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A sobrevivência e o sucesso reprodutivo de um indivíduo estão diretamente relacionados à sua capacidade de atender à demanda por nutrientes e energia. Estudos em Ecologia Nutricional, que adotam o Modelo Geométrico para nutrição, evidenciam que, mais do que maximizar a ingestão de energia, os animais procuram regular a ingestão de múltiplos nutrientes de maneira independente, almejando um balanço específico entre esses nutrientes. Neste trabalho, empregamos análises de conteúdo nutricional e o Modelo Geométrico para investigar os parâmetros nutricionais da dieta de duas populações de macacos-prego (Sapajus spp.), em dois biomas distintos: Sapajus libidinosus, numa área de ecótono Cerrado/Caatinga, a Fazenda Boa Vista (FBV), no Estado do Piauí; e Sapajus nigritus, numa área de Mata Atlântica, o Parque Estadual Carlos Botelho (PECB), no Estado de São Paulo. Estudos anteriores sugeriram que a qualidade e a disponibilidade dos alimentos explorados por S. libidinosus no semi-árido são superiores aos dos explorados por S. nigritus na Mata Atlântica. Porém, afirmações sobre a qualidade de dieta dependem de investigações sobre o balanço nutricional obtido pelo organismo e quais as estratégias nutricionais adotadas para atingir esse balanço. Assim, os objetivos desta tese foram (1) analisar as estratégias nutricionais de um grupo de Sapajus libidinosus na FBV e (2) comparar a qualidade nutricional da dieta dessa população com a da população de Sapajus nigritus do PECB. Foram analisadas amostras de 176 itens alimentares. Para estimar o consumo de nutrientes, foram realizados 134 focais contínuos diários em que foram registrados os itens alimentares consumidos pelo indivíduo e a quantidade ingerida. Os macacos da FBV regulam a ingestão de um balanço específico entre energia proteica e energia não proteica (advinda de lipídeos e/ou carboidratos), priorizando a ingestão de energia não proteica quando a disponibilidade desse nutriente nos itens alimentares diminui. Assumindo que o objetivo nutricional de S. nigritus é o mesmo de S. libidinosus, os resultados confirmaram a hipótese de que a qualidade da dieta na FBV é superior à do PECB, área em que os alimentos dos macacos apresentam menores concentrações de energia não proteica. O Modelo Geométrico para nutrição permitiu rever afirmações feitas em estudos anteriores de que na FBV a sazonalidade dos recursos mais consumidos não é um fator limitante para esses animais. Neste estudo, mostramos que há variações sazonais na qualidade nutricional dos alimentos, o que tem um impacto na ingestão de nutrientes pelos macacos da FBV / The survival and reproductive success of an individual are directly related to its ability to meet the demand for nutrients and energy. Studies in Nutritional Ecology, adopting the Geometric Framework for Nutrition, show that more than maximize energy intake, the animals seek to regulate the intake of multiple nutrients independently, targeting a specific balance between these nutrients. In this study, we used nutritional content analysis and the Geometric Framework to investigate the diet nutritional parameters of two populations of capuchin monkeys (Sapajus spp.) in two different biomes: Sapajus libidinosus in the ecotone Cerrado/Caatinga, at Fazenda Boa Vista (FBV) in Piauí state; and Sapajus nigritus in an Atlantic Forest area, at Parque Estadual Carlos Botelho (PECB), in São Paulo state. Previous studies suggest that the quality and availability of food exploited by S. libidinosus in the semi-arid region are higher than those of the exploited by S. nigritus in the Atlantic Forest. However, statements about diet quality depend on investigating the nutrient balance obtained by individuals and which nutrition strategies are adopted to achieve this balance. Therefore, the aims of this thesis were (1) to analyze the nutritional strategies of a group of Sapajus libidinosus in the FBV and (2) to compare the nutritional quality of the diet of this population with that of Sapajus nigritus in PECB. We analyzed samples of 176 food items eaten by the animals. To estimate nutrient daily intake, we conducted 134 continuous focal follows recording the item consumed and the amount ingested by individuals. The monkeys in FBV regulate the intake of a specific balance between protein energy and non-protein energy (lipids and / or carbohydrates), and prioritize non-protein energy intake when the availability of this nutrient in food items decreases. Assuming that the nutritional goal of S. nigritus is the same as S. libidinosus, the results confirmed the hypothesis that diet quality in FBV is superior than in PECB, where foods contain lower non-protein energy concentrations. The Geometric Framework allowed to review previous studies claims that in FBV seasonality of resources availability is not a limiting factor for these animals. In this study we have shown seasonal variations in the nutritional quality of foods, which impact nutrient intake by monkeys in FBV

