A new combination and a new variety in Artemisia tridentata

The combination Artemisia tridentata Nutt. ssp. spiciformis (Osterhout) Goodrich & McArthur comb. nov. is made. This high elevation taxon was originally described at the species level and more recently has generally been treated as a form of A. tridentata ssp. vaseyana . The subspecies designation is supported by its parallel nature to the other A. tridentata  subspecies and by its relatively widespread and locally abundant populations. Also, a new variety of A. tridentata ssp. vaseyana is proposed.         

Correlation of neighborhood relationships, carbon assimilation, and water status of sagebrush seedlings establishing after fire

Interactions of Artemisia tridentata ssp. vaseyana (mountain big sagebrush) and neighboring herbs may affect community development following fire in sagebrush steppe. Dry mass, photosynthesis, and water relations were measured for seedlings of A. tridentata ssp. vaseyana occurring at different distances from neighboring herbs in the initial growing seasons following fire, when herbs dominate plant community cover. Seedling mass significantly increased as distance to neighboring herbs increased, although a low r 2 indicated that mass was also affected by other sources of variation. Carbon assimilation ( A net ) was also greater for A. tridentata ssp. vaseyana seedlings in microsites farther from herbs, except during those sampling dates when A net and water availability were at low levels. Contrary to our expectations, water status of seedlings was not correlated with their distances to neighboring herbs, and supplemental watering did not affect the slope of the relationship between seedling mass and distance to neighboring herbs. These findings suggest that negative relationships between A. tridentata ssp. vaseyana seedlings and herbs establishing after fire are not likely due to competition over water and may instead result from interactions over factors not measured here (e.g., nutrients).     

Effects of nitrogen availability on growth and photosynthesis of Artemisia tridentata ssp. wyomingensis

This study examined the effects of alterations in soil nitrogen on the growth of Artemessia tridentata ssp. wyomingensis Nutt. Soil nitrogen content was altered by applying sugar (45 g/m 2 ), nitrate (4.5 g/m 2 ), or ammonium (4.5 g/m 2 ), and the results were compared with a control treatment (no soil amendments). Addition of either form of nitrogen significantly increased leaf nitrogen content, mean maximum length of ephemeral leaves, number of ephemeral leaves per terminal shoot, and current year's vegetative stem length over the control and sugar treatments. Both soil water and predawn xylem potentials during active growth were lower in the nitrogen-treated plots. The higher growth activity and greater leaf mass of A. tridentata in the nitrogen treatments may have been responsible for this result. Higher photosynthetic rates observed in the nitrogen treatments during an early June sampling period also lend support to this observation. This study suggests A. tridentata ssp. wyomingensis would opportunistically take advantage of increased availability of soil nitrogen. The ability of this species to respond positively to increased soil nitrogen may enhance its competitiveness over associated perennial species.     

Fertile island development around perennial shrubs across a Mojave Desert chronosequence

Spatial heterogeneity has been examined only recently as a factor in studies of ecosystem processes. The effect of this factor on desert organisms is exemplified in the ""fertile island"" a concentration of essential soil nutrients under perennial shrubs. In this study we examined fertile island patterns in undisturbed desert areas and on abandoned roadways to see at what rate and magnitude these fertile patches reappeared after disturbance. Small-scale examination of patterns of soil P, organic matter, moisture, and bulk density showed that soils beneath Larrea tridentata (DC.) Cov. shrubs on the roads lacked the tight circular gradient in these variables that was characteristic of soils beneath control shrubs, even 88 years after road abandonment. The nature of the initial soil disturbance altered both spatial patterns of soil N and temporal patterns of fertile island development. Fertile island patterns for total soil N, available P, and organic matter were more circular than patterns for bulk density, texture, or pH. We suggest that patterns of soil heterogeneity may develop first for elements that may be limiting to desert shrub growth (N, P, organic matter), followed by spatial development in other less limiting soil factors (bulk density, texture, pH).     

