Establishing perennial grasses from seed in postdisturbance Wyoming big sagebrush (Artemesia tridentata subsp. wyomingensis Welsh) communities is often unsuccessful, due in part to a lack of knowledge of the seedling ecology of perennial grasses. We examined the influence of planting timing on germination and seedling demography of bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] A. Love) in the northern Great Basin. In 2008 (year 1) and 2009 (year 2) we planted seeds monthly, September–December, in 1-m² plots (500 seeds per plot) using a randomized block design with five replications. Germination timing was indexed using seed bags placed adjacent to 1-m² plots and retrieved at 2-wk intervals in fall and 1-mo intervals in spring. Seedlings were marked in March–June of the year following planting; seedlings alive in July were considered initially established. Planting in September and October had up to 80% germination prior to winter, whereas December plantings germinated mainly in spring and at reduced rates (15–35%). Seeds planted in September and October emerged approximately a month earlier than November–December plantings. The percentage of germinated seeds that emerged was highest for September–October plantings but the percentage of emergent seeds surviving to the end of the first growing season was highest for later plantings. Final seedling density was lowest for November planting in year 1 and highest for September and October planting in year 2. Our data indicate that timing of and performance at critical stages of seedling development were affected by planting month. We suggest that it may be possible to use emerging technologies (e.g., seed coatings or germplasm manipulations) to produce variable chronologies of seedling development with single plantings and allow managers to exploit multiple temporal windows of opportunity for seedling establishment.

The microclimatic requirements for successful seedling establishment are much more restrictive than those required for adult plant survival. The purpose of the current study was to use hydrothermal germination models and a soil energy and water flux model to evaluate intra- and interannual variability in seedbed microclimate relative to potential germination response of six perennial grasses and cheatgrass. We used a 44-yr weather record to parameterize a seedbed microclimate model for estimation of hourly temperature and moisture at seeding depth for a sandy loam soil type at the Orchard Field Test Site in southwestern Ada County, Idaho. Hydrothermal germination response was measured in the laboratory for two seed lots of cheatgrass (Bromus tectorum L.), four seed lots of bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] Löve), three seed lots of bottlebrush squirreltail (Elymus elymoides [Raf] Swezey), and one seed lot each of Sandberg bluegrass (Poa secunda J. Presl.), big squirreltail (Elymus multisetus [J.G. Smith] M.E. Jones), thickspike wheatgrass (Elymus lanceolatus [Scribn. And J.G. Smith] Gould) and Idaho fescue (Festuca idahoensis Elmer). Germination response models were developed to estimate potential germination rate for 13 subpopulations of each seed lot for every hour of the 44-yr simulation. Seedbed microclimate was assessed seasonally and for each day, month, and year, and germination rate-sum estimates integrated for a numerical index of relative site favorability for germination for each time period. The rate-sum favorability index showed a consistent pattern among seed lots for different years, and provides a relatively sensitive indicator of annual and seasonal variability in seedbed microclimate. This index could be used with field data to define minimum weather thresholds for successful establishment of alternative plant materials, in conjunction with weather forecast models for making restoration and fire-rehabilitation management decisions in the fall season, for evaluation of potential climate-change impacts on plant community trajectories, and in optimization schemes for selecting among alternative restoration/rehabilitation management scenarios.

In sagebrush rangelands perennial bunchgrasses are typically seeded in fall and a high proportion of planted seeds germinate prior to winter onset but fail to emerge in spring. Our objectives were to evaluate freezing tolerance of germinated but nonemergent bluebunch wheatgrass seeds under laboratory conditions. We used data from a 2-yr pilot study to determine overwinter freezing temperature and duration for soils in southeastern Oregon. We then conducted two experiments to assess freezing tolerance. In experiment 1, bluebunch wheatgrass seeds were planted in control pots and compared to seeds planted at early, mid, or late postgermination stages. Pots from each treatment were placed in a grow room maintained at 12 h 40 min light/11 h 20 min dark photoperiod, with a constant temperature of 22°C for 30 d either immediately or following a 30-d freeze. In experiment 2, germinated bluebunch wheatgrass seeds were planted in pots that were left nonfrozen or were frozen for a specified duration prior to a 30-d period in the grow room. Emergence density and tillers · seedling⁻¹ were quantified for both experiments. The number of days per year for freezing soil conditions in the pilot study ranged yearly from 25 to 51; maximum duration of continuous freezing was 16.5 and 11.2 d. Freezing reduced or eliminated seedling emergence at all postgermination stages (P < 0.001) and tiller density was reduced by at least 50% (P < 0.001). Maximum reduction in seedling density (P < 0.001) was realized within 4 d of initiation of freezing and tillers · seedling⁻¹ were reduced 30–70% with > 6 d of freezing (P = 0.001). Our data indicate that freezing-associated mortality of germinated but nonemergent bluebunch wheatgrass seedlings can be extremely high and suggest that management practices to reduce prewinter germination of seeds could improve subsequent emergence and seeding success.

