Humans are creating significant global environmental change, including shifts in climate, increased nitrogen (N) deposition, and the facilitation of species invasions. A multi-factorial field experiment is being performed in an arid grassland within the Sevilleta National Wildlife Refuge (NWR) to simulate increased nighttime temperature, higher N deposition, and heightened El Niño frequency (which increases winter precipitation by an average of 50%). The purpose of the experiment is to better understand the potential effects of environmental drivers on grassland community composition, aboveground net primary production and soil respiration. The focus is on the response of two dominant grasses (Bouteloua gracilis and B eriopoda), in an ecotone near their range margins and thus these species may be particularly susceptible to global environmental change.
It is hypothesized that warmer summer temperatures and increased evaporation will favor growth of black grama (Bouteloua eriopoda), a desert grass, but that increased winter precipitation and/or available nitrogen will favor the growth of blue grama (Bouteloua gracilis), a shortgrass prairie species. Treatment effects on limiting resources (soil moisture, nitrogen availability, species abundance, and net primary production (NPP) are all being measured to determine the interactive effects of key global change drivers on arid grassland plant community dynamics and ecosystem processes. This dataset shows values of soil moisture, soil temperature, and the CO2 flux of the amount of CO2 that has moved from soil to air.
On 4 August 2009 lightning ignited a ~3300 ha wildfire that burned through the experiment and its surroundings. Because desert grassland fires are patchy, not all of the replicate plots burned in the wildfire. Therefore, seven days after the wildfire was extinguished, the Sevilleta NWR Fire Crew thoroughly burned the remaining plots allowing us to assess experimentally the effects of interactions among multiple global change presses and a pulse disturbance on post-fire grassland dynamics.
Our experimental design consists of three fully crossed factors (warming, increased winter precipitation, and N addition) in a completely randomized design, for a total of eight treatment combinations, with five replicates of each treatment combination, for a total of 40 plots. Each plot is 3 x 3.5 m. All plots contain B. eriopoda, B. gracilis and G. sarothrae. Our nighttime warming treatment is imposed using lightweight aluminum fabric shelters (mounted on rollers similar to a window shade) that are drawn across the warming plots each night to trap outgoing longwave radiation. The dataloggers controlling shelter movements are programmed to retract the shelters on nights when wind speeds exceed a threshold value (to prevent damage to shelters) and when rain is detected by a rain gauge or snow is detected by a leaf wetness sensor (to prevent an unintended rainout effect).
Each winter we impose an El Nino-like rainfall regime (50% increase over long-term average for non-El Nino years) using an irrigation system and RO water. El Nino rains are added in 6 experimental storm events that mimic actual El Nino winter-storm event size and frequency. During El Nino years we use ambient rainfall and do not impose experimental rainfall events. For N deposition, we add 2.0 g m-2 y-1 of N in the form of NH4NO3 because NH4 and NO3 contribute approximately equally to N deposition at SNWR (57% NH4 and 43% NO3; Bez et al., 2007). The NH4NO3 is dissolved in 12 liters of deionized water, equivalent to a 1 mm rainfall event, and applied with a backpack sprayer prior to the summer monsoon. Control plots receive the same amount of deionized water.
Soil temperature is measured with Campbell Scientific CS107 temperature probes buried at 2 and 8 cm In the soil. Soil volume water content, measured with Campbell Scientific CS616 TDR probes is an integrated measure of soil water availability from 0-15 cm deep in the soil. Soil CO2 is measured with Vaisala GM222 solid state CO2 sensors. For each plot, soil sensors are placed under the canopy of B. eriopoda at three depths: 2, 8, and 16 cm. Measurements are recorded every 15 minutes.
CO2 fluxes are calculated using the CO2, temperature, and moisture data, along with ancillary variables following the methods of Vargas et al (2012) Global Change Biology
Values of CO2 concentration are corrected for temperature and pressure using the ideal gas law according to the manufacturer (Vaisala). We calculate soil respiration using the flux-gradient method (Vargas et al. 2010) based on Fick’s law of diffusion where the diffusivity of CO2 is corrected for temperature and pressure (Jones 1992) and calculated as a function of soil moisture, porosity and texture (Moldrup et al. 1999).
Instrument Name: Solid State Soil CO2 sensor
Model Number: GM222
Instrument Name: Temperature Probe
Manufacturer: Campbell Scientific
Model Number: CS107
Instrument Name: Water Content Reflectometer Probe
Manufacturer: Campbell Scientific
Model Number: CS616
Net primary production is a fundamental ecological variable that quantifies rates of carbon consumption and fixation. Estimates of NPP are important in understanding energy flow at a community level as well as spatial and temporal responses to a range of ecological processes. While measures of both below- and above-ground biomass are important in estimating total NPP, this study focuses on above-ground net primary production (ANPP). Above-ground net primary production is the change in plant biomass, including loss to death and decomposition, over a given period of time. Volumetric measurements are made using vegetation data from permanent plots collected in SEV297, "Extreme Drought in Grassland Ecosystems (EDGE) Net Primary Production Quadrat Data" and regressions correlating biomass and volume constructed using seasonal harvest weights from SEV157, "Net Primary Productivity (NPP) Weight Data."
Derivation of Biomass and Net primary Production:
Data from SEV297 and SEV157 are used to calculate the seasonal and annual production (i.e., biomass) of each species in each quadrat for a given year. Allometric equations derived from harvested samples of each species for each season are applied to the measured cover, height, and count of each species in each quadrat. This provides seasonal biomass for winter, spring, and fall.
Seasonal net primary production (NPP) is derived by subtracting the previous season's biomass from the biomass for the current season. For example, spring NPP is calculated by subtracting the winter weight from the spring weight for each species in a given quadrat. Negative differences are considered to be 0. Likewise, fall production is computed by subtracting spring biomass from fall biomass. Annual biomass is taken as the sum of spring and fall NPP.
