Urbanization increases grassland carbon pools: effects of landscaping in Colorado's front range.

During the past few decades, urban and suburban developments have grown at unprecedented rates and extents with unknown consequences for ecosystem function. Carbon pools of soil and vegetation on landscaped properties were examined in the Front Range of Colorado, USA, in order to characterize vegetation and soils found in urban green spaces; analyze their aboveground biomass, vegetative C storage, and soil C storage; and compare these suburban ecosystem properties to their counterparts in native grassland and cultivated fields. Anthropogenic activities leave clear signatures on all three C compartments measured. Management level dominates the response of grass production, biomass, and N tissue concentration. This, in turn, influences the amount of C and N both stored in and harvested from sites. The site age dominates the amount of woody biomass as well as soil C and N. Soil texture only secondarily affects total soil carbon and total bulk density. Established urban green spaces harbor larger C pools, more than double in some cases, than native grasslands or agricultural fields on a per-area basis. Lawn grass produces more biomass and stores more C than local prairie or agricultural fields. Introduced woody vegetation comprises a substantial C pool in urban green spaces and represents a new ecosystem feature. After an initial decrease with site development, soil organic carbon (SOC) pools surpass those in grasslands within two decades. In addition to the marked increase of C pools through time, a shift in storage from belowground to aboveground occurs. Whereas grasslands store approximately 90% of C belowground, urban green spaces store a decreasing proportion of the total C belowground in soils through time, reaching approximately 70% 30-40 years after construction. Despite the substantial increase in C pools in this urban area, it is important to recognize that this shift is distinct from C sequestration since it does not account for a total C budget, including increased anthropogenic C emissions from these sites.     

Soil feedback of exotic savanna grass relates to pathogen absence and mycorrhizal selectivity

Enemy release of exotic plants from soil pathogens has been tested by examining plant-soil feedback effects in repetitive growth cycles. However, positive soil feedback may also be due to enhanced benefit from the local arbuscular mycorrhizal fungi (AMF). Few studies actually have tested pathogen effects, and none of them did so in arid savannas. In the Kalahari savanna in Botswana, we compared the soil feedback of the exotic grass Cenchrus biflorus with that of two dominant native grasses, Eragrostis lehmanniana and Aristida meridionalis. The exotic grass had neutral to positive soil feedback, whereas both native grasses showed neutral to negative feedback effects. Isolation and testing of root-inhabiting fungi of E. lehmanniana yielded two host-specific pathogens that did not influence the exotic C. biflorus or the other native grass, A. meridionalis. None of the grasses was affected by the fungi that were isolated from the roots of the exotic C. biflorus. We isolated and compared the AMF community of the native and exotic grasses by polymerase chain reaction-denaturing gradient gel elecrophoresis (PCR-DGGE), targeting AMF 18S rRNA. We used roots from monospecific field stands and from plants grown in pots with mixtures of soils from the monospecific field stands. Three-quarters of the root samples of the exotic grass had two nearly identical sequences, showing 99% similarity with Glomus versiforme. The two native grasses were also associated with distinct bands, but each of these bands occurred in only a fraction of the root samples. The native grasses contained a higher diversity of AMF bands than the exotic grass. Canonical correspondence analyses of the AMF band patterns revealed almost as much difference between the native and exotic grasses as between the native grasses. In conclusion, our results support the hypothesis that release from soil-borne enemies may facilitate local abundance of exotic plants, and we provide the first evidence that these processes may occur in arid savanna ecosystems. Pathogenicity tests implicated the involvement of soil pathogens in the soil feedback responses, and further studies should reveal the functional consequences of the observed high infection with a low diversity of AMF in the roots of exotic plants.     

Are soil mite assemblages structured by the identity of native and invasive alien grasses?

