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.     

Geographic patterns of simulated establishment of two Bouteloua species: implications for distributions of dominants and ecotones

Abstract. Our overall objective was to use a soil water model to predict spatial patterns in germination and establishment of two important perennial C₄-bunchgrasses across the North American shortgrass steppe and desert grassland regions. We also predicted changes in establishment patterns under climate change scenarios. Bouteloua gracilis dominates the shortgrass steppe from northeastern Colorado to southeastern New Mexico. Bouteloua eriopoda dominates desert grasslands in central and southern New Mexico. Germination and establishment for each species were predicted at 16 sites along the gradient using a daily time step, multi-layer soil water model (SOILWAT) to determine the percentage of years that temperature and soil water criteria for germination and establishment were met. Percentage of years with predicted establishment decreased from north to south for B. gracilis, but increased from north to south for B. eriopoda, comparable to observed dominance patterns. The 95 % confidence interval around the point at which simulated establishment were equal for the two species was near the location of the shortgrass steppe-desert grassland ecotone where both species are abundant. The intersection in percentage of years with establishment for the two species was predicted to move further north when climate was scaled using three Global Circulation Models (GCMs), indicating a possible northward expansion of B. eriopoda. Our results suggest that recruitment by seed may be an important process in determining, at least in part, the geographic distribution of these two species. Changes in climate that affect establishment constraints could result in shifts of species dominance that may or may not be accompanied by changes in species composition.     

Woody plant invasion at a semi‐arid/arid transition zone: importance of ecosystem type to colonization and patch expansion

Question: How do patterns in colonization and patch expansion of an invasive woody plant (Larrea tridentata, Zygo‐phyllaceae) differ between two grassland ecosystems at a biome transition zone? Location: Semi‐arid/arid transition zone in central New Mexico. Methods: Frequency of occurrence, height, and surface area of saplings (n= 134) and patches of adult plants (n= 247) of the invasive shrub, L. tridentata, were measured within a mosaic of ecosystems dominated either by the Chihuahuan Desert species, Bouteloua eriopoda (Poaceae), or the shortgrass steppe species, B. gracilis, located within 1 km of the L. tridentata‐dominated ecosystem. Distances between L. tridentata patches and patch area were used to estimate connectivity as a measure of propagule pressure. Sapling age (estimated from height using previously established relationships) and distance to the L. tridentata‐dominated ecosystem was used to evaluate patterns in dispersal. Cover by species or functional group inside each L. tridentata patch was compared with surrounding vegetation to estimate changes in species composition with patch expansion. Results: L. tridentata saplings (< 1%) and adult patches (15%) occurred less frequently in B. gracilis‐dominated ecosystems than expected based on areal extent of this ecosystem type. Propagule pressure did not differ with distance from the core ecosystem dominated by L. tridentata. Evidence for both local and long‐distance dispersal events was found. Similar relationships between number of plants and patch area in both grassland types indicate similar patterns in patch expansion. Cover of perennial forbs was higher and cover of dominant grasses was lower in L. tridentata patches compared with the surrounding vegetation for both ecosystem types. Conclusions Spatial variation in L. tridentata saplings and patches at this biome transition zone is related to the different susceptibilities to invasion by two grassland ecosystems. The persistence of grasslands at this site despite region‐wide expansion by L. tridentata may be related to the spatial distribution of B. gracilis‐dominated ecosystems that resist or deter invasion by this woody plant.     

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.     

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.     

Vegetation of ant-hills in a mountain grassland:effects of mound history and of dominant ant species

