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Niche variation among sexes and life stages within a population has been documented in many species, yet few studies have investigated niche variation within demographic groups or across ecological contexts. We examined the extent to which pregnant California sea lions (Zalophus californianus) at each of three breeding colonies target alternative

Niche variation among sexes and life stages within a population has been documented in many species, yet few studies have investigated niche variation within demographic groups or across ecological contexts. We examined the extent to which pregnant California sea lions (Zalophus californianus) at each of three breeding colonies target alternative prey resources and habitats. The three colonies are distributed across distinct regions of the Gulf of California, Mexico and have divergent population dynamics. We compared the nature of niche variation among colonies and investigated the fitness consequences of different foraging strategies within each colony. We analyzed the δ13C and δ15N values from fur collected from 206 suckling pups to characterize relative maternal foraging locations (δ13C) and trophic levels (δ15N) during the metabolically demanding late stages of gestation and lactation that occur simultaneously in California sea lions. The δ13C and δ15N values were regressed against pup body condition index values to compare the relative individual-level fitness benefits of different maternal foraging strategies. We found that the nature and extent of niche variation differed among colonies. Niche variation was most pronounced at the two largest colonies that appear to experience the highest levels of intraspecific competition and the variation was consistent with habitat features. One colony (Granito) displayed two distinct foraging groups with indistinguishable median pup body condition values, whereas the second (San Jorge) exhibited continuous niche variation and pup body condition varied in relation to maternal foraging location and trophic level, suggesting disparities among alternative foraging strategies. For the smallest colony (Los Islotes), females occupy similar niches with a few outliers. Body condition values of pups at this colony were most variable, but did not vary with maternal foraging strategy. Our results provide evidence for intrapopulation niche variation among demographically similar individuals during a period of high metabolic stress and reproductive importance. This work suggests possible fitness benefits conferred by alternative foraging strategies, and calls into question the common assumption that members of a population are ecologically equivalent. Future research aimed at understanding animal foraging strategies should consider the nature and extent of niche variation in the context of local ecological conditions.
ContributorsCrawford, Tara Gancos (Author) / Gerber, Leah R. (Thesis advisor) / Ogle, Kiona (Committee member) / Kurle, Carolyn M (Committee member) / Arizona State University (Publisher)
Created2013
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Description
This study examines almost 5000 lithic artifacts from three rockshelters in the Belize, part of the Maya Lowlands. These sites bear evidence of human occupation throughout the last 14,000 years, a period over which I assess change in four lithic traits: 1) cortex, or the weathered outer surface of a

This study examines almost 5000 lithic artifacts from three rockshelters in the Belize, part of the Maya Lowlands. These sites bear evidence of human occupation throughout the last 14,000 years, a period over which I assess change in four lithic traits: 1) cortex, or the weathered outer surface of a toolstone cobble; 2) platform type, or the degree of preparation of striking platforms; 3) bifaciality, or whether tools were flaked on one or both faces; and 4) retouch, or the removal of small flakes from tool edges. These traits are differentially associated with two modes of technological organization: curation and expedience. Curation involves greater effort spent creating tools with longer use-lives and is associated with low levels of cortex, more complex platforms, more bifacial flaking, and higher amounts of retouch. It is also more typical of highly mobile hunter-gatherers who move their residential base often. Expedience, which entails less effort creating lithics with shorter use-lives, is associated with higher amounts of cortex, simpler platforms, less bifacial flaking, and lower levels of retouch; it is also associated with more sedentary hunter-gatherers. My results indicate that, during the Late Pleistocene in Belize, groups favored curation and were likely highly mobile foragers inhabiting an open grassland landscape. However, not long after the Pleistocene-ending Younger Dryas climatic event (~12,600 to 11,700 cal BP), these groups began to favor expedience, indicating they’d begun to adopt a more sedentary lifestyle. This shift towards expedience and sedentism continued throughout the Early and Middle Holocene, when closed-canopy tropical forests came to dominate the landscape. I conclude that these foragers began to adopt lower levels of residential mobility much earlier than generally thought. Because the sparse, unpredictable nature of wild tropical forest flora and fauna favors high residential mobility, this means they were likely manipulating their landscapes and experimenting with cultivars several millennia before the first appearance of sedentary farming villages. This further implies that the origins of sedentism and agriculture in tropical forests must be sought in the changing lifeways of pre-agricultural, semi-sedentary forager-horticulturalists of the deep past.
ContributorsDennehy, Timothy J. (Author) / Smith, Michael E. (Thesis advisor) / Barton, C. M. (Committee member) / Marean, Curtis W. (Committee member) / Prufer, Keith M. (Committee member) / Arizona State University (Publisher)
Created2021
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Description
Foraging has complex effects on whole-organism homeostasis, and there is considerable evidence that foraging behavior is influenced by both environmental factors (e.g., food availability, predation risk) and the physiological condition of an organism. The optimization of foraging behavior to balance costs and benefits is termed state-dependent foraging (SDF) while behavior

