Distribution of plant species at a biome transition zone in New Mexico

Question: Is there a difference in plant species and life form composition between two major patch types at a biome transition zone? Are subordinate species associated with different patch types at the shortgrass steppe — Chihuahuan desert grassland transition zone? Is this association related to differences in soil texture between patch types and the geographic range of associated species? Location: central New Mexico, USA. Methods: Patches dominated by either Bouteloua gracilis, the dominant species in the shortgrass steppe, or Bouteloua eriopoda, dominant species in the Chihuahuan desert grasslands, were sampled for the occurrence of subordinate species and soil texture within a 1500‐ha transitional mosaic of patches. Results: Of the 52 subordinate species analysed, 16 species were associated with B. gracilis‐dominated patches and 12 species with B. eriopoda‐dominated patches. Patches dominated by B. gracilis were richer in annual grasses and forbs, whereas patches dominated by B. eriopoda contained more perennials forbs and shrubs. Soils of B. gracilis‐dominated patches had higher clay and lower rock contents compared with soils of B. eriopoda‐dominated patches. Differences in species characteristics of the dominant species as well as differences in soil texture between patch types contribute to patch‐scale variation in composition. The association of species to patch types was not related to their geographic range and occurrence in the adjacent biomes. Conclusions: Patch types at this biome transition zone have characteristic life‐form and species composition, but species are associated to patch types due to local constraints, independently from their affinity to the adjacent biomes.     

Burrowing activities of kangaroo rats and patterns in plant species dominance at a shortgrass steppe-desert grassland ecotone

Abstract. Our objective was to evaluate the effects of burrowing activities by banner-tail kangaroo rats (Dipodomys spectabilis Merriam) on plant community structure and species dominance for two patch types at the ecotone between shortgrass steppe and desert grassland in New Mexico, USA. 10 mounds produced by kangaroo rats were selected in patches dominated by Bouteloua gracilis (the dominant in shortgrass steppe communities) and 10 mounds were selected in patches dominated by B. eriopoda (the dominant in Chihuahuan desert grasslands). Plant cover and density by species were sampled from three locations associated with each mound: the mound proper, the edge of the mound in the transition area, and the off-mound vegetation. Similar cover of B. eriopoda for the edges of mounds in both patch types indicates the ability of this species to respond to animal disturbances regardless of the amount of cover in the surrounding undisturbed vegetation. By contrast, cover of B. gracilis was low for all mounds and mound edges in patches dominated by this species. Much higher cover of B. eriopoda on mound edges compared to the undisturbed vegetation in B. gracilis-dominated patches indicates that kangaroo rats have important positive effects on this species. Lower cover of perennial grasses and higher cover of forbs, shrubs, and succulents on the edges of mounds in B. eriopoda-dominated patches compared to patches dominated by B. gracilis indicate the importance of surrounding vegetation to plant responses on disturbed areas. Our results show that kangaroo rats have important effects on both species dominance and composition for different patch types, and may provide a mechanism for small-scale dominance patterns at an ecotone; thus providing further support for their role as keystone species in desert grasslands.     

Recruitment potential of two perennial grasses with different growth forms at a semiarid-arid transition zone

The objective of this study was to quantify differences in recruitment potential (seed production, seed presence in the soil) for two congeneric perennial grasses (Bouteloua gracilis, Bouteloua eriopoda [Poaceae]) that dominate adjacent arid and semiarid grassland biomes. It was hypothesized that these species have different recruitment strategies at the biome transition zone that are related to differences in their growth form and longevity. Recruitment potential for each Bouteloua species was compared in patches dominated by one or both species or codominated by the invasive shrub, Larrea tridentata (Zygophyllaceae). Regional variation in recruitment was examined for B. gracilis for cases in which comparable data were available in the literature for a site located within the semiarid grassland biome. The short-lived stoloniferous species B. eriopoda produced more seeds per plant than the long-lived bunchgrass B. gracilis, yet seed viability (<60%) and presence in the soil were lower. Mean viability of B. gracilis was higher (>90%) than that of B. eriopoda, and a greater percentage of seeds produced on a square meter basis was found in the soil (10-25%). Similar patterns were found for both species in all grass-dominated patches. Bouteloua eriopoda plants growing in patches codominated by L. tridentata produced fewer seeds per plant with lower viability, and fewer seeds were found in the soil compared to grass-dominated patches. Regional comparisons found greater seed production per square meter and more seeds in the soil for B. gracilis at the transitional site compared with a cooler, wetter site located within the semiarid grassland biome. These differences in recruitment potential along with published differences in rates of seedling establishment and vegetative spread may explain, at least in part, localized patterns in species dominance.     

