Competitive abilities of native grasses and non-native (Bothriochloa spp.) grasses

Old World Bluestems (OWB), introduced from Europe and Asia in the 1920s, recently have begun to raise concerns in the Great Plains. Despite suggestion in the late 1950s that OWB were weedy and negatively impacted biological diversity, they were widely introduced throughout the Great Plains for agricultural purposes. Anecdotal evidence suggests that OWB exhibit invasive characteristics that promote competitive exclusion of native species. The objective of our study was to quantify the competitive abilities of two OWB species (Caucasian bluestem; Bothriochloa bladhii (Retz.) S.T. Blake (= Bothriochloa caucasica (Trin.) C.E. Hubb.) and yellow bluestem; Bothriochloa ischaemum (L.) Keng) with three native grass species (big bluestem (Andropogon gerardii Vitman), little bluestem (Schizachyrium scoparium (Michx.) Nash), and sideoats grama (Bouteloua curtipendula (Michx.) Torr.)). A greenhouse target-neighbor study was conducted to assess both interspecific and intraspecific competition. A total of 480 pots (4.4 l) filled with native soil was used with all pair-wise combinations of species and four density treatments (six replications). Vegetative tiller height, above- and belowground biomass were measured at the end of 16 weeks. Both of the OWB significantly inhibited at least one growth parameter of the three native grass species, while most of the native species did not inhibit growth of either OWB species. Growth of B. ischaemum was enhanced when grown in association with S. scoparium. Based upon the results of our study of OWB competitive superiority and previous research, many of the characteristics possessed by OWB are found to be in common with known invasive species. Hence, we propose that two OWB are competitively superior to three common native prairie species providing them with the ability to invade and threaten the native grasslands of the Central and Southern Great Plains.     

Comparative demography of co-occurring introduced and native tussock grasses: persistence and potential expansion

Summary Demographic characteristics associated with the maintenance and growth of populations, such as seed dynamics, seedling emergence, survival, and tiller dynamics were examined for two tussock grasses, the native Agropyron spicatum and the introduced Agropyron desertorum in a 30-month field study. The introduced grass was aerially sown onto a native grassland site. Seed production of the introduced grass was greater than the native grass in both above- and below-average precipitation years. Seeds of A. spicatum were dispersed when they mature, while A. desertorum retained some seeds in inflorescences, and dispersed them slowly throughout the year. This seed retention allowed some seeds of the introduced grass to escape peak periods of seed predation during the summer and allowed seeds to be deposited constantly into the seed bank. Carryover of seeds in the seed bank beyond one year occurred in the introduced grass but not in the native species. For both species, seedling emergence occurred in both autumn or spring. Survival rates for A. desertorum were higher than A. spicatum when seedlings emerged between November and March. Survival rates of cohorts emerging before November favored A. spicatum whereas survival rates did not differ between species for cohorts emerging after March. Individuals of both species emerging after April were unable to survive the summer drought. Demographic factors associated with seeds of A. desertorum seemed to favor the maintenance and spread of this introduced grass into native stands formerly dominated by A. spicatum.     

Water relations of native and introduced C4 grasses in a neotropical savanna

Introduced African grasses are invading Neotropical savannas and displacing the native herbaceous community. This work, which is part of a program to understand the success of the African grasses, specifically investigates whether introduced and native grasses differ in their water relations. The water relations of the native Trachypogon plumosus and the successful invader Hyparrhenia rufa were studied in the field during two consecutive years in the seasonal savannas of Venezuela. The two C₄ grasses differed clearly in their responses to water stress. H. rufa consistently had higher stomatal conductance, transpiration rate, leaf water and osmotic potential and osmotic adjustment than the native T. plumosus. Also, leaf senescence occurred much earlier during the dry season in H. rufa. Both grasses showed a combination of water stress evasion and tolerance mechanisms such as stomatal sensitivity to atmospheric or soil water stress, decreased transpiring area and osmotic adjustment. Evasion mechanisms are more conspicuous in H. rufa whereas T. plumosus is more drought tolerant and uses water more conservatively. The evasion mechanisms and oportunistic use of water by H. rufa, characteristic of invading species, contribute to, but only partially explain, the success of this grass in the Neotropical savannas where it displaces native plants from sites with better water and nutrient status. Conversely, the higher water stress tolerance of t. plumosus is consistent with its capacity to resist invasion by alien grasses on shallow soils and sites with poorer nutrient and water status.     

