Nichole N. Barger

Changes in Plant Functional Groups, Litter Quality, and Soil Carbon and Nitrogen Mineralization With Sheep Grazing in an Inner Mongolian Grassland

Abstract This study reports on changes in plant functional group composition, litter quality, and soil C and N mineralization dynamics from a 9-year sheep grazing study in Inner Mongolia. Addressed are these questions: 1) How does increasing grazing intensity affect plant community composition? 2) How does increasing grazing intensity alter soil C and N mineralization dynamics? 3) Do changes in soil C and N mineralization dynamics relate to changes in plant community composition via inputs of the quality or quantity of litter? Grazing plots were set up near the Inner Mongolia Grassland Ecosystem Research Station (IMGERS) with 5 grazing intensities: 1.3, 2.7, 4.0, 5.3, and 6.7 sheep ha−1·yr−1. Plant cover was lower with increasing grazing intensity, which was primarily due to a dramatic decline in grasses, Carex duriuscula, and Artemisia frigida. Changes in litter mass and percentage organic C resulted in lower total C in the litter layer at 4.0 and 5.3 sheep ha−1·yr−1 compared with 2.7 sheep ha−1·yr−1. Total litter N was lower at 5.3 sheep ha−1·yr−1 compared with 2.7 sheep ha−1·yr−1. Litter C:N ratios, an index of litter quality, were significantly lower at 4.0 sheep ha−1·yr−1 relative to 1.3 and 5.3 sheep ha−1·yr−1. Cumulative C mineralized after 16 days decreased with increasing grazing intensity. In contrast, net N mineralization (NH+4 + NO−3) after a 12-day incubation increased with increasing grazing intensity. Changes in C and N mineralization resulted in a narrowing of CO2-C:net Nmin ratios with increasing grazing intensity. Grazing explained 31% of the variability in the ratio of CO2-C:net Nmin. The ratio of CO2-C:net Nmin was positively correlated with litter mass. Furthermore, there was a positive correlation between litter mass and A. frigida cover. Results suggest that as grazing intensity increases, microbes become more C limited resulting in decreased microbial growth and demand for N.

Soil carbon storage responses to expanding pinyon-juniper populations in southern Utah

Over the past several decades, the expansion and thickening of woodlands in the western United States has caused a range of ecological changes. Woody expansion often leads to increases in soil organic matter (SOM) pools with implications for both biogeochemical cycling and ecological responses to management strategies aimed at restoration of rangeland ecosystems. Here we directly measure C and N stocks and use simple non-steady-state models to quantify the dynamics of soil C accumulation under and around trees of varied ages in southern Utah woodlands. In the two pinyon-juniper forests of Grand Staircase Escalante National Monument studied here, we found approximately 3 kg C/m2 and approximately 0.12 kg N/m2 larger C and N stocks in soils under pinyon canopies compared to interspace sites. These apparent increases in soil C and N stocks under woody plant species were dominated by elevated SOM in the surface 10 cm of soil, particularly within non-mineral-associated organic fractions. The most significant accumulation of C was in the >850 microm fraction, which had an estimated C residence time of <20 yr. Rates of carbon accumulation following pinyon-juniper expansion appear to be dominated by changes in this fast-cycling surface soil fraction. In contrast, we found that after separating >850 microm organic matter from the remaining light fraction (LF), C had residence times of approximately 400 yr and mineral-associated (MA) soil C had residence times of approximately 600 yr. As a result, we calculate that input rates to the LF and MA pools to be 10 +/- 1 and 0.68 +/- 0.15 g m(-2) yr(-1) (mean +/- SE), respectively. These findings suggest that one consequence of management activities aimed at the reduction of pinyon-juniper biomass may be a relatively rapid loss of soil C and N pools associated with the >850 microm fraction. The temporal dynamics of the <850 microm pools suggest that carbon and nitrogen continue to accumulate in these fractions, albeit at very slow rates, and suggest that multidecadal storage of C following tree recruitment is limited to relatively small, subsurface fractions of the total soil C pool.

