Boyd Deep Canyon Desert Research Center

01 Jun, 06 AM - 17 Jun, 10 AM

Ben Wissinger Thesis Proposal May 1, 2011 Sonoran Desert Harvester Ant Response to Atmospheric Nitrogen Deposition Abstract Due to increases in airborne nitrogen pollution, excess nitrogen deposition has been affecting soil biogeochemical processes and plant community composition across the West. A number of studies have been conducted in forested systems and arid systems near urban centers; however, these investigations typically end with soil and plant tissue analysis. This study will build upon the work performed by Allen et al. 2009 and Rao et al. 2009, whose efforts revealed an atmospheric nitrogen deposition gradient from the San Bernardino Mountains through Joshua Tree National Park. I propose to bridge the gap between the effects of increased nitrogen on plants and harvester ants by examining stable isotope ratios in seeds and ants along the deposition gradient in the Sonoran Desert. Harvester ant community response to seeds produced by plants along the gradient will then be investigated through ant nest density surveys and seed preference measurements. Introduction Airborne nitrogen has increased over the last several decades as car exhaust, power plant emissions, industrial emissions, agricultural field fertilization and feedlots have dramatically increased (Vitousek et. al. 1997 and Fenn et al. 2003b,). This nitrogen is carried in many forms, including nitric oxide (NO), nitrogen dioxide (NO2), ammonia (NH3), and nitric acid (HNO3), via air currents and deposited on the soil affecting biogeochemical processes (Rao et al. 2009). Nitrogen deposition occurs in either wet or dry form. The arid West, including the mountains and deserts east of Los Angeles, experiences primarily dry deposition (Fenn et al. 2003b). Not only do emissions pollute air reducing air quality, but they also pollute the soil by increasing soil nitrogen levels (Galloway 1998, Hooper and Johnson 1999, and Krupa 2003). Atmospheric nitrogen deposition can have dramatic effects on soil and plant communities in various ecosystems (Sirulnik et al. 2007a and Sirulnik et al. 2007b). After water, nitrogen is typically the most limiting nutrient in arid systems (Burke 1989 and Hooper and Johnson 1999). Excess nitrogen in soil reduces the carbon to nitrogen ratio, which allows soil microbes to expand populations (Nave et al. 2009). Following excess nitrogen depletions, those larger microbial populations take up the majority of the nitrogen in the system and leave little for plants before dying. These cyclical swings of nitrogen availability can cause weedy plant invasions in desert systems, altering fire regimes and plant community composition (D?Antonio and Vitousek 1992, Evans et al. 2001, Brooks 2003, Schwinning et al. 2005). Many national parks and other government agencies are already monitoring atmospheric deposition, including NO, NO2, NH3, and HNH3 (Fenn et al. 2003b and Porter and Johnson 2007). Regions close to and downwind from high-density population centers and confined animal feeding operations are especially vulnerable to nitrogen deposition; thus, those areas are heavily monitored (Rao et al. 2009, Fenn et al. 2003a, and Junknys et al. 2007). The Park Service is concerned with a loss of biodiversity due to plant community shifts in part caused by increased nitrogen (Phoenix et al. 2006, and Fischlin et al. 2007, Porter and Johnson 2007). The effects of nitrogen deposition on plants and insects are interwoven with elevated carbon dioxide (CO2) and altered precipitation due to global climate change (Ellsworth et al. 2004; Fischlin et al. 2007). In some instances elevated CO2 may cause net primary productivity to increase; however, with a predicted change to more summer rain, primary productivity in the Mojave Desert may be more affected by precipitation rather than CO2 (Ellsworth et al. 2004). In addition, Newingham et al. (unpublished) found summer monsoons had more of an effect on insect abundance in the Mojave than either CO2 or nitrogen. The majority of studies suggest that plants will increase production (Drake et al. 1997; Jablonski et al. 2002) with predicted rising CO2 levels (Meehl et al. 2007); however, the effects of nitrogen deposition are more varied and not as well documented. A synthesis by Bobbink et al. (2010) found that nitrogen is more critical in determining community composition than previously realized. Understanding the interactions between elevated CO2 and nitrogen deposition is key in determining the carbon to nitrogen ratio (C:N) in plants, which