Arbuscular mycorrhizae of Mojave Desert plants

Roots of 15 Mojave Desert plant species were assessed for arbuscular mycorrhizal (AM) colonization in spring and autumn 1999. Another 19 species were assessed in autumn only. Perennial species were AM colonized, whereas annual species contained very low levels of AM hyphae or were nonmycorrhizal. The nonnative tree Tamarix ramosissima was also nonmycorrhizal. Levels of hyphae changed over the season in 2 species, arbuscle levels increased in 1 species, and vesicles decreased in 7 species. Mycorrhizal inoculum potential (MIP) was assessed in the soils associated with spring-collected plants. All soils were found to contain MIP. MIP values were not correlated with AM root colonization.     

Controls on radial growth of mountain big sagebrush and implications for climate change

Mountain big sagebrush ( Artemisia tridentata Nutt. ssp. vaseyana ) covers large areas in arid regions of western North America. Climate-change models predict a decrease in the range of sagebrush, but few studies have examined details of predicted changes on sagebrush growth and the potential impacts of these changes on the community. We analyzed effects of temperature, precipitation, and snow depth on sagebrush annual ring width for 1969 to 2007 in the Gunnison Basin of Colorado. Temperature at all times of year except winter had negative correlations with ring widths; summer temperature had the strongest negative relationship. Ring widths correlated positively with precipitation in various seasons except summer; winter precipitation had the strongest relationship with growth. Maximum snow depth also correlated positively and strongly with ring width. Multiple regressions showed that summer temperature and either winter precipitation or maximum snow depth, which recharges deeper soil horizons, are both important in controlling growth. Overall, water stress and perhaps especially maximum snow depth appear to limit growth of this species. With predicted increases in temperature and probable reduced snow depth, sagebrush growth rates are likely to decrease. If so, sagebrush populations and cover may decline, which may have substantial effects on community composition and carbon balance.     

Ecological impacts of seed harvester ants on soil attributes in a Larrea -dominated shrubland

The influence of seed harvester ant ( Pogonomyrmex rugosus ) colonies on soil properties and soil surface and moisture characteristics was investigated through comparison of adjacent, nonnest (reference, 4 m beyond ant colony) areas in Las Vegas, Nevada. Effects of ant colonies on both terrace and slope sites were investigated. Soil moisture content and soil bulk density in a creosote bush ( Larrea tridentata )--dominated shrubland were significantly lower, while soil temperature, soil organic matter, and percent pore space were significantly higher in soils with ant nests relative to adjacent reference soils. Soil pH and texture did not differ significantly between nest and reference soils. Among soil surface characteristics, percent bare soil and rock (gravel, cobble, and boulder) cover were not significantly different between nest and reference soils. In evaluating soil moisture characteristics, soils with ant nests had a significantly higher water infiltrability and greater depth of water penetration, but a significantly lower area of water spread (surface-water runoff) at both terrace and slope sites. Between the 2 geomorphic surfaces, water infiltrability and depth of water penetration were significantly greater at the terrace than at the slope. Water-borne soil movement (fluvial erosion) was significantly greater at the slope than terrace but did not differ significantly between nest and reference soils. The presence of active P. rugosus colonies in the L. tridentata -dominated shrubland altered certain soil properties and appeared to have a protective influence on the soil by fostering more infiltration and less runoff of surface water in southern Nevada.     

