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There is increasing evidence that ovarian status influcences behavioral phenotype in workers of the honey bee Apis mellifera. Honey bee workers demonstrate a complex division of labor. Young workers perform in-hive tasks (e.g. brood care), while older bees perform outside tasks (e.g. foraging for food). This age correlated division of

There is increasing evidence that ovarian status influcences behavioral phenotype in workers of the honey bee Apis mellifera. Honey bee workers demonstrate a complex division of labor. Young workers perform in-hive tasks (e.g. brood care), while older bees perform outside tasks (e.g. foraging for food). This age correlated division of labor is known as temporal polyethism. Foragers demonstrate further division of labor with some bees biasing collection towards protein (pollen) and others towards carbohydrates (nectar). The Reproductive Ground-plan Hypothesis proposes that the ovary plays a regulatory role in foraging division of labor. European honey bee workers that have been selectively bred to store larger amounts of pollen (High strain) also have a higher number of ovarioles per ovary than workers from strains bred to store less pollen (Low strain). High strain bees also initiate foraging earlier than Low strain bees. The relationship between ovariole number and foraging behavior is also observed in wild-type Apis mellifera and Apis cerana: pollen-biased foragers have more ovarioles than nectar-biased foragers. In my first study, I investigated the pre-foraging behavioral patterns of the High and Low strain bees. I found that High strain bees progress through the temporal polyethism at a faster rate than Low strain bees. To ensure that the observed relationship between the ovary and foraging bias is not due to associated separate genes for ovary size and foraging behavior, I investigated foraging behavior of African-European backcross bees. The backcross breeding program was designed to break potential gene associations. The results from this study demonstrated the relationship between the ovary and foraging behavior, supporting the proposed causal linkage between reproductive development and behavioral phenotype. The final study was designed to elucidate a regulatory mechanism that links ovariole number with sucrose sensitivity, and loading decisions. I measured ovariole number, sucrose sensitivity and sucrose solution load size using a rate-controlled sucrose delivery system. I found an interaction effect between ovariole number and sucrose sensitivity for sucrose solution load size. This suggests that the ovary impacts carbohydrate collection through modulation of sucrose sensitivity. Because nectar and pollen collection are not independent, this would also impact protein collection.
ContributorsSiegel, Adam J (Author) / Page, Jr., Robert E (Thesis advisor) / Hamilton, Andrew L. (Committee member) / Brent, Colin S (Committee member) / Amdam, Gro V (Committee member) / McGraw, Kevin J. (Committee member) / Arizona State University (Publisher)
Created2011
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Description
Aboveground net primary production (ANPP) and belowground net primary production (BNPP) may not be influenced equally by the same factors in arid grasslands. Precipitation is known to affect ANPP and BNPP, while soil fauna such as nematodes affect the BNPP through herbivory and predation. This study on black grama grass

Aboveground net primary production (ANPP) and belowground net primary production (BNPP) may not be influenced equally by the same factors in arid grasslands. Precipitation is known to affect ANPP and BNPP, while soil fauna such as nematodes affect the BNPP through herbivory and predation. This study on black grama grass (Bouteloua eriopoda) in the Chihuahuan Desert investigates the effects of precipitation and nematode presence or absence on net primary production (NPP) as well as the partitioning between the aboveground and belowground components, in this case, the fraction of total net primary production occurring belowground (fBNPP). I used a factorial experiment to investigate the effects of both precipitation and nematode presence on the components of NPP. I used rainout shelters and an irrigation system to alter precipitation totals, while I used defaunated and re-inoculated soil for the nematode treatments. Precipitation treatment and seasonal soil moisture had no effect on the BNPP and a nonsignificant positive effect on the ANPP. The fBNPP decreased with increasing precipitation and seasonal soil moisture, though without a significant effect. No predator nematodes were found in any of the microcosms at the end of the experiment, though other functional groups of nematodes, including herbivores, were found in the microcosms. Total nematode numbers did not vary significantly between nematode treatments, indicating that the inoculation process did not last for the whole experiment or that nematodes had little plant material to eat and resulted in low population density. Nematode presence did not affect the BNPP, ANPP, or the fBNPP. There were no significant interactions between precipitation and nematode treatment. The results are inconclusive, possibly as a result of ecosystem trends during an unusually high precipitation year, as well as the very low NPP values in the experiment that correlated with low nematode community numbers.
ContributorsWiedenfeld, Amy (Author) / Sala, Osvaldo (Thesis advisor) / Gerber, Leah (Committee member) / Hall, Sharon (Committee member) / Arizona State University (Publisher)
Created2018
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Description
Rangelands are an extensive land cover type that cover about 40% of earth’s ice-free surface, expanding into many biomes. Moreover, managing rangelands is crucial for long-term sustainability of the vital ecosystem services they provide including carbon (C) storage via soil organic carbon (SOC) and animal agriculture. Arid rangelands are particularly