affects herbivory, granivory, and detritivory. Higher carbon concentrations in leaf tissue and seeds provide more energy to herbivores and granivores, while higher nitrogen levels increase the capability to produce amino acids. As nitrogen is more limiting in most systems, plants producing more seed or seed with low C:N ratios could cause an increase in granivore populations, including harvester ants (Throop and Lerdau 2004). Differences in seed production, size, and nutrient content could have dramatic effects on granivores (Kelrick et al. 1986). Harvester ants along with birds and rodents move, store, and consume mass quantities of seed (MacMahon et al. 2000; White and Robertson 2009). Granivory and movement of seeds can deplete plant populations and/or shift species composition (MacMahon et al. 2000; White and Robertson 2010). However, little is known about seed production and subsequent predation under elevated nitrogen and how seed movement may affect ecosystem dynamics (Lewis and Gripenberg 2008). This study examines the connection between plants, seeds, and ants under different nitrogen pollution regimes. Study Objectives, Questions, and Predictions This project will examine the response of Messor pergandei to the effects of atmospheric nitrogen deposition in Joshua Tree National Park and surrounding desert in California. We will measure harvester ant nest densities and seed preferences along an established atmospheric nitrogen deposition gradient in the region. Study Questions Does atmospheric nitrogen deposition affect Messor pergandei harvester ants? Harvester ant distributions and nest characteristics 1. Are harvester ant nest densities greater in areas with higher atmospheric N deposition? Prediction ? Messsor pergandei nest densities will increase in high N deposition sites. 2. Do harvester ant nest characteristics vary along the depositional gradient? Prediction ? Changes in nest characteristics will occur between sites with high and low N deposition. Harvester ant ecological stoichiometry 3. Does increased N from atmospheric deposition alter C:N ratios in seeds and ants? Prediction ? C:N ratios will be lower in seeds and ants from high N deposition sites. Harvester ant food quality 4. Are harvester ants directly benefitting by consuming higher quality seeds created by atmospheric N deposition? Prediction ? Ants will have preferential selection for high quality (=high N) seed in low deposition sites. In high deposition sites where N seed content is overall high among spcecies, ants have less specific selection. (note ? This prediction is problematic as I am not necessarily investigating the process of seed selection but rather how the ants balance their stoichiometry. Advice on this one would be helpful and appreciated) Study Sites Sampling will be conducted in vegetation communities dominated by Larrea tridentata and Ambrosia dumosa. This vegetation type was chosen because of its vast land coverage; Larrea tridentata occurs in all the warm deserts in southwestern United States, as well into Mexico. The majority of Joshua Tree National Park and the Coachella Valley in California are covered by this vegetation community. Messor pergandei are the dominate harvester ant species in this region. Allen and co-workers detected an atmospheric nitrogen deposition gradient in southern California caused by air pollution from the Los Angeles Basin, Interstate 10 corridor, and the Inland Empire (Allen et al. 2009). The gradient stretches from the eastern slopes of the San Bernardino Mountains through Joshua Tree National Park (JOTR) to the eastern boundary of the park (Allen et al. 2009 and Rao et al. 2009). Allen and co-workers established 16 study plots (two outside of the park and 14 inside the park) in 2004. Many of these plots lie within Larrea tridentata and Ambrosia dumosa dominated vegetation communities. In collaboration with researchers at UC Riverside 15 sites (a combination of some of the Allen et al. sites and new sites) are being monitored for atmospheric nitrogen deposition. Based on prior site inspection, we will inventory eight of the monitored sites within the park and two sites in the Coachella Valley Preserve (CVP). More sites may be added in summer 2011. They are as follows (Figure 1): High Deposition Sites - TNC 1 (0563991 E, 3746465 N) North CVP Willis Palms (0562282 E, 3742667 N) South CVP Fan Canyon (0570020 E, 3749990 N) Southwest JOTR Mid Deposition Sites ? Cottonwood (0608773 E, 3735329 N) South Central JOTR Pinto Wash (0614916 E, 3744165 N) South Central JOTR Magic Circle (0607650 E, 3752092 