Lesley A. DeFalco

EFFECTS OF DESERT WILDFIRES ON DESERT TORTOISE (GOPHERUS AGASSIZII) AND OTHER SMALL VERTEBRATES

Abstract We report the results of standardized surveys to determine the effects of wildfires on desert tortoises (Gopherus agassizii) and their habitats in the northeastern Mojave Desert and northeastern Sonoran Desert. Portions of 6 burned areas (118 to 1,750 ha) were examined for signs of mortality of vertebrates. Direct effects of fire in desert habitats included animal mortality and loss of vegetation cover. A range of 0 to 7 tortoises was encountered during surveys, and live tortoises were found on all transects. In addition to desert tortoises, only small (<1 kg) mammals and reptiles (11 taxa) were found dead on the study areas. We hypothesize that indirect effects of fire on desert habitats might result in changes in the composition of diets and loss of vegetation cover, resulting in an increase in predation and loss of protection from temperature extremes. These changes in habitat also might cause changes in vertebrate communities in burned areas.

Desert wildfire and severe drought diminish survivorship of the long-lived Joshua Tree (Yucca brevifolia; Agavaceae)

Extreme climate events are transforming plant communities in the desert Southwest of the United States. Abundant precipitation in 1998 associated with El Niño Southern Oscillation (ENSO) stimulated exceptional alien annual plant production in the Mojave Desert that fueled wildfires in 1999. Exacerbated by protracted drought, 80% of the burned Yucca brevifolia, a long-lived arborescent monocot, and 26% of unburned plants died at Joshua Tree National Park by 2004. Many burned plants <1 m tall died immediately, and survival of all but the tallest, oldest plants declined to the same low level by 2004. Postfire sprouting prolonged survival, but only at the wetter, high-elevation sites. During succeeding dry years, herbaceous plants were scarce, and individuals of Thomomys bottae (pocket gopher) gnawed the periderm and hollowed stems of Y. brevifolia causing many of them to topple. Thomomys bottae damage reduced plant survivorship at low-elevation, unburned sites and diminished survival of burned plants in all but the driest site, which already had low survival. Accentuated ENSO episodes and more frequent wildfires are expected for the desert Southwest and will likely shift Y. brevifolia population structure toward tall, old adults with fewer opportunities for plant recruitment, thus imperiling the persistence of this unique plant community.

Seed Reserves Diluted During Surface Soil Reclamation in Eastern Mojave Desert

Surface soil reclamation is used to increase the re-establishment of native vegetation following disturbance through preservation and eventual replacement of the indigenous seed reserves. Employed widely in the mining industry, soil reclamation has had variable success in re-establishing native vegetation in arid and semi-arid regions. We tested whether variable success could be due in part to a decrease of seed reserves during the reclamation process by measuring the change in abundance of germinable seed when surface soil was mechanically collected, stored in a soil pile for 4 months, and reapplied upon completion of a roadway. Overall seed reserve declines amounted to 86% of the original germinable seed in the soil. The greatest decrease in seed reserves occurred during soil collection (79% of original reserves), compared to the storage and reapplication stages. At nearby sites where stored surface soil had been reapplied, no perennial plant cover occurred from 0.5 to 5 years after application and < 1% cover after 7 years compared to 5% cover in nearby undisturbed areas. The reduction in abundance of germinable seed during reclamation was primarily due to dilution of seed reserves when deeper soil fractions without seed were mixed with the surface soil during collection. Unless more precise techniques of surface soil collection are utilized, soil reclamation alone as a means for preserving native seed reserves is a method ill-suited for revegetating disturbed soils with a shallow seed bank, such as those found in the Mojave Desert.

Life‐history traits predict perennial species response to fire in a desert ecosystem

The Mojave Desert of North America has become fire-prone in recent decades due to invasive annual grasses that fuel wildfires following years of high rainfall. Perennial species are poorly adapted to fire in this system, and post-fire shifts in species composition have been substantial but variable across community types. To generalize across a range of conditions, we investigated whether simple life-history traits could predict how species responded to fire. Further, we classified species into plant functional types (PFTs) based on combinations of life-history traits and evaluated whether these groups exhibited a consistent fire-response. Six life-history traits varied significantly between burned and unburned areas in short (up to 4 years) or long-term (up to 52 years) post-fire datasets, including growth form, lifespan, seed size, seed dispersal, height, and leaf longevity. Forbs and grasses consistently increased in abundance after fire, while cacti were reduced and woody species exhibited a variable response. Woody species were classified into three PFTs based on combinations of life-history traits. Species in Group 1 increased in abundance after fire and were characterized by short lifespans, small, wind-dispersed seeds, low height, and deciduous leaves. Species in Group 2 were reduced by fire and distinguished from Group 1 by longer lifespans and evergreen leaves. Group 3 species, which also decreased after fire, were characterized by long lifespans, large non-wind dispersed seeds, and taller heights. Our results show that PFTs based on life-history traits can reliably predict the responses of most species to fire in the Mojave Desert. Dominant, long-lived species of this region possess a combination of traits limiting their ability to recover, presenting a clear example of how a novel disturbance regime may shift selective environmental pressures to favor alternative life-history strategies.