Different facets of tree sapling diversity influence browsing intensity by deer dependent on spatial scale

Browsing of tree saplings by deer hampers forest regeneration in mixed forests across Europe and North America. It is well known that tree species are differentially affected by deer browsing, but little is known about how different facets of diversity, such as species richness, identity, and composition, affect browsing intensity at different spatial scales. Using forest inventory data from the Hainich National Park, a mixed deciduous forest in central Germany, we applied a hierarchical approach to model the browsing probability of patches (regional scale) as well as the species‐specific proportion of saplings browsed within patches (patch scale). We found that, at the regional scale, the probability that a patch was browsed increased with certain species composition, namely with low abundance of European beech (Fagus sylvatica L.) and high abundance of European ash (Fraxinus excelsior L.), whereas at the patch scale, the proportion of saplings browsed per species was mainly determined by the species’ identity, providing a “preference ranking” of the 11 tree species under study. Interestingly, at the regional scale, species‐rich patches were more likely to be browsed; however, at the patch scale, species‐rich patches showed a lower proportion of saplings per species browsed. Presumably, diverse patches attract deer, but satisfy nutritional needs faster, such that fewer saplings need to be browsed. Some forest stand parameters, such as more open canopies, increased the browsing intensity at either scale. By showing the effects that various facets of diversity, as well as environmental parameters, exerted on browsing intensity at the regional as well as patch scale, our study advances the understanding of mammalian herbivore–plant interactions across scales. Our results also indicate which regeneration patches and species are (least) prone to browsing and show the importance of different facets of diversity for the prediction and management of browsing intensity and regeneration dynamics.     

Restoring Structure in Late-Successional Sagebrush Communities by Thinning with Tebuthiuron

Late successional, dense Artemisia tridentata (big sagebrush) stands restrict associated plant species production, resulting in a monotypic, shrub‐dominated community that threatens biodiversity preservation and ecosystem function. Traditional practices to control A. tridentata can severely reduce or temporarily eliminate A. tridentata and other plant species. Thinning A. tridentata with low rates of the herbicide tebuthiuron enhances herbaceous plant production, community structure, ecosystem functioning, and biodiversity. Tebuthiuron was applied at rates of 0.11–1.0 kg ai/ha to A. tridentata‐dominated plots at Ten Sleep (1979), Lander (1993), and Waltman (1993), Wyoming. Changes in A. tridentata canopy cover, associated plant species biomass, and community composition were evaluated 13 and 14 years post‐treatment at Ten Sleep, and 2 and 4 years post‐treatment at Lander and Waltman. At all sites A. tridentata canopy cover decreased proportionally with increased tebuthiuron rate. Biomass of grasses increased as shrub biomass and cover decreased with increasing tebuthiuron rate. Forb biomass varied between treatments across sample year and site. Shifts from shrub‐dominated control to grass‐dominated treatment plots were attributable to biomass differences of A. tridentata and 2–3 grass species among treatments. The number of plant species was not significantly different between treatments at any site. Similarity indexes revealed progressively greater dissimilar plant composition between the control and sequential treatment plots of increased tebuthiuron rate. Incremental rates of tebuthiuron produce gradual changes in plant species composition without reducing species richness, which may have utility in certain restoration projects.     

Climatic variation and simulated patterns in seedling establishment of two dominant grasses at a semi‐arid‐arid grassland ecotone

