Small-stone content of Mima mounds of the Columbia Plateau and Rocky Mountain regions: implications for mound origin

Mima moundfields were investigated at the Lawrence Memorial Grassland Preserve, located on the Columbia Plateau in southern Wasco County, Oregon, and at three locations in the San Luis Valley and Sangre de Cristo Mountains, southern Colorado, to test the alternative hypotheses of mound origins by erosion, frost action, and soil translocation by geomyid pocket gophers. The concentrations of two size classes of small stones, gravel (8-15 mm diameter) and pebbles (15-50 mm diameter), were sampled along mound-to-intermound transects and at different depths within the mounds. Numbers and masses of small stones per unit soil volume increased from intermounds to mounds tops at the Colorado sites and from mound edge to mound top at the Oregon site, where thin intermound soils lay directly on the weathering surface of basalt bedrock. Numbers and masses of small stones in the surface soil of mound tops were greater than or similar to concentrations in deeper layers. Mean masses of individual pebbles were greater in the intermound zone than in mound soils at the Oregon site, but did not differ along mound-intermound gradient at the Oregon site and at one Colorado site, being highest at mound edges or in intermounds. These observations support the hypothesis that mounds are formed by centripetal translocation of soil by geomyid pocket gophers, and are contrary to predictions based on theories assuming erosion or frost action to be the mechanism of mound formation.     

Microtopography of microbiotic crusts on the Colorado Plateau, and distribution of component organisms

We analyzed the microtopography of microbiotic soil crusts at 3 sites on the Colorado Plateau of southern Utah and investigated distributions of cyanobacteria and several lichens in distinctive microhabitats created by this topography. At all 3 sites the long axes of linear soil mounds were oriented nonrandomly in a NNW—SSE direction. The conspicuous and consistent orientation of soil mounds may result from a combination of physical and biotic processes. Subtle differences across sites in mound orientation and organismal distribution suggest that these variables may be useful in comparing disturbance histories of crusts retrospectively. Differences in colonization frequencies, abundances, and distributions of microorganisms comprising the crusts, as a function of mound aspect or exposure, suggest that these organisms are associated with particular aspects due to distinctive and favorable microhabitats on these exposures. Polysaccharide sheath material, deposited by cyanobacteria, and associated filaments occurred in greater quantities on ENE than WSW mound exposures, and cover by Collema spp. lichens exhibited the same pattern. Colonization of mounds by common lichen species occurred significantly more frequently on ENE than WSW mound aspects at 2 of 3 sites. In contrast, the 3 most common lichen species, aside from Collema spp., did not exhibit a tendency for greater cover on ENE than WSW mound aspects. Physiological differences between gelatinous cyanolichens and green-algal lichens may explain the different distributional patterns of Collema spp. and the 3 other lichens.     

Negative effects of the western thatching ant ( Formica obscuripes ) on spiders (Araneae) inhabiting big sagebrush ( Artemisia tridentata )

The effect of thatching ants ( Formica obscuripes ) on number of spiders found on sagebrush ( Artemisia tridentata ) was determined by comparing spider abundance on shrubs at different distances from ant mounds. Spider abundance was inversely related to proximity to mounds. Spider abundance on shrubs abound abandoned mounds revealed no effects of mound proximity. Two mounds in which ants were exterminated showed a declining effect of mound proximity 1 month after treatment. One year after treatment there was no effect.     

Predator-prey interactions between the digger wasp Clypeadon laticinctus and the harvester ant Pogonomyrmex occidentalis

Females of the digger wasp Clypeadon laticinctus prey exclusively on workers of the western harvester ant Pogonomyrmex occidentalis. There is considerable variability in the size of ant workers both within and between ant mounds, and wasps tend to select larger workers as prey. Worker ants were most likely to be outside the nest at surface temperatures of 25–35°C. Wasps were seen at ant mounds most frequently when surface temperatures were between 40 and 50°C. Wasps captured ants outside and inside ant nests. Predation attempts were equally likely to result in a successful capture in either location, but entering the ant nests was more time consuming. Approximately 63% of visits by wasps to ant mounds resulted in a successful prey capture. In general, ant mounds that were visually conspicuous or close to wasp nest aggregations were more likely to be visited by wasps. However, frequency of visitation and rates of predation at a given mound varied considerably from week to week. Agonistic interactions between female wasps involving chases and occasional fights were likely to occur whenever two wasps were simultaneously present at an ant mound, especially when the only access to prey was by entering the mound.     

