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The repression of reproductive competition and the enforcement of altruism are key components to the success of animal societies. Eusocial insects are defined by having a reproductive division of labor, in which reproduction is relegated to one or few individuals while the rest of the group members maintain the colony

The repression of reproductive competition and the enforcement of altruism are key components to the success of animal societies. Eusocial insects are defined by having a reproductive division of labor, in which reproduction is relegated to one or few individuals while the rest of the group members maintain the colony and help raise offspring. However, workers have retained the ability to reproduce in most insect societies. In the social Hymenoptera, due to haplodiploidy, workers can lay unfertilized male destined eggs without mating. Potential conflict between workers and queens can arise over male production, and policing behaviors performed by nestmate workers and queens are a means of repressing worker reproduction. This work describes the means and results of the regulation of worker reproduction in the ant species Aphaenogaster cockerelli. Through manipulative laboratory studies on mature colonies, the lack of egg policing and the presence of physical policing by both workers and queens of this species are described. Through chemical analysis and artificial chemical treatments, the role of cuticular hydrocarbons as indicators of fertility status and the informational basis of policing in this species is demonstrated. An additional queen-specific chemical signal in the Dufour's gland is discovered to be used to direct nestmate aggression towards reproductive competitors. Finally, the level of actual worker-derived males in field colonies is measured. Together, these studies demonstrate the effectiveness of policing behaviors on the suppression of worker reproduction in a social insect species, and provide an example of how punishment and the threat of punishment is a powerful force in maintaining cooperative societies.
ContributorsSmith, Adrian A. (Author) / Liebig, Juergen (Thesis advisor) / Hoelldobler, Bert (Thesis advisor) / Gadau, Juergen (Committee member) / Johnson, Robert A. (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2011
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Description
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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Description
Social insects collectively exploit food sources by recruiting nestmates, creating positive feedback that steers foraging effort to the best locations. The nature of this positive feedback varies among species, with implications for collective foraging. The mass recruitment trails of many ants are nonlinear, meaning that small increases in recruitment effort

Social insects collectively exploit food sources by recruiting nestmates, creating positive feedback that steers foraging effort to the best locations. The nature of this positive feedback varies among species, with implications for collective foraging. The mass recruitment trails of many ants are nonlinear, meaning that small increases in recruitment effort yield disproportionately large increases in recruitment success. The waggle dance of honeybees, in contrast, is believed to be linear, meaning that success increases proportionately to effort. However, the implications of this presumed linearityhave never been tested. One such implication is the prediction that linear recruiters will equally exploit two identical food sources, in contrast to nonlinear recruiters, who randomly choose only one of them. I tested this prediction in colonies of honeybees that were isolated in flight cages and presented with two identical sucrose feeders. The results from 15 trials were consistent with linearity, with many cases of equal exploitation of the feeders. In addition, I tested the prediction that linear recruiters can reallocate their forager distribution when unequal feeders are swapped in position. Results from 15 trials were consistent with linearity, with many cases of clear choice for a stronger food source, followed by a subsequent switch with reallocation of foragers to the new location of the stronger food source. These findings show evidence of a linear pattern of nestmate recruitment, with implications for how colonies effectively distribute their foragers across available resources.
ContributorsAlam, Showmik (Author) / Shaffer, Zachary (Thesis advisor) / Pratt, Stephen C (Thesis advisor) / Ozturk, Cahit (Committee member) / Pavlic, Theodore (Committee member) / Arizona State University (Publisher)
Created2022
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Description
Eusocial insect colonies have often been imagined as “superorganisms” exhibiting tight homeostasis at the colony level. However, colonies lack the tight spatial and organizational integration that many multicellular, unitary organisms exhibit. Precise regulation requires rapid feedback, which is often not possible when nestmates are distributed across space, making decisions asynchronously.