A sobrevivência e o sucesso reprodutivo de um indivíduo estão diretamente relacionados à sua capacidade de atender à demanda por nutrientes e energia. Estudos em Ecologia Nutricional, que adotam o Modelo Geométrico para nutrição, evidenciam que, mais do que maximizar a ingestão de energia, os animais procuram regular a ingestão de múltiplos nutrientes de maneira independente, almejando um balanço específico entre esses nutrientes. Neste trabalho, empregamos análises de conteúdo nutricional e o Modelo Geométrico para investigar os parâmetros nutricionais da dieta de duas populações de macacos-prego (Sapajus spp.), em dois biomas distintos: Sapajus libidinosus, numa área de ecótono Cerrado/Caatinga, a Fazenda Boa Vista (FBV), no Estado do Piauí; e Sapajus nigritus, numa área de Mata Atlântica, o Parque Estadual Carlos Botelho (PECB), no Estado de São Paulo. Estudos anteriores sugeriram que a qualidade e a disponibilidade dos alimentos explorados por S. libidinosus no semi-árido são superiores aos dos explorados por S. nigritus na Mata Atlântica. Porém, afirmações sobre a qualidade de dieta dependem de investigações sobre o balanço nutricional obtido pelo organismo e quais as estratégias nutricionais adotadas para atingir esse balanço. Assim, os objetivos desta tese foram (1) analisar as estratégias nutricionais de um grupo de Sapajus libidinosus na FBV e (2) comparar a qualidade nutricional da dieta dessa população com a da população de Sapajus nigritus do PECB. Foram analisadas amostras de 176 itens alimentares. Para estimar o consumo de nutrientes, foram realizados 134 focais contínuos diários em que foram registrados os itens alimentares consumidos pelo indivíduo e a quantidade ingerida. Os macacos da FBV regulam a ingestão de um balanço específico entre energia proteica e energia não proteica (advinda de lipídeos e/ou carboidratos), priorizando a ingestão de energia não proteica quando a disponibilidade desse nutriente nos itens alimentares diminui. Assumindo que o objetivo nutricional de S. nigritus é o mesmo de S. libidinosus, os resultados confirmaram a hipótese de que a qualidade da dieta na FBV é superior à do PECB, área em que os alimentos dos macacos apresentam menores concentrações de energia não proteica. O Modelo Geométrico para nutrição permitiu rever afirmações feitas em estudos anteriores de que na FBV a sazonalidade dos recursos mais consumidos não é um fator limitante para esses animais. Neste estudo, mostramos que há variações sazonais na qualidade nutricional dos alimentos, o que tem um impacto na ingestão de nutrientes pelos macacos da FBV / The survival and reproductive success of an individual are directly related to its ability to meet the demand for nutrients and energy. Studies in Nutritional Ecology, adopting the Geometric Framework for Nutrition, show that more than maximize energy intake, the animals seek to regulate the intake of multiple nutrients independently, targeting a specific balance between these nutrients. In this study, we used nutritional content analysis and the Geometric Framework to investigate the diet nutritional parameters of two populations of capuchin monkeys (Sapajus spp.) in two different biomes: Sapajus libidinosus in the ecotone Cerrado/Caatinga, at Fazenda Boa Vista (FBV) in Piauí state; and Sapajus nigritus in an Atlantic Forest area, at Parque Estadual Carlos Botelho (PECB), in São Paulo state. Previous studies suggest that the quality and availability of food exploited by S. libidinosus in the semi-arid region are higher than those of the exploited by S. nigritus in the Atlantic Forest. However, statements about diet quality depend on investigating the nutrient balance obtained by individuals and which nutrition strategies are adopted to achieve this balance. Therefore, the aims of this thesis were (1) to analyze the nutritional strategies of a group of Sapajus libidinosus