The bounding box coordinates for the corners of the polygon which encompasses the full EDGE black site are:NW: -106.729227 34.337913 Decimal DegreesNE: -106.728434 34.337937 Decimal DegreesSW: -106.729144 34.337298 Decimal DegreesSE: -106.728392 34.337310 Decimal DegreesEDGE blue:NW: -106.622610 34.342141 Decimal DegreesNE: -106.621689 34.342079 Decimal DegreesSW: -106.623365 34.341518 Decimal DegreesSE: -106.622711 34.341015 Decimal Degrees
EDGE is located at six grassland sites that encompass a range of ecosystems in the Central US - from desert grasslands to short-, mixed-, and tallgrass prairie. We envision EDGE as a research platform that will not only advance our understanding of patterns and mechanisms of ecosystem sensitivity to climate change, but also will benefit the broader scientific community. Identical infrastructure for manipulating growing season precipitation will be deployed at all sites. Within the relatively large treatment plots (36 m2), we will measure with comparable methods, a broad spectrum of ecological responses particularly related to the interaction between carbon fluxes (NPP, soil respiration) and species response traits, as well as environmental parameters that are critical for the integrated experiment-modeling framework, as well as for site-based analyses. By designing EDGE as a research platform open to the broader scientific community, with subplots in all replicates (n = 180 plots) set-aside for additional studies, and by making data available to the broader ecological community EDGE will have value beyond what we envision here.
The six sites were selected to capture the key environmental and ecological gradients of Central US grasslands and represent the major grassland ecosystem types (desert, shortgrass, mixedgrass, and tallgrass) of the region. Site selection criteria included: site characteristics (mean annual precipitation and temperature, dominant vegetation), access and site security, permission to build experimental infrastructure, participation in an existing or future network (e.g., LTER, NEON), and available site support and supporting data (e.g., LTER, USFWS or ARS).
Experimental Treatments and Plots
Our approach will be to impose a significant reduction in growing season precipitation (-66 % of ambient) over a 4-yr period. This is the equivalent of a ca. 50% reduction in annual precipitation because at all sites about 60-75% of annual precipitation falls in the growing season. We will impose this long-term drought either by reducing the size of each rainfall event (event size reduction, E) or by reducing the number of events (delayed rainfall treatment, D).
The control (C) treatment is included for comparison. At each site, the ambient (C) rainfall pattern will be reduced in two ways to impose a severe drought over a 4-yr period.
For the event size reduction treatment (E), each rainfall event will be passively reduced by a fixed proportion. Note that rain event number and the average number of days between events does not differ from ambient treatment.
For the reduced event number (D) treatment, shelters roofs will be removable to permit periods of complete rain exclusion alternating with periods of ambient rainfall inputs. Here, a + 10 mm rule is used to determine when roofs are on or off. When the cumulative precipitation amount in this D treatment falls 10 mm below the E treatment, the roofs are removed until the cumulative precipitation total is 10 mm greater than the E treatment. In this way, total precipitation amounts will be similar at the end of the growing season, but event number will be reduced and the average number of days between events increased, with no change in event size compared to the C treatment.
At each site, we will establish replicate 6 x 6 m experimental plots (n = 10 per treatment, including the control treatment) in a relatively homogeneous area (similar soils, vegetation, etc.) that is representative of the overall site. Plots will be arrayed such that each treatment will be co-located in a single block (n=10 blocks per site), with each block located at least 5 m apart.
The blocking will help control for environmental gradients if present. For each site, all plots within a block (including the control) will be located at least 2 m apart and trenched to 1-1.5 m and surrounded by a 6 mil plastic barrier to hydrologically isolate them from the adjacent soil, and each plot will be covered by the rainfall manipulation infrastructure. The 6 x 6 m plot size includes a 0.5 m external buffer to allow access to the plots and minimize edge effects associated with the infrastructure. The resulting 5 x 5 m area will be divided into 4 2.5 x 2.5 m subplots. One subplot will be designated for plant species composition sampling, two will for destructive sampling (ANPP, belowground productivity, soil sampling, etc.), and the fourth set aside for opportunistic studies.
Rainfall Manipulation Infrastructure
We will passively alter rainfall reaching the plots by using a version of a rainfall reduction shelter (Fig. 6) designed by Yahdjian and Sala (2002). Versions of these shelters (ranging from ~2 to 100 m2 ) are being used by the co-PIs at the Sevilleta, Konza Prairie and Shortgrass Steppe LTERs, as well as by many other ecologists, and thus, they are proven technology. The most significant environmental artifacts of these shelters are a 5- 10% reduction in light due to the acrylic Vshaped shingles and a ~ 20 cm edge effect (Yahdjian and Sala 2002). Shelters will consist of a steel frame that supports a roof. To cover the 36 m2 plots, the shelters will be constructed as modular 3 x 3 m units, with four units per plot. The roof of each modular unit will be slanted at 15° toward the edge of the plot, creating a 6 m long peak along the mid-line of the plot, with two lower 6 m long edges with gutters to move rainwater away from the plots. The peaked roof will facilitate run-off of rainfall and access to the plot, and the lower edge will be oriented to the prevailing wind direction to minimize blow-in. Average leaf canopy height varies among the desert/short-, midand tallgrass prairie sites (~0.2 to 0.6 m), and to maintain a consistent roof-to-canopy distance, peak height of the shelters will be 1.3, 1.55 and 1.8 m, with lower edges of the shelters at 0.5, 0.75 and 1.0 m, respectively, for the four grassland types. Construction of the shelters will begin in Yr 1 (after pretreatment measurements are taken) and treatments will be operational by the early spring of YR 2. For the ESR treatment, the roof will consist of clear acrylic (high light transmission, low yellowness index, UV transparent) v-shaped shingles arrayed at a density to passively reducing each rainfall event by ~66% (Fig. 6). For the REN treatment, the roof will consist of clear, corrugated polycarbonate (high light transmission, low yellowness index, UV transparent) to completely exclude rainfall. For both treatments, the roofs will be constructed to facilitate easy removal via a clamping system. The REN treatment roofs will then be manually deployed and removed at intermittent intervals (see Fig. 6 for more detail). Ambient plots will have a deer netting roof to achieve an average reduction in light similar to the rainfall reduction roofs.
Plant species composition, species traits, stem density, and light availability
In the subplot designated for species composition, we will establish a permanent 2 x 2 m sampling plots, which will be divided into four 1 x 1m quadrats in which canopy cover of each species will be visually estimated to the nearest 1%. For each site, these measures will be repeated at least twice during the growing season of each year to sample early and late season species. Maximum cover values of each species will be used to determine richness, diversity and dominance and changes in composition, species turnover, and species associations over time.