The effect of plant and soil disturbances on seed density, species richness, and seed longevity of the soil seed bank was quantified for a semi-arid rangeland, over a 5-yr period (2002/2003–2006/2007 growing seasons). The different soil and plant treatments included fire, tillage (intended as a trampling surrogate), and blocked seed rain (simulating heavy grazing). These three experimental factors were combined in a factorial arrangement. Seed responses were evaluated in the soil seed bank before the new seed set, after the first seed production event, and after the second seed production event. Before disturbance (physical impact on the plant and soil), soil seed bank was dominated by early successional species: conversely, aboveground vegetation was mainly dominated by perennial grasses. After only 4 yr of blocked seed rain, seedling emergence of Decreaser grass species ceased totally both in the field and seed bank, with lower effect on Increaser grass species. Emergence of both Decreaser and Increaser grass species decreased in the seed bank with tillage, whereas the opposite occurred in the field. By contrast, tillage increased the emergence of weeds in the seed bank. The decrease in emergence of Decreaser grass species in both seed bank and field was still evident 4 yr after the rangeland was burnt. The grass species Themeda triandra was the most sensitive to fire in terms of seedling emergence. Blocked seed rain treatment significantly decreased (P < 0.05) species richness. Regardless of treatments applied, there was poor similarity between aboveground vegetation and the associated seed bank. Differences in the soil seed bank are likely to reflect manifested properties rather than short-term changes. Several characteristics of seed banks (species composition, seed abundance, and longevity) must be considered in order to understand the dynamics of plant communities following disturbances.

Red brome (Bromus rubens), an exotic annual grass, can dominate soil seed banks and poses serious threats to mature native plant communities in the Mojave Desert by competing with native species and providing fine fuels that facilitate widespread wildfire. By exploring how seed bank density and composition in burned areas change over time since fire (TSF), we can improve our understanding of how the seed banks are affected by fire. Samples of the 5-cm-deep soil seed bank were collected from two microsites (under shrubs, in open interspaces) within paired burned and unburned areas on 12 fires ranging from 5–31 yr TSF. Seed bank samples were assayed using the emergence method and seed densities were compared among TSFs, burn status (burned, unburned), and microsites for the species that emerged. Red brome soil seed bank density was spatially variable and TSF rarely predicted abundance. Overall, undershrub seed densities did not differ between burned and unburned areas. However, at some fire sites, seed densities in interspaces were greater in burned than unburned areas. Although native seed densities were low overall, they did not appear to differ according to burn status. Studies have shown that red brome plant and seed bank densities can be greatly reduced immediately after fire. Management efforts that focus on this initial colonization window may be able to take advantage of diminished red brome seed densities to limit its reestablishment while facilitating the establishment of native species. However, this window is brief, as our findings indicate that once reestablished, red brome soil seed densities in burned areas can be similar to those in unburned areas within 5 yr.

The importance of sexual reproduction in tussock grasses that regenerate through vegetative growth is unclear. Festuca gracillima Hook. f. was studied as a model because it is a perennial tussock-forming grass that produces abundant seed but rarely regenerates through seedlings. The Study area was the Magellanic Steppe, Patagonia, Argentina (182 mm rainfall), managed with sheep-grazing regimes of 0.65 (high), 0.21 (low), and 0 (exclosure) ewe equivalents · ha⁻¹ · yr⁻¹. Tussock size and spikelet production of 358 individuals were recorded over 5 yr. Yearly models of reproductive effort in relation to plant size were tested using a maximum likelihood procedure. Seed was collected and soil cores were tested for germination and viability. Survival and growth of cohorts of seedlings sown in nylon bags were recorded. Eighteen experimental plots were cleared, and seed establishment under protected and grazed conditions was registered. Reproductive effort varied with years and plant size, with a mean of 2.41%. Florets were produced at mean density of 544 ± 217 · m⁻². Predispersal losses reduced viable seed production to 187 ± 48 seeds · m⁻². Seed weighed 2–2.5 mg, with 65–95% germination. Postdispersal losses reduced the seed bank in spring to 33 ± 1.3 seeds · m⁻². Seedling survival curves were negatively exponential, with 95% mortality in the first year. Up to 5% of resources were used for sexual reproduction in favorable years and a recruitment of 1–3 new seedlings · m⁻² · yr⁻¹ was expected. These new plants were not observed in undisturbed plots, but established naturally in cleared plots and reached a density of 1 plant · m⁻² after 10 yr, together with 44 plants · m⁻² of other species. Competition might block the final establishment in these grasslands. Grazing does not appear to interfere in any stage of seed reproduction. Seed production may not maintain population numbers but could enhance genetic variation in these clonal plant populations and enable dispersal and recolonization of disturbed areas.