in the FBV and (2) to compare the nutritional quality of the diet of this population with that of Sapajus nigritus in PECB. We analyzed samples of 176 food items eaten by the animals. To estimate nutrient daily intake, we conducted 134 continuous focal follows recording the item consumed and the amount ingested by individuals. The monkeys in FBV regulate the intake of a specific balance between protein energy and non-protein energy (lipids and / or carbohydrates), and prioritize non-protein energy intake when the availability of this nutrient in food items decreases. Assuming that the nutritional goal of S. nigritus is the same as S. libidinosus, the results confirmed the hypothesis that diet quality in FBV is superior than in PECB, where foods contain lower non-protein energy concentrations. The Geometric Framework allowed to review previous studies claims that in FBV seasonality of resources availability is not a limiting factor for these animals. In this study we have shown seasonal variations in the nutritional quality of foods, which impact nutrient intake by monkeys in FBV

Non-consumptive effects that predators have on prey are important to ecosystems. The perceived risk of predation can alter feeding behavior. Giving-up density (GUD) experiments have been a foundational method to evaluate perceived predation risk, but rely on the assumption that food preferences are absolute. However, nutritional preferences are context dependent and can change with risk. In my first chapter, I used spiders and grasshoppers to test the hypothesis that covariance in nutritional preferences and risk may confound the interpretation of GUD experiments. My results demonstrate that predation risk and nutritional preferences covary and can confound interpretation of GUD experiments. In my second chapter, I use a behavioral observation experiment to further explore non-consumptive effects, as well as the movement of prey in response to predation risk.

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Master of Science / Department of Biology / Anthony Joern / Temperature and food quality vary across broad latitudinal gradients, greatly affecting performance by insect herbivores. The contribution of each varies latitudinally so that geographically distinct populations are challenged by differences in nutritional needs and energetic demands. While there has been extensive work studying diet selectivity and nutritional ecology of insect herbivores, few studies have focused on how insect herbivores adapt across such vast environmental gradients. The generalist-feeding grasshopper, Melanoplus femurrubrum (DeGreer), has a broad geographic range that extends across much of North America, making this species ideal for comparative investigations of intrinsic performance responses to extensive but predictable patterns of environmental variation. I compared responses by six populations collected from populations located from Texas to North Dakota (USA) using a common garden experimental design to investigate clinal responses in grasshopper performance. I examined responses in: (1) body size, (2) thermoregulation and adaptive coloration, (3) developmental and growth rates, (4) metabolic rates, (5) total consumption and rates, (6) diet ratio selection, and (7) digestive processing efficiencies across the latitudinal gradient. Grasshopper body size followed the Converse Bergmann’s Rule with decreasing body size as latitude increased. Temperature influenced all other responses, but responses to diet were not always significant or directional. Latitudinal trends for development and growth rates were observed but mass-specific metabolic rates were similar for all populations. Total consumption was body size dependent but independent of diet type. Mass-specific consumption varied but no single directional trend was detected. There was a shift in carbohydrate-biased diet preference at low latitude toward protein-biased diet ratios at higher latitudes, suggesting adaptations to different energetic demands by these populations. However, post-ingestive (digestive) efficiencies demonstrated variable responses with northern populations observing highest efficiencies for some indices but not all. Overall, this research documents phenotypic plasticity to environmental variability to some degree for digestive efficiencies, but ecotypic responses in body size and diet preference among M. femurrubrum populations were observed.