Collecting the Data:
Net primary production data is collected twice each year, spring and fall, for both sites. Spring measurements are taken in April or May when shrubs and spring annuals have reached peak biomass. Fall measurements are taken in either September or October when summer annuals have reached peak biomass but prior to killing frosts. Winter measurements are taken in February before the onset of spring growth.
Vegetation data is collected on a palm top computer. A 1-m2 PVC-frame is placed over the fiberglass stakes that mark the diagonal corners of each quadrat. When measuring cover it is important to stay centered over the vegetation in the quadrat to prevent errors caused by angle of view (parallax). Each PVC-frame is divided into 100 squares with nylon string. The dimensions of each square are 10cm x 10cm and represent 1 percent of the total area.
The cover (area) and height of each individual live (green) vegetative unit that falls within the one square meter quadrat is measured. A vegetative unit consists of an individual size class (as defined by a unique cover and height) of a particular species within a quadrat. Cover is quantified by counting the number of 10cm x 10cm squares filled by each vegetative unit.
Niners and plexidecs are additional tools that help accurately determine the cover a vegetative unit. A niner is a small, hand-held PVC frame that can be used to measure canopies. Like the larger PVC frame it is divided into 10cm x 10cm squares, each square representing 1% of the total cover. However, there are only nine squares within the frame, hence the name “niner.” A plexidec can help determine the cover of vegetative units with covers less than 1%. Plexidecs are clear plastic squares that are held above vegetation. Each plexidec represents a cover of 0.5% and has smaller dimensions etched onto the surface that correspond to 0.01%, 0.05%, 0.1%, and 0.25% cover.
It is extremely important that cover and height measurements remain consistent over time to ensure that regressions based on this data remain valid. Field crew members should calibrate with each other to ensure that observer bias does not influence data collection.
Grasses-To determine the cover of a grass clump, envision a perimeter around the central mass or densest portion of the plant, excluding individual long leaves, wispy ends, or more open upper regions of the plant. Live foliage is frequently mixed with dead foliage in grass clumps and this must be kept in mind during measurement as our goal is to measure only plant biomass for the current season. In general, recently dead foliage is yellow and dead foliage is gray. Within reason, try to include only yellow or green portions of the plant in cover measurement while excluding portions of the plant that are gray. This is particularly important for measurements made in the winter when there is little or no green foliage present. In winter, sometimes measurements will be based mainly on yellow foliage. Stoloniferous stems of grasses that are not rooted should be ignored. If a stem is rooted it should be recorded as a separate observation from the parent plant.
Forbs, shrubs and sub-shrubs (non-creosote)-The cover of forbs, shrubs and sub-shrubs is measured as the horizontal area of the plant. If the species is an annual it is acceptable to include the inflorescence in this measurement if it increases cover. If the species is a perennial, do not include the inflorescence as part of the cover measurement. Measure all foliage that was produced during the current season, including any recently dead (yellow) foliage. Avoid measuring gray foliage that died in a previous season.
Cacti-For cacti that consist of a series of pads or jointed stems (Opuntia phaecantha, Opuntia imbricata) measure the length and width of each pad to the nearest cm instead of cover and height. Cacti that occur as a dense ball/clump of stems (Opuntia leptocaulis) are measured using the same protocol as shrubs. Pincushion or hedgehog cacti (Escobaria vivipara, Schlerocactus intertextus, Echinocereus fendleri) that occur as single (or clustered) cylindrical stems are measured as a single cover.
Yuccas-Make separate observations for the leaves and caudex (thick basal stem). Break the observations into sections of leaves that are approximately the same height and record the cover as the perimeter around this group of leaf blades. The caudex is measured as a single cover. The thick leaves of yuccas make it difficult to make a cover measurement by centering yourself over the caudex of the plant. The cover of the caudex may be estimated by holding a niner next to it or using a tape measure to measure to approximate the area.
Height is recorded as a whole number in centimeters. All heights are vertical heights but they are not necessarily perpendicular to the ground if the ground is sloping.
Annual grasses and all forbs-Measure the height from the base of the plant to the top of the inflorescence (if present). Otherwise, measure to the top of the green foliage.
Perennial grasses-Measure the height from the base of the plant to the top of the live green foliage. Do not include the inflorescence in the height measurement. The presence of live green foliage may be difficult to see in the winter. Check carefully at the base of the plant for the presence of green foliage. If none is found it may be necessary to pull the leaf sheaths off of several plants outside the quadrat. From this you may be able to make some observations about where green foliage is likely to occur.
Perennial shrubs and sub-shrubs (non-creosote)-Measure the height from the base of the green foliage to the top of the green foliage, ignoring all bare stems. Do not measure to the ground unless the foliage reaches the ground.
Plants rooted outside but hanging into a quadrat-Do not measure the height from the ground. Measure only the height of the portion of the plant that is within the quadrat.
Additional Information on the personnel associated with the Data Collection / Data Processing
Nathan Gehres 2014-present; Michell Thomey 2012-2014
This dataset contains pinon-juniper woodland biomass data and is part of a long-term study at the Sevilleta LTER measuring net primary production (NPP) across four distinct ecosystems: creosote-dominant shrubland (Site C, est. winter 1999), black grama-dominant grassland (Site G, est. winter 1999), blue grama-dominant grassland (Site B, est. winter 2002), and pinon-juniper woodland (Site P, est. winter 2003). Net primary production is a fundamental ecological variable that quantifies rates of carbon consumption and fixation. Estimates of NPP are important in understanding energy flow at a community level as well as spatial and temporal responses to a range of ecological processes.
Above-ground net primary production is the change in plant biomass, represented by stems, flowers, fruit and and foliage, over time and incoporates growth as well as loss to death and decomposition. To measure this change the vegetation variables in this dataset, including species composition and the cover and height of individuals, are sampled twice yearly (spring and fall) at permanent 1m x 1m plots within each site. A third sampling at Site C is performed in the winter. Volumetric measurements are made using vegetation data from permanent plots (SEV278, "Pinon-Juniper (Core Site) Quadrat Data for the Net Primary Production Study") and regressions correlating species biomass and volume constructed using seasonal harvest weights from SEV157, "Net Primary Productivity (NPP) Weight Data."
