Importance of Soil and Plant Community Disturbance for Establishment of Bromus tectorum in the Intermountain West, USA

The annual grass Bromus tectorum has invaded millions of hectares in western North America and has transformed former perennial grass and shrub-dominated communities into annual grasslands. Fire plays a key role in the maintenance of B. tectorum on the landscape but the type of disturbance responsible for initial invasion is less well understood. We conducted an experiment in a perennial shrub/grass/forb community in eastern Idaho, USA to examine the roles of plant community and soil disturbance on B. tectorum emergence and establishment prior to state-changing fires. Our experiment consisted of a plant community disturbance treatment where we (1) removed the shrub component, (2) removed the grass/forb component, or (3) removed all shrubs, grasses, and forbs. We followed this treatment with seeding of B. tectorum onto the soil surface that was (1) intact, or (2) disturbed. Each experimental plot had an associated control with no plant community disturbance but was seeded in the same manner. The experiment was replicated 20 times in two sites (high and low aboveground biomass). We measured emergence by counting seedlings in late spring and establishment by counting, removing, and weighing B. tectorum individuals in mid-summer. We also examined the influence of plant community disturbance on the soil environment by measuring extractable NH4 + and NO3 – four times each summer. Soil disturbance greatly influenced the number of B. tectorum individuals that emerged each spring. Plant community disturbance, specifically disturbance of the grass/forb component, increased N availability in the late growing season and biomass of B. tectorum the following summer. We conclude that soil disturbance and plant community disturbance interact to promote the initial invasion of B. tectorum in Intermountain West valley ecosystems

Morphological and Physiological Traits Account for Similar Nitrate Uptake by Crested Wheatgrass and Cheatgrass

Millions of hectares throughout the Intermountain West are either dominated or threatened by the invasive annual grass Bromus tectorum (cheatgrass). This invasion is largely linked to disturbance and few regions appear immune. Disturbance liberates resources in a community and cheatgrass appears exceptionally able to capitalize on these resources. One species, however, is consistently competitive with cheatgrass. Agropyron cristatum (crested wheatgrass), an improved plant material developed from several populations in central Asia, is drought resistant, grazing tolerant, and largely excludes cheatgrass in stands established within the Great Basin. While previous studies document high resource uptake ability by crested wheatgrass, it remains unknown if high uptake in this species is due to morphological or physiological adaptation. We examined N uptake and tissue morphology of four grasses common in the Intermountain West, including cheatgrass and crested wheatgrass. We also included two native grasses, Pseudoroegneria spicata (bluebunch wheatgrass) and Elymus elymoides (bottlebrush squirreltail). We observed similar rates of N uptake by cheatgrass and crested wheatgrass and their uptake was greater than the native perennial species. A multivariate analysis suggests that, of the three perennial grasses examined here, crested wheatgrass is morphologically most similar to cheatgrass, but that morphology only accounts for 57 percent of the variation in N uptake capacity among species. Consequently, physiological traits such as induction of N uptake or N efflux likely play a role in the ability of crested wheatgrass to achieve N uptake rates similar to cheatgrass

Temperature and Functional Traits Influence Differences in Nitrogen Uptake Capacity Between Native and Invasive Grasses

Performance differences between native and exotic invasive plants are often considered static, but invasive grasses may achieve growth advantages in western North America shrublands and steppe under only optimal growing conditions. We examine differences in N uptake and several morphological variables that influence uptake at temperatures between 5 and 25 C. We contrast two native perennial grasses in western North America: Elymus elymoides and Pseudoroegneria spicata; two invasive annual grasses: Bromus tectorum and Taeniatherum caputmedusae; and one highly selected non-native perennial grass: Agropyron cristatum. The influence of temperature on N uptake is poorly characterized, yet these invasive annual grasses are known to germinate in warm soils in the autumn, and both experience cool soils during the short growing season following snowmelt in the spring. To further explore the influence of temperature on the correlation between morphological variables and N uptake, our data are applied to a previously published path model and one proposed here. Differences in N uptake between native and invasive grasses were small at the lowest temperature, but were large at the highest temperature. At lower temperatures, uptake of N by annuals and perennials was correlated with leaf N and mass. At higher temperatures, uptake by annuals was correlated only with these leaf traits, but uptake by perennials was correlated with these leaf traits as well as root N and mass. Consequently, our results imply that annual grasses face fewer morphological constraints on N uptake than perennial grasses, and annual grasses may gain further advantage in warmer temperature conditions or during more frequent warm periods

