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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
Climate change has the potential to affect vegetation via changes in temperature and precipitation. In the semi-arid southwestern United States, heightened temperatures will likely lead to accelerated groundwater pumping to meet human needs, and altered storm patterns may lead to changes in flood regimes. All of these hydrologic changes have

Climate change has the potential to affect vegetation via changes in temperature and precipitation. In the semi-arid southwestern United States, heightened temperatures will likely lead to accelerated groundwater pumping to meet human needs, and altered storm patterns may lead to changes in flood regimes. All of these hydrologic changes have the potential to alter riparian vegetation. This research, consisting of two papers, examines relationships between hydrology and riparian vegetation along the Verde River in central Arizona, from applied and theoretical perspectives. One paper investigates how dominance of tree and shrub species and cover of certain functional groups change along hydrologic gradients. The other paper uses the Verde River flora along with that river's flood and moisture gradients to answer the question of whether functional groups can be defined universally. Drying of the Verde River would lead to a shift from cottonwood-willow streamside forest to more drought adapted desert willow or saltcedar, a decline in streamside marsh species, and decreased species richness. Effects drying will have on one dominant forest tree, velvet ash, is unclear. Increase in the frequency of large floods would potentially increase forest density and decrease average tree age and diameter. Correlations between functional traits of Verde River plants and hydrologic gradients are consistent with "leaf economics," or the axis of resource capture, use, and release, as the primary strategic trade-off for plants. This corresponds to the competitor-stress tolerator gradient in Grime's life history strategy theory. Plant height was also a strong indicator of hydrologic condition, though it is not clear from the literature if plant height is independent enough of leaf characteristics on a global scale to be considered a second axis. Though the ecohydrologic relationships are approached from different perspectives, the results of the two papers are consistent if interpreted together. The species that are currently dominant in the near-channel Verde River floodplain are tall, broad-leaf trees, and the species that are predicted to become more dominant in the case of the river drying are shorter trees or shrubs with smaller leaves. These results have implications for river and water management, as well as theoretical ecology.
ContributorsHazelton, Andrea Florence (Author) / Stromberg, Juliet C. (Thesis advisor) / Schmeeckle, Mark W (Committee member) / Franklin, Janet (Committee member) / Arizona State University (Publisher)
Created2011
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Description
Over the past century in the southwestern United States human actions have altered hydrological processes that shape riparian ecosystems. One change, release of treated wastewater into waterways, has created perennial base flows and increased nutrient availability in ephemeral or intermittent channels. While there are benefits to utilizing treated wastewater for

Over the past century in the southwestern United States human actions have altered hydrological processes that shape riparian ecosystems. One change, release of treated wastewater into waterways, has created perennial base flows and increased nutrient availability in ephemeral or intermittent channels. While there are benefits to utilizing treated wastewater for environmental flows, there are numerous unresolved ecohydrological issues regarding the efficacy of effluent to sustain groundwater-dependent riparian ecosystems. This research examined how nutrient-rich effluent, released into waterways with varying depths to groundwater, influences riparian plant community development. Statewide analysis of spatial and temporal patterns of effluent generation and release revealed that hydrogeomorphic setting significantly influences downstream riparian response. Approximately 70% of effluent released is into deep groundwater systems, which produced the lowest riparian development. A greenhouse study assessed how varying concentrations of nitrogen and phosphorus, emulating levels in effluent, influenced plant community response. With increasing nitrogen concentrations, vegetation emerging from riparian seed banks had greater biomass, reduced species richness, and greater abundance of nitrophilic species. The effluent-dominated Santa Cruz River in southern Arizona, with a shallow groundwater upper reach and deep groundwater lower reach, served as a study river while the San Pedro River provided a control. Analysis revealed that woody species richness and composition were similar between the two systems. Hydric pioneers (Populus fremontii, Salix gooddingii) were dominant at perennial sites on both rivers. Nitrophilic species (Conium maculatum, Polygonum lapathifolium) dominated herbaceous plant communities and plant heights were greatest in effluent-dominated reaches. Riparian vegetation declined with increasing downstream distance in the upper Santa Cruz, while patterns in the lower Santa Cruz were confounded by additional downstream agricultural input and a channelized floodplain. There were distinct longitudinal and lateral shifts toward more xeric species with increasing downstream distance and increasing lateral distance from the low-flow channel. Patterns in the upper and lower Santa Cruz reaches indicate that water availability drives riparian vegetation outcomes below treatment facilities. Ultimately, this research informs decision processes and increases adaptive capacity for water resources policy and management through the integration of ecological data in decision frameworks regarding the release of effluent for environmental flows.
ContributorsWhite, Margaret Susan (Author) / Stromberg, Juliet C. (Thesis advisor) / Fisher, Stuart G. (Committee member) / White, Dave (Committee member) / Holway, James (Committee member) / Wu, Jianguo (Committee member) / Arizona State University (Publisher)
Created2011
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A notable feature of advanced eusocial insect groups is a division of labor within the sterile worker caste. However, the physiological aspects underlying the differentiation of behavioral phenotypes are poorly understood in one of the most successful social taxa, the ants. By starting to understand the foundations on which social