Data Processing Techniques to Derive Biomass and NPP:
Data from SEV278 and SEV157 are used used to calculate seasonal and annual production of each species in each quadrat for a given year. Allometric equations derived from harvested samples of each species for each season are applied to the measured cover, height, and count of each species in each quadrat. This provides seasonal biomass for winter, spring, and fall.
Seasonal NPP is derived by subtracting the previous season's biomass from the biomass for the current season. For example, spring NPP is calculated by subtracting the winter weight from the spring weight for each species in a given quadrat. Negative differences are considered to be 0. Likewise, fall production is computed by subtracting spring biomass from fall biomass. Annual biomass is taken as the sum of spring and fall NPP.
Other researchers involved with collecting samples/data: Chandra Tucker (CAT; 04/2014-present), Megan McClung (MAM; 04/2013-present), Stephanie Baker (SRB; 10/2010-present), John Mulhouse (JMM; 08/2009-06/2013), Amaris Swann (ALS; 08/2008-present), Maya Kapoor (MLK; 08/2003 - 01/2005, 05/2010 - 03/2011), Terri Koontz (TLK; 02/2000 - 08/2003, 08/2006 - 08/2010), Yang Xia (YX; 01/2005 - 03/2010), Karen Wetherill (KRW; 02/2000 - 08/2009); Michell Thomey (MLT; 09/2005 - 08/2008), Heather Simpson (HLS; 08/2000 - 08/2002), Chris Roberts (CR; 09/2001- 08/2002), Shana Penington (SBP; 01/2000 - 08/2000), Seth Munson (SMM; 09/2002 - 06/2004), Jay McLeod (JRM; 01/2006 - 08/2006); Caleb Hickman (CRH; 09/2002 - 11/2004), Charity Hall (CLH; 01/2005 - 01/2006), Tessa Edelen (MTE, 08/2004 - 08/2005).
Begun in spring 2013, this project is part of a long-term study at the Sevilleta LTER measuring net primary production (NPP) across three distinct ecosystems: creosote-dominant shrubland (Site C), black grama-dominant grassland (Site G), and blue grama-dominant grassland (Site B). Net primary production is a fundamental ecological variable that quantifies rates of carbon consumption and fixation. Estimates of NPP are important in understanding energy flow at a community level as well as spatial and temporal responses to a range of ecological processes.
Above-ground net primary production is the change in plant biomass, represented by stems, flowers, fruit and and foliage, over time and incoporates growth as well as loss to death and decomposition. To measure this change the vegetation variables in this dataset, including species composition and the cover and height of individuals, are sampled twice yearly (spring and fall) at permanent 1m x 1m plots within each site. A third sampling at Site C is performed in the winter. The data from these plots is used to build regressions correlating biomass and volume via weights of select harvested species obtained in SEV999, "Net Primary Productivity (NPP) Weight Data." This biomass data is included in SEV999, "Seasonal Biomass and Seasonal and Annual NPP for Core Grid Research Sites."
Each sampling grid contains 40 1x1m quadrats in a 5x8 array. However, only 30 quadrats are sampled at each. These are quadrats 1-15 and 26-40. Thus, the middle two rows (i.e., 10 quadrats) are not sampled. Locating the Sampling Quadrats: Three core sites (B, G, and C) contain five rodent trapping and vegetation sampling webs. The vegetation grids are near these webs at each core site. At the blue grama site, the grid is located at the southern end of web 5, between webs 2 and 4. At the creosote site, the grid is east of web 3, near the road. At the black grama site, the grid is just northeast of web 5.
Net primary production data is collected twice each year, spring and fall, for all sites. The Five Points Creosote Core Site is also sampled in winter. Spring measurements are taken in April or May when shrubs and spring annuals have reached peak biomass. Fall measurements are taken in either September or October when summer annuals have reached peak biomass but prior to killing frosts. Winter measurements are taken in February before the onset of spring growth.
Height is recorded as a whole number in centimeters. All heights are vertical heights but they are not necessarily perpendicular to the ground if the ground is sloping.
Plants rooted outside but hanging into a quadrat-Do not measure the height from the ground. Measure only the height of the portion of the plant that is within the quadrat.
Creosote Measurements till 2013:
To measure creosote (i.e., Larrea tridenta) break the observations into two categories:
1.)Small, individual clusters of foliage on a branch (i.e., branch systems): Measure the horizontal cover of each live (i.e., green) foliage cluster, ignoring small open spaces (keeping in mind the 15% guideline stated above). Then measure the vertical "height" of each cluster from the top of the foliage to a plane created by extending a line horizontally from the bottom of the foliage. Each individual foliage cluster within a bush is considered a separate observation.
2.) Stems: Measure the length of each stem from the base to the beginning of live (i.e., green) foliage. Calculate the cumulative total of all stem measurements. This value is entered under "height" with the species as "stem" for each quadrat containing creosote. All other variable receive a default entry of "1" for creosote stem measurements. Do not measure dead stems or areas of dead foliage. If in doubt about whether a stem is alive, scrape the stem with your fingernail and check for the presence of green cambium.
Creosote Measurements 2013 and after:
Each creosote is only measured as one total cover. Each quad that contains creosote will have one cover observation for each creosote canopy in quad.
Recording the Data:
Excel spreadsheets are used for data entry and file names should begin with the overall study (npp), followed by the date (mm.dd.yy) and the initials of the recorder (.abc). Finally, "g" for "grid," along with the site abbreviation, should be added (i.e., gc, gg, gb). The final format for sites B, G, and C should be as follows: npp_core.mm.dd.yy.abgg.xls. File names should be in lowercase.
Other researchers involved with collecting samples/data: Chandra Tucker (CAT; 04/2014-present), Megan McClung (MAM; 04/2013-present), Stephanie Baker (SRB; 2013-present), John Mulhouse (JMM; 08/2009-06/2013).