Foraging has complex effects on whole-organism homeostasis, and there is considerable evidence that foraging behavior is influenced by both environmental factors (e.g., food availability, predation risk) and the physiological condition of an organism. The optimization of foraging behavior to balance costs and benefits is termed state-dependent foraging (SDF) while behavior that seeks to protect assets of fitness is termed the asset protection principle (APP). A majority of studies examining SDF have focused on the role that energy balance has on the foraging of organisms with high metabolism and high energy demands ("high-energy systems" such as endotherms). In contrast, limited work has examined whether species with low energy use ("low-energy systems" such as vertebrate ectotherms) use an SDF strategy. Additionally, there is a paucity of evidence demonstrating how physiological and environmental factors other than energy balance influence foraging behavior (e.g. hydration state and free-standing water availability). Given these gaps in our understanding of SDF behavior and the APP, I examined the state-dependency and consequences of foraging in a low-energy system occupying a resource-limited environment - the Gila monster (Heloderma suspectum, Cope 1869). In contrast to what has been observed in a wide variety of taxa, I found that Gila monsters do not use a SDF strategy to manage their energy reserves and that Gila monsters do not defend their energetic assets. However, hydration state and free-standing water availability do affect foraging behavior of Gila monsters. Additionally, as Gila monsters become increasingly dehydrated, they reduce activity to defend hydration state. The SDF behavior of Gila monsters appears to be largely driven by the fact that Gila monsters must separately satisfy energy and water demands with food and free-standing water, respectively, in conjunction with the timescale within which Gila monsters balance their energy and water budgets (supra-annually versus annually, respectively). Given these findings, the impact of anticipated changes in temperature and rainfall patterns in the Sonoran Desert are most likely going to pose their greatest risks to Gila monsters through the direct and indirect effects on water balance.
ContributorsWright, Christian (Author) / Denardo, Dale F. (Thesis advisor) / Harrison, Jon (Committee member) / McGraw, Kevin (Committee member) / Sullivan, Brian (Committee member) / Wolf, Blair (Committee member) / Arizona State University (Publisher)
Created2014
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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
An insect society needs to share information about important resources in order to collectively exploit them. This task poses a dilemma if the colony must consider multiple resource types, such as food and nest sites. How does it allocate workers appropriately to each resource, and how does it adapt its

An insect society needs to share information about important resources in order to collectively exploit them. This task poses a dilemma if the colony must consider multiple resource types, such as food and nest sites. How does it allocate workers appropriately to each resource, and how does it adapt its recruitment communication to the specific needs of each resource type? In this dissertation, I investigate these questions in the ant Temnothorax rugatulus.

In Chapter 1, I summarize relevant past work on food and nest recruitment. Then I describe T. rugatulus and its recruitment behavior, tandem running, and I explain its suitability for these questions. In Chapter 2, I investigate whether food and nest recruiters behave differently. I report two novel behaviors used by recruiters during their interaction with nestmates. Food recruiters perform these behaviors more often than nest recruiters, suggesting that they convey information about target type. In Chapter 3, I investigate whether colonies respond to a tradeoff between foraging and emigration by allocating their workforce adaptively. I describe how colonies responded when I posed a tradeoff by manipulating colony need for food and shelter and presenting both resources simultaneously. Recruitment and visitation to each target partially matched the predictions of the tradeoff hypothesis. In Chapter 4, I address the tuned error hypothesis, which states that the error rate in recruitment is adaptively tuned to the patch area of the target. Food tandem leaders lost followers at a higher rate than nest tandem leaders. This supports the tuned error hypothesis, because food targets generally have larger patch areas than nest targets with small entrances.

This work shows that animal groups face tradeoffs as individual animals do. It also suggests that colonies spatially allocate their workforce according to resource type. Investigating recruitment for multiple resource types gives a better understanding of exploitation of each resource type, how colonies make collective decisions under conflicting goals, as well as how colonies manage the exploitation of multiple types of resources differently. This has implications for managing the health of economically important social insects such as honeybees or invasive fire ants.
ContributorsCho, John Yohan (Author) / Pratt, Stephen C (Thesis advisor) / Hölldobler, Bert (Committee member) / Liebig, Jürgen R (Committee member) / Amazeen, Polemnia G (Committee member) / Rutowski, Ronald L (Committee member) / Arizona State University (Publisher)
Created2019