Structural changes in vegetation coincident with annual grass invasion negatively impacts sprint velocity of small vertebrates

Sagebrush ecosystems in the intermountain west of the United States are being threatened by conversion to the non-native grass, cheatgrass (Bromus tectorum). The dramatic shift in the physical structure of vegetation coincident with cheatgrass invasion likely has negative impacts on animal communities, yet these structural impacts have not been well-studied. In a previous study, dense cheatgrass stems reduced sprint velocity for the flattened, wide-bodied desert horned lizard (Phrynosoma platyrhinos). Here, we asked if a decrease in sprint velocity due to cheatgrass impediment can be generalized to the suite of small vertebrates inhabiting the sagebrush ecosystems of western Utah. We evaluated sprint performance of the common rodent (n = 3) and lizard (n = 4) species on two raceway types, cheatgrass and no-cheatgrass, and hypothesized that body size, body shape, and form of movement are important factors influencing sprint velocity through dense cheatgrass stems. All species showed significant reductions in speed on cheatgrass versus no-cheatgrass raceways, with percent reduction greatest for larger, wider, or hopping organisms compared to smaller, more slender, or running organisms. Of concern, surveys for rodents and lizards at our study areas support a common pattern: lower abundances of small vertebrates, as well as a loss of rodent species richness, in areas infested with cheatgrass compared to intact, native sagebrush communities. By extension, we expect a negative impact on animal communities in other semi-arid regions experiencing dramatic shifts in vegetation structure upon invasion by non-native grasses that are capable of forming dense stands in the interspaces of native desert plants [e.g., Sonoran desert invaded by buffelgrass (Pennisetum ciliare)].     

Seed harvesting by a generalist consumer is context‐dependent: Interactive effects across multiple spatial scales

Granivore foraging decisions affect consumer success and determine the quantity and spatial pattern of seed survival. These decisions are influenced by environmental variation at spatial scales ranging from landscapes to local foraging patches. In a field experiment, the effects of seed patch variation across three spatial scales on seed removal by western harvester ants Pogonomyrmex occidentalis were evaluated. At the largest scale we assessed harvesting in different plant communities, at the intermediate scale we assessed harvesting at different distances from ant mounds, and at the smallest scale we assessed the effects of interactions among seed species in local seed neighborhoods on seed harvesting (i.e. resource–consumer interface). Selected seed species were presented alone (monospecific treatment) and in mixture with Bromus tectorum (cheatgrass; mixture treatment) at four distances from P. occidentalis mounds in adjacent intact sagebrush and non‐native cheatgrass‐dominated communities in the Great Basin, Utah, USA. Seed species differed in harvest, with B. tectorum being least preferred. Large and intermediate scale variation influenced harvest. More seeds were harvested in sagebrush than in cheatgrass‐dominated communities (largest scale), and the quantity of seed harvested varied with distance from mounds (intermediate‐scale), although the form of the distance effect differed between plant communities. At the smallest scale, seed neighborhood affected harvest, but the patterns differed among seed species considered. Ants harvested fewer seeds from mixed‐seed neighborhoods than from monospecific neighborhoods, suggesting context dependence and potential associational resistance. Further, the effects of plant community and distance from mound on seed harvest in mixtures differed from their effects in monospecific treatments. Beyond the local seed neighborhood, selection of seed resources is better understood by simultaneously evaluating removal at multiple scales. Associational effects provide a useful theoretical basis for better understanding harvester ant foraging decisions. These results demonstrate the importance of ecological context for seed removal, which has implications for seed pools, plant populations and communities.     

