Grass-Shrub Associations over a Precipitation Gradient and Their Implications for Restoration in the Great Basin, USA.
ABSTRACT: As environmental stress increases positive (facilitative) plant interactions often predominate. Plant-plant associations (or lack thereof) can indicate whether certain plant species favor particular types of microsites (e.g., shrub canopies or plant-free interspaces) and can provide valuable insights into whether "nurse plants" will contribute to seeding or planting success during ecological restoration. It can be difficult, however, to anticipate how relationships between nurse plants and plants used for restoration may change over large-ranging, regional stress gradients. We investigated associations between the shrub, Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis), and three common native grasses (Poa secunda, Elymus elymoides, and Pseudoroegneria spicata), representing short-, medium-, and deep-rooted growth forms, respectively, across an annual rainfall gradient (220-350 mm) in the Great Basin, USA. We hypothesized that positive shrub-grass relationships would become more frequent at lower rainfall levels, as indicated by greater cover of grasses in shrub canopies than vegetation-free interspaces. We sampled aerial cover, density, height, basal width, grazing status, and reproductive status of perennial grasses in canopies and interspaces of 25-33 sagebrush individuals at 32 sites along a rainfall gradient. We found that aerial cover of the shallow rooted grass, P. secunda, was higher in sagebrush canopy than interspace microsites at lower levels of rainfall. Cover and density of the medium-rooted grass, E. elymoides were higher in sagebrush canopies than interspaces at all but the highest rainfall levels. Neither annual rainfall nor sagebrush canopy microsite significantly affected P. spicata cover. E. elymoides and P. spicata plants were taller, narrower, and less likely to be grazed in shrub canopy microsites than interspaces. Our results suggest that exploring sagebrush canopy microsites for restoration of native perennial grasses might improve plant establishment, growth, or survival (or some combination thereof), particularly in drier areas. We suggest that land managers consider the nurse plant approach as a way to increase perennial grass abundance in the Great Basin. Controlled experimentation will provide further insights into the life stage-specific effectiveness and practicality of a nurse plant approach for ecological restoration in this region.
Project description:The mechanisms supporting positive ecological interactions are important. Foundation species can structure desert biodiversity by facilitating seedbanks of annual plants, but the direct and indirect mechanisms of shrub effects on seedbank have not been experimentally decoupled. We conducted the first test of shrubs increasing seedbank densities through direct effects on the seedbank (i.e. shrub seed-trapping, animal-mediated dispersal) and indirect effects by facilitating the annual plant community (i.e. seed deposition, annual seed-trapping). Two distinct desert ecosystems were used to contrast transient seedbank densities in shrub and open microsites by manipulating annual plant density and the presence of the persistent seedbank. We measured transient seedbank densities at the end of the growing season by collecting soil samples and extracting seeds from each respective treatment. Transient seedbank densities were greatest in shrub canopies and with relatively higher annual plant densities. The persistent seedbank contributed to transient seedbank densities only in one desert and in the open microsite. Shrubs indirectly increased seedbank densities by facilitation the seed production of the annual plants. Therefore, shrubs are increasing seedbank independently of the annual plant community, likely through trapping effects, and dependently by facilitating seed production of the annuals. These findings provide evidence for a previously undescribed mechanism that supports annual seedbanks and thus desert biodiversity. We also identify shrubs as being significant drivers of desert plant communities and emphasize the need to consider multiple mechanisms to improve our ability to predict the response of ecosystems to change.
Project description: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.
Project description:The formation of fertility islands by shrubs increases soil resources heterogeneity in thicketization-grasslands. Clonal plants, especially rhizomatous or stoloniferous clonal plants, can form large clonal networks and use heterogeneously distributed resources effectively. In addition, shrubs, especially spiny shrubs, may also provide herbaceous plants with protection from herbivores, acting as 'shelters'. The interaction between pre-dominated clonal herbaceous plants and encroaching shrubs remains unclear in thicketization-grassland under grazing pressure. We hypothesized that clonal herbaceous plants can be facilitated by encroached shrubs as a 'shelter from herbivores' and/or as an 'increased soil resources' under grazing pressure. To test this hypothesis, a total of 60 quadrats were chosen in a thicket-grassland in northern China that was previously dominated by Leymus chinensis and was encroached upon by the spiny leguminous plant Caragana intermedia. The soil and plant traits beneath and outside the shrub canopies were sampled, investigated and contrasted with an enclosure. The soil organic matter, soil total nitrogen and soil water content were significantly higher in the soil beneath the shrub canopies than in the soil outside the canopies. L. chinensis beneath the shrub canopies had significantly higher plant height, single shoot biomass, leaf length and width than outside the shrub canopies. There were no significantly differences between plant growth in enclosure and outside the shrub canopies. These results suggested that under grazing pressure in a grassland undergoing thicketization, the growth of the rhizomatous clonal herbaceous plant L. chinensis was facilitated by the spiny shrub C. intermedia as a 'shelter from herbivores' more than through 'increased soil resources'. We propose that future studies should focus on the community- and ecosystem-level impacts of plant clonality.
