Postfire vegetation dynamics in a sagebrush steppe in southeastern Idaho,USA

Postburn vegetation in an Artemisia tripartita and A. tridentata sagebrush community one year after the burn was compared with unburned vegetation. While the vegetal cover amounted to 38.1% of the total area on the burned site, it was 91.1% on the unburned site. Dominance-diversity curves for plant communities on both sites approach the niche pre-emption hypothesis or geometric series. Cover values and soil residual propagule data were used to suggest mechanisms of persistence of the more prominent species through fire using Noble & Slatyer's (1980) Vital Attributes model. The first year postfire vegetation was dominated by forbs and grasses with vegetative and propagule storage mechanisms of persistence. Such information on succession mechanisms should be of benefit to range managers.     

Large herbivores in sagebrush steppe ecosystems: livestock and wild ungulates influence structure and function

Improving understanding of the connections between vegetation, herbivory, and ecosystem function offers a fundamental challenge in contemporary terrestrial ecology. Using exclosures constructed during the late 1950s, we examined effects of grazing by wild and domestic herbivores on plant community structure, aboveground herbaceous primary production, and nutrient cycling at six sites in semi-arid, sagebrush rangelands during 2001-2002 in Colorado, USA. Enclosures provided three treatments: no grazing, grazing by wild ungulates only, and grazing by wild and domestic ungulates. Excluding all grazing caused an increase in shrub cover (F = 4.97, P = 0.033) and decrease in bare ground (F = 4.74, P = 0.037), but also a decrease in plant species richness (F = 6.19, P = 0.018) and plant diversity (F = 7.93, P = 0.008). Effects of wild ungulate grazing on plant cover and diversity were intermediate to the effects of combined domestic and wild grazing. Aboveground net primary production was higher in both grazed treatments than in the ungrazed one (F(wild + domestic) = 2.98, P = 0.0936 and F(wild only) = 3.55, P = 0.0684). We were unable to detect significant effects of grazing on other ecosystem states and processes including C:N ratios of standing crops, N mineralization potential, or nitrification potential. Best approximating models revealed positive correlation between N availability and herbaceous cover and a negative correlation between herbaceous primary production and the ratio of shrub-herb cover and plant diversity. We conclude that ungulate herbivory, including both wild and domestic ungulates, had significant effects on plant community structure and ecosystem function during this 42-year span. Responses to the wild ungulate treatment were consistently intermediate to responses to the no grazing and wild + domestic grazing treatments. However, we were unable to detect statistical difference between effects of wild ungulates alone and wild ungulates in combination with livestock.     

Variable responses of insects to hybrid versus parental sagebrush in common gardens

Both ecological and genetic mechanisms have been proposed to explain patterns of herbivore attack on interspecific plant hybrids, but distinguishing among them can be difficult in natural hybrid zones. We performed a common-garden experiment to evaluate four genetic hypotheses: dominance, additivity, elevated hybrid susceptibility, and elevated hybrid resistance. Censuses and cage experiments were used to compare insect responses to basin big sagebrush (Artemisia tridentata spp. tridentata), mountain big sagebrush (A. t. vaseyana), and their F₂ progeny. After two growing seasons, hybrid shrubs resembled mountain big sagenbrush in size, but were more similar to basin big sagebrush in flower production. Censuses of naturally colonizing insects (the gall midge Rhopalomyia obovata, the bagworm moth Apterona helix, and the aphid Obtusicauda coweni) tended to support the dominance hypothesis: if the insect clearly discriminated between the two parents, its frequency on hybrids closely resembled that on one of the parents. Moreover, colonization of hybrids in all three cases suggested a dominance deviation toward the susceptible parent rather than toward the resistant parent. In contrast to the censuses, cage experiments involving two insects supported the hybrid-susceptibility hypothesis; both survival and growth of the grasshopper Melanoplus sanguinipes and growth of the leaf beetle Trirhabda pilosa were higher on hybrid shrubs than on either parent. Because many secondary compounds have been determined to occur at intermediate concentrations in F₂ shrubs, dominance for susceptibility may indicate that insects respond to plant traits (e.g., oviposition stimulants and deterrents) in a threshold manner. Mechanisms underlying increased hybrid susceptibility are less clear, but our experimental design makes environmental explanations (e.g., the plant-stress hypothesis) unlikely. Although we eliminated several confounding factors, our results agree with the conclusion from natural hybrid zones that insect responses to hybrid plants are likely to be idiosyncratic; even congeneric species did not respond similarly to hybrid and parental plants.     