Influence of an exotic species,Acroptilon repens (L.) DC. on seedling emergence and growth of native grasses

Our objective was to evaluate the effects of an invasive perennial forb, Acroptilon repens (Russian knapweed), on seed emergence and seedling survival of four native grass species that are important in semiarid grasslands of North America. Greenhouse experiments and field studies conducted at three sites in Colorado, USA were used to examine the response by four perennial grasses: Bouteloua gracilis, Koelaria cristata, Sporobolus cryptandrus, and Agropyon smithii to A. repens. In the greenhouse, seeds of each species were sown in pots that contained either live A. repens roots, A. repens litter on the soil surface, or both roots and litter. Field transects were placed inside stands of A. repens with adjacent control transects in the surrounding grass-dominated community. Emergence and initial survival were decreased by the presence of A. repens roots for K. cristata (35%), B. gracilis (31%), and S. cryptandrus (44%) in the greenhouse, and 57, 32, and 36%, respectively, in the field. Root weight was decreased by the presence of A. repens roots for both B. gracilis and K. cristata by more than 55% in the greenhouse. A. smithii survival and growth were unaffected by A. repens in either the greenhouse or the field. These species-specific responses to the presence of A. repens may explain, at least in part, differences in susceptibility and recovery of sites with different native grass species composition. This revised version was published online in August 2006 with corrections to the Cover Date.     

Short-term effects of fire frequency on vegetation composition and biomass in mixed prairie in south-western Manitoba

The net effects of one, two, and three spring burns in consecutive yearson the aboveground biomass, species composition, and soil variables wereassessed in two different mixed-grass prairie sites in south-western Manitoba.Precipitation in the first year was greater than the 30-year average but lowerduring the next three years. The first site (Area 6) was characterized byBouteloua gracilis, Stipa spartea,Selaginella densa and lichens, while the second, somewhatdrier site (Area 10) was dominated by B. gracilis andCarex spp. Each burn treatment was applied to 15 plots inarandom block design. Vegetation and soil data collected following the thirdburnare presented. In general, the effects of repeated burning were more pronouncedin the drier Area 10, where litter was significantly reduced with eachadditional burn. In Area 6, litter was significantly reduced only after threeyears of burning. In both sites, the cumulative effect of fire had nosignificant effect on total standing crop, even after three consecutive yearsofburning. In the drier Area 10, however, the biomass of several componentsshifted such that B. gracilis biomass increasedsignificantly and forb biomass decreased significantly after three burns. Plantcover was also affected more in the drier Area 10. The cover of B.gracilis, the dominant C₄ grass, significantly increasedand Carex spp. cover decreased after two burns in thedriersite. In Area 6, the most significant effect of fire was a reduction inSelaginella and lichen cover. Bare ground increased inbothsites as the number of burns increased. There was little change in soilnutrients with burning, but soil moisture somewhat decreased and surfacetemperature significantly increased after three burns.     

Competitive relationships between two contrasting but coexisting grasses

Leymus chinensis (Trin.) Tzvelev and Phragmites communis Trin. are co-dominant grasses in both nutrient-poor and rich steppes in the Songnen Plains of northeastern China. A replacement series experiment was conducted along a five-level nutrient gradient to test for changes in the competitive relationships of the species with nutrient availability. There were significant effects of nutrient level and species proportion on mean plant height, biomass and rhizome length. Facilitation was found in these plant attributes for both species when grown in mixture compared with monoculture, especially at high nutrient levels. For example, L. chinensis grew taller and intercepted more light when in mixture. The aboveground dominance of P. communis did not decrease in mixture, possibly because it benefited from belowground interactions with L. chinensis. Biomass allocation was apparently influenced by both above and belowground competition. For example, L. chinensis allocated much more biomass to shoots and less to roots when neighbor number was increased, suggesting that aboveground competition occurred. Root: shoot ratios of L. chinensis in monoculture decreased gradually with increasing nutrient availability, whereas the ratio in mixture declined rapidly, implying intense competition for light at high levels. Our results support the hypothesis that nutrient competition is more important than light competition when soil nutrients are scarce. Although the interactions between these species benefit both, P. communis is more likely to displace L. chinensis at high nutrient availability.     