Associations between plants and animals in aboveground communities are often predictable and specific. This has been exploited for the purposes of estimating the diversity of animal species based on the diversity of plant species. The introduction of invasive alien plants into an ecosystem can result in dramatic changes in both the native plant and animal assemblages. Few data exist at the species level to determine whether belowground animal assemblages share the same degree of association to plants. The hypotheses that soil mites (Acari) form assemblages specifically associated with different native grass species in an unmanipulated natural ecosystem and that invasive alien grasses will impact soil mite assemblage composition in this setting were tested. Soil mites sampled beneath five native and two invasive alien species of grasses at the Konza Prairie Biological Station, Kansas, USA, were similarly abundant, species rich, diverse, and taxonomically distinct. No mite species had affinities for a specific grass species. There was no evidence from analysis of similarity, canonical correspondence analysis, or a nonparametric assemblage analysis that the assemblage composition of soil mites was specific to grass species. Results suggest that soil mite assemblages were more related to characteristics of the plant assemblage as a whole or prevailing soil conditions. The most recent invasive alien grass did not support a successionally younger mite fauna, based on the ratio of mesostigmatid to oribatid mites, and neither of the two invasive grasses influenced mite assemblage structure, possibly because they had not yet substantially altered the soil environment. Our results suggest that extrapolations of soil mite diversity based on assumptions of plant specificity would be invalid.     

Bromus tectorum invasion alters nitrogen dynamics in an undisturbed arid grassland ecosystem

The nonnative annual grass Bromus tectorum has successfully replaced native vegetation in many arid and semiarid ecosystems. Initial introductions accompanied grazing and agriculture, making it difficult to separate the effects of invasion from physical disturbance. This study examined N dynamics in two recently invaded, undisturbed vegetation associations (C3 and C4). The response of these communities was compared to an invaded/ disturbed grassland. The invaded/disturbed communities had higher surface NH4+ input in spring, whereas there were no differences for surface input of NO3-. Soil inorganic N was dominated by NH4+, but invaded sites had greater subsurface soil NO3-. Invaded sites had greater total soil N at the surface four years post-invasion in undisturbed communities, but total N was lower in the invaded/disturbed communities. Soil delta15N increased with depth in the noninvaded and recently invaded communities, whereas the invaded/disturbed communities exhibited the opposite pattern. Enriched foliar delta15N values suggest that Bromus assimilated subsurface NO3-, whereas the native grasses were restricted to surface N. A Rayleigh distillation model accurately described decomposition patterns in the noninvaded communities where soil N loss is accompanied by increasing soil delta15N; however, the invaded/ disturbed communities exhibited the opposite pattern, suggesting redistribution of N within the soil profile. This study suggests that invasion has altered the mechanisms driving nitrogen dynamics. Bromus litter decomposition and soil NO3- concentrations were greater in the invaded communities during periods of ample precipitation, and NO3- leached from the surface litter, where it was assimilated by Bromus. The primary source of N input in these communities is a biological soil crust that is removed with disturbance, and the lack of N input by the biological soil crust did not balance N loss, resulting in reduced total N in the invaded/disturbed communities. Bromus produced a positive feedback loop by leaching NO3- from decomposing Bromus litter to subsurface soil layers, accessing that deepsoil N pool with deep roots and returning that N to the surface as biomass and subsequent litter. Lack of new inputs combined with continued loss will result in lower total soil N, evidenced by the lower total soil N in the invaded/disturbed communities.     

Grassland establishment under varying resource availability: a test of positive and negative feedback

The traditional logic of carbon (C) and nitrogen (N) interactions in ecosystems predicts further increases or decreases in productivity (positive feedback) in response to high and low fertility in the soil, respectively; but the potential for development of feedback in ecosystems recovering from disturbance is less well understood. Furthermore, this logic has been challenged in grassland ecosystems where frequent fires or grazing may reduce the contribution of aboveground litter inputs to soil organic matter pools and nutrient supply for plant growth, relative to forest ecosystems. Further, if increases in plant productivity increase soil C content more than soil N content, negative feedback may result from increased microbial demand for N making less available for plant growth. We used a field experiment to test for feedback in an establishing grassland by comparing aboveground net primary productivity (ANPP) and belowground pools and fluxes of C and N in soil with enriched, ambient, and reduced N availability. For eight years annual N enrichment increased ANPP, root N, and root tissue quality, but root C:N ratios remained well above the threshold for net mineralization of N. There was no evidence that N enrichment increased root biomass, soil C or N accrual rates, or storage of C in total, microbial, or mineralizable pools within this time frame. However, the net nitrogen mineralization potential (NMP) rate was greater following eight years of N enrichment, and we attributed this to N saturation of the microbial biomass. Grassland developing under experimentally imposed N limitation through C addition to the soil exhibited ANPP, root biomass and quality, and net NMP rate similar to the ambient soil. Similarity in productivity and roots in the reduced and ambient N treatments was attributed to the potentially high nitrogen-use efficiency (NUE) of the dominant C4 grasses, and increasing cover of legumes over time in the C-amended soil. Thus, in a developing ecosystem, positive feedback between soil N supply and plant productivity may promote enhanced long-term N availability and override progressive N limitation as C accrues in plant and soil pools. However, experimentally imposed reduction in N availability did not feed back to reduce ANPP, possibly due to shifts in NUE and functional group composition.     

