Squirreltail Research Articles

Seed Production and Seedling Fitness Are Uncoupled from Maternal Plant Productivity in Three Aridland Bunchgrasses

Maintaining self-sustaining populations of desired plants is fundamental to rangeland management, and understanding the relationships among plant growth, seed production, and seedling recruitment is critical to these efforts. Our objective was to evaluate how changes in maternal plant soil resource environment influences maternal plant biomass and seed production and seedling fitness in three widespread perennial bunchgrass species (Elymus elymoides [Raf.] Sezey, bottlebrush squirreltail; Festuca idahoensis Elmer, Idaho fescue; and Pseudoroegneria spicata [Pursh] A. Löve, bluebunch wheatgrass). We supplied water and nutrients to adult plants growing in the field and measured their productivity and fecundity. Then, in the laboratory, growth chamber, and field we assessed effects of the maternal water and nutrient additions on offspring performance. Across the three study species, vegetative traits were more plastic than reproductive traits, with resource addition measurably increasing plant growth but not seed production. Germination was high in both the laboratory and field across treatments, although seeds from irrigated maternal plants tended to have higher field germination. Seedling relative growth rate, leaf mass ratio, and relative root elongation rate (RRER) were highly variable, although RRER tended to be higher in seedlings derived from irrigated maternal plants. In the field, seedling survivorship was low across all species, but survivorship doubled in seedlings produced by P. spicata plants that received additional water through the growing season. Overall, our results suggest that biomass production and fecundity responses to nutrients are decoupled in the species and environment tested but maternal effects can have significant, although variable, impacts in some grassland species. As a result, biomass responses to natural and anthropogenic changes in resource availability may not be strong predictors of how altered resource supply may ultimately influence plant community dynamics in aridland systems.

Notice of release of Turkey Lake germplasm of bottlebrush squirreltail

Turkey Lake Germplasm of bottlebrush squirreltail ( Elymus elymoides (Raf.) Swezey ssp. californicus (J.G. Smith) Barkworth [Poaceae]) was released by USDA Agricultural Research Service in 2015 as a selected class of pre-variety germplasm (natural track). This new plant material originates in Gooding County in southern Idaho’s Snake River Plain. In replicated rangeland trials at 3 locations (Juab County, Utah; Sanpete County, Utah; White Pine County, Nevada), Turkey Lake Germplasm displayed substantially better establishment frequency than Toe Jam Creek Germplasm, a previous release of E. elymoides ssp. californicus (origin: Elko County, Nevada), as well as other released squirreltail plant materials. Comparative trials were also conducted in Cache County, Utah, and Bingham County, Idaho.

Temporal Variability in Microclimatic Conditions for Grass Germination and Emergence in the Sagebrush Steppe

Sagebrush steppe ecosystems in the western United States are characterized by harsh environmental conditions with high annual and seasonal variability in both precipitation and temperature. Environmental variability contributes to widespread failure in establishing stands of desired species on degraded and invaded landscapes. To characterize seasonal microclimatic patterns and planting date effects on restoration outcomes, we evaluated long-term simulations of seed germination response of cheatgrass (Bromus tectorum L.), bottlebrush squirreltail (Elymus elymoides [Raf] Swezey), and Idaho fescue (Festuca idahoensis Elmer) to annual patterns of soil temperature and moisture. Extremely high annual variability in both the conditions favorable for germination and patterns of post-germination drought and thermal stress make it difficult to justify general inferences about seedbed treatment and planting date effects from individual, short-term field studies. We discuss the interpretation of individual-year and seasonal plant establishment factors and offer a mechanistic model for interpreting planting date and year effects on initial seedling establishment. Historical ranking and mechanistic descriptions of individual-year seedbed conditions may allow for expanded inferences through meta-analysis of limited-term field experiments.