Re-establishing native communities that resist exotic weed invasion and provide diverse habitat for wildlife are high priorities for restoration in sagebrush ecosystems. Native forbs are an important component of healthy rangelands in this system, but they are rarely included in seedings. Understanding competitive interactions between forb and grass seedlings is required to devise seeding strategies that can enhance establishment of diverse native species assemblages in degraded sagebrush communities. We conducted a greenhouse experiment to examine seedling biomass and relative growth rate of common native forb species when grown alone or in the presence of a native bunchgrass or an exotic annual grass. Forb species included bigseed biscuitroot (Lomatium macrocarpum [Nutt. ex Torr. & A. Gray] J.M. Coult. & Rose), sulphur-flower buckwheat (Eriogonum umbellatum Torr.), hoary aster (Machaeranthera canescens [Pursh] Gray), royal penstemon (Penstemon speciosus Douglas ex Lindl.), and Munro's globemallow (Sphaeralcea munroana [Douglas ex Lindl.] Spach ex Gray); and neighboring grass species included bottlebrush squirreltail (Elymus elymoides [Raf.] Swezey), Sandberg bluegrass (Poa secunda J. Presl); and cheatgrass (Bromus tectorum L.). Forbs and grasses were harvested after 6, 9, or 12 wk of growth for biomass determination and calculation of relative growth rates (RGR) of forbs. Neither bunchgrass reduced biomass of any forb. RGR was reduced for royal penstemon when grown with either native grass and for Munro's globemallow when grown with bottlebrush squirreltail. Although only assessed qualitatively, forbs with vertically oriented root morphologies exhibited no reduction in RGR when grown with native grasses, compared to forbs with dense lateral branching, similar to the root morphology of native grasses. Biomass of forbs was reduced by 50% to 91% and RGR by 37% to 80% when grown with cheatgrass. Understanding native forb interactions with native grasses and cheatgrass will aid land managers in selecting effective seed mixes and making better use of costly seed.

The choice of plant materials is an important component of revegetation following disturbance. To determine the utility and effectiveness of various perennial grass species for revegetation on varied landscapes, a meta analysis was used to evaluate the stand establishment and persistence of 18 perennial cool-season grass species in 34 field studies in the Intermountain and Great Plains regions of the United States under monoculture conditions. Combined across the 34 studies, stand establishment values ranged from 79% to 43% and stand persistence values ranged from 70% to 0%. Intermediate wheatgrass (Thinopyrum intermedium [Host] Barkworth & D. R. Dewey), tall wheatgrass (Thinopyrum ponticum [Podp.] Z.-W. Liu & R.-C. Wang), crested wheatgrass (Agropyron spp.), Siberian wheatgrass (Agropyron fragile [Roth] P. Candargy), and meadow brome (Bromus riparius Rehmann) possessed the highest stand establishment (≥ 69%). There were no significant differences among the 12 species with the largest stand persistence values. Basin wildrye (Leymus cinereus (Scribn. & Merr.) Á. Löve), Altai wildrye (Leymus angustus [Trin.] Pilg.), slender wheatgrass (Elymus trachycaulus [Link] Gould ex Shinners), squirreltail (Elymus spp.), and Indian ricegrass (Achnatherum hymenoides [Roem. & Schult.] Barkworth) possessed lower stand persistence (≤ 32%) than the majority of the other species, and Indian ricegrass (0%) possessed the lowest stand persistence of any of the species. Correlations between environmental conditions and stand establishment and persistence showed mean annual study precipitation to have the most consistent, although moderate effect (r = ∼0.40) for establishment and persistence. This relationship was shown by the relatively poor stand establishment and persistence of most species at sites receiving less than 310 mm of annual precipitation. These results will be a tool for land managers to make decisions concerning the importance of stand establishment, stand persistence, and annual precipitation for revegetation projects on disturbed sites.

