Selective herbivory by the desert woodrat ( Neotoma lepida ) on Joshua trees ( Yucca brevifolia )

We studied herbivory by the desert woodrat ( Neotoma lepida ) on Joshua trees ( Yucca brevifolia ) in the Mojave Desert to determine whether N. lepida fed selectively based on leaf nitrogen content. We measured leaf nitrogen content and the location and amount of herbivory by woodrats on Y. brevifolia trees in southwestern Utah, USA. Neotoma lepida removed the outer tips of leaves (1/3 to 2/3 of total leaf length) but left leaf bases, which had lower nitrogen content, uneaten. Herbivory by N. lepida also was concentrated on rosettes that were oriented south, and these had significantly higher nitrogen content than rosettes that were oriented north. Finally, N. lepida fed more on trees that had higher mean leaf nitrogen contents. Thus, N. lepida selectively foraged among leaf parts, among rosettes of leaves, and among trees of Y. brevifolia , and disproportionate foraging was correlated with nitrogen content of leaves.     

Facilitation of Yucca brevifolia recruitment by Mojave Desert shrubs

Patterns of recruitment for Yucca brevifolia (Joshua tree) were investigated on 3 elevational transects, 1000-2000 m, in the Spring and Sheep Mountain ranges of southern Nevada. Yucca brevifolia is distributed throughout a broad range of plant communities dominated by Larrea tridentata and Ambrosia dumosa at low elevations, Coleogyne ramosissima at middle elevations, and an Artemisia-Pinus-Juniperus community at upper elevations. The density of Y. brevifolia gradually increased from the lowest elevations, peaked at 1600 m, and remained at intermediate levels at high elevations until reaching an abrupt upper elevational limit at 2000 m. Open substrate dominated the study areas; however, a large majority of Y. brevifolia seedlings were found growing under the canopy of other woody shrubs. This pattern of recruitment did not vary by site or elevation. Thirty-five species of perennial shrubs were identified in the study areas, 16 of which were found in association with at least 1 Y. brevifolia seedling. However, 4 shrubs were found in a nurse plant relationship with Y. brevifolia above the frequency predicted by either their canopy area or numerical dominance. Seedlings exhibited significant variation in aspect, relative to the center of the nurse shrub. In Lee and Lucky Strike canyons, recruitment occurred predominantly on the east and west sides of nurse shrubs, indicating the importance of specific microhabitats. Local presence of specific perennial shrubs resulted in higher levels of recruitment, causing a distinct pattern of community development, resumably through amelioration of abiotic stresses.     

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.     

Orientation and slope preference in barrel cactus ( Ferocactus acanthodes ) at its northern distribution limit

Microsite distribution and diurnal tissue temperature fluctuations were measured in Ferocactus acanthodes (barrel cactus) at its northern distribution limit in the Beaver Dam Mountains of southwestern Utah. At this location, barrel cacti were limited to south-facing slopes. Orientation (azimuth and angle) of both the cactus stem and of the apical region at the top of the cactus were nonrandom, strongly facing south. The potential adaptive significance of this orientation in minimizing apical meristematic tissue damage under low wintertime conditions and minimizing high temperature damage under summertime conditions is discussed.     

Differential responses to nitrogen form and concentration for Oryzopsis hymenoides and Elymus lanceolatus

In a greenhouse experiment, effects of nitrogen form and concentration on productivity and dry matter allocation differed between two species native to semiarid ecosystems of the Great Basin. Aboveground production of green surface area and of dry matter were consistently enhanced by increased nitrogen for the rhizomatous grass Elymus lanceolatus , but not for the bunchgrass Oryzopsis hymenoides . These differences were likely due to inherently low growth rates of O. hymenoides . Aboveground dry matter allocation also differed between the two species. O. hymenoides had more leaves per tiller with increased nitrogen, whereas leaf size but not number increased for E. lanceolatus . Furthermore, increases in tiller density with increased nitrogen for E. lanceolatus were almost three times greater than those for O. hymenoides . E. lanceolatus , but not O. hymenoides , was sensitive to the form of nitrogen supplied to the plants. When NH 4 -N was the only form of nitrogen supplied, high concentrations of NH 4 -N inhibited aboveground production of E. lanceolatus .     