Experimental manipulations of precipitation seasonality: effects on oak ( Quercus ) seedling demography and physiology

Predicted changes in regional precipitation patterns and soil moisture caused by anthropogenic trace gas emissions may affect the distribution and abundance of woody plants in arid and semiarid regions. To test the response of woody plants to potential changes in precipitation regimes, we manipulated summer and winter precipitation on plots that contained seedlings of Quercus emoryi Torr. (Emory oak), the dominant tree in oak savannas of the southwestern United States. Throughout the growing season, we monitored seedling survival and physiology (predawn leaf water potential, midday instantaneous gas exchange, and leaf carbon isotope discrimination). Seedling survival and physiological performance differed little between treatments, which embodied 50% changes to quantities of summer and winter precipitation, and encompassed a continuum of precipitation from 359 mm · year -1 to 846 mm · year -1 . However, survival and physiological performance of seedlings were negatively impacted by seasonal environmental conditions common to all treatments, especially during the annual pre-'monsoon' drought. Seedling predawn leaf water potentials, net CO 2 assimilation, and stomatal conductance indicate that growing conditions for Q. emoryi seedlings at this site are generally restricted to periods with adequate soil moisture (i.e., April and August). Results contrast with an assumption implicit to the ""two-layer"" soil water resource partitioning hypothesis that woody plants in all life history stages are more dependent upon winter than summer precipitation. In fact, summer precipitation appears more important than winter precipitation for Q. emoryi seedling recruitment and growth.     

Summer habitat use and selection by female Sage Grouse ( Centrocercus urophasianus ) in Oregon

Cover types and vegetative characteristics (e.g., grasses, forbs, shrubs) used by female Sage Grouse ( Centrocercus urophasianus ) during summer were compared with available habitat on two study areas in southeastern Oregon. Broodless hens, which constituted 114 of the 125 (91%) radio-marked hens studied, selected big ( Artemisia tridentata ) subspp.) and low sagebrush ( A. arbuscular ) cover types at both study areas. At Hart Mountain, broodless hens did not select specific vegetative characteristics within cover types. However, at Jackass Creek, forb cover was greater ( P = .004) at broodless hen sites than at random locations. Differences in habitat use by broodless hens between study areas were associated with differences in forb availability. Broodless hens used a greater diversity of cover types than hens with broods. Broodless hens gathered in flocks and remained separate from but near hens with broods during early summer. By early July broodless hens moved to meadows while hens with broods remained in upland habitats.     

Does the presence of Wyethia mollis affect growth of Pinus jeffreyi seedlings?

Regeneration of Pinus jeffreyi in the Sierra Nevada is often limited on sites dominated by Wyethia mollis . Allelopathic chemicals and competition for soil moisture have been suggested as possible mechanisms for limiting regeneration. We tested the hypothesis that soil chemical and microbial properties from sites in different stages of succession influence seedling growth of Pinus jeffreyi . Soil was collected from an early-seral site dominated by Wyethia mollis , a mid-seral site dominated by the shrubs Arctostaphylos patula, Ceanothus prostratus, C. velutinus , and Purshia tridentata , and a late-seral site dominated by mature Pinus . These sites were compared for nutrient content, Pinus seedling growth capacity, and microbial population size. Soil (0-33 cm) from the early-seral site had the lowest C, microbial biomass, and fungal and bacterial populations. There were no consistent trends in soil nutrient content among sites. The early-seral site had the lowest soil Ca and Mg contents but also had a lower C/N ratio and more than twofold greater P content than either the mid- or late-seral site. Pinus seedling growth and foliar nutrient concentrations were compared at 3 harvest dates (220, 314, and 417 days after germination) in a greenhouse bioassay. The treatment design was a 3 × 2 factorial with soil from each of the 3 sites either with or without Pinus seedlings. Pots without seedlings were used as controls to assess the effects of seedlings on microbial biomass. Seedling growth in the early-seral soil was initially suppressed in comparison to growth in the mid-seral soil, but by the final harvest total seedling weight was similar between these 2 treatments. The most obvious treatment effect was a reduction in growth for seedlings planted in lateseral soil, probably due to a nutrient imbalance in the soil. Seedlings grown in late-seral soil had Fe and Al levels that were nearly twice those of seedlings grown in early- and mid-seral soils. Microbial biomass followed a temporal pattern similar to that found for seedling growth. Differences in microbial biomass between the early- and mid-seral soils, although initially large, were not detected by the final harvest. We interpret these results to indicate that allelopathy or soil nutrient deficiencies resulting from the presence of Wyethia are unlikely to be responsible for limited growth of Pinus seedlings in Wyethia -dominated stands.     