Revegetation of gopher mounds on aspen range in Utah

The colonization of the pocket - gopher ( Thomomys talpoides ) mounds by annual and perennial species of the understory of aspen woodland was observed over a four - year period. New and old gopher mounds exist as a mosaic of sites in one of three surface conditions: bare, dominated by annuals with a few seedlings of perennials, and dominated by perennials with annuals in peripheral areas. The regular creation of these new sites for plant colonization appears to favor the maintenance of aggressive perennials at high densities in the understory vegetation.     

Influence of soil and site characteristics on Palouse prairie plant communties

The structure of grassland communities can vary widely in response to heterogeneous habitat variables. In this study we document plant communities, soil types, and site characteristics for 12 Palouse prairie remnants in southeastern Washington and northern Idaho. We used general linear models to test the predictive value of 6 biophysical variables (slope, aspect, distance from edge, edge type, vegetation structure, and soil type) on 3 plant community metrics: species richness, Simpson’s diversity index, and the dominance of exotic species. From full models including all variables, we used Akaike’s information criterion (AIC) to select the best model for each metric. Aspect and vegetation structure were significant predictors of species richness ( R 2 = 0.08) and diversity ( R 2 = 0.09), while aspect, soil type, distance from edge, and edge type (type of adjacent matrix habitat) influenced the dominance of exotic species ( R 2 = 0.28). Additional soil sampling determined that the depth to a restrictive layer also had significant impacts on the dominance of exotic species ( R 2 = 0.31). This study confirms that biophysical characteristics influence Palouse prairie plant communities and may help set research and conservation priorities for isolated and uninventoried remnants.     

Natural variability of vegetation, soils, and physiography in the bristlecone pine forest of the Rocky Mountains

Pinus aristata Engelm. forest vegetation in Colorado was studied to determine vegetation composition and the relationship between vegetation and environment. Species percent cover, tree size class, and environmental variables were recorded for 49 plots. Previously collected data for 4 plots form New Mexico were included. Environmental variables included latitude, longitude, elevation, slope, aspect, topographic position, parent material, percent rock cover, mean rock size, litter depth, estimated plot age class, and evidence of anthropogenic disturbance. Soils were analyzed for texture, depth, and percent carbon and nitrogen. Pinus aristata foliage was analyzed for percent nitrogen and phosphorus. Direct and indirect gradient analyses (CANOCO) were used to determine environmental factors associated with community composition. Vegetation in P. aristata forests is influenced primarily by elevation and soil pH. Substrate, soil texture, topographic position, and geographic location are secondary factors. Six plant associations were identified using cluster analysis (listed in an elevational sequence from low to high): Pinus aristata/ Festuca arizonica Vasey, Pinus aristata / Festuca thurberi Vasey, Pinus aristata / Juniperus communis L., Pinus aristata / Vaccinium myrtilus L., Pinus aristata / Ribes montigenum McClatchie, and Pinus aristata / Trifolium dasyphyllum Torr. & Gray.     

Effects of prescribed fall burning on a wetland plant community, with implications for management of plants and herbivores

An important contemporary challenge for adaptive resource management is assessing both the direct and indirect effects of management activities by designing appropriate monitoring programs and sound analysis methods. Here we evaluate the effects of prescribed fall burning on a wetland plant community that is managed primarily for spring-migrating geese. During late fall in 2 consecutive years, we burned vegetation in 4 replicate blocks (2.3 ha each) that traversed a natural moisture and associated vegetation gradient. We used ordination, gradient analysis, and contingency table analysis to evaluate how annual changes in relative abundance of plants were affected by burning as well as other important ecological factors. Burning increased species diversity of plants, especially in the 2 wetter vegetation zones, but had no effect on species richness or on the proportion of native plant species. Wetland plant species responded to prescribed burning independently, and their response often differed by vegetation zone and with annual variation in flooding. Burning enhanced the abundance of native foxtail barley ( Hordeum jubatum ) and reduced the abundance of introduced swamp timothy ( Crypsis shoenoides ). Saltgrass ( Distichlis spicata ), a native plant species, was usually less abundant following burning, although the level of response was different for each of the 3 vegetation zones. Two other introduced plant species, quackgrass ( Elytrigia repens ) and reed canarygrass ( Phalaris arundinaceae ), were less abundant after fall burning, especially when spring flooding was more extensive. Wild geese using the experimental blocks for feeding clearly preferred burned sites, suggesting that fall burning can enhance wetland use by geese during spring. Given that simple manipulations such as burning and flooding of a wetland system may often produce complex results, we suggest that on-going management schemes be regularly evaluated with field experiments such as those conducted in this study.     