Grass-Shrub Associations over a Precipitation Gradient and Their Implications for Restoration in the Great Basin, USA

As environmental stress increases positive (facilitative) plant interactions often predominate. Plant-plant associations (or lack thereof) can indicate whether certain plant species favor particular types of microsites (e.g., shrub canopies or plant-free interspaces) and can provide valuable insights into whether “nurse plants” will contribute to seeding or planting success during ecological restoration. It can be difficult, however, to anticipate how relationships between nurse plants and plants used for restoration may change over large-ranging, regional stress gradients. We investigated associations between the shrub, Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis), and three common native grasses (Poa secunda, Elymus elymoides, and Pseudoroegneria spicata), representing short-, medium-, and deep-rooted growth forms, respectively, across an annual rainfall gradient (220–350 mm) in the Great Basin, USA. We hypothesized that positive shrub-grass relationships would become more frequent at lower rainfall levels, as indicated by greater cover of grasses in shrub canopies than vegetation-free interspaces. We sampled aerial cover, density, height, basal width, grazing status, and reproductive status of perennial grasses in canopies and interspaces of 25–33 sagebrush individuals at 32 sites along a rainfall gradient. We found that aerial cover of the shallow rooted grass, P. secunda, was higher in sagebrush canopy than interspace microsites at lower levels of rainfall. Cover and density of the medium-rooted grass, E. elymoides were higher in sagebrush canopies than interspaces at all but the highest rainfall levels. Neither annual rainfall nor sagebrush canopy microsite significantly affected P. spicata cover. E. elymoides and P. spicata plants were taller, narrower, and less likely to be grazed in shrub canopy microsites than interspaces. Our results suggest that exploring sagebrush canopy microsites for restoration of native perennial grasses might improve plant establishment, growth, or survival (or some combination thereof), particularly in drier areas. We suggest that land managers consider the nurse plant approach as a way to increase perennial grass abundance in the Great Basin. Controlled experimentation will provide further insights into the life stage-specific effectiveness and practicality of a nurse plant approach for ecological restoration in this region.

Cheatgrass die-offs as an opportunity for restoration in the Great Basin, USA: Will local or commercial native plants succeed where exotic invaders fail?

Bromus tectorum (cheatgrass) has widely invaded the Great Basin, U.S.A. The sporadic natural phenomenon of complete stand failure (‘die-off’) of this invader may present opportunities to restore native plants. A recent die-off in Nevada was precision-planted with seeds of the native grasses Poa secunda (Sandberg bluegrass) and Elymus elymoides (bottlebrush squirreltail), of both local and nonlocal origin, to ask: 1) Can native species be restored in recent B. tectorum die-offs? And 2) Do local and nonlocal seeds differ in performance? Additionally, we asked how litter removal and water addition affected responses. Although emergence and growth of native seeds was lower in die-off than control plots early in year one, in year two, seedlings in die-offs had increased vigor and growth, at equal or higher densities, than control plots. Local seeds consistently outperformed nonlocal seeds for P. secunda, whereas for E. elymoides, nonlocal showed an advantage in the first season, but in the second season, there were more local seeds present under die-off and unraked conditions. Seedbed treatments affected performance, but did not notably improve establishment or modify other results. Our results warrant further investigation into die-off restoration as well as recognition of the importance of seed source selection in restoration.

Rhizosphere-mediated effects of the invasive grass Bromus tectorum L. and native Elymus elymoides on nitrogen cycling in Great Basin Desert soils

Background and aims There is evidence that the invasive grass Bromus tectorum can affect soil nitrogen (N) cycling, possibly leading to a positive plant-soil feedback. Rhizosphere priming of N mineralization could provide a mechanistic explanation for such a feedback. Methods We conducted a greenhouse study to isolate rhizosphere effects on N cycling by the invasive annual grass, Bromus tectorum L., and the native perennial grass, Elymus elymoides (Raf.) Swezey, in invaded and uninvaded soils. We compared the rhizosphere priming effect (RPE) on N mineralization by species and the distribution of N in various pools by planting treatment and soil type. Results B. tectorum had a negative RPE (�23 and �22 % in invaded and uninvaded soils, respectively), while E. elymoides had no significant RPE. B. tectorum was more competitive over E. elymoides in invaded compared to uninvaded soil. Conclusions B. tectorumhad a negative effect on soil N availability via root-mediated processes, even though its growth and competitiveness increased in invaded soils. Positive plant-soil feedback effects of B. tectorum may be mediated by aboveground inputs rather than belowground and/or depend on site-specific conditions.