Abstract. The objective of this study was to predict the effects of climatic variation at multiple temporal frequencies on seedling establishment by two congeneric C₄ perennial grasses (Bouteloua gracilis and B. eriopoda) at the ecotone between shortgrass steppe grassland and Chihuahuan desert grassland in central New Mexico, USA. The approach was to use a daily time‐step simulation model to determine the occurrence of a recruitment event in each year based upon the amount and timing of soil water required for establishment. Historical weather data were used to predict effects of seasonal and inter‐decadal variation in climate on establishment. A sensitivity analysis was used to predict effects of directional climate change on establishment. Bouteloua gracilis had a broad pattern of simulated establishment from May through September that included periods with high year‐to‐year variation in precipitation. B. eriopoda establishment events occurred primarily in July when precipitation amounts were most reliable. Climatic conditions from 1949 through 1968 were more favorable for B. eriopoda establishment compared to the cooler, wetter conditions from 1969 through 1988 that favored B. gracilis. Establishment of B. eriopoda was lowest in El Niño years whereas B. gracilis establishment was highest in La Niña years. Establishment of B. gracilis was most sensitive to temperature when precipitation was higher than current amounts. The greatest response to temperature by B. eriopoda for all precipitation amounts occurred at cooler temperatures than found currently. These results indicate that climatic variation at multiple frequencies has differential effects on seedling establishment for these two perennial grasses, and may account at least in part for patterns in dominance at this biome transition zone.     

Patch dynamics in grazed subtropical native pastures in south‐east Queensland

Abstract Patch formation is common in grazed grasslands but the mechanisms involved in the formation and maintenance of patches are not clear. To increase our knowledge on this subject we examined possible reasons for patch formation and the influence of management on changes between patch states in three experiments in native pasture communities in the Crows Nest district, south‐east Queensland. In these communities, small‐scale patches (tall grassland (dominated by large and medium tussock grasses), short swards (dominated by short tussock grasses and sedges), and lawns (dominated by stoloniferous and/or rhizomatous grasses)) are readily apparent. We hypothesized that the formation of short sward and lawn patches in areas of tall grassland was due to combinations of grazing and soil fertility effects. This was tested in Experiment 1 by applying a factorial combination of defoliation, nutrient application and transplants of short tussock and stoloniferous species to a uniform area of tall grassland. Total species density declined during the experiment, was lower with high nutrient applications, but was not affected by defoliation. There were significant changes in abundance of species that provided support for our hypotheses. With light defoliation and low nutrients, the tall grassland remained dominated by large tussock grasses and contained considerable amounts of forbs. With heavy defoliation, the pastures were dominated by medium tussock grasses and there were significant decreases in forbs and increases in sedges (mainly with low nutrients) and stoloniferous grasses (mainly with high nutrients). Total germinable seed densities and those of most species groups were significantly lower in the heavy defoliation than the light defoliation plots. Total soil seed numbers were not affected by nutrient application but there were fewer seeds of the erect forbs and more sedge seeds in plots with high nutrients. The use of resting from grazing and fire to manage transitions between patches was tested. In Experiment 2 , changes in species density and abundance were measured for 5 years in the three patch types with and without grazing. Experiment 3 examined the effects of fire, grazing and resting on short sward patches over 4 years. In Experiment 2 , total species density was lower in lawn than short sward or tall grassland patches, and there were more species of erect forbs than other plant groups in all patch types. The lawn patches were originally dominated by Cynodon spp. This dominance continued with grazing but in ungrazed patches the abundance of Cynodon spp. declined and that of forbs increased. In the short sward patches, dominance of short tussock grasses continued with grazing but in ungrazed plots their abundance declined while that of large tussock grasses increased. The tall grassland patches remained dominated by large and medium tussock species. In Experiment 3 , fire had no effect on species abundance. On the grazed plots the short tussock grasses remained dominant but where the plots were rested from grazing the small tussock grasses declined and the large tussock grasses increased in abundance. The slow and relatively small changes in these experiments over 4 or 5 years showed how stable the composition of these pastures is, and that rapid changes between patch types are unlikely.     

Plant dispersal strategies and the colonization of Araucaria forest patches in a grassland‐forest mosaic

