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The coordination of group behavior in the social insects is representative of a broader phenomenon in nature, emergent biological complexity. In such systems, it is believed that large-scale patterns result from the interaction of relatively simple subunits. This dissertation involved the study of one such system: the social foraging of

The coordination of group behavior in the social insects is representative of a broader phenomenon in nature, emergent biological complexity. In such systems, it is believed that large-scale patterns result from the interaction of relatively simple subunits. This dissertation involved the study of one such system: the social foraging of the ant Temnothorax rugatulus. Physically tiny with small population sizes, these cavity-dwelling ants provide a good model system to explore the mechanisms and ultimate origins of collective behavior in insect societies. My studies showed that colonies robustly exploit sugar water. Given a choice between feeders unequal in quality, colonies allocate more foragers to the better feeder. If the feeders change in quality, colonies are able to reallocate their foragers to the new location of the better feeder. These qualities of flexibility and allocation could be explained by the nature of positive feedback (tandem run recruitment) that these ants use. By observing foraging colonies with paint-marked ants, I was able to determine the `rules' that individuals follow: foragers recruit more and give up less when they find a better food source. By altering the nutritional condition of colonies, I found that these rules are flexible - attuned to the colony state. In starved colonies, individual ants are more likely to explore and recruit to food sources than in well-fed colonies. Similar to honeybees, Temmnothorax foragers appear to modulate their exploitation and recruitment behavior in response to environmental and social cues. Finally, I explored the influence of ecology (resource distribution) on the foraging success of colonies. Larger colonies showed increased consistency and a greater rate of harvest than smaller colonies, but this advantage was mediated by the distribution of resources. While patchy or rare food sources exaggerated the relative success of large colonies, regularly (or easily found) distributions leveled the playing field for smaller colonies. Social foraging in ant societies can best be understood when we view the colony as a single organism and the phenotype - group size, communication, and individual behavior - as integrated components of a homeostatic unit.
ContributorsShaffer, Zachary (Author) / Pratt, Stephen C (Thesis advisor) / Hölldobler, Bert (Committee member) / Janssen, Marco (Committee member) / Fewell, Jennifer (Committee member) / Liebig, Juergen (Committee member) / Arizona State University (Publisher)
Created2014
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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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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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Communication amongst eusocial insect is key to their success. Ants rely on signaling to mediate many different functions within a colony such as policing and nest mate recognition. Camponotus floridanus uses chemosensory signaling in the form of cuticular hydrocarbons to regulate these functions. Each cuticular hydrocarbon profile contains numerous hydrocarbons,

Communication amongst eusocial insect is key to their success. Ants rely on signaling to mediate many different functions within a colony such as policing and nest mate recognition. Camponotus floridanus uses chemosensory signaling in the form of cuticular hydrocarbons to regulate these functions. Each cuticular hydrocarbon profile contains numerous hydrocarbons, however it is yet to be seen if Camponotus floridanus can discriminate between linear hydrocarbons of similar length. Individual specimens were conditioned in three different ways: 5 conditioning with high concentration of sugar water (1;1 ratio), 1 conditioning with high concentration of sugar water, and 5 conditioning with low concentration of sugar water (1;4). Two linear hydrocarbons were use, C23 and C24, with C23 always being the conditioned stimulus. Specimens who were conditioned 5 times with high concentration of sugar water were the only group to show a significant response to the conditioned stimulus with a p-value of .008 and exhibited discrimination behavior 46% of the time. When compared 5 conditioning with high concentration to the other two testing conditioning groups, 1 conditioning with high concentration produced an insignificant p-value of .13 was obtained whereas when comparing it with 5 conditioning low concentration of sugar a significant p-value of .0132 was obtained. This indiciates that Camponotus floridanus are capable of discrimination however must be conditioned with high concentration of sugar water, while number of conditioning is insignificant.
ContributorsDamari, Ben Aviv (Author) / Liebig, Juergen (Thesis director) / Ghaninia, Majid (Committee member) / Pratt, Stephen (Committee member) / Barrett, The Honors College (Contributor) / School of Life Sciences (Contributor)
Created2014-05
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ABSTRACT Communication is vital in the context of everyday life for all organisms, but particularly so in social insects, such as Z. nevadensis. The overall lifestyle and need for altruistic acts of individuals within a colony depends primarily on intracolony chemical communication, with a focus on odorants. The perception of

ABSTRACT Communication is vital in the context of everyday life for all organisms, but particularly so in social insects, such as Z. nevadensis. The overall lifestyle and need for altruistic acts of individuals within a colony depends primarily on intracolony chemical communication, with a focus on odorants. The perception of these odorants is made possible by the chemoreceptive functions of sensilla basiconica and sensilla trichoid which exist on the antennal structure. The present study consists of both a morphological analysis and electrophysiological experiment concerning sensilla basiconica. It attempts to characterize the function of neurons present in sensilla basiconica through single sensillum recordings and contributes to existing literature by determining if a social insect, such as the dampwood termite, is able to perceive a wide spectrum of odorants despite having significantly fewer olfactory receptors than most other social insect species. Results indicated that sensilla basiconica presence significantly out-paced that of sensilla trichoid and sensilla chaetica combined, on all flagellomeres. Analysis demonstrated significant responses to all general odorants and several cuticular hydrocarbons. Combined with the knowledge of fewer olfactory receptors present in this species and their lifestyle, results may indicate a positive association between the the social complexity of an insect's lifestyle and the number of ORs the individuals within that colony possess.
ContributorsMcGlone, Taylor (Author) / Liebig, Juergen (Thesis director) / Ghaninia, Majid (Committee member) / Barrett, The Honors College (Contributor)
Created2015-05
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Olfactory discrimination tasks can provide useful information about how olfaction may have evolved by demonstrating which types of compounds animals will detect and respond to. Ants discriminate between nestmates and non-nestmates by using olfaction to detect the cuticular hydrocarbons on other ants, and Camponotus floridanus have particularly clear and aggressive

