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

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

Greater sage‐grouse respond positively to intensive post‐fire restoration treatments

Habitat loss is the most prevalent threat to biodiversity in North America. One of the most threatened landscapes in the United States is the sagebrush (Artemisia spp.) ecosystem, much of which has been fragmented or converted to non‐native grasslands via the cheatgrass‐fire cycle. Like many sagebrush obligates, greater sage‐grouse (Centrocercus urophasianus) depend upon sagebrush for food and cover and are affected by changes to this ecosystem. We investigated habitat selection by 28 male greater sage‐grouse during each of 3 years after a 113,000‐ha wildfire in a sagebrush steppe ecosystem in Idaho and Oregon. During the study period, seeding and herbicide treatments were applied for habitat restoration. We evaluated sage‐grouse responses to vegetation and post‐fire restoration treatments. Throughout the 3 years post‐fire, sage‐grouse avoided areas with high exotic annual grass cover but selected strongly for recovering sagebrush and moderately strongly for perennial grasses. By the third year post‐fire, they preferred high‐density sagebrush, especially in winter when sagebrush is the primary component of the sage‐grouse diet. Sage‐grouse preferred forb habitat immediately post‐fire, especially in summer, but this selection preference was less strong in later years. They also selected areas that were intensively treated with herbicide and seeded with sagebrush, grasses, and forbs, although these responses varied with time since treatment. Wildfire can have severe consequences for sagebrush‐obligate species due to loss of large sagebrush plants used for food and for protection from predators and thermal extremes. Our results show that management efforts, including herbicide application and seeding of plants, directed at controlling exotic annual grasses after a wildfire can positively affect habitat selection by sage‐grouse.     

Improving restoration success through microsite selection: an example with planting sagebrush seedlings after wildfire

Post‐fire restoration of foundation plant species, particularly non‐sprouting shrubs, is critically needed in arid and semi‐arid rangeland, but is hampered by low success. Expensive and labor‐intensive methods, including planting seedlings, can improve restoration success. Prioritizing where these more intensive methods are applied may improve restoration efficiency. Shrubs in arid and semi‐arid environments can create resource islands under their canopies that may remain after fire. Seedlings planted post‐fire in former canopy and between canopies (interspace) may have different survival and growth. We compared planting Wyoming big sagebrush (Artemisia tridentata Nutt. ssp. wyomingensis Beetle & Young) seedlings post‐fire in former sagebrush canopy and interspace microsites at five locations. Four growing seasons after planting, seedling survival was 46 and 7% in canopy and interspace microsites, respectively. Sagebrush cover was 5.8 times greater in canopy compared to interspace microsites. Sagebrush survival and cover were likely greater because of less competition from herbaceous vegetation as well as benefiting from resource island effects in canopy microsites. Initially, post‐fire abundance of exotic annual grasses was less in canopy microsites, but by the third year post‐fire it was substantially greater in canopy microsites, indicating that resource availability to seedlings was greater, at least initially, in canopy microsites. These results suggest microsites with greater likelihood of success should be identified and then utilized to improve restoration success and efficiency. This is important as the need for restoration greatly exceeds resources available for restoration.     

Restoration of Native Perennials in a California Annual Grassland after Prescribed Spring Burning and Solarization

Grasslands dominated by exotic annual grasses have replaced native perennial vegetation types in vast areas of California. Prescribed spring fires can cause a temporary replacement of exotic annual grasses by native and non‐native forbs, but generally do not lead to recovery of native perennials, especially where these have been entirely displaced for many years. Successful reintroduction of perennial species after fire depends on establishment in the postfire environment. We studied the effects of vegetation changes after an April fire on competition for soil moisture, a key factor in exotic annual grass dominance. As an alternative to fire, solarization effectively kills seeds of most plant species but with a high labor investment per area. We compared the burn to solarization in a study of establishment and growth of seeds and transplants of the native perennial grass Purple needlegrass (Nassella pulchra) and coastal sage species California sagebrush (Artemisia californica). After the fire, initial seed bank and seedling densities and regular percent cover and soil moisture (0–20 cm) data were collected in burned and unburned areas. Burned areas had 96% fewer viable seeds of the dominant annual grass, Ripgut brome (Bromus diandrus), leading to replacement by forbs from the seed bank, especially non‐native Black mustard (Brassica nigra). In the early growing season, B. diandrus dominating unburned areas consistently depleted soil moisture to a greater extent between rains than forbs in burned areas. However, B. diandrus senesced early, leaving more moisture available in unburned areas after late‐season rains. Nassella pulchra and A. californica established better on plots treated with fire and/or solarization than on untreated plots. We conclude that both spring burns and solarization can produce conditions where native perennials can establish in annual grasslands. However, the relative contribution of these treatments to restoration appears to depend on the native species being reintroduced, and the long‐term success of these initial restoration experiments remains to be determined.     