Arbuscular mycorrhizae in thermal-influenced soils in Yellowstone National Park

Mycorrhizae are common plant-fungal symbioses occurring in most land plants. Despite their ubiquity, little is known about the distribution of arbuscular mycorrhizae (AM) in extreme environments. We surveyed for the presence of AM in thermal sites in Yellowstone National Park (YNP) where soils are characterized by extreme pHs, elevated temperatures, and toxic element concentrations. Plants at 5 sites, growing in soils with rooting-zone temperatures up to 48°C and soil pH values as low as 3.4, were mycorrhizal (colonization levels from 4% to 34%). Soils from a sparsely vegetated thermal area and an adjacent, continuously vegetated transition area differed significantly in rooting-zone temperature (35°C vs. 26°C), acidity (pH 3.8 vs. 5.4), electrical conductivity (2.22 vs. 0.49 mmhos cm -1 ), Fe (181.3 vs. 48.5 mg kg -1 ),Mn (7.2 vs. 98.2 mg kg -1 ), and Zn (2.3 vs. 4.5 mg kg -1 ). Mycorrhizal infectivity potential (MIP) was 77% greater in the transition soils, with colonization levels of 26% and 46% in thermal and transition soils, respectively. Furthermore, colonization of Agrostis scabra , Dichanthelium lanuginosum , and Mimulus guttatus was found to be consistently high throughout the growing season (from 48% to 72%). It is possible that AM are essential for plant life on the edge of thermal areas, and that either or both symbionts are specifically adapted to their environment. Further research is required to elucidate AM function in and specific adaptations to YNP's thermal areas.     

Nitrogen acquisition from different spatial distributions by six Great Basin plant species

Plants of different growth form may utilize soil nutrients in various spatial distributions through different scales of foraging. In this study we evaluated the ability of 6 species commonly found in the Great Basin to utilize nitrogen (N) distributed in different patterns. Three growth forms were represented by these 6 species. We applied 15 N-labeled nitrogen in concentrated patches and over broader uniform areas (at approximately 1% the concentration of the patches) in large, outdoor sand-culture plots. Six weeks after N was applied, 2 plants adjacent to the patch (Patch Treatment) and 2 plants within the uniform application (Uniform Treatment) were harvested. One plant 35-45 cm from both applications (Distant Treatment) was also harvested. The proportion of application-derived N in the leaf N pool was calculated and the mass of N this represented was estimated. Winter annual species Aegilops cylindrical and Bromus tectorum utilized the concentrated patches to a greater extent than did perennial species. The mass of N acquired by Patch-Treatment annual plants was significantly greater than by Uniform- and Distant-Treatment plants. Annual plants in the Distant Treatment had very little application-derived N in their leaf tissue. The perennial tussock grasses Agropyron desertorum and Pseudoroegneria spicata differed in utilization of the N applications. Agropyron acquired a greater quantity of N from patches than from uniform applications, and Distant- Treatment plants acquired very little from treatment applications. On the other hand, Pseudoroegneria utilized N in the 3 treatments equally. The shrub species Artemisia tridentata ssp. vaseyana and Chrysothamnus nauseosus also differed in their pattern of N acquisition. There were no differences in quantity of N acquired by plants from different treatments for Chrysothamnus ; all treatment plants acquired appreciable amounts of N from the applications. In contrast, Artemisia tridentata was very effective at acquiring large quantities of N from patches relative to Uniform- and Distant-Treatment plants, and yet there was still appreciable acquisition of applied N by Distant-Treatment Artemisia plants. We compared our results for these species in utilizing N patches with their ability to utilize N pulses (Bilbrough and Caldwell 1997). The annual grasses, Artemisia , and Agropyron were capable of effectively acquiring N from both pulses and patches, whereas Pseudoroegneria was effective in exploiting pulses but not patches. Chrysothamnus was generally not responsive to either patches or pulses. Our results suggest that the 2 shrubs and 2 perennial grasses differed in the scale at which they foraged for nutrients. Some species exhibited a coarse-scale utilization of nutrients while others were clearly capable of fine-scale utilization of spatially distributed nutrient sources. This suggests the potential for at least some spatial niche separation among these species.     

