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.     

Field establishment of fourwing saltbush in processed oil shale and disturbed native soil as influenced by vesicular-arbuscular mycorrhizae

Seedlings of fourwing saltbush ( Atriplex canescens (Pursh) Nutt.) were inoculated with indigenous vesicular-arbuscular mycorrhizal (VAM) fungi in a containerized system and transplanted into processed oil shale and disturbed native soil in a semiarid rangeland environment in northwestern Colorado. After two growing seasons in the field, plants inoculated with VAM had greater aboveground biomass, cover, and height than noninoculated plants. Mycorrhizal plants were more effective in the uptake of water and phosphorus. Infection levels of inoculated plants were greatly reduced in processed shale (from 13.0 at outplanting to 3.8 at harvest), but functional VAM associations could be found after two growing seasons. Results indicate that VAM help make processed oil shale a more tractable medium for the establishment of plants representative of later successional stages by allowing these plants to make effective use of the natural resources that are limiting under conditions of high stress.        

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 –0.2 MPa.     

New taxa and combinations in the Utah flora

New taxa include: Cryptantha cinerea (Torr.) Cronq. var. arenicola Higgins & Welsh; Physaria chambersii Rollins var. sobolifera Welsh (Cruciferae); Phacelia demissa Gray var. minor N. D. Atwood (Hydrophyllaceae); Iris pariensis Welsh (Iridaceae); Astragalus preussii var. cutleri Barneby and Pediomelum aromaticum (Payson) Welsh var. tuhyi Welsh (Leguminosae); Abronia nana Wats. var. harrisii Welsh (Nyctaginaceae); Camissonia atwoodii Cronq. (Onagraceae); Habenaria zothecina Higgins & Welsh (Orchidaceae); Aqiiilegia formosa Fisch. in DC. var. fosteri Welsh (Ranunculaceae). New nomenclatural combinations include: Rhus aromatica Ait. var. simplicifolia (Greene) Cronq. (Anacardiaceae); Lomatium kingii (Wats.) Cronq., L. kingii var. alpinum (Wats.) Cronq. (Apiaceae); Cryptantha cinerea (Torr.) Cronq. var. laxa (Macbr.) Higgins; Mertensia lanceolata (Pursh) DC. var. nivalis (Wats.) Higgins (Boraginaceae); Opuntia erinacea Engelm. var. aurea (Baxter) Welsh (Cactaceae); Arenaria fendleri Gray var. aculeata (Wats.) Welsh, A. fendleri var. eastwoodiae (Rydb.) Welsh, Lychnis apetala L. var. kingii (Wats.) Welsh, Stellaria longipes Goldie var. monantha (Hulten) Welsh (Caryophyllaceae); Draba densifolia Nutt. ex T. & G. var. apiculata (C. L. Hitchc.) Welsh, D. oligosperma Hook. var. juniperina (Dorn) Welsh, Physaria acutifolia Rydb. var. stylosa (Rollins) Welsh, Thelypodiopsis sagittata (Nutt.) Schulz var. ovalifolia (Rydb.) Welsh (Cruciferae); Lotus plebeius (T. Brandg.) Barneby, Lupinus polyphyllus Lindl. in Edwards var. ammophilus (Greene) Barneby, L polyphyllus var. humicola (A. Nels.) Barneby, L. argenteus Pursh var. fulvomaculatus (Payson) Barneby, L. argenteus var. palmeri (Wats.) Barneby, Pediomelum aromaticum (Payson) Welsh, P. epipsilum (Barneby) Welsh, Psoralidium lanceolatum (Pursh) Rydb. var. stenophyllum (Rydb.) Welsh, and P. lanceolatum var. stenostachys (Rydb.) Welsh (Leguminosae); Mirabilis linearis (Pursh) Hiemerl var. decipiens (Standl.) Welsh (Nyctaginaceae); Camissonia boothii var. condensata (Munz) Cronq., C. boothii var. villosa (Wats.) Cronq., C. clavaeformis (Torr. & Frem.) Raven var. purpurascens (Wats.) Cronq., C. scapoidea (T. & G.) var. utahensis (Raven) Welsh, Oenothera caespitosa var. macroglottis (Rydb.) Cronq., Oe. caespitosa var. navajoensis (Wagner, Stockhouse, & Klein) Cronq., Oe. flava (A. Nels.) Garrett var. acutissima (W. L. Wagner) Welsh, and Oe. primiveris Gray var. bufonis (Jones) Cronq. (Onagraceae); Papaver radicatum Rottb. var. pygmaeum (Rydb.) Welsh (Papaveraceae); Dodecatheon pulchellum (Raf.) Merr. var. zionense (Eastw.) Welsh (Primulaceae); Aquilegia flavescens Wats. var. rubicunda (Tidestr.) Welsh, Delphinium andersonii Gray var. scaposum (Greene) Welsh, D. occidentalis (Wats.) Wats. var. barbeyi (Huth) Welsh, and Ranunculus andersonii Gray var. juniperinus (Jones) Welsh (Ranunculaceae); Purshia mexicana (D. Don) Welsh and P. mexicana var. stansburyi (Torr.) Welsh (Rosaceae); Galium mexicanum H.B.K. var. asperrimum (Gray) Higgins & Welsh (Rubiaceae); Castilleja parvula Rydb. var. revealii (N. Holmgren) N. D. Atwood and C. rhexifolia Rydb. var. sulphurea (Rydb.) N. D. Atwood (Scrophulariaceae).     

