Invasive Annual Grass Research Articles

Repeated mowing to restore remnant native grasslands invaded by nonnative annual grasses: upsides and downsides above and below ground

California grasslands have been severely impacted by the invasion of nonnative annual grasses, which often limit restoration of this important ecosystem. In this study, we explored the use of mowing as a restoration tool for native perennial grasslands at the Santa Rosa Plateau Ecological Reserve in southern California. We sought to evaluate if, over time, mowing would reduce nonnative annual grass cover and benefit native species, especially the native bunchgrassStipa pulchra. We hypothesized that repeated mowing, carefully timed to target nonnative annual grasses prior to seed maturation, would reduce nonnative seed inputs into the soil and eventually lead to diminished abundance of these species. We monitored vegetation in mowed and unmowed plots for 4 years, and conducted a seed bank study after 5 years to better understand the cumulative effects of mowing on native and nonnative seed inputs. Consistent with our hypotheses, we found that mowing successfully reduced nonnative annual grass cover and benefitted some native species, includingS. pulchra. However, we also found that nonnative forb species showed progressive increases in mowed plots over time. We observed similar patterns of species composition in the soil seed bank. Together, these results suggest that mowing can be used to control nonnative annual grasses and increase the abundance of native bunchgrasses, but that this method may also have the unintended consequence of increasing nonnative forb species.

Effects of Changing Climate on the Hydrological Cycle in Cold Desert Ecosystems of the Great Basin and Columbia Plateau

Climate change is already resulting in changes in cold desert ecosystems, lending urgency to the need to understand climate change effects and develop effective adaptation strategies. In this review, we synthesize information on changes in climate and hydrologic processes during the past century for the Great Basin and Columbia Plateau and discuss future projections for the 21st century. We develop midcentury projections of temperature and climate for the Great Basin and Columbia Plateau at timescales relevant to managers (2020−2050) and discuss concepts and strategies for adapting to the projected changes. For the instrumented record in the Great Basin and Columbia Plateau (1985−2011), a temperature increase of 0.7−1.4°C has been documented, but changes in precipitation have been relatively minor with no clear trends. Climate projections for 2020−2050 indicate that temperatures will continue to increase, especially in winter and during the night. Precipitation is more difficult to project, and estimates range from an 11% decrease to 25% increase depending on location. Recent records indicate that the Great Basin and Columbia Plateau are becoming more arid, a trend that is projected to continue. Droughts are likely to become more frequent and last longer, invasive annual grasses are likely to continue to expand, and the duration and severity of wildfire seasons are likely to increase. Climate projections can help in developing adaptive management strategies for actual or expected changes in climate. Strategies include reducing the risks of nonnative invasive plant spread and wildfires that result in undesirable transitions, planning for drought, and where necessary, facilitating the transition of populations, communities, and ecosystems to new climatic conditions. A proactive approach to planning for and adapting to climate change is needed, and publicly available Internet-based resources on climate data and planning strategies are available to help meet that need.

A proposed mechanism for high pathogen-caused mortality in the seed bank of an invasive annual grass

Abstract Pyrenophora semeniperda can infect nondormant Bromus tectorum seeds under optimal germination conditions, but most escape mortality. This reduces pathogen fitness relative to infection of dormant seeds, which are almost always killed. However, field experiments showed that a large proportion of seeds killed following inoculum addition were not accounted for as dormant seeds, but instead were likely nondormant seeds that would have germinated without inoculum addition. We hypothesized that widely fluctuating water availability to seeds would favor pathogenesis by delaying germination and allowing disease progression at water potentials below those that permit radicle emergence. To test this, nondormant host seeds were inoculated, imbibed for 8 or 24 h, subjected to controlled dehydration for 1–21 d, rehydrated, and scored for mortality. With dehydration at −4 MPa, mortality increased with dehydration duration after short or long imbibition. At −40 MPa, mortality increased with dehydration duration only after long imbibition. At −150 MPa, there was no effect of dehydration duration on generally low mortality. These results illustrate that fluctuating moisture can cause high nondormant seed mortality, explaining how this pathogen kills nondormant seeds.