Rangelands are an extensive land cover type that cover about 40% of earth’s ice-free surface, expanding into many biomes. Moreover, managing rangelands is crucial for long-term sustainability of the vital ecosystem services they provide including carbon (C) storage via soil organic carbon (SOC) and animal agriculture. Arid rangelands are particularly susceptible to dramatic shifts in vegetation cover, physical and chemical soil properties, and erosion due to grazing pressure. Many studies have documented these effects, but studies focusing on grazing impacts on soil properties, namely SOC, are less common. Furthermore, studies testing effects of different levels of grazing intensities on SOC pools and distribution yield mixed results with little alignment. The primary objective of this thesis was to have a better understanding of the role of grazing intensity on arid rangeland soil C storage. I conducted research in long established pastures in Jornada Experimental Range (JER). I established a 1500m transect in three pastures originating at water points and analyzed vegetation cover and SOC on points along these transects to see the effect of grazing on C storage on a grazing gradient. I used the line-point intercept method to measure and categorize vegetation into grass, bare, and shrub. Since soil adjacent to each of these three cover types will likely contain differing SOC content, I then used this vegetation cover data to calculate the contribution of each cover type to SOC. I found shrub cover and total vegetation cover to decrease, while grass and bare cover increased with decreasing proximity to the water source. I found areal (g/m2) and percent (go SOC to be highest in the first 200m of the transects when accounting for the contribution of the three vegetation cover types. I concluded that SOC is being redistributed toward the water source via foraging and defecation and foraging, due to a negative trend of both total vegetation cover and percent SOC (g/g). With the decreasing trends of vegetation cover and SOC further from pasture water sources, my thesis research contributes to the understanding of storage and distribution of SOC stocks in arid rangelands.
ContributorsBoydston, Aaron (Author) / Sala, Osvaldo (Thesis advisor) / Throop, Heather (Committee member) / Hall, Sharon (Committee member) / Arizona State University (Publisher)
Created2018
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Description
Water is the main driver of net primary productivity (NPP) in arid ecosystems, followed by nitrogen and phosphorous. Precipitation is the primary factor in determining water availability to plants, but other factors such as surface rocks could also have an impact. Surface rocks may positively affect water availability by preventing

Water is the main driver of net primary productivity (NPP) in arid ecosystems, followed by nitrogen and phosphorous. Precipitation is the primary factor in determining water availability to plants, but other factors such as surface rocks could also have an impact. Surface rocks may positively affect water availability by preventing evaporation from soil, but at higher densities, surface rocks may also have a negative impact on water availability by limiting water infiltration or light availability. However, the direct relationship between rock cover and aboveground net primary productivity (ANPP), a proxy for NPP, is not well understood. In this research we explore the relationship between rock cover, ANPP, and soil nutrient availability. We conducted a rock cover survey on long-term fertilized plots at fifteen sites in the Sonoran Desert and used 4 years of data from annual plant biomass surveys to determine the relationship between peak plant biomass and surface rock cover. We performed factorial ANCOVA to assess the relationship among annual plant biomass, surface rocks, precipitation, and fertilization treatment. Overall we found that precipitation, nutrients, and rock cover influence growth of Sonoran Desert annual plants. Rock cover had an overall negative relationship with annual plant biomass, but did not show a consistent pattern of significance over four years of study and with varying average winter precipitation.
ContributorsShaw, Julea Anne (Author) / Hall, Sharon (Thesis director) / Sala, Osvaldo (Committee member) / Cook, Elizabeth (Committee member) / Barrett, The Honors College (Contributor) / School of Life Sciences (Contributor)
Created2015-05
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Description
Hydrological models in arid and semi-arid ecosystems can be subject to high uncertainties. Spatial variability in soil moisture and evapotranspiration, key components of the water cycle, can contribute to model uncertainty. In particular, an understudied source of spatial variation is the effect of plant-plant interactions on water fluxes. At patch

