It is generally accepted that the Marginal Value Theorem (MVT) describes optimal foraging strategies. Some research findings, however, indicate that in natural conditions foragers not always behave according to the MVT. To address this inconsistency, in a series of computer simulations, I examined the behaviour of four types of foragers having specific foraging efficiencies and using the MVT and alternative strategies in 16 simulated landscapes in an ideal environment (no intra- and inter-species interactions). I used data on elk (Cervus elaphus) to construct the virtual forager. Contrary to the widely accepted understanding of the MVT, I found that in environments with the same average patch quality and varying average travel times between patches, patch residence times of some foragers were not affected by travel times. I propose a mechanism responsible for this observation and formulate the perfect forager theorem (PFT). I also introduce the concepts of a foraging coefficient (F) and foragers hub (), and formulate a model to describe the relationship between the perfect forager and other forager types. I identify situations where a forager aiming to choose an optimal foraging strategy and maximize its cumulative consumption should not follow the MVT. I describe these situations in a form of a mathematical model. I also demonstrate that the lack of biological realism and environmental noise are not required to explain the deviations from the MVT observed in field research, and explain the importance of scale in optimal foraging behaviour. I also demonstrate that smart foraging, which is a set of rules based on key ecological concepts: the functional response curve (FRC), satisficing, the MVT, and incorporates time limitations, should allow for fitness maximization. Thus, it should be an optimal behavior in the context of natural selection. I also demonstrate the importance of the FRC as a driver for foraging behaviors and argue that animals should focus more on increasing the slope of their FRC than on choosing a specific foraging strategy. Natural selection should, therefore, favor foragers with steep FRC. My findings introduce new concepts in behavioural ecology, have implications for animal ecology and inform wildlife management.

The New Zealand fur seal (Arctocephalus forsteri) is the most abundant fur seal species in the Australian-New Zealand region. Approximately 85 % of Australia’s population of New Zealand fur seals reside in the state of South Australia. As a result of their abundance and size, it has been estimated that the New Zealand fur seal population in South Australia consumes the greatest biomass of resources of all marine mammal and seabird species. However, despite the importance of New Zealand fur seals as top predators, our understanding of their foraging ecology in South Australia is limited. In order to better understand the habitat utilized and the diet of New Zealand fur seals, this study explores the foraging ecology of lactating seals from four primary colonies in South Australia, which account for ~ 78 % of the Australian population. These colonies are Cape Gantheaume (36о04’S, 137о27’E) and Cape du Couedic (36о03’S, 136о42’E) on Kangaroo Island; North Neptune Island (35о13’S, 136о03’E) and Liguanea Island (34о59’S, 135о37’E). I start this study by assessing the seasonal variation in foraging location and dive behaviour of lactating New Zealand fur seals from Cape Gantheaume. 18 seals were fitted with satellite transmitters and time depth recorders (TDRs). The presence of thermoclines (derived from TDRs), were used as a surrogate measure of upwelling activity in continental shelf habitats. During the austral autumn 80 % of lactating fur seals foraged on the continental shelf (114 ± 44 km from the colony), in a region associated with a seasonal coastal upwelling system, the Bonney upwelling. In contrast, during winter months seals predominantly foraged in oceanic waters (62 %), in a region associated with the Subtropical Front (460 ± 138 km from the colony). Results suggested that lactating New Zealand fur seals shift their foraging location from continental shelf to oceanic habitats, in response to a seasonal decline in continental shelf productivity, attributed to the cessation of the Bonney upwelling in autumn. To study inter-colony differences in foraging locations, 21 New Zealand fur seals were satellite tracked from four colonies within close proximity (46 km – 200km apart). Seals initiated foraging trips on a colony-specific bearing (Cape Gantheaume 141 ± 33º, Cape du Couedic 186 ± 12º, North Neptune Island 200 ± 23º and Liguanea Island 234 ± 69º), and recorded little overlap between colony-specific foraging areas. The distribution of colony-specific foraging grounds appeared to be influenced by the proximity of colonies to predictable local upwelling features, as well as a distant oceanic frontal zone, the Subtropical Front. Foraging site fidelity and route-choice was further assessed by comparing site fidelity between continental shelf and oceanic habitats. Data from 31 lactating females, satellite tracked over 107 consecutive foraging trips indicated that females foraging on the continental shelf recorded a significantly greater overlap in foraging area between consecutive