Attempting to restore mountain big sagebrush (Artemisia tridentata ssp. vaseyana) four years after fire

Restoration of shrubs is needed throughout the world because of altered fire regimes, anthropogenic disturbance, and overutilization. The native shrub mountain big sagebrush (Artemisia tridentata Nutt. ssp. vaseyana (Rydb.) Beetle) is a restoration priority because of its value to wildlife in western North America. One of the principal threats to mountain big sagebrush is encroachment by western juniper (Juniperus occidentalis ssp. occidentalis Hook) and other conifers. Fire is frequently applied to control juniper; however, sagebrush recovery after fire can be variable. Seeding sagebrush postfire can hasten sagebrush recovery; however, seeding is not always necessary. Therefore, it may be advantageous to monitor postfire recovery to determine if seeding is needed. The effect of seeding sagebrush several years after fire is unknown. We evaluated the efficiency of seeding mountain big sagebrush four years after fire‐controlled junipers at five sites. Sagebrush cover (<0.5%) and density (<0.07 plants/m²) was low in seeded plots and did not differ from unseeded controls in the three postseeding years. We conclude that seeding sagebrush four years after fire did not accelerate sagebrush recovery. We speculate that seeded sagebrush failed to establish because of competition from herbaceous vegetation that had four years to recover after fire. Although it would be beneficial to seed sagebrush only when needed, our results suggest postponing seeding until monitoring has determined that recovery is inadequate may not be advisable. We suggest researchers investigate methods to improve predicting sagebrush recovery to allow for seeding, when needed, before the first postfire growing season.     

California native and exotic perennial grasses differ in their response to soil nitrogen,exotic annual grass density,and order of emergence

Early emergence of plant seedlings can offer strong competitive advantages over later-germinating neighbors through the preemption of limiting resources. This phenomenon may have contributed to the persistent dominance of European annual grasses over native perennial grasses in California grasslands, since the former species typically germinate earlier in the growing season than the latter and grow rapidly after establishing. Recently, European perennial grasses have been spreading into both non-native annual and native perennial coastal grass stands in California. These exotic perennials appear to be less affected by the priority effects arising from earlier germination by European annual grasses. In addition, these species interactions in California grasslands may be mediated by increasing anthropogenic or natural soil nitrogen inputs. We conducted a greenhouse experiment to test the effects of order of emergence and annual grass seedling density on native and exotic perennial grass seedling performance across different levels of nitrogen availability. We manipulated the order of emergence and density of an exotic annual grass (Bromus diandrus) grown with either Nassella pulchra (native perennial grass), Festuca rubra (native perennial grass), or Holcus lanatus (exotic perennial grass), with and without added nitrogen. Earlier B. diandrus emergence and higher B. diandrus density resulted in greater reduction in the aboveground productivity of the perennial grasses. However, B. diandrus suppressed both native perennials to a greater extent than it did H. lanatus. Nitrogen addition had no effect on the productivity of native perennials, but greatly increased the growth of the exotic perennial H. lanatus, grown with B. diandrus. These results suggest that the order of emergence of exotic annual versus native perennial grass seedlings could play an important role in the continued dominance of exotic annual grasses in California. The expansion of the exotic perennial grass H. lanatus in coastal California may be linked to its higher tolerance of earlier-emerging annual grasses and its ability to access soil resources amidst high densities of annual grasses.     

Effectiveness of repeated autumn and spring fires for understorey restoration in weed‐invaded temperate eucalypt woodlands