Phytotoxic Activity of Metabolites Isolated from Rutstroemia sp.n., the Causal Agent of Bleach Blonde Syndrome on Cheatgrass (Bromus tectorum).

A fungal pathogen soon to be described as Rutstroemia capillus-albis (Rutstroemiaceae, Helotiales, Leotiomycetes) has been identified as the causal agent of ‘bleach blonde syndrome’ on the invasive annual grass weed Bromus tectorum (cheatgrass) in western North America. This apparently common but previously undescribed disease causes premature senescence and sterility, but does not affect seed germination or seedling emergence and growth. This study investigated whether the new species produces phytotoxins that could be implicated in pathogenesis. The compounds 9-O-methylfusarubin, 9-O-methylbostrycoidin, 5-O-methylnectriafurone, trans-methyl-p-coumarate and terpestacin were isolated from the solid culture of this fungus. The undescribed absolute stereochemistry at C-3 of 9-O-methylfusarubin and at C-1’ of 5-O-methylnectriafurone were assigned by applying electronic and vibrational circular dichroism (ECD and VCD) combined with computational methods and the advanced Mosher’s method, respectively. The first three listed compounds are naphtoquinone pigments, while terpestacin is a sesterterpene, and trans-methyl-p-coumarate could be the product of an unusual fungal phenylpropanoid biosynthesis pathway. In a juvenile plant immersion bioassay, both 9-O-methylfusarubin and terpestacin proved to be highly toxic at 10−4 M, causing wilting and plant death within 10 days. This finding suggests that these two compounds could play a role in pathogenesis on B. tectorum.

Spider community responses to grassland restoration: balancing trade‐offs between abundance and diversity

Spiders (Araneae) play key roles in ecosystems, not only as common and abundant generalist predators, but also as major contributors to biodiversity in many areas. In addition, due to their short generation times and high mobility, spiders respond rapidly to small changes in their environment, potentially making them useful indicators for restoration monitoring. However, few studies have focused on spider responses to grassland restoration in the United States. We compared degraded, native, and restored grassland sites to examine how spider communities and habitat respond to arid grassland restoration. We also examined how responses varied with the age of the restoration project. Spider communities in native sites differed from those in restored and degraded sites in several ways: native sites had fewer spiders and a different community composition than degraded and restored sites. However, native and restored sites had more species than degraded sites. Chronosequence data showed trends for lower abundance, higher species richness, and changing community composition as restoration projects mature. Several habitat variables were closely linked to variation in spider communities including cover of invasive annual grasses, litter, and biological soil crusts. Our data suggest that spider and vegetation responses to grassland restoration efforts can be successful in the long term—with resulting communities becoming more similar to native ones—and that spiders are useful indictors of grassland restoration. Our results also suggest that restoration may involve balancing trade‐offs between ecosystem services, with potential losses in predatory control offset by increases in biodiversity with restoration effort.

Viewpoint: An Alternative Management Paradigm for Plant Communities Affected by Invasive Annual Grass in the Intermountain West

On the Ground • Over 400,000 km2 of the Intermountain West is colonized by cheatgrass and other annual grasses. • Planning and management actions designed to foster perennial grass health throughout the region have never addressed how annual grasses would respond. • For decades, the most significant landscape-level management approach toward invasive annual grasses has been to complain. • We now know how to begin the process of taking the Intermountain West back from the domination of invasive annual grasses: through the management of standing dead litter. • Sustaining perennial bunchgrasses at landscape scales will require an integrated ecological approach to fuels management.

Niche opportunities for invasive annual plants in dryland ecosystems are controlled by disturbance, trophic interactions, and rainfall.