Nitrogen mineralization in native cultivated and abandoned fields in shortgrass steppe

We conducted a set of in situ incubations to evaluate patterns of N availability among dominant land uses in the shortgrass steppe region of Colorado, USA, and to assess recovery of soil fertility in abandoned fields. Replicated 30 d incubations were performed in 3 sets of native (never cultivated), abandoned (cultivated until 1937), and currently cultivated, fallow fields. Net N mineralization and the percentage of total N that was mineralized increased in the order: native, abandoned, cultivated. Higher soil water content in fallow fields is the most likely reason for greater mineralization in cultivated fields, while higher total organic C and C/N ratios in native and abandoned fields may explain differences in mineralization between these land uses. Recovery of soil organic matter in abandoned fields appears to involve accumulation of soil C and N under perennial plants, but probable methodological artifacts complicate evaluation of the role of individual plants in recovery of N availability. Higher N mineralization and turnover in cultivated fields may make them more susceptible to N losses; recovery of N cycling in abandoned fields appears to involve a return to slower N turnover and tighter N cycling similar to native shortgrass steppe.     

土壤水分及其垂直分布对内蒙古草原冰草和冷蒿竞争的影响

内蒙古退化冷蒿草原围封恢复演替3-4年,植物群落更替出现半灌木冷蒿（Artemisia frigida）向禾草冰草（Agropyron cristatum）群落突然转变的现象,这对内蒙古草原植被恢复具有积极的作用,但目前关于冰草-冷蒿演替和竞争机制的研究鲜见报道。水分作为草原植被生长的主要限制因子,是影响群落演替的重要因素。气候干旱和过度放牧导致草原土壤水分垂直分布发生明显变化。由于冰草和冷蒿的根系分布深浅不同,土壤水分垂直变化可能会影响冰草-冷蒿的竞争。为探讨土壤水分及其垂直分布对冰草和冷蒿竞争的影响,本盆栽试验设置冰草、冷蒿单种以及混种处理,并进行不同土壤深度（0-30cm,30-60cm）的水分处理（上干下湿、上湿下干、上干下干）,结果表明：（1）与冷蒿相比,冰草受土壤水分变化影响显著,相对于上干下干处理,冰草在湿润（上干下湿或者上湿下干）处理的地上、地下生物量均显著增加,而冷蒿没有显著变化;（2）上干下湿和上湿下干处理间,冰草、冷蒿的地上、地下总生物量和根长差异均不显著;（3）不同水分处理,冰草和冷蒿的根系均可分布在30-60cm土壤中,且土壤30-60cm层冰草单种的根生物量和根长显著高于冷蒿单种;（4）相对竞争强度和竞争攻击力系数表明：湿润处理冰草的竞争力大于冷蒿;上干下干处理冷蒿的竞争力大于冰草。土壤水分变化引起冰草、冷蒿的明显竞争,上湿下干与上干下湿处理间冰草与冷蒿竞争力差异不显著。由于冰草和冷蒿均为多年生物种,冰草-冷蒿的竞争实验仍需在今后的研究中反复地验证。     

Edge effects and intraguild predation in native and introduced centipedes: evidence from the field and from laboratory microcosms

Human alteration of habitat has increased the proportion of forest edge in areas of previously continuous forest. This edge habitat facilitates invasion of exotic species into remaining fragments. The ability of native species to resist invasion varies and may depend on intrinsic variables such as dispersal and reproductive rates as well as external factors such as rate of habitat change and the density of populations of introduced species in edge habitat. We examined the distributional and competitive relationships of two members of the class Chilopoda, Scolopocryptops sexspinosus, a centipede native to the eastern US, and Lithobius forficatus, an exotic centipede introduced from Europe. We found that L. forficatus was most abundant in edge habitat and S. sexspinosus was most abundant in the interior habitat at our field sites. Although L. forficatus was present in habitat interiors at 11 of 12 sites, there was no correlation between fragment size and numbers of L. forficatus in interior habitat. The native centipede was rarely found occupying fragment edges. We used laboratory microcosms to examine potential competitive interactions and to indirectly assess prey preferences of the two species. In microcosms both species consumed similar prey, but the native centipede, S. sexspinosus, acted as an intraguild predator on the introduced centipede. Native centipedes were competitively superior in both intraspecific and interspecific pairings. Our results suggest that intraguild predation may aid native centipedes in resisting invasion of introduced centipedes from edge habitat.     

