:150–153. Google Scholar


Lesica, P., and F. Allendorf. 1999. Ecological genetics and the restoration of plant communities: Mix or match? Restoration Ecology 7:42–50. Google Scholar


Leu, M., S.E. Hanser, and S.T. Knick. 2008. The human footprint in the West: A large-scale analysis of anthropogenic impacts. Ecological Applications 18:1119–1139. Google Scholar


Luckenbach, R. 1982. Ecology and management of the desert tortoise (Gopherus agassizii) in California. Pp. 1–37 in R.B. Bury, ed., North American Tortoises: Their Conservation and Ecology. U.S. Fish and Wildlife Service Report 12. Government Printing Office. Google Scholar


MacMahon, J.A., and F.H. Wagner. 1985. The Mojave, Sonoran and Chihuahuan deserts of North America. Pp. 139–174 in M. Evenari, I. Noy-Meir, and D.W. Goodall, eds., Ecosystems of the World 12A: Hot Deserts and Arid Shrublands. Elsevier Scientific Publishing Company, New York. Google Scholar


Maron, J.L., K.C. Baer, A.L. Angert, and J. Lau. 2014. Disentangling the drivers of context-dependent plant–animal interactions. Journal of Ecology 102:1485–1496. Google Scholar


McArthur, E.D., S.C. Sanderson, and B.L. Webb. 1994. Nutritive quality and mineral content of potential desert tortoise food plants. Research Paper INT-473. USDA-Forest Service, Intermountain Research Station, Ogden, UT. Google Scholar


Medica, P.A., and S.E. Eckert. 2007. Gopherus agassizii, desert tortoise. Food/Mechanical Injury. Natural History Notes. Herpetological Review 38:446–448. Google Scholar


Medica, P.A., C.L. Lyons, and F.B. Turner. 1981. A comparison of populations of the desert tortoise (Gopherus agassizii) in grazed and ungrazed areas in Ivanpah Valley, California. California Desert District Final Report, Bureau of Land Management. Google Scholar


Michener, C.D. 1979. Biogeography of the bees. Annals of the Missouri Botanical Garden 66:277–347. Google Scholar


Miller, D.M., D.L. Hughson, and R.H. Webb. 2009. Part IV Recovery, restoration, and ecosystem monitoring. Pp. 339–342 in R.H. Webb, L.F. Fenstermaker, J.S. Heaton, D.L. Hughson, E.V. McDonald, and D.M. Miller, eds., The Mojave Desert: Ecosystem Processes and Sustainability. University of Nevada Press, Reno. Google Scholar


Minckley, R.L., J.H. Cane, and L. Kervin. 2000. Origins and ecological consequences of pollen specialization among desert bees. Proceedings of the Royal Society of London B 267:265–271. Google Scholar


Minnich, R.A. 2007. Climate, paleoclimate, and paleovegetation. Pp. 43–70 in M.G. Barbour, T. Keeler-Wolf, and A.A. Schoenherr, eds., Terrestrial Vegetation of California. 3rd ed. University of California Press, Berkeley. Google Scholar


Moldenke, A.R., and J.L. Neff. 1974. Studies on pollination ecology and species diversity of natural California plant communities. International Biological Programme Technical Report 74-13, Volume II. Google Scholar


Nussear, K.E., and T. Tuberville. 2014. Habitats of North American tortoises. Pp. 77–84 in D. Rostal, H. Mushinsky, and E. McCoy, eds., Biology and Conservation of North American Tortoises. Johns Hopkins University Press, Baltimore, MD. Google Scholar


Ollerton, J., R. Alarcon, N.M. Waser, M.V. Price, S. Watts, L. Cranmer, A. Hingston, C.I. Peter, and J. Rotenberry. 2009. A global test of the pollination syndrome hypothesis. Annals of Botany 103:1471–1480. Google Scholar


Olwell, P., and L. Riibe. 2016. National Seed Strategy: Restoring pollinator habitat begins with the right seed in the right place at the right time. Natural Areas Journal 36:363–365. Google Scholar


Raguso, R.A., and M.A. Willis. 2003. Hawkmoth pollination in Arizona's Sonoran Desert: Behavioral responses to floral traits. Pp. 43–65 in C.L. Boggs, W.B. Watt, and P.R. Ehrlich, eds., Butterflies: Evolution and Ecology Taking Flight: Butterflies as Model Systems. University of Chicago Press, Chicago, IL. Google Scholar


Raven, P.H. 1979. A survey of reproductive biology in Onagraceae. New Zealand Journal of Botany 17:575–593. Google Scholar


Real, L.A. 1981. Uncertainty and pollinator–plant interactions: The foraging behavior of bees and wasps on artificial flowers. Ecology 62:20–26. Google Scholar


Reubens, B., C. Moeremans, J. Poesen, J. Nyssen, S. Tewoldeberhan, S. Franzel, J. Deckers, C. Orwa, and B. Muys. 2011. Tree species selection for land rehabilitation in Ethiopia: From fragmented knowledge to an integrated multi-criteria decision approach. Agroforestry Systems 82:303–330. Google Scholar


Rice, K.J., and N.C. Emery. 2003. Managing microevolution: Restoration in the face of global change. Frontiers in Ecology and the Environment 1:469–478. Google Scholar


Scoles-Sciulla, S.J., and L.A. DeFalco. 2009. Seed reserves diluted during surface soil reclamation in eastern Mojave Desert. Arid Land Research and Management 23:1–13. Google Scholar