Net carbon exchange and evapotranspiration in postfire and intact sagebrush communities in the Great Basin

Invasion of non-native annuals across the Intermountain West is causing a widespread transition from perennial sagebrush communities to fire-prone annual herbaceous communities and grasslands. To determine how this invasion affects ecosystem function, carbon and water fluxes were quantified in three, paired sagebrush and adjacent postfire communities in the northern Great Basin using a 1-m³ gas exchange chamber. Most of the plant cover in the postfire communities was invasive species including Bromus tectorum L., Agropyron cristatum (L.) Gaertn and Sisymbrium altissimum L. Instantaneous morning net carbon exchange (NCE) and evapotranspiration (ET) in native shrub plots were greater than either intershrub or postfire plots. Native sagebrush communities were net carbon sinks (mean NCE 0.2–4.3 μmol m⁻² s⁻¹) throughout the growing season. The magnitude and seasonal variation of NCE in the postfire communities were controlled by the dominant species and availability of soil moisture. Net C exchange in postfire communities dominated by perennial bunchgrasses was similar to sagebrush. However, communities dominated by annuals (cheatgrass and mustard) had significantly lower NCE than sagebrush and became net sources of carbon to the atmosphere (NCE declined to −0.5 μmol m⁻² s⁻¹) with increased severity of the summer drought. Differences in the patterns of ET led to lower surface soil moisture content and increased soil temperatures during summer in the cheatgrass-dominated community compared to the adjacent sagebrush community. Intensive measurements at one site revealed that temporal and spatial patterns of NCE and ET were correlated most closely with changes in leaf area in each community. By altering the patterns of carbon and water exchange, conversion of native sagebrush to postfire invasive communities may disrupt surface-atmosphere exchange and degrade the carbon storage capacity of these systems.     

Plant age, communication, and resistance to herbivores: young sagebrush plants are better emitters and receivers

Plants progress through a series of distinct stages during development, although the role of plant ontogeny in their defenses against herbivores is poorly understood. Recent work indicates that many plants activate systemic induced resistance after herbivore attack, although the relationship between resistance and ontogeny has not been a focus of this work. In addition, for sagebrush and a few other species, individuals near neighbors that experience simulated herbivory become more resistant to subsequent attack. Volatile, airborne cues are required for both systemic induced resistance among branches and for communication among individuals. We conducted experiments in stands of sagebrush of mixed ages to determine effects of plant age on volatile signaling between branches and individuals. Young and old control plants did not differ in levels of chewing damage that they experienced. Systemic induced resistance among branches was only observed for young plants. Young plants showed strong evidence of systemic resistance only if airflow was permitted among branches; plants with only vascular connections showed no systemic resistance. We also found evidence for volatile communication between individuals. For airborne communication, young plants were more effective emitters of cues as well as more responsive receivers of volatile cues.     

Secondary successional patterns in a sagebrush (Artemisia tridentata) community as they relate to soil disturbance and soil biological activity