A notable feature of advanced eusocial insect groups is a division of labor within the sterile worker caste. However, the physiological aspects underlying the differentiation of behavioral phenotypes are poorly understood in one of the most successful social taxa, the ants. By starting to understand the foundations on which social behaviors are built, it also becomes possible to better evaluate hypothetical explanations regarding the mechanisms behind the evolution of insect eusociality, such as the argument that the reproductive regulatory infrastructure of solitary ancestors was co-opted and modified to produce distinct castes. This dissertation provides new information regarding the internal factors that could underlie the division of labor observed in both founding queens and workers of Pogonomyrmex californicus ants, and shows that changes in task performance are correlated with differences in reproductive physiology in both castes. In queens and workers, foraging behavior is linked to elevated levels of the reproductively-associated juvenile hormone (JH), and, in workers, this behavioral change is accompanied by depressed levels of ecdysteroid hormones. In both castes, the transition to foraging is also associated with reduced ovarian activity. Further investigation shows that queens remain behaviorally plastic, even after worker emergence, but the association between JH and behavioral bias remains the same, suggesting that this hormone is an important component of behavioral development in these ants. In addition to these reproductive factors, treatment with an inhibitor of the nutrient-sensing pathway Target of Rapamycin (TOR) also causes queens to become biased towards foraging, suggesting an additional sensory component that could play an important role in division of labor. Overall, this work provides novel identification of the possible regulators behind ant division of labor, and suggests how reproductive physiology could play an important role in the evolution and regulation of non-reproductive social behaviors.
ContributorsDolezal, Adam G (Author) / Amdam, Gro V (Thesis advisor) / Brent, Colin S. (Committee member) / Gadau, Juergen (Committee member) / Hoelldobler, Bert (Committee member) / Liebig, Juergen (Committee member) / Arizona State University (Publisher)
Created2012
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Novel resource inputs represent an increasingly common phenomenon in ecological systems as global change alters environmental factors and species distributions. In semiarid riparian areas, hydric pioneer tree species are being replaced by drought-tolerant species as water availability decreases. Additionally, introduced omnivorous crayfish, which feed upon primary producers, allochthonous detritus, and

Novel resource inputs represent an increasingly common phenomenon in ecological systems as global change alters environmental factors and species distributions. In semiarid riparian areas, hydric pioneer tree species are being replaced by drought-tolerant species as water availability decreases. Additionally, introduced omnivorous crayfish, which feed upon primary producers, allochthonous detritus, and benthic invertebrates, can impact communities at multiple levels through both direct and indirect effects. In arid and semiarid systems of the American Southwest, crayfish may be especially important as detrital processors due to the lack of specialized detritivores. I tested the impact of virile crayfish (Orconectes virilis) on benthic invertebrates and detrital resources across a gradient of riparian vegetation drought-tolerance using field cages with leaf litter bags in the San Pedro River in Southeastern Arizona. Virile crayfish increased breakdown rate of drought-tolerant saltcedar (Tamarix ramosissima), but did not impact breakdown of Fremont cottonwood (Populus fremontii), Gooding's willow (Salix goodingii), or seepwillow (Baccharis salicifolia). The density and composition of the invertebrate community colonizing leaf litter bags were both heavily influenced by litter species but not directly by crayfish presence. As drought-tolerant species become more abundant in riparian zones, their litter will become a larger component of the organic matter budget of desert streams. By increasing breakdown rates of saltcedar, crayfish shift the composition of leaf litter in streams, which in turn may affect the composition and biomass of colonizing invertebrate communities. More research is needed to determine the full extent to which these alterations change community composition over time.
ContributorsMoody, Eric Kellan (Author) / Sabo, John L (Thesis advisor) / Collins, James P. (Committee member) / Stromberg, Juliet C. (Committee member) / Arizona State University (Publisher)
Created2012
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Description

