Classification and ordination of Coleogyne communities in southern Nevada

Woody plant community composition was analyzed throughout the range of Coleogyne ramosissima in the Spring and Sheep Mountain ranges of southern Nevada. The lower Coleogyne elevational boundary was analyzed in detail in Lucky Strike Canyon, on the eastern edge of the Spring Mountains. TWINSPAN (2-way indicator species analysis) identified 4 primary species and stand groups from the 2 mountain ranges (extensive survey). Analysis of DECORANA (detrended correspondence analysis) results indicated that elevation and soil depth were the environmental factors most significantly associated with distribution of species and stand groups in the extensive survey. Elevation was the only significant physical factor associated with distribution of species and stand groups in the intensive survey. Five vegetation zones form the 2 mountain ranges were identified based on their dominant species in 15 extensive transects. Coleogyne were subdivided into pure stands and upper and lower ecotones for further investigation of species distribution and environmental factors. Selected environmental factors appear to play an important role in structuring the Mojave Desert vegetation zones in southern Nevada.     

Environmental attributes correlating with density of blackbrush ( Coleogyne ramosissima ) shurbs in the Spring Mountains of southern Nevada

Discovery of distinct mid-elevational bands of blackbrush ( Coleogyne ramosissima Torr.) shrublands on desert mountain slopes in the Mojave Desert caused an investigation of the relationships between environmental factors and Coleogyne distribution. Environmental factors were quantitatively examined to determine which were significant predictors of Coleogyne density at upper-elevational limits (ecotones) in the Spring Mountains of southern Nevada. Path analysis revealed significant, direct causal effects of air temperature, soil moisture, soil depth, and percent litter cover on the distribution of Coleogyne . Specifically, air temperature was a significant positive predictor, while soil moisture, soil depth, and percent litter cover were significant negative predictors of Coleogyne density, with the effects of other environmental variables parceled out. Path analysis also indicated that indirect effects of soil pH, bulk density, compaction, percent pore space, organic matter, soil temperature, salinity, cryptogam, and percent bare soil and rock cover on Coleogyne density were substantially more potent than their direct casual effects. Environmental attributes associated with elevational changes correlate with and may determine the density of Coleogyne shrubs at upper ecotones in southern Nevada.     

Ecological impacts of Pogonomyrmex on woody vegetation of a Larrea-Ambrosia shrubland

Discovery of distinct mid-elevational bands of blackbrush ( Coleogyne ramosissima Torr.) shrublands on desert mountain slopes in the Mojave Desert caused an investigation of the relationships between environmental factors and Coleogyne distribution. Environmental factors were quantitatively examined to determine which were significant predictors of Coleogyne density at upper-elevational limits (ecotones) in the Spring Mountains of southern Nevada. Path analysis revealed significant, direct causal effects of air temperature, soil moisture, soil depth, and percent litter cover on the distribution of Coleogyne . Specifically, air temperature was a significant positive predictor, while soil moisture, soil depth, and percent litter cover were significant negative predictors of Coleogyne density, with the effects of other environmental variables parceled out. Path analysis also indicated that indirect effects of soil pH, bulk density, compaction, percent pore space, organic matter, soil temperature, salinity, cryptogam, and percent bare soil and rock cover on Coleogyne density were substantially more potent than their direct casual effects. Environmental attributes associated with elevational changes correlate with and may determine the density of Coleogyne shrubs at upper ecotones in southern Nevada.     

Survival and development of Phoradendron californicum and Acacia greggii during a drought

Soil compaction from human trampling, biking, and off-road motor vehicle traffic was quantitatively investigated in a blackbrush ( Coleogyne ramosissima ) shrubland in Kyle Canyon of the Spring Mountains in southern Nevada. A significant difference was detected in soil compaction, bulk density, and percent pore space at a particular frequency of visits in each of 4 disturbance types. On average a single vehicle pass was equivalent to 10 human footprints. Ten and 100 footprints were equivalent to 1 motorcycle pass and 10 vehicle passes, respectively. Soil compaction is a product of increased bulk density and decreased pore space. The degree of soil compaction is a function of disturbance type and visit frequency when examining these 2 factors independently. However, interactive effects of disturbance type and visit frequency on soil bulk density, compaction, and percent pore space were not significantly different. The greatest effects occurred during the first few passes, with changes per pass decreasing as the number of passes increased in all 4 trails. Results of this study suggest that the effects of hiking and biking slowly increase over time relative to the effects of motor vehicle traffic in the Coleogyne shrubland of Kyle Canyon in southern Nevada.     