The relationship between secondary succession, soil disturbance, and soil biological activity were studied on a sagebrush community (Artemisia tridentata) in the Piceance Basin of northwestern Colorado, U.S.A. Four levels of disturbance were imposed. I: the vegetation was mechanically removed and as much topsoil as possible was left; 2: the vegetation was mechanically removed and the topsoil scarified to a depth of 30 cm; 3: topsoil and subsoil were removed to a depth of 1 m, mixed and replaced; 4: topsoil and subsoil were removed to a depth of 2 m and replaced in a reverse order. Plant species composition, dehydrogenase and phosphatase enzymatic activity, mycorrhizae infection potentials, and percent organic matter were the variables measured.Treatment 4 drastically altered the pattern of vegetation succession. Treatments 2, 3, and 4 started with Salsola iberica as the dominant species but six years later, 3 and to lesser extent 2 changed in the direction of the species composition of 1, dominated by perennial grasses and perennial forbs. Treatment 4 developed a shrub dominated community. The rate of succession was not decreased by the increased levels of disturbance. Both dehydrogenase enzyme activity and mycorrhizae infection potential (MIP) increased with the change from Salsola iberica to a vegetation dominated by either perennial grasses and forbs or shrubs. The intensity of disturbance in 2, 3, and 4 reduced drastically dehydrogenase activity and MIP, but in six years they recovered to levels comparable to 1. Phosphatase enzyme activity and organic matter were unrelated to species composition but related to treatment and time elapsed. In both cases a significant decrease was observed throughout the six-year period.Nomenclature followsThis study was funded by the United States Department of Energy under Contract No. DE-AS02-76EV04018.     

Organic matter turnover in a sagebrush steppe landscape

Laboratory incubations of¹⁵N-amended soils from a sagebrush steppe in south-central Wyoming indicate that nutrient turnover and availability have complex patterns across the landscape and between microsites. Total and available N and P and microbial C and N were highest in topographic depressions characterized by tall shrub communities. Net and gross N mineralization rates and respiration were also highest in these areas, but microbial efficiencies expressing growth relative to respiration cost were highest in soils of exposed ridgetop sites (prostrate shrub communities). Similar patterns occurred between shrub and intershrub soils, with greater nutrient availability under shrubs, but lower microbial efficiencies under shrubs than between. Surface soils had higher soil nutrient pools and N mineralization rates than subsurface soils, but N and C turnover and microbial efficiencies were lower in those surface soils. All soils decreased in respiration, mineralization, and immobilization rates during the 30-day incubation period, apparently approaching a steady-state substrate use. Soil microbial activity of the high organic matter accumulation areas was apparently more limited by labile substrate.     

Fire mediated patterns of population densities in mountain big sagebrush bird communities

We employed a chronosequence approach to evaluate patterns of bird abundance in relation to post-fire vegetation recovery in mountain big sagebrush (Artemisia tridentata vaseyana). We estimated population density for 12 species of birds within the perimeters of 4 fires that had undergone 8–20 years of vegetation recovery and on adjacent unburned areas in the northwestern Great Basin, USA. Six species showed negative responses to fire persisting up to 20 years. Two species showed positive responses with effects persisting for <20 years. Understory vegetation was similar between burned and unburned areas irrespective of recovery time, and shrub canopy cover was similar between burned and unburned sites after 20 years of recovery. Persistent reductions in bird densities lead us to conclude that shrub canopy cover alone is not a sufficient metric for predicting recovery of songbird abundances following disturbance in mountain big sagebrush. © 2013 The Wildlife Society.     

Effects of local extinction of the plains vizcacha (Lagostomus maximus) on vegetation patterns in semi-arid scrub

We studied spatial and temporal effects of local extinction of the plains vizcacha (Lagostomus maximus) on plant communities following widespread, natural extinctions of vizcachas in semi-arid scrub of Argentina. Spatial patterns in vegetation were examined along transects extending outward from active and extinct vizcacha burrow systems. Responses of vegetation to removal of vizcachas were assessed experimentally with exclosures and by documenting vegetation dynamics for 6 years following extinctions. Transect data demonstrated clear spatial patterns in plant cover, particularly an increase in perennial grasses, outward from active vizcacha burrows. These patterns were consistent with predictions based on foraging theory and studies that document grasses as the preferred food of vizcachas. Removal of vizcachas, experimentally and with extinctions, resulted in an immediate increase in perennial and annual forbs indicating that intense herbivory can depress forb cover, as well as grasses. After a 1-year lag following cessation of herbivory, cover of grasses increased. Forbs declined as grasses increased. The long-term effect of extinction of vizcachas was a conversion of colony sites from open patches dominated by forbs to dense bunch grass characteristic of the matrix. Major changes in vegetation occurred within 2–3 years after extinction, resulting in a large pulse of landscape change. However, some species of grasses were uncommon until 5–6 years after the vizcacha extinction. With extinction and colonization, vizcachas generate a dynamic mosaic of patches on the landscape and create temporal, as well as spatial, heterogeneity in semi-arid scrub.     