The purpose of this project is to test the hypothesis that the smallest 50% of precipitation events during the monsoon season affect microbial functioning and grassland productivity in mixed grasslands of B.eriopoda and B. gracilis at the SNWR. At the SNWR, the summer monsoon season accounts for 60% of total annual precipitation and drives the majority of vegetation productivity during the year; the largest 25% of precipitation events account for the majority of this precipitation. I predict that important ecological variables such as nutrient and soil moisture availability are disproportionately influenced by smaller events. The proposed project will help tease apart the importance of precipitation event classes on nutrient availability and grassland aboveground net primary production (ANPP). This research will also provide a basis for understanding how increased aridity in the U.S. southwest due to increasing global surface temperature and altered precipitation could affect grassland communities at the SNWR.
We will implement 10 open plots (control) and 10 precipitation exclosure plots(treatment; 20 total plots) at a mixed blue and black grama grassland site at the SNWR. In this experiment, treatment plots will only receive the largest 50% of precipitation events. This will maintain statistically similar total precipitation between control and treatment plots because the smallest 50% of events have an insignificant effect on total seasonal precipitation. How these small events are linked to microbial activity and vegetation productivity is still very much unknown. I predict that soil microbial activity and nutrient availability will differ between control and treatment plots and will result in differing vegetation ANPP between them. These effects may become more distinct as time progresses, which is the reason for conducting this research for a series of monsoon seasons.
Existing precipitation exclosures (2.45 m x 2.45 m) will be employed at the mixed grassland site. We will implement 20 total plots (10 control, 10 treatment; approx. 500 m2 total area). Temporary site infrastructure will include 10 precipitation exclosures, a water tank (1100 gal.) and soil moisture probes. This infrastructure currently exists at the mixed grassland site and will be adopted from Michell Thomey's project entitled, "Soil moisture extremes and soil water dynamics across a semiarid grassland ecotone."
Precipitation is the only independent variable in this experiment. Using precipitation exclosures, I will remove all ambient precipitation from treatment plots from DOY 182-273. Ambient daily precipitation thatexceeds the estimated 50% threshold will be delivered to the plots within 24 hours of an event. Delivered precipitation will be adjusted for atmospheric demand differences.
Dependent variables in this experiment are vegetation ANPP, soil nitrogen content, soil enzymatic activityand soil moisture content. Vegetation biomass will be collected from the sites on DOY 181 and 274. Soil enzymatic activity will be determined approximately 4 times per monsoon season using plot soil samples. Soil nitrogen content will be measured under vegetation using nitrogen probes. Volumetric soil moisture content [m3 m-3] will be measured continuously using soil moisture probes (30 cm depth).
This dataset consists of profiles of soil water potential measured via in situ thermocouple psychrometers located within a rainfall manipulation experiment in a piñon-juniper woodland. The sensors are centrally located within 40 m x 40 m water addition, water removal, infrastructure control, and ambient control plots. The profiles are installed under each of ten target piñon and juniper trees (five of each species) which were also used for other physiological measurements, as well as at five intercanopy areas. The raw sensor output (in μV) has been temperature-corrected and individual calibration equations applied.
Background: This sensor array is part of a larger experiment investigating the mechanisms of drought-related mortality in the piñon-juniper woodland. Briefly, one hypothesis is that, during periods of extended, very-negative, soil water potential (“drought”) trees experience xylem tensions greater than their threshold for cavitation, lose their hydraulic connection to the soil, dessicate and die. A second hypothesis is that in order to avoid this hydraulic failure, trees restrict water loss via reduction in stomatal conductance which also limits the diffusion of CO2 for photosynthesis, and eventually may starve to death depending on the drought duration.
The goal of the project is to apply drought stress on an area significantly larger than the scale of individual trees to determine whether hydraulic failure or carbon starvation is a more likely mechanism for mortality under drought conditions. The cover control treatment replicates the microenvironment created under the plastic rainfall removal troughs (slightly elevated soil and air temperatures and relative humidity) without removing ambient precipitation. The water addition treatment is intended to simulate 150% of the 30-yr average rainfall via n = 6 19-mm super-canopy applications during the growing season (April-October). More details on the experiment can be found in Pangle et al. 2012 Ecosphere 3(4) 28 (http://dx.doi.org/10.1890/ES11-00369.1). These three treatments, in addition to an ambient control with no infrastructure, are applied to three replicated blocks, one on relatively level terrain, one on a southeast-facing slope, and one on a north-facing slope. The soil psychrometers were installed in the southeast-facing block only, to measure the effectiveness of the treatments on plant-available soil moisture.
For the purposes of comparing soil water potential under various cover types (piñon, juniper, intercanopy), it should be noted that significant tree mortality has occurred on Plot 10. As described below, on 5 August 2008 the site was struck by lightening and many of the soil psychrometers were rendered inoperable. At approximately the same time, four of the five target piñon trees in the southeast-facing drought plot (Plot 10) started browning and it was discovered that they had bark beetle (Ips confusus) galleries and were infected with Ophiostoma fungi. By October 2008 those trees (T1, T2, T4, and T5) had dropped all their needles. Therefore the psychrometers buried under them were no longer located “under trees” after that time. By June 2009 the remaining target piñon (T3) died. By March 2010, one of the target juniper trees (T10) had died. At the time of this writing (April 2011) it is anticipated that more of the juniper trees on P10 will die during this year.
Methods: Within Plots 9-12 of the larger PJ rainfall manipulation experiment, thermocouple psychrometers (Wescor Inc., Logan, UT, USA) were installed. Soil psychrometers profiles were placed under each of the initial ten target trees in each plot, and at the same five intercanopy areas that were instrumented to measure soil and air temperature and soil volumetric water at 5 cm depth. Each profile consisted of sensors at (1) 15 cm; (20) cm; and (3) as deep as could be augered and installed by hand, generally 50-100 cm depth. Sensors were calibrated with four NaCl solutions of known water potential before field deployment.
Sensors are controlled via a Campbell Scientific CR-7 datalogger (Campbell Scientific, Logan, UT, USA). The datalogger takes measurements every 3 h but soil water potential does not change that fast and the daytime measurements are generally unusable because of thermal gradients between the datalogger and the sensors, therefore the data presented here are only the 3:00 AM timepoints.