Long-term population dynamics of seeded plants in invaded grasslands

In recent decades, dozens of studies have involved attempts to introduce native and desirable nonnative plant species into grasslands dominated by invasive weeds. The newly introduced plants have proved capable of establishing, but because they are rarely monitored for more than four years, it is unknown if they have a high likelihood of persisting and suppressing invaders for the long-term. Beyond invaded grasslands, this lack of long-term monitoring is a general problem plaguing efforts to reintroduce a range of taxa into a range of ecosystems. We introduced species from seed and then periodically measured plant abundances for nine years at one site and 15 years at a second site. To our knowledge, our 15-year data are the longest to date from a seeding experiment in invaded, never-cultivated grassland. At one site, three seeded grasses maintained high densities for three or more years, but then all or nearly all individuals died. At the second site, one grass performed similarly, but two other grasses proliferated and at least one greatly suppressed the dominant invader (Centaurea maculosa). In one study, our point estimate suggests that the seeded grass Thinopyrum intermedium reduced C. maculosa biomass by 93% 15 years after seeding. In some cases, data from three and fewer years after seeding falsely suggested that seeded species were capable of persisting within the invaded grassland. In other cases, data from as late as nine years after seeding falsely suggested seeded populations would not become large enough to suppress the invader. These results show that seeded species sometimes persist and suppress invaders for long periods, but short-term data cannot predict if, when, or where this will occur. Because short-term data are not predictive of long-term seeded species performances, additional long-term data are needed to identify effective practices, traits, and species for revegetating invaded grasslands.     

Carbon stocks across a chronosequence of thinned and unmanaged red pine (Pinus resinosa) stands

Forests function as a major global C sink, and forest management strategies that maximize C stocks offer one possible means of mitigating the impacts of increasing anthropogenic CO2 emissions. We studied the effects of thinning, a common management technique in many forest types, on age-related trends in C stocks using a chronosequence of thinned and unmanaged red pine (Pinus resinosa) stands ranging from 9 to 306 years old. Live tree C stocks increased with age to a maximum near the middle of the chronosequence in unmanaged stands, and increased across the entire chronosequence in thinned stands. C in live understory vegetation and C in the mineral soil each declined rapidly with age in young stands but changed relatively little in middle-aged to older stands regardless of management. Forest floor C stocks increased with age in unmanaged stands, but forest floor C decreased with age after the onset of thinning around age 40 in thinned stands. Deadwood C was highly variable, but decreased with age in thinned stands. Total ecosystem C increased with stand age until approaching an asymptote around age 150. The increase in total ecosystem C was paralleled by an age-related increase in total aboveground C, but relatively little change in total belowground C. Thinning had surprisingly little impact on total ecosystem C stocks, but it did modestly alter age-related trends in total ecosystem C allocation between aboveground and belowground pools. In addition to characterizing the subtle differences in C dynamics between thinned and unmanaged stands, these results suggest that C accrual in red pine stands continues well beyond the 60-100 year management rotations typical for this system. Management plans that incorporate longer rotations and thinning in some stands could play an important role in maximizing C stocks in red pine forests while meeting other objectives including timber extraction, biodiversity conservation, restoration, and fuel reduction goals.     

Effect of rhizosphere on soil microbial community and pyrene biodegradation

To access the influence of a vegetation on soil microorganisms toward organic pollutant biogegration, this study examined the rhizospheric effects of four plant species (sudan grass, white clover, alfalfa, and fescue) on the soil microbial community and in-situ pyrene (PYR) biodegradation. The results indicated that the spiked PYR levels in soils decreased substantially compared to the control soil without planting. With equal planted densities, the efficiencies of PYR degradation in rhizosphere with sudan grass, white clover, alfalfa and fescue were 34.0%, 28.4%, 27.7%, and 9.9%, respectively. However, on the basis of equal root biomass the efficiencies were in order of white clover >> alfalfa > sudan > fescue. The increased PYR biodegradation was attributed to the enhanced bacterial population and activity induced by plant roots in the rhizosphere. Soil microbial species and biomasses were elucidated in terms of microbial phospholipid ester-linked fatty acid (PLFA) biomarkers. The principal component analysis (PCA) revealed significant changes in PLFA pattern in planted and non-planted soils spiked with PYR. Total PLFAs in planted soils were all higher than those in non-planted soils. PLFA assemblages indicated that bacteria were the primary PYR degrading microorganisms, and that Gram-positive bacteria exhibited higher tolerance to PYR than Gram-negative bacteria did.     