Questions: In a natural grassland‐forest mosaic: What is the influence of phylogeny and diaspore traits related to disperser attraction (DAT) on (1) seed size/number trade‐off (SSNT) in woody species colonizing forest patches; (2) on the frequency of the species? 3. What is the influence of forest patch area on mean seed size and number. 4. Do phylogeny and DAT expressed at the species level affect this relationship? Location: Campos grassland and Araucaria forest in São Francisco de Paula, RS, Brazil, at ca. 29°28’ S; 50° 13’ W. Methods: Forest patches of different sizes in a grassland site recovering for ten years since human disturbances were surveyed by the relative abundance of vertebrate‐dispersed woody saplings. We described colonizer species according to taxonomic phylogenetic relationships and diaspore type, size and color. We analyzed with a variation partitioning method their influence on SSNT and on species frequency in the patches. At the community level we regressed mean seed size and number on forest patch area and evaluated how these relationships were affected by phylogeny and DAT at the species level. Results: 1. Phylogeny and DAT mostly explained seed size and seed number per diaspore variation. 2. By controlling phylogeny and DAT influence the frequency of species in forest patches was positively associated with their seed number in the diaspores, and negatively associated with their seed size. 3. Mean seed size and seed number at the community level were positively associated with patch area. 4. When phylogeny and DAT influences on seed size were removed this relationship was stronger for seed size and weaker for seed number. Conclusions: 1. Energy allocation to dispersal in detriment of offspring survival increased the successful establishment of colonizer species in forest patches, despite phylogenetic relationships and DAT variation in their diaspores. 2. Although patch area exerted a selective pressure on seed size, habitat preferences of dispersers may also influence patch colonization.     

Livestock exclusion increases the spatial heterogeneity of vegetation in Colorado shortgrass steppe

Abstract. Spatial heterogeneity, an important characteristic in semi‐arid grassland vegetation, may be altered through grazing by large herbivores. We used Moran's I, a measure of autocorrelation, to test the effect of livestock grazing on the fine scale spatial heterogeneity of dominant plant species in the shortgrass steppe of northeastern Colorado. Autocorrelation in ungrazed plots was significantly higher than in grazed plots for the cover of the dominant species Bouteloua gracilis, litter cover and density of other bunchgrasses. No species had higher autocorrelation in grazed compared to ungrazed sites. B. gracilis cover was significantly auto‐correlated in seven of eight 60‐yr ungrazed exclosures, four of six 8‐yr exclosures, and only three of eight grazed sites. Autocorrelograms showed that B. gracilis cover in ungrazed sites was frequently and positively spatially correlated at lag distances less than 5 m. B. gracilis cover was rarely autocorrelated at any sampled lag distance in grazed sites. The greater spatial heterogeneity in ungrazed sites appeared linked to patches characterized by uniformly low cover of B. gracilis and high cover of C₃ grasses. This interpretation was supported by simple simulations that modified data from grazed sites by reducing the cover of B. gracilis in patches of ca. 8 m diameter and produced patterns quite similar to those observed in ungrazed sites. In the one exclosure where we intensively sampled soil texture, autocorrelation coefficients for sand content and B. gracilis cover were similar at lag distances up to 12 m. We suggest that the negative effect of sand content on B. gracilis generates spatial heterogeneity, but only in the absence of grazing. An additional source of heterogeneity in ungrazed sites may be the negative interaction between livestock exclusion and B. gracilis recovery following patchy disturbance.     

Desertification alters patterns of aboveground net primary production in Chihuahuan ecosystems

Abstract The Chihuahuan desert of New Mexico, USA, has changed in historical times from semiarid grassland to desert shrublands dominated by Larrea tridentata and Prosopis glandulosa. Similar displacement of perennial grasslands by shrubs typifies desertification in many regions. Such structural vegetation change could alter average values of net primary productivity, as well as spatial and temporal patterns of production. We investigated patterns of aboveground plant biomass and net primary production in five ecosystem types of the Jornada Basin Long‐Term Ecological Research (LTER) site. Comparisons of shrub‐dominated desertified systems and remnant grass‐dominated systems allowed us to test the prediction that shrublands are more heterogeneous spatially, but less variable over time, than grasslands. We measured aboveground plant biomass and aboveground net primary productivity (ANPP) by species, three times per year for 10 years, in 15 sites of five ecosystem types (three each in Larrea shrubland, Bouteloua eriopoda grassland, Prosopis dune systems, Flourensia cernua alluvial flats, and grass‐dominated dry lakes or playas). Spatial heterogeneity of biomass at the scale of our measurements was significantly greater in shrub‐dominated systems than in grass‐dominated vegetation. ANPP was homogeneous across space in grass‐dominated systems, and in most growing seasons was significantly more patchy in shrub vegetation. Substantial interannual variability in ANPP complicates comparison of mean values across ecosystem types, but grasslands tended to support higher ANPP values than did shrub‐dominated systems. There were significant interactions between ecosystem type and season. Grasslands demonstrated higher interannual variation than did shrub systems. Desertification has apparently altered the seasonality of productivity in these systems; grasslands were dominated by summer growth, while sites dominated by Larrea or Prosopis tended to have higher spring ANPP. Production was frequently uncorrelated across sites of an ecosystem type, suggesting that factors other than season, regional climate, or dominant vegetation may be significant determinants of actual NPP.     