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.     

Leaf area ratios for selected rangeland plant species

Leaf area estimates are required by hydrologic erosion, and growth/yield simulation models and are important to the understanding of transpiration interception, CO 2 fixation, and the energy balance for native plant communities. Leaf biomass (g) to leaf area (mm 2 ) linear regression relationships were evaluated for 15 perennial grasses, 12 shrubs, and 1 tree. The slope coefficient (β 0 ) of the linear regression equation is a ratio of leaf area to leaf biomass and is defined as the leaf area ratio [LAR = one-sided leaf area (mm 2 )/oven-dry leaf weight (g)]. LAR represents β 0 in each regression equation, where Y = β 0 (X). Linear regression relationships for leaf area were computed ( r 2 = .84-.98) for all 28 native range species after full leaf extension. Within-plant estimates of leaf area for mesquite ( Prosopis glandulosae Torr. var. glandulosae [Torr.] Cockll.) or lime prickly ash ( Zanthoxylum fagara [L.] Sarg.) were not significantly different ( P ≤ .05). LARs for three of the shrubs and the tree were established at four different phenological stages. There were no significant differences ( P ≤ .05) in LARs for lime prickly ash, mesquite, and Texas persimmon ( Diospyros texana Scheele) after full leaf extension during the growing season. The LAR relationship for Texas persimmon changed significantly after full leaf extension. LAR relationships for Texas colubrina ( Colubrina texensis [T. & G.] Gray) changed in response to water stress.     

Feeding behavior and performance of a rabbitbrush leaf-beetle ( Trirhabda lewisii ) feeding on Chrysothamnus nauseosus regrwoth after fire

Fire often positively affects the growth and nutrient content of plants regrowing after a burn. These changes have been associated with preferential feeding by herbivores in burned areas. In this study in southeastern Wyoming, Chrysothamnus nauseosus Pursh (rubber rabbitbrush) regrowing after a fire produced new shoots with a distinct growth form. Shoots were longer than those on unburned control sites and had longer leaves with longer internodes between leaves. We conducted feeding trials to detect whether C. nauseosus shoots regrowing after fire were nutritionally superior to shoots from unburned plants for the specialist leaf beetle, Trirhabda lewisii Crotch (Coleoptera: Chrysomelidae). We also measured C:N ratios and nitrogen and water contents of leaves from burned and unburned plants. Trirhabda lewisii adults preferred shoots from burned plants when given a choice. The beetles ate similar amounts of burned and unburned plants when fed only a single type. Females that were fed either burned or unburned plants did not differ in number of eggs laid. Chemical analyses revealed no significant differences in nitrogen or water content of leaves from burned versus unburned plants. Carbon-to-nitrogen ratio of burned plants was marginally lower compared with unburned plants. In contrast to previous studies, which suggest that herbivore attraction to burned areas leads to enhanced performance, our study shows that performance is not necessarily enhanced after fire.     

"Curing" of Nicotiana attenuata leaves by small mammals does not decrease nicotine contents

Basal leaves of Nicotiana attenuata are frequently found neatly excised at the petiole and piled on rocks or soil in the sun until dry, after which they disappear, sometimes to be found again in the nests of Neotoma lepida . In response to herbivore attack, N. attenuata increases the concentration of nicotine in its leaves, where it functions as an induced defense. Since excision of leaves at the petiole allows for leaf removal without substantially activating this induced defense, and air-drying at high temperatures can volatilize nicotine, we examined the hypothesis that the observed leaf ""curing"" behavior decreased nicotine contents. In a natural population, replicate bundles of excised leaves were allowed to dry in the sun for up to 96 hours and harvested in 10 intervals. Even though surface temperatures reached 63° C during drying, no significant loss of nicotine was observed. In the laboratory, significant losses of nicotine were not observed until leaves were dried at 100° C. Nicotine contents of naturally ""cured"" leaf piles at 4 populations were found to be marginally higher than those of neighboring intact plants from which the leaves were likely harvested. We conclude that mammalian ""curing"" behavior does not reduce nicotine contents and may allow the leaves to be used for insect repellant purposes.     