Project description:BACKGROUND:Carrizo Plain National Monument (San Joaquin Desert, California, USA) is home to many threatened and endangered species including the blunt-nosed leopard lizard (Gambelia sila). Vegetation is dominated by annual grasses, and shrubs such as Mormon tea (Ephedra californica), which is of relevance to our target species, the federally listed blunt-nosed leopard lizard, and likely also provides key ecosystem services. We used relatively nonintrusive camera traps, or trail cameras, to capture interactions between animals and these shrubs using a paired shrub-open deployment. Cameras were placed within the shrub understory and in open microhabitats at ground level to estimate animal activity and determine species presence. FINDINGS:Twenty cameras were deployed from April 1st, 2015 to July 5th, 2015 at paired shrub-open microsites at three locations. Over 425,000 pictures were taken during this time, of which 0.4 % detected mammals, birds, insects, and reptiles including the blunt-nosed leopard lizard. Trigger rate was very high on the medium sensitivity camera setting in this desert ecosystem, and rates did not differ between microsites. CONCLUSIONS:Camera traps are an effective, less invasive survey method for collecting data on the presence or absence of desert animals in shrub and open microhabitats. A more extensive array of cameras within an arid region would thus be an effective tool to estimate the presence of desert animals and potentially detect habitat use patterns.
Project description: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.
Project description:Herbivory by both grazing and browsing ungulates shapes the structure and functioning of terrestrial ecosystems worldwide, and both types of herbivory have been implicated in major ecosystem state changes. Despite the ecological consequences of differences in diets and feeding habits among herbivores, studies that experimentally distinguish effects of grazing from spatially co-occurring, but temporally segregated browsing are extremely rare. Here we use a set of long-term exclosures in northern Utah, USA, to determine how domestic grazers vs. wild ungulate herbivores (including browsers and mixed feeders) affect sagebrush-dominated plant communities that historically covered ~62 million ha in North America. We sampled plant community properties and found that after 22 years grazing and browsing elicited perceptible changes in overall plant community composition and distinct responses by individual plant species. In the woody layer of the plant community, release from winter and spring wild ungulate herbivory increased densities of larger Wyoming big sagebrush (Artemisia tridentata, ssp. wyomingensis) at the expense of small sagebrush, while disturbance associated with either cattle or wild ungulate activity alone was sufficient to increase bare ground and reduce cover of biological soil crusts. The perennial bunchgrass, bottlebrush squirretail (Elymus elymoides), responded positively to release from summer cattle grazing, and in turn appeared to competitively suppress another more grazing tolerant perennial grass, Sandberg's blue grass (Poa secunda). Grazing by domestic cattle also was associated with increased non-native species biomass. Together, these results illustrate that ungulate herbivory has not caused sagebrush plant communities to undergo dramatic state shifts; however clear, herbivore-driven shifts are evident. In a dry, perennial-dominated system where plant community changes can occur very slowly, our results provide insights into potential long-term trajectories of these plant communities under different large herbivore regimes. Our results can be used to guide long-term management strategies for sagebrush systems and improve habitat for endemic wildlife species such as sage-grouse (Centrocercus spp.).
Project description:Invasive species, local plant communities and invaded ecosystems change over space and time. Quantifying this change may lead to a better understanding of the ecology and the effective management of invasive species. We used data on density of the highly invasive shrub Lantana camara (lantana) for the period 1990-2008 from a 50 ha permanent plot in a seasonally dry tropical forest of Mudumalai in southern India. We used a cumulative link mixed-effects regression approach to model the transition of lantana from one qualitative density state to another as a function of biotic factors such as indicators of competition from local species (lantana itself, perennial grasses, invasive Chromolaena odorata, the native shrub Helicteres isora and basal area of native trees) and abiotic factors such as fire frequency, inter-annual variability of rainfall and relative soil moisture. The density of lantana increased substantially during the study period. Lantana density was negatively associated with the density of H. isora, positively associated with basal area of native trees, but not affected by the presence of grasses or other invasive species. In the absence of fire, lantana density increased with increasing rainfall. When fires occurred, transitions to higher densities occurred at low rainfall values. In drier regions, lantana changed from low to high density as rainfall increased while in wetter regions of the plot, lantana persisted in the dense category irrespective of rainfall. Lantana seems to effectively utilize resources distributed in space and time to its advantage, thus outcompeting local species and maintaining a population that is not yet self-limiting. High-risk areas and years could potentially be identified based on inferences from this study for facilitating management of lantana in tropical dry forests.