Cone and seed production in Pinus ponderosa : a review

Factors associated with seed cone production in Pinus ponderosa were reviewed to identify broad patterns and potential effectiveness of restoration activities. Cone and seed production are quite variable, with differences between (1) years, (2) sites, and (3) individual trees. Between-year, population-wide crop failures suggest large-scale triggers for cone and seed production, perhaps high temperatures and dry weather. Stem diameter is the most important determinant for cone production at the tree level, with other factors such as genetic disposition, moisture, soil nutrients, and insect pests and disease playing a smaller role. Some extrinsic factors affect growth rate, indirectly affecting cone production. For example, less competition and lower stand densities result in P. ponderosa trees that increase in diameter more quickly, possibly because of more light, and produce seeds earlier. This literature suggests that restoration activities, especially thinning, will result in trees better able to produce larger seed crops. The effect of prescribed fire is less clear, with contradictory effects depending on site conditions, burn severity, and nutrient status of the site.     

Response of sagebrush steppe species to elevated CO 2 and soil temperature

Elevated atmospheric CO 2 may cause long-term changes in the productivity and species composition of the sagebrush steppe. Few studies, however, have evaluated the effects of increased CO 2 on growth and physiology of species important to this ecosystem. Since the response of plants to elevated CO 2 may be limited by environmental factors, soil temperature was also examined to determine if low soil temperatures limit CO 2 response. To determine how CO 2 and soil temperature affect the growth of species native to the sagebrush steppe, bottlebrush squirreltail [ Elymus elymoides (Raf.) Swezey], Thurber needlegrass ( Stipa thurberiana Piper), and Wyoming big sagebrush ( Artemisia tridentata ssp. wyomingensis Beetle) were grown in ambient (374 mL L -1 ) or high (567 mL L -1 ) CO 2 and low (13° C) or high (18° C) soil temperature for approximately 4 months. Although soil temperature affected the growth of squirreltail and needlegrass, temperature did not modify their response to elevated CO 2 . Total biomass of sagebrush was consistent across soil temperature and CO 2 treatments, reflecting its slow-growing strategy. All 3 species had higher leaf water-use efficiency at elevated CO 2 due to higher net photosynthesis and lower transpiration rates. We conclude that elevated CO 2 and soil warming may increase the growth of grasses more than shrubs. Field studies in the sagebrush steppe are necessary to determine if differences in biomass, resulting from changes in CO 2 and soil temperature, are exhibited in the field.     

Cyanobacteria and cyanolichens: Can they enhance availability of essential minerals for higher plants?

In both field and greenhouse studies, cyanobacterial and cyanolichens of cold-temperature deserts often enhance growth and essential uptake by associated herbs. That effect is associated with better seedling establishment and larger seedlings. The following are possible mechanisms for these effects: (1) the microbiota concentrate essential elements in available forms in soil surface layers, (2) the microbial surface covers are usually darker colored than the soil itself and produce warmer soils during cool seasons when soil water is most available, (3) the gelatinous sheaths of several cyanobacterial genera common on alkaline deserts contain chelating compounds, and (4) conditions that favor persistent microbial growths on soil surfaces also favor maintenance of larger populations of microorganisms that form mycorrhizal and/or rhizosheath associations with seed plants. There is evidence that associated animals may be nutritionally benefited by the enhanced mineral content of forage plants growing in well-developed cyanobacterial crusts.     

