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Of all the signals and cues that orchestrate the activities of a social insect colony, the reproductives' fertility pheromones are perhaps the most fundamental. These pheromones regulate reproductive division of labor, a defining characteristic of eusociality. Despite their critical role, reproductive fertility pheromones are not evenly expressed across the development

Of all the signals and cues that orchestrate the activities of a social insect colony, the reproductives' fertility pheromones are perhaps the most fundamental. These pheromones regulate reproductive division of labor, a defining characteristic of eusociality. Despite their critical role, reproductive fertility pheromones are not evenly expressed across the development of a social insect colony and may even be absent in the earliest colony stages. In the ant Camponotus floridanus, queens of incipient colonies do not produce the cuticular hydrocarbons that serve as fertility and egg-marking signals in this species. My dissertation investigates the consequences of the dramatic change in the quantity of these pheromones that occurs as the colony grows. C. floridanus workers from large, established colonies use egg surface hydrocarbons to discriminate among eggs. Eggs with surface hydrocarbons typical of eggs laid by established queens are nurtured, whereas eggs lacking these signals (i.e., eggs laid by workers and incipient queens) are destroyed. I characterized how workers from incipient colonies responded to eggs lacking queen fertility hydrocarbons. I found that established-queen-laid eggs, incipient-queen-laid eggs, and worker-laid eggs were not destroyed by workers at this colony stage. Destruction of worker-laid eggs is a form of policing, and theoretical models predict that policing should be strongest in incipient colonies. Since there was no evidence of policing by egg-eating in incipient C. floridanus colonies, I searched for evidence of another policing mechanism at this colony stage. Finding none, I discuss reasons why policing behavior may not be expressed in incipient colonies. I then considered the mechanism that accounts for the change in workers' response to eggs. By manipulating ants' egg experience and testing their egg-policing decisions, I found that ants use a combination of learned and innate criteria to discriminate between targets of care and destruction. Finally, I investigated how the increasing strength of queen-fertility hydrocarbons affects nestmate recognition, which also relies on cuticular hydrocarbons. I found that queens with strong fertility hydrocarbons can be transferred between established colonies without aggression, but they cannot be introduced into incipient colonies. Queens from incipient colonies cannot be transferred into incipient or established colonies.
ContributorsMoore, Dani (Author) / Liebig, Juergen (Thesis advisor) / Gadau, Juergen (Committee member) / Pratt, Stephen (Committee member) / Smith, Brian (Committee member) / Rutowski, Ronald (Committee member) / Arizona State University (Publisher)
Created2012
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Description
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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Description
Warning coloration deters predators from attacking prey that are defended, usually by being distasteful, toxic, or otherwise costly for predators to pursue and consume. Predators may have an innate response to warning colors or learn to associate them with a defense through trial and error. In general, predators should select

Warning coloration deters predators from attacking prey that are defended, usually by being distasteful, toxic, or otherwise costly for predators to pursue and consume. Predators may have an innate response to warning colors or learn to associate them with a defense through trial and error. In general, predators should select for warning signals that are easy to learn and recognize. Previous research demonstrates long-wavelength colors (e.g. red and yellow) are effective because they are readily detected and learned. However, a number of defended animals display short-wavelength coloration (e.g. blue and violet), such as the pipevine swallowtail butterfly (Battus philenor). The role of blue coloration in warning signals had not previously been explicitly tested. My research showed in laboratory experiments that curve-billed thrashers (Toxostoma curvirostre) and Gambel's quail (Callipepla gambelii) can learn and recognize the iridescent blue of B. philenor as a warning signal and that it is innately avoided. I tested the attack rates of these colors in the field and blue was not as effective as orange. I concluded that blue colors may function as warning signals, but the effectiveness is likely dependent on the context and predator.

Blue colors are often iridescent in nature and the effect of iridescence on warning signal function was unknown. I reared B. philenor larvae under varied food deprivation treatments. Iridescent colors did not have more variation than pigment-based colors under these conditions; variation which could affect predator learning. Learning could also be affected by changes in appearance, as iridescent colors change in both hue and brightness as the angle of illuminating light and viewer change in relation to the color surface. Iridescent colors can also be much brighter than pigment-based colors and iridescent animals can statically display different hues. I tested these potential effects on warning signal learning by domestic chickens (Gallus gallus domesticus) and found that variation due to the directionality of iridescence and a brighter warning signal did not influence learning. However, blue-violet was learned more readily than blue-green. These experiments revealed that the directionality of iridescent coloration does not likely negatively affect its potential effectiveness as a warning signal.
ContributorsPegram, Kimberly Vann (Author) / Rutowski, Ronald L (Thesis advisor) / Hoelldobler, Berthold (Committee member) / Liebig, Juergen (Committee member) / McGraw, Kevin (Committee member) / Smith, Brian H. (Committee member) / Arizona State University (Publisher)
Created2015
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Description
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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Description

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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Description
Complex animal societies consist of a plethora of interactions between members. To successfully thrive they must be able to recognize members and their kin, and to understand how they do this we need sufficient and reliable methods of testing. Eusocial insects are especially good at recognizing their nestmates, but the

Complex animal societies consist of a plethora of interactions between members. To successfully thrive they must be able to recognize members and their kin, and to understand how they do this we need sufficient and reliable methods of testing. Eusocial insects are especially good at recognizing their nestmates, but the exact mechanism or how well they can discriminate is unknown. Ants achieve nestmate recognition by identifying varying proportions of cuticular hydrocarbons. Previous studies have shown ants can be trained to discriminate between pairs of hydrocarbons. This study aims to compare two methodologies previously shown to demonstrate odor learning to identify which one is the most promising to use for future odor learning experiments. The two methods tested were adapted from Sharma et al. (2015) and Guerrieri and d’Ettorre (2010). The results showed that the Guerrieri method demonstrated learning better and was more reliable and faster than the Sharma method. The Guerrieri method should be used in future experiments regarding odor learning discrimination
ContributorsDavis, Cole (Author) / Liebig, Juergen (Thesis director) / Stephen, Pratt (Committee member) / School of Life Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2020-05
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Description
Across the animal kingdom, communication serves a vital purpose. The transfer of information between and among species is often paramount to many behaviors including mating, collaboration, and defense. While research has provided tremendous insight into animal communication and interaction, there is still much that we have yet to understand. Due

Across the animal kingdom, communication serves a vital purpose. The transfer of information between and among species is often paramount to many behaviors including mating, collaboration, and defense. While research has provided tremendous insight into animal communication and interaction, there is still much that we have yet to understand. Due to their reliance on interactions that maximize efficiency within their complicated colony structure and array of member roles, eusocial insects serve as an excellent model for animal communication. Among eusocial insects, ants are some of the most heavily researched, with a tremendous amount of literature focused on their cuticular hydrocarbons. Along with serving as a waterproofing agent, cuticular hydrocarbons also play a major role in recognition and communication in these insects. By studying the importance of hydrocarbons in ant social structure, their tremendously specialized olfactory system, and the use of learning assays in its study, parallels between communication in ants and other animals are revealed, demonstrating how ants serve as a relevant model for animal communication as a whole.
ContributorsSpirek, Benton Forest Ensminger (Author) / Liebig, Juergen (Thesis director) / Pratt, Stephen (Committee member) / School of Life Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2020-12