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.     

Vegetation zones and soil characteristics in vernal pools in the Channeled Scabland of eastern Washington

Vernal pools are seasonal pools occurring in Mediterranean-type climates within which grow concentric zones of vegetation. We studied two vernal pools that lie within an Artemisia tridentata/Festuca idahoensis shrub-steppe landscape in the Channeled Scabland of eastern Washington to determine the relationship between vegetation zonation and soil characteristics. Abundant plant species in the pools include Elymus cinereus, Poa scabrella, Lomatium grayi, Allium geyeri, Eleocharis palustris, Epilobium minutum, Myosurus aristatis, Deschampsia danthonioides, and Psilocarphus oregonus . We surveyed topography, measured plant species frequency and cover to describe the vegetation zones, and used Sorenson's index of percent to similarity to verify our designation of plant zones as communities. In one pool we described soil profiles and sampled soils throughout the growing season according to plant communities. We analyzed soils for pH; electrical conductivity; sodium, calcium, and magnesium ions; sodium absorption ratio; particle size; organic carbon; and water matric potential. ANOVA tests of soil characteristics and topography among plant communities showed that only differences in topography are statistically significant. There are, however, trends in particle size, some soil chemical parameters, and soil moisture potential among plant communities along the topographic gradient. Electrical conductivity decreased with increasing dryness of the soil through spring and summer. Seasonal changes in soil moisture potential showed that shallower soils in the centers of pools were wetter during the wet season and drier during the dry season than are deeper soils. These changes in moisture may be the most important influence on vegetation distribution within the vernal pools.     

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.     

Carbon isotope discrimination and water relations of oak hybrid populations in southwestern Utah

The evergreen oak Quercus turbinella and the deciduous Q. gambelii form natural hybrids in southwestern Utah and northern Arizona. Hybrid individuals also are found in northern Utah in a region where only Q. gambelii currently exists, indicating that Q. turbinella has recently retreated southward. Our objectives were to (1) examine the ecophysiology of parental taxa and hybrids under natural conditions in southeastern Utah, and (2) investigate the level of integration between leaf carbon isotope discrimination (a synthetic gas exchange trait) and structural and chemical traits of leaves in morphologically variable hybrid populations. Leaf length, width, mass-to-area ratio (LMA, g m -2 ), and nitrogen concentration (N, g g -1 ) within 2 hybrid populations near New Harmony, Utah, were highly intercorrelated. Variation within the hybrid populations spanned mean values for these traits observed in parental taxa from adjacent ""pure"" populations of each species. Carbon isotope discrimination (Δ), an integrated measure of the ratio of intercellular to ambient CO 2 concentration, ranged from 16.1‰ to 19.6‰ within the 2 hybrid populations and was positively correlated with leaf nitrogen concentration and negatively correlated with LMA; individuals in hybrid populations with leaves resembling Q. gambelii had the highest leaf Δ and N concentrations and lowest LMA compared with leaves from plants that resembled Q. turbinella . CO 2 uptake is limited by stomatal conductance and possibly by mesophyll resistance to a greater extent in Q. turbinella phenotypes than in intermediate or Q. gambelii phenotypes. δD of stem xylem water (an indication of active rooting depth) and predawn water potential during the peak monsoon period in August were not correlated to leaf Δ values within the hybrid populations. Several individuals that were morphologically similar to Q. turbinella in the hybrid populations maintained high predawn water potentials and derived moisture from winter recharge that presumably was taken from deep soil layers. Apparently, a few adult individuals of the Q. turbinella phenotype in hybrid populations accessed water from deep in the soil profile, which enabled them to avoid summer drought. Reduced monsoonal activity may have been an important, but not the single, determinant of Q. turbinella s retreat from northern Utah during the recent Holocene.     

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.     