Nutrient Limitation to Primary Productivity in a Secondary Savanna in Venezuela1

We examined nutrient limitation to primary productivity in a secondary savanna in the interior branch of the Coastal Range of Venezuela, which was converted from forest to savanna more than 100 years ago. We manipulated soil nutrients by adding nitrogen (+N), phosphorus and potassium (+PK), and nitrogen, phosphorus, and potassium (+NPK) to intact savanna. Eleven months after fertilization, we measured aboveground biomass and belowground biomass as live fine roots in the top 20 cm of soil, and species and functional group composition in response to nutrient additions. Aboveground biomass was highest in the NPK treatment ([mean g/m2]; control = 402, +N = 718, +PK = 490, +NPK = 949). Aboveground production, however, appeared to be limited primarily by N. Aboveground biomass increased 78 percent when N was added alone but did not significantly respond to PK additions when compared to controls. In contrast to aboveground biomass, belowground biomass increased with PK additions but showed no significant increase with N (depth 0-20 cm; [mean g/m2]; control = 685, +N = 443, +PK = 827, +NPK = 832). There was also a 36 percent increase in root length with PK additions when compared to controls. Whole savanna shootxoot ratios were similar for control and +PK (0.6), while those for + N or +NPK fertilization were significantly higher (1.7 and 1.2, respectively). Total biomass response (above + belowground) to nutrient additions showed a strong N and PK co-limitation ([mean g/m2]; control = 1073, +N = 1111, +PK = 1258, +NPK = 1713). Aboveground biomass of all monocots increased with N additions, whereas dicots showed no response to nutrient additions. Trachypogon spp. ( T plumosw + 7: vestitus) and Axonopw canescens, the two dominant grasses, made up more than 89 percent of the total aboveground biomass in these sites. Tracbpogon spp. responded to NPK, whereas A. canescens, sedges, and the remaining monocots only responded to N. Even though nutrient additions resulted in higher aboveground biomass in N and NPK fertilized plots, this had little effect on plant community composition. With the exception of sedges, which responded positively to N additions and increased from 4 to 8 percent of the plant community, no changes were observed in plant community composition after 11 months.

Influence of Livestock Grazing and Climate on Pinyon Pine (Pinus edulis) Dynamics

Abstract Over the last century there has been marked expansion and infilling of pinyon (Pinus spp.)–juniper (Juniperus spp.) woodlands into grassland and shrubland ecosystems across the western United States. Although range expansions in pinyon-juniper populations have been documented with changing climate throughout the Holocene, over the last century, local scale impacts such as livestock grazing, changes in fire regimes, and increasing atmospheric CO2 concentrations are thought to be more recent drivers of pinyon-juniper woodland distribution. Our objective was to examine the role of historical livestock grazing relative to past climate in regulating pinyon (Pinus edulis Engelm.) recruitment and growth over the last century in a persistent pinyon-juniper woodland. We compared pinyon dynamics on a remote mesa site, minimally grazed by domestic livestock, to a nearby historically grazed site in southeastern Utah. The presence of a significant number of old trees (> 200 yr) at both sites suggests that these populations were well established prior to the 20th century. No differences in pinyon density or basal area were observed between grazed and ungrazed sites. Stand age structure data showed that pinyon recruitment across these sites was highly synchronous, with a large recruitment event occurring during the early 1900s; 17% of the pinyons on both sites dated to the 1920s. Climatic conditions during this decade were consistently cool and wet—conditions known to support enhanced recruitment and growth in pinyon pines. Pinyon growth was also strongly synchronous across sites (r  =  0.96). Pinyon growth was significantly correlated with winter/spring precipitation and negatively correlated with June temperature. Taken together, our results suggest that past climate may be more important in structuring pinyon populations than historical land use in these persistent pinyon-juniper woodlands. Given future climate projections of increasing temperature and more extended drought periods, regeneration of pinyon populations following the recent regional-scale dieback may be slow. Moreover, prolonged drought combined with potentially slow regeneration times for pinyon under future climate scenarios could result in substantial declines in pinyon populations across the region, a result that land managers should consider when planning for future restoration treatments in persistent pinyon-juniper woodlands.

Are Namibian “Fairy Circles” the Consequence of Self-Organizing Spatial Vegetation Patterning?

Causes of over-dispersed barren “fairy circles” that are often surrounded by ca. 0.5 m tall peripheral grasses in a matrix of shorter (ca. 0.2 m tall) grasses in Namibian grasslands remain mysterious. It was hypothesized that the fairy circles are the consequence of self-organizing spatial vegetation patterning arising from resource competition and facilitation. We examined the edaphic properties of fairy circles and variation in fairy circle size, density and landscape occupancy (% land surface) with edaphic properties and water availability at a local scale (<50 km) and with climate and vegetation characteristics at a regional scale. Soil moisture in the barren fairy circles declines from the center towards the periphery and is inversely correlated with soil organic carbon, possibly indicating that the peripheral grass roots access soil moisture that persists into the dry season within fairy circles. Fairy circle landscape occupancy is negatively correlated with precipitation and soil [N], consistent with fairy circles being the product of resource-competition. Regional fairy circle presence/absence is highly predictable using an empirical model that includes narrow ranges of vegetation biomass, precipitation and temperature seasonality as predictor variables, indicating that fairy circles are likely a climate-dependent emergent phenomenon. This dependence of fairy circle occurrence on climate explains why fairy circles in some locations may appear and disappear over time. Fairy circles are only over-dispersed at high landscape occupancies, indicating that inter-circle competition may determine their spacing. We conclude that fairy circles are likely to be an emergent arid-grassland phenomenon that forms as a consequence of peripheral grass resource-competition and that the consequent barren circle may provide a resource-reservoir essential for the survival of the larger peripheral grasses and provides a habitat for fossicking fauna.