Edaphic limitations to growth and photosynthesis in Sierran and Great Basin vegetation

Summary Soils derived from hydrothermally altered andesite support unique communities of Sierran conifers (Pinus ponderosa Laws. and P. jeffreyi Grev. and Balf.) amongst sagebrush (Artemisia tridentata Nutt.) vegetation in the western Great Basin. Plants grown in soil derived from hydrothermally altered bedrock had lower growth rates, total biomass, and net photosynthetic rates than plants grown in soil derived from unaltered andesite of the same formation. Total dry mass was 10 to 28% lower for conifers grown in altered soil whereas dry mass of Artemisia tridentata and Bromus tectorum L. was reduced by over 90%. Results from a nutrient amendment experiment indicated that low phosphorus was the dominant limitation in altered soil, and phosphorus-deficiency affected growth primarily by limiting leaf area development rather than direct inhibition of photosynthesis. The proportionately greater reduction of biomass for Artemisia and Bromus grown in altered soil supports our hypothesis that Great Basin vegetation is excluded from altered soil by intolerance to nutrient deficiency. The Sierran conifers growing on this rock type are therefore free of competition for water with Great Basin vegetation and are able to persist in an exceptionally dry climate.     

Pollen dispersal by <Emphasis Type="Italic">Artemisia tridentata</Emphasis> (Asteraceae)

While the biophysics of anemophilous pollen dispersal is understood in principle, empirical studies for testing such principles are rare, particularly in native ecosystems. This paper describes mechanisms underlying the dispersal of Artemisia pollen in a Wyoming sagebrush steppe. The relationships between meteorological variables and pollen flux were defined during the 1999 Artemisia flowering season, and detailed processes at the individual plant level were experimentally tested in the field in 2000. Results indicated that Artemisia pollen presentation is continuous but with early morning maxima. Atmospheric pollen concentrations and potential dispersal rates are controlled at diurnal time scales by individual flower development together with characteristic changes in temperature/humidity and wind speeds, at multi-day scales by frontal weather patterns, and at week-long scales by flowering phenology.     

Spatial patterns of surface soil properties and vegetation in a Mediterranean semi-arid steppe

In arid and semi-arid areas with sparse vegetation cover, the spatial pattern of surface soil properties affects water and nutrient flows, and is a question of considerable interest for understanding degradation processes and establishing adequate management measures. In this study, we investigate the spatial distribution of vegetation and surface soil properties (biological crusts, physical crusts, mosses, rock fragments, earthworm casts, fine root accumulation and below-ground stones) in a semi-arid Stipa tenacissima L. steppe in SE Spain. We applied the combination of spatial analysis by distance indices (SADIE) and geostatistics to assess the spatial pattern of soil properties and vegetation, and correlation analyses to explore how these patterns were related. SADIE analysis detected significant clumped patterns in the spatial distribution of vegetation, mosses, fine root accumulation and below-ground stone content. Contoured SADIE index of clustering maps suggested the presence of patchiness in the distribution of earthworm casts, fine roots, below-ground stone content, mosses and biological crusts. Correlation analyses suggested that spatial pattern of some soil properties such as biological crusts, moss cover, surface rock fragments, physical crusts and fine roots were significantly related with above-ground plant distribution. We discuss the spatial arrangement of surface soil properties and suggest mechanistic explanations for the observed spatial patterns and relationships.     