Resource availability and biotic interactions control opportunities for the establishment and expansion of invasive species. Studies on biotic resistance to plant invasions have typically focused on competition and occasionally on herbivory, while resource-oriented studies have focused on water or nutrient pulses. Through synthesizing these approaches, we identify conditions that create invasion opportunities. In a nested fully factorial experiment, we examined how chronic alterations in water availability and rodent density influenced the density of invasive species in both the Mojave Desert and the Great Basin Desert after fire. We used structural equation modeling to examine the direct and mediated effects controlling the density of invasives in both deserts. In the first 2 years after our controlled burn in the Great Basin, we observed that fire had a direct effect on increasing the invasive forb Halogeton glomeratus as well as a mediated effect through reducing rodent densities and herbivory. 4 years after the burn, the invasive annual grass Bromus tectorum was suppressing Halogeton glomeratus in mammal exclusion plots. There was a clear transition from years where invasives were controlled by disturbance and trophic interactions to years were resource availability and competition controlled invasive density. Similarly, in the Mojave Desert we observed a strong early influence of trophic processes on invasives, with Schismus arabicus benefitted by rodents and Bromus rubens negatively influenced by rodents. In the Mojave Desert, post-fire conditions became less important in controlling the abundance of invasives over time, while Bromus rubens was consistently benefitted by increases in fall rainfall.

Seeding plants for long-term multiple ecosystem service goals

The historical management of agroecological systems, such as California's rangelands, have received criticism for a singular focus on agricultural production goals, while society has shifting expectations to the supply of multiple ecosystem services from these working landscapes. The sustainability and the multiple benefits derived from these complex social-ecological systems is increasingly threatened by weed invasion, extreme disturbance, urban development, and the impacts of a rapidly changing and increasingly variable climate. California's grasslands, oak savannas, and oak woodlands are among the most invaded ecosystems in the world. Weed eradication efforts are rarely combined with seeding on these landscapes despite support for the inclusion of the practice in a weed management program. Depending on seed mix choice, cost and long-term uncertainty, especially for native seed, is an impediment to adoption by land managers. We investigated four seeding mixes (forage annual, native perennial, exotic perennial, and exotic-native perennial) to evaluate how these treatments resist reinvasion and support the delivery of simultaneous multiple ecosystem services (invasion resistance, native richness, nitrogen fixing plants, pollinator food sources, plant community diversity, forage quality, and productivity). We found the increase of exotic and native perennial cover will drive resistance to an invading weedy summer flowering forb Centaurea solstitialis but provides a mixed response to resisting invasive annual grasses. The resistance to invasion is coupled with little tradeoff in forage productivity and quality and gains in plant diversity and native cover.

Competition between cheatgrass and bluebunch wheatgrass is altered by temperature, resource availability, and atmospheric CO2 concentration

Global change drivers (elevated atmospheric CO2, rising surface temperatures, and changes in resource availability) have significant consequences for global plant communities. In the northern sagebrush steppe of North America, the invasive annual grass Bromus tectorum (cheatgrass) is expected to benefit from projected warmer and drier conditions, as well as increased CO2 and nutrient availability. In growth chambers, we addressed this expectation using two replacement series experiments designed to test competition between B. tectorum and the native perennial bunchgrass Pseudoroegneria spicata. In the first experiment, we tested the effects of elevated temperature, decreased water and increased nutrient availability, on competition between the two species. In the second, we tested the effects of elevated atmospheric CO2 and decreased water availability on the competitive dynamic. In both experiments, under all conditions, P. spicata suppressed B. tectorum, though, in experiment one, warmer and drier conditions and elevated nutrient availability increased B. tectorum’s competitiveness. In experiment two, when grown in monoculture, both species responded positively to elevated CO2. However, when grown in competition, elevated CO2 increased P. spicata’s suppressive effect, and the combination of dry soil conditions and elevated CO2 enhanced this effect. Our findings demonstrate that B. tectorum competitiveness with P. spicata responds differently to global change drivers; thus, future conditions are unlikely to facilitate B. tectorum invasion into established P. spicata communities of the northern sagebrush steppe. However, disturbance (e.g., fire) to these communities, and the associated increase in soil nutrients, elevates the risk of B. tectorum invasion.