Purposeful introductions of exotic species for rehabilitation efforts following wildfire are common on rangelands in the western United States, though ecological impacts of exotic species in novel environments are often poorly understood. One such introduced species, Kochia prostrata (L.) Schrad (forage kochia) has been seeded on over 200 000 ha throughout the Intermountain West to provide fuel breaks and forage, and to compete with invasive plants. Despite its potential benefits, K. prostrata has been reported to spread from some seeded areas, and no studies have addressed its potential interactions with native species. A systematic investigation is needed to increase understanding of the extent to which K. prostrata spreads from seeded areas, the environmental conditions under which it spreads, and its interactions with the associated plant communities. We sampled 28 K. prostrata postfire rehabilitation and greenstrip seedings in southwestern Idaho, which ranged from 3 to 24 yr since seeding. We analyzed cover of K. prostrata and the associated plant community in adjacent seeded and unseeded areas, and quantified extent of spread from seeded areas. Abundance of K. prostrata was negatively associated with that of most plant functional groups, including native species, but was positively associated with abundance of exotic annual forbs. Kochia prostrata spread to unseeded areas on 89% of sampled sites; distances of the farthest individual from the seeding boundary were greater than those previously reported, ranging from 0 to 710 m, with a mean distance of 208 m. Further, although the area covered by K. prostrata increased with time since seeding, we found no evidence that plant community composition affected spread of K. prostrata. Results contribute to current understanding of potential ecological implications of seeding K. prostrata and will enhance the ability of land managers to make scientifically based management decisions about its use.

In the southwestern United States, redberry (Juniperus pinchotii Sudw.) and ashe (Juniperus ashei Buchholz) juniper are two invasive species that dominate some rangelands. Goats will consume up to 30% of their diet in juniper, but it is unknown if sheep will accept juniper to the same extent. The objectives of this study were to determine if sheep can be conditioned to consume juniper and to compare intake among different breeds. Rambouillet (n = 10), Suffolk (n = 10), and Dorper-cross (n = 10) lambs were randomly placed in individual pens for 31 d. A basal diet of alfalfa pellets (2.5% body weight [BW]) and juniper were fed. Juniper was fed each morning from 0800 to 0830 hours. The basal diet was fed for the remainder of the day. Intake of each was measured daily. Following the first 17 d, the basal diet was reduced to 2% BW for 7 d and then reduced to 1.5% BW for the final 7 d. Serum aspartate transaminase (AST), gamma glutamyltransferase (GGT), blood urea nitrogen (BUN), creatinine, and bilirubin levels, and live body weight were measured to assess any adverse physiological effects from juniper consumption. In a second trial, lambs were again fed alfalfa (2.5% BW) and juniper. One half of the lambs were also fed a 36% protein supplement to determine if supplementation with protein sources that escape rumen degradation would improve juniper consumption. Lambs received alfalfa, juniper, and protein supplement for 22 d with intake of each recorded daily. Intake of juniper was similar (P > 0.05) among breeds of sheep. Lambs readily consumed juniper and increased (P < 0.05) intake of juniper as the amount of alfalfa fed was reduced. Weight change was also similar among treatments. Protein supplementation did not improve juniper consumption. We contend that sheep will consume a diet consisting of 24% juniper without experiencing any adverse effects.

We conducted this study to describe the intermediate-term periodicities in percentage juniper (Juniperus spp.) in goat diets and to develop optimal sampling schemes to estimate individual animal variation in juniper consumption. Fecal samples were collected from 12 multiparous female Angora goats on Monday and Thursday for a 24-mo period. Percentage juniper in the diet was determined using fecal near-infrared reflectance spectroscopy. Spectral analysis was used to determine the presence and length of cyclic variation in juniper consumption during growing and dormant season periods. Significant periodicities were found for 37% and 68% of the goats in the dormant and growing seasons, respectively. Cycle lengths varied from 9 d in the dormant season to 7 or 8 d in the growing season. The simple coefficient of determination between a two-sample moving average and the mean of all observations on individual goats was highest during a 3-mo period in the spring, which indicates that samples collected in the spring provided the best estimate of the yearlong percentage juniper in the diet. Monte Carlo simulations for 7-d cycles showed the root mean squared difference between estimated and population mean for two samples with 2 or 3 d between samples was only 1% greater than the root mean square difference for three or four samples collected every other day. The optimal sampling strategy for determining the dietary percentage of a species is to collect two samples separated by one-half of the cycle length.