Question. Can strategic burning, targeting differing ecological characteristics of native and exotic species, facilitate restoration of native understorey in weed‐invaded temperate grassy eucalypt woodlands? Location. Gippsland Plains, eastern Victoria, Australia. Methods. In a replicated, 5‐year experimental trial, the effects of repeated spring or autumn burning were evaluated for native and exotic plants in a representative, degraded Eucalyptus tereticornis grassy woodland. Treatments aimed to reduce seed banks and modify establishment conditions of exotic annual grasses, and to exhaust vegetative reserves of exotic perennial grasses. Treatments were applied to three grassland patch types, dominated by the native grass Austrodanthonia caespitosa, ubiquitous exotic annuals, or the common exotic perennial grass Paspalum dilatatum. Results. The dominant native grass Austrodanthonia caespitosa and native forbs were resilient to repeated fires, and target exotic annuals and perennials were suppressed differentially by autumn and spring fires. Exotic annuals were also suppressed by drought, reducing the overall treatment effects but indicating important opportunities for restoration. The initially sparse exotic geophyte Romulea rosea increased in cover with fire and the impact of this species on native forbs requires further investigation. There was minimal increase in diversity of subsidiary natives with fire, probably owing to lack of propagules. Conclusions. While fire is often considered to increase ecosystem invasibility, our study showed that strategic use of fire, informed by the relative responses of available native and exotic taxa, is potentially an effective step towards restoration of weed‐invaded temperate eucalypt woodlands.     

The Mark of Zorro: Effects of the Exotic Annual Grass Vulpia myuros on California Native Perennial Grasses

Native perennial grasses were once common in California prairies that are now dominated by annual grasses introduced from Europe. Competition from exotics may be a principal impediment to reestablishment of native perennial grasses. Introduced annual grasses, such as Vulpia myuros (zorro fescue), are often included with native perennial species in revegetation seed mixtures used in California. To examine the potential suppressive effect of this graminoid, we evaluated the growth and performance of a mixture of California native perennial grasses and resident weeds when grown with varying densities of V. myuros. The annual fescue exhibited a strongly plastic growth response to plant density, producing similar amounts of above‐ground biomass at all seeding densities. Perennial grass seedling survival and above‐ ground biomass decreased and individuals became thinner (i.e., reduced weight‐to‐height ratio) with increasing V. myuros seeding density. V. myuros also significantly suppressed above‐ground biomass and densities of weeds and had a more negative effect on weed densities than on native perennial grass densities. Biomass of native grasses and weeds was not differentially affected by increasing densities of V. myuros. Overall, because V. myuros significantly reduced the survival and performance of the mixture of native perennial grasses and this effect increased with increasing V. myuros density, we conclude that including this exotic annual in native seed mixtures is counterproductive to restoration efforts.     

Physical disturbance shapes vascular plant diversity more profoundly than fire in the sagebrush steppe of southeastern Idaho,U.S.A

Fire is thought to profoundly change the ecology of the sagebrush steppe. The Idaho National Laboratory provides an ideal setting to compare the effects of fire and physical disturbance on plant diversity in high‐native‐cover sagebrush steppe. Seventy‐eight 1‐hectare transects were established along paved, green‐striped, gravel, and two‐track roads, in overgrazed rangeland, and within sagebrush steppe involving different fire histories. Transects were sampled for the diversity and abundance of all vascular plants. Alpha, beta, and phylogenetic beta diversity were analyzed as a response to fire and physical disturbance. Postfire vegetation readily rebounds to prefire levels of alpha plant diversity. Physical disturbance, in contrast, strongly shapes patterns of alpha, beta, and especially phylogenetic beta diversity much more profoundly than fire disturbance. If fire is a concern in the sagebrush steppe then the degree of physical‐disturbance should be more so. This finding is probably not specific to the study area but applicable to the northern and eastern portions of the sagebrush biome, which is characterized by a pulse of spring moisture and cold mean minimum winter temperatures. The distinction of sagebrush steppe from Great Basin sagebrush should be revised especially with regard to reseeding efforts and the control of annual grasses.     

Restoring big sagebrush after controlling encroaching western juniper with fire: aspect and subspecies effects

The need for restoration of shrubs is increasingly recognized around the world. In the western United States, restoration of mountain big sagebrush (Artemisia tridentata Nutt. ssp. vaseyana [Rydb.] Beetle) after controlling encroaching conifers is a priority to improve sagebrush‐associated wildlife habitat. Conifers can be cost effectively removed with prescribed burning when sagebrush is codominant; however, burning removes sagebrush and natural recovery may be slow. We evaluated seeding mountain and Wyoming big sagebrush (A. tridentata Nutt. ssp. wyomingensis Beetle & Young) on north and south aspects after western juniper (Juniperus occidentalis ssp. occidentalis Hook) control with prescribed burning. We included seeding Wyoming big sagebrush, a more drought tolerant subspecies of big sagebrush, because it might grow better than mountain big sagebrush on hot, dry south slopes, during drought, or after juniper encroachment. Seeding mountain big sagebrush increased sagebrush cover and density compared to unseeded controls. In mountain big sagebrush‐seeded plots, sagebrush cover was 19 times greater on north compared to south aspects in the fourth year after seeding. At this time, sagebrush cover was also greater on mountain compared to Wyoming big sagebrush‐seeded plots. Natural recovery (i.e. unseeded) of sagebrush was occurring on north aspects with sagebrush cover averaging 3% 4 years after fire. Sagebrush was not detected on unseeded south aspects at the end of the study. These results suggest that postfire sagebrush recovery, with and without seeding, will be variable across the landscape based on topography. This study suggests seeding sagebrush after controlling junipers with burning may accelerate sagebrush recovery.     