River and riparian areas are important foraging habitat for insectivorous bats. Numerous studies have shown that aquatic insects provide an important trophic resource to terrestrial consumers, including bats, and are key in regulating population size and species interactions in terrestrial food webs. Yet these studies have generally ignored how structural

River and riparian areas are important foraging habitat for insectivorous bats. Numerous studies have shown that aquatic insects provide an important trophic resource to terrestrial consumers, including bats, and are key in regulating population size and species interactions in terrestrial food webs. Yet these studies have generally ignored how structural characteristics of the riverine landscape influence trophic resource availability or how terrestrial consumers respond to ensuing spatial and temporal patterns of trophic resources. Moreover, few studies have examined linkages between a stream's hydrologic regime and the timing and magnitude of aquatic insect availability. The main objective of my dissertation is to understand the causes of bat distributions in space and time. Specifically, I examine how trophic resource availability, structural components of riverine landscapes (channel confinement and riparian vegetation structure), and hydrologic regimes (flow permanence and timing of floods) mediate spatial and temporal patterns in bat activity. First, I show that river channel confinement determines bat activity along a river's longitudinal axis (directly above the river), while trophic resources appear to have stronger effects across a river's lateral (with distance from the river) axis. Second, I show that flow intermittency affects bat foraging activity indirectly via its effects on trophic resource availability. Seasonal river drying appears to have complex effects on bat foraging activity, initially causing imperfect tracking by consumers of localized concentrations of resources but later resulting in disappearance of both insects and bats after complete river drying. Third, I show that resource tracking by bats varies among streams with contrasting patterns of trophic resource availability and this variation appears to be in response to differences in the timing of aquatic insect emergence, duration and magnitude of emergence, and adult body size of emergent aquatic insects. Finally, I show that aquatic insects directly influence bat activity along a desert stream and that riparian vegetation composition affects bat activity, but only indirectly, via effects on aquatic insect availability. Overall, my results show river channel confinement, riparian vegetation structure, flow permanence, and the timing of floods influence spatial and temporal patterns in bat distributions; but these effects are indirect by influencing the ability of bats to track trophic resources in space and time.

ContributorsHagen, Elizabeth M (Author) / Sabo, John L (Thesis advisor) / Fisher, Stuart G. (Committee member) / Grimm, Nancy (Committee member) / Schmeeckle, Mark W (Committee member) / Stromberg, Juliet C. (Committee member) / Arizona State University (Publisher)
Created2010
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General ecological thought pertaining to plant biology, conservation, and urban areas has rested on two potentially contradictory underlying assumptions. The first is that non-native plants can spread easily from human developments to “pristine” areas. The second is that native plants cannot disperse through developed areas. Both assume anthropogenic changes to

General ecological thought pertaining to plant biology, conservation, and urban areas has rested on two potentially contradictory underlying assumptions. The first is that non-native plants can spread easily from human developments to “pristine” areas. The second is that native plants cannot disperse through developed areas. Both assume anthropogenic changes to ecosystems create conditions that favor non-native plants and hinder native species. However, it is just as likely that anthropogenic alterations of habitats will favor certain groups of plant species with similar functional traits, whether native or not. Migration of plants can be divided into the following stages: dispersal, germination, establishment, reproduction and spread. Functional traits of species determine which are most successful at each of the stages of invasion or range enlargement. I studied the traits that allow both native and non-native plant species to disperse into freeway corridors, germinate, establish, reproduce, and then disperse along those corridors in Phoenix, Arizona. Field methods included seed bank sample collection and germination, vegetation surveys, and seed trapping. I also evaluated concentrations of plant-available nitrate as a result of localized nitrogen deposition. While many plant species found on the roadsides are either landscape varieties or typical weedy species, some uncommon native species and unexpected non-native species were also encountered. Maintenance regimes greatly influence the amount of vegetative cover and species composition along roadsides. Understanding which traits permit success at various stages of the invasion process indicates whether it is native, non-native, or species with particular traits that are likely to move through the city and establish in the desert. In a related case study conducted in Victoria, Australia, transportation professionals and ecologists were surveyed regarding preferences for roadside landscape design. Roadside design and maintenance projects are typically influenced by different groups of transportation professionals at various stages in a linear project cycle. Landscape architects and design professionals have distinct preferences and priorities compared to other transportation professionals and trained ecologists. The case study reveals the need for collaboration throughout the stages of design, construction and maintenance in order to efficiently manage roadsides for multiple priorities.

