organisms

Stress Response from Male-Male Competition in Varying Thermal Environments at the Sevilleta National Wildlife Refuge, New Mexico

Abstract: 

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.

Additional Project roles: 

32
33

Data set ID: 

284

Core Areas: 

Keywords: 

Methods: 

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).

Fungal Thermophile Survey at the Sevilleta National Wildlife Refuge, New Mexico

Abstract: 

Biological soil crusts (BSCs) are complex assemblages of fungi, lichens, bacteria, mosses and green algae that stabilize surface soils and manage and traffic photosynthate, nutrients and water to diverse microbial and producer communities in arid environments worldwide.  In Sevilleta grasslands, BSCs occupy much of the open space between clumps of vegetation and vary substantially in terms of structure. 

BSCs have important biological and physical roles.  They have been termed ‘mantles of fertility’ because of their general importance in biogeochemical cycling and net primary production in arid ecosystems.  It has been proposed that BSCs play a role in the rapid movement of N, C and water from open areas to plants (see below).  BSCs stabilize soils, and physical and chemical disturbances of BSCs lead to topsoil loss and dust storms.  BSCs are therefore critical components in efforts to understand implications of both climate change and physical disturbance.  Related to this, it has been suggested that BSC diversity can be used to inform conservation policies.

BSCs have been the subject of several previous Sevilleta LTER studies.  Green et al. showed that stable-isotope carbon and nitrogen could be transferred bi-directionally between BSCs and adjacent plants.  This led Collins et al. to propose that fungal hyphae provide connections between plant roots and BSCs that allow for transport between the two, a proposal known as the “fungal loop hypothesis.”  Porras-Alfaro et al. have surveyed the diversity of fungi in BSCs from Sevilleta grasslands using molecular methods.  We have also shown that thermophilic fungi are common in BSCs (unpublished results), a result that is not unexpected given the high summer temperatures attained in Sevilleta surface soils.  Yet, many questions remain regarding the organisms present in BSCs, their biological roles and how long it takes for BSCs to re-establish after disturbance.  Long-term, we are interested in the types of fungi present in BSCs and in how fungi function in transporting nutrients between BSCs and adjacent plants.  We are also interested in the extent to which specific fungi provide structure to BSCs and in how they help protect from stress agents such as desiccation.  We are interested in the extent to which fungi might help BSCs tolerate high summer soil temperatures, which often reach ≥ 60C.  We therefore have a special interest in thermophilic fungi present in the BSCs.  To date, little has been done to actually culture fungi from Sevilleta BSCs, hence the need for the current study.  

In summary, BSCs are one of the most important features of aridland ecosystems and form a critical interface between physical and biological domains.  Understanding the roles of BSCs in protecting soil structure, and in the cycling of carbon, water and nitrogen, is fundamental to aridland ecology.  The work proposed here continues efforts to characterize the specific fungi associated with Sevilleta BSCs.  It is a modest but important step toward addressing the long-term goals mentioned above.

Core Areas: 

Additional Project roles: 

31

Data set ID: 

281

Keywords: 

Methods: 

For each sampling site and sampling period a small amount of surface crust (approx. one teaspoon per sample) was taken from each of 10 locations at approximately 1 meter intervals across a transect.  Samples were transported back to the laboratory in plastic bags.

On rare occasions, a larger sample of 0.5 liter volume or less may have been removed at one or two sampling stations.

Additional information: 

Data was collected at: LTER PJ site (N 34 23’ 08.7” W 106 31’ 27.0”), a sand dune above the railroad tracks near the Sevilleta wetlands (N34 18' 06.5"  W106 51' 14.1"), gypsum outcroppings (N34 12' 40.5"  W106 45' 35.5"), grasslands near the Sev LTER warming and monsoon sites (N34 21' 34.3" W106 41' 29.4" and N34 20' 38.1"  W106 43' 34.5"), and the Rio Grande Bosque (N34 19'45" W106 51'40").

 

The Contribution of Biological Soil Crust Carbon and Nitrogen Exchange to the Net Ecosystem Exchange Along an Elevation Gradient at the Sevilleta National Wildlife Refuge, New Mexico

Abstract: 

The purpose of this project is to: 1.) determine how biological soil crust (BSC) cover changes along an elevation gradient and across seasons, 2.) determine how carbon and nitrogen exchanges of BSC communities vary with temperature along an elevation gradient in arid and semi-arid environments and, 3.) use photosynthetic and respiration rates of BSC communities to determine how the contribution of the BSC communities to whole ecosystem carbon exchange varies across the same gradient and over seasons.

Core Areas: 

Additional Project roles: 

247

Data set ID: 

280

Keywords: 

Methods: 

At each sampling site and sampling period a small amount of surface crust (approx. one teaspoonful per sample) was taken from each of 10 locations at approximately 1 meter intervals over a transect.  Samples were transported back to the laboratory in plastic bags.

On rare occasions we removed a larger sample, 0.5 liter volume or less, at one or two sampling stations.