Interannual climate variability mediates changes in carbon and nitrogen pools caused by annual grass invasion in a semiarid shrubland

Exotic plant invasions alter ecosystem properties and threaten ecosystem functions globally. Interannual climate variability (ICV) influences both plant community composition (PCC) and soil properties, and interactions between ICV and PCC may influence nitrogen (N) and carbon (C) pools. We asked how ICV and non-native annual grass invasion covary to influence soil and plant N and C in a semiarid shrubland undergoing widespread ecosystem transformation due to invasions and altered fire regimes. We sampled four progressive stages of annual grass invasion at 20 sites across a large (25,000 km²) landscape for plant community composition, plant tissue N and C, and soil total N and C in 2013 and 2016, which followed 2 years of dry and wet conditions, respectively. Multivariate analyses and ANOVAs showed that in invasion stages where native shrub and perennial grass and forb communities were replaced by annual grass-dominated communities, the ecosystem lost more soil N and C in wet years. Path analysis showed that high water availability led to higher herbaceous cover in all invasion stages. In stages with native shrubs and perennial grasses, higher perennial grass cover was associated with increased soil C and N, while in annual-dominated stages, higher annual grass cover was associated with losses of soil C and N. Also, soil total C and C:N ratios were more homogeneous in annual-dominated invasion stages as indicated by within-site standard deviations. Loss of native shrubs and perennial grasses and forbs coupled with annual grass invasion may lead to long-term declines in soil N and C and hamper restoration efforts. Restoration strategies that use innovative techniques and novel species to address increasing temperatures and ICV and emphasize maintaining plant community structure—shrubs, grasses, and forbs—will allow sagebrush ecosystems to maintain C sequestration, soil fertility, and soil heterogeneity.     

Post-pastoral changes in composition and guilds in a semi-arid conservation area, Central Otago, New Zealand

Changes in the vegetation of Flat Top Hill, a highly modified conservation area in semi;arid Central Otago, New Zealand, are described four years after the cessation of sheep and rabbit grazing. Unusually moist weather conditions coincide with the four-year period of change in response to the cessation of grazing. Between 1993 and 1997, the average richness and diversity (H') of species increased, and the average proportion of native species decreased significantly. The vegetation was significantly richer in exotic annual and perennial grass species, exotic perennial forbs, exotic woody species and native tussock grasses in 1997 than in 1993. Eight response guilds of species are identified. Most "remnant" native shrubs and forbs were stable, in that they remained restricted to local refugia and showed little change in local frequency. However, taller native grass species increased, some locally, and others over wide environmental ranges. Rare native annual forbs and several native perennial species from "induced" xeric communities decreased, and this may be a consequence of competition from exotic perennial grasses in the absence of grazing. The invasive exotic herb Sedum acre decreased in abundance between 1993 and 1997, but several other prominent exotic species increased substantially in range and local frequency over a wide range of sites. Exotic woody species, and dense, sward-forming grasses are identified as potential threats to native vegetation recovery.     