Project description:Woody vegetation has increased on rangelands worldwide for the past 100-200 years, often because of reduced fire frequency. However, there is a general aversion to reintroducing fire, and therefore, fire surrogates are often used in its place to reverse woody plant encroachment. Determining the conservation effectiveness of reintroducing fire compared with fire surrogates over different time scales is needed to improve conservation efforts. We evaluated the conservation effectiveness of reintroducing fire with a fire surrogate (cutting) applied over the last ~30 years to control juniper (Juniperus occidentalis Hook.) encroachment on 77 sagebrush-steppe sites. Critical to conservation of this imperiled ecosystem is to limit juniper, not encourage exotic annual grasses, and promote sagebrush dominance of the overstory. Reintroducing fire was more effective than cutting at reducing juniper abundance and extending the period of time that juniper was not dominating the plant community. Sagebrush was reduced more with burning than cutting. Sagebrush, however, was predicted to be a substantial component of the overstory longer in burned than cut areas because of more effective juniper control. Variation in exotic annual grass cover was explained by environmental variables and perennial grass abundance, but not treatment, with annual grasses being problematic on hotter and drier sites with less perennial grass. This suggests that ecological memory varies along an environmental gradient. Reintroducing fire was more effective than cutting at conserving sagebrush-steppe encroached by juniper over extended time frames; however, cutting was more effective for short-term conservation. This suggests fire and fire surrogates both have critical roles in conservation of imperiled ecosystems.
Project description:Shrub encroachment induced by global change and human disturbance strongly affects ecosystem structure and function. In this study, we explore the degree to which invading leguminous shrubs affected neighboring grasses, including via the transfer of fixed nitrogen (N). We measured N concentrations and natural abundance (15)N of shoot tissues from three dominant grasses from different plant functional groups across seven distances along a local transect (up to 500 cm) to the leguminous shrub, Caragana microphylla. C. microphylla did transfer fixed N to neighboring grasses, but the amount and distance of N transferred were strongly species-specific. Shoot N concentrations decreased significantly with distance from C. microphylla, for a rhizomatous grass, Leymus chinensis, and a bunchgrass, Achnatherum sibiricum. However, N concentrations of another bunchgrass, Stipa grandis, were higher only directly underneath the shrub canopy. Shoot ?(15)N values of L. chinensis were enriched up to 500 cm from the shrub, but for S. grandis were enriched only below the shrub canopy. In contrast, ?(15)N of A. sibiricum did not change along the 500-cm transect. Our results indicated the rhizomatous grass transferred fixed N over long distances while bunchgrasses did not. The presence of C. microphylla increased the shoot biomass of L. chinensis but decreased that of S. grandis and A. sibiricum. These findings highlight the potential role of nutrient-acquisition strategies of neighboring grasses in moderating the interspecific variation of fixed N transfer from the leguminous shrub. Overall, leguminous shrubs have either positive or negative effects on the neighboring grasses and dramatically affect plant community composition and structure.
Project description:The facilitation mechanism maintains ecosystem richness by increasing seedling recruitment. Overgrazed grasslands of northwestern Patagonia are invaded by shrubs that could promote the seedling recruitment of forage species. We investigated the role of Acaena splendens shrubs on the maintenance of diversity and its usefulness as a nurse shrub in the recruitment of Festuca pallescens, a grass of high forage value present with a low cover in degraded grasslands. To test the performance of A.splendens as a nurse plant in non-degraded grassland, we recorded the species richness four years inside of A. splendens senescent shrubs and in gaps among dominant tussock grasses. Species were grouped in four functional groups: annual and biannual herbs and grasses, perennial herbs, perennial grasses and shrubs. To test the usefulness of A. splendens in the restoration of degraded grassland, we monitored the seedling emergence and survival of F. pallescens inside A. splendens and in gaps. We related seedling survival to meteorological and microenvironmental conditions. Species richness was higher in Acaena nurse plants than in gaps. The frequency of functional groups, with exception of annual and biannual herbs and grasses, were higher in Acaena than in gaps. Seedling emergence and survival of F. pallescens were higher in Acaena, but the seedlings died in summer in both microsites. Mean maximum temperature was higher and mean minimum humidity lower in gaps than in Acaena during spring. However, the spring-summer season in which we monitored F. pallescens survival, was exceptionally dry and hot, affecting the survival of F. pallescens seedlings. Our results show that A. splendens act as a nurse species increasing the richness in the non-degraded grassland and facilitating the seedling recruitment of an important forage species in the degraded grassland. Nevertheless, the facilitation mechanism will fail in drought conditions, indicating that this restoration tool is limited by climate.