Gas exchange, δ 13 C, and heterotrophy for Castilleja linariifolia and Orthocarpus tolmiei , facultative root hemiparsites on Artemisia Tridentata

Gas exchange and carbon isotope ratios were measured on 2 facultative hemiparasites, Castilleja linariifolia Benth. (Indian paintbrush); Scrophulariaceae) and Othrocarpus tomiei H. & A. (Tolmie owl clover, Scrophulariaceae), and their Artemisia tridentata L. (big sagebrush; Asteraceae) hosts. Photosynthetic rates differed greatly between years; rates in 1995 were more than double those in 1994, likely due to more precipitation and less water stress during 1995. Despite the difference in precipitation, photosynthetic rates for C. linariifolia were not different from those of their hosts for either year. However, carbon isotope ratios of C. linariifolia and O. tolmiei were up to 3% more negative than those of their A. tridentata hosts. Using measured δ 13 C ratios in conjunction with δ 13 C predicted from gas exchange measurements, we calculated that C. linariifolia derived, on average, 40% of its leaf carbon heterotrophically. Contrary to current suggestions that high photosynthetic rates of hemiparasites are an indication of reduced heterotrophy, C. linariifolia exhibited photosynthetic rates similar to autotrophic plants and used a substantial amount of host-derived carbon. Moreover, this evidence shows that manipulation of a heterotrophic carbon supply transcends obligate hemiparasites to include those plants whose parasitism is facultative.     

Nonresponse of native cottonwood trees to water additions during summer drought

Studies have demonstrated that some riparian trees may switch their reliance on surface soil water (unsaturated or vadose zone) to groundwater (saturated zone) sources during the growing season in association with changes in moisture availability. A closely related question is: How do these trees respond to pulse increases in water availability in previously dry zones? We tested the whole-tree physiological response of 6 natural Populus genotypes to water additions during the peak of summer drought in northern Utah, USA. We found clear evidence that trees were insensitive to water additions to the surface soil that were twice the magnitude of whole-tree transpiration rates. Our results suggest that some cottonwoods may have little immediate transpiration or leaf conductance response to pulse soil moisture increases. This lack of response may be related to a water-use strategy associated with regional climate patterns (i.e., genetic or environmental programming), cavitation recovery, or other physical determinants of water use such as depth to groundwater. Our data suggest that it is important to consider potential nonresponsiveness to changes in soil water availability when evaluating the impact of climate change on these important and productive ecosystems.     

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.     

Woody riparian vegetation response to different alluvial water table regimes

Woody riparian vegetation in western North American riparian ecosystems is commonly dependent on alluvial groundwater. Various natural and anthropogenic mechanisms can cause groundwater declines that stress riparian vegetation, but little quantitative information exists on the nature of plant response to different magnitudes, rates, and durations of groundwater decline. We observed groundwater dynamics and the response of Populus fremontii , Salix gooddingii , and Tamarix ramosissima saplings at 3 sites between 1995 and 1997 along the Bill Williams River, Arizona. At a site where the lowest observed groundwater level in 1996 (-1.97 m) was 1.11 m lower than that in 1995 (-0.86 m), 92-100% of Populus and Salix saplings died, whereas 0-13% of Tamarix stems died. A site with greater absolute water table depths in 1996 (-2.55 m), but less change from the 1995 condition (0.55 m), showed less Populus and Salix mortality and increased basal area. Excavations of sapling roots suggest that root distribution is related to groundwater history. Therefore, a decline in water table relative to the condition under which roots developed may strand plant roots where they cannot obtain sufficient moisture. Plant response is likely mediated by other factors such as soil texture and stratigraphy, availability of precipitation-derived soil moisture, physiological and morphological adaptations to water stress, and tree age. An understanding of the relationships between water table declines and plant response may enable land and water managers to avoid activities that are likely to stress desirable riparian vegetation.     

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.     

Selenium geochemical relationships of some northern Nevada soils

Soil samples, one from each of 10 locations in northern Nevada, were evaluated for redox potential, total and extractable selenium, phosphate, free iron oxide, total and ferrous iron. Mole fractions for extractable selenium species were calculated from redox potentials. Data were used to extrapolate general geochemical relationships for soil selenium at the sample sites. Results obtained from one sample per location allowed only the most general conclusions to be drawn. Soil phosphate levels, which affect the adsorption of selenite species on iron oxide by competing for adsorption sites, were not correlated with levels of extractable selenium in this study. This would suggest that selenium would exist in solution, having been displaced from adsorption sites by phosphorous. Ferrous iron, iron oxides, and redox potential had a combined effect on the level of extractable selenium at all sites. Soils in this study support selenite species that are not readily available to plants and therefore could not support vegetation adequate in Se.     