Cryptogamic soil crusts: seasonal variation in algal populations in the Tintic Mountains, Juab County, Utah

The soil algae in the Tintic Mountains, Juab County, Utah, was studied over a one-year period in 1982 and 1983. Fluorescence microscopy was used to measure algal density in samples directly from the field. A total of 30 algal taxa was observed, blue-green algae being most abundant both in terms of density and number of species. Algal density showed peaks in late fall and late spring. Minima were present in September 1982 and July 1983. Several weak correlations between algal density and climatic data existed. In general algae correlated positively with precipitation and negatively with temperature. A combination of low precipitation and hot temperatures was likely responsible for the low density observed in July. The Chrysophyta followed slightly different trends than the other algal groups, having minima in early October 1982 and late August 1983. Field observations indicated that the degree of algal crusting varied noticeably over a period of one year, with highest abundance of hummocking in spring. Heavy summer thunderstorms destroyed algal crusting during July and August of 1983, though absolute density of algae increased during this time in response to the extra moisture.     

Long-term fire history in Great Basin sagebrush reconstructed from macroscopic charcoal in spring sediments, Newark Valley, Nevada

We use macroscopic charcoal analysis to reconstruct fire history in sagebrush ( Artemisia tridentata var. wyomingensis and A. tridentata var. tridentate ), in Newark Valley, Nevada. We analyzed charcoal at continuous 1-cm intervals (～7-127 years), and pollen at 2- to 10-cm intervals (～70-263 years) in a core spanning the last 5500 cal yr BP (calendar years before present). A charcoal peak in the historic period was associated with a >1400-ha fire dated to 1986 that burned in the watershed. We reconstructed the prehistoric fire history by inferring fires from similar charcoal peaks that were significantly greater than the background charcoal accumulation. Our results suggest the fire regime is climate and fuel driven. During periods of wetter climate, sagebrush increased and fires were more abundant, and during extended dry periods when sagebrush decreased, fires were less frequent. Our method does not allow calculation of a fire-return interval; however, our results support models that estimate a mean fire-return interval of up to a century in Artemisia tridentata var. wyomingensis . The charcoal record indicates that fires have increased within the historic period. This contrasts with pinyon/juniper studies that indicate an expansion of woodland associated with fewer fires in the historic period. We suggest that in the central Great Basin, a regime of frequent fires in sagebrush that limits woodland expansion is true for the sagebrush-woodland ecotone, but in sagebrush-dominated valleys with lower fuel loads, fires have always been less frequent. Protecting sagebrush-dominated valleys from frequent fire would appear to be consistent with the prehistoric fire regime.     

Experimental manipulations of fertile islands and nurse plant effects in the Mojave Desert, USA

In a mixed desert shrub community we removed and added shrub canopies to examine above- and belowground influences of 3 species of shrubs on islands of soil fertility and the survival of transplanted Ambrosia dumosa seedlings. Soils sampled under shrubs in the wet season had higher pH, water content, organic matter, and both total and mineralizable nitrogen than soils in adjacent open areas, confirming a widely established pattern in arid lands. However, we also found species differences in soil parameters. Soils under Coleogyne ramosissima had highest pH, soils under A. dumosa had highest water content and nitrogen mineralization rates, and soils under Larrea tridentata had lowest water content. Soils sampled under shrubs in the dry season, 7 months after experimental shrub removal, maintained higher organic matter and total and mineralizable nitrogen content than adjacent open soils, but pH and water were altered by shrub manipulations. Species differences persisted only in soil water levels ( A. dumosa soils were driest). Over a 1-year period, transplanted A. dumosa seedlings had highest survivorship in shrub removal and open treatments and died most rapidly under control shrubs of all 3 species, suggesting that shrubs had a strong negative effect on seedling survival, even in the presence of higher organic matter, nutrients, and (initially) higher water content of fertile islands. Our results suggest that nurse plants and islands of soil fertility have the potential to facilitate growth of other species by nutrient additions, but that the net effect of nurse plants can be negative due to shading and/or root competition.