Mycorrhizae transfer carbon from a native grass to an invasive weed: evidence from stable isotopes and physiology

Invasive exotic weeds pose one of the earth's most pressing environmental problems. Although many invaders completely eliminate native plant species from some communities, ecologists know little about the mechanisms by which these exotics competitively exclude other species. Mycorrhizal fungi radically alter competitive interactions between plants within natural communities, and a recent study has shown that arbuscular mycorrhizal (AM) fungi provide a substantial competitive advantage to spotted knapweed, Centaurea maculosa, a noxious perennial plant that has spread throughout much of the native prairie in the northwestern U.S. Here we present evidence that this advantage is potentially due to mycorrhizally mediated transfer of carbon from a native bunchgrass, Festuca idahoensis, to Centaurea. Centaurea maculosa, Festuca idahoensis (Idaho fescue, C₃), and Bouteloua gracilis (blue gramma, C₄) were grown in the greenhouse either alone or with Centaurea in an incomplete factorial design with and without AM fungi. Centaurea biomass was 87–168% greater in all treatments when mycorrhizae were present in the soil (P < 0.0001). However, Centaurea biomass was significantly higher in the treatment with both mycorrhizae and Festuca present together than in any other treatment combination (P < 0.0001). This high biomass was attained even though Centaurea photosynthetic rates were 14% lower when grown with Festuca and mycorrhizae together than when grown with Festuca without mycorrhizae. Neither biomass nor photosynthetic rates of Centaurea were affected by competition with the C₄ grass Bouteloua either with or without mycorrhizae. The stable isotope signature of Centaurea leaves grown with Festuca and mycorrhizae was more similar to that of Festuca, than when Centaurea was grown alone with mycorrhizae (P = 0.06), or with Festuca but without mycorrhizae (P = 0.09). This suggests that carbon was transferred from Festuca to the invasive weed. We estimated that carbon transferred from Festuca by mycorrhizae contributed up to 15% of the aboveground carbon in Centaurea plants. Our results indicate that carbon parasitism via AM soil fungi may be an important mechanism by which invasive plants out compete their neighbors, but that this interaction is highly species-specific.     

Soil modification by invasive plants: effects on native and invasive species of mixed-grass prairies

Invasive plants are capable of modifying attributes of soil to facilitate further invasion by conspecifics and other invasive species. We assessed this capability in three important plant invaders of grasslands in the Great Plains region of North America: leafy spurge (Euphorbia esula), smooth brome (Bromus inermis) and crested wheatgrass (Agropyron cristatum). In a glasshouse, these three invasives or a group of native species were grown separately through three cycles of growth and soil conditioning in both steam-pasteurized and non-pasteurized soils, after which we assessed seedling growth in these soils. Two of the three invasive species, Bromus and Agropyron, exhibited significant self-facilitation via soil modification. Bromus and Agropyron also had significant facilitative effects on other invasives via soil modification, while Euphorbia had significant antagonistic effects on the other invasives. Both Agropyron and Euphorbia consistently suppressed growth of two of three native forbs, while three native grasses were generally less affected. Almost all intra- and interspecific effects of invasive soil conditioning were dependent upon presence of soil biota from field sites where these species were successful invaders. Overall, these results suggest that that invasive modification of soil microbiota can facilitate plant invasion directly or via ‘cross-facilitation’ of other invasive species, and moreover has potential to impede restoration of native communities after removal of an invasive species. However, certain native species that are relatively insensitive to altered soil biota (as we observed in the case of the forb Linum lewisii and the native grasses), may be valuable as ‘nurse’species in restoration efforts.     