Testing hypothesized evolutionary shifts toward stress tolerance in hybrid Helianthus species

We examined how plant traits related to growth and resource use have evolved during hybrid speciation and specialization into stressful habitats. Two desert sunflower species of homoploid hybrid origin are endemic to habitats with lower soil nutrient levels than those of their ancestral parent species. We hypothesized that the hybrid species would exhibit greater tolerance to low levels of soil nutrients than their parental species. The 2 hybrid species, Helianthus anomalus and H. deserticola , and their parental species, H. annuus and H. petiolaris , were compared for plant traits and growth through reproduction under 3 nutrient levels in a greenhouse study. An additional seedling study compared species for maximum seedling relative growth rate under optimum conditions. The hybrid species did have greater tolerance of nutrient limitation than the parental species, demonstrated by a resistance to change in stem height and diameter growth across treatments. A similar trend was observed in total biomass at final harvest. This ability to maintain growth may be partially explained by maintained investment in photosynthetic enzymes regardless of nutrient treatment. Though the hybrid species were more tolerant of nutrient stress, differences in the hybrid response to nutrient stress compared to the parental species' response were much smaller than expected from habitat comparisons. Helianthus anomalus has evolved a classic stress-tolerant phenotype, having long leaf lifespan, tough leaves, and slower early seedling relative growth rate. While both hybrid species have a conservative growth strategy, which confers greater stress tolerance than the parental species possess, functional trait differences among the hybrids suggest that the 2 species have experienced vastly different selective pressures.     

Ecophysiology of the temperate desert halophytes: Allenrolfea occidentalis and Sarcobatus vermiculatus

Numerous basins of the intermountain area often have extensive playa surfaces that are nearly devoid of vegetation. Margins of these playas support sparse communities dominated by chenopod shrubs Allenrolfea occidentalis (iodine bush) and Sacrobatus vermiculatus (black greasewood). These plants establish and persist in an environment where halomorphic soils induce extreme osmotic stress and atmospheric precipitation is very low and erratic and occurs largely during the winter when temperatures are too low for growth. We measured net CO 2 assimilation rates, leaf conductances, transpiration rates, water-use efficiencies, and stem xylem potentials for these two C3 species. Data were collected in above-average (1991) and below-average (1992) precipitation years. Net CO 2 assimilation rates for Allenrolfea were statistically similar in 1991 and 1992 but in general declined for Sarcobatus in 1992. For both species, leaf conductances and leaf transpiration rates declined significantly from 1991 to 1992, with the decline greater for Sarcobatus . Water-use efficiencies doubled from 1991 to 1992 for both plant species. Predawn xylem water potentials were -2.2 and -3.3 MPa for Allenrolfea and -1.8 and -2.6 MPa for Sarcobatus by September 1991 and 1992, respectively. Afternoon xlem water potentials were -3.1 and -2.0 MPa for Allrolfea and -2.6 and -2.2 for Sarcobatus beginning in May 1991 and 1992, respectively. Xylem water potentials dropped to -5.0 MPa for Allenrolfea and -3.4 MPa for Sarcobatus by September for both 1991 and 1992. For Allenrolfea , in general, the total soil water potential within the zone of maximum root activity is more negative than the plant's predawn xylem potential, which suggests that the plant is partially phreatophytic and/or has a large capacitance due to its extensive woody root system.     

Growth of Plecoptera (stonefly) nymphs at constant, abnormally high temperatures

Six species of Plecoptera were maintained at four different temperatures, which were constant and higher than occurred in the natural habitat, and three species at two different day lengths. Each animal was weighed each day or each week. Weight of two species in the wild was monitored from periodic collection. The weight of each animal fluctuated rhythmically, changing about five percent every five days. These short - term fluctuations probably resulted from changes in water content. Molting occurred when a peak weight was predicted from the cycle and involved temporary gain of about 20 percent in weight. Growth probably stopped for some time before molt and was most rapid just afterward. Many animals died at molt. The time before death was less for univoltine species than for those with longer life cycles. Plecoptera collected in winter from water near 0 C lived for shorter times than did those collected in autumn from water near 10 C. Two species died sooner at higher temperatures and one died sooner with shorter day lengths. Growth in the laboratory was generally slower than in nature. One species grew faster, while three grew more slowly at higher temperatures. One species grew faster under long - than short - day conditions. Premature emergence, expected at the higher temperatures, did not occur, except in one animal.     