N) Central JOTR (not on map) 29 Palms (0588686 E, 3771090 N) North Central JOTR Low Deposition Sites ? Cadiz Valley (0645792 E, 3773238 N) Northeast JOTR (not on map) Cox Cell (0654028 E, 3770708 N) Northeast JOTR Aqueduct (0659744 E, 3763225 N) East JOTR Figure 1. Atmospheric HNO3 Concentration gradient across the Coachella Valley and Joshua Tree National Park region of southern California. Concentrations were measured with passive samplers at each location from August 26 to September 23, 2010. These concentrations are multiplied by the depositional velocities from the EPA?s CASTNET monitoring system in the north end of the park to determine nitrogen deposition. Note: Magic Circle is in between Cottonwood and Pinto Basin and Cadiz is to the west of Cox Cell ? both not marked on map. Whitewater and Snow Creek will be sampled in 2011 and CVARS and Salton Sea will not be inventoried. (From Mike Bell UC Riverside) Study Design Atmospheric Deposition Passive samplers will be used to measure atmospheric nitrogen deposition (Rao et al. 2009 and Shen et al. 2009) along the atmospheric nitrogen deposition gradient as described in Allen et al. 2009 and Rao et al. 2009. Atmospheric nitric acid (HNO3) will be collected via nylon filters (Bytnerowicz et al. 2005). Nitrogen deposition will be measured for six week periods in the winter and summer to compare wet deposition during rainy winter months and dry deposition in the arid summer months (Rao et al. 2009). Following sample processing, the inferential method (multiplying deposition velocities by the atmospheric concentration measured) will be used to determine an estimate for the annual nitrogen deposition (Rao et al. 2009 and Shen et al. 2009). The Environmental Protection Agency (EPA) operates a Clean Air Status and Trends Network monitoring station in the Park and deposition velocities will be obtained from its published data on the EPA website (Rao et al. 2009). Atmospheric deposition sampling and analysis will be conducted in conjunction with researchers at UC Riverside. Harvester Ant Distributions and Nest Characteristics Q1. Are harvester ant nest densities greater in areas with higher atmospheric N deposition? Q2. Do harvester ant nest characteristics vary along the depositional gradient? In the aforementioned plots, harvester ant nest density will be determined. All ant mounds will be counted in the half hectare plots. At each nest encountered, the species of harvester ant will be identified and the size of the cleared disk surrounding the nest entry measured. A small sample of ants will be collected to verify identification of species and to determine the isotopic signature. The distance from each nest to the nearest shrub (canopy and base) will be measured to ascertain site selection. Ants could be building nests under or near shrubs to take advantage of the islands of fertility surrounding the shrubs. The number of abandoned nests/entrances surrounding each active nest will be counted to represent ant colony activity. In addition, at each nest soil samples will be collected to measure the following soil parameters: pH, total carbon, and total nitrogen. As nitrogen deposition can lead to acidification of soils (Vitousek et al. 1997), pH will be measured to determine if ants are choosing nest sites within a certain pH range. Harvester Ant Ecological Stoichiometry Q3. Does increased N from atmospheric deposition lower C:N ratios in seeds and ants? Q4. Are harvester ants directly benefitting by consuming higher quality seeds created by atmospheric N deposition? Pollution, seeds, and ants will all be analyzed for their specific nitrogen isotopic signature to determine quantitatively if the pollution is directly affecting trophic levels in the park. Each successive step in the chain will have a particular concentration of the δ15 N/ δ14 N ratios, which would reveal how much of the nitrogen pollution is being concentrated within the plant and harvester ant communities (Ehleringer and Rundel 1989). In addition, recent work has furthered the use of stable isotopes ratios to determine sources of nitrogen pollution (Yeatman et al. 2001, Li et al. 2007, Toyoda et al. 2008, and Zhao et al. 2009); therefore, it might be possible to categorize the sources of nitrogen deposition in the park. A subsample of the nitrogen deposition collected with the passive samplers will analyzed for δ15 N/ δ14 N ratio enrichment at the University of California Riverside. To determine if the plants are utilizing the nitrogen from pollution, selected seeds will also be examined for δ15 N enrichment. Finally, harvester