Eusocial insect colonies have often been imagined as “superorganisms” exhibiting tight homeostasis at the colony level. However, colonies lack the tight spatial and organizational integration that many multicellular, unitary organisms exhibit. Precise regulation requires rapid feedback, which is often not possible when nestmates are distributed across space, making decisions asynchronously. Thus, one should expect poorer regulation in superorganisms than unitary organisms.Here, I investigate aspects of regulation in collective foraging behaviors that involve both slow and rapid feedback processes. In Chapter 2, I examine a tightly coupled system with near-instantaneous signaling: teams of weaver ants cooperating to transport massive prey items back to their nest. I discover that over an extreme range of scenarios—even up vertical surfaces—the efficiency per transporter remains constant. My results suggest that weaver ant colonies are maximizing their total intake rate by regulating the allocation of transporters among loads. This is an exception that “proves the rule;” the ant teams are recapitulating the physical integration of unitary organisms. Next, I focus on a process with greater informational constraints, with loose temporal and spatial integration. In Chapter 3, I measure the ability of solitarily foraging Ectatomma ruidum colonies to balance their collection of protein and carbohydrates given different nutritional environments. Previous research has found that ant species can precisely collect a near-constant ratio between these two macronutrients, but I discover these studies were using flawed statistical approaches. By developing a quantitative measure of regulatory effect size, I show that colonies of E. ruidum are relatively insensitive to small differences in food source nutritional content, contrary to previously published claims. In Chapter 4, I design an automated, micro-RFID ant tracking system to investigate how the foraging behavior of individuals integrates into colony-level nutrient collection. I discover that spatial fidelity to food resources, not individual specialization on particular nutrient types, best predicts individual forager behavior. These findings contradict previously published experiments that did not use rigorous quantitative measures of specialization and confounded the effects of task type and resource location.
ContributorsBurchill, Andrew Taylor (Author) / Pavlic, Theodore P (Thesis advisor) / Pratt, Stephen C (Thesis advisor) / Hölldobler, Bert (Committee member) / Cease, Arianne (Committee member) / Berman, Spring (Committee member) / Arizona State University (Publisher)
Created2022
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Description
Insecticide resistance is a continuing issue that negatively affects both public health and agriculture and allows vector-borne diseases to spread throughout the globe. To improve resistance management strategies (RMS), robust susceptibility bioassays need to be performed in order to fill the gap of the relationship between resistant and susceptible genotype

Insecticide resistance is a continuing issue that negatively affects both public health and agriculture and allows vector-borne diseases to spread throughout the globe. To improve resistance management strategies (RMS), robust susceptibility bioassays need to be performed in order to fill the gap of the relationship between resistant and susceptible genotype and phenotype, and a deeper knowledge of how bioassay data relates to vector control success or failure is imperative. A bioassay method that is infrequently used but yields robust results is the topical application bioassay, where the insect is directly treated with a constant volume and concentration of an insecticide via a syringe. To bring more attention to this method, my colleagues and I published a paper in the Journal of Visualized Experiments where the optimized protocol of the topical application bioassay for mosquitoes and fruit flies is described, and the strengths and limitations to the method are explained. To further investigate insecticide susceptibility tests, I set up my individual project where I used Aedes aegypti mosquitoes to compare the topical application bioassay to the commonly used Centers for Disease Control and Prevention (CDC) bottle bioassay and World Health Organization (WHO) tube test. The objective of this study was to test which method exhibited the most variability in mortality results, which would guide the choice of assay to determine the link between resistant and susceptible genotype and phenotype. The results showed that the topical application method did indeed exhibit the least amount of variation, followed by the CDC bottle bioassay (WHO data is currently being collected). This suggests that the topical application bioassay could be a useful tool in insecticide resistance surveillance studies, and, depending on the goal, may be better than the CDC and WHO tube tests for assessing resistance levels at a given site. This study challenges the value of the widely used CDC and WHO assays and provides a discussion on the importance of technical and practical resistance assays. This will help vector control specialists to collect accurate surveillance data that will inform effective RMS.
ContributorsAlthoff, Rachel (Author) / Huijben, Silvie (Thesis advisor) / Harris, Robin (Committee member) / Collins, James (Committee member) / Arizona State University (Publisher)
Created2022
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Description
The weevil genus Pachnaeus Schoenherr, 1826 (Coleoptera: Curculionidae: Entiminae: Eustylini Lacordaire) is revised to accommodate 21 species, including the following 10 new species from the northern Caribbean region: Pachnaeus andersoni sp. nov. (Little Cayman), Pachnaeus eisenbergi sp. nov. (Jamaica), Pachnaeus godivae sp. nov. (Cayman Brac), Pachnaeus gordoni sp. nov. (Jamaica),