Soil compaction from human trampling, biking, and off-road motor vehicle activity in a blackbrush ( Colleogyne ramosissima ) shrubland

Soil compaction from human trampling, biking, and off-road motor vehicle traffic was quantitatively investigated in a blackbrush ( Coleogyne ramosissima ) shrubland in Kyle Canyon of the Spring Mountains in southern Nevada. A significant difference was detected in soil compaction, bulk density, and percent pore space at a particular frequency of visits in each of 4 disturbance types. On average a single vehicle pass was equivalent to 10 human footprints. Ten and 100 footprints were equivalent to 1 motorcycle pass and 10 vehicle passes, respectively. Soil compaction is a product of increased bulk density and decreased pore space. The degree of soil compaction is a function of disturbance type and visit frequency when examining these 2 factors independently. However, interactive effects of disturbance type and visit frequency on soil bulk density, compaction, and percent pore space were not significantly different. The greatest effects occurred during the first few passes, with changes per pass decreasing as the number of passes increased in all 4 trails. Results of this study suggest that the effects of hiking and biking slowly increase over time relative to the effects of motor vehicle traffic in the Coleogyne shrubland of Kyle Canyon in southern Nevada.     

Marmot distribution and habitat associations in the Great Basin

In this study I describe the distribution and habitat associations of yellow-bellied marmots ( Marmota flaviventris ) in the Great Basin, compare my findings with those of E.R. Hall during his 1929-1936 survey and later surveys, and discuss potential reasons for changes in marmot distribution over time. I found 62 marmot burrow sites in 18 mountain ranges, mostly in rocky meadows situated on well-drained slopes between 2100 m and 3000 m elevation. Marmots were generally found near burrows dug within talus slopes, talus-like rock piles, or clusters of massive boulders. Oceanspray ( Holodiscus discolor ) was the shrub most commonly associated with occupied rock formations. Marmots were most abundant in the Ruby/East Humboldt Range and were common in the Desatoya, Shoshone, Toiyabe, Toquima, Cherry Creek, Schell Creek, Deep Creek, and Stansbury Ranges. Marmots appeared to be uncommon in the Monitor Range and rare in the Clan Alpine, Roberts, and Snake Ranges. I was unable to find marmots in the Diamond, Egan, Spruce-Pequop, White Pine, and Oquirrh Ranges, although I located old, weathered marmot scats in all but the latter 2 ranges. Other evidence confirms that marmots do actually occur in the Oquirrh Range, but extensive searches of the White Pine Range, including some of the same rock formations where E.R. Hall collected marmots, revealed no sign of marmots. My distribution data suggest that marmots may have gone extinct in some Great Basin mountain ranges during the last century. These disappearances may represent a natural extinction-recolonization dynamic, but a more alarming possibility is a recent die-off linked to climate change, which is predicted to force montane vegetation zones further upslope, shrinking the habitat of associated faunas. However, marmots in this study were observed as low as 1550 m elevation, indicating an altitudinal flexibility that may allow this species to survive climatic change better than more specialized boreal species such as pikas ( Ochotona princeps ) and water shrews ( Sorex palustris ).     

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.     

Variation in germination response to temperature and water availability in blackbrush ( Coleogyne ramosissima ) and its ecological significance

Blackbursh ( Coleogyne ramosissima Torr.) is a dominant desert shrub in a distinct mid-elevations vegetation belt between creosote bush-bursage ( Larrea tridentata-Ambrosia dumosa ) shrubland below and big sagebrush-pinyon pine- Utah juniper ( Artemisia tridentata-Pinus monophylla-Juniperus osteosperma ) woodland above in the Mojave Desert. Seed germination patterns of blackbrush seeds collected from 2 elevations (1200 and 1550 m) in 5 mountain ranges within the blackbrush shrublands were investigated. Morphological features of blackbrush seeds, including weight, length, and width, were not significantly different ( P > 0.05) among elevations and mountain ranges in the Mojave Desert. Germination of blackbrush seeds was optimal when preceded by a prechill period of 4-6 wk. Seeds incubated at room temperature germinated poorly. Seeds collected at warm, low-elevation sites appeared to be less dormant (required less prechill time), germinated faster, and showed a higher overall germination response; watering at 2-wk intervals revealed the greatest germination. Some ecotypic variation among populations establishing at different elevations was evident with regard to dormancy duration and germination response at certain constant temperatures.     