Note that on 5 August 2008 the site was struck by lightening. Many of the soil psychrometers were destroyed by ground current, with the worst damage on the drought and cover control plots where the metal support posts for the infrastructure may have helped to propagate ground current. Some of those sensors were eventually replaced but in the case of the infrastructure plots we were limited to installing new sensors between the plastic troughs because it was impossible to auger under the plastic. Therefore, while the original installation was random with regard to the pattern of plastic domes and troughs, the replacement installation was exclusively outside the plastic and the data may therefore be biased towards wetter microsites.
Instrument Name: thermocouple psychrometer with stainless steel screen
Manufacturer: Wescor, Inc, Logan, UT, USA
Model Number: PST-55
Environmental temperature influences virtually all aspects of organismal performance, including fitness. And since temperature varies throughout space and time, organisms must regularly compete for optimal thermal habitats, much as they do for other resources (e.g. territory, food, or females). However, competition for thermal resources imposes costs, often in the form of a stress response (i.e. increased corticosterone production). Elevated corticosterone promotes physiological and behavioral responses that can increase an organism’s chance of survival, but if left in an organism’s system for too long, it will reduce immunity, degenerate neurons, and lower fitness. Previous theoretical and empirical work indicates that, all else being equal, patchy thermal landscapes reduce the energetic cost of thermoregulation. Therefore, I hypothesize that lizards exposed to patchy distributions of preferred temperatures will have less stress (and thus lower levels of corticosterone) than those exposed to clumped distributions. Furthermore, patchily distributed resources are more difficult for territorial males to monopolize, and thus, subordinate males in patchy thermal landscapes should experience less stress than subordinate males in clumped thermal landscapes.
Experimental design: Starting in the July of 2012, I will initiate this project as part of a continuing large-scale field study at Sevilleta LTER site in collaboration with PI Michael Angilletta’s Spatially Explicit Theory of Thermoregulation project. As in past research conducted in 2008, 2009 and 2011, I will use male Yarrow’s spiny lizard. This lizard thermoregulates accurately in the absence of predators14,15 and aggressively defends resources from conspecific males14,16.
Nine outdoor arenas (20 x 20 m), consisting of sheet metal walls and a canopy of shade cloth, will be used to manipulate the thermal environments. Among the arenas, three patterns of shade patches will be replicated three times each to generate distinct thermal landscapes (see Figure 1). Lizards will be paired by size: large dominant (22-30 g) with a small subordinate (15-21 g). Each pair (n = 12) will be randomly assigned one of the thermal environments. Prior to each trial, males will be habituated to their arenas for 10 days. During this period, each male will be exposed to the thermal arena every other day (for a 24-h period) in the absence of a competitor (total of 5 days per animal). After the habituation period, males will be placed in arenas for a 4-day testing period. Males will spend two of these days in isolation and the other two in competition. Half the pairs will start the trial in isolation (solitary treatment), and the other half of the pairs will start the trial in competition (social treatment). A matched pair of lizards will be placed together in one arena, and the other two arenas will each have one individual (either small or large) placed into it. After two days, all lizards will be captured and blood samples will be collected within three minutes (speed of collection is necessary to prevent handling stress from affecting plasma corticosterone levels17). Blood will be taken from the orbital sinus with a glass capillary tube and then taken back to the lab where the plasma will be obtained through centrifugation. Plasma will be stored at -80˚C for hormone assays18. After bleeding, solitary lizards will be placed together in one arena, and the previously paired individuals will be separated and split between the two remaining arenas. Thus, a completed habituation and observation set for six pairs (two pairs per type of thermal environment) will take 14 days. And 3 sets will be conducted per season giving a total of 18 pairs per season in each thermal environment (54 pairs in isolation and competition per season). Mixed modeling procedures in the statistical software R will be used to quantify the effects of competition and thermal patchiness on the corticosterone levels of lizards19.
Fig. 1. Patterns of shade patches for arenas (each replicated 3x).
In the southwestern United States two important seasons influence stream flow: snowmelt in spring and summer monsoonal rainfall events. Flow patterns exhibit peak discharge from snowmelt runoff in the spring followed by pulsed increases in stream discharge during late summer monsoons. Molles and Dahm showed the intensity of the snowmelt discharge is linked to El Nino-Southern Oscillation (ENSO) conditions in the tropical Pacific. El Nino and La Nina climate patterns also may affect late summer monsoonal precipitation in New Mexico by intensifying the monsoon during La Nina years and weakening monsoons during El Nino years. Stage gage data show seasonal and interannual variability in the intensity of snowmelt and monsoonal runoof events in montane catchments in New Mexico. Further, in-situ YSI sonde, Satlantic Submersible Ultraviolet Nitrate Analyzer (SUNA) and CycleP instrumentation show physical and chemical constituents respond to higher flow events driven by climate variability, and the constituents these instruments measure can be used as a proxy to estimate whole stream metabolism and nutrient cycling processes.
Data Selection: Historical flux tower data from 2007 to 2011 was provided by UNM Marcy Litvak for two locations near our study site on the EFJR. Los Alamos National Lab provided flux data from 2005 to 2006. Flux towers provided photosynthetic active radiation (PAR) and barometric pressure in 30 minute time intervals. In-stream YSI sondes continuously monitored the Jemez and East Fork Jemez Rivers in 15 minute time intervals collecting water quality data (dissolved oxygen, pH, turbidity, specific conductance, water temperature).
Instrument Name: YSI Sonde Manufacturer: YSI IncorporatedModel Number: 6920V2-0
Sonde and flux data were QAQC'd using Aquarius software to delete suspicious data (or outliers) and to correct for drift from biofouling on probes.
Study Area Name: East Fork Jemez River
Study Area Location: Valles Caldera National Preserve, New Mexico
Study Area Description:
Elevation: 2582 meters
Landform: Montane grassland, caldera
Soils: Rich organic soils; Mollisols
Hydrology: snowpack(winter) and monsoonal rainfall (summer)
Vegetation: grassland, meadow
Site history: Domestic grazing of sheep from mid-1800's to 1940's, then cattle by 1940's.