Patterns and Controls on Nitrogen Cycling of Biological Soil Crusts

In low-nutrient environments with few vascular plant symbiotic N fixers, biocrusts play an important role in ecosystem N cycling. A large number of studies across a wide range of biomes clearly confirm that not only the presence of biocrusts but biocrust community composition strongly influences N-fixation activity, with N fixation increasing with level of development (cyanobacterial-lichen biocrusts > dark cyanobacterial biocrust (e.g., Nostoc spp. and Collema spp.) > light Microcoleus-dominated biocrust). Nitrogen fixation by biocrusts results in N release to the soil in a variety of N forms (inorganic and organic N), thus elevating soil inorganic N pools in the top few millimeters of soil. The influence of N release on the bulk soil at greater soil depths is less clear, with biocrusts either elevating or having no influence on bulk soil inorganic N pools. The fate of N fixed and released by biocrusts, and whether this N is retained in the ecosystem in either soils or plants, determines ecosystem N balance over longer time scales, and results on the influence of biocrusts are mixed. Whereas we have multiple studies that examine a single compartment of N budgets, we lack studies that simultaneously address N inputs, losses, and soil and plant pools, thus precluding the construction of definitive N balances. One of the most consistent impact biocrusts have on ecosystem N is reducing N loss via wind and water erosion, with such losses consistently decreasing with increasing biocrust development.

Tree regeneration following drought‐ and insect‐induced mortality in piñon–juniper woodlands

Widespread piñon (Pinus edulis) mortality occurred across the southwestern USA during 2002-2003 in response to drought and bark beetle infestations. Given the recent mortality and changes in regional climate over the past several decades, there is a keen interest in post-mortality regeneration dynamics in piñon-juniper woodlands. Here, we examined piñon and Utah juniper (Juniperus osteosperma) recruitment at 30 sites across southwestern Colorado, USA that spanned a gradient of adult piñon mortality levels (10-100%) to understand current regeneration dynamics. Piñon and juniper recruitment was greater at sites with more tree and shrub cover. Piñon recruitment was more strongly facilitated than juniper recruitment by trees and shrubs. New (post-mortality) piñon recruitment was negatively affected by recent mortality. However, mortality had no effect on piñon advanced regeneration (juveniles established pre-mortality) and did not shift juvenile piñon dominance. Our results highlight the importance of shrubs and juniper trees for the facilitation of piñon establishment and survival. Regardless of adult piñon mortality levels, areas with low tree and shrub cover may become increasingly juniper dominated as a result of the few suitable microsites for piñon establishment and survival. In areas with high piñon mortality and high tree and shrub cover, our results suggest that piñon is regenerating via advanced regeneration.

Denitrification from nitrogen-fixing biologically crusted soils in a cool desert environment, southeast Utah, USA

IntroductionNitrogen fixation by microorganisms within biological soil crust (“biocrust”) communities provides an important pathway for N inputs in cool desert environments where soil nutrients are low and symbiotic N-fixing plants may be rare. Estimates of N fixation in biocrusts often greatly exceed that of N accretion rates leading to uncertainty regarding N loss pathways.MethodsIn this study we examined nitrogen fixation and denitrification rates in biocrust communities that differed in N fixation potential (low N fixation = light cyanobacterial biocrust, high N fixation = dark cyanolichen crust) at four temperature levels (10, 20, 30, 40°C) and four simulated rainfall levels (0.05, 0.2, 0.6, 1 cm rain events) under controlled laboratory conditions.ResultsAcetylene reduction rates (AR, an index of N fixation activity) were over six-fold higher in dark crusts relative to light crusts. Dark biocrusts also exhibited eight-fold higher denitrification rates. There was no consistent effect of temperature on denitrification rates, but there was an interactive effect of water addition and crust type. In light crusts, denitrification rates increased with increasing water addition, whereas the highest denitrification rates in dark crusts were observed at the lowest level of water addition.ConclusionsThese results suggest that there are no clear and consistent environmental controls on short-term denitrification rates in these biologically crusted soils. Taken together, estimates of denitrification from light and dark biocrusts constituted 3 and 4% of N fixation rates, respectively suggesting that losses as denitrification are not significant relative to N inputs via fixation. This estimate is based on a previously published conversion ratio of ethylene produced to N fixed that is low (0.295), resulting in high estimates of N fixation. If future N fixation studies in biologically crusted soils show that these ratios are closer to the theoretical 3:1 ratio, denitrification may constitute a more significant loss pathway relative to N fixed.