Ces dernières années ont vu le développement du commerce international entraînant une accélération des mouvements intercontinentaux d'espèces. Ces multiples introductions sont à l'origine de nombreux cas d'invasion, où les espèces s'établissent et se dispersent. Dans le nouvel environnement, étudier les facteurs liés à la niche écologique permet de comprendre comment se déroule l'établissement des espèces introduites au sein de la communauté d'espèces résidentes. L'hypothèse de la niche vide, qui prédit que l'invasion est possible car un espace vide existe au sein des niches écologiques de la communauté résidente, est un des facteurs facilitants de nombreux cas d'invasion. Par ailleurs, la stratégie d'exploitation de la ressource par l'espèce invasive au sein de la niche écologique peut aussi expliquer la propension d'une espèce à envahir ou non le nouvel environnement. L'objectif de cette thèse est d'explorer l'importance de ces hypothèses dans le succès invasif, au travers de l'exemple de Drosophila suzukii. Introduite en 2008 en Europe et aux Etats-Unis, cette espèce a envahi chacun des continents en moins de 7 ans. Elle est la seule espèce européenne de drosophiles capable de pondre à l'intérieur de fruits sains omniprésents, au sein d'une niche écologique vide de tout compétiteur. Elle est à l'origine de gros problèmes pour la culture de fruits cultivés et se retrouve être l'un des ravageurs majeurs en production de fruits rouges. Mieux comprendre son invasion, sa niche écologique et sa stratégie de sélection d'habitat permet de mieux lutter contre l'insecte, notamment par des méthodes permettant de perturber la localisation des fruits dans l'environnement. Cet aspect appliqué est aussi un objectif important de cette thèse CIFRE, financée pour optimiser la lutte par piégeage massif / In the last few decades, the development of international trade has seen resulting in an acceleration of intercontinental movements of species. These multiple introductions are the cause of many cases of invasion, where species are established and spread. In the new environment, studying factors related to the ecological niche help understanding what happens during the establishment of introduced species within the community resident species. The “empty niche hypothesis”, which predicted that the invasion is possible because a gap exists in the ecological niches of the resident community, is one of the facilitating factors of many cases of invasion. In addition, the strategy of utilization of the resource by the invasive species in the ecological niche may also explain the propensity of a species to invade or not the new environment. The objective of this thesis is to explore the significance of these assumptions in the invasive success, through the example of Drosophila suzukii. Introduced in 2008 in Europe and the US, this species has invaded every continent in less than 7 years. It is the only European species of fruit flies that can lay eggs inside healthy fruits, which are omnipresent, in an ecological niche free of any competitor. It is the cause of dreadful issues for grown fruit and is one of the major pests in production of berries. Better understand its invasion, its ecological niche and habitat selection strategy help better fighting the insect, including methods to disrupt the location of fruit in the environment. This applied context is also an important aspect of this thesis, funded also to optimize the fight by mass trapping

Na presente tese estudei a biologia de Centris (Paracentris) burgdorfi, uma abelha solitária de ampla distribuição no Brasil. O único local de nidificação conhecido até o momento fica em uma área de dunas, no nordeste do Brasil, à 25 km da cidade de Natal. Neste local as fêmeas nidificam exclusivamente em paleodunas à cerca de 1 km do mar. Estudei o sistema de acasalamento da espécie, onde pude constatar que a alta competição por fêmeas próxima ao ninho faz com que o macho tenha que retirar a fêmea do local para conseguir o contato genital. Nenhuma fêmea foi vista acasalando mais de uma vez e, logo após a cópula a fêmea dá início à construção do ninho. O ninho é constituído de um túnel raso com acesso às células de cria, construídas uma ao lado da outra. Avaliei o número de viagens que a fêmea faz para a coleta de óleo e pólen e o tempo despendido em cada ação dentro do ninho (deposição de pólen, óleo, ovoposição e operculação). Durante o período de construção das células as fêmeas não dormem dentro do ninho, e sim, em ramos de Krameria tomentosa, a planta