Forest productivity decline caused by successional paludification of boreal soils.

Long-term forest productivity decline in boreal forests has been extensively studied in the last decades, yet its causes are still unclear. Soil conditions associated with soil organic matter accumulation are thought to be responsible for site productivity decline. The objectives of this study were to determine if paludification of boreal soils resulted in reduced forest productivity, and to identify changes in the physical and chemical properties of soils associated with reduction in productivity. We used a chronosequence of 23 black spruce stands ranging in postfire age from 50 to 2350 years and calculated three different stand productivity indices, including site index. We assessed changes in forest productivity with time using two complementary approaches: (1) by comparing productivity among the chronosequence stands and (2) by comparing the productivity of successive cohorts of trees within the same stands to determine the influence of time independently of other site factors. Charcoal stratigraphy indicates that the forest stands differ in their fire history and originated either from high- or low-severity soil burns. Both chronosequence and cohort approaches demonstrate declines in black spruce productivity of 50-80% with increased paludification, particularly during the first centuries after fire. Paludification alters bryophyte abundance and succession, increases soil moisture, reduces soil temperature and nutrient availability, and alters the vertical distribution of roots. Low-severity soil burns significantly accelerate rates of paludification and productivity decline compared with high-severity fires and ultimately reduce nutrient content in black spruce needles. The two combined approaches indicate that paludification can be driven by forest succession only, independently of site factors such as position on slope. This successional paludification contrasts with edaphic paludification, where topography and drainage primarily control the extent and rate of paludification. At the landscape scale, the fire regime (frequency and severity) controls paludification and forest productivity through its effect on soil organic layers. Implications for global carbon budgets and sustainable forestry are discussed.     

Above- and belowground responses of Arctic tundra ecosystems to altered soil nutrients and mammalian herbivory

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The exploitation ecosystem hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.     

Effect of rhizosphere on soil microbial community and in-situ pyrene biodegradation

To access the influence of a vegetation on soil microorganisms toward organic pollutant biogegration, this study examined the rhizospheric effects of four plant species (sudan grass, white clover, alfalfa, and fescue) on the soil microbial community and in-situ pyrene (PYR) biodegradation. The results indicated that the spiked PYR levels in soils decreased substantially compared to the control soil without planting. With equal planted densities, the efficiencies of PYR degradation in rhizosphere with sudan grass, white clover, alfalfa and fescue were 34.0%, 28.4%, 27.7%, and 9.9%, respectively. However, on the basis of equal root biomass the efficiencies were in order of white clover >> alfalfa > sudan > fescue. The increased PYR biodegradation was attributed to the enhanced bacterial population and activity induced by plant roots in the rhizosphere. Soil microbial species and biomasses were elucidated in terms of microbial phospholipid ester-linked fatty acid (PLFA) biomarkers. The principal component analysis (PCA) revealed significant changes in PLFA pattern in planted and non-planted soils spiked with PYR. Total PLFAs in planted soils were all higher than those in non-planted soils. PLFA assemblages indicated that bacteria were the primary PYR degrading microorganisms, and that Gram-positive bacteria exhibited higher tolerance to PYR than Gram-negative bacteria did.     

Inversion of plant dominance-diversity relationships along a latitudinal stress gradient