Response of greater sage-grouse to sagebrush reduction treatments in Wyoming big sagebrush

Vegetation treatments have been widely implemented in efforts to enhance conditions for wildlife populations. Yet the effectiveness of such efforts often lack rigorous evaluations to determine whether these practices are effective for targeted species. This is particularly important when manipulating wildlife habitats in ecosystems that are faced with multiple stressors. The sagebrush (Artemisia spp.) ecosystem has been altered extensively over the last century leading to declines of many associated species. Wyoming big sagebrush (A. tridentata wyomingensis) is the most widely distributed subspecies, providing important habitats for sagebrush-obligate and associated wildlife. Sagebrush often has been treated with chemicals, mechanical treatments, and prescribed burning to increase herbaceous forage species released from competition with sagebrush overstory. Despite many studies documenting negative effects of sagebrush control on greater sage-grouse (Centrocercus urophasianus) habitat, treatments are still proposed as a means of improving habitat for sage-grouse and other sagebrush-dependent species. Furthermore, most studies have focused on vegetation response and none have rigorously evaluated the direct influence of these treatments on sage-grouse. We initiated a 9-year (2011–2019) experimental study in central Wyoming, USA, to better understand how greater sage-grouse respond to sagebrush reduction treatments in Wyoming big sagebrush communities. We evaluated the influence of 2 common sagebrush treatments on greater sage-grouse demography and resource selection. We implemented mowing and tebuthiuron application in winter and spring 2014 and evaluated the pre- (2011–2013) and post-treatment (2014–2019) responses of sage-grouse relative to these management actions. We evaluated responses to treatments using demographic and behavioral data collected from 620 radio-marked female greater sage-grouse. Our specific objectives were to evaluate how treatments influenced 1) sage-grouse reproductive success and female survival; 2) sage-grouse nesting, brood-rearing, and female resource selection; 3) vegetation responses; and 4) forbs and invertebrates. Our results generally suggested neutral demographic responses and slight avoidance by greater sage-grouse in response to Wyoming big sagebrush treated by mowing and tebuthiuron. Neither mowing nor tebuthiuron treatments influenced nest survival, brood survival, or female survival. Selection for nest and brood-rearing sites did not differ before and after treatments. Females selected habitats near treatments before and after they were implemented; however, the strength of selection was lower after treatments compared with pre-treatment periods, which may be explained by a lack of response in vegetation and invertebrates following treatments. Perennial grass cover and height varied temporally yet did not vary systematically between treatment and control plots. Forb cover and species richness varied annually but not in relation to either treatment type. Perennial grass cover and height, forb cover, and forb species richness did not increase within mowed or tebuthiuron-treated areas that received 2 or 6 years of grazing rest compared with areas that received no grazing rest. Finally, forb and invertebrate dry mass did not differ between treated plots and control plots at mowing or tebuthiuron sites in any years following treatments. Results from our study add to a large body of evidence that sage-grouse using Wyoming big sagebrush vegetation communities do not respond positively to sagebrush manipulation treatments. Management practices that focus on the maintenance of large, undisturbed tracts of sagebrush will best facilitate the persistence of sage-grouse populations and other species reliant on the sagebrush steppe.     

Creosotebush vegetation after 50 years of lagomorph exclusion

In 1939, an experiment was established on the Jornada Experimental Range to evaluate the effects of shrub removal, rabbit exclusion, furrowing, and seeding in creosotebush [Larrea tridentata (DC.) Cov] vegetation. Sixteen plots (21.3×21.3 m) were laid out in four rows of four plots per row with a buffer zone of 7.6 m between plots and rows. A barbed wire fence excluded cattle and poultry wire fencing excluded lagomorphs. Treatments were factorially applied at two levels. Plant cover in the plots was sampled in 1938 (before treatment), 1947, 1956, 1960, 1967 and 1989 with randomly located, line-intercept transects. Data from all sampling dates were analyzed as a split plot in time and main effects for 1989 tested by analysis of variance for a 2×4 factorial experiment. There were significant (P<0.10) year x treatment interactions. Seeding and furrowing treatments were ineffective but lagomorph exclusion and shrub clearing treatments resulted in significant treatment differences for several species. In 1989, basal area of spike dropseed (Sporobolus contractus A.S. Hitchc.) was 30-fold greater on the lagomorph excluded than on the lagomorph unexcluded treatment. Canopy cover of honey mesquite (Prosopis glandulosa Torr. var. glandulosa), tarbush (Flourensia cernua DC.) and mariola (Parthenium incanum H.B.K.) were affected by lagomorph exclusion. None of the responses were viewed as successional in nature. They principally represented individual species sensitivities to either absence of a primary herbivore or removal of aboveground shrub biomass. Though the physical treatments could be regarded as relatively severe disturbances of the system, the impacts on community vegetation dynamics were relatively insignificant.     