New experimental hosts of Barley yellow dwarf virus among wild grasses, with implications for grassland habitats

Barley yellow dwarf virus (BYDV), an economically important virus, infects small grain cereal crops and over 150 other Poaceae species. BYDV infection plays an important role in competition among grasses in non‐managed systems, but many grasses remain unexamined as potential BYDV hosts. This study examined grass species that have not been reported as BYDV hosts but are commonly encountered in non‐managed grasslands throughout the United States and Canada. Laboratory inoculations with BYDV‐PAV using the aphid vector Rhopalosiphum padi were performed to examine the ability of 13 grass species and barley to be infected with the virus; eight of the grass species were not documented previously as virus hosts. Serological and molecular assays were used to confirm BYDV‐PAV infection. Plant height, number of leaves, number of tillers and weight were recorded to evaluate susceptibility or sensitivity to BYDV. Infection with BYDV was experimentally achieved for the first time on Achnatherum lettermanii, Achnatherum occidentale, Achnatherum thurberianum, Danthonia intermedia, Poa fendleriana, Sporobolus airoides and Sporobolus cryptandrus, but not on Alopecurus pratensis and Elymus wawawaiensis. Infection was confirmed in Bromus inermis, Elymus elymoides, Poa bulbosa, Poa secunda and Hordeum vulgare, which served as controls. BYDV infection caused reductions in plant height on P. bulbosa and P. fendleriana. BYDV‐infected P. secunda had more leaves per plant compared to healthy plants of the same species. BYDV‐infected A. lettermanii exhibited reduced dry weight in both below‐ground and above‐ground tissue. These findings have implications for the management and conservation of grassland habitats.

Bidirectional recovery patterns of Mojave Desert vegetation in an aqueduct pipeline corridor after 36 years: I. Perennial shrubs and grasses

We studied recovery of 21 perennial plant species along a severely disturbed aqueduct corridor in a Larrea tridentata-Ambrosia dumosa plant alliance in the Mojave Desert 36 years after construction. The 97-m wide corridor contained a central dirt road and buried aqueduct pipeline. We established transects at 0 m (road verge), 20 m and 40 m into the disturbance corridor, and at 100 m in undisturbed habitat (the control). Although total numbers of shrubs per transect did not vary significantly with distance from the verge, canopy cover of shrubs, species richness, and species diversity were higher in the control than at the verge and other distances. Canopy cover of common shrubs (Ericameria nauseosa, Ambrosia salsola, A. dumosa, L. tridentata, Grayia spinosa) and perennial grasses (Elymus elymoides, Poa secunda) also varied significantly by location. Discriminant analysis clearly separated the four distances based on plant composition. Patterns of recovery were bidirectional: secondary succession from the control into the disturbance corridor and inhibition from the verge in the direction of the control. Time estimated for species composition to resemble the control is dependent on location within the disturbance corridor and could be centuries at the road verge. Our findings have applications to other deserts.

Abundances of Coplanted Native Bunchgrasses and Crested Wheatgrass after 13 Years

Crested wheatgrass (Agropyron cristatum [L] Gaertm) has been seeded on more than 5 million hectares in western North America because it establishes more readily than native bunchgrasses. Currently, there is substantial interest in reestablishing native species in sagebrush steppe, but efforts to reintroduce native grasses into crested wheatgrass stands have been largely unsuccessful, and little is known about the long-term dynamics of crested wheatgrass/native species mixes. We examined the abundance of crested wheatgrass and seven native sagebrush steppe bunchgrasses planted concurrently at equal low densities in nongrazed and unburned plots. Thirteen years post establishment, crested wheatgrass was the dominant bunchgrass, with a 10-fold increase in density. Idaho fescue (Festuca idahoensis Elmer), Thurber’s needlegrass (Achnatherum thurberianum (Piper) Barkworth), basin wildrye (Leymus cinereus [Scribn. & Merr.] A. Löve), and Sandberg bluegrass (Poa secunda J. Presl) maintained their low planting density, whereas bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] A. Löve), needle-and-thread (Hesperostipa comata [Trin. & Rupr.] Barkworth), and squirreltail (Elymus elymoides [Raf.] Swezey) densities declined. Our results suggest that densities of native bunchgrasses planted with crested wheatgrass are unlikely to increase and that some species may only persist at low levels. The high recruitment of crested wheatgrass suggests that coplanting of some native bunchgrasses may be a viable way of avoiding crested wheatgrass monocultures when this species is necessary for rehabilitation or restoration.

