Tradicionalmente, nichos populacionais são descritos como a somatória de todos os recursos utilizados por uma população. Entretanto, diversos estudos mostram que indivíduos dentro de uma população podem usar recursos de forma distinta. Investigamos três maneiras pelas quais indivíduos podem variar quanto ao uso do recurso. Indivíduos podem apresentar a mesma preferência por presas, mas diferir na propensão à adição de novos itens alimentares em sua dieta (Preferências Compartilhadas); indivíduos podem apresentar a mesma presa preferida mas diferirem em suas presas alternativas (Refúgio Competitivo); ou indivíduos podem apresentar presas preferidas distintas (Preferências Distintas). Estudamos os padrões de interação que emergem sob os pressupostos de cada um dos modelos usando redes de interação entre indivíduos e os recursos que eles consomem. Dessa forma, para derivarmos as previsões de cada um dos modelos de uso de recurso, desenvolvemos modelos simples que geram redes de interação segundo regras que seguem os pressupostos dos modelos e confrontamos essas previsões com dados empíricos, comparando a estrutura dessa redes de interação. Encontramos que o modelo que menos se assemelha ao padrão de uso de recurso observado para as populações estudadas foi o modelo de Preferências Compartilhadas. Para as populações estudadas, a variação intrapopulacional na escolha de presas parece estar mais associada a diferenças nas sequências de preferências por presas entre indivíduos e não à propensão desses indivíduos em adicionarem novos recursos às suas dietas. / Traditionally, a population\'s niche is described as the sum of all resources consumed by a population. However, several studies have highlighted that individuals within a population can use resources differently. We investigate three ways in which individuals can vary in their resource use. Individuals can show the same preference for prey, but differ in their likelihood of adding new prey to their diets (Shared Preferences); individuals can share the same top-ranked prey but differ in their alternative prey (Competitive Refuge); or individuals can have different top-ranked prey (Distinct Preferences). We studied the pattern of interaction that emerges under each model\'s assumption using interaction networks between individuals and the resources they consume. In this sense, to derive the predictions associated with each model of resource use, we developed simple models that generates interaction networks according to a set of rules that represent the assumptions of each model and then confronted these predictions with empirical data on interaction networks, by looking at the structure of these interaction networks. We found that the model that least resembles the pattern of resource use observed in the populations studied was the Shared Preferences model. For the studied populations, intrapopulation variation is not associated with individuals sharing the same rank sequence and differing in their willingness to add new resources to their diets. Instead, it seems that differences in the rank sequence of prey choice are more important in structuring the pattern of resource use in these populations.

Tradicionalmente, nichos populacionais são descritos como a somatória de todos os recursos utilizados por uma população. Entretanto, diversos estudos mostram que indivíduos dentro de uma população podem usar recursos de forma distinta. Investigamos três maneiras pelas quais indivíduos podem variar quanto ao uso do recurso. Indivíduos podem apresentar a mesma preferência por presas, mas diferir na propensão à adição de novos itens alimentares em sua dieta (Preferências Compartilhadas); indivíduos podem apresentar a mesma presa preferida mas diferirem em suas presas alternativas (Refúgio Competitivo); ou indivíduos podem apresentar presas preferidas distintas (Preferências Distintas). Estudamos os padrões de interação que emergem sob os pressupostos de cada um dos modelos usando redes de interação entre indivíduos e os recursos que eles consomem. Dessa forma, para derivarmos as previsões de cada um dos modelos de uso de recurso, desenvolvemos modelos simples que geram redes de interação segundo regras que seguem os pressupostos dos modelos e confrontamos essas previsões com dados empíricos, comparando a estrutura dessa redes de interação. Encontramos que o modelo que menos se assemelha ao padrão de uso de recurso observado para as populações estudadas foi o modelo de Preferências Compartilhadas. Para as populações estudadas, a variação intrapopulacional na escolha de presas parece estar mais