Alternative pathways to landscape transformation: invasive grasses, burn severity and fire frequency in arid ecosystems

Summary Arid ecosystems are often vulnerable to transformation to invasive‐dominated states following fire, but data on persistence of these states are sparse. The grass/fire cycle is a feedback process between invasive annual grasses and fire frequency that often leads to the formation of alternative vegetation states dominated by the invasive grasses. However, other components of fire regimes, such as burn severity, also have the potential to produce long‐term vegetation transformations. Our goal was to evaluate the influence of both fire frequency and burn severity on the transformation of woody‐dominated communities to communities dominated by invasive grasses in major elevation zones of the Mojave Desert of western North America. We used a chronosequence design to collect data on herbaceous and woody cover at 229 unburned reference plots and 578 plots that burned between 1972 and 2010. We stratified the plots by elevation zone (low, mid, high), fire frequency (1–3 times) and years post‐fire (YPF; 1–5, 6–10, 11–20 and 21–40 YPF). Burn severity for each plot was estimated by the difference normalized burn ratio. We identified two broad post‐fire successional pathways. One was an outcome of fire frequency, resulting in a strong potential transformation via the grass/fire cycle. The second pathway was driven by burn severity, the critical aspect being that long‐term transformation of a community could occur from just one fire in areas that burned at high or sometimes moderate severity. Dominance by invasive grasses was most likely to occur in low‐and high‐elevation communities; cover of native herbaceous species was often greater than that of invasive grasses in the mid‐elevation zone. Synthesis. Invasive grasses can dominate a site that burned only one time in many decades at high severity, or a site that burned at low severity but multiple times in the same time period. However, high burn severity may predispose areas to more frequent fire because they have relatively high cover of invasive annual grass, suggesting burn severity and fire frequency have both independent and synergistic effects. Resilience in vegetation structure following fire in many arid communities may be limited to a narrow window of low burn severity in areas that have not burned in many decades.

A warmer and drier climate in the northern sagebrush biome does not promote cheatgrass invasion or change its response to fire

Dryland shrub communities have been degraded by a range of disturbances and now face additional stress from global climate change. The spring/summer growing season of the North American sagebrush biome is projected to become warmer and drier, which is expected to facilitate the expansion of the invasive annual grass Bromus tectorum (cheatgrass) and alter its response to fire in the northern extent of the biome. We tested these predictions with a factorial experiment with two levels of burning (spring burn and none) and three climate treatments (warming, warming + drying, and control) that was repeated over 3 years in a Montana sagebrush steppe. We expected the climate treatments to make B. tectorum more competitive with the native perennial grass community, especially Pseudoroegneria spicata, and alter its response to fire. Experimental warming and warming + drying reduced B. tectorum cover, biomass, and fecundity, but there was no response to fire except for fecundity, which increased; the native grass community was the most significant factor that affected B. tectorum metrics. The experimental climate treatments also negatively affected P. spicata, total native grass cover, and community biodiversity, while fire negatively affected total native grass cover, particularly when climate conditions were warmer and drier. Our short-term results indicate that without sufficient antecedent moisture and a significant disruption to the native perennial grass community, a change in climate to a warmer and drier spring/summer growing season in the northern sagebrush biome will not facilitate B. tectorum invasion or alter its response to fire.

Novel fine-scale aerial mapping approach quantifies grassland weed cover dynamics and response to management.