foraging routes, when compared to females that foraged in oceanic waters (55.9 ± 20.4 % and 13.4 ± 7.6 %, respectively). Findings suggest that seals learn the direction of travel to a predictable foraging region, and initiate a foraging trip on that bearing. However, actual foraging routes are likely to be influenced by a number of factors including previous foraging trip experience and prey encounter rate, which is related to prey density and the spatial scale of the patch exploited. The final chapter integrates scat analysis with milk fatty acid (FA) analysis to investigate dietary differences between continental shelf and oceanic waters. Milk FA composition was determined for 29 satellite-tracked fur seals, that were known to forage in either shelf or oceanic habitats. Based on FA compositions, I predicted the likelihood that milk samples collected at random (n = 131) represented individual seals having foraged either on the continental shelf or in distant oceanic waters. FA analysis and satellite tracking results contrasted with scat analyses, from which only 6 % of scats by frequency of occurrence contained prey remains from oceanic waters. The results suggest that scats were biased toward females foraging on the continental shelf. This study highlights the importance of two predictable ocean features utilised by New Zealand fur seals; (1) a nearby and seasonally predictable coastal upwelling system, the Bonney upwelling and; (2) a distant but permanent oceanic front, the Subtropical Front. / http://proxy.library.adelaide.edu.au/login?url= http://library.adelaide.edu.au/cgi-bin/Pwebrecon.cgi?BBID=1347312 / Thesis (Ph.D.) - University of Adelaide, School of Earth and Environmental Studies, 2008

The question of group-living in animals has attracted a lot of research attention. Today, we have a nuanced understanding of ultimate explanations and proximate mechanisms for group-living, as well as the genetic basis for seemingly puzzling aspects of sociality such as altruism. However, our current view of sociality has been obtained almost entirely in an intraspecific context, i.e. based on groups consisting of individuals of a single species. On the other hand, heterospecific sociality, i.e. groups formed by individuals of multiple species, has been largely ignored in group-living theory and empirical research. This is despite the fact that heterospecific sociality is widely prevalent in nature and is known to occur in many different contexts, in a variety of taxa and in both marine and terrestrial habitats. My dissertation focuses on one of the best-known examples of heterospecific sociality, namely heterospecific foraging associations of terrestrial insectivorous birds (commonly called mixed-species flocks; flocks hereon). Flocks are found all over the world, in a variety of habitats, and include a unique suite of species in each area. Although flocks are the best-studied among different types of heterospecific associations known, most of the research has been descriptive. In other words, while we have descriptions of the richness, size and composition of flocks from many areas, we know little about why and how flocks form. Therefore, one broad objective of this dissertation was to better understand the causes of flock formation. Birds are thought to join flocks either to obtain direct foraging benefits or better protection from predators. I used different approaches to understand which of these two reasons is likely to be more important. Additionally, to situate heterospecific sociality within sociality in general, I also examined whether the mechanisms underlying flocks are similar to single-species groups or unique to a mixed-species condition. In other words, are flocks a different route to obtain the same benefits as single-species groups, or a route to obtain benefits that conspecifics cannot provide? In the previous section, I described why understanding flocks is important from the point of view of sociality. Flocks are also important for the consequences they might have on community organization. While each flock is a collection of interacting individuals of different species, populations of species are linked in a network of interactions across multiple flocks in an area. The theoretical framework guiding our understanding of community organization is currently dominated by the idea of interspecific competition. Mutualisms or positive interactions, if any, are only expected to occur between members of different trophic levels. Only recently has the need to incorporate positive interactions between members of the same trophic level into ecological theory been emphasized. Flocks and other heterospecific associations present an important example of within-trophic level positive interactions. Therefore, the second broad objective of this dissertation was to examine the consequences of flock formation on community organization. The objectives described above were addressed using multiple approaches in this dissertation. “Why” questions in ecology and evolution are generally tackled using manipulative experiments. Given that experiments are neither feasible nor ethical in the case of multispecies flocks in the wild, I addressed the “why” question by focusing on “who” instead. To understand “why” flocks