utilizada como fonte de óleo no local. Buscando ampliar nosso conhecimento sobre a espécie, procurei outras populações no Brasil para que pudesse avaliar o nicho trófico. Utilizei o pólen encontrado no corpo das fêmeas para inferir sobre a dieta parcial de adultos e imaturos. Além da área de dunas, no Nordeste, encontramos C. burgdorfi no cerrado em Cavalcante, Goiás e em áreas de campo de altitude, em Ponta Grossa, Paraná. Através da análise polínica, constatei a importância de plantas do gênero Chamaecrista como fonte de pólen, sendo este um dos poucos gêneros compartilhados entre os três locais. Krameria grandiflora foi a fonte de óleo em Cavalcante e em Natal, e Angelonia integérrima, foi a fonte de óleo no Paraná. Isso revela a amplitude de nicho trófico em C. burgdorfi não apenas em relação às espécies de plantas utilizadas na dieta mas, também, quanto ao tipo de elaióforo que a fêmea consegue acessar (epitelial e tricromático). Esta plasticidade possibilita a ampla distribuição da espécie em diferentes fitofisionomias. Analisei nutricionalmente a dieta na provisão da cria de C. burgdorfi do nordeste do Brasil e de células de cria de Centris (Paracentris) pallida, coletadas no sudoeste dos Estados Unidos. Centris pallida é uma espécie de Centridini que perdeu o comportamento da coleta de óleo. Centris burgdorfi possui uma dieta mais rica em lipídeos e proteínas, sendo a dieta de C. pallida mais rica em carboidratos. A dieta mais protéica de C. burgdorfi se deve à presença de Chamaecrista já que plantas que anteras poricidas contem mais proteínas em comparação às não-poricidas, como é o caso das fontes de pólen de C. pallida (Parkinsonia e Olneya) / In the present thesis, I studied the biology of Centris (Paracentris) burgdorfi, a solitary bee of wide distribution in Brazil. The only nesting site known is in a dune area in northeastern Brazil, 25 km from the city of Natal. In this place, the females nest exclusively in petrified dunes about 1 km from the sea. I studied the mating system of the species, where I could verify that the high competition by females near the nest forces the male to carry the female to another place to mate. No female was seen mating more than once and, soon after copulation, the female begins to construct the nest. The nest consists of a shallow tunnel with access to the brood cells. I evaluated the number of trips that the female made to the collection of oil and pollen and the time spent in each action within the nest (depositions of pollen, oil, oviposition and operculation). During the period of construction of the cells the females do not sleep inside the nest, but in branches of Krameria tomentosa, the plant used as source of oil. In order to increase our knowledge about the species, I searched for other populations in Brazil so that I could evaluate the trophic niche. I used pollen found in the body of females to infer about the partial diet of adults and immature. In addition to the dune area in the Northeast, I found C. burgdorfi in the tropical Savanna (Cerrado) in Cavalcante, state of Goiás and in high altitude field areas, in Ponta Grossa, state of Paraná. Through pollen analysis, I verified the importance of plants of the genus Chamaecrista as a source of pollen, being this one of the few genera shared among the three sites. Krameria grandiflora was the source of oil in Cavalcante and in the Natal, and Angelonia integerrima, was the source of oil in the Ponta Grossa. This reveals the trophic niche amplitude in C. burgdorfi not only in relation to the species of plants used in the diet, but also to the type of elaiophore that the female can access (epithelial and trichromatic). This plasticity makes possible the wide distribution of the species in different phytophysiognomies. Nutritionally I analyzed the diet in the provision of C. burgdorfi offspring from northeastern Brazil and from the breeding cells of Centris (Paracentris) pallida collected in the southwestern United States. Centris pallida is a species of Centridini that has lost the behavior of the oil-collection. Centris burgdorfi has a diet richer in lipids and proteins, and the diet of C. pallida is richer in carbohydrates. The most proteinic diet of C. burgdorfi is due to the presence of Chamaecrista, since plants with poricidal anthers contain more proteins compared to non-poricidal ones, such as C. pallida (Parkinsonia and Olneya) pollen sources