Differential seedling regeneration patterns across forest–grassland ecotones in two tropical treeline species (Polylepis spp.)

Successful forest expansion into grassland can be limited by seed dispersal and adverse conditions for tree seedlings in the grassland environment. In the high‐elevation Andes, human‐induced fragmentation has exacerbated the patchy distribution of Polylepis forests, threatening their unique biological communities and spurring restoration interest. Studies of Polylepis forest extent in Peru suggest that forest borders have remained stable over the past century despite decreasing anthropogenic disturbance, suggesting that tree seedling recruitment is being limited in the open grassland habitat. We studied natural seedling dispersion patterns of Polylepis sericea and Polylepis weberbaueri (Rosaceae) at forest–grassland edges across a range of environmental conditions to examine seedling recruitment and colonization of grasslands in Huascaran National Park (Peru). Using data from 2367 seedlings found in 48 forest–grassland edge plots (15 m × 15 m) at forest patches between 3900–4500 masl, we employed generalized mixed modelling to identify the significant associations of seedling densities with environmental covariates. In addition, we compared these associations to patterns of adult presence on the landscape. Seedling densities were associated with a combination of variables varying within (distance to forest edge) and among (elevation and dry season solar irradiation) plots across the landscape. For both species, seedling densities decreased with increasing distance away from the forest in a manner consistent with short‐distance seed dispersal by wind. Our results suggest that such short‐distance dispersal may slow forest expansion, but that there also appear to be substantial post‐dispersal limitations to seedling establishment in the grassland. Polylepis sericea densities decreased with elevation, while P. weberbaueri increased with elevation and decreased with solar irradiation. Associations of adult presence with elevation and solar irradiation mirrored those of seedling densities. Management of areas with forest patches dominated by these species should consider these differences in their environmental tolerances, particularly during species selection and zonation for reforestation.     

Forage patch use by grazing herbivores in a South African grazing ecosystem

Understanding how different herbivores make forage patch use choices explains how they maintain an adequate nutritional status, which is important for effective conservation management of grazing ecosystems. Using telemetry data, we investigated nonruminant zebra (Equus burchelli) and ruminant red hartebeest (Alcelaphus buselaphus subspecies camaa), use of burnt patches in a landscape mosaic of nutrient-poor, old grassland interspersed with young, recently burnt, nutrient-rich grass patches. The Mkambati Nature Reserve landscape on the east coast of South Africa provided large grazers with a challenge in finding and using appropriate patches in which to forage to meet their nutritional requirements. In Mkambati, grassland fires, mostly ignited by poachers, induce regrowth of young nutrient-rich grass, which subsequently attract grazers. We tested if the study animals foraged more in burnt patches than in the unburned grassland and whether burnt patch use was related to the distance to the previously visited burnt patch, burnt patch size, burnt patch age, and distance to areas with high poaching risk using MANOVA. In general, zebra moved faster than red hartebeest, and both species moved faster in unburnt grassland than in burnt patches. Red hartebeest and zebra patch selection were influenced by interpatch distance, patch age, patch size, and poaching risk. A limited set of intrinsic traits, i.e., body mass, digestion strategy, and muzzle width, yielded different patch use rules for the two species. Large ungulates patch use behaviour varied among species and across conditions and was influenced by anthropogenic impacts such as poaching and changed fire regimes. This could potentially affect biodiversity negatively and needs to be factored into management of conservation areas.     