form, I examined what kinds of species participate in flocks, who associates with whom, and who provides and who receives the benefits. This dissertation uses a combination of methods and data, including questions addressed at a global scale using descriptive information on flocks available from across the world. For other questions that were based on entirely new approaches developed in this dissertation, data were collected in a field site in the Western Ghats. In general, given the two broad objectives of this dissertation, the approaches I used were drawn from both behavioural and community ecology. The dissertation is organized in the following way: Chapter 1: General Introduction The first chapter provides the background to the two broad objectives of this thesis, namely understanding the causes and consequences of heterospecific foraging associations in terrestrial bird communities. Chapter 2 (published in The American Naturalist) Based on a global dataset (55 presence-absence matrices from 24 locations in multiple continents) on the composition of flocks, I asked if flocks largely consist of ecologically similar or dissimilar species. Using null models and randomization tests followed by meta-analysis, I found that the association strength of species in flocks was strongly related to similarity in body size and foraging behaviour, and higher for congeneric compared with non-congeneric species pairs. In other words, flocks seem to consist largely of similar species. Extending group-living and social information use theory to a heterospecific context, I discuss potential behavioral mechanisms leading to positive interactions among similar species in flocks as well as ways in which competition costs are reduced. These findings highlight the need to consider positive interactions along with competition when seeking to explain community organisation. Chapter 3 (published in Animal Behaviour) Two kinds of participants are recognized in flocks: those that join other species (‘followers’) and are therefore likely to be the recipients of the benefits of flock participation and those that are joined (‘leaders’). Through comparative analyses, using a large sample of flocks from around the world, I found (1) ‘followers’ tend to be smaller, more insectivorous, and feed in higher strata than matched species that participate in flocks to a lesser extent and (2) ‘leaders’ tend to be cooperative breeders (which are known to have well-developed anti-predatory systems) more often than matched species that are not known to lead flocks. Furthermore, meta-analyses of published results from across the world showed that bird species in terrestrial mixed-species flocks increased foraging rates and reduced vigilance compared to when they were solitary or in conspecific groups. Moreover, the increase in foraging rates was seen only in the case of flock followers and not flock leaders. These findings suggest a role for predation in the evolution of mixed-species flocking. Species that are vulnerable to predation follow species whose vigilance they can exploit. By doing so, they are able to reduce their own vigilance and forage at higher rates. Chapter 4 (to be submitted to Behavioural Ecology and Sociobiology) In this study, conducted in a tropical evergreen forest in the Western Ghats of India, I used intraflock association patterns to generate a community-wide assessment of benefits of flock participation for different species. I assumed that individuals needed to be physically proximate to particular heterospecific individuals within flocks to obtain any direct foraging benefit (flushed prey, kleptoparasitism, copying foraging locations). Alternatively, for all anti-predation benefits, physical proximity to particular heterospecifics is not required, i.e. just being in the flock vicinity will suffice. Therefore, I used the choice of locations within flocks to infer whether individual species are obtaining direct foraging or antipredation benefits. A small subset of the bird community (5/29 species), composed of all members of the sallying guild, showed non-random physical proximity to heterospecifics within flocks. All preferred associates were from non-sallying guilds, suggesting that the sallying species were likely obtaining direct foraging benefits, either in the form of flushed or snatched prey. The majority of species (24/29) chose locations randomly with respect to heterospecifics within flocks, and thus were likely obtaining anti-predation benefits. In summary, my study indicates that direct foraging benefits are important for only a small proportion of species in flocks and therefore that predation is likely to be the main driver of flocking. Chapter 5 (to be submitted to Oecologia) Two types of species – intraspecifically gregarious and sallying species – are thought to play important roles in flocks because studies have shown they attract other flock participants. However, it is not clear why these types of species are attractive, i.e. are they cues for flocks or do they directly provide benefits to other species? It is also not known whether these types are essential for flock formation. In this study, in a tropical evergreen forest in the Western Ghats of India, I used a novel approach to address these questions. Flocks