ants from the plots where the seeds were sampled from will also be analyzed for their stable isotopic signature. Except for the nitrogen deposition, all stable isotope analysis will be done at the University of Idaho Stable Isotopes Laboratory. This line of analysis will explore the pathways the nitrogen pollution takes following deposition in the park and link effects of pollution on the atmosphere to flora and fauna. Seed Collection and Analysis Q4. Are harvester ants directly benefitting by consuming higher quality seeds created by atmospheric N deposition? Seeds will be collected from selected native and invasive plant species along the atmospheric nitrogen deposition gradient as described in Allen et al. 2009 and Rao et al. 2009. The study plots as detailed above are one-half hectare in dimension (Rao et al. 2009). Following harvesting, the seeds will be analyzed for total carbon and nitrogen to determine carbon to nitrogen ratios in seeds along the gradient. At selected sites along the gradient, seeds will be collected from ants returning to their nests. Harvester ants follow trunk trails that stretch from the nest to various seed sources (H?lldobler and Wilson 1970). By removing the seeds these ants carry, a picture of what they are consuming will be painted. This measurement will also determine if the ants are harvesting seeds from native or non-native plant species. These seeds will be identified to genus or species. Summary of parameters that will be measured ? Messor pergandei nest densities at 10 sites along an atmospheric N deposition gradient (0.5 hectare plots) Q1 ? Distance from each nest to the nearest shrub (canopy and base) Q2 ? Small sample of ants from all nests counted (for identification) Q1 ? Large sample of ants from 3 nests at each of the 10 sites (for identification, %C and %N, and stable isotope analysis) Q1,Q3,Q4 ? Number of abandoned nests/entrances surrounding each active nest Q2 ? Seeds from LATR, AMDU, and selected annuals (%C and %N and stable isotope analysis) Q3,Q4 ? Leaves/small branch from LATR from almost each site (%C and %N and stable isotope analysis) Q3,Q4 ? Soil samples from the three nests where a large ant sample was collected (%C and %N and stable isotope analysis) Q1,Q3,Q4 ? Seeds taken from ants returning to the three nests where a large ant sample was collected (seeds plucked with forceps for 20 minutes at each nest) Q3,Q4 Data Analysis All parameters measured will be analyzed with ANOVAs to determine differences among sites. Then each parameter will be correlated with the atmospheric N deposition gradient via regression analysis to establish patterns of harvester ant response to elevated nitrogen. Stable isotope ratios from atmospheric, soil, plant, seed, and ant samples will be modeled for connections. Conclusion Air pollution from the Los Angeles Basin, blowing over the San Bernardino Mountains, is affecting ecological processes in fragile desert ecosystems. By combining previous work on atmospheric nitrogen deposition gradients with new research on seed characteristics and harvester ant responses, I will attempt to link global change pollution with insect ecology. This study will be one part of the increasing knowledge on insect responses to global change, as the interactions of increased CO2, nitrogen deposition, and changes in precipitation will all likely play a role in the future of desert systems. Literature Cited Allen, E.B., L.E. Rao, R.J. Steers, A. Bytnerowicz, and M.E. Fenn. 2009. Impacts of atmospheric nitrogen deposition on vegetation and soils in Joshua Tree National Park. In: Webb, R.H., L.F. Fenstermaker, J.S. Heaton, D.L. Hughson, E.V. McDonald, and D.M. Miller (Eds.). The Mojave Desert: Ecosystem Processes and Sustainability. University of Nevada Press, Las Vegas, USA. Bobbink, R., K. Hicks, J. Galloway, T. Spranger, R. Alkemade, M. Ashmore, M. Bustamante, S. Cinderby, E. Davidson, F. Dentener, B. Emmett, J.-W. 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Atmospheric Environment 35:1307-1320. Zhao, X., X. Yan, Z. Xiong, Y. Xie, G. Xing, S. Shi, and Z. Zhu. 2009. Spatial and temporal variation of inorganic nitrogen wet deposition to the Yangtze River Delta Region, China. Water, Air, and Soil Pollution 203:277-289.

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Mojave Desert Harvester Ant Response to Atmospheric Nitrogen Deposition

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Benjamin Wissinger Jun 1 - 17, 2011 (17 days)
Benjamin Wissinger Jun 1 - 17, 2011 (17 days)

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