Soil water dynamics,transpiration, and water losses in a crested wheatgrass and native shortgrass ecosystem

The status of water in soil and vegetation was monitored in a stand of crested wheatgrass (Agropyron cristatum) and a nearby shortgrass steppe during a growing season. This was done to determine if water use and losses were similar among two very different communities in a similar climate. Precipitation was similar throughout the study period for both the crested wheatgrass and native shortgrass communities. However, the native shortgrass community with greater root biomass had consistently greater soil water depletion in the deeper soil horizons than was found in the crested wheatgrass community. Greater depletion of soil water by native shortgrass species suggests that they might be more competitive than crested wheatgrass in a water-limited environment.Crested wheatgrass maintained high leaf water potential early in the season, but lower water potential during the latter part of the growing season as compared with the major species of the shortgrass steppe, blue grama (Bouteloua gracilis) and western wheatgrass (Agropyron smithii). Leaf conductance was lower for crested wheatgrass than for the native grasses during the later part of the growing season. Consequently, seasonal transpiration for crested wheatgrass was lower when compared with blue grama or western wheatgrass. Lower conductance allowed crested wheatgrass to maintain relatively high internal water potential and may have accounted for less soil water use at deeper soil depths during the latter part of the growing season.Water loss through transpiration was less for western wheatgrass than for either blue grama or crested wheatgrass because western wheatgrass had less leaf area. However, western wheatgrass was as efficient as the other species in its use of water. Crested wheatgrass transpired more water than blue grama early in the growing season, but less than either native species for the remainder of the growing season. Estimated seasonal transpiration loss was greater in the shortgrass ecosystem than in the established crested wheatgrass stand.     

Drought response of a native and introduced Hawaiian grass

The alien grass, Pennisetum setaceum, dominates many of the lowland arid regions that once supported native Heteropogon contortus grassland on the island of Hawaii. Response to drought in a glasshouse was compared between these C₄ grasses to test if success as an invader is related to drought tolerance or plasticity for traits that confer drought tolerance. Pennisetum produced 51% more total biomass, allocated 49% more biomass to leaves, and had higher net photosynthetic rates (P _n) on a leaf area basis than Heteropogon. Plants of both species under drought produced less total biomass and increased their allocation to roots compared to well-watered plants, but there was no difference between the two species in the magnitude of these responses. The decline in P _n with decreasing leaf water potential (₁) was greater for Pennisetum compared to Heteropogon. Plasticity in the response of P _n to ₁, osmotic potentials, and the water potentials at turgor loss in response to drought were not different between the two species. Stomata were more responsive to w in Heteropogon than in Pennisetum and for well-watered plants compared to droughted plants. Plasticity for the stomatal response to w, however, was not different between the species. There was no evidence that the alien, Pennisetum, had greater plasticity for traits related to drought tolerance compared to the native, Heteropogon. Higher P _n and greater biomass allocation to leaves resulted in greater growth for Pennisetum compared to Heteropogon and may explain the success of Pennisetum as an invader of lowland arid zones on Hawaii.     

Competitive ability is linked to rates of water extraction

Summary The relative competitive abilities of Agropyron desertorum and Agropyron spicatum under rangeland conditions were compared using Artemisia tridentata ssp. wyomingensis transplants as indicator plants. We found A. desertorum to have substantially greater competitive ability than A. spicatum as manifested by the responses of Artemisia shrubs that were transplanted into nearly monospecific stands of these grass species. The Artemisia indicator plants had lower survival, growth, reproduction, and late-season water potential in the neighborhoods dominated by A. desertorum than in those dominated by A. spicatum. In similar, essentially monospecific grass stands, neutron probe soil moisture measurements showed that stands of A. desertorum extracted water more rapidly from the soil profile than did those of A. spicatum. These differences in extraction rates correlate clearly with the differences in indicator plant success in the respective grass stands. Nitrogen and phosphorus concentrations in Artemisia tissues suggested these nutrients were not limiting indicator plant growth and survival in the A. desertorum plots.     