Species interactions affect plant diversity through the net effects of competition and facilitation, with the latter more prevalent in physically stressful environments when plant cover ameliorates abiotic stress. One explanation for species loss in invader-dominated systems is a shift in the competition-facilitation balance, with competition intensifying in areas formerly structured by facilitation. We test this possibility with a 10-site prairie meta-experiment along a 500-km latitudinal stress gradient, quantifying the relationships among abiotic stress, exotic dominance, and native plant recruitment over five years. The latitudinal gradient is inversely correlated with abiotic stress, with lower latitudes more moisture- and nutrient-limited. We observed strong negative effects by invasive dominant grasses on plant establishment, but only in northern sites with lower-stress environments. At these locations, disturbance was critical for recruitment by reducing the suppressive dominant (invasive) canopy. In more stressful environments to the south, the impacts of the dominant invaders on plant establishment became facilitative, and diversity was more limited by seed availability. Disturbance prevented recruitment because seedling survival depended on a protective plant canopy, presumably because the canopy reduced temperature or moisture stress. Seed limitation was similarly prevalent in all sites. Our work confirms the importance of facilitation as an organizing process for plants in higher-stress environments, even with transformations of species composition and dominance. It also demonstrates that the mechanisms regulating diversity, including invader impacts, can vary within the same plant community depending on environmental context. Because limits on native plant recruitment are environmentally contingent, management strategies that seek to increase diversity, including invader eradication, must account for site-level variations in the balance between biotic and abiotic constraints.     

An invader differentially affects leaf physiology of two natives across a gradient in diversity

Little is known about how exotics influence the ecophysiology of co-occurring native plants or how invader impact on plant physiology may be mediated by community diversity or resource levels. We measured the effect of the widespread invasive forb spotted knapweed (Centaurea maculosa) on leaf traits (leaf dry matter content, specific leaf area, leaf nitrogen percentage, leaf C:N ratios, and delta13C as a proxy for water use efficiency) of two co-occurring native perennial grassland species, Monarda fistulosa (bee balm) and Koeleria macrantha (Junegrass). The impact of spotted knapweed was assessed across plots that varied in functional diversity and that either experienced ambient rainfall or received supplemental water. Impact was determined by comparing leaf traits between identical knapweed-invaded and noninvaded assemblages. Virtually all M. fistulosa leaf traits were affected by spotted knapweed. Knapweed impact, however, did not scale with its abundance; the impact of knapweed on M. fistulosa was similar across heavily invaded low-diversity assemblages and lightly invaded high-diversity assemblages. In uninvaded assemblages, M. fistulosa delta13C, leaf nitrogen, and C:N ratios were unaffected by native functional group richness, whereas leaf dry matter content significantly increased and specific leaf area significantly decreased across the diversity gradient. The effects of spotted knapweed on K. macrantha were weak; instead native functional group richness strongly affected K. macrantha leaf C:N ratio, delta13C, and specific leaf area, but not leaf dry matter content. Leaf traits for both species changed in response to spotted knapweed or functional richness, and in a manner that may promote slower biomass accumulation and efficient conservation of resources. Taken together, our results show that an invader can alter native plant physiology, but that these effects are not a simple function of how many invaders exist in the community.     

盐碱草原土壤特性及生物量空间异质性研究

土壤盐分是土壤特性中最活跃和复杂的一部分。受土壤性质、气象、地形、地下水文及长期地球化学过程等自然条件和耕作、灌溉等人类活动因子的影响,土壤水分、盐分的变异状态在一定程度上反映了土壤各层的盐渍化程度和状态。而土壤盐分与水分、土壤物理性质等有十分密切的联系,因此,运用地统计学研究采用振动深松集成技术改良盐碱化草原前后土壤水分、盐分等特性的空间变异规律,以地理信息系统ArcGIS为平台,建立土壤水分、盐分等图形数据和属性数据相结合的信息数据库,建立其空间分异模型和空间分布图,直观的表现各特性的空间变异规律。结果表明,采用振动深松集成技术后土壤盐分含量在空间上有递减,平均ρ（土壤盐）由5.60g.L-1下降到3.73 g.L-1;土壤容重、土壤硬度分别降低13%和24%,利于牧草根系生长,生物量变异结果显示,改良前后的草层高度和产量空间分布逐渐达到了均一化,牧草产量达到了2738.1 kg.hm-2,实现了植被全覆盖,盐碱化草原得到了恢复。     

Variation in resource availability changes the impact of invasive thistles on native bunchgrasses.