High rates of photosynthesis in the desert shrub Chrysothamnus nauseosus ssp. albicaulis

  Basic aspects of photosynthesis were investigated in white rubber rabbitbrush ( Chrysothamnus nauseosus (Pallas) Britt. ssp. albicaulis ), a common C 3 deciduous shrub native to arid regions of the western U.S. Under favorable field conditions, net photosynthesis (P n ) ranged from 36 to 73 mgCO 2 · dm – 2 · hr – 1 , which is relatively high for a woody species. The leaves from the actively growing flowering shoots exhibited higher P n than those on the vegetative shoots. P n also varied according to the age of the leaves and the location of the plants. P n did not light saturate even at quantum flux densities (QFD) equivalent to full sunlight. The light compensation point was relatively high (ca 100 μ mol · m – 2 · S – 1 ), perhaps due to the presence of a tomentose vestiture on the leaf surface. At high QFD's, the stomatal conductance was high (ca 520 mmol · m – 2 · s – 1 ) for a woody species. RUBP - carboxylase content of the leaves ranged from 20 to 22 mg per gram F.W., which is similar to that found in most C 3 crop species. These results suggest that rabbitbrush is able to maintain high rates of P n , at least under nonstressed conditions.           

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.     

Computer analysis of cross sections of leaves of Chrysothamnus taxa and their relation to environmental conditions

Thirty-eight accessions of 20 taxa (species and subspecies) of Chrysothamnus from a range of environments were grown in a uniform garden. During mid-June, terminal and lateral leaves were removed, fixed, and processed for light-microscopy studies. Area and perimeter measurements of ink tracings of midleaf vein cross sections were measured using computer-scan techniques. Leaf area and perimeter were compared with vein perimeter and area measurements. Area and perimeter of these Chrysothamnus leaves varied in response to the environment at the collection location. Plants native to hot, dry conditions had small, thick leaves, whereas plants native to cooler, more mesic conditions had relatively large, thin leaves. Similarly, leaves with round veins were adapted to hot, dry environments and leaves with elliptical veins were adapted to cooler, more mesic environments. Both terminal (young) and lateral (older) leaves were sampled. Terminal leaves showed significant (p < .05) correlational responses to environmental parameters, whereas the lateral leaves, while fronding in the predicted morphological directions, did not generally show significant responses. The different responses of terminal and lateral leaves may have been due to canopy position and uniform environmental conditions of the garden. Leaves with large cross sections characteristically had large veins, and leaves with small cross sections characteristically had small veins. Large leaf cross sections were normally flat. Small leaf cross sections were normally round. Decreasing leaf size and increasing leaf thickness were correlated with aridity.     

Seasonal microhabitat relationships of blue grouse in southeastern Idaho

Microhabitat characteristics of blue grouse ( Dendragapus obscurus ) were analyzed in breeding and wintering habitats in southeastern Idaho. Breeding habitats typically were open sagebrush ( Artemisia spp.), mixed shrub, mountain mahogany ( Cercocarpus ledifolius ) , and maple ( Acer grandidentatum ) stands on east to south facing aspects of slopes below 2100 m elevation. Breeding blue grouse selected areas with approximately a 50:50 or greater open to cover ratio. Blue grouse selected areas with higher tree coverage than available on average within the mixed shrub vegetation type. Hens with broods preferred sites with relatively tall (>50 cm) herbaceous vegetation. During autumn and winter, blue grouse preferred high elevation (>2285 m) stands of open (50% tree cover) conifer. Douglas-fir ( Pseudotsuga menziesii ) were preferred as winter roost trees. Sites selected in winter had significantly more Douglas-fir than those selected in autumn.&nbsp;     

Influence of the holly leaf miner,Phytomyza ilicis (Diptera Agromyzidae), on leaf abscission

Holly trees were sampled at two heights for 11 annual generations of Phytomyza ilicis Curtis. The number of mines per 1000 leaves, mine intensity, was consistently more at the lower height. Mine-induced leaf abscission led to a reduction in mine intensity from 1- to 2-year-old leaves. Abscission was shown to have been greater in samples taken from higher in the tree. There was a suggestion of a density-dependent effect on the probability of mine-induced abscission.     