Utah plant types—historical perspective 1840 to 1981—annotated list, and bibliography

New taxa include: Cryptantha cinerea (Torr.) Cronq. var. arenicola Higgins & Welsh; Physaria chambersii Rollins var. sobolifera Welsh (Cruciferae); Phacelia demissa Gray var. minor N. D. Atwood (Hydrophyllaceae); Iris pariensis Welsh (Iridaceae); Astragalus preussii var. cutleri Barneby and Pediomelum aromaticum (Payson) Welsh var. tuhyi Welsh (Leguminosae); Abronia nana Wats. var. harrisii Welsh (Nyctaginaceae); Camissonia atwoodii Cronq. (Onagraceae); Habenaria zothecina Higgins & Welsh (Orchidaceae); Aqiiilegia formosa Fisch. in DC. var. fosteri Welsh (Ranunculaceae). New nomenclatural combinations include: Rhus aromatica Ait. var. simplicifolia (Greene) Cronq. (Anacardiaceae); Lomatium kingii (Wats.) Cronq., L. kingii var. alpinum (Wats.) Cronq. (Apiaceae); Cryptantha cinerea (Torr.) Cronq. var. laxa (Macbr.) Higgins; Mertensia lanceolata (Pursh) DC. var. nivalis (Wats.) Higgins (Boraginaceae); Opuntia erinacea Engelm. var. aurea (Baxter) Welsh (Cactaceae); Arenaria fendleri Gray var. aculeata (Wats.) Welsh, A. fendleri var. eastwoodiae (Rydb.) Welsh, Lychnis apetala L. var. kingii (Wats.) Welsh, Stellaria longipes Goldie var. monantha (Hulten) Welsh (Caryophyllaceae); Draba densifolia Nutt. ex T. & G. var. apiculata (C. L. Hitchc.) Welsh, D. oligosperma Hook. var. juniperina (Dorn) Welsh, Physaria acutifolia Rydb. var. stylosa (Rollins) Welsh, Thelypodiopsis sagittata (Nutt.) Schulz var. ovalifolia (Rydb.) Welsh (Cruciferae); Lotus plebeius (T. Brandg.) Barneby, Lupinus polyphyllus Lindl. in Edwards var. ammophilus (Greene) Barneby, L polyphyllus var. humicola (A. Nels.) Barneby, L. argenteus Pursh var. fulvomaculatus (Payson) Barneby, L. argenteus var. palmeri (Wats.) Barneby, Pediomelum aromaticum (Payson) Welsh, P. epipsilum (Barneby) Welsh, Psoralidium lanceolatum (Pursh) Rydb. var. stenophyllum (Rydb.) Welsh, and P. lanceolatum var. stenostachys (Rydb.) Welsh (Leguminosae); Mirabilis linearis (Pursh) Hiemerl var. decipiens (Standl.) Welsh (Nyctaginaceae); Camissonia boothii var. condensata (Munz) Cronq., C. boothii var. villosa (Wats.) Cronq., C. clavaeformis (Torr. & Frem.) Raven var. purpurascens (Wats.) Cronq., C. scapoidea (T. & G.) var. utahensis (Raven) Welsh, Oenothera caespitosa var. macroglottis (Rydb.) Cronq., Oe. caespitosa var. navajoensis (Wagner, Stockhouse, & Klein) Cronq., Oe. flava (A. Nels.) Garrett var. acutissima (W. L. Wagner) Welsh, and Oe. primiveris Gray var. bufonis (Jones) Cronq. (Onagraceae); Papaver radicatum Rottb. var. pygmaeum (Rydb.) Welsh (Papaveraceae); Dodecatheon pulchellum (Raf.) Merr. var. zionense (Eastw.) Welsh (Primulaceae); Aquilegia flavescens Wats. var. rubicunda (Tidestr.) Welsh, Delphinium andersonii Gray var. scaposum (Greene) Welsh, D. occidentalis (Wats.) Wats. var. barbeyi (Huth) Welsh, and Ranunculus andersonii Gray var. juniperinus (Jones) Welsh (Ranunculaceae); Purshia mexicana (D. Don) Welsh and P. mexicana var. stansburyi (Torr.) Welsh (Rosaceae); Galium mexicanum H.B.K. var. asperrimum (Gray) Higgins & Welsh (Rubiaceae); Castilleja parvula Rydb. var. revealii (N. Holmgren) N. D. Atwood and C. rhexifolia Rydb. var. sulphurea (Rydb.) N. D. Atwood (Scrophulariaceae).     