Net carbon exchange and evapotranspiration in postfire and intact sagebrush communities in the Great Basin

Invasion of non-native annuals across the Intermountain West is causing a widespread transition from perennial sagebrush communities to fire-prone annual herbaceous communities and grasslands. To determine how this invasion affects ecosystem function, carbon and water fluxes were quantified in three, paired sagebrush and adjacent postfire communities in the northern Great Basin using a 1-m³ gas exchange chamber. Most of the plant cover in the postfire communities was invasive species including Bromus tectorum L., Agropyron cristatum (L.) Gaertn and Sisymbrium altissimum L. Instantaneous morning net carbon exchange (NCE) and evapotranspiration (ET) in native shrub plots were greater than either intershrub or postfire plots. Native sagebrush communities were net carbon sinks (mean NCE 0.2–4.3 μmol m⁻² s⁻¹) throughout the growing season. The magnitude and seasonal variation of NCE in the postfire communities were controlled by the dominant species and availability of soil moisture. Net C exchange in postfire communities dominated by perennial bunchgrasses was similar to sagebrush. However, communities dominated by annuals (cheatgrass and mustard) had significantly lower NCE than sagebrush and became net sources of carbon to the atmosphere (NCE declined to −0.5 μmol m⁻² s⁻¹) with increased severity of the summer drought. Differences in the patterns of ET led to lower surface soil moisture content and increased soil temperatures during summer in the cheatgrass-dominated community compared to the adjacent sagebrush community. Intensive measurements at one site revealed that temporal and spatial patterns of NCE and ET were correlated most closely with changes in leaf area in each community. By altering the patterns of carbon and water exchange, conversion of native sagebrush to postfire invasive communities may disrupt surface-atmosphere exchange and degrade the carbon storage capacity of these systems.     

Enemy release from the effects of generalist granivores can facilitate Bromus tectorum invasion in the Great Basin Desert

The enemy release hypothesis (ERH) of plant invasion asserts that natural enemies limit populations of invasive plants more strongly in native ranges than in non‐native ranges. Despite considerable empirical attention, few studies have directly tested this idea, especially with respect to generalist herbivores. This knowledge gap is important because escaping the effects of generalists is a critical aspect of the ERH that may help explain successful plant invasions. Here, we used consumer exclosures and seed addition experiments to contrast the effects of granivorous rodents (an important guild of generalists) on the establishment of cheatgrass (Bromus tectorum) in western Asia, where cheatgrass is native, versus the Great Basin Desert, USA, where cheatgrass is exotic and highly invasive. Consistent with the ERH, rodent foraging reduced cheatgrass establishment by nearly 60% in western Asia but had no effect in the Great Basin. This main result corresponded with a region‐specific foraging pattern: rodents in the Great Basin but not western Asia generally avoided seeds from cheatgrass relative to seeds from native competitors. Our results suggest that enemy release from the effects of an important guild of generalists may contribute to the explosive success of cheatgrass in the Great Basin. These findings corroborate classic theory on enemy release and expand our understanding of how generalists can influence the trajectory of exotic plant invasions.     

Negative Effects of an Exotic Grass Invasion on Small-Mammal Communities

Exotic invasive species can directly and indirectly influence natural ecological communities. Cheatgrass (Bromus tectorum) is non-native to the western United States and has invaded large areas of the Great Basin. Changes to the structure and composition of plant communities invaded by cheatgrass likely have effects at higher trophic levels. As a keystone guild in North American deserts, granivorous small mammals drive and maintain plant diversity. Our objective was to assess potential effects of invasion by cheatgrass on small-mammal communities. We sampled small-mammal and plant communities at 70 sites (Great Basin, Utah). We assessed abundance and diversity of the small-mammal community, diversity of the plant community, and the percentage of cheatgrass cover and shrub species. Abundance and diversity of the small-mammal community decreased with increasing abundance of cheatgrass. Similarly, cover of cheatgrass remained a significant predictor of small-mammal abundance even after accounting for the loss of the shrub layer and plant diversity, suggesting that there are direct and indirect effects of cheatgrass. The change in the small-mammal communities associated with invasion of cheatgrass likely has effects through higher and lower trophic levels and has the potential to cause major changes in ecosystem structure and function.     