Vegetation in grasslands with well-developed long-lastingant-hills in the Slovenské Rudohorie Mts., Slovakia, was studiedin relation to (i) position on the mound, (ii) ant speciesforming the mound, and (iii) history of the mound. Permanent plotrecordings of mound size and dominant ant species started fifteen years priorthe study began provided information on the history of individual mounds.The mound vegetation bears a striking similarity to vegetation insimilar habitats across a large part of Europe due to presence of species suchas Agrostis capillaris, Dianthusdeltoides, Polytrichum commune agg.,Thymus pulegioides, and Veronicaofficinalis. Out of the three major ant species-groups presentat the site (Lasius flavus, Tetramoriumcaespitum and Formica spp.), L.flavus had the most pronounced and the most lasting effect on themound vegetation. The dominance of the plant species listed above increased withthe time span over which the mound was inhabited by L.flavus. The effects of other species on vegetation composition,though discernible from short-term observation, disappeared over severalyears. The mounds proper did not differ from the undisturbed grassland in theproportion of myrmecochorous plants or plants with specific seed size ordormancy type. However, there was a highly significant concentration ofmyrmecochorous plants in the grassland patches immediately neighbouring themounds; this is likely to be due to seeds deposited there by the workers fromthe nest after the elaiosomes had been consumed. The mound vegetation wascomposed mainly of species with long stolons or rhizomes; however, there was nosignificant difference in formation type or length of stolons/rhizomesbetween mounds and the rest of the grassland or among mounds formed by differentant species.     

Structure in a desert rodent community: use of space around Dipodomys spectabilis mounds

Summary In an earlier paper (Bowers et al. 1987) we reported patterns of microhabitat use by desert rodents among 0.25-ha plots where seeds were added or certain rodent species removed. We used the results to make inferences about the spatial organization of the whole rodent community. Here we change our focus to test for spatial usage patterns at a smaller (within-plot) scale. Specifically, we examine to what extent spatial use varies with proximity to mounded burrows of the large kangaroo rat, Dipodomys spectabilis. Capture frequency of five of nine rodent species was correlated with distance from D. spectabilis mounds, while six species showed correlations with vegetative cover that also increased with distance from mounds. In general, sites nearer mounds were underutilized, and sites further away overutilized by the rodent community as a whole. Logistic regression analyses showed for six species that vegetative cover and particularly, distance to nearest mound accounted for more of the variation in whether a station captured a species than did plot-to-plot (i.e., treatment) effects. Similar analyses using two plots where D. spectabilis was removed (where vacant mounds persited for years) suggested that the selection of microhabitats with contrasting vegetation structure was more pronounced in the absence than in the presence of D. spectabilis, and that in most cases distance from mounds was important only if mounds were occupied. Spatial structure in this community appears to revolve around the occupation of space by dominant individuals that partially modify/obscure large scale patterns involving the selection of particular structural microhabitats.     

Short-Term Vegetation and Soil Responses to Mechanical Destruction of Rabbit (Oryctolagus cuniculus L.) Warrens in an Australian Box Woodland

We studied the impact of disturbance by rabbits on plants and soils along a gradient out from the center of ripped rabbit warrens in an Australian semiarid woodland. Five years after the warrens were ripped, the impact of rabbits was still apparent. The cover of bare soil declined, and the cryptogam cover increased with increasing distance from the warren mound. However, litter cover, plant cover, and plant diversity remained unchanged with increasing distance from the mounds. Differences in plant composition were apparent with increasing distance from the mounds, with three species, Schismus barbatus, Salsola kali var. kali, and Chenopodium melanocarpum dominating the mounds, whereas the perennial grass Austrostipa scabra dominated the nonwarren control surfaces. Two species, Crassula sieberana and S. barbatus, dominated the active soil seed bank on ripped warrens. The mounds had the lowest number of species in the soil seed bank, whereas the warren edge microsite had the greatest. Ripped and unripped warrens differed substantially in their complement of species, and ripped warrens contained an order of magnitude fewer active warren entrances compared with unripped warrens. Ripped warrens also had significantly more plant cover than unripped warrens. Taken together, our results reinforce the view that rabbits have a destructive effect on surface soils and vegetation in semiarid woodlands and suggest that restoration of the original woodland vegetation after warren ripping is likely to be a slow and ongoing process.     

Effects of experimental rainfall manipulations on Chihuahuan Desert grassland and shrubland plant communities