Vegetation response to prescribed fire in Dinosaur National Monument

Much of western North America is dominated by dense, monotypic, late seral stands of big sagebrush ( Artemisia tridentata Nutt.). These stands often have depauperate understories with limited species richness, diversity, and herbaceous cover. The National Park Service at Dinosaur National Monument, Colorado, is using both strategic and natural prescribed fire in Wyoming big sagebrush ( Artemisia tridentata ssp. wyomingensis Beetle and Young) communities to foster intra-community (α -scale) and landscape diversity. This study analyzed an accumulated foliar cover data set between paired burn and control areas on 6 different sites during the last 20 years. Across the monitoring period, mean total vegetation cover of all combined sites was 44% control and 42% burn. Total vegetation cover in burn areas was higher than or equal to paired control areas within 2-3 years post-burn. Shrubs were essentially eliminated in burn areas, but perennial grass cover was 10-35% higher. Mean number of species on all sites and years combined was 17 control and 18 burn. Species richness was different on only 1 site-year, Dry Woman 1995 ( P = 0.001, 15 control, 9 burn). Species similarity by site and between treatments ranged from 44% to 75%. Differences in Shannon-Weiner diversity index values between paired sites occurred in 6 of 20 years ( P < 0.05). Index value differences on these 6 sites were due to a large annual grass component in burn areas. Prescribed burning successfully shifted late successional sagebrushdominated communities to earlier herbaceous-dominated successional stages without lowering total vegetation cover, while maintaining -scale diversity and species richness.     

Bitterbrush ( Purshia tridentata Pursh) growth in relation to browsing

The objectives of this study were to compare vegetative and reproductive growth characters of bitterbrush ( Purshia tridentata Pursh) stands as they relate to browsing levels. Growth characters were measured on 10 ecologically diverse stands in southwestern Montana on which browsing ranged from 0% to 60% of all current annual long shoot (LS) growth. Bitterbrush plants exhibited both twig-level and plant-level responses to browsing. Total bud density per plant was similar for browsed and unbrowsed sites, but differed ( P P P P P < 0.05). Our data suggest that long-term unbrowsed plants should not be used as a standard for comparison with normally browsed plants.     

Suitability of shrub establishment on Wyoming mined lands reclaimed for wildlife habitat