Hydrological models in arid and semi-arid ecosystems can be subject to high uncertainties. Spatial variability in soil moisture and evapotranspiration, key components of the water cycle, can contribute to model uncertainty. In particular, an understudied source of spatial variation is the effect of plant-plant interactions on water fluxes. At patch scales (plant and associated soil), plant neighbors can either negatively or positively affect soil water availability via competition or hydraulic redistribution, respectively. The aboveground microclimate can also be altered via canopy shading effects by neighbors. Across longer timescales (years), plants may adjust their physiological (water-use) traits in response to the neighbor-altered microclimate, which subsequently affects transpiration rates. The influence of physiological adjustments and neighbor-altered microclimate on water fluxes was assessed around Larrea tridentata in the Sonoran Desert. Field measurements of Larrea’s stomatal behavior and vertical root distributions were used to examine the effects of neighbors on Larrea’s physiological controls on transpiration. A modeling based approach was implemented to explore the sensitivity of evapotranspiration and soil moisture to neighbor effects. Neighbors significantly altered both above- and belowground physiological controls on evapotranspiration. Compared to Larrea growing alone, neighbors increased Larrea’s annual transpiration by up to 75% and 30% at the patch and stand scales, respectively. Estimates of annual transpiration were highly sensitive to the presence/absence of competition for water, and on seasonal timescales, physiological adjustments significantly influenced transpiration estimates. Plant-plant interactions can be a significant source of spatial variation in ecohydrological models, and both physiological adjustments to neighbors and neighbor effects on microclimate affect small scale (patch to ecosystem) water fluxes.
ContributorsKropp, Heather (Author) / Ogle, Kiona (Thesis advisor) / Hultine, Kevin (Committee member) / Sala, Osvaldo (Committee member) / Vivoni, Enrique (Committee member) / Wojciechowski, Martin (Committee member) / Arizona State University (Publisher)
Created2015
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Description
Drylands cover over 40% of the Earth’s surface, account for one third of global carbon cycling, and are hotspots for climate change, with more frequent and severe droughts coupled with deluges of novel magnitude and frequency. Because of their large terrestrial extent, elucidating dryland ecosystem responses to changes in water

Drylands cover over 40% of the Earth’s surface, account for one third of global carbon cycling, and are hotspots for climate change, with more frequent and severe droughts coupled with deluges of novel magnitude and frequency. Because of their large terrestrial extent, elucidating dryland ecosystem responses to changes in water availability is critical for a comprehensive understanding of controls on global aboveground net primary productivity (ANPP), an important ecosystem service. The focus of this dissertation is to investigate cause-effect mechanisms between altered water availability and ecosystem processes in dryland ecosystems. Across a network of experimental rainfall manipulations within a semiarid Chihuahuan Desert grassland, I examined short- and long-term dynamics of multiple ecosystem processes—from plant phenology to nitrogen cycling—in response to directional precipitation extremes. Aboveground, I found herbaceous plant phenology to be more sensitive in greenup timing compared to deep-rooted, woody shrubs, implying that precipitation extremes will disproportionately affect grass-dominated compared to woody ecosystems. Surprisingly, after 14 years of experimentally adding water and N, I observed no effect on ANPP. Belowground, bulk soil N dynamics remained stable with differing precipitation amounts. However, mineral associated organic N (MAOM-N) significantly increased under chronic N inputs, indicating potential for dryland soil N sequestration. Conversely, the difference between low- and high-N soil N content may increase a drawdown of N from all soil N pools under low-N conditions whereas plants source N from fertilizer input under high-N conditions. Finally, I considered ecosystem-level acclimation to climate change. I found that N availability decreased with annual precipitation in space across continents, but it posed initially increasing trends in response to rainfall extremes at the Jornada that decreased after 14 years. Mechanisms for the acclimation process are thus likely associated with differential lags to changes in precipitation between plants and microorganisms. Overall, my dissertation demonstrates that examining linkages between multiple ecosystem processes, from aboveground phenological cycles to belowground N cycling dynamics, can provide a more integrative understanding of dryland response to climate change. Because dryland range is potentially expanding globally, water limited systems provide a unique and critical focus area for future research that revisit and revise current ecological paradigms.
ContributorsCurrier, Courtney (Author) / Sala, Osvaldo (Thesis advisor) / Collins, Scott (Committee member) / Reed, Sasha (Committee member) / Throop, Heather (Committee member) / Arizona State University (Publisher)
Created2023
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Climate change is increasing global surface temperatures, intensifying droughts and increasing rainfall variation, particularly in drylands. Understanding how dryland plant communities respond to climate change-induced rainfall changes is crucial for implementing effective conservation strategies. Concurrent with climate change impacts on drylands is woody encroachment: an increase in abundance of woody