Single Point: EFJR at Hidden Valley (from VCNP)
North Coordinate: 35.83666667
West Coordinate: -106.5013833
The distribution, structure and function of mesic savanna grasslands are strongly driven by fire regimes, grazing by large herbivores, and their interactions. This research addresses a general question about our understanding of savanna grasslands globally: Is our knowledge of fire and grazing sufficiently general to enable us to make accurate predictions of how these ecosystems will respond to changes in these drivers over time? Some evidence suggests that fire and grazing influence savanna grassland structure and function differently in South Africa (SA) compared to North America (NA). These differences have been attributed to the contingent factors of greater biome age, longer evolutionary history with fire and grazing, reduced soil fertility, and greater diversity of plants and large herbivores in SA. An alternative hypothesis is that differences in methods and approaches used to study these systems have led to differing perspectives on the role of these drivers. If the impacts of shared ecosystem drivers truly differ between NA and SA, this calls into question the generality of our understanding of these ecosystems and our ability to forecast how changes in key drivers will affect savanna grasslands globally. Since 2006, an explicitly comparative research program has been conducted to determine the degree of convergence in ecosystem (productivity, N and C cycling) and plant community (composition, diversity, dynamics) responses to fire and grazing in SA and NA.
Thus far, initial support has been found for convergence at the ecosystem level and divergence at the community level in response to alterations in both fire regimes and grazing. However, there have also been two unexpected findings (1) the ways in which fire and grazing interact differed between NA and SA, and (2) the rate of change in communities when grazers were removed was much greater in NA than in SA. These unexpected findings raise a number of important new questions: (Q1) Will exclusion of grazing eventually affect community structure and composition across all fire regimes in SA? (Q2) Will these effects differ from those observed in NA? (Q3) What are the determinants of the different rates of community change? (Q4) How will these determinants influence future trajectories of change? (Q5) Will the different rates and trajectories of community change be mirrored by responses in ecosystem function over time? This project is based on a large herbivore exclusion study established within the context of long-term (25-50+ yr) experimental manipulations of fire frequency at the Konza Prairie Biological Station (KPBS) in NA and the Kruger National Park (KNP) in SA. The suite of core studies and measurements include plant community composition, ANPP, and herbivore abundance and distribution at both study sites to answer these research questions.
This data set was added to the Sevilleta LTER archive at the request of the SEV Principal Investigator Scott Collins.
Data are collected twice each year at each site. Sample periods are equivalent to spring and late summer at each study site (December/January and March/April in South Africa, May and September in North America.
Where the Data were Collected:
Ukulinga Research Farm, Pietermaritzburg, South Africa; Satara Region of Kruger National Park, South Africa; Konza Prairie Biological Station, North America
Additional Geographic Metadata:
Ukulinga Research Farm (URF), South Africa. The URF of the University of KwaZulu-Natal is located in Pietermaritzburg, in southeastern South Africa (30o 24’ S, 29o 24’ E). The site is dominated by native perennial C4 grasses, such as Themeda triandra and Heteropogon contortus, that account for much of the herbaceous aboveground net primary production (ANPP). Mean annual precipitation is 790 mm, coming mostly as convective storms during summer (Oct-Apr). Summers are warm with a mean monthly maximum of 26.4oC in February, and winters are mild with occasional frost. Soils are fine-textured and derived from shales. There has been no grazing at this site for >60 years. Long-term experimental plots were established at URF in 1950 with the objective of determining the optimal fire and/or summer cutting regime to maximize hay production. The experiment is a randomized block (three replicates) split-plot design with four whole-plot haying treatments and 11 subplot fire or mowing treatments. Subplot sizes are 13.7 x 18.3 m.
Kruger National Park (KNP), South Africa. The KNP is a 2 million ha protected area of savanna grassland that includes many of the large herbivores common to southern Africa (22º 25' to 25º 2 32' S, 30º 50' to 32º 2' E). The extant abundance and grazing intensity of herbivores in KNP is considered moderate for regional savanna grasslands. In the south-central region of KNP where our research takes place, average rainfall is 537 mm with most falling during the growing season (Oct-Apr). The dormant season is mild, dry and frost free, and summers are warm with mean monthly maximum air temperature of 28.9oC in January. Because of the importance of fire in savanna grassland ecosystems, the Experimental Burn Plot (EBP) experiment was initiated in 1954 to examine the effects of fire frequency (control-no fire, 1-, 2-, 3-, 4- and 6-yr return interval) and season [early spring (Aug), spring (Oct), mid-summer (Dec), late summer (Feb), and fall (Apr)] on vegetation communities in the park. Four blocks of 12 plots (two were later split for the 4- and 6-yr trts), each ~7 ha (370 x 180 m) in size, were established in four primary vegetation types covering the two major soil types (granites and basalts) and spanning the precipitation gradient in the park. Each plot has 50+ years of known fire history, and native herbivores have had unrestricted access, thus fire and grazing effects are combined. This research focuses on the EBPs located near Satara where precipitation, soil type, and the mix of herbaceous and woody plants are similar to KPBS. Vegetation on the blocks is co-dominated by C4 grasses, such as Bothriochloa radicans, Panicum coloratum and Digiteria eriantha, and woody plants, such as Acacia nigrescens and Sclerocarya birrea. Soils are fine-textured and derived from basalts. Adjacent to one of the Satara blocks is the Cape buffalo enclosure, erected in 2000 for veterinary purposes. The 200 ha permanent enclosure contains 65-80 animals and is divided into 4 blocks burned on a rotational basis. The grazing intensity inside is comparable to the moderate levels imposed in the park and at KPBS. Two blocks are burned annually while others are burned infrequently (approximately once every 4-yr).
Konza Prairie Biological Station (KPBS), North America. The KPBS is a 3,487 ha savanna grassland in northeastern Kansas, USA (39o 05’ N, 96o 35’ W) dominated by native perennial C4 grasses such as Andropogon gerardii and Sorghastrum nutans that account for the majority of ANPP. Scattered shrub and tree species include Cornus drummondii, Gleditsia triacanthos, and Prunus spp. Numerous sub-dominant grasses and forbs contribute to the floristic diversity of the site. The climate is continental, with mean July air temperature of 27°C. Annual precipitation is ca. 820 mm/year, with 75% falling as rain during the Apr-Oct growing season. Soils are fine textured, silty clay loams derived from limestone and shales. KPBS includes fully replicated watershed-level fire and fire/grazing treatments, in place since 1977 and 1987, respectively. Replicate watersheds (mean size ~60ha) are burned at 1-, 2-, 4-, 10- and 20-yr intervals, mainly in April, to encompass a range of likely natural fire frequencies and management practices. A subset of watersheds has not been grazed for more than 30 years. To address the role of native grazers and fire/grazing interactions, bison (~260 individuals) were reintroduced to KPBS in a 1000-ha fenced area that includes replicate watersheds burned in the spring at 1-, 2-, 4- and 20-year intervals. The overall grazing intensity is considered moderate.