Potential for woody plant control by Spanish goats in the sagebrush steppe

Botanical composition of the diet of eight Spanish goats was studied on the Northern Great Basin Experimental Range during two stages of plant phenology (active growth in early July and cured forage in mid-August) to assess their potential for the control of sagebrush (Artemisia tridentata sspp. wyomingensis Nutt.) and western juniper (Juniperus occidentalis Hook.) and their nutritional status on sagebrush-steppe rangelands. Diets were quantified by documenting the number of visits to each species, the bites harvested and time expended grazing each forage over 4 consecutive days in each period. Forage chemical characteristics evaluated included: CP, NDF, ADF, ADL and IVDMD. Diet composition and forage quality indices varied significantly (P < 0.05) with changes in plant phenology. Vegetation cover averaged 52% in the pasture and consisted of 36% grasses, 8% forbs, 7% shrubs and 0.6% trees. Available herbage (excluding woody plants) was 534 kg ha⁻¹ during active growth trials and 572 kg ha⁻¹ when forages had cured. When forages were green goats acquired 28% of their total bites from grasses, 71% from forbs, 0.3% from shrubs and 0.9% from juniper trees. After forages had cured values were 35% from grasses, 56% from forbs, 0.1% from shrubs and 8.8% from juniper. The browsing of juniper (both foliage and bark), after herbaceous forages had cured, was the only substantive use of woody plants. Sagebrush was only lightly used (0.2% of total bites) when herbaceous forages were actively growing. Available herbage was of relatively high quality during both trials. When forages were actively growing, CP of grasses ranged from 8.9 to 5.6%, forbs from 17.7 to 8.5%, sagebrush scored at 8.5%, juniper foliage at 8.1% and juniper bark at 3.2%. After herbaceous forages had cured, grass CPs ranged between 6.2 and 3.1%, forbs from 10.4 to 4.4%, shrubs were not sampled and juniper foliage averaged 7.6%. Given the low levels of browsing exhibited by goats on sagebrush and juniper we see little opportunity for control of these woody plants when pastures provide a diverse (N = 25 species) array of readily available (534–572 kg ha⁻¹) and nutritious forages and pastures are lightly stocked (0.63 goats ha⁻¹ month⁻¹). We do need, however, to further explore their potential for control of these species under several other regimes. These include extended trials on more deteriorated rangelands, trials during seasons or conditions where forage is limited and the animal's selective opportunity is restricted and trials in high quality environments on newly established sagebrush and juniper seedlings.     

Fire Effects on Cover and Dietary Resources of Sage-Grouse Habitat

Abstract: We evaluated 6 years of vegetation response following prescribed fire in Wyoming big sagebrush (Artemisia tridentata spp. wyomingensis) steppe on vegetation cover, productivity, and nutritional quality of forbs preferred by greater sage-grouse (Centrocercus urophasianus), and abundance of common arthropod orders. Habitat cover (shrubs and tall herbaceous cover [>18 cm ht]) was about 50% lower after burning compared to unburned controls because of the loss of sagebrush. Perennial grasses and an invasive annual forb, pale alyssum (Alyssum alyssoides), increased in cover or yield after fire. There were no increases in yield or nutritional quality of forb species important in diets of sage-grouse. Abundance of ants (Hymenoptera), a significant component in the diet of young sage-grouse, decreased after fire. These results suggest that prescribed fire will not improve habitat characteristics for sage-grouse in Wyoming big sagebrush steppe where the community consists of shrubs, native grasses, and native forbs.     

A Spatial Model to Prioritize Sagebrush Landscapes in the Intermountain West (U.S.A.) for Restoration

The ecological integrity of Sagebrush ( Artemisia spp.) ecosystems in the Intermountain West (U.S.A.) has been diminished by synergistic relationships among human activities, spread of invasive plants, and altered disturbance regimes. An aggressive effort to restore Sagebrush habitats is necessary if we are to stabilize or improve current habitat trajectories and reverse declining population trends of dependent wildlife. Existing economic resources, technical impediments, and logistic difficulties limit our efforts to a fraction of the extensive area undergoing fragmentation, degradation, and loss. We prioritized landscapes for restoring Sagebrush habitats within the intermountain western region of the United States using geographic information system (GIS) modeling techniques to identify areas meeting a set of conditions based on (1) optimum abiotic and biotic conditions favorable for revegetation of Sagebrush; (2) potential to increase connectivity of Sagebrush habitats in the landscape to benefit wildlife; (3) location of population strongholds for Greater Sage-Grouse ( Centrocercus urophasianus , a species of conservation concern); and (4) potential impediments to successful restoration created by Cheatgrass ( Bromus tectorum , an invasive exotic annual grass). Approximately 5.8 million ha in southwestern Idaho, northern Nevada, and eastern Oregon met our criteria for restoring Wyoming big sagebrush ( Artemisia tridentata ssp. wyomingensis ) and 5.1 million ha had high priority for restoring Mountain big sagebrush ( A. tridentata ssp. vaseyana ). Our results represent an integral component in a hierarchical framework after which site-specific locations for treatments can be focused within high-priority areas. Using this approach, long-term restoration strategies can be implemented that combine local-scale treatments and objectives with large-scale ecological processes and priorities.     