Invasive weeds threaten the biodiversity and forage productivity of grasslands worldwide. However, management of these weeds is constrained by the practical difficulty of detecting small-scale infestations across large landscapes and by limits in understanding of landscape-scale invasion dynamics, including mechanisms that enable patches to expand, contract, or remain stable. While high-end hyperspectral remote sensing systems can effectively map vegetation cover, these systems are currently too costly and limited in availability for most land managers. We demonstrate application of a more accessible and cost-effective remote sensing approach, based on simple aerial imagery, for quantifying weed cover dynamics over time. In California annual grasslands, the target communities of interest include invasive weedy grasses (Aegilops triuncialis and Elymus caput-medusae) and desirable forage grass species (primarily Avena spp. and Bromus spp.). Detecting invasion of annual grasses into an annual-dominated community is particularly challenging, but we were able to consistently characterize these two communities based on their phenological differences in peak growth and senescence using maximum likelihood supervised classification of imagery acquired twice per year (in mid- and end-of season). This approach permitted us to map weed-dominated cover at a 1-m scale (correctly detecting 93% of weed patches across the landscape) and to evaluate weed cover change over time. We found that weed cover was more pervasive and persistent in management units that had no significant grazing for several years than in those that were grazed, whereas forage cover was more abundant and stable in the grazed units. This application demonstrates the power of this method for assessing fine-scale vegetation transitions across heterogeneous landscapes. It thus provides means for small-scale early detection of invasive species and for testing fundamental questions about landscape dynamics.

Next‐generation restoration for sage‐grouse: a framework for visualizing local conifer cuts within a landscape context

AbstractThe expansion of coniferous trees into sagebrush ecosystems is a major driver of habitat loss and fragmentation, resulting in negative impacts to wildlife. Greater sage‐grouse (Centrocercus urophasianus) respond directly to conifer expansion through decreased breeding activity, nesting, and overall survival; thus, small amounts of conifer expansion can have significant impacts on sage‐grouse habitat and populations. To this end, conservation partners have collaborated across private and public lands to reduce the threat of conifer expansion through targeted removal of conifer trees. Here, we demonstrate the use of the Marxan framework to incorporate important ecosystem attributes in the prioritization of conifer removal within the Oregon range of sage‐grouse. We prioritized conifer removal relative to three separate goals: (1) enhancement of existing sage‐grouse breeding, nesting, and early brood‐rearing habitats; (2) facilitation of sage‐grouse movement between breeding and brood‐rearing habitats; and (3) improvement of connectivity among sage‐grouse priority areas for conservation (PACs). Optimization models successfully identified areas with low conifer canopy cover, high resilience and resistance to wildfire and annual grass invasion, and high bird abundance to enhance sage‐grouse habitat. The inclusion of mesic resources resulted in further prioritization of areas that were closer to such resources, but also identified potential pathways that connected breeding habitats to the late brood‐rearing habitats associated with mesic areas. Examining areas outside of PACs resulted in the selection of potential corridors to facilitate connectivity; although areas with low conifer cover were selected similarly to the other optimization models, areas with high cover were also chosen to be able to enhance connectivity. Areas identified by optimization models were largely consistent with and overlapped ongoing conifer removal efforts in the Warner Mountains of south‐central Oregon. Land ownership of preferential areas selected by models varied with priority goals and followed general ownership patterns of the region, with public lands managed by the Bureau of Land Management and private lands being selected the most. The increased availability of landscape‐level datasets and assessment tools in sagebrush ecosystems can reduce the time and cost of both planning and implementation of habitat projects involving conifer removal. Most importantly, incorporating these new datasets and tools can supplement expert‐based knowledge to maximize benefits to sagebrush and sage‐grouse conservation.