Longevity and turnover of roots in the shortgrass steppe: influence of diameter and depth

We used minirhizotrons to determine patterns of root longevity andturnover for the perennial bunchgrass Bouteloua gracilisinthe shortgrass steppe of eastern Colorado, USA. We hypothesized that rootlongevity would be partially controlled by root diameter, following previouslyobserved patterns in woody plants. In addition, we hypothesized that rootturnover would be greatest in surface soil horizons and decrease with depth dueto variation in soil moisture availability and temperature. Root longevity wascorrelated with root diameter. Median life span of roots > 0.4mm was approximately 320 days, while roots < 0.2mmhad a median life span of 180 days. There was approximately a 6%decreasein the likelihood of mortality with a 0.10-mm increase inroot diameter, controlling for the effect of depth in the soil profile. Rootlength production and mortality were highest in the upper20 cm of the soil profile and decreased with depth.However,because root length density also decreased with depth, there were nosignificantdifferences in turnover rate of root length among sampling intervals. Turnoverwas approximately 0.86 yr^–1 based on root length production,while turnover was 0.35 yr^–1 using root length mortality as ameasurement of flux. The imbalance between turnover estimates may be aconsequence of the time the minirhizotrons were in place prior to imaging or mayresult from our lack of over-winter measures of mortality. Our worksuggests that Bouteloua gracilis roots have complex lifehistory strategies, similar to woody species. Some portion of the root systemishighly ephemeral, while slightly larger roots persist much longer. Thesedifferences have implications for belowground carbon and nitrogen cycles in theshortgrass steppe.     

Balance of Competitive and Facilitative Effects of Exotic Trees on a Native Patagonian Grass

The balance between facilitation and competition in plants changes with species characteristics and environmental conditions. Facilitative effects are common in natural ecosystems, particularly in stressful environments or years. Contrarily, in artificial associations of plants, such as agroforestry systems, some authors have suggested that even when facilitative effects may occur, net balance of tree effects on grasses is usually negative, particularly in dry environments. The aim of this study was to determine the net effect of the exotic ponderosa pine on the native grass Festuca pallescens (St. Ives) Parodi in agroforestry systems in Patagonia. Soil water content, plant water status, and relative growth were measured in the grass growing in different treatments (determined by tree cover level) during two growing seasons with contrasting climatic conditions. Facilitative effects of trees over grass water status were recorded only when water availability was high. A net negative effect was detected on dates when soil water content was very low and evaporative demand was high. The strength of these negative effects depended on tree density and climatic conditions, being higher in treatments with lower tree canopy cover. These results indicate that the positive effect of trees could only be expected under relatively low stress conditions. However, relative growth of grasses was always similar in plants growing in forested plots than in open grassland. Differences in biomass allocation for grasses growing in shade and open habitats may reconcile these contrary results. Our results highlight the importance of the physiology of a species (relative drought and shade tolerance) in determining the response of a plant to a particular interacting species.     

Successional dynamics of a semiarid grassland: effects of soil texture and disturbance size

We evaluated effects of soil texture and disturbance size on the successional dynamics of a semiarid grassland dominated by the perennial bunchgrass, Bouteloua gracilis (H.B.K.) Lag. ex Griffiths. A spatially-explicit gap dynamics simulation model was used to evaluate recovery patterns. The model simulates establishment, growth, and mortality of individual plants on an array of small plots through time at an annual time step. Each simulated disturbance consisted of a grid of plots of the same soil texture interconnected by processes associated with dispersal of B. gracilis seeds. Soil texture was incorporated into the model as effects on seed germination, seedling establishment, and subsequent growth of B. gracilis. Five soil texture classes and five disturbance sizes were simulated.Soil texture was more important to recovery of B. gracilis than either size of a disturbance or location of plots within a disturbance. Constraints on recruitment of seedlings had a greater effect on recovery than constraints associated with plant growth. Fastest recovery occurred on soils with the largest silt content, the variable that affects seedling establishment. Disturbances with slowest recovery were on soils with low silt contents, and either high or low water-holding capacity, the variable that affects plant growth. Biomass and recovery decreased as disturbance size increased, and as distance from a disturbed plot to the edge of the disturbance increased. In most cases, important interactions between soil texture and disturbance size on recovery were not found.     