The termites of the Mayombe Forest Reserve,Congo (Brazzaville): transect sampling reveals an extremely high diversity of ground-nesting soil feeders

Sampling using a replicated standard transect method in an extensive undisturbed primary hill rain forest in Congo (Brazzaville) revealed a very high species richness of termites, especially soil-feeding soldierless termites (Apicotermitinae). The assemblage, as estimated by the transects, resembled that previously characterized in comparable forest in southern Cameroon, but with some species turnover reflecting the gamma diversity of the Guinean-Congolese block as a whole. Species richness was three or four times that shown by a previous study addressed to mound populations alone, emphasizing the importance of sampling soil-feeding termites neither building nor living in mounds.     

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.     

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.     

Classification of the riparian vegetation along a 6-km reach of the Animas River, southwestern Colorado

Riparian ecosystems are important components of landscapes, particularly because of their role in biodiversity. A first step in using a ""coarse-filter"" approach to riparian biodiversity conservation is to determine the kinds of riparian ecosystems. These ecosystems vary substantially in plant species composition along a single river reach, as well as between rivers, and yet the river-reach scale has received little attention. We sampled the vascular plant composition of 67 contiguous patches of riparian vegetation along the reach of the Animas River, in southwestern Colorado's San Juan Mountains, that is relatively undisturbed by human land uses. Using cluster analysis and detrended correspondence analysis, we identified eight riparian community types along the reach. Using a new technique, we combined overstory size-class data and understory cover data to identify community types. The eight community types, which are in part the products of past floods, are spatially arranged along the reach in relation to variation in valley morphology, tributary location, and geomorphic landforms. These eight community types do not necessarily represent successional stages of a single potential vegetation type. This study at the river-reach scale suggests that sampling and analysis, as well as conservation, may need to be turned to the scale of patchiness produced by flood disturbances in the riverine landscape, since vegetation varies significantly at this scale.     

Transverse and longitudinal variation in woody riparian vegetation along a montane river

This study explores how the relationship between flow and riparian vegetation varies along a montane river. We mapped occurrence of woody riparian plant communities along 58 km of the San Miguel River in southwestern Colorado. We determined the recurrence interval of inundation for each plant community by combining step-backwater hydraulic modeling at 4 representative reaches with Log-Pearson analysis of 4 stream gaging stations. Finally, we mapped bottomland surficial geology and used a Geographic Information System to overlay the coverages of geology and vegetation. Plant communities were distinctly arrayed along the hydrologic gradient. The Salix exigua Nuttall (sandbar willow) community occurred mostly on surfaces with a recurrence interval of inundation shorter than 2.2 years; the Betula occidentalis Hooker (river birch) community peaked on sites with recurrence intervals of inundation between 2.2 and 4.6 years. The hydrologic position occupied by communities dominated by Populus angustifolia James (narrowleaf cottonwood) was strongly related to age of trees and species composition of understory shrubs. The fraction of riparian vegetation on surfaces historically inundated by the river decreased in the upstream direction from almost 100% near Uravan to <50% along the South Fork of the San Miguel River. In upstream reaches much of the physical disturbance necessary to maintain riparian vegetation is provided by valley-side processes including debris flows, floods from minor tributaries, landslides, and beaver activity. Where valley-side processes are important, prediction of riparian vegetation change based on alterations of river flow will be incomplete.     

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.     

A field study of the nesting ecology of the thatching ant, Fomica obscuripes Forel, at high altitude in Colorado

A field study of the thatching ant, Formica obscuripes Forel, at 2560 m elevation in Colorado provided information on mound density, composition, dimensions, and temperatures; worker longevity; and mite parasitization. Density was 115 mounds/ha. Mounds had 1-52 entrances and Peromyscus fecal pellets in the thatch. Mounds conserved heat and exhibited thermal stratification. Excavations of 4 nests revealed depths of 0.3 m to almost 1 m, novel myrmecophiles, and 0-198 wingless queens per nest. Marking experiments demonstrated that some workers overwinter and live more than a year.     