Native granivores reduce the establishment of native grasses but not invasive <Emphasis Type="Italic">Bromus tectorum</Emphasis>

Seed predation can structure plant communities by imposing strong population controls on some species but not others. In this context, studies from various ecosystems report that native granivores selectively forage for seeds from native species over seeds from exotic invaders, which could disproportionately favor the establishment of invaders and facilitate their dominance in communities. However, few studies have connected selective foraging for native seeds to differential patterns of establishment among native and invasive species. Thus, the extent to which preferential foraging for native seeds favors the establishment of invasive plants is unclear. Here, we used experimental seed additions and exclosure treatments at five field sites distributed across?≈?80,000 km² of the Great Basin Desert, USA to compare the effects of rodent foraging on the establishment of less-preferred cheatgrass (Bromus tectorum—an annual species native to Eurasia that is exotic and highly invasive across the Great Basin) and four species of more-preferred native grasses that commonly co-occur with cheatgrass. Rodent foraging reduced the establishment of each native species by at least 80% but had no effect on the establishment of cheatgrass, and this finding was consistent across study sites. Our results suggest that selective foraging for native species may favor the establishment of cheatgrass over native grasses, potentially exacerbating one of the most extensive plant invasions in North America.     

Introduced annual grass increases regional fire activity across the arid western USA (1980–2009)

Non‐native, invasive grasses have been linked to altered grass‐fire cycles worldwide. Although a few studies have quantified resulting changes in fire activity at local scales, and many have speculated about larger scales, regional alterations to fire regimes remain poorly documented. We assessed the influence of large‐scale Bromus tectorum (hereafter cheatgrass) invasion on fire size, duration, spread rate, and interannual variability in comparison to other prominent land cover classes across the Great Basin, USA. We compared regional land cover maps to burned area measured using the Moderate Resolution Imaging Spectroradiometer (MODIS) for 2000–2009 and to fire extents recorded by the USGS registry of fires from 1980 to 2009. Cheatgrass dominates at least 6% of the central Great Basin (650 000 km²). MODIS records show that 13% of these cheatgrass‐dominated lands burned, resulting in a fire return interval of 78 years for any given location within cheatgrass. This proportion was more than double the amount burned across all other vegetation types (range: 0.5–6% burned). During the 1990s, this difference was even more extreme, with cheatgrass burning nearly four times more frequently than any native vegetation type (16% of cheatgrass burned compared to 1–5% of native vegetation). Cheatgrass was also disproportionately represented in the largest fires, comprising 24% of the land area of the 50 largest fires recorded by MODIS during the 2000s. Furthermore, multi‐date fires that burned across multiple vegetation types were significantly more likely to have started in cheatgrass. Finally, cheatgrass fires showed a strong interannual response to wet years, a trend only weakly observed in native vegetation types. These results demonstrate that cheatgrass invasion has substantially altered the regional fire regime. Although this result has been suspected by managers for decades, this study is the first to document recent cheatgrass‐driven fire regimes at a regional scale.     

Fire effects on the mobilization and uptake of nitrogen by cheatgrass (Bromus tectorum L.)

Cheatgrass (Bromus tectorum L.), an invasive annual grass, is displacing native species and causing increased fire frequency in the Great Basin of the southwestern United States. Growth and nitrogen uptake patterns by cheatgrass were examined in a greenhouse study using soils from sites with the same soil type but different fire histories: 1) an area that burned in 1999 that is now completely invaded with cheatgrass (CG); 2) an area that has not burned recently and is now dominated by Wyoming big sagebrush (Artemisia tridentatassp.wyomingensis Beetle and Young) and Sandberg’s bluegrass (Poa secunda J. Presl) (WBS); and 3) a Wyoming big sagebrush area that burned in August of 2008 just prior to soil collection (NB). Cheatgrass seedlings had higher leaf numbers, height and mass in the NB soil. Ammonium-N mobilized by fire in the NB soil had significantly enriched ¹⁵N than soils from CG or WBS sites and this pattern was reflected in the isotopic signatures of the plants. Fire-mobilized mineral N accounted for only 58% of N taken up by cheatgrass in the NB soil, suggesting fire enhanced the ability of cheatgrass to assimilate more recalcitrant soil organic N.     