associada a diferenças nas sequências de preferências por presas entre indivíduos e não à propensão desses indivíduos em adicionarem novos recursos às suas dietas. / Traditionally, a population\'s niche is described as the sum of all resources consumed by a population. However, several studies have highlighted that individuals within a population can use resources differently. We investigate three ways in which individuals can vary in their resource use. Individuals can show the same preference for prey, but differ in their likelihood of adding new prey to their diets (Shared Preferences); individuals can share the same top-ranked prey but differ in their alternative prey (Competitive Refuge); or individuals can have different top-ranked prey (Distinct Preferences). We studied the pattern of interaction that emerges under each model\'s assumption using interaction networks between individuals and the resources they consume. In this sense, to derive the predictions associated with each model of resource use, we developed simple models that generates interaction networks according to a set of rules that represent the assumptions of each model and then confronted these predictions with empirical data on interaction networks, by looking at the structure of these interaction networks. We found that the model that least resembles the pattern of resource use observed in the populations studied was the Shared Preferences model. For the studied populations, intrapopulation variation is not associated with individuals sharing the same rank sequence and differing in their willingness to add new resources to their diets. Instead, it seems that differences in the rank sequence of prey choice are more important in structuring the pattern of resource use in these populations.

Anthropogenic landscape changes threaten our ecologically and economically critical honey bees by decreasing the availability of quality foraging resources. Importantly, waggle dance analysis provides a versatile and relatively cost-effective tool for investigating the obstacles that honey bees face, such as habitat loss, in our changing landscapes. While this emerging tool has improved our understanding of honey bee foraging in specific landscape contexts, additional research is needed to identify broad trends that span across landscapes. For this dissertation, I used waggle dance decoding and analysis to investigate honey bee foraging, and sublethal stress responses, across three ecologically distinct landscapes in Virginia. In Chapter 1, I introduce waggle dances as a model study system for investigating honey bee foraging and sublethal stress responses by summarizing modern methodological advances in its analysis and emerging research gaps. In Chapter 2, I tested the effects of sublethal imidacloprid exposure on honey bee foraging and recruitment using a semi-field feeder experiment. In doing so, I report that honey bees decreased their foraging, but not recruitment, to an imidacloprid-laced sucrose solution, compared to a control solution. Together, these effects could potentially harm honey bee health by increasing their exposure to pesticides and decreasing their food intake. In Chapter 3, I compared the foraging distances communicated by waggle dancing nectar and pollen foragers across landscapes to explore the economic forces driving foraging to these resources. I observed higher overall and monthly nectar foraging distances compared to pollen foraging distances. Such results suggest that nectar foraging cost dynamics are driven by supply, while pollen foraging cost dynamics are driven by demand. In Chapter 4, I used waggle dance decoding to map and quantify foraging to agricultural grasslands in a mixed-use landscape. In doing so, I demonstrate that honey bees recruit to agricultural grasslands throughout the season, but that this land type was not more attractive than the broader landscape after correcting for foraging distance, which is a relevant cost that flying bees must consider. Additionally, I qualitatively observe a foraging hot spot, representing high honey bee interest, over a highly heterogenous section of the landscape. The collective results of this chapter identify agricultural grasslands as a potential management target and support the importance of landscape heterogeneity to honey bees/pollinators. In Chapter 5, I used waggle dance decoding to investigate honey bee foraging spatial patterns in the context of optimal foraging theory. In particular, I explore whether co-localized honey bee colonies forage optimally by converging on the same resource