Natural selection favors root investment in native grasses during restoration of invaded fields

Competitive interactions with invasive species can cause evolutionary changes in native populations. In the Great Basin, USA, the invasive annual grass Bromus tectorum pre-emptively accesses limiting soil resources through extensive root production. Here, we determined whether evolutionary changes in root traits occurred during re-seeding of Elymus elymoides, a native bunchgrass. We grew plants from seeds collected from survivors in two restoration sites and compared them with plants grown from the original seed, with and without competition from B. tectorum. We quantified competition, root allocation, and morphology over time. Relative to the original seed source, E. elymoides plants that established in restoration sites showed small but significant increases in root biomass fraction, and plants were smaller, both aboveground and belowground, with parallel results seen in both field sites. In contrast, E. elymoides seed sources were not significantly different in competitive response to or in competitive effect on B. tectorum. Thus, results are consistent with natural selection favoring traits that allowed plants to establish in B. tectorum invaded areas but not traits that increased competitive ability. Selecting or breeding restoration plants for smaller size but increased relative root investment could improve establishment under reduced soil resource conditions and hence native plant restoration.

Seedling root traits strongly influence field survival and performance of a common bunchgrass

Seedling survival is a limiting factor in arid-land restoration. We investigated how variation in the root traits of glasshouse-reared seedlings related to the field performance of different genotypes from two populations of Elymus elymoides (squirreltail), a common bunchgrass native to the Western United States. Seeds from 100 E. elymoides individuals were collected from two sites in northern Nevada. We planted offspring of these 100 individuals in the glasshouse to characterize 10-day root traits of each maternal family. Root traits of glasshouse-reared plants and seed size measures were correlated with the performance of siblings grown in field plots close to the seed collection sites. Seedling root traits were related to performance of siblings at both sites. We estimate that within-population variation in root traits was associated with a more than six- to nine-fold increase in seedling survival probability and a two-fold increase in height at the less productive site, and a two-fold increase in survival and 1.2-fold increase in size at the more productive site. At both sites, effects of root traits were complex, with extreme values of some traits favoured and intermediate values of other traits favoured. There is a recognized need to integrate understanding of plant functional traits into larger conceptual frameworks of ecological restoration. Here we show that within-population variation in a suite of root functional traits relates to large variation in seedling survival and size of an arid-land grass species, improving our understanding of how trait variation affects performance in the field. Understanding such variation may be used to positively impact restoration outcomes.

The fear of unseen predators: ground squirrel tail flagging in the absence of snakes signals vigilance

Rather than fleeing from predators, many animals exhibit conspicuous displays that are thought to deter attacks by advertising quality (physical capabilities), or awareness of the predator. However, prey commonly exhibit the same antipredator display when predators are absent. Signaling in this context could either dishonestly indicate predator detection, or honestly indicate vigilance, but no study has tested these hypotheses experimentally. We tested the vigilance advertisement function of tail flagging, a signal given by California ground squirrels (Otospermophilus beecheyi) to rattlesnakes (Crotalus oreganus), by simulating snake strikes and quantifying strike avoidance behaviors in signaling and non-signaling squirrels. We removed the effect of snake presence on signaling by simulating strikes 1) in the presence of a rattlesnake, 2) with no snake present, or 3) in an area where squirrels had recently encountered a snake that was no longer present. Squirrels tail flagged most when interacting with a snake. Squirrels also signaled in areas of recent snake encounters, and in this context, they responded faster to simulated strikes and were more likely to leap vertically or horizontally away from the attack. Tail flagging rate was also related to squirrels’ body displacement time and flee modality. Squirrels rarely signaled or exhibited evasive leaping with no snake present. These results indicate that displays in the absence of predators can honestly signal vigilance, or the readiness to avoid an attack, even if predators are undetected. Such antipredator displays may be widespread as many species could benefit from advertising alertness when uncertain of the location of predators.