Relative Abundance of and Composition within Fungal Orders Differ between Cheatgrass (Bromus tectorum) and Sagebrush (Artemisia tridentata)-Associated Soils

Nonnative Bromus tectorum (cheatgrass) is decimating sagebrush steppe, one of the largest ecosystems in the Western United States, and is causing regional-scale shifts in the predominant plant-fungal interactions. Sagebrush, a native perennial, hosts arbuscular mycorrhizal fungi (AMF), whereas cheatgrass, a winter annual, is a relatively poor host of AMF. This shift is likely intertwined with decreased carbon (C)-sequestration in cheatgrass-invaded soils and alterations in overall soil fungal community composition and structure, but the latter remain unresolved. We examined soil fungal communities using high throughput amplicon sequencing (ribosomal large subunit gene) in the 0–4 cm and 4–8 cm depth intervals of six cores from cheatgrass- and six cores from sagebrush-dominated soils. Sagebrush core surfaces (0–4 cm) contained higher nitrogen and total C than cheatgrass core surfaces; these differences mirrored the presence of glomalin related soil proteins (GRSP), which has been associated with AMF activity and increased C-sequestration. Fungal richness was not significantly affected by vegetation type, depth or an interaction of the two factors. However, the relative abundance of seven taxonomic orders was significantly affected by vegetation type or the interaction between vegetation type and depth. Teloschistales, Spizellomycetales, Pezizales and Cantharellales were more abundant in sagebrush libraries and contain mycorrhizal, lichenized and basal lineages of fungi. Only two orders (Coniochaetales and Sordariales), which contain numerous economically important pathogens and opportunistic saprotrophs, were more abundant in cheatgrass libraries. Pleosporales, Agaricales, Helotiales and Hypocreales were most abundant across all libraries, but the number of genera detected within these orders was as much as 29 times lower in cheatgrass relative to sagebrush libraries. These compositional differences between fungal communities associated with cheatgrass- and sagebrush-dominated soils warrant future research to examine soil fungal community composition across more sites and time points as well as in association with native grass species that also occupy cheatgrass- invaded ecosystems.     

Biosolids Application and Long‐Term Noxious Weed Dominance in the Western United States

Vegetation characteristics were assessed on three sets of 10‐year‐old test plots and one set of 5‐year‐old plots that received 0, 34, 45, and 67 tons/ha (0, 15, 20, and 30 short tons/acre) of biosolids at a semiarid mine reclamation site in Utah. On average, noxious weed species such as Bromus tectorum L. (cheatgrass) provided two‐thirds of the cover on the biosolids test plots, but only one‐tenth of the cover on adjacent control plots that received no biosolids. Cheatgrass provided more than half of the total cover on every biosolids test plot. Seeded species provided about two times more cover at the control plots than at the biosolids plots. Surfaces treated with 45 tons/ha composted biosolids (one part biosolids and two parts wood chips) had a much lower percentage of noxious weed cover compared to biosolids alone. The relatively heavy initial nitrogen load associated with biosolids application may have promoted cheatgrass dominance. Although the available nitrogen eventually declines, once cheatgrass is established it may maintain its dominance indefinitely. Given the risk of weed invasion, heavy biosolids applications should be used with caution for reclamation projects in semiarid climates if perennial species establishment is desired. Consideration should instead be given to light applications (<45 tons/ha) of biosolids/wood chip compost or forgoing the use of biosolids entirely. The underapplication of nutrients may provide a slower, but ultimately more reliable, strategy for the establishment of a healthy, native perennial vegetation community.     