Tree densities on pinyon-juniper woodland sites in Nevada and California

The densities of sing le leaf pinyon and Utah juniper trees in four diameter classes (1&ndash;9, 10&ndash;19, 20&ndash;29, and &ge; 30 cm) were measured on 522 plots of 1/10 ha each throughout the Great Basin. Density distribution patterns of pinyon and juniper varied with aspect, elevation, and eastern (EGB) versus western Great Basin (WGB) locations. On most locations north and, to a lesser extent, west slopes supported higher densities of pinyon than south and east slopes, with high relative densities of small diameter trees on north slopes and large diameter trees on west slopes. Pinyon densities were higher on EGB than on WGB sites and on higher elevation than on lower elevation sites. Juniper densities were higher on EGB than on WGB sites and on lower elevation than on higher elevation sites. Juniper densities on low-elevation WGB sites were higher on south and west aspects than on north and east, with higher relative densities in the 20&ndash;29 cm diameter class than in other diameter classes. On low elevation EGB sites, east and south slopes supported higher juniper densities than did north and west slopes, with comparatively higher relative densities in the 10&ndash;19 cm diameter class. Differences in relative densities between diameter classes were not significant among aspects on high elevation sites.     

Depletion of soil moisture by two cold-desert bunchgrasses and effects on photoshythetic performance

This study compared the abilities of two cool-season bunchgrasses to extract moisture from a drying soil and compared photosynthetic and stomatal responses of the two species as soil moisture supplies were depleted. When grown in 49-L pots in a greenhouse, Leymus cinereus extracted more water from the soil and maintained higher gas exchange rates to lower absolute amounts of soil water than did Agropyron desertorum . The soil water content at the lower limit of extraction was 10.3% for L. cinereus and 13.3% for A. desertorum . When soil moisture was expressed as extractable soil water, there was little difference between the species in pattern of water use. Both species maintained high stomatal conductances (g w ) and photosynthetic rates (A) until extractable soil moisture was reduced to about 15%. For field-grown plants under severe water stress, A was higher in L. cinereus than in A. desertorum at comparable leaf water potentials. The relationship between A and g w was similar for the two species; higher A in L. cinereus was a consequence of higher g w . Thus, higher A in L. cinereus is achieved through some sacrifice of water-use efficiency.     

Development and longevity of ephemeral and perennial leaves on Artemisia tridentata Nutt. ssp. wyomingensis

Big sagebrush ( Artemisia tridentata Nutt.) is one of the most successful plants in the Great Basin based on its abundance and wide distribution. The development of dimorphic leaves may be an important mechanism attributing to its adaptive and competitive abilities. Development, persistence, and proportions of ephemeral and perennial leaves on Wyoming big sagebrush ( Artemisia tridentata Nutt. ssp. wyomingensis ) were studied for two years. The large ephemeral leaves are the first to develop in early spring. As early developing ephemerals mature and stems elongate, new ephemeral and perennial leaves develop in the axes of these large ephemerals. Perennial leaves expanded in the summer of their first growing season, persisting on the shrub until their abscission during summer drought of the second growing season. Plants maintained 33% of their leaf weight through the winters of 1985 and 1986. Active leaf and stem growth occurred at soil water potentials above &ndash;0.2 MPa.     

Relationship of western juniper stem conducting tissue and basal circumference to leaf area and biomass

The ability to measure leaf area and biomass on a plant community basis has many important ecological applications. These include quantification of gas exchange, use of water resources on the site, nutrient pools, and construction of models simulating production and resource allocation. To test a nondestructive technique for estimating leaf area and leaf biomass of western juniper ( Juniperus occidentalis Hook.), sapwood area and basal circumference were evaluated as predictors of total leaf biomass and leaf area. Nineteen trees, ranging in size from 9.0 to 263 cm in circumference, were destructively sampled. The entire leaf biomass was harvested and measured, and regression equations were developed. Both sapwood area and basal circumference significantly (P < .01) correlated with projected leaf area and leaf biomass (r values = 0.98).     

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.