The weevil genus Pachnaeus Schoenherr, 1826 (Coleoptera: Curculionidae: Entiminae: Eustylini Lacordaire) is revised to accommodate 21 species, including the following 10 new species from the northern Caribbean region: Pachnaeus andersoni sp. nov. (Little Cayman), Pachnaeus eisenbergi sp. nov. (Jamaica), Pachnaeus godivae sp. nov. (Cayman Brac), Pachnaeus gordoni sp. nov. (Jamaica), Pachnaeus howdenae sp. nov. (Bahamas), Pachnaeus ivieorum sp. nov. (Bahamas with adventive records from Florida), Pachnaeus maestrensis sp. nov. (Cuba), Pachnaeus morelli sp. nov. (Haiti), Pachnaeus obrienorum sp. nov. (Cuba and Bahamas), and Pachnaeus quadrilineatus sp. nov. (Jamaica).Pachnaeus can be distinguished from similar, co-occurring taxa such as Exophthalmus quadrivittatus (Olivier, 1807), Exophthalmus roseipes (Chevrolat, 1876), Exophthalmus vittatus (Linnaeus, 1758), and Diaprepes abbreviatus (Linnaeus, 1758) by (1) the presence of postocular vibrissae, (2) endophallus primarily membranous and sac-like proximally, and long (>3 × width), tubular, and sclerotized distally, (3) additional endophallic sclerites typically absent, (4) a never bicarinate, typically tricarinate, rostrum, and several additional characteristics of the pedon, endophallus, pronotal structure, rostral structure, and scaling. Based on these characters, Pachnaeus sommeri (Munck af Rosenschoeld in Schoenherr, 1840) comb, nov. and Pachnaeus gowdeyi (Marshall, 1926) comb. nov. are transferred into the genus from Exophthalmus Schoenherr and Lachnopus Schoenherr respectively. This revision provides genus and species redescriptions, diagnoses, illustrations, and the first comprehensive key to all 21 species within the present circumscription of Pachnaeus, in addition to reviewing the known biology and observed intraspecific variation within species. The complex taxonomic history of the genus is reviewed, and the evolutionary relationships of its presumed constituent clades are proposed through the construction of informal species groups and subgroups based on diagnosable shared traits. Lectotypes for Pachnaeus citri Marshall, Pachnaeus costatus Perroud, and Exophthalmus sommeri Munck af Rosenschoeld in Schoenherr and paralectotypes of P. citri (3 specimens) and E. sommeri (4 specimens) are designated. New state and national records are reported for Pachnaeus azurescens Gyllenhal in Schoenherr for Florida, U.S.A. and new national records are reported for Pachnaeus litus (Germar) for the Bahamas. Validity of the names Docorhinus Schoenherr, 1823 and Pachnaeus Schoenherr, 1826 is treated. Generic placement of Pachnaeus roseipes Chevrolat, 1876 is explored.
ContributorsReily, Brian Herndon (Author) / Franz, Nico M (Thesis advisor) / Taylor, Jesse (Committee member) / Pigg, Kathleen (Committee member) / Johnston, Murray A (Committee member) / Arizona State University (Publisher)
Created2022
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Description
The wide-spread use of insecticides has contributed to the rapid decline of insect diversity and abundance. In light of recent guidance from international and governmental organizations, other non-chemical control methods are necessary to control insect pest populations. In my study, I used occupancy modeling techniques and found that environmental variables