Landscape genomics of Sphaeralcea ambigua in the Mojave Desert: a multivariate, spatially-explicit approach to guide ecological restoration

Abstract Local adaptation influences plant species’ responses to climate change and their performance in ecological restoration. Fine-scale physiological or phenological adaptations that direct demographic processes may drive intraspecific variability when baseline environmental conditions change. Landscape genomics characterize adaptive differentiation by identifying environmental drivers of adaptive genetic variability and mapping the associated landscape patterns. We applied such an approach to Sphaeralcea ambigua, an important restoration plant in the arid southwestern United States, by analyzing variation at 153 amplified fragment length polymorphism loci in the context of environmental gradients separating 47 Mojave Desert populations. We identified 37 potentially adaptive loci through a combination of genome scan approaches. We then used a generalized dissimilarity model (GDM) to relate variability in potentially adaptive loci with spatial gradients in temperature, precipitation, and topography. We identified non-linear thresholds in loci frequencies driven by summer maximum temperature and water stress, along with continuous variation corresponding to temperature seasonality. Two GDM-based approaches for mapping predicted patterns of local adaptation are compared. Additionally, we assess uncertainty in spatial interpolations through a novel spatial bootstrapping approach. Our study presents robust, accessible methods for deriving spatially-explicit models of adaptive genetic variability in non-model species that will inform climate change modelling and ecological restoration.

Short seed longevity, variable germination conditions, and infrequent establishment events provide a narrow window for Yucca brevifolia (Agavaceae) recruitment

UNLABELLED PREMISE OF THE STUDY The future of long-lived stand-forming desert plants such as Yucca brevifolia (Joshua tree) has come into question in light of climate variation and landscape-scale disturbances such as wildfire. Understanding plant establishment dynamics is important for mitigating the impacts of disturbances and promoting revegetation. • METHODS We placed Y. brevifolia seeds in shallow caches and manipulated granivore access, nurse shrub effects, and the season of cache placement to determine conditions for seed germination and seedling establishment. • KEY RESULTS Greatest seedling emergence occurred during spring and summer, when increased soil moisture was accompanied by warm soil temperatures. Late winter-spring emergence for cached seeds was enhanced beneath shrub canopies, but seedling survival declined beneath shrubs as temperatures increased in spring. Germinability of seed remaining in the soil was reduced from 50-68% after 12 mo residence time in soil and declined to <3% after 40 mo. Following dispersal from parent plants, seeds are either removed by granivores or lose germinability, imposing substantial losses of potential germinants. • CONCLUSIONS Specific germination and establishment requirements impose stringent limits on recruitment rates for Y. brevifolia. Coupled with infrequent seed availability, the return rates to prefire densities and demographic structure may require decades to centuries, especially in light of potential changes to regional desert climate in combination with the potential for fire recurrence. Demographic patterns are predicted to vary spatially in response to environmental variability that limits recruitment and may already be apparent among extant populations.

Dust Deposition Effects on Growth and Physiology of the Endangered Astragalus jaegerianus (Fabaceae)

Abstract Human expansion into the Mojave Desert is a significant threat to rare desert plants. While immediate habitat loss is often the greatest concern, rare plants situated near areas where soil surfaces experience frequent disturbance may be indirectly impacted when fine particulate dust accumulates on leaf surfaces. Remaining populations of the federally listed Astragalus jaegerianus (Lane Mountain milkvetch) occur on land open to expanding military activities and on adjacent public land with increasing recreational use. This study was initiated to determine whether dust accumulation could decrease the vigor and fitness of A. jaegerianus through reduced growth. Beginning in early May 2004, plants located on Bureau of Land Management (BLM) land were dusted bimonthly at canopy-level dust concentrations ranging from 0 to 32 g/m2, and physiology and growth were monitored until late June when plants senesced. The maximum experimental dust level simulates dust concentrations of Mojave Desert perennials neighboring military activities at a nearby army training center. Average shoot growth declined with increasing dust accumulation, but seasonal net photosynthesis increased. Further investigation of plants grown in a greenhouse supported similar trends. This pattern of greater net photosynthesis with increasing dust accumulation may be explained by higher leaf temperatures of dusted individuals. Ambient dust deposition measured in traps near field plants (May 2004–July 2004) ranged from 0.04–0.17 g/m2/ d, which was well below the lowest level of dust on experimental plants (3.95 g/m2/d). With this low level of ambient deposition, we expect that A. jaegerianus plants in this population were not greatly affected by the dust they receive at the level of recreational use during the study.