Aridland ecosystems are predicted to be responsive to both increases and decreases in precipitation. In addition, chronic droughts may contribute to encroachment of native C₃ shrubs into C₄-dominated grasslands. We conducted a long-term rainfall manipulation experiment in native grassland, shrubland and the grass–shrub ecotone in the northern Chihuahuan Desert, USA. We evaluated the effects of 5 years of experimental drought and 4 years of water addition on plant community structure and dynamics. We assessed the effects of altered rainfall regimes on the abundance of dominant species as well as on species richness and subdominant grasses, forbs and shrubs. Nonmetric multidimensional scaling and MANOVA were used to quantify changes in species composition in response to chronic addition or reduction of rainfall. We found that drought consistently and strongly decreased cover of Bouteloua eriopoda, the dominant C₄ grass in this system, whereas water addition slightly increased cover, with little variation between years. In contrast, neither chronic drought nor increased rainfall had consistent effects on the cover of Larrea tridentata, the dominant C₃ shrub. Species richness declined in shrub-dominated vegetation in response to drought whereas richness increased or was unaffected by water addition or drought in mixed- and grass-dominated vegetation. Cover of subdominant shrubs, grasses and forbs changed significantly over time, primarily in response to interannual rainfall variability more so than to our experimental rainfall treatments. Nevertheless, drought and water addition shifted the species composition of plant communities in all three vegetation types. Overall, we found that B. eriopoda responded strongly to drought and less so to irrigation, whereas L. tridentata showed limited response to either treatment. The strong decline in grass cover and the resistance of shrub cover to rainfall reduction suggest that chronic drought may be a key factor promoting shrub dominance during encroachment into desert grassland.     

Impact of Formica exsecta Nyl. on seed bank and vegetation patterns in a subalpine grassland ecosystem

The mound building ant Formica exsecta Nyl. is widely distributed in grassland ecosystems of the Central European Alps. We studied the impact of these ants on seed bank and vegetation patterns in a 11 ha subalpine grassland, where we counted over 700 active ant mounds. The mounds showed a distinct spatial distribution with most of them being located in tall‐grass, which was rarely visited by ungulates (red deer; Cervus elaphus L.). Heavily grazed short‐grass, in contrast, seemed to be completely avoided by ants as only few mounds were found in this vegetation type. The species composition of the ant mound and grassland seed banks was quite similar, i.e. from 15 common plant species 12 were found in both seed bank types. We found the same proportions of myrmecochorous seeds in ant mound and grassland soil samples. In contrast, the number of seeds was 15 times higher in mound compared with the grassland soil samples. Also, the vegetation growing on ant mounds significantly differed from the vegetation outside the mounds: graminoids dominated on ant mounds, herbaceous and myrmecochorous species in the grassland vegetation. We found significant continuous changes in vegetation composition on gradients from the ant mound centre to 1 m away from the mound edge. Overall, F. exsecta was found to have a considerable impact on seed bank and vegetation patterns in the grassland ecosystem studied. These insects not only altered grassland characteristics in the close surrounding of their mounds, but also seem to affect the entire ecosystem including, for example, the spatial use of the grassland by red deer.     

Effects of ants on plant diversity in semi-natural grasslands

Ants (L. niger and L. flavus) build conspicuous mounds that are covered with vegetation. The aim of this study was to investigate whether the vegetation on ant mounds in semi-natural grasslands differed from that around the mounds. Another aim was to investigate whether the changes in the vegetation on ant mounds were influenced by grazing management or by habitat characteristics, semi-dry versus moist. Here, the total number of plant species and total plant cover were lower on ant mounds than in patches off-mound. The plant cover of perennials that form rosettes was twice as high on mounds inhabited by L. niger than on those inhabited by L. flavus. Only a few plant species were restricted to either ant mounds or adjacent field and the effects of ants on the plant diversity in semi-natural grasslands seemed to be low. Grazing management did not affect the differences in the vegetation on ant mounds and in equal-sized patches off-mound, whereas habitat characteristics affected ant-induced changes in vegetation cover of some plant species.     

A mast‐seeding desert shrub regulates population dynamics and behavior of its heteromyid dispersers