C 3 and C 4 Species Changes Identified by δ 13 C values of soil organic matter in a Colorado prairie

We measured carbon isotope signatures (δ 13 C) from 0-10 cm and 10-20 cm soil depth intervals for grassland soils near Boulder, Colorado. These grasslands included tall-, short-, and mixed-grass prairies that were grazed, ungrazed, or hayed. Soils exhibited δ 13 C signatures consistent with observations that current sites are a mix of C 3 and C 4 species, with C 3 plants more abundant in mixed-grass than in native tall- or shortgrass prairies. The δ 13 C signatures were not significantly different for grassland types; however, management treatments (grazing, no grazing, haying) significantly influenced changes in soil δ 13 C signatures from the 0-10 cm to 10-20 cm soil depth intervals. We observed a correlation ( r = 0.63) between isotopic values of surface soils and percent native species in total vegetation cover. Overall, the community type with the lowest percentage of nonindigenous species cover had the most enriched δ 13 C signature. Sites currently grazed by prairie dogs, cattle, or both herbivores had stronger C 3 signatures, indicating that grazing may have increased C 3 plant productivity in these communities at the expense of C 4 grasses. This finding differs from studies of native shortgrass steppe where grazing has the opposite effect on the relative abundance of these 2 functional groups of plants. This result, along with the correlation between C 3 isotopic values and nonnative vegetation abundance, provides evidence that management practices that maintain dominance of C 4 grasses should be encouraged.     

Leafhopper assemblages on native and reseeded grasslands in southwestern Montana

Using sweep samples, we surveyed leafhoppers (Homoptera: Cicadellidae) on grassland sites in the Gallatin Valley of Montana during 1988 and 1991. We sampled 12 sites representing 2 habitat types defined by their dominant plant species in an undisturbed state ( Stipa comata / Bouteloua gracilis and Festuca idahoensis / Agropyron spicatum ). At half of the sites the native plant communities were present, whereas the remainder had been reseeded with either Agropyron spicatum (to replace the S. comata / B. gracilis assemblage ) or Bromus inermis (to replace the F. idahoensis / A. spicatum assemblage). We found at least 66 species of leafhoppers among 44,428 adults collected. Seven taxa comprised 83% of all individuals collected: Doratura stylata (26%), Ceratagallia spp. (18%), Endria inimica (17%), Orocastus perpusillus (7%), Sorhoanus spp. (6%), Athysanella spp. (5%), and Psammotettix lividellus (4%). Sites with similar vegetation had broadly similar leafhopper assemblages, and assemblages differed most between the relatively xeric Stipa comata / Bouteloua gracilis sites and the more mesic sites dominated by Bromus inermis . The composition of a leafhopper assemblage at a site tended to be more similar to those on noncontiguous sites with the same overall vegetation than to those on contiguous sites with different vegetation. These patterns are likely related to the fact that many Cicadellidae are host specialists. In fact, variation in abundance of some of the most common leafhopper taxa on our sites was correlated with the percent cover of their known host plants. Our analyses of the leafhopper assemblages generally support the contention that terrestrial plant associations are among the more useful indicators of insect community composition.     

Form and dispersion of Mima mounds in relation to slope steepness and aspect on the Columbia Plateau

Patterned ground consisting of Mima-type earth mounds and associated sorted stone circles and nets is widespread on the Columbia Plateau of western North America. Studies of the geometric relationships of mounds and stone nets to slope aspect and steepness were carried out at the Lawrence Memorial Grassland Preserve, north central Oregon in June 1987. Mound and moundfield characteristics were sampled on randomly chosen 1-ha plots on slopes of different aspect and steepness. Mounds were largest, most circular and symmetrical in form, and most fully encircled by beds of size-sorted stones on level sites. On slopes of increasing steepness, mounds decreased in size, showed increasing asymmetry and downslope elongation, and became connected into lines oriented up- and downslope. Encircling stone beds became more weakly developed or disappeared on upslope and downslope sides of the mounds, and the lateral beds developed downslope extensions that eventually merged with those of adjacent upslope and downslope mounds. These patterns are interpreted as reflecting changes in the manner of soil translocation by northern pocket gophers, Thomomys talpoides , due to their responses to tunneling on slopes and to the modification of the flow of water across the slope because of the presence of mounds.     

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).     

A sagebrush community type classification for mountainous northeastern Nevada rangelands

Vegetation characteristics of 15 sagebrush community types identified on the Humboldt National Forest, northeastern Nevada, are described. A total of 218 plant species were found over the 372 relatively undisturbed rangeland communities sampled. The dominant plant families encountered were the Asteraceae (45 taxa), Poaceae (32 taxa), Scrophulariaceae (14 taxa), and the Fabaceae (12 taxa). Average annual dry weight production of the community types ranged from about 400 kg/ha/yr on types with Artemisia nova as the dominant sagebrush species to 1,200 kg/ha/yr on some A. tridentata ssp. vaseyana community types. A general increase in species richness and vegetation plus litter ground cover was observed within community types as the dominant sagebrush species changed from A. nova to A. arbuscula to A. longiloba to A. tridentata spp. tridentata to A. tridentata ssp. vaseyana . Major differences in plant species production and constancy exist between the sagebrush community types studied.