The wide-spread use of insecticides has contributed to the rapid decline of insect diversity and abundance. In light of recent guidance from international and governmental organizations, other non-chemical control methods are necessary to control insect pest populations. In my study, I used occupancy modeling techniques and found that environmental variables could predict the presence of Rhaphidophoridae, in Hidalgo, Mexico. The results showed that variables associated with forested habitats increase the probability of Rhaphidophoridae detection, and higher elevation increases the probability of Rhaphidophoridae occupancy. Understanding the specific habitat variables associated with human detection and occupancy of Rhaphidophoridae give people the ability to utilize the Integrative Pest Management (IPM) strategy of cultural control to prevent Rhaphidophoridae pest populations in my study region.
ContributorsPalmrose-Krieger, Christina (Author) / Martins, Emilia P (Thesis advisor) / Harrison, Jon F (Committee member) / Zúñiga-Vega, J Jaime (Committee member) / Arizona State University (Publisher)
Created2024
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Description
The migratory grasshopper (Melanoplus sanguinipes) is one of the most economically important grasshoppers in the western rangelands of the United States (US), capable of causing incredible amounts of damage to crops and rangelands. While M. sanguinipes has been the focus of many research studies, areas like field nutritional physiology and

The migratory grasshopper (Melanoplus sanguinipes) is one of the most economically important grasshoppers in the western rangelands of the United States (US), capable of causing incredible amounts of damage to crops and rangelands. While M. sanguinipes has been the focus of many research studies, areas like field nutritional physiology and ecology, and interactions between nutritional physiology and biopesticide resistance have very little research. This dissertation presents a multifaceted approach through three research-driven chapters that examine the nutritional physiology of M. sanguinipes and how it interacts with an entomopathogenic fungus for grasshopper management, as well as the challenges of using biopesticides for grasshopper management. Using the Geometric Framework for Nutrition (GFN), I established baseline macronutrient intake for M. sanguinipes, both in laboratory and field populations. Through this work, I found that field and lab populations can exhibit different protein (p) to carbohydrate (c) ratios, or Intake Targets (ITs), but that the field populations had ITs that matched the nutrients available in their environment. I also used the GFN to show that infections with the fungal entomopathogen Metarhizium robertsii DWR2009 did not alter ITs in M. sanguinipes. Although, when confined to carbohydrate- or protein-biased diets, infected grasshoppers had a slightly extended lifespan relative to grasshoppers fed balanced protein:carbohydrate diets. Interestingly, in a postmortem for the grasshopper, the fungus was only able to effectively sporulate on grasshoppers fed the 1p:1c diets, suggesting that grasshopper diet can have substantial impacts on the spread of fungal biopesticides throughout a population, in the absence of any inhibitory abiotic factors. Lastly, I examined the major barriers to fungal and microsporidian biopesticide usage in the United States, including low efficacy, thermal and environmental sensitivity, non-target effects, unregistered or restricted use, and economic or accessibility barriers. I also explored potential solutions to these challenges. This dissertation's focus on Melanoplus sanguinipes and Metarhizium roberstii Strain DWR2009, generates new information about how nutritional physiology and immunology intersect to impact M. sanguinipes performance. The methodology in each of the experimental chapters provides a framework for examining other problematic grasshopper species, by determining baseline nutritional physiology, and coupling nutrition with immunology to maximize the effectiveness of biological pesticides.
ContributorsZembrzuski, Deanna (Author) / Cease, Arianne (Thesis advisor) / Harrison, Jon (Committee member) / Angilletta, Michael (Committee member) / Jaronski, Stefan (Committee member) / Arizona State University (Publisher)
Created2023
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Description
The diverse weevil genus Rhyssomatus Schoenherr, 1837 (Curculionidae: Molytinae: Cleoginini) is currently composed of 175 species throughout the New World (O’Brien et. al 1982; Wibmer et. al 1986). The majority of species are generalist feeders and the group contains many notorious agricultural pests, such as Rhyssomatus nigerimus Fahraeus 1837 and