Invasion resistance and persistence: established plants win,even with disturbance and high propagule pressure

Disturbances and propagule pressure are key mechanisms in plant community resistance to invasion, as well as persistence of invasions. Few studies, however, have experimentally tested the interaction of these two mechanisms. We initiated a study in a southwestern ponderosa pine (Pinus ponderosa Laws.)/bunch grass system to determine the susceptibility of remnant native plant communities to cheatgrass (Bromus tectorum L.) invasion, and persistence of cheatgrass in invaded areas. We used a 2 × 2 factorial design consisting of two levels of aboveground biomass removal and two levels of reciprocal seeding. We seeded cheatgrass seeds in native plots and a native seed mixture in cheatgrass plots. Two biomass removal disturbances and sowing seeds over 3 years did not reverse cheatgrass dominance in invaded plots or native grass dominance in non-invaded native plots. Our results suggest that two factors dictated the persistence of the resident communities. First, bottlebrush squirreltail (Elymus elymoides (Raf.) Swezey) was the dominant native herbaceous species on the study site. This species is typically a poor competitor with cheatgrass as a seedling, but is a strong competitor when mature. Second, differences in pretreatment levels of plant-available soil nitrogen and phosphorus may have favored the dominant species in each community. Annual species typically require higher levels of plant-available soil nutrients than perennial plants. This trend was observed in the annual cheatgrass community and perennial native community. Our study shows that established plants and soil properties can buffer the influences of disturbance and elevated propagule pressure on cheatgrass invasion.     

Postwildfire seeding to restore native vegetation and limit exotic annuals: an evaluation in juniper‐dominated sagebrush steppe

Reestablishment of perennial vegetation is often needed after wildfires to limit exotic species and restore ecosystem services. However, there is a growing body of evidence that questions if seeding after wildfires increases perennial vegetation and reduces exotic plants. The concern that seeding may not meet restoration goals is even more prevalent when native perennial vegetation is seeded after fire. We evaluated vegetation cover and density responses to broadcast seeding native perennial grasses and mountain big sagebrush (Artemisia tridentata Nutt. spp. vaseyana [Rydb.] Beetle) after wildfires in the western United States in six juniper (Juniperus occidentalis ssp. occidentalis Hook)‐dominated mountain big sagebrush communities for 3 years postfire. Seeding native perennial species compared to not seeding increased perennial grass and sagebrush cover and density. Perennial grass cover was 4.3 times greater in seeded compared to nonseeded areas. Sagebrush cover averaged 24 and less than 0.1% in seeded and nonseeded areas at the conclusion of the study, respectively. Seeding perennial species reduced exotic annual grass and annual forb cover and density. Exotic annual grass cover was 8.6 times greater in nonseeded compared to seeded areas 3 years postfire. Exotic annual grass cover increased over time in nonseeded areas but decreased in seeded areas by the third‐year postfire. Seeded areas were perennial‐dominated and nonseeded areas were annual‐dominated at the end of the study. Establishing perennial vegetation may be critical after wildfires in juniper‐dominated sagebrush steppe to prevent the development of annual‐dominated communities. Postwildfire seeding increased perennial vegetation and reduced exotic plants and justifies its use.     

Evaluating strategies for facilitating native plant establishment in northern Nevada crested wheatgrass seedings

Non‐native crested wheatgrasses (Agropyron cristatum and A. desertorum) were used historically within the Great Basin for the purpose of competing with weed species and increasing livestock forage. These species continue to be used in some areas, especially after wildfires occurring in low elevation/precipitation, formerly Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis)/herbaceous communities. Seeding native species in these sites is often unsuccessful, and lack of establishment results in invasion and site dominance by exotic annuals. However, crested wheatgrass often forms dense monocultures that interfere competitively with the establishment of desirable native vegetation and do not provide the plant structure and habitat diversity for wildlife species equivalent to native‐dominated sagebrush plant communities. During a 5‐year study, we conducted trials to evaluate chemical and mechanical methods for reducing crested wheatgrass and the effectiveness of seeding native species into these sites after crested wheatgrass suppression. We determined that discing treatments were ineffective in reducing crested wheatgrass cover and even increased crested wheatgrass density in some cases. Glyphosate treatments initially reduced crested wheatgrass cover, but weeds increased in many treated plots and seeded species diminished over time as crested wheatgrass recovered. We concluded that, although increases in native species could possibly be obtained by repeating crested wheatgrass control treatments, reducing crested wheatgrass opens a window for invasion by exotic weed species.     