Climate change is increasing global surface temperatures, intensifying droughts and increasing rainfall variation, particularly in drylands. Understanding how dryland plant communities respond to climate change-induced rainfall changes is crucial for implementing effective conservation strategies. Concurrent with climate change impacts on drylands is woody encroachment: an increase in abundance of woody plant species in areas formerly dominated by grasslands or savannahs. For example, the woody plant, Prosopis velutina (velvet mesquite), has encroached into grasslands regionally over the past century. From an agricultural perspective, P. velutina is an invasive weed that hinders cattle forage. Understanding how P. velutina will respond to climate change-induced rainfall changes can be useful for management and conservation efforts. Prosopis velutina was used to answer the following question: Is there a significant interactive effect of mean soil water moisture content and pulse frequency on woody seedling survival and growth in dryland ecosystems? There were 256 P. velutina seedlings sourced from the Santa Rita Experimental Range in southern Arizona grown under four watering treatments where mean and pulse frequency were manipulated over two months. Data were collected on mortality, stem height, number of leaves, instantaneous gas exchange, chlorophyll fluorescence, biomass, and the leaf carbon to nitrogen (C:N) ratio. Mortality was low across treatments. Pulse frequency had less impact across response variables than the mean amount of water received. This may indicate that P. velutina seedlings are relatively insensitive to rainfall timing and are more responsive to rainfall amount. Prosopis velutina in the low mean soil moisture treatments lost a majority of their leaves and had greater biomass allocation to roots. Prosopis velutina’s ability to survive in low soil moisture conditions and invest in root biomass can allow it to persist as drylands are further affected by climate change. Prosopis velutina could benefit ecosystems where native plants are at risk due to rainfall variation if P. velutina occupies a similar niche space. Due to conflicting viewpoints of P. velutina as an invasive species, it’s important to examine P. velutina from both agricultural and conservation perspectives. Further analysis on the benefits to P. velutina in these ecosystems is recommended.
ContributorsDavis, Ashley R. (Author) / Throop, Heather (Thesis advisor) / Hultine, Kevin (Committee member) / Sala, Osvaldo (Committee member) / Arizona State University (Publisher)
Created2020
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Description
Desert organisms lead harsh lives owing to the extreme, often unpredictable environmental conditions they endure. Climate change will likely make their existence even harsher. Predicting the ecological consequences of future climate scenarios thus requires understanding how the biota will be affected by climatic shifts. Biological soil crusts (biocrusts) are