Effects of conifer treatments on soil nutrient availability and plant composition in sagebrush steppe

Piñon-juniper woodlands of the western United States have expanded 2 to 10-fold since the late 1800’s. Tree control measures using chainsaws, heavy equipment and prescribed fire have been used to reduce woodlands and restore big sagebrush steppe and decrease woody fuel loading. We evaluated nutrient availability and herbaceous recovery following various cutting and prescribed fire treatments in late succession western juniper woodlands on two sites in southeast Oregon from 2007 to 2012. Sites were a cool, wet big sagebrush-Idaho fescue association (FESCUE), highly resistant to exotic annual grasses and a warm dry big sagebrush-bluebunch wheatgrass association (BLUEBUNCH), moderately resistant to annual grass invasion. Treatments were untreated controls, partial cutting followed by fall broadcast burning (SEP), cut and leave (CUT), and cut and burn in winter (JAN) and spring (APR). Soil inorganic N (NO3−, NH4+), phosphorus (H2PO4−), potassium (K+), and cover of herbaceous species were measured in three zones; interspace, litter mats around the tree canopy (canopy), and beneath felled trees (debris). Following woodland cutting, the results of the various slash treatments measured significant differences through time in the availability of inorganic N, P, and K and vegetation composition. Peak nutrient availability tended to occur within the first two years after treatment. The increases in N, P, and K were greatest in severely burned debris and canopy zones of the SEP and APR treatments. Invasive annual grass cover was positively correlated to soil inorganic N concentrations. Herbaceous composition at the FESCUE site was generally resistant to annual grasses after juniper treatments and native plants dominating post-treatment even in highly impacted debris and canopy zones of the SEP treatment. The BLUEBUNCH site was less resistance and resilient, thus, exotic annual grasses were a major component of the understory especially when tree and slash burning was of high fire severity. To lessen these impacts requires slash burning be applied from late fall to early spring, when fuel moisture and relative humidity are higher, to maintain an adequate perennial herbaceous composition for recovery.

Livestock grazing supports native plants and songbirds in a California annual grassland.

Over eight years we measured the effects of plant community composition, vegetation structure, and livestock grazing on occurrence of three grassland bird species—Western Meadowlark (Sturnella neglecta), Horned Lark (Eremophila alpestris), and Grasshopper Sparrow (Ammodramus savannarum)—at sites in central California during breeding season. In California’s Mediterranean-type climatic region, coastal and inland grassland vegetation is dominated by exotic annual grasses with occasional patches of native bunchgrass and forbs. Livestock grazing, primarily with beef cattle, is the most widely used management tool. Compared with ungrazed plots, grazed plots had higher bare ground, native plant cover, and vertically heterogeneous vegetation. Grazed plots also had less plant litter and shorter vegetation. Higher native plant cover, which is predominantly composed of bunchgrasses in our study area, was associated with livestock grazing and north-facing aspects. Using an information theoretic approach, we found that all three bird species had positive associations with native plant abundance and neutral (Western Meadowlark, Grasshopper Sparrow) or positive (Horned Lark) association with livestock grazing. All species favored flatter areas. Horned Larks and Western Meadowlark occurred more often where there were patches of bare ground. Western Meadowlarks and Grasshopper Sparrows were most common on north-facing slopes, suggesting that these species may be at risk from projected climate change. These findings demonstrate that livestock grazing is compatible with or supports grassland bird conservation in Mediterranean-type grasslands, including areas with high levels of exotic annual grass invasion, in part because grazing supports the persistence of native plants and heterogeneity in vegetation structure. However, conservation of low-lying grasslands with high native species presence, and active management to increase the abundance of native plant species are also likely to be important for sustaining grassland birds long-term.

Population genetic structure of Bromus tectorum in the mountains of western North America.

Invasive species are often initially restricted to a narrow range and may then expand through any of multiple mechanisms including phenotypic plasticity, in situ evolution, or selection on traits preadapted for new habitats. Our study used population genetics to explore possible processes by which the highly selfing invasive annual grass Bromus tectorum has expanded into montane environments. We used 69 single nucleotide polymorphic (SNP) markers to genotype ca. 20 individuals from each of 38 montane cheatgrass populations from throughout the Intermountain West and to identify characteristic SNP haplotypes and examine their distribution. Five invariant SNP haplotypes were dominant in montane cheatgrass populations, making up 59% of genotyped individuals, with each haplotype present in 12 to 21 populations. Four of these were absent or present at low frequency in low elevation populations, while the fifth was also sometimes dominant at low elevation. Sixteen haplotypes made up 78% of all genotyped individuals. These haplotypes were distributed across several haplogroups within the clade that also includes most sagebrush steppe lineages. The wide geographic distribution of several common haplotypes almost completely restricted to montane habitats suggests that dominant lineages in montane populations may possess adaptive syndromes that are preserved through reduced outcrossing rates or negative selection on outcrossed progeny. However, conclusive evidence of such local adaptation requires reciprocal seeding experiments and further characterization of adaptive traits and breeding system characteristics. Other lineages have likely risen to dominance in montane populations through selectively neutral processes.