Simulating cheatgrass (<Emphasis Type="Italic">Bromus tectorum</Emphasis>) invasion decreases access to food resources for small mammals in sagebrush steppe

Invasions by nonnative plants can alter the abundance of native animals, yet we know little about the mechanisms driving these changes. Shifts in vegetation characteristics resulting from nonnative plants can alter availability of food resources, predation risk, and foraging efficiency (both the access to and ability to find food), each providing a potential mechanism for documented changes in animal communities and populations in invaded systems. Cheatgrass (Bromus tectorum) is a nonnative grass that invades sagebrush steppe, resulting in declines in some small mammal populations. We examined whether changes in structural characteristics associated with cheatgrass invasion could alter foraging by small mammals, providing a potential mechanism for documented population declines. We quantified differences in vegetation structure between native and cheatgrass-invaded sagebrush steppe, then experimentally added artificial structure in native areas to simulate these differences. We placed grain at foraging stations and measured the amount removed by small mammals nightly. Adding litter at depths approximating invasion by cheatgrass reduced the average amount of grain removed in 2 of 3 study areas, but increasing stem density did not. Based on this experiment, the deeper litter created by cheatgrass invasion may increase costs to small mammals by decreasing foraging efficiency and access to existing food resources, which may explain population-level declines in small mammals documented in other studies. By isolating and identifying which structural attributes of cheatgrass invasion are most problematic for small mammals, land managers may be able to design treatments to efficiently mitigate impacts and restore invaded ecosystems.     

Fire, native species, and soil resource interactions influence the spatio-temporal invasion pattern of Bromus tectorum

Bromus tectorum (cheatgrass) is an invasive annual that occupies perennial grass and shrub communities throughout the western United States. Bronus tectorum exhibits an intriguing spatio‐temporal pattern of invasion in low elevation ponderosa pine Pinus ponderosa/bunchgrass communities in western Montana where it forms dense rings beneath solitary pines following fire. This pattern provides a unique opportunity to investigate several indirect effects of native vegetation that influence the invasion pattern of B. tectorum, and specifically how native species, disturbance, and soil resources interact to influence the spatio‐temporal pattern of invasion. We established four replicate field sites, each containing burned‐tree, burned‐grass, unburned‐tree, and unburned‐grass sampling locations, and initiated a series of field sampling and greenhouse experiments utilizing these locations. The objective of our first greenhouse experiment was to identify whether belowground factors contributed to the pattern of B. tectorum biomass observed in these field locations. This experiment generated a B. tectorum biomass response that was nearly identical to the invasion pattern observed in the field, suggesting further investigation of belowground factors was necessary. We measured resin‐sorbed NH₄⁺ and NO₃⁻ during one generation of B. tectorum, and measured a suite of P fractions through a sequential extraction procedure from these soils. These data revealed that a resource island of high N and P exists beneath pine trees. Through a second greenhouse experiment, we determined that N limited B. tectorum biomass in tree soil, whereas P limited biomass in bunchgrass soil. Finally, through a germination experiment we determined that pine litter strongly inhibited B. tectorum germination. These data suggest B. tectorum is regulated by P in bunchgrass soil, and by N and inhibition by pine litter beneath trees, effects that are likely alleviated by fire. These data demonstrate the combined role of direct and indirect interactions between native and invasive species in regulating biological invasions.     