patches, or by partitioning the landscape in to distinct foraging territories. Spatial analysis revealed that the colonies widely distributed their foraging at the landscape-scale, with dances from the same and different colonies being similarly distributed, while also establishing distinct, patch-scale, colony-specific, foraging aggregations. Together, these results suggest that the honey bee foraging system produces an emergent foraging pattern that may decrease both within- and among-colony foraging competition. Finally, in Chapter 6, I place my research findings in the context of historical and current trends in honey bee behavioral ecology. Overall, my dissertation improves our understanding of honey bee foraging ecology across landscape contexts using waggle dance analysis, while demonstrating its versatility and effectiveness as a tool for ecologists. / Doctor of Philosophy / Honey bees collect nectar (carbohydrate source) and pollen (protein source) from flowers as their food for survival and reproduction. Human activities, such urbanization, change landscapes and threaten our critically important honey bees by decreasing the availability of flower-rich habitats. Importantly, honey bees share the location of good food sources with their nest mates using a communication behavior called the waggle dance. Interestingly, scientists can estimate the approximate location of the food sources communicated by waggle dancing bees through close observation and cutting-edge analysis. Therefore, we can "decode" honey bees' waggle dances to map their food collection, or foraging, patterns and investigate the obstacles that they face in our changing landscapes. For this dissertation, I used waggle dance decoding and analysis to investigate honey bee foraging across three different landscapes in Virginia. In Chapter 1, I introduce waggle dances as a tool for investigating honey bee behavior by summarizing the modern improvements in its analysis and areas where research is needed. In Chapter 2, I tested the effects of a sublethal exposure to a pesticide, imidacloprid, by observing the foraging and waggle dance behavior of bees visiting feeders with artificial food. I report that honey bees decreased their foraging, but not recruitment, while collecting an imidacloprid-laced sugar solution, compared to a solution without imidacloprid. In Chapter 3, I compared the foraging distances communicated by waggle dancing nectar and pollen foragers across landscapes to explore the economic forces driving foraging to these resources. I observed higher overall and monthly nectar foraging distances compared to pollen foraging distances. Such results suggest that nectar foraging is driven by supply, while pollen foraging is more driven by demand. In Chapter 4, I used waggle dance decoding to map and quantify foraging to agricultural grasslands (pastures and hay fields) in a landscape characterized by diverse land uses. In doing so, I demonstrate that honey bees recruit to agricultural grasslands throughout the season, but that this land type was not more attractive than the broader landscape after correcting for foraging distance. Additionally, I qualitatively observe a foraging hot spot, representing high honey bee interest, over a highly heterogenous section of the landscape. The collective results of this chapter identify agricultural grasslands as a potential management target and support the importance of landscape heterogeneity to honey bees/pollinators. In Chapter 5, I used waggle dance decoding to investigate the spatial patterns of honey bee foraging in the context of optimal foraging theory, which attempts to explain efficient resource collection strategies. In particular, I explore whether neighboring honey bee colonies forage optimally by converging on the same resource patches, or by dividing the landscape in to distinct foraging territories. We found that colonies distributed their foraging widely at the landscape-scale, with dances locations from the same and different colonies being similarly distributed, while also establishing distinct, patch-scale, colony-specific, foraging areas. Together, these results suggest that honey bees use a foraging strategy that decreases both within- and among-colony foraging competition. Finally, in Chapter 6, I place my research findings in the context of historical and current trends in honey bee behavioral ecology. Overall, my dissertation uses waggle dance analysis to improve our understanding of honey bee foraging behavior, while demonstrating its versatility and effectiveness as a tool for ecologists.