Woodland recovery following drought-induced tree mortality across an environmental stress gradient

Recent droughts and increasing temperatures have resulted in extensive tree mortality across the globe. Understanding the environmental controls on tree regeneration following these drought events will allow for better predictions of how these ecosystems may shift under a warmer, drier climate. Within the widely distributed piñon-juniper woodlands of the southwestern USA, a multiyear drought in 2002-2004 resulted in extensive adult piñon mortality and shifted adult woodland composition to a juniper-dominated, more savannah-type ecosystem. Here, we used pre- (1998-2001) and 10-year post- (2014) drought stand structure data of individually mapped trees at 42 sites to assess the effects of this drought on tree regeneration across a gradient of environmental stress. We found declines in piñon juvenile densities since the multiyear drought due to limited new recruitment and high (>50%) juvenile mortality. This is in contrast to juniper juvenile densities, which increased over this time period. Across the landscape, piñon recruitment was positively associated with live adult piñon densities and soil available water capacity, likely due to their respective effects on seed and water availability. Juvenile piñon survival was strongly facilitated by certain types of nurse trees and shrubs. These nurse plants also moderated the effects of environmental stress on piñon survival: Survival of interspace piñon juveniles was positively associated with soil available water capacity, whereas survival of nursed piñon juveniles was negatively associated with perennial grass cover. Thus, nurse plants had a greater facilitative effect on survival at sites with higher soil available water capacity and perennial grass cover. Notably, mean annual climatic water deficit and elevation were not associated with piñon recruitment or survival across the landscape. Our findings reveal a clear shift in successional trajectories toward a more juniper-dominated woodland and highlight the importance of incorporating biotic interactions and soil properties into species distribution modeling approaches.

Microbial Nursery Production of High-Quality Biological Soil Crust Biomass for Restoration of Degraded Dryland Soils

ABSTRACT Biological soil crusts (biocrusts) are slow-growing, phototroph-based microbial assemblages that develop on the topsoils of drylands. Biocrusts help maintain soil fertility and reduce erosion. Because their loss through human activities has negative ecological and environmental health consequences, biocrust restoration is of interest. Active soil inoculation with biocrust microorganisms can be an important tool in this endeavor. We present a culture-independent, two-step process to grow multispecies biocrusts in open greenhouse nursery facilities, based on the inoculation of local soils with local biocrust remnants and incubation under seminatural conditions that maintain the essence of the habitat but lessen its harshness. In each of four U.S. Southwest sites, we tested and deployed combinations of factors that maximized growth (gauged as chlorophyll a content) while minimizing microbial community shifts (assessed by 16S rRNA sequencing and bioinformatics), particularly for crust-forming cyanobacteria. Generally, doubling the frequency of natural wetting events, a 60% reduction in sunlight, and inoculation by slurry were optimal. Nutrient addition effects were site specific. In 4 months, our approach yielded crusts of high inoculum quality reared on local soil exposed to locally matched climates, acclimated to desiccation, and containing communities minimally shifted in composition from local ones. Our inoculum contained abundant crust-forming cyanobacteria and no significant numbers of allochthonous phototrophs, and it was sufficient to treat ca. 6,000 m2 of degraded dryland soils at 1 to 5% of the typical crust biomass concentration, having started from a natural crust remnant as small as 6 to 30 cm2. IMPORTANCE Soil surface crusts can protect dryland soils from erosion, but they are often negatively impacted by human activities. Their degradation causes a loss of fertility, increased production of fugitive dust and intensity of dust storms with associated traffic problems, and provokes general public health hazards. Our results constitute an advance in the quest to actively restore biological soil covers by providing a means to obtain high-quality inoculum within a reasonable time (a few months), thereby allowing land managers to recover essential, but damaged, ecosystem services in a sustainable, self-perpetuating way as provided by biocrust communities.