Additional information: 

Study sites included: Flux tower sites, desert grassland, desert shrubland, juniper savanna, piñon-juniper woodland, ponderosa pine forest, and mixed conifer forest.

Konza Species Composition: Fire by Nitrogen Project

Abstract: 

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.

Data set ID: 

268

Core Areas: 

Keywords: 

Methods: 

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. 

Additional information: 

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

Geology: Basalts

Soils: Rhodic nitisols, Haplic luvisols, Leptic phaeozems

Vegetation: Native grassland

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

Vegetation: Native grassland

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

Ukulinga Species Composition: Fire by Nitrogen Project

Abstract: 

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.

Data set ID: 

263

Core Areas: 

Keywords: 

Methods: 

Konza-Ukulinga fire by nitrogen project: We used comparable experimental designs and sampling procedures at both URF and KPBS (Figure 1). 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. 

Konza-Kruger fire by grazing project: For this study, we are utilizing the long-term experiments at KPBS and KNP in which native megaherbivore grazers are present and fire frequency is directly manipulated. To assess the effects of grazing and fire-grazing interactions, we constructed seven sets of permanent exclosures and adjacent control plots in three blocks at both sites. The exclosures and matching paired open plots were established in 2005 in the Satara EBPs that are burned every 1 and 3 years in the spring or left unburned and at KPBS in watersheds that are burned every 1 and 4 years or left unburned. (N=63 exclosures/site; Fig. 1). Within each exclosure and paired open plot, we sample plant community composition and light availability in permanent 2x2 m subplots. We collect ANPP at the end of each growing season from each exclosure, and throughout the growing season in grazed areas adjacent to the unexclosed plots using 1x1 m moveable exclosures (Fig. 1).

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. 

Additional information: 

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

Geology: Basalts

Soils: Rhodic nitisols, Haplic luvisols, Leptic phaeozems

Vegetation: Native grassland

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

Vegetation: Native grassland

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

Soil microbial response to altered precipitation regimes at the Sevilleta National Wildlife Refuge, New Mexico

Abstract: 

Microbes substantially control many biogeochemical processes in semiarid systems, including carbon and nitrogen fixation and carbon mineralization. Bacteria and fungi are osmotrophs that release enzymes into the environment to generate assimilable carbon and nutrients from organic particles. These enzymes are also the first agents to respond to pulses of soil moisture. The capacity to stabilize extracellular enzymes on soil particles preserves the utility of these nutrient-generating agents during extended dry periods. Enzyme stability can relate to environmental conditions and increase with clay mineral and humic compound concentrations. To better understand microbial response to precipitation variability, our objective was to determine the stability of extracellular enzymes under various monsoon precipitation regimes. During summer 2010, soil enzyme activity was measured in a rainfall manipulation study within a mixed-grass semiarid grassland in New Mexico, USA. Plots received either one large rain event or three evenly spaced small rain events per month. Before and after the first rain of each month, soil samples from the rhizosphere and from interspaces between plants were collected and analyzed for activity of four hydrolases; beta-glucosidase, beta-N-acetylglucosaminidase, leucine aminopeptidase, and alkaline phosphatase. 

Additional Project roles: 

303

Data set ID: 

245

Core Areas: 

Keywords: 

Methods: 

For experimental design and precipitation manipulations see SEV218.

Before the first rain of each month, soil samples were collected from the rhizosphere and interspaces between plants. Four soil cores (1cm wide, 3 cm deep) were taken across the plot, with rhizoshpere samples from under B. eriopoda and B. gracilis, and mixed together for each sample. Enzyme activity in the rhizosphere and interspace were analyzed separately. Two hours after the rain event, soil samples were again collected in the same manner. Microbial response to precipitation is quick therefore 2 hours was ample time to assess microbial response. Samples were refrigerated and processed within 48 hours of collection to prevent enzyme degradation. Soils were subsampled for organic matter and water content. Field soil moisture was calculated by comparing weights of freshly collected soil and soil dried at 60 °C. A subsample was also burned at 500 °C for 4 hours to determine percent organic matter. The potential activity levels of beta-glucosidase, beta-N-acetylglucosaminidase, leucine aminopeptidase, alkaline phosphatase, and phenol oxidase were measured in the lab following the methods of Stursova et al. (2006).

The varied topography and large elevation gradients that characterize the arid and semi-arid Southwest create a wide range of climatic conditions - and associated biomes - within relatively short distances. This creates an ideal experimental system in which to study the effects of climate on ecosystems. Such studies are critical givien that the Southwestern U.S. has already experienced changes in climate that have altered precipitation patterns (Mote et al. 2005), and stands to experience dramatic climate change in the coming decades (Seager et al. 2007; Ting et al. 2007).