Two closely-related, different-sized species of geese nest sympatrically south of the Queen Maud Gulf (QMG) in Canadas central arctic. Following a period of high population growth rate in both species within the QMG, the population growth rate of larger-bodied lesser snow geese (Chen caerulescens caerulescens; hereafter snow geese) has slowed most recently to roughly half that observed in smaller-bodied Rosss geese (Chen rossii). I focused on factors that influence productivity and recruitment in these two species, to improve our understanding of life history variation associated with interspecific differences in body size, and to test for density-dependent population responses. I used long-term data (1991 to 2008) to compare spring nutrient reserves, breeding strategies, clutch sizes, nest success, and juvenile survival in Rosss and snow geese breeding at Karrak Lake, Nunavut; a large breeding colony located within the QMG.<p> Long-term patterns of spring body condition (i.e., fat and protein reserves) diverged in prospective breeding female Rosss and snow geese implying that differences in food acquisition ability had become more acute. Snow geese displayed larger reductions in protein and fat reserves through time compared to Rosss geese thereby suggesting a differential density-dependent response in the ability to store nutrient reserves, a prerequisite for breeding in both species. Decreased per capita food availability influenced the timing of reproduction in both species. Nesting phenologies of Rosss and snow geese, adjusted for variation in phenology of local spring climate, have become later by 6.5 and 5.0 days, respectively, since 1991. Nutritional strategies (i.e., reliance on reserves versus local food) used for clutch formation differed between species. Rosss geese displayed greater reliance on stored reserves (i.e., capital breeding) than did snow geese, though both used endogenous reserves (> 62% of yolk protein, > 48% of albumen, and > 73% of yolk lipid) for clutch formation. Rosss and snow geese experienced declines of 28% and 23% in body masses from arrival to post-laying and also until hatch demonstrating that endogenous reserves are the main nutrient sources for incubation. Still, constraints of small size forced Rosss geese to use a mixture of local food plants and reserves for incubation metabolism.<p> I then examined differences in clutch size, nest success, and juvenile survival to understand of the role of recruitment in the interspecific divergence of population trajectories. I did not find strong interspecific differences in clutch size and nest success. Overall, snow geese had a larger mean clutch size, which was expected based on benefits of a larger-body size. Clutch sizes decreased with delays in breeding and decreasing protein reserves of arriving females, although Rosss geese displayed larger declines with decreasing protein reserves. Mean apparent nest success for Rosss geese was 4.5% higher compared to snow geese. Nest success showed large declines (11%) in both species with increasing population size at the breeding colony. However, nest success of snow geese decreased twice as fast with delays in breeding compared to Rosss geese. Last, I found no evidence of negative density dependence in juvenile survival over time. Juvenile survival was higher in snow geese (48%) compared to Rosss geese (38%), consistent with a life history prediction based on body size differences. Despite lower juvenile survival, recruitment by Rosss geese is likely greater than that of snow geese because of earlier sexual maturity, higher breeding probability and/or greater nest success.<p> Ultimately, small body size of Rosss geese may produce an ideal life history schedule under resource limitation at this colony i.e., one that maximizes fitness compared to larger snow geese. Life history characteristics of Rosss geese (e.g., absolutely lower energy requirement, have a flexible breeding strategy, higher reproductive effort, an earlier age of sexual maturity, a shorter breeding cycle allowing delayed arrival and nest initiation on arctic breeding areas, and shorter time required by goslings to attain adult size), in addition to their smaller bill morphology may allow exploitation of a wider niche space (i.e., one that includes marginal quality and low quantity vegetation) relative to snow geese. Because there were no large differences in components of recruitment considered here, other components of recruitment (age of sexual maturity, breeding probability) may be affected more strongly by diminished spring nutrition in snow geese and thus have a larger influence on local population dynamics.