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Bumble bees (bombus spp.) are significant pollinators of many plants, and are particularly attracted to mass-flowering crops such as Oilseed Rape (Brassica Napus), which they cross-pollinate. B. napus is both wind and insect-pollinated, and whilst it has been found that wind is its most significant pollen vector, the influence of bumble bee pollination could be non-trivial when bee densities are large. Therefore, the assessment of pollinator-mediated cross-pollination events could be important when considering containment strategies of genetically modified (GM) crops, such as GM varieties of B. napus, but requires a landscape-scale understanding of pollinator movements, which is currently unknown for bumble bees. I developed an in silico model, entitled HARVEST, which simulates the foraging and consequential inter-patch movements of bumble bees. The model is based on principles from Reinforcement Learning and Individual Based Modelling, and uses a Linear Operator Learning Rule to guide agent learning. The model incoproates one or more agents, or bees, that learn by ‘trial-and-error’, with a gradual preference shown for patch choice actions that provide increased rewards. To validate the model, I verified its ability to replicate certain iconic patterns of bee-mediated gene flow, and assessed its accuracy in predicting the flower visits and inter-patch movement frequencies of real bees in a small-scale system. The model successfully replicated the iconic patterns, but failed to accurately predict outputs from the real system. It did, however, qualitatively replicate the high levels of inter-patch traffic found in the real small-scale system, and its quantitative discrepancies could likely be explained by inaccurate parameterisations. I also found that HARVEST bees are extremely efficient foragers, which agrees with evidence of powerful learning capabilities and risk-aversion in real bumble bees. When applying the model to the landscape-scale, HARVEST predicts that overall levels of bee-mediated gene flow are extremely low. Nonetheless, I identified an effective containment strategy in which a ‘shield’ comprised of sacrificed crops is placed between GM and conventional crop populations. This strategy could be useful for scenarios in which the tolerance for GM seed set is exceptionally low.

Whether species interactions are static or change over time has wide-reaching ecological and evolutionary consequences. However, species interaction networks are typically constructed from temporally aggregated interaction data, thereby implicitly assuming that interactions are fixed. This approach has advanced our understanding of communities, but it obscures the timescale at which interactions form (or dissolve) and the drivers and consequences of such dynamics. We address this knowledge gap by quantifying the within-season turnover of plant-pollinator interactions from weekly censuses across 3years in a subalpine ecosystem. Week-to-week turnover of interactions (1) was high, (2) followed a consistent seasonal progression in all years of study and (3) was dominated by interaction rewiring (the reassembly of interactions among species). Simulation models revealed that species' phenologies and relative abundances constrained both total interaction turnover and rewiring. Our findings reveal the diversity of species interactions that may be missed when the temporal dynamics of networks are ignored.

Optimal Foraging Theory (OFT) has its origin in processualistic ideas in 1960s with traces back to the dawn of the archaeological science in the 19th century. The OFT model is based on the construction of an individual’s food item selection understood as an evolutionary construct that maximizes the net energy gained per unit feeding time. The most common variants are diet patch choice, diet breadth/prey choice models and Marginal Value Theorem (MVT). The theory introduced experimental studies combined with mathematically data analyses and computer simulations. The results visualized in the experimental diagrammed curve are possible to compare with the archaeological records. What is “optimal” is an empirical question not possible to know but still useful as a benchmark for measuring culture. The theory is common in USA but still not in Europe. OFT seems to be useful in hunter-gatherer research looking at human decisions, energy flow, depression of resources and extinction. This literature review concludes that the prey-choice/diet-breadth model seems to be useful for hunter-gatherer research on Gotland focusing on possible causes of the hiatus in archaeological records between 5000-4500 BC. / Optimal Foraging Theory (OFT) har sitt ursprung i de processualistiska ideérna under 1960-talet med spår tillbaka till arkeologins början som vetenskap under 1800-talet. OFT modellen baseras på konstruktionen av en individs födoämnesval som förstås som en evolutionär konstruktion som maximerar nettoenergiintaget per tidsenhet som gått åt för försörjningen. De vanligaste varianterna är patch-choice, diet breadth/prey choice modellerna och Marginal Value Theorem (MVT). Experimentella studier genomförs och data bearbetas matematiskt och visar datorsimulerade kurvdiagram möjliga att jämföra med arkeologiska källmaterial. Vad som är ”optimalt” är en empirisk fråga omöjlig att veta men användbar ändå som en slag referens för att mäta kultur. Teorin är vanlig i USA men ännu inte i Europa. OFT förefaller användbar inom forskning av jägare-samlare om man fokuserar på beslutsfattande, energiflöde, depression av resurser och utrotning av arter. Slutsatsen i denna litteraturöversikt är att prey choice/diet breadth modellen tycks vara användbar för gotländsk jägare-samlare-forskning som fokuserar på möjliga orsaker till de arkeologiska fyndens hiatus mellan 5000-4500 BC.