Olfactory discrimination tasks can provide useful information about how olfaction may have evolved by demonstrating which types of compounds animals will detect and respond to. Ants discriminate between nestmates and non-nestmates by using olfaction to detect the cuticular hydrocarbons on other ants, and Camponotus floridanus have particularly clear and aggressive responses to non-nestmates. A new method of adding hydrocarbons to ants, the “Snow Globe” method was further optimized and tested on C. floridanus. It involves adding hydrocarbons and a solvent to a vial of water, vortexing it, suspending hydrocarbon droplets throughout the solution, and then dipping a narcotized ant in. It is hoped this method can evenly coat ants in hydrocarbon. Ants were treated with heptacosane (C27), nonacosane (C29), hentriacontane (C31), a mixture of C27/C29/C31, 2-methyltriacontane (2MeC30), S-3-methylhentriacontane (SMeC31), and R-3-methylhentriacontane (RMeC31). These were chosen to see how ants reacted in a nestmate recognition context to methyl-branched hydrocarbons, R and S enantiomers, and to multiple added alkanes. Behavior assays were performed on treated ants, as well as two untreated controls, a foreign ant and a nestmate ant. There were 15 replicates of each condition, using 15 different queenright colonies. The Snow Globe method successfully transfers hydrocarbons, as confirmed by solid phase microextraction (SPME) done on treated ants, and the behavior assay data shows the foreign control, SMeC31, and the mixture of C27/29/31 were all statistically significant in their differences from the native control. The multiple alkane mixture received a significant response while single alkanes did not, which supports the idea that larger variations in hydrocarbon profile are needed for an ant to be perceived as foreign. The response to SMeC31 shows C. floridanus can respond during nestmate recognition to hydrocarbons that are not naturally occurring, and it indicates the nestmate recognition process may simply be responding to any compounds not found in the colony profile and rather than detecting particular foreign compounds.

ContributorsNoss, Serena Marie (Author) / Liebig, Juergen (Thesis director) / Pratt, Stephen (Committee member) / Haight, Kevin (Committee member) / School of Life Sciences (Contributor) / Department of Psychology (Contributor) / Barrett, The Honors College (Contributor)
Created2021-05
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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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Weevils are one of the most diverse groups of animals with thousands of species suspected to remain undiscovered. The Conoderinae Schoenherr, 1833 are no exception, being especially diverse and unknown in the Neotropics where they are recognizable for their unique behaviors and color patterns among weevils. Despite these peculiarities, the

Weevils are one of the most diverse groups of animals with thousands of species suspected to remain undiscovered. The Conoderinae Schoenherr, 1833 are no exception, being especially diverse and unknown in the Neotropics where they are recognizable for their unique behaviors and color patterns among weevils. Despite these peculiarities, the group has received little attention from researchers in the past century, with almost nothing known about their evolution. This dissertation presents a series of three studies that begin to elucidate the evolutionary history of these bizarre and fascinating weevils, commencing with an overview of their biology and classificatory history (Chapter 1).

Chapter 2 presents the first formal cladistic analysis on the group to redefine the New World tribes Lechriopini Lacordaire, 1865 and Zygopini, Lacordaire, 1865. An analysis of 75 taxa (65 ingroup) with 75 morphological characters yielded six equally parsimonious trees and synapomorphies that are used to reconstitute the tribes, resulting in the transfer of sixteen genera from the Zygopini to the Lechriopini and four generic transfers out of the Lechriopini to elsewhere in the Conoderinae.

Chapter 3 constitutes a taxonomic revision of the genus Trichodocerus Chevrolat, 1879, the sole genus in the tribe Trichodocerini Champion, 1906, which has had an uncertain phylogenetic placement in the Curculionidae but has most recently been treated in the Conoderinae. In addition to redescriptions of the three previously described species placed in the genus, twenty-four species are newly described and an identification key is provided for all recognized species groups and species.

Chapter 4 quantitatively tests the similarity in color pattern among species hypothesized to belong to several different mimicry complexes. The patterns of 160 species of conoderine weevils were evaluated for 15 categorical and continuous characters. Non-metric multidimensional scaling (NMDS) is used to visualize similarity by the proximity of individual species and clusters of species assigned to a mimicry complex in ordination space with clusters being statistically tested using permutational multivariate analysis of variance (PERMANOVA).
ContributorsAnzaldo, Salvatore (Author) / Franz, Nico (Thesis advisor) / Martins, Emilia (Committee member) / Rabeling, Christian (Committee member) / Pigg, Kathleen (Committee member) / Arizona State University (Publisher)
Created2019