Post‐fire vegetation response at the woodland–shrubland interface is mediated by the pre‐fire community

AbstractUnderstanding the drivers of ecosystem responses to disturbance is essential for management aimed at maintaining or restoring ecosystem processes and services, especially where invasive species respond strongly to disturbance. In this study, we used repeat vegetation surveys from a network of prescribed fire treatments at the woodland–shrubland interface in the Shoshone Mountains, Nevada, USA, to investigate post‐fire responses of the understory plant community. First, we characterized post‐fire community trajectories for sites located along an elevational gradient, hypothesizing a positive relationship between elevation and resistance to invasion. Unburned understory communities and their responses to disturbance differed along the gradient. As hypothesized, lower elevations experienced greater species turnover and higher invasive annual grass (Bromus tectorum) cover after fire. Second, we characterized post‐fire plant community trajectories for sites with varying levels of pre‐fire pinyon and juniper cover, hypothesizing that pre‐fire tree dominance and associated reductions in perennial understory species lead to less predictable post‐fire trajectories. Among sites with low‐to‐moderate tree cover, burning largely eliminated differences in understory composition, suggesting that biotic legacies were sufficient to result in predictable trajectories. In contrast, sites with high pre‐fire tree cover transitioned into an annual forb‐dominated community with sparse vegetation cover, suggesting that the loss of the understory community initiated unpredictable and divergent post‐fire trajectories. Because plant communities were still changing four years after fire, it is unclear whether the alternate trajectories in sites with high tree cover will result in the formation of alternate states, or whether community composition will eventually converge with other sites at the same elevation. Results indicate that careful evaluation of site characteristics can be used to predict treatment outcomes at the woodland–shrubland interface, and to guide the appropriate use of prescribed fire or other management practices.

The seed ecology ofAegilops triuncialis: linking trait variation to growing conditions

AbstractAdaptive plastic responses in invasive species allow for establishment and persistence despite the lack of genetic matching to new environments. The capacity of annual species to invade habitats to which they are not adapted is likely to be predicated on post-invasion seed trait variation correlated with conditions in the new habitat. To test this, I compared variation in seed traits and germination patterns ofAegilops triuncialis, an invasive annual grass, from 69 sample populations from 24 sites in California (USA) across 13 years. Seed mass, germination fractions, seed viability, and strength of induced dormancy between dimorphic seed pairs were used to investigate plastic variation within and among populations, across two general soil types, and among two widespread maternal genotypes. I found that seed mass variation was constrained although both seed types showed a 3-fold range among populations, 20–25% variation within populations between years, and was positively correlated with longer growing seasons. However, induced seed dormancy in the small seed type was correlated with large seed mass and to late-season precipitation, suggesting that longer growing seasons influence the strength of maternal and sibling chemical signals that induce dormancy in the small seeds. Thus variation in small seed germination fractions varies with the growing conditions experienced by the maternal plant. The greater longevity of the small seed type suggests that plasticity in germination in this species may contribute to seed bank formation, which is atypical of invasive annual grasses in California, and this may contribute to the persistence and spread ofA. triuncialisin invaded ranges.