Impact of Cultivation Legacies on Rehabilitation Seedings and Native Species Re-Establishment in Great Basin Shrublands

Little is known about how cultivation legacies affect the outcome of rehabilitation seedings in the Great Basin, even though both frequently co-occur on the same lands. Similarly, there is little known about how these legacies affect native species re-establishment into these seedings. We examined these legacy effects by comparing areas historically cultivated and seeded to adjacent areas that were seeded but never cultivated, for density of seeded crested wheatgrass (Agropyron cristatum [L.] Gaertn.) and native perennial grasses, vegetation cover, and ground cover. At half of the sites, historically cultivated areas had lower crested wheatgrass density (P < 0.05), and only one site had a higher density of crested wheatgrass (P < 0.05). Likewise, the native shrub Wyoming big sagebrush (Artemisia tridentata Nutt. subsp. wyomingensis Beetle & Young) had lower cover (P < 0.05) in historically cultivated areas at half the sites. Sandberg bluegrass (Poa secunda J. Presl.) density was consistently lower in historically cultivated areas relative to those seeded-only. At sites where black greasewood (Sarcobatus vermiculatus [Hook.] Torr.) and bottlebrush squirreltail (Elymus elymoides [Raf.] Swezey) were encountered, there was either no difference or a higher density and cover within historically cultivated areas (P < 0.05). Likewise, cover of exotic forbs, especially halogeton (Halogeton glomeratus [M. Bieb.] C. A. Mey.), was either not different or higher in historically cultivated areas (P < 0.05). Bare ground was greater in historically cultivated areas at three sites (P < 0.05). These results suggest that cultivation legacies can affect seeding success and re-establishment of native vegetation, and therefore should not be overlooked when selecting research sites or planning land treatments that include seeding and or management to achieve greater native species diversity.

Notice of Release of Antelope Creek and Pleasant Valley Germplasms of Bottlebrush Squirreltail

Two new natural-track pre-variety germplasms (selected class), Antelope Creek and Pleasant Valley bottlebrush squirreltail ( Elymus elymoides (Raf.) Swezey “ssp. C” [Poaceae]), were released by the USDA Agricultural Research Service and the Utah Agricultural Experiment Station in 2009 and 2010, respectively. Antelope Creek Germplasm is recommended for use in the western Blue Mountains of Oregon and the eastern Cascade slopes and foothills of Oregon and northeastern California. Pleasant Valley Germplasm is recommended for use in the eastern Blue Mountains of Oregon, Washington, and Idaho. Both germ -plasms are members of the unofficially designated bottlebrush squirreltail “subspecies C,” which possesses a distribution that is centered in central and eastern Oregon. Jones TA, Parsons MC, Larson SR, Mott IW. 2014. Notice of release of Antelope Creek and Pleasant Valley germplasms of bottlebrush squirreltail. Native Plants Journal 15(1):57–64.

A nomenclatural guide and simplified key to the squirreltail taxa

The squirreltails, Elymus elymoides (Raf.) Swezey (bottlebrush squirreltail) (Poaceae) and E. multisetus (J.G. Smith) Burtt Davy (big squirreltail), occupy rangelands in the arid and semiarid portions of the northwestern US. Bottlebrush squirreltail exhibits multiple subspecies, while big squirreltail does not. Commercially available plant materials of squirreltail have been keyed and are listed by their taxon. A dichotomous key is presented to identify the correct squirreltail taxon that is based on the number of glumes per node and the cleavage displayed by those glumes. This key was developed from a previous key published in 1963 that was based on a now-obsolete systematic treatment. The new key better reflects phylogenetic relationships, as it is based on Dewey’s genomic system of classification rather than on morphological characters. . A nomenclatural guide and simplified key to the squirreltail taxa. Native Plants Journal 15(1):51–55.