contain anywhere between two and tens of species. My approach focused on the simplest, i.e. two-species, flocks. In two-species flocks, attraction between species must be based on direct benefit to at least one of the species. Therefore, only species combinations that result in benefit to at least one species will occur as two species flocks. Further, by examining leading and following behaviour in these combinations, I could also determine the direction of benefit-flow. Finally, given that all flocks pass through a two-species step, examining which two-species combinations are joined by other species helped understand species roles in flock formation. I found that intraspecifically gregarious species, but not sallying species, were disproportionately represented in two-species flocks, always provided the benefits when present, and that flocks containing them were joined significantly more often. Therefore, intraspecifically gregarious species, but not sallying species, play a role in providing flock benefits and flock formation. Moreover, given that most (7/8) intraspecifically gregarious species in my study site played these roles, the intraspecifically gregarious species guild can be considered an example of a within-trophic level “keystone”. Chapter 6 (“in press” in Oikos) There is a growing recognition of the need to integrate non-trophic interactions into ecological networks for a better understanding of whole-community organization. To achieve this, the first step is to build networks of individual non-trophic interactions. In this study, I analyzed the network of interdependencies among bird species that participated in flocks in an evergreen forest site in the Western Ghats, India. I found that flock networks contain a small core of highly important species that other species are strongly dependent on, a pattern seen in many other biological networks. Further, I found that structural importance of species in the network was strongly correlated to functional importance of species at the individual flock level. Finally, comparisons with flock networks from other Asian forests showed that the same taxonomic groups were important in general, suggesting that species importance was an intrinsic trait and not dependent on local ecological conditions. Hence, given a list of species in an area, it may be possible to predict which ones are likely to be important. Chapter 7: Conclusions In this section, I provide a summary of the main findings of this dissertation, discuss the main conclusions with regard to each of the two broad objectives and finally suggest future lines of investigation to further understand the causes and consequences of flock formation and heterospecific sociality in general. In summary, the work presented in this dissertation provides a picture of how and why flocks form and their likely consequences for community organization. The main driver of flock formation seems to be protection from predators, although a few species do obtain direct foraging benefits. Additionally, given that I found flocks to largely be groupings of similar species, the mechanisms through which benefits are obtained are likely to be similar to those in single-species groups. In other words, flocks are possibly a way for species that are unable to group with conspecifics, to obtain group-living benefits. My finding, that flocks largely consisted of similar species, also questions the stereotyping of interactions between similar species in communities as competitive. Other results from this study demonstrate the importance of intraspecifically gregarious species in flocks. Such species are the main providers of benefits to other flock participants and seem essential for flock formation. Finally, network analysis showed that a small core of species, mainly including species known to provide benefits at the individual flock level, are disproportionately important in the emergent community-level network of interdependencies. Such species are possibly one of the first examples of within-trophic level “keystones” discovered. Apart from the findings, this study also provides a set of new approaches and analytical frameworks that can be used to examine other multispecies foraging groups and heterospecific foraging associations in other contexts.

La variation des comportements sociaux est une problématique très importante pour appréhender l’évolution de la socialité dans le règne animal. Par l’étude des interactions entre les paramètres génétiques et environnementaux sur les comportements sociaux de la drosophile (Drosophila melanogaster), cette thèse s’inscrit dans cette problématique et cherche à mettre en évidence les paramètres déterminants pour la expliquer la variation des comportements sociaux des individus. Au cours de ce travail de recherche, on a pu étudier et mettre en évidence l’existence de comportements sociaux chez la drosophile tels que l’apprentissage et l’utilisation des informations sociales dans un contexte de recherche spatiale, le comportement d’agrégation et l’établissement de nombreuses interactions. Ces comportements sont les premiers pas vers plus de socialité et leur étude est indispensable pour comprendre l’évolution vers plus de coopération et de communication entre les individus. L’utilisation