Stonefly (Plecoptera) records from the basin ranges of Nevada and Utah

Distributional records are given for 40 stonefly species on 15 isolated mountain ranges in Nevada and Utah.     

Zoogeographic affinities of the stoneflies (Plecoptera) of the Raft River Mountains, Utah

We examined faunal affinities of the Raft River Mountains using stoneflies (Plecoptera) as indicators. This island-like mountain range is isolated from other major mountain ranges in the Intermountain West by low-elevation, arid regions. Thirty-seven species were recorded from collections from 19 sites in the Raft River Mountains. Cluster analysis demonstrated the Raft River Mountain stonefly assemblage to be most similar to faunas of the Sawtooth and Wasatch mountains, and quite different from that of the Sierra Nevada. An analysis of the distribution patterns of each species, on a family-by-family basis, showed that the Raft River Mountains fauna consists mostly of species widespread in western North America. Most families were represented by at least 1 species whose distribution supports faunal affinities with regions to the north and west. Logistic regression of 6 long-distance dispersal factors against stonefly presence-absence data did not support long-distance dispersal as a viable means of colonization for the Raft River Mountains. This suggest that stonefly distribution patterns may be attributed to expansion and subsequent vicariance of suitable stonefly habitats during Pleistocene climatic oscillations.     

Factors influencing anuran distribution in coastal dune wetlands in southern Brazil

We investigated the potential role of hydroperiod and habitat structural complexity as explanatory factors defining richness, abundance and spatial and temporal distribution of anurans in wetlands of coastal dunes. This survey was performed in 15 wetlands along the Atlantic coastal zone of southern Brazil. We identified 10 anuran species (nine in the adult and eight in the tadpole stage) distributed in seven genera from five families. The adult richness and abundance varied among the hydroperiod classes, but without temporal variations. Both tadpole richness and abundance varied temporally and both were influenced by hydroperiod. Adult anuran composition was associated with dry months and wetland area, whereas tadpole composition was associated with emergent and floating macrophytes, wetland area and vegetation cover. This study identified the importance of habitat structural complexity and hydroperiod in spatial–temporal distribution models of anurans.     

Beetles from the environs of Lake Powell in southern Utah and northern Arizona

Sixty-eight species of beetles of 14 families were collected in pit traps in 12 major vegetative communities during environmental monitoring studies in the summers of 1971 to 1973 in southern Utah and northern Arizona. Seasonal and plant community differences and correlations in population and composition were noted for the 16 sites studied. Highest populations and the most species were found in a grass community, but occurred in different months in each of the three years. Lowest populations were found in a Coleogyne community, and fewest species in an Artemisia- grass community.     

Biogeographic and taxonomic relationships among the mountain snails (Gastropoda: Oreohelicidae) of the central Great Basin

Described here are 4 species of mountain snails, Oreohelix , isolated on mountains in the central Great Basin of Nevada and Utah since the end of the Pleistocene. Forty-three mountains were searched during an 18-year period, resulting in 24 mountains found with no oreohelicids present. One population, Oreohelix loisae (19 mm to 23 mm in shell diameter), is described here as a new species related to, but geographically isolated from, the species Oreohelix nevadensis (17 mm to 22 mm diameter). Oreohelix loisae is present only in the Goshute Mountains while O. nevadensis is represented in 3 geographically adjacent ranges in the central Great Basin. These 2 species are possibly related to the Oreohelix haydeni group from the northern Wasatch Range. The subspecies Oreohelix strigosa depressa (15 mm to 21 mm diameter) is present on 11 ranges from western Utah west to east central Nevada. This subspecies is closely related to populations found today in the northern Wasatch Mountains of Utah. The smallest species in diameter (8 mm to 14 mm), Oreohelix hemphilli , is centered in the central Great Basin and found on 16 ranges often in sympatry with 1 or 2 of the larger conspecifics. Both qualitative and quantitative information on shell characters and soft anatomy is provided here for these 4 species. Shell characters, soft anatomy, geographical isolation, and statistical analysis suggest that 4 distinct species inhabit the central Great Basin today. Xeric and calciphilic species include O. hemphilli and O. loisae , while O. strigosa and O. nevadensis typically are associated with permanent water and both metamorphic and limestone mountains.     