Response of the invasive Centaurea maculosa and two native grasses to N-pulses

Nitrogen is often a limiting resource on semi-arid grasslands. During the growing season, N is often only available during short-term pulses associated with wetting events. The Eurasian forb Centaurea maculosa Lam. has invaded millions of hectares of semi-arid grasslands in western North America. C. maculosa's success could be attributed to greater use of N-pulses, or more efficient use of N supplied in those pulses compared with native grasses. In a glasshouse, C. maculosa and two native grasses, the caespitose Pseudoroegneria spicata [Scribn. and Smith] A. Love and the rhizomatous Pascopyrum smithii [Rybd.] A. Love, were established in mixed- and monoculture combinations, and then conditioned to weekly N-pulses of 8, 24, or 72 h for 8 weeks. These pulse durations are typical on semi-arid grasslands. At the end of the 8 weeks, plants were exposed to ¹⁵N-labeled nitrate (¹⁵NO₃ ^–) for 8 h and harvested 16 h later to compare short-term root uptake of ¹⁵NO₃ ^–. C. maculosa did not have greater enrichment (atom % ¹⁵N), rate of ¹⁵N-uptake (mol g^–1 h^–1), or ¹⁵N acquired (relative to ¹⁵N applied) than the grasses. C. maculosa's ¹⁵N-uptake per unit mass was relatively consistent across pulse durations, whereas ¹⁵N-uptake was lower at the longer pulse durations for the grasses. In general, C. maculosa acquired more of the applied ¹⁵N than P. spicata but less than P. smithii. ¹⁵N acquired was often influenced by the neighbour's identity. Regarding growth responses, C. maculosa produced more total biomass than the grasses, except for P. smithii plants growing with C. maculosa conditioned to 72 h pulses of N. Root mass ratios varied depending on the neighbor. Overall, C. maculosa used nitrogen less efficiently than the grasses. C. maculosa's success as an invasive species cannot be explained wholly by a greater response to N-pulses or more efficient use of N-pulses compared with native grasses with which it competes.     

Local and regional differences in abundance of co-dominant grasses in the shortgrass steppe: a modeling analysis of potential causes

The shortgrass steppe is co-dominated by two C₄perennial grasses, Bouteloua gracilis andBuchloë dactyloides. At our site and throughouteastern Colorado Bouteloua gracilis has higher cover andbiomass than Buchloë dactyloides. We hypothesizedthatthe interaction between seedling recruitment differences and disturbance regimeand tolerance to drought conditions were the most likely causes of the observeddifferences in relative abundances. We used a simulation model to investigatethe relative importance of the three factors. We studied plant biomass of thesetwo species in 18 simulated treatments that resulted from a factorialcombination of seedling recruitment differences, disturbance regime, andtolerance to drought conditions. Analysis of the simulation outputs with ANOVAindicated that biomass of each species responded positively to increases inrecruitment probability. Increasing disturbance frequency favoredBuchloë dactyloides over Boutelouagracills, whereas the susceptibility Buchloëdactyloides to drought favored Boutelouagracilis. The results indicated that differences in droughttoleranceand seedling recruitment probabilities along with their interactions withdisturbance regimes exert a major control on the biomass of the species. In theabsence of or with intermediate disturbance, a higher recruitment probabilityand greater tolerance to drought of Bouteloua gracilisthanBuchloë dactyloides yielded patterns of relativebiomass similar to the patterns observed in the shortgrass steppe.     

Influence of soil depth on the decomposition of Bouteloua gracilis roots in the shortgrass steppe

The distribution and turnover of plant litter contribute to soil structure, the availability of plant nutrients, and regional budgets of greenhouse gasses. Traditionally, studies of decomposition have focused on the upper soil profile. Other work has shown that temperature, precipitation, and soil texture are important determinates of patterns of decomposition. Since these factors all vary through a soil profile, it has been suggested that decomposition rates may vary with depth in a soil profile. In this work, we examine patterns of root decomposition through a shortgrass steppe soil profile. We buried fresh root litter from Bouteloua gracilis plants in litterbags at 10, 40, 70, and 100 cm. Litterbags were retrieved six times between July 1996 and May 1999. We found that the decomposition rate for fresh root litter was approximately 50% slower at 1 m than it was at 10 cm. After 33 months, 55% of the root mass buried at 10 cm remained, while 72% of the root mass buried at 1 m was still present. This corresponds to a 19-year residence time for roots at 10 cm and a 36-year residence time for roots at 1 m. Mass loss rates decreased linearly from 10 cm to 1 m. Patterns of total carbon and cellulose loss rates followed those of mass loss rates. Roots at 1 m tended to accumulate lignin-like compounds over the course of the experiment. Differences in the stabilization of lignin may be a consequence of differences in microbial community through a shortgrass steppe soil profile.