Bromus tectorum Response to Fire Varies with Climate Conditions

The invasive annual grass Bromus tectorum (cheatgrass) forms a positive feedback with fire in some areas of western North America’s sagebrush biome by increasing fire frequency and size, which then increases B. tectorum abundance post-fire and dramatically alters ecosystem structure and processes. However, this positive response to fire is not consistent across the sagebrush steppe. Here, we ask whether different climate conditions across the sagebrush biome can explain B. tectorum’s variable response to fire. We found that climate variables differed significantly between 18 sites where B. tectorum does and does not respond positively to fire. A positive response was most likely in areas with higher annual temperatures and lower summer precipitation. We then chose a climatically intermediate site, with intact sagebrush vegetation, to evaluate whether a positive feedback had formed between B. tectorum and fire. A chronosequence of recent fires (1–15 years) at the site created a natural replicated experiment to assess abundance of B. tectorum and native plants. B. tectorum cover did not differ between burned and unburned plots but native grass cover was higher in recently burned plots. Therefore, we found no evidence for a positive feedback between B. tectorum and fire at the study site. Our results suggest that formation of a positive B. tectorum-fire feedback depends on climate; however, other drivers such as disturbance and native plant cover are likely to further influence local responses of B. tectorum. The dependence of B. tectorum’s response to fire on climate suggests that climate change may expand B. tectorum’s role as a transformative invasive species within the sagebrush biome.     

Expanding and invading plant species in sagebrush steppe affect multiple aspects of small-mammal ecology

Invasion and expansion of non-native and native plants have altered vegetation structure in many terrestrial ecosystems. Small mammals influence multiple ecosystem processes through their roles as ecosystem engineers, predators, and prey, and changes to vegetation structure can affect habitat use, community composition, and predator-prey interactions for this assemblage of wildlife. In the sagebrush (Artemisia spp.) shrublands of the western United States, invasion by non-native grasses and expansion of native conifer trees beyond their historical range has altered vegetation structure. These changes may potentially affect distributions and interactions of deer mice (Peromyscus maniculatus), which are generalist omnivores, and Columbia Plateau pocket mice (Perognathus parvus), more specialized granivores. To assess the extent to which altered habitat affects small-mammal density, survival, and home-range size, we examined these aspects of small-mammal ecology along a gradient of cheatgrass (Bromus tectorum) invasion and western juniper (Juniperus occidentalis) establishment in sagebrush shrublands in southwestern Idaho, USA. From 2017–2019, we used a spatially explicit mark-recapture design to examine attributes of small-mammal ecology along an invasion gradient. We did not find support for an effect of cheatgrass cover on density or survival of either species. Home-range size of deer mice was 2.3 times smaller in heavier cheatgrass cover (60%) compared to areas with little or no cheatgrass cover. Density of deer mice was highest (5 individuals/ha) in areas with 10% juniper cover and decreased with increasing juniper cover, whereas density of pocket mice was positively influenced by shrub cover. Survival of deer mice declined as juniper stem density increased. Conversely, survival of pocket mice increased with increasing juniper stem density. We found evidence for interspecific interactions between these 2 species, in the form of a density-dependent effect of deer mice on pocket mouse home-range size. Home-range size for pocket mice was 2 times smaller in areas with the highest estimated density of deer mice compared to areas with low densities of deer mice. Our data provides unique information about how small mammals in the sagebrush steppe are affected by expanding and invasive plant species and potential ways that habitat restoration efforts, in the form of conifer removal, may influence small mammals. Understanding the response of small mammals to conifer expansion or removal may shed light on the demographic and numerical responses of other wildlife associated with the sagebrush biome, including predators.     