Community Structure Affects Annual Grass Weed Invasion During Restoration of a Shrub–Steppe Ecosystem

AbstractEcological restoration of shrub–steppe communities in the western United States is often hampered by invasion of exotic annual grasses during the process. An important question is how to create restored communities that can better resist reinvasion by these weeds. One hypothesis is that communities comprised of species that are functionally similar to the invader will best resist invasion, while an alternative hypothesis is that structurally more complex and diverse communities will result in more effective competitive exclusion. In this field experiment, we examined the effects of restored community structure on the invasion success of three annual grass weeds (downy brome, jointed goatgrass, and cereal rye). We created replicated community plots that varied in species composition, structural complexity and density, then seeded in annual grass weeds and measured their biomass and seed production the following year, and their cover after 1 and 3 yr. Annual grass weeds were not strongly suppressed by any of the restored communities, indicating that it was difficult for native species to completely capture available resources and exclude annual grass weeds in the first years after planting. Perennial grass monocultures, particularly of the early seral grass bottlebrush squirreltail, were the most highly invaded communities, while structurally complex and diverse mixtures of shrubs (big sagebrush, rubber rabbitbrush), perennial grasses (bluebunch wheatgrass and bottlebrush squirreltail) and forbs (Lewis flax, Utah sweetvetch, hairy golden aster, gooseberryleaf globemallow) were more resistant to invasion. These results suggest that restoration of sagebrush steppe communities resistant to annual grass invasion benefits from higher species diversity; significant reduction of weed propagule pressure prior to restoration may be required.

Indirect effects of an invasive annual grass on seed fates of two native perennial grass species

Invasive plants exhibit both direct and indirect negative effects on recruitment of natives following invasion. We examined indirect effects of the invader Bromus tectorum (cheatgrass) on seed fates of two native grass species, Elymus elymoides and Pseudoroegneria spicata, by removing B. tectorum and by adding inoculum of the shared seed pathogen Pyrenophora semeniperda in factorial experiments at xeric and mesic field sites. We also included a supplemental watering treatment to increase emergence and also the potential for pathogen escape. We recorded emergence and survival of native seedlings and also determined the fate of unemerged seeds. At the xeric site, Pyrenophora-caused mortality was high (34%), and effects of other pathogens and failed emergence of germinants were smaller. Cheatgrass removal negatively affected both emergence (35 vs. 25%) and spring survival (69 vs. 42%). Pyrenophora-caused seed mortality increased with inoculum augmentation for both species (22 vs. 47% overall), but emergence was negatively impacted only for P. spicata (20 vs. 34%). At the mesic site, Pyrenophora-caused mortality was low (6%). Cheatgrass removal doubled emergence (26 vs. 14%). Seed mortality increased significantly with inoculum augmentation for P. spicata (12 vs. 5%) but not E. elymoides, while emergence was not significantly affected in either species. A large fraction of seeds produced germinants that failed to emerge (37%), while another large fraction (35%) was killed by other pathogens. We conclude that facilitation by cheatgrass at the xeric site but interference at the mesic site was probably mediated through litter effects that could be ameliorative or suppressive. Apparent competition between cheatgrass and native grasses could occur through Pyrenophora, especially in a xeric environment, but effects were weak or absent at emergence. This was probably because Pyrenophora attacks the same slow-germinating fraction that is subject to pre-emergence mortality from other causes, including attack by other pathogens such as Fusarium.