The threat posed by invasive nonnative plants to native plant populations is one of the largest challenges facing both conservation biology and restoration ecology. California has been highly impacted by invaders, although many relict stands of native plants are found on shallow, rocky soils with limited resources. The abiotic conditions of these sites may strongly influence the performance of an invasive plant and its effect on resident native species. In addition, the maturity of native plants in these sites may modulate an invader's impact; larger, well-established plants may be better able to resist invaders. In this study we examined how the impact of an invasive thistle (Centaurea solstitialis) on a native perennial bunchgrass (Nassella pulchra) changed in response to variation in soil depth, soil water availability, and bunchgrass maturity. We measured plant performance in terms of survival, growth, reproduction, and predawn water potential. We found that soil depth, water availability, and bunchgrass maturity acted in concert to influence the impact of the invasive thistle on the native bunchgrass. Both species performed better in deep soils, especially during dry years. The combination of shallow soil and low water availability reduced C. solstitialis performance and ameliorated its negative effect on N. pulchra growth and reproduction. Higher water availability resulted in a stronger negative effect of C. solstitialis on N. pulchra in both shallow and deep soils. However, as N. pulchra matured and increased in size, we saw a steady decline in C. solstitialis growth and reproductive output. Higher water availability increased the performance of C. solstitialis in shallow soils. C. solstitialis may thus have a stronger impact on N. pulchra and be more able to invade relict stands of N. pulchra in shallow soils during high-rainfall years. However, established stands of N. pulchra appear to be more resistant to invasion by C. solstitialis as N. pulchra plants grow older and larger.     

Native fish diversity alters the effects of an invasive species on food webs

Aquatic communities have been altered by invasive species, with impacts on native biodiversity and ecosystem function. At the same time, native biodiversity may mitigate the effects of an invader. Common carp (Cyprinus carpio) is a ubiquitous, invasive fish species that strongly influences community and ecosystem processes. We used common carp to test whether the potential effects of an invasive species are altered across a range of species diversity in native communities. In mesocosms, treatments of zero, one, three, and six native fish species were used to represent the nested subset patterns observed in fish communities of lakes in Illinois, USA. The effect of the invader was tested across fish richness treatments by adding common carp to the native community and substituting native biomass with common carp. Native species and intraspecific effects reduced invader growth. The invader reduced native fish growth; however, the negative effect was minimized with increasing native richness. The zooplankton grazer community was modified by a top-down effect from the invader that increased the amount of phytoplankton. Neither the invader nor richness treatments influenced total phosphorus or community metabolism. Overall, the invader reduced resources for native species; and the effect scaled with how the invader was incorporated into the community. Higher native diversity mitigated the impact of the invader, confirming the need to consider biodiversity when predicting the impacts of invasive species.     

Trajectories of change in sagebrush steppe vegetation communities in relation to multiple wildfires

Repeated perturbations, both biotic and abiotic, can lead to fundamental changes in the nature of ecosystems, including changes in state. Sagebrush steppe communities provide important habitat for wildlife and grazing for livestock. Fire is an integral part of these systems, but there is concern that increased ignition frequencies and invasive species are fundamentally altering them. Despite these issues, the majority of studies of fire effects in systems dominated by Artemisia tridentata wyomingensis have focused on the effects of single burns. The Arid Lands Ecology Reserve (ALE), in south-central Washington (U.S.A.), was one of the largest contiguous areas of sagebrush steppe habitat in the state until large wildfires burned the majority of it in 2000 and 2007. We analyzed data from permanent vegetation transects established in 1996 and resampled in 2002 and 2009. Our objective was to describe how the fires, and subsequent postfire restoration efforts, affected communities' successional pathways. Plant communities differed in response to repeated fire and restoration; these differences could largely be ascribed to the functional traits of the dominant species. Low-elevation communities, previously dominated by obligate seeders, moved furthest from their initial composition and were dominated by weedy, early-successional species in 2009. Higher-elevation sites with resprouting shrubs, native bunchgrasses, and few invasive species were generally more resilient to the effects of repeated disturbances. Shrub cover has been almost entirely removed from ALE, although there was some recovery where communities were dominated by resprouters. Bromus tectorum dominance was reduced by herbicide application in areas where it was previously abundant, but it increased significantly in untreated areas. Several resprouting species, notably Phlox longifolia and Poa secunda, expanded remarkably following competitive release from shrub canopies and/or abundant B. tectorum. Our results suggest that community dynamics can be understood through a state and transition model with two axes (shrub/grass and native/invasive abundance), although such models also need to account for differences in plant functional traits and disturbance regimes. We use our results to develop a conceptual model that will be validated with further research.     