Desert organisms lead harsh lives owing to the extreme, often unpredictable environmental conditions they endure. Climate change will likely make their existence even harsher. Predicting the ecological consequences of future climate scenarios thus requires understanding how the biota will be affected by climatic shifts. Biological soil crusts (biocrusts) are an important ecosystem component in arid lands, one that covers large portions of the landscape, improving soil stability and fertility. Because cyanobacteria are biocrust’s preeminent primary producers, eking out an existence during short pulses of precipitation, they represent a relevant global change object of study. I assessed how climate scenarios predicted for the Southwestern United States (US) will affect biocrusts using long-term, rainfall-modifying experimental set-ups that imposed either more intense drought, a seasonally delayed monsoon season, or a shift to smaller but more frequent precipitation events. I expected drought to be detrimental, but not a delay in the monsoon season. Surprisingly, both treatments showed similar effects on cyanobacterial community composition and population size after four years. While successionally incipient biocrusts were unaffected, mature biocrusts lost biomass and diversity with treatment, especially among nitrogen-fixing cyanobacteria. In separate experiments, I assessed the effect of rainfall with modified event size and frequency after a decade of treatment. Small, frequent rainfall events surprisingly enhanced the diversity and biomass of bacteria and cyanobacteria, with clear winners and losers: nitrogen-fixing Scytonema sp. benefited, while Microcoleus vaginatus lost its dominance. As an additional finding, I could also show that water addition is not always beneficial to biocrusts, calling into question the notion that these are strictly water-limited systems.

Finally, results interpretation was severely hampered by a lack of appropriate systematic treatment for an important group of biocrust cyanobacteria, the “Microcoleus steenstrupii complex”. I characterized the complex using a polyphasic approach, leading to the formal description of a new family (Porphyrosiphonaceae) of desiccation resistant cyanobacteria that includes 11 genera, of which 5 had to be newly described. Under the new framework, the distribution and abundance of biocrust cyanobacteria with respect to environmental conditions can now be understood. This body of work contributes significantly to explain current distributional patterns of biocrust cyanobacteria and to predict their fate in the face of climate change.
ContributorsMoreira Camara Fernandes, Vanessa (Author) / Garcia-Pichel, Ferran (Thesis advisor) / Rudgers, Jennifer (Committee member) / Sala, Osvaldo (Committee member) / Penton, Christopher (Committee member) / Arizona State University (Publisher)
Created2020
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Description
In many social groups, reproduction is shared between group members, whocompete for position in the social hierarchy for reproductive dominance. This reproductive conflict can lead to different means of enforcing reproductive differences, such as dominance displays or limited control of social hierarchy through antagonistic encounters. In eusocial insects, archetypal colonies contain a single,

In many social groups, reproduction is shared between group members, whocompete for position in the social hierarchy for reproductive dominance. This reproductive conflict can lead to different means of enforcing reproductive differences, such as dominance displays or limited control of social hierarchy through antagonistic encounters. In eusocial insects, archetypal colonies contain a single, singly-mated fertile queen, such that no reproductive conflict exists within a colony. However, many eusocial insects deviate from this archetype and have multiply-mated queens (polyandry), multiple queens in a single colony (polygyny), or both. In these cases, reproductive conflict exists between the matrilines and patrilines represented in a colony, specifically over the production of sexual offspring. A possible outcome of reproductive conflict may be the emergence of cheating lineages, which favor the production of sexual offspring, taking advantage of the worker force produced by nestmate queens and/or patrilines. In extreme examples, inquiline social parasites may be an evolutionary consequence of reproductive conflict between nestmate queens. Inquiline social parasitism is a type of social parasitism that is usually defined by a partial or total loss of the worker caste, and the “infiltration” of host colonies to take advantage of the host worker force for reproduction. It has been hypothesized that these inquiline social parasites evolve through the speciation of cheating queen lineages from within their incipient host species. This “intra- specific” origin model involves a foundational hypothesis that the common ancestor of host and parasite (and thus, putatively, the host at the time of speciation) should be functionally polygynous, and that parasitism evolves as a “resolution” of reproductive conflict in colonies. In this dissertation, I investigate the hypothesized role of polygyny in the evolution of inquiline social parasites. I use molecular ecology and statistical approaches to validate the role of polygyny in the evolution of some inquiline social parasites. I further discuss potential mechanisms for the evolution and speciation of social parasites, and discuss future directions to elucidate these mechanisms.
ContributorsDahan, Romain Arvid (Author) / Rabeling, Christian (Thesis advisor) / Amdam, Gro V (Committee member) / Fewell, Jennifer H (Committee member) / Pratt, Stephen C (Committee member) / Rüppell, Olav (Committee member) / Arizona State University (Publisher)
Created2021