ContributorsGade, Kristin Joan (Author) / Kinzig, Ann P (Thesis advisor) / Grimm, Nancy (Committee member) / Perrings, Charles (Committee member) / Robbins, Paul (Committee member) / Stromberg, Juliet C. (Committee member) / Arizona State University (Publisher)
Created2010
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The highly-social plateau pika (Lagomorpha: Ochotona curzoniae) excavates vast burrow complexes in alpine meadows on the Tibetan Plateau. Colonies of over 300 individuals/ha have been reported. As an ecosystem engineer, their burrowing may positively impact ecosystem health by increasing plant species diversity, enhancing soil mixing, and boosting water infiltration. However,

The highly-social plateau pika (Lagomorpha: Ochotona curzoniae) excavates vast burrow complexes in alpine meadows on the Tibetan Plateau. Colonies of over 300 individuals/ha have been reported. As an ecosystem engineer, their burrowing may positively impact ecosystem health by increasing plant species diversity, enhancing soil mixing, and boosting water infiltration. However, pikas are commonly regarded as pests, and are heavily poisoned throughout their range. The underlying assumption of eradication programs is that eliminating pikas will improve rangeland quality and decrease soil erosion. This dissertation explores the link between plateau pikas and the alpine meadow ecosystem in Qinghai Province, PRC. This research uses both comparative field studies and theoretical modeling to clarify the role of pika disturbance. Specifically, these studies quantify the impact of pikas on nutrient cycling (via nutrient concentrations of vegetation and soil), hydrology (via water infiltration), local landscape properties (via spatial pattern description), and vascular plant communities (via species richness and composition). The competitive relationship between livestock and pikas is examined with a mathematical model. Results of this research indicate that pika colonies have both local and community level effects on water infiltration and plant species richness. A major contribution of pika disturbance is increased spatial heterogeneity, which likely underlies differences in the plant community. These findings suggest that the positive impact of plateau pikas on rangeland resources has been undervalued. In concurrence with other studies, this work concludes that plateau pikas provide valuable ecosystem services on the Tibetan Plateau.
ContributorsHogan, Brigitte Wieshofer (Author) / Smith, Andrew T. (Thesis advisor) / Anderies, J. Marty (Committee member) / Briggs, John M. (Committee member) / Stromberg, Juliet C. (Committee member) / Wu, Jianguo (Committee member) / Arizona State University (Publisher)
Created2010
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As an industrial society, humans have increasingly separated agricultural processes from natural ecosystems. Many areas of the Southwestern US, however, maintain traditional practices that link agricultural systems to the natural environment. One such practice, diverting river water into fields via earthen irrigation canals, allows ditch water to recharge

As an industrial society, humans have increasingly separated agricultural processes from natural ecosystems. Many areas of the Southwestern US, however, maintain traditional practices that link agricultural systems to the natural environment. One such practice, diverting river water into fields via earthen irrigation canals, allows ditch water to recharge groundwater and riparian vegetation to prosper along canal banks. As there is growing interest in managing landscapes for multiple ecosystem services, this study was undertaken to determine if irrigation canals function as an extension of the riparian corridor. I was specifically interested in determining if the processes within semi-arid streams that drive riparian plant community structure are manifested in earthen irrigation ditches. I examined herbaceous and woody vegetation along the middle Verde River, AZ, USA and three adjacent irrigation ditches across six months. I also collected sieved hydrochores--seeds dispersing through water--within ditches and the river twelve times. Results indicate that ditch vegetation was similar to streamside river vegetation in abundance (cover and basal area) due to surface water availability but more diverse than river streamside vegetation due to high heterogeneity. Compositionally, herbaceous vegetation along the ditch was most similar to the river banks, while low disturbance fostered woody vegetation along the ditches similar to high floodplain and river terrace vegetation. Hydrochore richness and abundance within the river was dependent on seasonality and stream discharge, but these relationships were dampened in the ditches. Species-specific strategies of hydrochory, however, did emerge in both systems. Strategies include pulse species, which disperse via hydrochory in strict accordance with their restricted dispersal windows, constant species, which are year round hydrochores, and combination species, which show characteristics of both. There was high overlap in the composition of hydrochores in the two systems, with obligate wetland species abundant in both. Upland species were more seasonally constant and abundant in the ditch water than the river. The consistency of river processes and similarity of vegetation suggest that earthen irrigation ditches do function as an extension of the riparian corridor. Thus, these man-made irrigation ditches should be considered by stakeholders for their multiple ecosystem services.
ContributorsBetsch, Jacqueline Michelle (Author) / Stromberg, Juliet C. (Thesis advisor) / Hall, Sharon J (Committee member) / Merritt, David M. (Committee member) / Arizona State University (Publisher)
Created2010
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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