Seed Menus: An integrated decision-support framework for native plant restoration in the Mojave Desert.

The combination of ecosystem stressors, rapid climate change, and increasing landscape-scale development has necessitated active restoration across large tracts of disturbed habitats in the arid southwestern United States. In this context, programmatic directives such as the National Seed Strategy for Rehabilitation and Restoration have increasingly emphasized improved restoration practices that promote resilient, diverse plant communities, and enhance native seed reserves. While decision-support tools have been implemented to support genetic diversity by guiding seed transfer decisions based on patterns in local adaptation, less emphasis has been placed on identifying priority seed mixes composed of native species assemblages. Well-designed seed mixes can provide foundational ecosystem services including resilience to disturbance, resistance to invasive species, plant canopy structure to facilitate natural seedling recruitment, and habitat to support wildlife and pollinator communities. Drawing from a newly developed dataset of species distribution models for priority native plant taxa in the Mojave Desert, we created a novel decision support tool by pairing spatial predictions of species habitat with a database of key species traits including life history, flowering characteristics, pollinator relationships, and propagation methods. This publicly available web application, Mojave Seed Menus, helps restoration practitioners generate customized seed mixes for native plant restoration in the Mojave Desert based on project locations. Our application forms part of an integrated Mojave Desert restoration program designed to help practitioners identify species to include in local seed mixes and nursery stock development while accounting for local adaptation by identifying appropriate seed source locations from key restoration species.

Harnessing landscape genomics to identify future climate resilient genotypes in a desert annual.

Local adaptation features critically in shaping species responses to changing environments, complicating efforts to revegetate degraded areas. Rapid climate change poses an additional challenge that could reduce fitness of even locally sourced seeds in restoration. Predictive restoration strategies that apply seeds with favourable adaptations to future climate may promote long-term resilience. Landscape genomics is increasingly used to assess spatial patterns in local adaption and may represent a cost-efficient approach for identifying future-adapted genotypes. To demonstrate such an approach, we genotyped 760 plants from 64 Mojave Desert populations of the desert annual Plantago ovata. Genome scans on 5,960 SNPs identified 184 potentially adaptive loci related to climate and satellite vegetation metrics. Causal modelling indicated that variation in potentially adaptive loci was not confounded by isolation by distance or isolation by habitat resistance. A generalized dissimilarity model (GDM) attributed spatial turnover in potentially adaptive loci to temperature, precipitation and NDVI amplitude, a measure of vegetation green-up potential. By integrating a species distribution model (SDM), we find evidence that summer maximum temperature may both constrain the range of P. ovata and drive adaptive divergence in populations exposed to higher temperatures. Within the species' current range, warm-adapted genotypes are predicted to experience a fivefold expansion in climate niche by midcentury and could harbour key adaptations to cope with future climate. We recommend eight seed transfer zones and project each zone into its relative position in future climate. Prioritizing seed collection efforts on genotypes with expanding future habitat represents a promising strategy for restoration practitioners to address rapidly changing climates.

Drawing a line in the sand: Effectiveness of off-highway vehicle management in California's Sonoran desert.

Public land policies manage multiple uses while striving to protect vulnerable plant and wildlife habitats from degradation; yet the effectiveness of such policies are infrequently evaluated, particularly for remote landscapes that are difficult to monitor. We assessed the use and impacts of recreational vehicles on Mojave Desert washes (intermittent streams) in the Chemehuevi Desert Wildlife Management Area (DWMA) of southern California. Wash zones designated as open and closed to off-highway vehicle (OHV) activity were designed in part to protect Mojave desert tortoise (Gopherus agassizii) habitat while allowing recreation in designated areas. OHV tracks were monitored in washes located near access roads during winter and early spring holidays - when recreation is typically high - and at randomly dispersed locations away from roads. Washes near access roads had fewer vehicle tracks within closed than open zones; further away from roads, OHV tracks were infrequent and their occurrence was not different between wash designations. Washes were in better condition in closed zones following major holidays as indicated by less vegetation damage, presence of trash, and wash bank damage. Furthermore, the frequency of washes with live tortoises and their sign was marginally greater in closed than open wash zones. Collectively, these results suggest that low impacts to habitats in designated closed wash zones reflect public compliance with federal OHV policy and regulations in the Chemehuevi DWMA during our study. Future monitoring to contrast wash use and impacts during other seasons as well as in other DWMAs will elucidate spatial and temporal patterns of recreation in these important conservation areas.

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.

Yucca brevifolia fruit production, predispersal seed predation, and fruit removal by rodents during two years of contrasting reproduction.

PREMISE OF THE STUDY: The distribution of Yucca brevifolia, a keystone species of the Mojave Desert, may contract with climate change, yet reproduction and dispersal are poorly understood. We tracked reproduction, seed predation, and fruit dispersal for two years and discuss whether Y. brevifolia is a masting species. METHODS: Fruit maturation, seed predation (larval yucca moths), and fruit dispersal (rodents) were monitored on a random sample of panicles during 2013 and 2014, which were years of high and low reproduction, respectively. Fates of fruits placed on the ground and in canopies were also tracked. Rodents were live-trapped to assess abundance and species composition. KEY RESULTS: In 2013, 66% of inflorescences produced fruit of which 53% escaped larval predation; 19.5% of seeds were destroyed in infested fruits. Total seed production was estimated to be >100 times greater in 2013 than 2014. One-third of the fruit crop fell to the ground and was removed by rodents over the course of 120 d. After ground fruits became scarce, rodents exploited canopy fruits. Rodent numbers were low in 2013, so fruits remained in canopies for 370 d. In 2014, fruit production was approximately 20% lower. Larvae infested the majority of fruits, and almost twice the number of seeds were damaged. Fruits were exploited by rodents within 65 d. CONCLUSIONS: High fertilization, prolific seed production, and low predispersal predation in 2013 suggests that pollinator attraction and satiation of seed predators influence masting in Y. brevifolia. Abundant, prolonged fruit availability to seed-dispersing rodents likely extends recruitment opportunities during mast years.