The diverse weevil genus Rhyssomatus Schoenherr, 1837 (Curculionidae: Molytinae: Cleoginini) is currently composed of 175 species throughout the New World (O’Brien et. al 1982; Wibmer et. al 1986). The majority of species are generalist feeders and the group contains many notorious agricultural pests, such as Rhyssomatus nigerimus Fahraeus 1837 and R. subtillis Fiedler 1937 that cause thousands of dollars’ worth of crop damage in South America, Central America, and West Indies (Cazado, 2016; Lopez-Guillen, et. al). Although most notable as a crop pest in the literature, the species Rhyssomatus maginatus Fahraeus has also proven to be a great success in an Integrated Pest Management (IPM), controlling the invasive leguminous tree Sesbenia punicea (Cav.) Benth., in South Africa. (Hoffman & Moran 1991; 1992; 1998; 1999). The last century has seen revisions of the Neotropical species with Central American species revised in 1904 by Champion and the South American taxa treated by Fiedler in the subsequent years of 1937 and 1942 (O’Brien & Wibmer, 1982; Wibmer & O’Brien, 1986). However, North American fauna have not been treated since Casey in 1895 and revision is needed as climate change and global trade have more than likely expanded the distributional range of previously lower latitude Rhyssomatus species northwards.
ContributorsArguez, Katherine MacKenzie (Author) / Franz, Nico (Thesis advisor) / Pigg, Kathleen (Committee member) / Gile, Gillian (Committee member) / Arizona State University (Publisher)
Created2023
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Description
Vector control plays an important role in the prevention and control of mosquito-borne diseases (MBDs). As there are no (prophylactic) drugs and/or vaccines available for many arboviral diseases (such as zika, chikungunya, Saint Louis encephalitis, Ross River virus), the frontline approach to prevent or reduce disease morbidity and mortality is

Vector control plays an important role in the prevention and control of mosquito-borne diseases (MBDs). As there are no (prophylactic) drugs and/or vaccines available for many arboviral diseases (such as zika, chikungunya, Saint Louis encephalitis, Ross River virus), the frontline approach to prevent or reduce disease morbidity and mortality is through the reduction of the mosquito vector population size and/or reducing vector-human contact using insecticides. Frontline tools in malaria (an MBD caused by a parasite) control and elimination have been drugs (targeting the malaria parasite) and insecticides (targeting the vectors) through indoor residual spraying (IRS) (spraying the internal walls and sometimes the roofs of dwellings with residual insecticides to kill adult mosquito vectors), and long-lasting insecticidal nets (LLINs), while arboviral vectors are frequently targeted using outdoor fogging and space spraying (indoor or outdoor spraying of insecticides to kill adult mosquito vectors). Integrative and novel vector control efforts are urgently needed since the aforementioned tools may not be as effective against those mosquito species that are resistant to insecticides and/or have a different (or changed) behavior allowing them to avoid existing tools. In Chapters 2 and 3, I investigate mosquito vector surveillance in Arizona by (i) discussing the species composition and public health implications of the State’s mosquito fauna, and (ii) comparing the effectiveness of 4 different carbon dioxide (CO2) sources in attracting different mosquito species on the Arizona State University Tempe Campus. In Chapters 4 and 5, I investigate a novel vector control tool by (i) completing a literature review on using electric fields (EFs) to control insects, and (ii) presenting novel data on using Insulated Conductor Wires (ICWs) to generate EFs that prevent host-seeking female Aedes aegypti from entering spaces. In Chapter 6, I discuss the non-target effects of chemical malaria control on other arthropods, including other biological and mechanical infectious disease vectors. Overall, this dissertation highlights the important role that the development of novel surveillance and vector control tools could play in improved mosquito control, which ultimately will reduce disease morbidity and mortality.
ContributorsJobe, Ndey Bassin (Author) / Paaijmans, Krijn (Thesis advisor) / Cease, Arianne (Committee member) / Hall, Sharon (Committee member) / Huijben, Silvie (Committee member) / Arizona State University (Publisher)
Created2024