Granivorous rodent populations in deserts are primarily regulated through precipitation‐driven resource pulses rather than pulses associated with mast‐seeding, a pattern more common in mesic habitats. We studied heteromyid responses to mast‐seeding in the desert shrub blackbrush (Coleogyne ramosissima), a regionally dominant species in the Mojave–Great Basin Desert transition zone. In a 5‐year study at Arches National Park, Utah, USA, we quantified spatiotemporal variation in seed resources in mast and intermast years in blackbrush‐dominated and mixed desert vegetation and measured responses of Dipodomys ordii (Ord's kangaroo rat) and Perognathus flavescens (plains pocket mouse). In blackbrush‐dominated vegetation, blackbrush seeds comprised >79% of seed production in a mast year, but 0% in the first postmast year. Kangaroo rat abundance in blackbrush‐dominated vegetation was highest in the mast year, declined sharply at the end of the first postmast summer, and then remained at low levels for 3 years. Pocket mouse abundance was not as strongly associated with blackbrush seed production. In mixed desert vegetation, kangaroo rat abundance was higher and more uniform through time. Kangaroo rats excluded the smaller pocket mice from resource‐rich patches including a pipeline disturbance and also moved their home range centers closer to this disturbance in a year of low blackbrush seed production. Home range size for kangaroo rats was unrelated to seed resource density in the mast year, but resource‐poor home ranges were larger (P < 0.001) in the first postmast year, when resources were limiting. Blackbrush seeds are higher in protein and fat but lower in carbohydrates than the more highly preferred seeds of Indian ricegrass (Achnatherum hymenoides) and have similar energy value per unit of handling time. Kangaroo rats cached seeds of these two species in similar spatial configurations, implying that they were equally valued as stored food resources. Blackbrush mast is a key resource regulating populations of kangaroo rats in this ecosystem.     

Effects of Erosion from Mounds of Different Termite Genera on Distinct Functional Grassland Types in an African Savannah

A key aspect of savannah vegetation heterogeneity is mosaics formed by two functional grassland types, bunch grasslands, and grazing lawns. We investigated the role of termites, important ecosystem engineers, in creating high-nutrient patches in the form of grazing lawns. Some of the ways termites can contribute to grazing lawn development is through erosion of soil from aboveground mounds to the surrounding soil surface. This may alter the nutrient status of the surrounding soils. We hypothesize that the importance of this erosion varies with termite genera, depending on feeding strategy and mound type. To test this, we simulated erosion by applying mound soil from three termite genera (Macrotermes, Odontotermes, and Trinervitermes) in both a field experiment and a greenhouse experiment. In the greenhouse experiment, we found soils with the highest macro nutrient levels (formed by Trinervitermes) promoted the quality and biomass of both a lawn (Digitaria longiflora) and a bunch (Sporobolus pyramidalis) grass species. In the field we found that soils with the highest micro nutrient levels (formed by Macrotermes) showed the largest increase in cover of grazing lawn species. By linking the different nutrient availability of the mounds to the development of different grassland states, we conclude that the presence of termite mounds influences grassland mosaics, but that the type of mound plays a crucial role in determining the nature of the effects.     

Influence of small-scale disturbances by kangaroo rats on Chihuahuan Desert ants

Banner-tailed kangaroo rats (Dipodomys spectabilis) are prominent ecosystem engineers that build large mounds that influence the spatial structuring of fungi, plants, and some ground-dwelling animals. Ants are diverse and functionally important components of arid ecosystems; some species are also ecosystem engineers. We investigated the effects of patch disturbances created by D. spectabilis mounds on ant assemblages in a Chihuahuan Desert grassland in southern New Mexico by using pitfall traps in a paired design (mound vs. matrix). Although the disturbances did not alter species richness or harbor unique ant communities relative to the matrix, they did alter species composition; the abundances of 6 of 26 species were affected. The disturbances might also act to disrupt spatial patterning of ants caused by other environmental gradients. In contrast to previous investigations of larger-scale disturbances, we detected no effects of the disturbances on ants at the functional-group level. Whether ant communities respond to disturbance at a functional-group or within-functional-group level may depend on the size and intensity of the disturbance. Useful functional-group schemes also may be scale-dependent, however, or species may respond idiosyncratically. Interactions between disturbance-generating mammals and ants may produce a nested spatial structure of patches. Received: 11 October 1999 / Accepted: 11 March 2000     

Interactive effects of soil-dwelling ants, ant mounds and simulated grazing on local plant community composition