Soil properties and root biomass responses to prescribed burning in young Corsican pine (Pinus nigra Arn.) stands

Fire is an important tool in the management of forest ecosystems. Although both prescribed and wildland fires are common in Turkey, few studies have addressed the influence of such disturbances on soil properties and root biomass dynamics. In this study, soil properties and root biomass responses to prescribed fire were investigated in 25-year-old corsican pine (Pinus nigra Arn.) stands in Kastamonu, Turkey. The stands were established by planting and were subjected to prescribed burning in July 2003. Soil respiration rates were determined every two months using soda-lime method over a two-year period. Fine (0-2 mm diameter) and small root (2-5 mm diameter) biomass were sampled approximately bimonthly using sequential coring method. Mean daily soil respiration ranged from 0.65 to 2.19 g Cm(-2) d(-1) among all sites. Soil respiration rates were significantly higher in burned sites than in controls. Soil respiration rates were correlated significantly with soil moisture and soil temperature. Fine root biomass was significantly lower in burned sites than in control sites. Mean fine root biomass values were 4940 kg ha(-1) for burned and 5450 kg ha(-1) for control sites. Soil pH was significantly higher in burned sites than in control sites in 15-35 cm soil depth. Soil organic matter content did not differ significantly between control and burned sites. Our results indicate that, depending on site conditions, fire could be used successfully as a tool in the management of forest stands in the study area.     

The effects of Pinus tabuliformis on soil detachment under different influencing factors in the Loess Plateau of China

In situ experiments were undertaken to examine the impacts of Pinus tabuliformis on soil detachment under different influencing factors. Experimental sites with slopes containing vegetation cover, plant roots and bare ground were investigated for slope gradients of 8.7%, 17.6%, 26.8%, 36.4% and 46.6% and flow discharges of 0.25, 0.5, 1.0, 1.5 and 2.0?m³?s^?1. Results from our study provide details on the relationship between soil detachment and both slope gradient and flow discharge. Soil detachment rates increased with an increase in slope and discharge, and discharge was identified to have a stronger influence on soil detachment than slope. In contrast, the combination of litter cover and roots played an important role in reducing soil detachment; detachment rate decreased by about 55% relative to bare slopes; and plant roots had a greater impact on detachment reduction. The presence of vegetation cover can decrease rill erodibility by 83% and increase critical shear stress by 224% compared with bare slopes. These results provide valuable information on the importance of woodland in soil detachment control, and may help to improve vegetation construction in seriously eroded regions of the Loess Plateau.     

Constitutive and induced defenses to herbivory in above- and belowground plant tissues

A recent surge in attention devoted to the ecology of soil biota has prompted interest in quantifying similarities and differences between interactions occurring in above- and belowground communities. Furthermore, linkages that interconnect the dynamics of these two spatially distinct ecosystems are increasingly documented. We use a similar approach in the context of understanding plant defenses to herbivory, including how they are allocated between leaves and roots (constitutive defenses), and potential cross-system linkages (induced defenses). To explore these issues we utilized three different empirical approaches. First, we manipulated foliar and root herbivory on tobacco (Nicotiana tabacum) and measured changes in the secondary chemistry of above- and belowground tissues. Second, we reviewed published studies that compared levels of secondary chemistry between leaves and roots to determine how plants distribute putative defense chemicals across the above- and belowground systems. Last, we used meta-analysis to quantify the impact of induced responses across plant tissue types. In the tobacco system, leaf-chewing insects strongly induced higher levels of secondary metabolites in leaves but had no impact on root chemistry. Nematode root herbivores, however, elicited changes in both leaves and roots. Virtually all secondary chemicals measured were elevated in nematode-induced galls, whereas the impact of root herbivory on foliar chemistry was highly variable and depended on where chemicals were produced within the plant. Importantly, nematodes interfered with aboveground metabolites that have biosynthetic sites located in roots (e.g., nicotine) but had the opposite effect (i.e., nematodes elevated foliar expression) on chemicals produced in shoots (e.g., phenolics and terpenoids). Results from our literature review suggest that, overall, constitutive defense levels are extremely similar when comparing leaves with roots, although certain chemical classes (e.g., alkaloids, glucosinolates) are differentially allocated between above- and belowground parts. Based on a meta-analysis of induced defense studies we conclude that: (1) foliar induction generates strong responses in leaves, but much weaker responses in roots, and (2) root induction elicits responses of equal magnitude in both leaves and roots. We discuss the importance of this asymmetry and the paradox of cross-system induction in relation to optimal defense theory and interactions between above- and belowground herbivory.