Grassland connectivity explains entomophilous plant species assemblages in an agricultural landscape of the Pampa Region,Argentina

The Pampa grassland of Argentina is one of the most highly threatened biomes in the world. A high proportion of the original grassland cover has been transformed into land for agriculture or degraded. In the southern part of the region, fragmented semi‐natural grasslands over exposed rock still persist and connectivity between them is assumed to be crucial for maintaining viable populations. We quantified overall connectivity of grassland patches in a sector of the Southern Pampa region, and investigated the degree to which landscape connectivity explains entomophilous plant species assemblages in a subset of patches. We characterized each of the 301 patches in the landscape by their degree of intra‐patch and inter‐patch connectivity based on graph theory, and considering threshold dispersal distances from 100 to 1000 m. We surveyed entomophilous plant species in 39 grassland patches and classified the species in three categories (annual herbs, perennial herbs and shrubs) considering their different growth form and longevity. The influence of connectivity variables on entomophilous plant species assemblages variation was explored using Canonical Correspondence Analysis. Although grassland patches were poorly connected at all threshold distances, some of them were found to be critical for global connectivity. Connectivity significantly explained total, annual‐biennial and shrub assemblages for all threshold dispersal distances (6–13% of total variation). Variation in annual species assemblages was associated with intra‐patch and inter‐patch connectivity at short distance (100 m), while variation in shrub species assemblages was explained by intra‐patch and inter‐patch connectivity for distances between 100 m and 1000 m. This study evidenced the low connectivity of the study system, allowed the identification of critical areas for conservation, and provided valuable information to develop management strategies in increasingly human‐dominated landscapes.     

Assessing the vulnerability of riparian vegetation to invasion by Mimulus guttatus: relative importance of biotic and abiotic variables in determining species occurrence and abundance

Riparian habitats are particularly susceptible to invasion by non‐native plants. At present, attempts to build consensus as to what the primary drivers of plant invasion in riparian ecosystems might be is hindered by the absence of common standards for data collected on plant species (e.g. occurrence, or relative abundance). Mimulus guttatus L., a non‐native riparian plant species, was used as a model to determine how environmental drivers influence two aspects of invasibility: species occurrence and abundance (assessed in relation to three variables number of patches, patch area and number of stems per patch). Mimulus occurrence and abundance, together with 20 environmental variables, were surveyed in almost 700 contiguous 50‐m‐long riverbank segments within a catchment in north‐east Scotland. More than half of the segments had been colonized by Mimulus. Occurrence and number of patches responded to similar environmental gradients, particularly bare sediment, boulders, high soil moisture, short‐statured ruderal communities, and open canopies, and tended to be highest downstream where the river was widest. In contrast to occurrence and patch number, patch area and stem number per patch were higher in the upper reaches of the catchment and were positively associated with low tree canopy and vegetation dominated by light‐demanding species and smaller‐statured species. Patch area and stem number per patch were also positively related to grazing. This study has highlighted the importance of assessing more than one measure of invasion success (occurrence or patch number and either patch area or stem number per patch), as they are each determined by a different suite of environmental variables. Abiotic factors, such as sediment availability and presence of boulders, appeared to be the major determinants of occurrence and patch number, whereas biotic factors, such as interspecific competition and grazing, were more important ecological determinants underlying area and stem number per patch.     

Resource partitioning between ungulate populations in arid environments

Herbivores are major drivers of ecosystem structure, diversity, and function. Resilient ecosystems therefore require viable herbivore populations in a sustainable balance with environmental resource availability. This balance is becoming harder to achieve, with increasingly threatened species reliant on small protected areas in increasingly harsh and unpredictable environments. Arid environments in North Africa exemplify this situation, featuring a biologically distinct species assemblage exposed to extreme and volatile conditions, including habitat loss and climate change‐associated threats. Here, we implement an integrated likelihood approach to relate scimitar‐horned oryx (Oryx dammah) and dorcas gazelle (Gazella dorcas) density, via dung distance sampling, to habitat, predator, and geographic correlates in Dghoumes National Park, Tunisia. We show how two threatened sympatric ungulates partition resources on the habitat axis, exhibiting nonuniform responses to the same vegetation gradient. Scimitar‐horned oryx were positively associated with plant species richness, selecting for vegetated ephemeral watercourses (wadis) dominated by herbaceous cover. Conversely, dorcas gazelle were negatively associated with vegetation density (herbaceous height, litter cover, and herbaceous cover), selecting instead for rocky plains with sparse vegetation. We suggest that adequate plant species richness should be a prerequisite for areas proposed for future ungulate reintroductions in arid and semi‐arid environments. This evidence will inform adaptive management of reintroduced ungulates in protected environments, helping managers and planners design sustainable ecosystems and effective conservation programs.     