Understory Plant Community Responses to Fuel-Reduction Treatments and Seeding in an Upland Piñon-Juniper Woodland

Woody plant expansion and infilling into nonwooded rangeland ecosystems have been observed worldwide. Such expansion may lead to declines in herbaceous understory plant communities and increased fuel loads in rangelands. Under the US National Fire Plan, fuel-reduction treatments have been implemented over vast expanses of western forest types to reduce the risk of catastrophic wildfire and restore historical ecosystem structure, function, and diversity. The benefits of fuel-reduction may, however, also carry inherent ecological risk such as promoting non-native species colonization. Here, we compare understory plant community responses to three commonly used fuel-reduction treatments with seeding applications in an upland piñon (Pinus edulis Engelm.)–juniper (Juniperus osteosperma [Torr.] Little) woodland on the Colorado Plateau: 1) mechanical mastication, 2) lop and slash piled then burned (pile burn), and 3) lop and scatter followed by a broadcast burn (broadcast burn). Data were collected pretreatment (2009) and one (2010), two (2011), and six (2015) growing seasons post treatment. We found while understory perennial herbaceous plant cover remained low 1 and 2 yr post treatment, it increased by > 700% in all fuel-reduction treatment plots six growing seasons post treatment. Furthermore, while we observed minor increases in invasive annual grass, Bromus tectorum L. (cheatgrass), colonization in 2010 and 2011, there were substantial increases in B. tectorum cover by 2015. B. tectorum cover varied among treatments with the greatest cover in the unseeded mastication plot at nearly 30%. Seeding applications did not increase overall seed mix species cover but enhanced seed mix species richness and, thus, may have increased resistance to B. tectorum invasion in seeded treatment plots. Our findings offer valuable insights to the ecological consequences of fuel-reduction activities in piñon-juniper woodlands through comparison of common fuel-reduction treatments and seeding applications and highlight differences in understory plant community responses to treatments across short to longer time scales.

Development of remote sensing indicators for mapping episodic die-off of an invasive annual grass (Bromus tectorum) from the Landsat archive

The exotic annual grass Bromus tectorum (cheatgrass) dominates vast acreages of rangeland in the western USA, leading to increased fire frequency and ecosystem degradation that is often irreversible. Episodic regeneration failure (“die-off”) has been observed in cheatgrass monocultures and can have negative ecosystem consequences, but can also provide an opportunity for restoration of native species and ecological function within the imperiled sagebrush steppe ecosystem. Proximate causes of cheatgrass die-off are uncertain, although several taxa of fungal soil pathogens have been implicated. Die-off occurrence is stochastic and can occur in remote areas. Thus, developing remote sensing indicators that are repeatable over long time periods, across extensive regions, and at relatively fine spatial resolution would be beneficial for accurately pinpointing events.We developed a remote sensing approach for mapping historical die-off from the Landsat archive (1985–2015), and used this to quantify spatial and temporal patterns of die-off occurrence annually at 30-m resolution. A Random Forests classification of image-derived spectral endmembers, trained on 2014 data, predicted die-off with 93% accuracy (κ=0.845) when applied to independent validation data from 2009. Die-off extent varied greatly across years, though some areas experienced die-off multiple times during the observation period. We found a strong correspondence between die-off occurrence and winter drought conditions, with strongest negative associations with current-year winter precipitation and previous years’ lagged winter and annual precipitation. Die-off duration was heavily skewed towards single-year die-off (81.5% of events), even in localized areas of more frequent occurrence (‘die-off hotspots’).Our retrospective classification of the Landsat archive suggests that cheatgrass die-off occurrence is predictable as the intersection of particular site environmental conditions with annual weather conditions, possibly because they are favorable for disease expression. Associations with previous year conditions suggest that die-off may also depend upon feedbacks between cheatgrass dominance and litter production, suitable sites for seedling establishment, and abundance and persistence of pathogen inoculum. Remote sensing indicators such as those developed in this study are needed to develop hypotheses about die-off causal factors, as well as support improved management of cheatgrass, mitigation for negative consequences associated with die-off (e.g. wind erosion), and successful restoration of native plant communities on die-off areas.