Interactions between aboveground vertebrate herbivores and subterranean yellow meadow ants (Lasius flavus) can drive plant community patterns in grassland ecosystems. Here, we study the relative importance of the presence of ants (L. flavus) and ant mounds under different simulated grazing regimes for biomass production and species composition in plant communities. We set up a greenhouse experiment using intact soil cores with their associated vegetation.We found that plant biomass production in the short term was affected by an interaction between simulated grazing (clipping) and ant mound presence. Clipping homogenized production on and off mounds, while in unclipped situations production was higher off than on mounds. During the experiment, these differences in unclipped situations disappeared, because production on unclipped mounds increased. Plant species richness was on average higher in clipped treatments and patterns did not change significantly over the experimental period. Plant community composition was mainly affected by clipping, which increased the cover of grazing-tolerant plant species. The actual presence of yellow meadow ants did not affect plant community composition and production.We conclude that the interaction between ant mounds and clipping determined plant community composition and biomass production, while the actual presence of ants themselves was not important. Moreover, clipping can overrule effects of ant mounds on biomass production. Only shortly after the cessation of clipping biomass production was affected by ant mound presence, suggesting that only under low intensity clipping ant mounds may become important determining plant production. Therefore, under low intensity grazing ant mounds may drive the formation of small-scale plant patches.     

Alpine and subalpine snow patch vegetation on the Bogong High Plains,SE Australia

Abstract. Snow patch vegetation in Australia is rare, being restricted to the relatively small area of alpine and subalpine country in the highlands of southeastern Australia. Snow patch vegetation occurs on steeper, sheltered southeastern slopes, where snow persists until well into the growing season (December/January). We surveyed the vegetation of 33 snow patch sites in the alpine and subalpine tracts of the Bogong High Plains, within the Alpine National Park, in Victoria. The vegetation was dominated by herbs and graminoids, with few shrubs and mosses. Major structural assemblages identified included closed herb‐fields dominated by Celmisia spp, and grasslands dominated by Poa fawcettiae or Poa costiniana. These assemblages occurred on mineral soils. Open herb‐fields dominated by Caltha introloba and several sedge species occurred on rocky and stony substrata. Vegetation‐environment relationships were explored by ordination and vector fitting. There was significant variation in the floristic composition of snow patch vegetation as a function of duration of snow cover, altitude, slope and site rockiness. Alpine sites were floristically distinct from subalpine sites, with a greater cover of Celmisia spp. and a lesser cover of low shrubs in the former. There was floristic variation within some snow patches as a function of slope position (upper, middle or lower slope) but this was not consistent across sites. The current condition of snow patch vegetation on the Bogong High Plains is degraded, with bare ground exceeding 20% cover at most sites. Snow patch vegetation is utilized preferentially by domestic cattle, which graze parts of the Bogong High Plains in summer. Such grazing is a potential threat to this rare vegetation type.     

Vegetation change following removal of keystone herbivores from desert grasslands in New Mexico

Abstract. Responses of plant communities to mammalian herbivores vary widely, due to variation in plant species composition, herbivore densities, forage preferences, soils, and climate. In this study, we evaluated vegetation changes on 30 sites within and adjacent to the Sevilleta National Wildlife Refuge (SNWR) in central New Mexico, USA, over a 20‐yr period following removal of the major herbivores (livestock and prairie dogs) in 1972–1975. The study sites were established in 1976, and were resampled in 1986 and 1996 using line transect methods. At the landscape scale, repeated measures ANOVA of percentage cover measurements showed no significant overall net changes in total perennial plant basal cover, either with or without herbivores present; however, there was an overall increase in annual forbs and plant litter from 1976 to 1996. At the site scale, significant changes in species composition and dominance were observed both through time and across the SNWR boundary. Site histories varied widely, with sites dominated by Bouteloua eriopoda being the most dynamic and sites dominated by Scleropogon brevifolius being the most persistent. Species‐specific changes also were observed across multiple sites: B. eriopoda cover increased while Gutierrezia sarothrae greatly decreased. The non‐uniform, multi‐directional changes of the sites' vegetation acted to prevent detection of overall changes in perennial vegetation at the landscape level. Some sites displayed significant changes after removal of herbivores, while others appeared to respond primarily to climate dynamics. Certain species that were not preferred by livestock or prairie dogs, showed overall declines during drought periods, while other preferred species exhibited widespread increases during wetter periods regardless of herbivore presence. Therefore, the vegetation dynamics cannot be attributed solely to removal of herbivores, and in some cases can be explained by short‐ and long‐term fluctuations in climate. These results emphasize the variety of responses of sites with differences in vegetation to mammalian herbivores under otherwise similar climatic conditions, and illustrate the value of site‐ and landscape‐scale approaches to understanding the impacts of plant‐herbivore interactions.     