de deux lignées issues du polymorphisme génétique du gène foraging (phénotypes Rover et Sitter) a permis de mettre en évidence des différences de comportements entre les individus et d’étudier les interactions entre paramètres génétiques et environnementaux. Les individus de phénotype Sitter témoignent d’un comportement plus social que les individus Rover, utilisant préférentiellement l’information sociale, ils forment aussi des agrégats plus importants semblant attirés par le plus grand nombre. On note aussi que la variation de la composition génétique du groupe entraine une modification du comportement d’agrégation du groupe. Ainsi, les variations interindividuelles influencent le comportement collectif. Le dernier volet de cette thèse s’intéresse aux interactions entre les individus au sein de 12 populations différentes de l’espèce Drosophila melanogaster. Cette approche a permis de révéler une grande variation de l’organisation sociale des individus au sein d’une même espèce et ouvre de nouvelles perspectives pour étudier l’évolution de la socialité.Ce travail de recherche permet d’apporter de nouveaux éléments sur les facteurs influençant la variation des comportements sociaux et de mettre en avant de nouvelles perspectives pour l’étude de leur évolution. / The social behavior variation is a very important issue for understanding evolution of sociality in animal kingdom. By studying genetic and environmental interactions influence on social behavior in Drosophila melanogaster, this thesis fits with this problematic and focus on the parameters which could explain the social behavior variations. In this research work, we studied and highlighted the existence of social behavior in Drosophila like learning and using social information in a spatial learning task, aggregation behavior and numerous social interactions. These behaviors are the first step before more sociality and their study with no-social species is essential to understand the evolution toward more sociality. The use of two lines from genetic polymorphism of the foraging gene (Rover and Sitter phenotypes) reveals differences in behavior between individuals and enable us to study the influence of genetic and environment parameters. Sitter individuals show more social behavior than Rover ones, using more social information and adopting more aggregation behavior. Variation of the genetic composition of the group influence the aggregation level of Rover individuals. They show more gregarious behavior with majority of Sitter individuals. These observations raise the question of the effects of the information source variations on the social behavior and their implication in interactions variations. The last part of this thesis focuses on the interactions between individuals in 12 different populations of Drosophila melanogaster. This approach revealed an important variation in social organization within species and opens new perspectives to study the evolution of sociality.This research bring new elements on factors influencing the change in social behavior and highlight new perspectives for the study of their evolution.

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / A range of behavioral strategies and sensory abilities allow animals to minimize costs involved in the search for food. Among the factors involved in the variation of foraging costs, the availability of resources represents a central role and it is recognized for modulating the animal home range. Some species of termites can exhibit characteristics that make the foraging process even more expensive. Species belonging to the genus Nasutitermes, for example, have an extra cost in foraging: in addition to build a network of tunnels to access resources, they have a large number of soldiers, a caste trophically dependent. In the present study: (i) we evaluated the foraging pattern in Nasutitermes aff. coxipoensis (Termitidae: Nasutitermitinae), including the searching strategy and the role of soldiers; and (ii) we analyzed whether colonies of this species responds to variations in the availability of food resources. Manipulative experiments were conducted in the field and in the laboratory to analyze the pattern of foraging in N. aff. coxipoensis. For this, 35 nests were transplanted into areas of dunes in Pirambu-SE. Seven plots were established in a continuous increament in the density of resources (sugarcane baits). Laboratory tests were conducted to examine the role of chemical signals (eg. trail pheromone) present in the sternal gland of workers and soldiers. During 10 consecutive days it was measured: the number of trails and tunnels, their total length, number of branches and the tunneling speed (cm/day). Data were analyzed using generalized linear models and mixed linear models. Nasutitermes aff. coxipoensis showed mainly nocturnal foraging. Soldiers were the first individuals to start the foraging, however, in established trails, the number of workers were always higher than soldiers. The number of active trails remained constant over the observation period, while the number of tunnels increased. In groups of soldiers and workers, the workers chose to follow the trail signals of soldiers. The number of trails, the total length and their branches decreased with increment