Direct and indirect effects of environmental variability on growth and survivorship of pre-reproductive Joshua trees, Yucca brevifolia Engelm. (Agavaceae).

UNLABELLED: • PREMISE OF STUDY: Accurate demographic information about long-lived plant species is important for understanding responses to large-scale disturbances, including climate change. It is challenging to obtain these data from desert perennial plants because seedling establishment is exceptionally rare, and estimates of survival are lacking for their vulnerable early stages. Desert wildfires, urbanization, and climate change influence the persistence of the long-lived Yucca brevifolia. Quantitative demographic attributes are crucial for understanding how populations will respond to disturbances and where populations will recede or advance under future climate scenarios.• METHODS: We measured survival in a cohort of 53 pre-reproductive Y. brevifolia at Yucca Flat, Nevada, USA, for 22 yr and recorded their growth, nurse-plant relationships, and herbivory.• KEY RESULTS: Herbivory by black-tailed jackrabbits (Lepus californicus) caused severe losses of plants during the first and second years (45% and 31%, respectively). Surviving plants experienced <2.5% annual mortality. Survival for the population was 19% over 22 yr. Plants <25 cm in height had lower life expectancy. Average growth rate (± SD) for plants that survived to the last census was 3.12 ± 1.96 cm yr(-1), and growth rates were positively associated with precipitation. Thirty-year-old Y. brevifolia had not yet reproduced.• CONCLUSIONS: A rare establishment event for Y. brevifolia during 1983-1984, triggered by above-average summer rainfall, provided a unique opportunity to track early survival and growth. Infrequent but acute episodes of herbivory during drought influenced demography for decades. Variability in survival among young Y. brevifolia indicates that size-dependent demographic variables will improve forecasts for this long-lived desert species under predicted regional climate change.

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.

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

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.

Short-term soil inorganic N pulse after experimental fire alters invasive and native annual plant production in a Mojave Desert shrubland.

Post-fire changes in desert vegetation patterns are known, but the mechanisms are poorly understood. Theory suggests that pulse dynamics of resource availability confer advantages to invasive annual species, and that pulse timing can influence survival and competition among species. Precipitation patterns in the American Southwest are predicted to shift toward a drier climate, potentially altering post-fire resource availability and consequent vegetation dynamics. We quantified post-fire inorganic N dynamics and determined how annual plants respond to soil inorganic nitrogen variability following experimental fires in a Mojave Desert shrub community. Soil inorganic N, soil net N mineralization, and production of annual plants were measured beneath shrubs and in interspaces during 6 months following fire. Soil inorganic N pools in burned plots were up to 1 g m(-2) greater than unburned plots for several weeks and increased under shrubs (0.5-1.0 g m(-2)) more than interspaces (0.1-0.2 g m(-2)). Soil NO(3) (-)-N (nitrate-N) increased more and persisted longer than soil NH(4) (+)-N (ammonium-N). Laboratory incubations simulating low soil moisture conditions, and consistent with field moisture during the study, suggest that soil net ammonification and net nitrification were low and mostly unaffected by shrub canopy or burning. After late season rains, and where soil inorganic N pools were elevated after fire, productivity of the predominant invasive Schismus spp. increased and native annuals declined. Results suggest that increased N availability following wildfire can favor invasive annuals over natives. Whether the short-term success of invasive species following fire will direct long-term species composition changes remains to be seen, yet predicted changes in precipitation variability will likely interact with N cycling to affect invasive annual plant dominance following wildfire.

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.

Are Mojave Desert annual species equal? Resource acquisition and allocation for the invasive grass Bromus madritensis subsp. rubens (Poaceae) and two native species.

Abundance of invasive plants is often attributed to their ability ot outcompete native species. We compared resource acquisition and allocation of the invasive annual grass Bromus madritensis subsp. rubens with that of two native Mojave Desert annuals, Vulpia octoflora and Descurainia pinnata, in a glasshouse experiment. Each species was grown in monoculture at two densities and two levels of N availability to compare how these annuals capture resources and to understand their relative sensitivities to environmental change. During >4 mo of growth, Bromus used water more rapidly and had greater biomass and N content than the natives, partly because of its greater root-surface area and its exploitation of deep soils. Bromus also had greater N uptake, net assimilation and transpiration rates, and canopy area than Vulpia. Resource use by Bromus was less sensitive to changes in N availability or density than were the natives. The two native species in this study produced numerous small seeds that tended to remain dormant, thus ensuring escape of offspring from unfavorable germination conditions; Bromus produced fewer but larger seeds that readily germinated. Collectively, these traits give Bromus the potential to rapidly establish in diverse habitats of the Mojave Desert, thereby gaining an advantage over coexisting native species.