This dissertation presents an evaluation of coprolite specimens from the Lower Pecos canyonlands as records of individual dietary decisions. Prior studies of coprolites from this region have greatly expanded our knowledge of Archaic subsistence patterns, but have not taken full advantage of the record of individual dietary decisions recorded in each coprolite specimen. The menu, or dietary combinations, reflected in individual coprolite specimens are assessed through the identification of several congruent botanical components derived from the same food resource, phytoliths, fiber ultimates, and epidermal sheets. The data is analyzed with hierarchical cluster analysis, an exploratory statistical technique. The resultant menus reflected in these clusters are evaluated with reference to the diet-breadth model developed for the known staple resources of the canyonlands as well as the seasonal subsistence patterns observed in the ethnohistoric record of modern-day Mexico and Texas. This same technique is also applied to the coprolite data available from previous studies in the Lower Pecos canyonlands. Overall, the combined dietary data available for the Lower Pecos canyonlands presents a similar dependence on desertic plant resources throughout the Archaic. Three main menus are apparent in the specimens. The first menu consists of prickly pear (Opuntia sp.) cladodes, or nopales, and was principally, although not exclusively, consumed in the late spring. This menu is primarily consumed when other resources were not readily available and may be considered a dependable but undesirable meal. The second menu consists of pit-baked lechuguilla (Agave lechuguilla) and sotol (Dasylirion sp.) caudices, or hearts, common throughout the cool season. This menu entails high processing costs, but would provide a reliable caloric return. The third menu exhibits a monolithic reliance on prickly pear fruits, or tunas, during the summer. The ease of harvest and consumption is reflected in the seasonal dominance of this resource, which was assuredly a highly desirable meal. The dietary patterns recorded in the coprolite specimens from the Lower Pecos canyonlands demonstrate a seasonally variable diet-breadth that incorporated low-ranked resources during times of seasonal scarcity as well as a monolithic dependence on high-ranked resources when they were available in the local landscape.

The Optimal Foraging Theory, rooted in the processual archaeology, uses a measuring methodology where the foraging strategy that gives the highest payoff measured as the highest ratio of energy gain per time unit is analysed (Mac Arthur & Pianca 1966, Emlen 1966). The theory is a branch of evolutionary ecology why much attention is paid to the interdependence of humans and preys and environmental conditions caused by climatologically and geographical changes or by overexploitation or other changes caused by humans. The analysis of Early Mesolithic pioneers onGotland, who settle in a transforming landscape, leaves indications of a Maglemose culture origin, probably from flooded original settlements in the south/southwest Baltic basin. The pioneers have to adapt to a seal-hunting economy dominated by grey seal which give the best cost-benefit outcome as big terrestrial mammals are missing and only mountain hare is available. The diet is narrow and there is a great risk for deficiency diseases as well as for acquiring hypervitaminosis and osteoporosis caused by excess of seal food. There is a hiatus c. 5000-4500 BC in the archaeological records on Gotland and the south-western Baltic region, and the master thesis hypothesises that Littorina Transgression I with a severe cold dip called the “8.2 ka BP cold event” has a delayed, but such a severe impact also on fauna and flora on Gotland, that the ecological system is destroyed. The possibility for humans to survive in a sustainable society is questionable. The extremely cold winters during this c. 400 years cold event, with glaciers moving southwards, delayed the blooming season, diminished the harvest and changed both flora and fauna. When the ecological