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.     

Distribution of ecosystem C and N within contrasting vegetation types in a semiarid rangeland in the Great Basin,USA

Semiarid sagebrush ecosystems are being transformed by wildfire, rangeland improvement techniques, and exotic plant invasions, but the effects on ecosystem C and N dynamics are poorly understood. We compared ecosystem C and N pools to 1 m depth among historically grazed Wyoming big sagebrush, introduced perennial crested wheatgrass, and invasive annual cheatgrass communities, to examine whether the quantity and quality of plant inputs to soil differs among vegetation types. Natural abundance δ¹⁵N isotope ratios were used to examine differences in ecosystem N balance. Sagebrush-dominated sites had greater C and N storage in plant biomass compared to perennial or annual grass systems, but this was predominantly due to woody biomass accumulation. Plant C and N inputs to soil were greatest for cheatgrass compared to sagebrush and crested wheatgrass systems, largely because of slower root turnover in perennial plants. The organic matter quality of roots and leaf litter (as C:N ratios) was similar among vegetation types, but lignin:N ratios were greater for sagebrush than grasses. While cheatgrass invasion has been predicted to result in net C loss and ecosystem degradation, we observed that surface soil organic C and N pools were greater in cheatgrass and crested wheatgrass than sagebrush-dominated sites. Greater biomass turnover in cheatgrass and crested wheatgrass versus sagebrush stands may result in faster rates of soil C and N cycling, with redistribution of actively cycled N towards the soil surface. Plant biomass and surface soil δ¹⁵N ratios were enriched in cheatgrass and crested wheatgrass relative to sagebrush-dominated sites. Source pools of plant available N could become ¹⁵N enriched if faster soil N cycling rates lead to greater N trace gas losses. In the absence of wildfire, if cheatgrass invasion does lead to degradation of ecosystem function, this may be due to faster nutrient cycling and greater nutrient losses, rather than reduced organic matter inputs.     

Prefire grazing by cattle increases postfire resistance to exotic annual grass (Bromus tectorum) invasion and dominance for decades

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Global change effects on Bromus tectorum L. (Poaceae) at its high‐elevation range margin

Global change is likely to affect invasive species distribution, especially at range margins. In the eastern Sierra Nevada, California, USA, the invasive annual grass, Bromus tectorum, is patchily distributed and its impacts have been minimal compared with other areas of the Intermountain West. We used a series of in situ field manipulations to determine how B. tectorum might respond to changing climatic conditions and increased nitrogen deposition at the high‐elevation edge of its invaded range. Over 3 years, we used snow fences to simulate changes in snowpack, irrigation to simulate increased frequency and magnitude of springtime precipitation, and added nitrogen (N) at three levels (0, 5, and 10 g m^?2) to natural patches of B. tectorum growing under the two dominant shrubs, Artemisia tridentata and Purshia tridentata, and in intershrub spaces (INTR). We found that B. tectorum seedling density in April was lower following deeper snowpack possibly due to delayed emergence, yet there was no change in spikelet production or biomass accumulation at the time of harvest. Additional spring rain events increased B. tectorum biomass and spikelet production in INTR plots only. Plants were primarily limited by water in 2009, but colimited by N and water in 2011, possibly due to differences in antecedent moisture conditions at the time of treatments. The threshold at which N had an effect varied with magnitude of water additions. Frequency of rain events was more influential than magnitude in driving B. tectorum growth and fecundity responses. Our results suggest that predicted shifts from snow to rain could facilitate expansion of B. tectorum at high elevation depending on timing of rain events and level of N deposition. We found evidence for P‐limitation at this site and an increase in P‐availability with N additions, suggesting that stoichiometric relationships may also influence B. tectorum spread.     

A Race for Survival: Can Bromus tectorum Seeds Escape Pyrenophora semeniperda-caused Mortality by Germinating Quickly?

BACKGROUND AND AIMS: Pathogen-seed interactions may involve a race for seed resources, so that seeds that germinate more quickly, mobilizing reserves, will be more likely to escape seed death than slow-germinating seeds. This race-for-survival hypothesis was tested for the North American seed pathogen Pyrenophora semeniperda on seeds of the annual grass Bromus tectorum, an invasive plant in North America. In this species, the seed germination rate varies as a function of dormancy status; dormant seeds germinate slowly if at all, whereas non-dormant seeds germinate quickly. METHODS: Three experimental approaches were utilized: (a) artificial inoculations of mature seeds that varied in primary dormancy status and wounding treatment; (b) naturally inoculated undispersed seeds that varied in primary dormancy status; and (c) naturally inoculated seeds from the carry-over seed bank that varied in degree of secondary dormancy, habitat of origin and seed age. KEY RESULTS: In all three approaches, seeds that germinated slowly were usually killed by the pathogen, whereas seeds that germinated quickly frequently escaped. Pyrenophora semeniperda reduced B. tectorum seed banks. Populations in drier habitats sustained 50 times more seed mortality than a population in a mesic habitat. Older carry-over seeds experienced 30 % more mortality than younger seeds. CONCLUSIONS: Given the dramatic levels of seed death and the ability of this pathogen to reduce seed carry-over, it is intriguing to consider whether P. semeniperda could be used to control B. tectorum through direct reduction of its seed bank.     