The study investigated the effect of general and homogeneous tree cover on grassland composition on an extensive Mediterranean rangeland with sparse oak trees in central Spain. We analyzed this effect together with other significant factors identified in this type of rangeland: topography and plowing. Data were collected in the 1984 growing season and they form part of a historical database on the characteristics of vegetation and livestock behavior; these data refer to grasslands below and away from the tree crowns of 91 individual trees, located in different topographical positions and in areas that were last plowed at different times. We used multivariate analyses to identify the main compositional trends of variation in pasture communities. The results indicate that the herbaceous community below tree crowns was more similar to that of the lowland areas than to the nearby areas away from the tree. This result supports the idea of tree cover in semiarid rangelands as a factor attenuating the effects on pastures of environmental conditions typical of high and intermediate topographical positions—generally presenting low soil moisture and fertility. Coupled with this, we also found effects of some individual trees related with the way livestock uses them as shelter and resting places. Our results indicate that the role played by dispersed trees in the management of this type of rangeland should be analyzed at two complementary spatial scales: the overall effect of tree cover as a factor acting at landscape scale and the specific effect of some individual trees acting at a more detailed scale.

Shrub encroachment can be explained by the abandonment of extensive livestock farming and changes to land use, and it is a common problem in the Mediterranean mountain pastures of Europe, with direct effects on biodiversity and landscape quality. In this paper, the effects of livestock exclusion vs. grazing on the dynamics of shrub and herbaceous vegetation were analyzed in a Spanish natural park located in a dry Mediterranean mountain area over a 5-yr period. Twelve 10 × 10 m exclosures were set up in six representative pasture areas of the park (with two replicates per location). Each year, the shrub number, volume, and biomass were measured in April, and the herbage height, biomass, and quality were measured in April and December (which represent the start and end of the vegetative growth season). A sustained increase of the shrub population and individual biomass was observed throughout the study, which was reflected in total shrub biomass per ha. Growth was greater in nongrazed exclosures (2 563 kg dry matter [DM] · ha⁻¹ · yr⁻¹), but it also happened in the grazed control areas (1 173 kg DM · ha⁻¹ · yr⁻¹), with different patterns depending on the location and shrub species. Herbage biomass did not change when grazing was maintained, but it did increase in places where grazing was excluded (291 kg DM · ha⁻¹ · yr⁻¹), mostly as a consequence of the accumulation of dead material, with a concomitant reduction in herbage quality. It was concluded that at the current stocking rates and management regimes, grazing alone is not enough to prevent the intense dynamics of shrub encroachment, and further reductions in grazing pressure should be avoided.

Medusahead (Taeniatherum caput-medusae [L.] Nevski) is an exotic annual grass invading western rangelands. Successful revegetation of invaded-plant communities can be prohibitively expensive because it often requires iterative applications of integrated control and revegetation treatments. Prescribed burning has been used to control medusahead and prepare seedbeds for revegetation, but burning has been constrained by liability concerns and has produced widely varying results. Capitalizing on naturally occurring wildfires could reduce revegetation costs and alleviate liability concerns. Thus, our objective was to determine if early summer wildfires and fall drill seeding could be used as a treatment combination to decrease medusahead and increase perennial and native vegetation. Treatments were evaluated pretreatment and for 3 yr postfire at six sites and included 1) an early summer wildfire combined with a seeding treatment (burn and seed) and 2) a nontreated (no burn, no seed) control. Perennial grass density was 4.6- to 10.0-fold greater in the burn-and-seed treatment compared to the control in the first 3 yr posttreatment (P < 0.05). Exotic annual grass density and cover in the third year posttreatment were lower in the burn-and-seed treatment than in the control (P < 0.05). However, exotic annual grass density was still > 130 individuals · m⁻² in the burn-and-seed treatment. The density of exotic annual grass is of concern because over time medusahead may displace perennial grasses and annual forbs that increased with the burn-and-seed treatment. Though not directly tested in this study, we suggest that, based on other research, the burn-and-seed treatment may need to incorporate a preemergent herbicide application to further suppress medusahead and increase the establishment of seeded vegetation. However, it appears that early summer wildfires may provide an opportunity to reduce the cost of integrated programs to revegetate medusahead-invaded plant communities.