Soil microstructure in soils of the Colorado Plateau: the role of the cyanobacterium Microcoleus vaginatus

The role of the cyanobacterium Microcoleus vaginatus in cold-desert soil crusts is investigated using scanning electron microscopy. Crusts from sandstone-, limestone-, and gypsum-derived soils are examined. When dry, polysaccharide sheath material from this cyanobacterium can be seen winding through and across all three types of soil surfaces, attaching to and binding soil particles together. When wet, sheaths and living filaments can be seen absorbing water, swelling and covering soil surfaces even more extensively. Addition of negatively charged material, found both as sheath material and attached clay particles, may affect cation exchange capacity of these soils as well. As a result of these observations, we propose that the presence of this cyanobacterium may significantly enhance soil stability, moisture retention, and fertility of cold-desert soils.     

Seed production in Gentiana newberryi (Gentianaceae)

Experimental manipulations and observations in one population of Gentiana newberryi Gray flowers over 2 years showed significant variation in seed production relative to pollinator and soil water availability. When pollinators were rare, there was a significant relationship between number of bees present and number of mature seeds produced, and supplemental hand cross-pollination (xenogamy) did improve seed set in Gentiana newberryi Gray. When pollinators were abundant, supplemental hand cross-pollination did not increase seed set. Self-fertilized seeds (autogamy) germinated at the same rate as cross-pollinated seeds. Seed production in unvisited flowers is probably limited anatomically and is not influenced by the type of fertilization. There was a significant relationship between soil moisture and flower size in G. newberryi , with larger flowers found in wetter areas.     

Effect of irrigation on survival of third-stage Haemonchus contortus larvae (Nematoda: Trichostrongylidae)

During May through October 1976, samples of sheep pellets containing laboratory - reared, thirdstage Haemonchus contortus larvae were placed on irrigated and nonirrigated pasture plots at Provo, Utah. Periodically thereafter, grass clippings, soil scrapings, and remaining pellets were collected and baermannized to determine their comparative survival from the two environments. Water was added to the irrigated plots in accordance with a weekly sprinkling regime designed to furnish sufficient moisture to maintain pasture grass in this semiarid region. Meteorologic measurements were collected daily from both irrigated and nonirrigated sections.   During the year the nonirrigated section received a total of 131 mm of precipitation, whereas an additional 979 mm of water were added via sprinkling to the irrigated section. The monthly mean maximum temperature at soil surface under grass cover for the six - month study period on the irrigated section averaged 17.7 C less than on the nonirrigated section, and the corresponding soil moisture content remained 14.4 percent higher. A bioclimatograph of conditions on the nonirrigated section showed that none of the months during the year had levels of temperature and moisture which fell within the prescribed limits for optimum pasture transmission of H. contortus; on the irrigated section only October of the six - month study period failed to have suitable conditions for optimum pasture transmission. Larvae placed on the plots survived significantly longer and also in significantly greater numbers on the irrigated section, and irrigation enhanced the ability of larvae to migrate from pellets onto vegetation.     

Habitat relationships of Glaux maritima in central Utah

Thirty-five study sites were established in the meadow communities surrounding Utah Lake in central Utah. The study sites ranged across several community types. Glaux maritima was found in all sites but varied as to its ecological importance in these communities. Sixteen soil factors were measured relative to the stands studied. Cover of Glaux maritima correlated with parts per million sodium and total soluble salts in the soil. No other factors correlated significantly with the cover of Glaux maritima. Glaux maritima occupied only those sites with high levels of moisture throughout the growing season. The high moisture levels came from springs, seeps, elevated water tables, and early seasonal inundation. High levels of Glaux maritima cover corresponded to low numbers of species in the habitat.