Examining ecological consequences of feral horse grazing using exclosures

Although feral horses have inhabited western North America since the end of the 16th century, relatively little synecological research has been conducted to quantitatively characterize how they interact with ecosystem components. Because feral horses exhibit watering behavior markedly different from that of domestic cattle, it is particularly important to evaluate response of ecosystem elements near water sources to horse use. To assess this response, we performed live-trapping of small mammals and 2-tiered vegetative sampling in 2 mountain ranges in central Nevada in the interior Great Basin, USA. At low elevations, plots around horse-excluded springs exhibited notably greater plant species richness, percent cover, and abundance of grasses and shrubs, as well as more small mammal burrow entrances than plots at horse-grazed springs. At high elevations, meadows protected from grazing exhibited maximum vegetation heights 2.8 times greater than vegetation grazed by horses only and 4.5 times greater than vegetation grazed by horses and cattle. Species richness in quadrats was most different between the horse-and-cattle-grazed meadow and its ungrazed counterpart, suggesting the possibility of synergistic effects of horse and cattle grazing in the same location. This study, the first in the Great Basin to investigate quantitatively ecosystem consequences of feral horse use with exclosures, represents a preliminary step in identifying factors that determine the magnitude of horse grazing impacts.     

Factors controlling structural and floristic variation of riparian zones in a mountainous landscape of the western United States

We examined landscape patterns in the physical conditions and vegetative composition of montane riparian zones to identify their most important sources of variation. Information on plant species cover and on physical characteristics that occur at coarse, medium, and fine scales was collected for 144 riparian plots located throughout the Lake Tahoe Basin, which straddles the California-Nevada border in the western United States. Constrained and unconstrained ordination analyses were used to identify the most important correlates of physical form and plant species composition. Through multivariate analysis of environmental variables (principal components analysis), vegetation data (detrended correspondence analysis), and the combined relationship between the environmental and vegetation data (canonical correspondence analysis), we consistently found that the greatest variation occurred along a gradient of decreasing valley width, decreasing stream sinuosity, and increasing stream slope. Although surface characteristics reflected a 2nd important source of variation in physical conditions, plant species distribution was not strongly correlated with riparian surface conditions. Strong correlations among physical variables that occur at different scales, such as between valley form and geofluvial surface and between geofluvial surface and surface conditions, support the use of a physically based hierarchical framework for organizing riparian zones within the landscape. Such a hierarchical framework would be useful for interpreting patterns in riparian structure and process at different scales and could be applied to riparian zones in other mountain landscapes of the western United States and elsewhere. Moreover, our finding that riparian plant species composition is most strongly correlated with environmental variables that occur at coarse to moderate scales, most of which can be derived from existing data, supports the idea that modeling montane riparian community distribution using topographic and remotely sensed data could be useful; however, a large degree of species variation, unexplained by the variables we collected, indicates that other variables, perhaps disturbance regime, should be included in such a venture. Nomenclature: Hickman (1993).     

Nesting ecology of Yellow Warblers ( Dendroica petechia ) in montane chaparral habitat in the northern Sierra Nevada

The Yellow Warbler ( Dendroica petechia ) is considered a riparian specialist in much of western North America, but in California it also breeds in a second habitat type: montane chaparral of the Sierra Nevada and southern Cascades. We monitored Yellow Warbler nests in montane chaparral and assessed their poorly known nesting ecology in this habitat. We also conducted point counts in upland habitat throughout the region. We determined habitat associations for Yellow Warblers based on nest site and point-count vegetation data; nests were predominantly in bush chinquapin ( Chrysolepis sempervirens ) and greenleaf manzanita ( Arctostaphylos patula ), and point-count abundance was most strongly associated with high overall shrub cover. The importance of montane chaparral for a number of shrub-dependent Sierran birds is well documented, yet the chaparral is threatened by various practices including fire suppression, closed-canopy-focused forest management, and understory fuels-reduction treatments. Although Yellow Warblers are far more abundant in wet mountain meadow riparian habitat in the region, we recommend that management of montane chaparral habitat consider this species and the requirements of other shrub-nesting birds.     