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.     

Species interactions at the level of fine roots in the field: influence of soil nutrient heterogeneity and plant size

Interference at the level of fine roots in the field was studied by detailed examination of fine root distribution in small soil patches. To capture roots as they occur in natural three-dimensional soil space, we used a freezing and slicing technique for microscale root mapping. The location of individual roots intersecting a sliced soil core surface was digitized and the identity of shrub and grass roots was established by a chemical technique. Soil patches were created midway between the shrub, Artemisia tridentata, and one of two tussock grasses, Pseudoroegneria spicata or Agropyron desertorum. Some soil patches were enriched with nutrients and others given only deionized water (control); in addition, patches were located between plants of different size combination (large shrubs with small tussock grasses and small shrubs with large tussock grasses). The abundance of shrub and grass roots sharing soil patches and the inter-root distances of individual fine roots were measured. Total average rooting density in patches varied among these different treatment combinations by only a factor of 2, but the proportion of shrub and grass roots in the patches varied sixfold. For the shrub, the species of grass roots sharing the patches had a pronounced influence on shrub root density; shrub roots were more abundant if the patch was shared with Pseudoroegneria roots than if shared with Agropyron roots. The relative size of plants whose roots shared the soil patches also influenced the proportion of shrub and grass roots; larger plants were able to place more roots in the patches than were the smaller plants. In the nutrient-enriched patches, these influences of grass species and size combination were amplified. At the millimeter- to centimeter-scale within patches, shrub and grass roots tended to segregate, i.e., avoid each other, based on nearest-neighbor distances. At this scale, there was no indication that the species-specific interactions were the result of resource competition, since there were no obvious patterns between the proportion of shrub and grass roots of the two species combinations with microsite nutrient concentrations. Other potential mechanisms are discussed. Interference at the fine-root level, and its species-specific character, is likely an influential component of competitive success, but one that is not easily assessed.     

Soil solution phosphate,root uptake kinetics and nutrient acquisition: implications for a patchy soil environment

Summary The importance of increased root phosphate (P) uptake kinetics, root proliferation and local increases of soil solution P (P₁) for P acquisition from fertile soil microsites was explored with a simulation model and calculated uptake was compared with experimental data. Based on the partitioning of added P in microsites to P₁ and P adsorbed on soil particles and the results of a dual-isotope-labeling experiment (Caldwell et al. 1991a), acquisition of P from the fertile microsites was some 20 X that of uptake from an equal volume of soil which received only water. Simulations were in general agreement and also showed that elevation of root P uptake kinetics could contribute more to the increased acquisition than did root proliferation under these circumstances. Although increased physiological uptake capacity for P has generally been considered to be of little benefit because of diffusion limitation, in patchy soil environments selective elevation of P uptake kinetics in fertile microsites may be of considerable benefit. These tests were conducted in calcareous soil which releases much less P into the soil solution than do many other soils. In many noncalcareous soils the benefits of selective elevation of root uptake kinetics would likely be greater.     

Multi-scale responses of vegetation to removal of horse grazing from Great Basin (USA) mountain ranges