niche for the grey seal is destroyed with flooded beaches close to the pioneers, human overexploitation is reinforced. With a diminishing population of mountain hare, which eventually gets extinct at the end of the Mesolithic, there are no alternatives but some birds and fish, hard to catch. Probably the pioneers abandon Gotland or move to a higher level on Gotland but no records are yet found why the period is called a hiatus. Extinction is the worst scenario or survival in such a small number that a sustainable society is lost. If so, new population groups repopulatedGotlandafter the Littorina transgressions. The origin is still unknown of the Pitted-ware and Funnelbeaker cultures that are populating Gotland after the transgressions. This master thesis can not confirm an affluent life style but rather a suffering starving society flooded by Littorina transgressions and struggling with the severe cold, caused by the “8.2 ka cold event” that makes the environmental conditions even worse. The subsistence economy is successively destroyed which probably causes the hiatus in archaeological records. The Littorina Transgression I with the “8.2 cold event” and the lack of terrestrial big animals are bottle necks. / Optimal Foraging Theory, med sina rötter i den processuella arkeologin, använder en metodik utgående från mätningar där insamlingsstrategin som ger den högsta avkastningen per tidsenhet analyseras (Mac Arthur & Pianca 1966, Emlen 1966). Teorin är en undergrupp inom den evolutionära ekologin och därför ägnas stor tid åt att uppmärksamma det ömsesidiga beroendet och påverkan som sker i miljön p.g.a. klimatologiska och geologiska orsaker men också p.g.a. mänsklig påverkan som exempelvis överförbrukning.  Analysen av tidigmesolitiska pionjärbosättare på Gotland, som möter ett landskap i förvandling, lämnar spår efter sig som tyder på ett ursprung i Maglemosekulturen i södra/sydvästra Östersjöregionen. De tvingas bli adapterade till en säljägarekonomi dominerad av gråsäl som ger det bästa energiutbytet, eftersom stora landdäggdjur saknas och endast bergshare finns tillgänglig. Dietvalet är smalt och det föreligger stor risk för både bristsjukdomar och A-vitaminförgiftning och osteoporos p.g.a. överkonsumtion av sälprodukter. Det finns ett uppehåll i de arkeologiska fynden c. 5000-4500 BC på Gotland liksom i södra Östersjöområdet. Magisteruppsatsens hypotes är att den kalla perioden med temperatursänkning som kallas ”8.2 ka BP cold event” under Littorinatransgression I har en fördröjd men så kraftigt övergripande effekt, på både djur- och växtliv på Gotland, att den förstör det ekologiska systemet och därmed möjligheten för människor att överleva i ett hållbart samhälle. De mycket hårda vintrarna under de c. 400 årens ”cold event” medför att glaciärerna dras sig söderut, blomningssäsongen fördröjs, skörden minskar och både fauna och flora förändras. När den ekologiska nischen för gråsälen förstörs av översvämmade stränder nära bosättarna förstärks överexploateringen, och då det inte finns någon alternativ föda utom en minskande harstam, svårfångade fåglar och fiskar, blir situationen fatal för de tidigmesolitiska bosättarna. Troligtvis flyttar de till andra platser inom Östersjönätverket eller till en högre nivå på Gotland, men fynd saknas hittills varför detta tomrum benämns ”hiatus”. Det värsta scenariot är att bosättarna dör ut eller överlever i ett så litet antal att det hållbara samhället går under.Om så är fallet återbefokas Gotland av gropkeramisk kultur och trattbägarkultur i anslutning till Littorinatrasgressionernas slut. Denna magisteruppsats kan inte konfirmera en livsstil i överflöd, utan snarare ett lidande svältande samhälle som översvämmas av Littorinatransgressioner med miljömässiga förhållanden som förvärras av den allvarliga kylan orsakad av ”8.2 ka cold event”.  Försörjningsmöjligheterna förstörs succesivt och befolkningen försvinner vilket troligen orsakar ett hiatus i de arkeologiska fynden. Littorina Transgression I med ”8.2 ka cold event” och bristen på stora landdjur är stora flaskhalsar.

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