Competition of Squirreltail with Cheatgrass at Three Nitrogen Levels

Squirreltail (Elymus elymoides [Raf] Swezey) can grow in cheatgrass (Bromus tectorum L.) stands, and has reduced the biomass production of cheatgrass in its immediate vicinity. A field experiment was conducted to determine: 1) if competition for nitrogen resources occurs between seedlings of squirreltail and seedlings of cheatgrass, under low, medium, and high nitrogen levels, and; 2) if competition for nitrogen resources in the seedling stage is a mechanism that allows squirreltail to establish in cheatgrass stands. Five accessions of squirreltail were each seeded with a single accession of cheatgrass in a cultivated field near Ephraim, Utah in the fall of 1995 and again in 1996. A completely randomized strip-plot design with 3 replications was used including the 5 joint seedings as well as pure seedings of each accession of both species. Three nitrogen levels representing high, medium (control), and low were applied. The study was replicated over 2 years in different areas of the same field. Harvests of above-ground biomass of squirreltail and cheatgrass within each treatment were conducted in July of 1996 and 1997. Using biomass production as a measure of efficient nitrogen use, cheatgrass competed for and used nitrogen resources more efficiently than squirreltail when nitrogen was not limiting. All squirreltail accessions were able to compete for and use nitrogen more efficiently than cheatgrass when there was low availability of nitrogen. Some accessions of squirreltail competed for nitrogen resources more efficiently than others both in the control and at the reduced nitrogen level. White Rocks and Sublette squirreltail accessions were the best competitors with cheatgrass at the low nitrogen level reducing the cheatgrasss biomass by as much as 75% and 67% respectively. An intermediate competitor with cheatgrass was the Gunnison accession. The Washakie and Pueblo accessions were poor competitors with cheatgrass at low nitrogen levels.

Despite spillover, a shared pathogen promotes native plant persistence in a cheatgrass‐invaded grassland

How pathogen spillover influences host community diversity and composition is poorly understood. Spillover occurs when transmission from a reservoir host species drives infection in another host species. In cheatgrass-invaded grasslands in the western United States, a fungal seed pathogen, black fingers of death (Pyrenophora semeniperda), spills over from exotic cheatgrass (Bromus tectorum) to native perennial bunchgrasses such as squirreltail (Elymus elymoides). Previous theoretical work based on this system predicts that pathogens that spill over can favor either host coexistence, the exclusion of either host species, or priority effects, depending on species-specific transmission rates and pathogen tolerance. Here, these model predictions were tested by parameterizing a population growth model with field data from Skull Valley, Utah, USA. The model suggests that, across the observed range of demographic variation, the pathogen is most likely to provide a net benefit to squirreltail and a net cost to cheatgrass, though both effects are relatively weak. Although cheatgrass (the reservoir host) is more tolerant, squirreltail is far less susceptible to infection, and its long-lived adult stage buffers population growth against seed losses to the pathogen. This work shows that, despite pathogen spillover, the shared pathogen promotes native grass persistence by reducing exotic grass competition. Counterintuitively, the reservoir host does not necessarily benefit from the presence of the pathogen, and may even suffer greater costs than the nonreservoir host. Understanding the consequences of shared pathogens for host communities requires weighing species differences in susceptibility, transmission, and tolerance using quantitative models.

Cultivation legacies alter soil nutrients and differentially affect plant species performance nearly a century after abandonment

Cultivation legacies affect native vegeta- tion in old fields of the Great Basin, USA for nearly a century after these fields are abandoned. We hypoth- esized that cultivation lowered soil nutrients and that this legacy would differentially impact plant perfor- mance of four representative Great Basin species. To test these hypotheses, we compared soil nutrients (C, N, P, K, Mg and Ca) between two formerly cultivated and adjacent noncultivated sites in two soil series. We then compared the plant growth and foliar nutrient content of an exotic grass (Bromus tectorum L.), two native grasses (Elymus elymoides (Raf.) Swezey and Achnatherum hymenoides (Roem. and Schult.) Barkworth), and a native forb (Sphaeralcea grossu- lariifolia (Hook. and Arn.) Rydb) grown in these soils in the greenhouse and in the field. Only one sampling site had reduced soil nutrients associated with culti- vation legacies, where most of the negative effects on plant performance were found. E. elymoides appeared to be less affected by cultivation legacies than did A. hymenoides, which had a reduced survivorship and 20 % less above-ground biomass in cultivated soils. No species, including B. tectorum, were favored by cultivation. Our findings suggest that cultivation legacies can affect plant performance of different species in different ways and that altered soil nutrients may interact with other abiotic and biotic cultivation legacies in complex ways.