Estimating wildfire risk on a Mojave Desert landscape using remote sensing and field sampling

Predicting wildfires that affect broad landscapes is important for allocating suppression resources and guiding land management. Wildfire prediction in the south-western United States is of specific concern because of the increasing prevalence and severe effects of fire on desert shrublands and the current lack of accurate fire prediction tools. We developed a fire risk model to predict fire occurrence in a north-eastern Mojave Desert landscape. First we developed a spatial model using remote sensing data to predict fuel loads based on field estimates of fuels. We then modelled fire risk (interactions of fuel characteristics and environmental conditions conducive to wildfire) using satellite imagery, our model of fuel loads, and spatial data on ignition potential (lightning strikes and distance to roads), topography (elevation and aspect) and climate (maximum and minimum temperatures). The risk model was developed during a fire year at our study landscape and validated at a nearby landscape; model performance was accurate and similar at both sites. This study demonstrates that remote sensing techniques used in combination with field surveys can accurately predict wildfire risk in the Mojave Desert and may be applicable to other arid and semiarid lands where wildfires are prevalent. Language: en

Landscape genetic approaches to guide native plant restoration in the Mojave Desert

Restoring dryland ecosystems is a global challenge due to synergistic drivers of disturbance coupled with unpredictable environmental conditions. Dryland plant species have evolved complex life-history strategies to cope with fluctuating resources and climatic extremes. Although rarely quantified, local adaptation is likely widespread among these species and potentially influences restoration outcomes. The common practice of reintroducing propagules to restore dryland ecosystems, often across large spatial scales, compels evaluation of adaptive divergence within these species. Such evaluations are critical to understanding the consequences of large-scale manipulation of gene flow and to predicting success of restoration efforts. However, genetic information for species of interest can be difficult and expensive to obtain through traditional common garden experiments. Recent advances in landscape genetics offer marker-based approaches for identifying environmental drivers of adaptive genetic variability in non-model species, but tools are still needed to link these approaches with practical aspects of ecological restoration. Here, we combine spatially explicit landscape genetics models with flexible visualization tools to demonstrate how cost-effective evaluations of adaptive genetic divergence can facilitate implementation of different seed sourcing strategies in ecological restoration. We apply these methods to Amplified Fragment Length Polymorphism (AFLP) markers genotyped in two Mojave Desert shrub species of high restoration importance: the long-lived, wind-pollinated gymnosperm Ephedra nevadensis, and the short-lived, insect-pollinated angiosperm Sphaeralcea ambigua. Mean annual temperature was identified as an important driver of adaptive genetic divergence for both species. Ephedra showed stronger adaptive divergence with respect to precipitation variability, while temperature variability and precipitation averages explained a larger fraction of adaptive divergence in Sphaeralcea. We describe multivariate statistical approaches for interpolating spatial patterns of adaptive divergence while accounting for potential bias due to neutral genetic structure. Through a spatial bootstrapping procedure, we also visualize patterns in the magnitude of model uncertainty. Finally, we introduce an interactive, distance-based mapping approach that explicitly links marker-based models of adaptive divergence with local or admixture seed sourcing strategies, promoting effective native plant restoration.

Contrasting Long-Term Survival of Two Outplanted Mojave Desert Perennials for Post-Fire Revegetation

Post-fire recovery of arid shrublands is typically slow, and planting greenhouse-raised seedlings may be a means of jump-starting this process. Recovery can be further accelerated by understanding the factors controlling post-planting survival. In fall 2007 and 2009, we outplanted seedlings of two contrasting native evergreen shrubs—fast-growing Nevada jointfir and slow-growing blackbrush—across five burned sites in the Mojave Desert. To increase soil moisture and optimize seedling survival, we experimentally applied and evaluated soil amendments and supplemental watering. We also evaluated two herbicides that reduce competitive invasive annual grasses and two types of herbivore protection. Survival of jointfir outplanted in 2007 was 61% after 43 months, and site largely influenced survival, while herbicide containing imazapic applied more than one year after outplanting reduced survival. Reduced survival of jointfir outplanted in 2009 coincided with delayed seasonal precipitation that intensified foliar damage by small mammals. In contrast, blackbrush survival was 4% after 43 months, and was influenced by site, type of herbivore protection, and greenhouse during the 2007 outplanting, and soil amendment during 2009. Counter to expectations, we found that supplemental watering and soil amendments did not influence long-term survival of either blackbrush or jointfir. Shrub species with rapid growth rates and broad environmental tolerances, such as jointfir, make ideal candidates for outplanting, provided that seedlings are protected from herbivores. Re-introduction of species with slow growth rates and narrow environmental tolerances, such as blackbrush, requires careful consideration to optimize pre- and post-planting conditions.