Controls over invasion of Bromus tectorum: The importance of climate,soil, disturbance and seed availability

Question: Predicting the future abundance and distribution of invasive plants requires knowing how they respond to environmental conditions. In arid and semi‐arid ecosystems where water is a limiting resource, environmental conditions and disturbance patterns influence invasions by altering acquisition and utilization of water over space and time. We ask: 1. How do variations in climatic and soil properties influence temporal soil water dynamics? 2. How does this variation affect the establishment of Bromus tectorum (cheatgrass), a cool‐season annual grass that has successfully colonized much of the U.S. Great Basin? Location: Short‐grass Steppe in northeastern Colorado, USA; Arid Lands Ecology reserve in southeastern Washington, USA; and the Patagonian steppe of the Chubut province in Argentina. Methods: We utilized a soil water model to simulate seasonal soil water dynamics in multiple combinations of climatic and soil properties. In addition, we utilized a gap dynamics model to simulate the impact of disturbance regime and seed availability on competition between B. tectorum and native plants. Results: Our results suggest that climate is very important, but that soil properties do not significantly influence the probability of observing conditions suitable for B. tectorum establishment. Results of the plant competition model indicate that frequent disturbance causes more Bromus tectorum in invaded areas and higher seed availability causes faster invasion. Conclusions: These results imply a general framework for understanding Bromus tectorum invasion in which climatic conditions dictate which areas are susceptible to invasion, disturbance regime dictates the severity of invasion and seed availability dictates the speed of invasion.     

Strong genetic differentiation in the invasive annual grass <Emphasis Type="Italic">Bromus tectorum</Emphasis> across the Mojave–Great Basin ecological transition zone

Bromus tectorum, an inbreeding annual grass, is a dominant invader in sagebrush steppe habitat in North America. It is also common in warm and salt deserts, displaying a larger environmental tolerance than most native species. We tested the hypothesis that a suite of habitat-specific B. tectorum lineages dominates warm desert habitats. We sampled 30 B. tectorum Mojave Desert and desert fringe populations and genotyped 10–26 individuals per population using 69 single nucleotide polymorphic (SNP) markers. We compared these populations to 11 Great Basin steppe and salt desert populations. Populations from warm desert habitats were dominated by members of two haplogroups (87 % of individuals) that were distinct from haplogroups common in Great Basin habitats. We conducted common garden studies comparing adaptive traits and field performance among haplogroups typically found in different habitats. In contrast to the haplogroup abundant in sagebrush steppe, warm desert haplogroups generally lacked a vernalization requirement for flowering. The most widespread warm desert haplogroup (Warm Desert 1) also had larger seeds and a higher root:shoot ratio than other haplogroups. In the field, performance of warm desert haplogroups was dramatically lower than the sagebrush steppe haplogroup at one steppe site, but one warm desert haplogroup performed as well as the steppe haplogroup under drought conditions at the other site. Our results suggest that B. tectorum succeeds in widely disparate environments through ecotypic variation displayed by distinct lineages of plants. Accounting for this ecotypic variation is essential in modeling its future distribution in response to climate change.     

Edaphic limitations to growth and photosynthesis in Sierran and Great Basin vegetation

Summary Soils derived from hydrothermally altered andesite support unique communities of Sierran conifers (Pinus ponderosa Laws. and P. jeffreyi Grev. and Balf.) amongst sagebrush (Artemisia tridentata Nutt.) vegetation in the western Great Basin. Plants grown in soil derived from hydrothermally altered bedrock had lower growth rates, total biomass, and net photosynthetic rates than plants grown in soil derived from unaltered andesite of the same formation. Total dry mass was 10 to 28% lower for conifers grown in altered soil whereas dry mass of Artemisia tridentata and Bromus tectorum L. was reduced by over 90%. Results from a nutrient amendment experiment indicated that low phosphorus was the dominant limitation in altered soil, and phosphorus-deficiency affected growth primarily by limiting leaf area development rather than direct inhibition of photosynthesis. The proportionately greater reduction of biomass for Artemisia and Bromus grown in altered soil supports our hypothesis that Great Basin vegetation is excluded from altered soil by intolerance to nutrient deficiency. The Sierran conifers growing on this rock type are therefore free of competition for water with Great Basin vegetation and are able to persist in an exceptionally dry climate.