Vertical distribution of adult fish in Pyramid Lake, Nevada

Inshore and limnetic vertical distribution of adult fish in Pyramid Lake, Nevada was determined from late spring to early fall 1977. The benthic (23 m) inshore area exhibited a relatively diverse composition of five species, while the inshore surface and offshore limnetic zones were composed of over 98 percent tui chubs ( Gila bicolor ). Vertical fish distribution was associated with temperature and zooplankton distribution.     

Tamarix aphylla : a newly invasive tree in southern Nevada

In the southwestern United States, the nonnative athel pine ( Tamarix aphylla ) was presumed to be sterile and therefore not as likely to spread as its widely distributed, nonnative congener, T. ramosissima . However, at Lake Mead National Recreation Area (LMNRA) in southern Nevada, populations of T. aphylla have recently spread beyond their limited pre-1990 distribution and now form extensive monospecific stands. Over a 3-year period, we quantified seed production and germination from 60 T. aphylla trees at LMNRA. The annual mean seed production period was 50.6 days, and the mean potential germination (under laboratory conditions) was 22%, indicating that T. aphylla trees at LMNRA are capable of sexual reproduction in southern Nevada. No seeds germinated in field experiments, apparently because of high soil salt levels. However, seedling regeneration is becoming increasingly common at LMNRA. Tamarix aphylla trees occupied a distinct zone along the shoreline of Lake Mead, above T. ramosissima in elevation and below native Larrea tridentata communities, suggesting either competitive exclusion or differential resource utilization. The T. aphylla zone tended to have the highest mean values for total vegetation cover, leaf litter depth, soil salinity, soil moisture, pH, total Kjeldahl nitrogen, and soil organic matter. The capacity for sexual reproduction of this alien plant, combined with a suite of characteristics shared with the invasive T. ramosissima (e.g., drought tolerance and copious saline leaf litter), makes T. aphylla a potentially invasive species along the shores of LMNRA and other mesic areas in the desert Southwest.     

Plant community patterns in unburned and burned blackbrush ( Coleogne ramosissima Torr.) shrublands in the Mojave Desert

The blackbrush vegetation type is dominated by Coleogyne ramossisima , which is thought to preclude the coexistence of many other plant species. Fire can remove blackbrush cover and possibly increase plant species richness and evenness. Fire also may increase the frequency and cover of alien annual grasses, thereby intensifying landscape flammability. We tested these predictions in unburned and burned (6-14 years postfire) blackbrush at 3 sites spanning the range of this vegetation type in the Mojave Desert. Species richness in unburned blackbrush was similar to published values for vegetation types in western North America, bur richness varied significantly among the 3 sites and 4 spatial scales (1, 10, 100, and 1000 m 2 ). Richness values declined in order from annual forbs, woody perennials, herbaceous perennials, annual grasses, cacti, to perennial grasses. Fire reduced Coleogyne cover, thus boosting species evenness. In contrast, species richness decreased after burning, although the results varied among spatial scales. Total cover was unaffected by fire because cover of wood perennials decreased, while cover of annual forbs, annual grasses, herbaceous perennials, and perennial grasses increased. Native species richness and cover decreased, whereas alien richness and cover increased after burning especially where the alien forb Erodium cicutarium was present. Fire had no effect on frequency and variable effects on cover of alien annual grasses. These results indicate that in blackbrush species richness can vary among sites and local spatial scales, and effects of fires can vary among plant life-forms and between natives and aliens.     

Coexistence of two species of sucker, Catostomus , in Sagehen Creek, California, and notes on their status in the western Lahontan Basin

The observed distribution and relative abundance of two morphologically similar species of sucker, Catostomus , have shifted dramatically over the past four decades in Sagehen Creek and nearby streams in eastern California. The mountain sucker, C. platyrhynchus , formerly abundant and more numerous than the Tahoe sucker, C. tahoensis , has become relatively rare and during this study was consistently less abundant than the Tahoe sucker at all eastern California sites in 1983. Similar shifts in abundance were not seen at the three Nevada sites. Behavioral observations and data on spatial and temporal patterns of habitat use, collected in Sagehen Creek between May and September 1982 and 1983 using a snorkel survey method, indicate nearly complete overlap between mountain and Tahoe sucker habitat use and an absence of any agonistic behavioral interaction between species. The decline of the mountain sucker in these areas is likely the result of an interaction of loss of habitat due to reservoir construction and destructive management practices. These changes may have led to the elimination of isolating mechanisms between the two species and may be increasing the opportunity for introgressive hybridization.