Plant population and community responses to removal of dominant species in the shortgrass steppe

Question: What are the plant population‐ and community‐level effects of removal of dominant plant species in the shortgrass steppe? Location: The Shortgrass Steppe Long‐Term Ecological Research site in northern Colorado, USA. Methods: We annually measured plant cover and density by species for 10 years after a one‐time aboveground removal of the dominant perennial grass, Bouteloua gracilis. Removal and control plots (3 m × 3 m) were within grazed and ungrazed locations to assess the influence of grazing on recovery dynamics. Our analyses examined plant species, functional type, and community responses to removal, paying special attention to the dynamics of subdominant and rare species. Results: Basal cover of B. gracilis increased by an average of 1% per year, but there was significantly less plant cover in treatment compared to control plots for 5 years following removal. In contrast to the lower cover in treatment plots, the plant density (number of plants m^?2) of certain subdominant perennial grasses, herbaceous perennial and annual forbs, a dwarf shrub, and cactus increased after removal of the dominant species, with no major change in species richness (number of species per 1 m × 1 m) or diversity. Subdominant species were more similar between years than rare species, but dominant removal resulted in significantly lower similarity of the subdominant species in the short term and increased the similarity of rare species in the long term. Conclusions: Removal of B. gracilis, the dominant perennial grass in the shortgrass steppe, increased the absolute density of subdominant plants, but caused little compensation of plant cover by other plants in the community and changes in species diversity.     

Positive interactions between desert granivores: localized facilitation of harvester ants by kangaroo rats

Facilitation, when one species enhances the environment or performance of another species, can be highly localized in space. While facilitation in plant communities has been intensely studied, the role of facilitation in shaping animal communities is less well understood. In the Chihuahuan Desert, both kangaroo rats and harvester ants depend on the abundant seeds of annual plants. Kangaroo rats, however, are hypothesized to facilitate harvester ants through soil disturbance and selective seed predation rather than competing with them. I used a spatially explicit approach to examine whether a positive or negative interaction exists between banner-tailed kangaroo rat (Dipodomys spectabilis) mounds and rough harvester ant (Pogonomyrmex rugosus) colonies. The presence of a scale-dependent interaction between mounds and colonies was tested by comparing fitted spatial point process models with and without interspecific effects. Also, the effect of proximity to a mound on colony mortality and spatial patterns of surviving colonies was examined. The spatial pattern of kangaroo rat mounds and harvester ant colonies was consistent with a positive interspecific interaction at small scales (<10 m). Mortality risk of vulnerable, recently founded harvester ant colonies was lower when located close to a kangaroo rat mound and proximity to a mound partly predicted the spatial pattern of surviving colonies. My findings support localized facilitation of harvester ants by kangaroo rats, likely mediated through ecosystem engineering and foraging effects on plant cover and composition. The scale-dependent effect of kangaroo rats on abiotic and biotic factors appears to result in greater founding and survivorship of young colonies near mounds. These results suggest that soil disturbance and foraging by rodents can have subtle impacts on the distribution and demography of other species.     

Multi-scale pattern analysis of a mound-building termite species

Termite mounds are a widespread feature in most African savannas. These structures exhibit high nutrient contents and often host a special vegetation composition. In this study, we analysed mound distribution patterns of a fungus-growing termite species, Macrotermes michaelseni, an important ecosystem engineer in the savannas of Namibia. Inhabited mounds taller than 0.7 m were regularly distributed. We view this pattern as a result of intraspecific competition. The heights of mounds taller than 0.7 m were correlated positively with their distance, such that mounds closer together, i.e. up to inter-mound distances of approximately 50 m, tended to be smaller than average. This indicates that intraspecific competition for foraging areas controls mound distribution pattern and colony size. Differences between mound heights increased on the spatial scale up to inter-mound distances of 80 m. We assume that the foundation of new colonies is only possible in unoccupied patches. In such patches, young colonies are able to occur close together as they have a relatively low foraging demand and therefore a low spatial demand. In contrast, their critical distance to taller colonies with higher foraging demands is rather large, which leads to the observed increasing difference of mound heights with increasing distances between them.