Restoring coal mined land to pre-mining shrub cover, density, height, community composition, and diversity to renew wildlife habitat quality is a priority for reclamation specialists. Long-term shrub reestablishment success on reclaimed mined land in Wyoming and suitability of these lands for wildlife habitat are unknown. Fourteen reclaimed study sites, 10 yr old or older, were selected on 8 mines in Wyoming to evaluate shrub reestablishment and wildlife habitat value for antelope ( Antilocapra americana ) and sage grouse ( Centrocercus urophasianus ). Five sites were categorized as fourwing saltbush ( Atriplex canescens ) sites and 9 as fourwing saltbush/big sagebrush ( A. canescens/Artemisia tridentata spp. wyomingensis ) sites. Published data describing antelope and sage grouse-preferred habitat requirements in sagebrush-grassland steppe ecosystems were used to evaluate shrub community value of sampled sites for wildlife habitat. Mean shrub canopy cover, density, and height for fourwing saltbush sites were 5.8%, 0.23 m -2 , and 41.6 cm, respectively, compared to 5.6%, 0.61 m -2 , and 31.1 cm for fourwing saltbush/big sagebrush sites. Two fourwing saltbush and 4 fourwing saltbush/big sagebrush sites provided sufficient cover for antelope, while 2 fourwing saltbush and 4 fourwing saltbush/big sagebrush sites were adequate for sage grouse. Only 1 fourwing saltbush/big sagebrush site provided high enough shrub densities for sage grouse. One fourwing saltbush and 7 fourwing saltbush/big sagebrush sites provided ample shrub heights for antelope, while 1 fourwing saltbush and 8 fourwing saltbush/big sagebrush sites were sufficient for sage grouse. One fourwing saltbush and 1 fourwing saltbush/big sagebrush site provided enough grass, forb, and shrub composition for antelope, while no site in either reclamation type was satisfactory for sage grouse. Shrub diversity was 3 times higher for fourwing saltbush/big sagebrush sites (0.984) than for fourwing saltbush sites (0.328). Individually, sites seeded with multiple shrub species had higher canopy cover, density, and diversity compared with single-species shrub seedings. Achieving premining shrub cover, density, height, community composition, and diversity within existing bond-release time frames is unrealistic, considering that some native shrublands require 30-60 yr to reach maturity.     

Forage quality of rillscale ( Atriplex suckleyi ) grown on amended bentonite mine spoil

At peak standing crop, rillscale ( Atriplex suckleyi ) foliage grown on amended bentonite mine spoil contained adequate digestible energy, crude protein, and all mineral elements except phosphorus necessary for cattle, sheep, antelope, and deer. Amendments (sawdust, NPK, gypsum) generally did not affect forage quality. Iron, manganese, aluminum, sodium, and potassium concentrations were high and may have adversely affected forage quality. Forage utility would be limited to a few months during the growing season.     

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.     

Controlling performance of bluebunch wheatgrass and spotted knapweed using nitrogen and sucrose amendments

Range- and wildlands are being invaded by nonindigenous plants, resulting in an unprecedented, rapid change in plant community composition across the United States. Successional management predicts that species performance may be modified by resource availability. The objective of this study was to determine whether species performance could be altered by modifying soil nitrate (NO 3 – ) and ammonium (NH 4 + ) concentrations within an Idaho fescue ( Festuca idahoensis )/bluebunch wheatgrass ( Pseudoroegneria spicata ) plant association. We planted bluebunch wheatgrass and spotted knapweed ( Centaurea maculosa ) in an addition series at 2 sites in southwestern Montana. Each plot in the addition series matrix was divided into thirds, and we applied nitrogen (N) to a subplot and sucrose to a 2nd subplot. The remaining subplot was not amended and considered a control. Nitrogen amendment tended to enhance the performance of spotted knapweed, while sucrose amendment had no effect. Bluebunch wheatgrass performance was not affected by either amendment. Sucrose treatments only decreased soil NO 3 – at the more productive site. Regression models for predicting bluebunch wheatgrass and spotted knapweed biomass accounted for only about 30% of the variation, suggesting other processes in addition to interference were responsible for explaining relative plant perfor - mance. We recommend that land managers prevent activities that increase soil N concentration while the effectiveness of carbon amendments as a means to decrease soil N concentrations and shift interference relationships is further investigated.     

Recovery and germinability of native seed fed to cattle

Using livestock as seed dispersal agents may be an effective method for increasing species diversity on degraded and previously seeded rangelands. We quantified seed passage and recovery rates, and post-passage germinability of Wyoming big sagebrush ( Artemisia tridentata Nutt. ssp. wyomingensis Beetle & Young), bottlebrush squirreltail ( Elymus elymoides [Raf.] Swezey), and gooseberry globemallow ( Sphaeralcea grossulariaefolia [H. & A.] Rydb.) by feeding Holstein heifers seeds of each species at 3 levels (15,000; 30,000; and 60,000 seeds) over a period of 3 weeks. One-kg fecal samples were collected 1, 2, 3, and 4 days after seed ingestion. Undamaged seeds were extracted from the samples and tested for germinability. Globemallow had the highest percentage of recovered, undamaged seed, followed by squirreltail and sagebrush. Globemallow and sagebrush seed passage was highest on Day 1, after which seed numbers dropped sharply. Squirreltail passage and recovery were more consistent through time, with higher seed recovery at lower seed feeding levels. Post-passage germinability was highest for squirreltail and globemallow on Day 1. Sagebrush germination was negligible. Differences in physical seed properties (size, shape, and seed coat) likely influenced interspecies variation in passage, recovery, and germinability. Globemallow and squirreltail seeds may be suited for livestock dispersal, but sagebrush seeds are not.     