A New Perspective on Trait Differences Between Native and Invasive Exotic Plants

Functional differences between native and exotic species potentially constitute one factor responsible for plant invasion. Differences in trait values between native and exotic invasive species, however, should not be considered fixed and may depend on the context of the comparison. Furthermore, the magnitude of difference between native and exotic species necessary to trigger invasion is unknown. We propose a criterion that differences in trait values between a native and exotic invasive species must be greater than differences between co-occurring natives for this difference to be ecologically meaningful and a contributing factor to plant invasion. We used a meta-analysis to quantify the difference between native and exotic invasive species for various traits examined in previous studies and compared this value to differences among native species reported in the same studies. The effect size between native and exotic invasive species was similar to the effect size between co-occurring natives except for studies conducted in the field; in most instances, our criterion was not met although overall differences between native and exotic invasive species were slightly larger than differences between natives. Consequently, trait differences may be important in certain contexts, but other mechanisms of invasion are likely more important in most cases. We suggest that using trait values as predictors of invasion will be challenging

Invasion is Contingent on Species Assemblage and Invasive Species Identity inExperimental Rehabilitation Plots

Ecological studies often suggest that diverse communities are most resistant to invasion by exotic plants, but relatively few local species may be available to a rehabilitation practitioner. We examine the ability of monocultures and diverse assemblages to resist invasion by an exotic annual grass (cheatgrass) and an exotic biennial forb (dyer’s woad) in experimental rehabilitation plots. We constructed seven assemblages that included three monocultures of grass, forb, or shrub; three four-species mixtures of grasses, forbs, or shrubs; and a three-species mixture of one species from each growth form in an experimental field setting to test resistance to invasion. Assemblages were seeded with cheatgrass and dyer’s woad for two consecutive years and quantified as biomass and density of individuals from each exotic species. Soil NO3 and leaf-area index were examined as predictors of invasive plant abundance. Cheatgrass invasion was greatest in forb and shrub assemblages, and least in mixed grass or grass monoculture; dyer’s woad invasion was greatest into mixed grass or grass monoculture, but least into monoculture or mixedspecies assemblages composed of forbs or shrubs. The community composed of grasses, forbs, and shrubs suppressed invasion by both species. Consequently, assemblages were most resistant to invasion by species of the same growth form. Moreover, these monocultures and mixtures were generally similar in conferring resistance to invasion, but a monoculture of big sagebrush was more resistant than a mixture of shrubs. Soil NO3 was correlated with invasion by cheatgrass, whereas LAI was correlated with invasion by dyer’s woad, suggesting these species were more limited by belowground and aboveground resources, respectively. Overall, increasing diversity with limited species did not necessarily enhance resistance to invasion

The Missing Angle: Ecosystem Consequences of Phenological Mismatch

Climate change leads to unequal shifts in the phenology of interacting species, such as consumers and their resources, leading to potential phenological mismatches. While studies have investigated how phenological mismatch affects wild populations, we still lack studies and a framework for investigating how phenological mismatch affects ecosystems, particularly nutrient cycling

Associations of Near-Surface Soil Moisture and Annual Plant Community Dynamics

Invasive species have become an increasingly large concern, particularly in already degraded ecosystems, such as sagebrush (Artemisia tridentata)-steppe of the Intermountain West. Much of this ecosystem is already infested with large cheatgrass (Bromus tectorum) stands and is potentially at risk for future invasions depending on biotic and abiotic conditions. In these ecosystems, the existing vegetation, whether native or non-native, may not effectively utilize the soil moisture resources in the upper portion of the soil, termed the growth pool. If the existing vegetation does not effectively utilize moisture in the growth pool, an open resource is left for the establishment of other plants, including invasives. Through a combination of soil moisture modeling and observational studies, we identified three potential invasion pathways, particularly by annual plants, into a cheatgrass-dominated system, all consistent with the fluctuating resource hypothesis, and all resulting from an available water resource in the growth pool. Results suggest these arid and semi-arid systems are likely to be protected from novel invasive species by complete utilization of growth pool soil water resources by any existing vegetation, whether native or non-native. Our results also suggest the same features which make the site more prone to novel annual invaders may also be useful in guiding establishment of desired vegetation during restoration efforts

Interactions among climate, topography and herbivory control greenhouse gas (CO2, CH4 and N2O) fluxes in a subarctic coastal wetland