Inferences for life history strategies of Artemisia tridentata subspecies

Communities dominated by Artemisia tridentata ssp. tridentata, A. tridentata ssp. wyomingensis, and A. tridentata ssp. vaseyana, found in the Piceance Basin of western Colorado, were evaluated for life history strategy. Species cover data were analyzed using Grime's (1984) triangular model. Then a canonical analysis was conducted to obtain an ordination of relative species cover. Results from these two analyses were then used to infer strategy differences among life histories of subspecies, based on the respective plant community associates. These differences were found to be consistent with the divergent evolution assumed to have occurred in this species. The primary ordination axis was interpreted as an elevation-moisture gradient. Further analysis of soil data by factor analysis also separated the three Artemisia subspecies along a soil texture gradient.     

Influence of Patch Factors and Connectivity on the Avifauna of Fragmented Polylepis Forest in the Ecuadorian Andes

Human‐induced alteration of habitat is a major threat to biodiversity worldwide, especially in areas of high biological diversity and endemism. Polylepis (Rosaceae) forest, a unique forest habitat in the high Andes of South America, presently occurs as small and isolated patches in grassland dominated landscapes. We examine how the avian community is likely influenced by patch characteristics (i.e., area, plant species composition) and connectivity in a landscape composed of patches of Polylepis forest surrounded by páramo grasslands in Cajas National Park in the Andes of southern Ecuador. We used generalized linear mixed models and an information‐theoretic approach to identify the most important variables probably influencing birds inhabiting 26 forest patches. Our results indicated that species richness was associated with area of a patch and floristic composition, particularly the presence of Gynoxys (Asteraceae). However, connectivity of patches probably influenced the abundance of forest and generalists species. Elsewhere, it has been proposed that effective management plans for birds using Polylepis should promote the conservation of mature Polylepis patches. Our results not only suggest this but also show that there are additional factors, such as the presence of Gynoxys plants, which will probably play a role in conservation of birds. More generally, these findings show that while easily measured attributes of the patch and landscape may provide some insights into what influences patch use by birds, knowledge of other factors, such as plant species composition, is essential for better understanding the distribution of birds in fragmented landscapes.     

Enhanced growth of sagebrush (Artemisia tridentata) in response to manipulated ecosystem warming

Global models project impending climate changes that could significantly alter plant species composition in ecosystems. Climate manipulation experiments provide an opportunity to investigate such effects. Here we describe and apply a method for extracting the age‐detrended growth rate of sagebrush (Artemisia tridentata Nutt.) and show that experimental ecosystem warming enhances the growth rate of this shrub. Snowmelt date, not soil temperature or moisture, is demonstrated to be the dominant climate variable controlling the observed effect. Our findings suggest that global climate change will result in increased growth and range expansion of sagebrush near northern or high‐elevation range boundaries in the Western United States.     

Restoring riparian meadows currently dominated by Artemisia using alternative state concepts ‐ the establishment component

Abstract. We evaluated the potential for restoring riparian grass and sedge meadows currently dominated by Artemisia tridentata var. tridentata with an experiment in which we burned sites with low, intermediate, and high water tables, i.e., dry, intermediate, and wet sites. To define the alternative states and thresholds for these ecosystems, we examined burning and water table effects on both abiotic variables and establishment of grasses adapted to relatively high (Poa se‐cunda ssp. juncifolia), intermediate (Leymus triticoides), or low (L. cinereus) water tables. Wet sites had lower soil temperatures and higher soil water contents than dry sites. Burning increased soil temperatures on all sites. Undershrub microsites on control plots had the lowest temperatures, while former undershrub microsites on burn plots had the highest temperatures. Surface soil water was low on burn plots early in the growing season due to desiccation, but higher at deeper depths after active plant growth began. Emergence was generally greater on wet sites, but survival was microsite‐ and species‐specific. Undershrub microsites on control plots facilitated emergence and first‐year survival, but seedlings that survived initially harsh conditions on burn plots had similar numbers alive at the end. In general, favorable environments and establishment of species adapted to mesic conditions indicate that wet sites represent an alternative state of the naturally occurring dry meadow ecosystem type, and can be restored to grass and sedge meadows. Harsh environments and lack of establishment of species adapted to mesic conditions indicate that dry sites have crossed a threshold and may represent a new ecosystem type. Understory vegetation and seed banks on dry sites have been depleted, and restoration will require burning and reseeding with species adapted to more xeric conditions.