Although free-roaming equids occur on all of the world’s continents except Antarctica, very few studies (and none in the Great Basin, USA) have either investigated their grazing effects on vegetation at more than one spatial scale or compared characteristics of areas from which grazing has been removed to those of currently grazed areas. We compared characteristics of vegetation at 19 sites in nine mountain ranges of the western Great Basin; sites were either grazed by feral horses (Equus caballus) or had had horses removed for the last 10–14 years. We selected horse-occupied and horse-removed sites with similar aspect, slope, fire history, grazing pressure by cattle (minimal to none), and dominant vegetation (Artemisia tridentata). During 1997 and 1998, line-intercept transects randomly located within sites revealed that horse-removed sites exhibited 1.1−1.9 times greater shrub cover, 1.2–1.5 times greater total plant cover, 2–12 species greater plant species richness, and 1.9–2.9 times greater cover and 1.1–2.4 times greater frequency of native grasses than did horse-occupied sites. In contrast, sites with horses tended to have more grazing-resistant forbs and exotic plants. Direction and magnitude of landscape-scale results were corroborated by smaller-scale comparisons within horse-occupied sites of horse-trail transects and (randomly located) transects that characterized overall site conditions. Information-theoretic analyses that incorporated various subsets of abiotic variables suggested that presence of horses was generally a strong determinant of those vegetation-related variables that differed significantly between treatments, especially frequency and cover of grasses, but also species richness and shrub cover and frequency. In contrast, abiotic variables such as precipitation, site elevation, and soil erodibility best predicted characteristics such as forb cover, shrub frequency, and continuity of the shrub canopy. We found species richness of plants monotonically decreased across sites as grazing disturbance increased, suggesting that either the bell-shaped diversity-disturbance curve of the intermediate-disturbance hypothesis does not apply in this system or that most sites are already all on the greater-disturbance slope of the curve. In our study, numerous vegetation properties of less-grazed areas and sites differed notably from horse-grazed sites at local and landscape scales during a wetter and an average-precipitation year. Electronic Supplementary Material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Short-term study of effects of fertilisation and cutting treatments on the vegetation dynamics of mountain heathlands in Spain

The influence of management and nutrient availability on the vegetation dynamics of heathlands characterised by Calluna vulgaris and Erica tetralix were studied in three mountain sites in Northern Spain. A total of 90 plots (1 m² each) received different combinations of cutting and twice the estimated background atmospheric deposition of nitrogen (56 kg ha⁻¹ yr⁻¹). One of the two dominant ericaceous species was selectively cut by hand at ground level and their regeneration compared in the presence or absence of the other. The results after 2 years showed significant effects of the fertiliser on the vegetation cover, mainly by favouring perennial herbaceous species. There were less noteworthy effects on the number of flowers and on the annual growth of the ericaceous species. It is concluded that, in the short term, increased nutrients alone, at twice the estimated current atmospheric deposition for the area, will not alter significantly the composition of the mountain heathlands. However, once the stands reach the mature phase, the capacity of the community to regenerate after a severe disturbance diminishes. A drastic impact, such as cutting may not result in re-growth of the same shrub species but in replacement by herbaceous species, which will also benefit from the increased nutrients.     

Sagebrush and grasshopper responses to atmospheric carbon dioxide concentration

Summary Seed- and clonally-propagated plants of Big Sagebrush (Artemisia tridentata var.tridentata) were grown under atmospheric carbon dioxide regimes of 270, 350 and 650 μl l⁻¹ and fed toMelanoplus differentialis andM. sanguinipes grasshoppers. Total shrub biomass significantly increased as carbon dioxide levels increased, as did the weight and area of individual leaves. Plants grown from seed collected in a single population exhibited a 3–5 fold variation in the concentration of leaf volatile mono- and sesquiterpenes, guaianolide sesquiterpene lactones, coumarins and flavones within each CO₂ treatment. The concentration of leaf allelochemicals did not differ significantly among CO₂ treatments for these seed-propagated plants. Further, when genotypic variation was controlled by vegetative propagation, allelochemical concentrations also did not differ among carbon dioxide treatments. On the other hand, overall leaf nitrogen concentration declined significantly with elevated CO₂. Carbon accumulation was seen to dilute leaf nitrogen as the balance of leaf carbon versus nitrogen progressively increased as CO₂ growth concentration increased. Grasshopper feeding was highest on sagebrush leaves grown under 270 and 650 μl l⁻¹ CO₂, but varied widely within treatments. Leaf nitrogen concentration was an important positive factor in grasshopper relative growth but had no overall effect on consumption. Potential compensatory consumption by these generalist grasshoppers was apparently limited by the sagebrush allelochemicals. Insects with a greater ability to feed on chemically defended host plants under carbon dioxide enrichment may ultimately consume leaves with a lower nitrogen concentration but the same concentration of allelochemicals. Compensatory feeding may potentially increase the amount of dietary allelochemicals ingested for each unit of nitrogen consumed.