High-latitude ecosystems are experiencing the most rapid climate changes globally, and in many areas these changes are concurrent with shifts in patterns of herbivory. Individually, climate and herbivory are known to influence biosphere-atmosphere greenhouse gas (GHG) exchange; however, the interactive effects of climate and herbivory in driving GHG fluxes have been poorly quantified, especially in coastal systems that support large populations of migratory waterfowl. We investigated the magnitude and the climatic and physical controls of GHG exchange within the Yukon-Kuskokwim Delta in western Alaska across four distinct vegetation communities formed by herbivory and local microtopography. Net CO2 flux was greatest in the ungrazed Carex meadow community (3.97 ± 0.58 [SE] µmol CO2 m−2 s−1), but CH4 flux was greatest in the grazed community (14.00 ± 6.56 nmol CH4 m−2 s−1). The grazed community is also the only vegetation type where CH4 was a larger contributor than CO2 to overall GHG forcing. We found that vegetation community was an important predictor of CO2 and CH4 exchange, demonstrating that variation in regional gas exchange is best explained when the effect of grazing, determined by the difference between grazed and ungrazed communities, is included. Further, we identified an interaction between temperature and vegetation community, indicating that grazed regions could experience the greatest increases in CH4 emissions with warming. These results suggest that future GHG fluxes could be influenced by both climate and by changes in herbivore population dynamics that expand or contract the vegetation community most responsive to future temperature change

Early Goose Arrival Increases Soil Nitrogen Availability More Than an Advancing Spring in Coastal Western Alaska

An understudied aspect of climate change-induced phenological mismatch is its effect on ecosystem functioning, such as nitrogen (N) cycling. Migratory herbivore arrival time may alter N inputs and plant–herbivore feedbacks, whereas earlier springs are predicted to increase N cycling rates through warmer temperatures. However, the relative importance of these shifts in timing and how they interact to affect N cycling are largely unknown. We conducted a 3-year factorial experiment in coastal western Alaska that simulated different timings of Pacific black brant (Branta bernicla nigricans) arrival (3 weeks early, typical, 3 weeks late, or no-grazing) and the growing season (ca. 3 weeks advanced and ambient) on adsorbed and mobile inorganic (NH4+–N, NO3-–N) and mobile organic N (amino acid) pools. Early grazing increased NH4+–N, NO3-–N, and amino acids by 103%, 119%, and 7%, respectively, whereas late grazing reduced adsorbed NH4+–N and NO3−–N by 16% and 17%, respectively. In comparison, the advanced growing season increased mobile NH4+–N by 26%. The arrival time by geese and the start of the season did not interact to influence soil N availability. While the onset of spring in our system is advancing at twice the rate of migratory goose arrival, earlier goose migration is likely to be more significant than the advances in springs in influencing soil N, although both early goose arrival and advanced springs are likely to increase N availability in the future. This increase in soil N resources can have a lasting impact on plant community composition and productivity in this N-limited ecosystem

Cloud Cover and Delayed Herbivory Relative to Timing of Spring Onset Interact to Dampen Climate Change Impacts on Net Ecosystem Exchange in a Coastal Alaskan Wetland

Rapid warming in northern ecosystems over the past four decades has resulted in earlier spring, increased precipitation, and altered timing of plant–animal interactions, such as herbivory. Advanced spring phenology can lead to longer growing seasons and increased carbon (C) uptake. Greater precipitation coincides with greater cloud cover possibly suppressing photosynthesis. Timing of herbivory relative to spring phenology influences plant biomass. None of these changes are mutually exclusive and their interactions could lead to unexpected consequences for Arctic ecosystem function. We examined the influence of advanced spring phenology, cloud cover, and timing of grazing on C exchange in the Yukon–Kuskokwim Delta of western Alaska for three years. We combined advancement of the growing season using passive-warming open-top chambers (OTC) with controlled timing of goose grazing (early, typical, and late season) and removal of grazing. We also monitored natural variation in incident sunlight to examine the C exchange consequences of these interacting forcings. We monitored net ecosystem exchange of C (NEE) hourly using an autochamber system. Data were used to construct daily light curves for each experimental plot and sunlight data coupled with a clear-sky model was used to quantify daily and seasonal NEE over a range of incident sunlight conditions. Cloudy days resulted in the largest suppression of NEE, reducing C uptake by approximately 2 g C m−2 d−1 regardless of the timing of the season or timing of grazing. Delaying grazing enhanced C uptake by approximately 3 g C m−2 d−1. Advancing spring phenology reduced C uptake by approximately 1.5 g C m−2 d−1, but only when plots were directly warmed by the OTCs; spring advancement did not have a long-term influence on NEE. Consequently, the two strongest drivers of NEE, cloud cover and grazing, can have opposing effects and thus future growing season NEE will depend on the magnitude of change in timing of grazing and incident sunlight