Gunnison’s Prairie Dog Use of Resource Pulses in a Chihuahuan Desert Grassland at the Sevilleta National Wildlife Refuge, New Mexico: Re-sight Scan Data

Abstract: 

Seasonal environments experience cyclical or unpredictable pulses in plant growth that provide important resources for animal populations, and may affect the diversity and persistence of animal communities that utilize these resources. The timing of breeding cycles and other biological activities must be compatible with the availability of critical resources for animal species to exploit these resource pulses; failure to match animal needs with available energy can cause population declines. Adult Gunnison’s prairie dogs emerge from hibernation and breed in early spring, when plant growth is linked to cool-season precipitation and is primarily represented by the more nutritious and digestible plants that utilize the C3 photosynthetic pathway. In contrast, summer rainfall stimulates growth of less nutritious plants using the C4 photosynthetic pathway. Prairie dogs should therefore produce young during times of increased productivity from C3 plants, while pre-hibernation accumulation of body fat should rely more heavily upon C4 plants.  If seasonal availability of high-quality food sources is important to Gunnison’s prairie dog population growth, projected changes in climate that alter the intensity or timing of these resource pulses could result in loss or decline of prairie dog populations.  This project will test the hypothesis that population characteristics of Gunnison's prairie dog, an imperiled grassland herbivore, are associated with climate-based influences on pulses of plant growth.

Data set ID: 

242

Core Areas: 

Additional Project roles: 

40
41

Keywords: 

Methods: 

Gunnison’s prairie dogs will be monitored at 6 colonies, with 3 colonies each occurring with the range of prairie and montane populations. Colonies for study within the prairie populations occur at Sevilleta National Wildlife Refuge (n = 3 prairie populations) and at Vermejo Park Ranch (n = 3 montane populations).  Live-trapping of prairie dogs will be conducted during 3 periods of the active seasons—following emergence (April), after juveniles have risen to the surface (mid-to-late June), and pre-immergence (beginning in August).  Trapping will occur for 3-day periods, following pre-baiting with open traps.  At capture, sex and body mass of each individual will be recorded.  Blood and subcutaneous body fat samples will be collected nondestructively for analysis of isotopic composition.  Prairie dogs will be marked with dye, and released on site immediately following processing.  After trapping periods at each site have concluded, population counts will be conducted during 2-3 re-sighting (or recapture) periods for each prairie dog colony.  Resighting observation periods will be ~3 hours in length, and consist of 2-6 systematic scans of the entire colony, beginning and ending from marked points outside of the colony boundary.  During each observation period, prairie dogs will be counted, recorded as marked or unmarked, and location on the colony noted.  

Vegetation cover and composition measurements will be collected (or obtained at Sevilleta, where such data is already being collected) during pre- and post-monsoon periods of the active season.  Total cover will be measured by plant species (or to genus if species is indeterminable). Total cover will be measured at 12 grid points per colony using Daubenmire frames (0.5 m x 0.5 m), and at 12 grid locations 200-800 m outside of each colony boundary.  Adjacent to each Daubenmire frame, a 20 cm x 30 cm sample of vegetation will be clipped and dried for determination of volumetric moisture content of vegetation.  

Primary productivity variables (cover, moisture content) will be tested for correlations to individual and population-level condition indicators in prairie dogs.  Carbon isotope ratios (δ13C) from prairie dog blood and fat samples will be analyzed on a continuous flow isotope ratio mass spectrometer.  The relative contribution of C3 and C4 plants to the diet of each individual will be determined based upon δ13C ratios for C3 and C4 plants in the study area and a 2-endpiont mixing model, and will be calculated for each individual animal, population and season.  Population estimates will be calculated using mark-resight estimates, and compared to maximum above-ground counts.  The influence of resource pulses on prairie dog population parameters will be tested by comparing the vegetation cover, moisture content, and ratio of total C3:C4 plant cover to the ratio of C3:C4 plants in prairie dog diets, population estimates, and juvenile:adult ratios as an index to population recruitment.   

Instrumentation: 

*Instrument Name: Continuous flow isotope ratio mass spectrometer

*Manufacturer: Thermo-Finnigan IRMS  Delta Plus 

*Instrument Name: Elemental Analyzer

*Manufacturer: Costech

*Model Number: ECS4010

Additional information: 

Other Field Crew Members: Talbot, William; Duran, Ricardo; Gilbert, Eliza; Donovan, Michael; Nichols, Erv; Sevilleta LTER prairie dog field crew led by Koontz, Terri; Sevilleta NWR prairie dog field crew led by Erz, Jon.

Tissue samples are analyzed for stable carbon isotope ratios in stable isotope laboratory operated by Dr. Zachary Sharp and Dr. Nicu-Viorel Atudorei of the Department of Earth and Planetary Sciences, University of New Mexico.

The primary objective of this study is to examine the control that substrate quality and climate have on patterns of long-term decomposition and nitrogen accumulation in above- and below-ground fine litter. Of particular interest will be to examine the degree these two factors control the formation of stable organic matter and nitrogen after extensive decay.

Plant phenology or life-history pattern changes seasonally as plants grow, mature, flower, and produce fruit and seeds. Plant phenology follows seasonal patterns, yet annual variation may occur due to annual differences in the timing of rainfall and ambient temperature shifts. Foliage growth and fruit and seed production are important aspects of plant population dynamics and food resource availability for animals.

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