Effects of exotic grass invasion on spatial heterogeneity in the ground-layer of grassy woodlands

We quantified the effects of exotic annual grass invasion on the ground-layer structure of grassy eucalypt woodlands, with the aim of determining if weed invasion decreased gap size and plant basal area leading to reduced spatial heterogeneity. We measured plant density, distance between plants and basal plant area in woodland sites which ranged from zero to 100% exotic plant cover in the ground-layer. The ground-layer in uninvaded woodlands was heterogeneous, with a large variation in basal plant area and distance between plants. Exotic annual grass density was positively correlated with total plant density, whereas native plant density was negatively correlated. Total plant basal area decreased as total plant density increased, with a lower total plant area in exotic dominated transects compared to native dominated. Variation in basal plant area decreased with increasing plant density. Exotic annual grasses were more closely spaced together (smaller gap size) and had a smaller basal area than the native grasses and rushes. There was also less variation in basal area and gap size with individual exotic annual grasses compared to the native grasses. Inter-plant distance was greater for both the native and exotic grasses when they had native grasses neighbouring them instead of exotic grasses. These findings show that woodlands invaded by exotic annual grasses have relatively less spatial heterogeneity in the ground-layer. These results have implications for other aspects of perennial grassy ecosystems invaded by annual grasses, including plant recruitment and restoration strategies.     

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

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

Exotic plants increase and native plants decrease with loss of foundation species in sagebrush steppe

Dominant plant species, or foundation species, are recognized to have a disproportionate control over resources in ecosystems, but few studies have evaluated their relationship to exotic invasions. Loss of foundation species could increase resource availability to the benefit of exotic plants, and could thereby facilitate invasion. The success of exotic plant invasions in sagebrush steppe was hypothesized to benefit from increased available soil water following removal of sagebrush (Artemisia tridentata), a foundation species. We examined the effects of sagebrush removal, with and without the extra soil water made available by exclusion of sagebrush, on abundance of exotic and native plants in the shrub steppe of southern Idaho, USA. We compared plant responses in three treatments: undisturbed sagebrush steppe; sagebrush removed; and sagebrush removed plus plots covered with “rainout” shelters that blocked winter-spring recharge of soil water. The third treatment allowed us to examine effects of sagebrush removal alone, without the associated increase in deep-soil water that is expected to accompany removal of sagebrush. Overall, exotic herbs (the grass Bromus tectorum and four forbs) were 3–4 times more abundant in shrub-removal and 2 times more abundant in shrub-removal + rainout-shelter treatments than in the control treatment, where sagebrush was undisturbed. Conversely, native forbs were only about half as abundant in shrub removal compared to control plots. These results indicate that removal of sagebrush facilitates invasion of exotic plants, and that increased soil water is one of the causes. Our findings suggest that sagebrush plays an important role in reducing invasions by exotic plants and maintaining native plant communities, in the cold desert we evaluated.     

Nitrogen limitation, 15N tracer retention, and growth response in intact and Bromus tectorum-invaded Artemisia tridentata ssp. wyomingensis communities

Annual grass invasion into shrub-dominated ecosystems is associated with changes in nutrient cycling that may alter nitrogen (N) limitation and retention. Carbon (C) applications that reduce plant-available N have been suggested to give native perennial vegetation a competitive advantage over exotic annual grasses, but plant community and N retention responses to C addition remain poorly understood in these ecosystems. The main objectives of this study were to (1) evaluate the degree of N limitation of plant biomass in intact versus B. tectorum-invaded sagebrush communities, (2) determine if plant N limitation patterns are reflected in the strength of tracer ¹⁵N retention over two growing seasons, and (3) assess if the strength of plant N limitation predicts the efficacy of carbon additions intended to reduce soil N availability and plant growth. Labile C additions reduced biomass of exotic annual species; however, growth of native A. tridentata shrubs also declined. Exotic annual and native perennial plant communities had divergent responses to added N, with B. tectorum displaying greater ability to use added N to rapidly increase aboveground biomass, and native perennials increasing their tissue N concentration but showing little growth response. Few differences in N pools between the annual and native communities were detected. In contrast to expectations, however, more ¹⁵N was retained over two growing seasons in the invaded annual grass than in the native shrub community. Our data suggest that N cycling in converted exotic annual grasslands of the northern Intermountain West, USA, may retain N more strongly than previously thought.