Plant age/size distributions in black sagebrush ( Artemisia nova ): effects on community structure

The demography of black sagebrush ( Artemisia nova Nelson) was investigated in the Buckskin Mountains of western Nevada to determine patterns of stand renewal in sagebrush communities currently free from wildfires. Biomass sampling was conducted to develop growth classes that reflected apparent age of the shrubs. The density of black sagebrush plants was twice that of basin big sagebrush ( A. tridentata ssp. tridentata Nutt.) in adjacent communities on contrasting soils (2.2 versus 1.1 plants per m 2 ). Black sagebrush accumulated only 75% as much woody biomass as big sagebrush. Regression equations were developed and tested for predicting total woody biomass, current annual growth (CAG), and leaf weight of black sagebrush plants. Apparent age classes were developed both for the black sagebrush plants and the sub-canopy mounds on which they grew. Discriminant analysis was used to test this classification system. Plant succession, apparently controlled by nitrate content of the surface soil, appeared to eliminate the successful establishment of black sagebrush seedlings on the mounds. After the shrubs die, the mounds eventually deflate. We propose that mounds reform around shrub seedlings, but because seedling establishment is so rare in these communities, this could not be verified.     

Long-term vegetation dynamics in a cut western juniper woodland

Western juniper ( Juniperus occidentalis spp. occidentalis Hook.) expansion in the northern Great Basin has reduced shrubsteppe productivity and diversity. Chainsaw cutting of western juniper woodlands is commonly applied to remove tree interference and restore sagebrush plant communities. Studies assessing understory response following cutting have been limited to early successional stages and have not evaluated the effects of western juniper debris on plant succession. Cutting western juniper produces a large amount of debris which is commonly left on site, occupying a significant portion of treated areas. This study evaluated successional dynamics spanning 13 years after western juniper cutting. Four 0.45-ha blocks were selected on Steens Mountain in southeastern Oregon. Western juniper cover averaged 26% and mature tree density averaged 250 trees ? ha –1 . Blocks were cut in late summer 1991. Understory standing crop, cover, and density were compared among 3 locations: old canopy litter mats (canopy), interspace, and area underneath cut western juniper (debris). In the interspace, perennial grasses increased in cover and in standing crop relative to other functional groups. In canopy and debris locations, species composition shifted in the 6th year after cutting as annual grass cover, density, and standing crop increased. However, by 2003, perennial grass biomass was 2 times greater than annual grass biomass in canopy and debris locations. Because annual grasses increased in areas of debris accumulation, managers need to be cognizant of western juniper treatments that create safe sites that are favorable to the establishment of weedy species. Retaining western juniper debris on this site did not increase establishment and growth of perennial grasses compared to the interspace. A shift in perennial grass dominance from Thurber's needlegrass ( Achnatherum thurberianum [Piper] Barkworth) to bottlebrush squirreltail ( Elymus hystrix [Nutt.] Smith) occurred in areas of debris accumulation. Our results demonstrated that long-term vegetation evaluations are necessary to properly assess management activities and disturbance.     

Pollination needs of arrowleaf balsamroot, Balsamorhiza sagittata (Heliantheae: Asteraceae)

Arrowleaf balsamroot, Balsamorhiza sagittata (Pursh) Nutt, is a common, sometimes dominant, longlived forb that flowers early in spring from the foothills to upper-montane areas of the northern Rocky Mountains and Intermountain West. Public land managers desire its seed for rangeland rehabilitation. Through manual pollination field trials, the species was found to have a mixed pollination system. It is primarily xenogamous (46% of ovules yielded plump achenes) but partially self-compatible (31% of achenes were plump). Unvisited flower heads formed virtually no mature achenes; only plump achenes contained seeds with endosperm. Freely visited flower heads in 2 populations produced as many achenes as manual outcross pollinations of flower heads, suggesting that seed production was not pollinator limited. Two species of Osmia bees rely mostly on Balsamorhiza and its close relative, Wyethia , for pollen. At least 165 females per hectare will need to be stocked to achieve thorough flower visitation in cultivated seed production fields.