Study Area 1:
Study Area Name: Ukulinga Research Farm
Study Area Location: Near Pietermaritzburg, South Africa
Elevation: 840 m above sea level
Landform: Colluvium fan
Geology: Marine shales and dolerite colluvium
Soils: Dystric leptosols, Chromic luvisols, Haplic plinthisols
Vegetation: Native grassland
Climate: Mean annual precipitation is 844 mm, Mean annual temperature 17.6C
Site history: Ungrazed since 1950
Single Point: 29o 40’ S / 30o 20’ E
Study Area 2: Kruger National Park, South Africa
Study Area Name: Satara Experimental Burn Plots and Cape Buffalo Exclosure
Study Area Location: Near Satara rest camp
Elevation: 240-320 meters above sea level
Landform: Level Upland
Soils: Rhodic nitisols, Haplic luvisols, Leptic phaeozems
Climate: Mean annual precipitation 544 mm; mean annual temperature 21.2–23.3C
Site history: Grazed by native herbivores
Single Point: 23–25o S /30-31o E
Study Area 3: Konza Prairie Biological Station
Study Area Name: Konza Prairie
Study Area Location: Watersheds N20B, N4D, N1B, N4B; 1D, 4F, 20B
Elevation: 320-444 meters above sea level
Landform: Alluvial terrace
Geology: Cherty limestone and shale
Soils: Udic argiustolls
Climate: Mean annual precipitation 835 mm; mean annual temperature 12.7C
Site history: Ungrazed watersheds (since 1971), watersheds grazed by native herbivores (since 1987)
Single Point: 39o 05.48’ N / 96o 34.12’ W
We used comparable experimental designs and sampling procedures at both URF and KPBS. At URF we used three replicate plots (not hayed or mowed) that have been burned every 1 and 3 years in the spring, and those left unburned (N=9 plots). At KPBS, we established replicate plots in experimental watersheds burned every 1 and 4 years in the spring, and those left unburned (N=9 plots). Thus, the only difference in design between NA and SA was the intermediate burn frequency. In 2005 at both sites we established four 2x2m areas in each replicate of the 1-yr, 3-4 yr burned, and unburned plots (N=36 subplots). We then randomly selected two of the subplots for the fertilization treatment and the other two subplots served as controls (Fig. 1). Starting in 2006 at KPBS and 2007 at URF, we began adding 10 gN/m2/yr as NH4+NO3- to assess the interactive effects of fire frequency and nitrogen limitation on plant community composition, structure and dynamics.
Fig. 1. Experimental design and sampling for the proposed studies: A) the role of long-term fire regimes (without megaherbivores), B) the importance of grazing and grazing/fire interactions, and C) the role of megaherbivore diversity. Moveable exclosures (3/plot) will be used to estimate ANPP in the grazed plots. N addition subplots (2 x 2 m) will be divided into 4 1 x 1 plots, with two designated for plant species composition sampling and the other two for destructive sampling. Soil samples will be collected from areas not designated for ANPP or plant composition sampling. Note that the same annually and infrequently burned plots at Kruger and Konza will be used in (B) and (C). In addition, similar plots will be established minus the N addition subplots in the 1-yr and 4-yr burned blocks of the Buffalo enclosure for (C).
Each of the 2x2m subplots was divided into four 1x1m quadrats. Annually since 2005 (prior to nitrogen addition) canopy cover of each species rooted in each quadrat was visually estimated twice during the growing season to sample early and late season species. As a surrogate for aboveground production, we measured light availability at the end of the growing season above the canopy at the ground surface in each quadrat (N=4 per subplot) using a Decagon ceptometer.
Net primary production measurements: Prior to the 2005 growing season we established plots (13.7 m by 18.3 m) in ungrazed areas burned annually, at 3–4-year intervals, and unburned (n = 3 per fire treatment) at both KBPS and URF. Areas with trees or large shrubs were avoided as our main goal was to evaluate responses in the herbaceous plant community. ANPP was estimated from end-of-season harvests starting in 2005 (September for KBPS, April for URF). In 10, 0.1-m2 (20 cm by 50 cm) quadrats randomly located in each plot (n = 30/treatment/site), we harvested the vegetation at ground level and separated it into grass, forb, and previous year’s dead biomass. Samples were dried at 60C to a constant weight. For annually burned plots, total biomass harvested represents ANPP. For the intermediate and unburned sites, we calculated ANPP by summing all but the previous year’s dead component.
To assess the impacts of fire on ANPP in grazed areas, we established herbivore exclusion treatments in KBPS in North America and KNP in South Africa. Herbivore exclosures in grazed areas in KPBS and KNP were erected prior to the 2006 growing season. The exclosures were 7 m in diameter, 2 m tall, and constructed of diamond mesh (5-cm diameter). Seven exclosures were established in each of three blocks of the three fire treatments— annually burned, intermediate burn (3- years for KNP or 4-years for KPBS), and unburned (n = 21 exclosures/treatment/site). As our focus was on ANPP responses of the herbaceous layer, exclosures were not located beneath trees or where dense shrub patches were present. Additionally, in the Satara region of KNP is a 900-ha permanent enclosure containing 80–90 adult African buffalo (S. caffer). This enclosure was erected in 2000 and was divided into six areas (100–200 ha each), with these burned on a rotational basis including plots burned annually and plots that were unburned. We used the unburned and annually burned areas in the buffalo enclosure to provide a direct comparison for determining the effects of a single-species large grazer in KNP and KPBS, and to assess the effects of large herbivore diversity at adjacent sites in KNP. Similar exclosures were built in the African buffalo enclosure at KNP. We placed 7 exclosures in the three blocks of each fire treatment (annually burned and unburned) resulting in 21 exclosures/treatment. We sampled ANPP by harvesting plant biomass from three 0.1 m2 quadrats per herbivore exclosure at the end of the growing season starting in 2006.