SEINet Portal Network. 2020. Accessed 24 April 2020 from < http//>. Google Scholar


Shryock, D.F., L.A. DeFalco, and T.C. Esque. 2014. Life-history traits predict perennial species response to fire in a desert ecosystem. Ecology and Evolution 4:3046–3059. Google Scholar


Shryock, D.F., L.A. DeFalco, and T.C. Esque. 2018. Spatial decision-support tools to guide restoration and seed-sourcing in the Desert Southwest. Ecosphere 9(10):e02453. < Scholar


Shryock, D.F., C.A. Havrilla, L.A. DeFalco, T.C. Esque, N.A. Custer, and T.E. Wood. 2017. Landscape genetic approaches to guide native plant restoration in the Mojave Desert. Ecological Applications 27:429–445. Google Scholar


Shryock, D.F., L.K. Washburn, L.A. DeFalco, and T.C. Esque. 2020. Harnessing landscape genomics to identify future climate resilient genotypes in a desert annual. Molecular Ecology < >. Google Scholar


Simpson, B.B., and J.L. Neff. 1987. Pollination ecology in the Southwest. Aliso: A Journal of Systematic and Evolutionary Botany 11:417–440. Google Scholar


Simpson, B.B. 1989. Krameriaceae. Flora Neotropica 49:1–109. Google Scholar


Spang, E.F., G.W. Lamb, F. Rowley, W.H. Radtkey, R.R. Olendorff, E.A. Dahlem, and S. Slone. 1988. Desert tortoise management on the public lands: A rangewide plan. Unpublished report to USDI Bureau of Land Management, Division of Wildlife and Fisheries. Google Scholar


Steers, R.J., and E.B. Allen. 2010. Post-fire control of invasive plants promotes native recovery in a burned desert shrubland. Restoration Ecology 18:334–343. Google Scholar


Tallamy, D.W. 2004. Do alien plants reduce insect biomass? Conservation Biology 18:1689–1692. Google Scholar


Tallis, H., P. Kareiva, M. Marvier, and A. Chang. 2008. An ecosystem services framework to support both practical conservation and economic development. Proceedings of the National Academy of Sciences 105:9457–9464. Google Scholar


Thorp, R.W., and W.E. LaBerge. 2005. A revision of the bees of the genus Andrena of the Western Hemisphere. Part XIV—Subgenus Onagrandrena. Illinois Natural History Survey Bulletins 37:1–64. Google Scholar


Tilman, D., P.B. Reich, and J.M. Knops. 2006. Biodiversity and ecosystem stability in a decade-long grassland experiment. Nature 441:629. Google Scholar


Tracy, C.R., and P.F. Brussard. 1994. Preserving biodiversity: Species in landscapes. Letters to the Editor. Ecological Applications 4:205–207. Google Scholar


Turner, F.B., P.A. Medica, and C.L. Lyons. 1984. Reproduction and survival of the desert tortoise (Scaptochelys agassizii) in Ivanpah Valley, California. Copeia 1984:811–820. Google Scholar


Turner, R.M. 1994. Warm-temperate desertlands, 153.1 Mohave desertscrub. Pp. 157–168 in D.E. Brown, ed., Biotic Communities of the Southwestern United States and Northwestern Mexico. University of Utah Press, Salt Lake City. Google Scholar


USFWS [United States Fish and Wildlife Service]. 2011. Revised recovery plan for the Mojave population of the desert tortoise (Gopherus agassizii). U.S. Fish and Wildlife Service, Pacific Southwest Region, Sacramento, CA. Google Scholar


Vandergast, A.G., R.D. Inman, K.R. Barr, K.E. Nussear, T.C. Esque, S.A. Hathaway, D.A. Wood, P.A. Medica, J.W. Breinholt, C.L. Stephen, et al. 2013. Evolutionary hotspots in the Mojave Desert. Diversity 5:293–319. Google Scholar


Van Devender, T.R., R.C. Averill-Murray, T.C. Esque, P.A. Holm, V.M. Dickinson, C.R. Schwalbe, E.B. Wirt, and S.L. Barrett. 2002. Grasses, mallows, desert vine, and more: Diet of the desert tortoise in Arizona and Sonora. Pp. 159–193 in T.R. Van Devender, ed., The Sonoran Desert Tortoise: Natural History, Biology, and Conservation. University of Arizona Press, Tucson. Google Scholar


Vilsack, T., and G. McCarthy. 2015. National strategy to promote the health of honeybees and other pollinators. Report Issued by the White House Pollinator Health Task Force on May 19, 2015. Google Scholar


Weigand, J., and J. Rodgers. 2009. Active restoration for the Mojave Desert. Pp. 378–409 in R.H. Webb, L.F. Fenstermaker, J.S. Heaton, D.L. Hughson, E.V. McDonald, and D.M. Miller, eds., The Mojave Desert: Ecosystem Processes and Sustainability. University of Nevada Press, Reno. Google Scholar


Wilson, E.O. 1987. The little things that run the world: The importance and conservation of invertebrates. Conservation Biology 1:344–346. Google Scholar
Todd C. Esque, Lesley A. DeFalco, Gayle L. Tyree, K. Kristina Drake, Kenneth E. Nussear, and Joseph S. Wilson "Priority Species Lists to Restore Desert Tortoise and Pollinator Habitats in Mojave Desert Shrublands," Natural Areas Journal 41(2), 145-158, (22 April 2021).
Published: 22 April 2021

aridland restoration
desert tortoise
Mojave Desert
Native species
Get copyright permission
Back to Top