Seedling Survival from Locally and Commercially Obtained Seeds on Two Semiarid Sites

Local populations of plants are likely to be better adapted to a site than populations from elsewhere. Thus, local seeds should yield higher survival in restoration attempts than commercial seed stocks. We compared seedling survival from locally and commercially obtained seeds of seven species, Pseudoroegneria spicata (bluebunch wheatgrass), Elymus elymoides (squirreltail), Pascopyrum smithii (western wheatgrass), Stipa hymenoides (Indian ricegrass), Stipa comata (needle‐and‐thread), Chrysothamnus nauseosus (rubber rabbitbrush) and Ephedra nevadensis (Mormon tea) over three years on two sites in Utah (Dugway and Tintic) that were dominated by the introduced annual Bromus tectorum (cheatgrass). At the Dugway site we included burned and unburned seedbed treatments. For all species at Dugway, seedling survival to the first summer was higher on burned plots where B. tectorum densities were greatly reduced. First‐year seedling survival was 20–30% for most species on the Dugway burned plots and at Tintic. At the drier Dugway site, only S. hymenoides and Ephedra had substantial third‐year survival. Elymus and Pascopyrum survived to the third year only at the moister Tintic site. Survival to the third year was less than 3% for all species except S. comata (6% survival), and densities were low (0.2–1.0 plants/m²). However, third‐year plants were well established and the grasses flowered. Pseudoroegneria and Ephedra at Dugway (on burned plots) and S. comata and Elymus at Tintic had higher first‐year survival or higher survival based on survival curves from local than from commercial seeds. However, final survival was never significantly higher, although such a trend was suggested. Seed dormancy traits could also provide advantages to local populations, and we observed differences in dormancy between local and commercial S. comata and S. hymenoides seeds that may be an example.     

Low temperature during infection limits Ustilago bullata (Ustilaginaceae, Ustilaginales) disease incidence on Bromus tectorum (Poaceae, Cyperales)

Ustilago bullata is frequently encountered on the exotic winter annual grass Bromus tectorum in western North America. To evaluate the biocontrol potential of this seedling-infecting pathogen, we examined the effect of temperature on the infection process. Teliospore germination rate increased linearly with temperature from 2.5 to 25°C, with significant among-population differences. It generally matched or exceeded host seed germination rate over the range 10-25°C, but lagged behind at lower temperatures. Inoculation trials demonstrated that the pathogen can achieve high disease incidence when temperatures during infection range 20-30°C. Disease incidence was drastically reduced at 2.5°C. Pathogen populations differed in their ability to infect at different temperatures, but none could infect in the cold. This may limit the use of this organism for biocontrol of B. tectorum to habitats with reliable autumn seedling emergence, because cold temperatures are likely to limit infection of later-emerging seedling cohorts.     

Streetlights positively affect the presence of an invasive grass species

Anthropogenic disturbances associated with urban ecosystems can create favorable conditions for populations of some invasive plant species. Light pollution is one of these disturbances, but how it affects the growth and establishment of invasive plant populations is unknown. Cheatgrass (Bromus tectorum) is a problematic invasive species where it has displaced native grassland communities in the United States, but to our knowledge, there have been no studies of the ecological factors that affect cheatgrass presence in urban ecosystems. We conducted field surveys in urban alleys in Denver, Colorado, to compare the presence of cheatgrass at sites with and without artificial light at night (hereafter artificial light) from streetlights. These streetlights are mounted on utility poles, which cause ground disturbance when installed in alleys; we were able to test the independent effect of poles on cheatgrass establishment because not all poles have streetlights on them. We found that cheatgrass was positively associated with the presence of streetlights and to a lesser extent poles. In addition to cheatgrass, we also found that other plants were positively associated with the presence of both poles and streetlights. Our results suggest that artificial light may benefit the occurrence of cheatgrass and other plant species in urban settings. While invasive populations of cheatgrass in wild habitats attract the most attention from managers, we suggest more consideration for this grass in urban environments where its growth and establishment benefit from anthropogenic changes.     

Invasive grass reduces aboveground carbon stocks in shrublands of the Western US

Understanding the terrestrial carbon budget, in particular the strength of the terrestrial carbon sink, is important in the context of global climate change. Considerable attention has been given to woody encroachment in the western US and the role it might play as a carbon sink; however, in many parts of the western US the reverse process is also occurring. The conversion of woody shrublands to annual grasslands involves the invasion of non-native cheatgrass ( Bromus tectorum ) which in turn leads to increased frequency and extent of fires. We compared carbon storage in adjacent plots of invasive grassland and native shrubland. We scaled-up the impact of this ecosystem shift using regional maps of the current invasion and of the risk of future invasion. The expansion of cheatgrass within the Great Basin has released an estimated 8±3 Tg C to the atmosphere, and will likely release another 50±20 Tg C in the coming decades. This ecosystem conversion has changed portions of the western US from a carbon sink to a source, making previous estimates of a western carbon sink almost certainly spurious. The growing importance of invasive species in driving land cover changes may substantially change future estimates of US terrestrial carbon storage.