of resource availability. The conversion of trails in tunnels increased in areas with higher resource density. Our results suggest that the costs involved in the production of soldiers in N. aff. coxipoensis seem to be compensated by their decisive role during the foraging process. Colonies of this species seem to optimize their foraging through a combined strategy of constructing trails and tunnels. In places with low resource availability, termites forage mainly on trails, avoiding to convert trails in tunnels. This strategy has not yet been reported in the literature. Thus, N. aff. coxipoensis presents strategies in order to optimize the foraging, minimizing costs involved in this process. These results can contribute to the key question about the evolution of termite foraging behavior, as well as to understand the mechanisms involved in the distribution patterns and structuring communities of these insects. / Uma gama de estratégias comportamentais e habilidades sensoriais permite aos animais minimizar os custos envolvidos na busca por alimento. Dentre os fatores envolvidos na variação dos custos de forrageio, a disponibilidade de recursos representa um papel central e é reconhecida por modular as áreas de uso dos animais. Algumas espécies de cupins exibem características que podem tornar o processo de forrageio ainda mais oneroso. Espécies do gênero Nasutitermes, por exemplo, apresentam gastos extras no forrageio, uma vez que além de construírem uma rede de túneis para acessar os recursos, ainda apresentam um elevado número de soldados, indivíduos troficamente dependentes. No presente estudo, avaliamos (i) o padrão de forrageio em Nasutitermes aff. coxipoensis (Termitidae: Nasutitermitinae), incluindo as estratégias de busca e o papel dos soldados; e (ii) analisamos se colônias desta espécie respondem à variação na disponibilidade de recursos. Experimentos manipulativos foram conduzidos em campo e em laboratório a fim de analisar o padrão de forrageio de N. aff. coxipoensis. Para isso, 35 ninhos foram transplantados em áreas de dunas, em Pirambu- SE. Foram estabelecidas sete parcelas em um contínuo de aumento da densidade de recursos (iscas de cana-de-açúcar). Testes em laboratório foram realizados a fim de analisar o papel dos sinais químicos (ex. feromônio de trilha) presente na glândula esternal de operários e soldados. Durante 10 dias consecutivos foram quantificados: o número de trilhas e de túneis, o comprimento total, o número de ramificações destes e a velocidade de construção dos túneis. Os dados foram analisados utilizando-se modelos lineares generalizados e modelos mistos. Nasutitermes aff. coxipoensis apresentou forrageio principalmente noturno. Soldados foram os primeiros indivíduos a iniciarem o forrageio, no entanto, em trilhas já estabelecidas, o número de operários foi sempre maior do que o de soldados. O número de trilhas ativas permaneceu constante ao longo do período de observação, enquanto o número de túneis aumentou de forma gradativa. Em grupos compostos por soldados e operários, os operários preferiram seguir os sinais de trilha dos soldados. O número de trilhas, o comprimento total e suas ramificações reduziram com o aumento da disponibilidade de recursos. A conversão de trilhas em túneis aumentou em locais com maior densidade de recursos. Nossos resultados sugerem que os custos envolvidos na produção de soldados de Nasutitermes aff. coxipoensis parecem ser compensados pelo seu papel decisivo no forrageio. Colônias desta espécie parecem otimizar o forrageio através de uma estratégia combinada de formação de trilhas e túneis. Em locais com baixa disponibilidade de recursos os cupins forrageiam preferencialmente em trilhas, evitando a conversão destas em túneis. Tal estratégia, ainda não foi relatada na literatura. Assim, N. aff. coxipoensis apresenta estratégias para otimização do forrageio, minimizando os custos envolvidos nesse processo. Os resultados deste estudo podem contribuir para a investigação de questões sobre evolução do comportamento de forrageio de cupins, assim como para desvendar os mecanismos envolvidos nos padrões de distribuição e estruturação de comunidades desses insetos.

Submitted by Fabio Sobreira Campos da Costa (fabio.sobreira@ufpe.br) on 2016-06-14T13:27:03Z No. of bitstreams: 2 license_rdf: 1232 bytes, checksum: 66e71c371cc565284e70f40736c94386 (MD5) Hugo_Andrade_Doutorado.pdf: 3065927 bytes, checksum: 2eeb9c1ecb93e60c146992117b01cbb6 (MD5) / Made available in DSpace on 2016-06-14T13:27:03Z (GMT). No. of bitstreams: 2 license_rdf: 1232 bytes, checksum: 66e71c371cc565284e70f40736c94386 (MD5) Hugo_Andrade_Doutorado.pdf: 3065927 bytes, checksum: 2eeb9c1ecb93e60c146992117b01cbb6 (MD5) Previous issue date: 2013-02-07 / CNPq / Nesta Dissertac¸ ˜ao estudamos a dinˆamica energ´etica das buscas aleat ´orias aplicadas ao problema de foraging, em que animais buscam por comida ou parceiros em ambientes escassos. Discutiremos, inicialmente, um modelo estat´ıstico de caminhadas aleat ´orias utilizando as distribuic¸ ˜oes de L´evy para os tamanhos dos passos de busca, as quais tˆem sido reportadas na literatura como estrat´egias de eficiˆencia ´otima para o problema. Em seguida vamos incluir no modelo ganhos e perdas de energia na caminhada aleat ´ oria de busca, e abordaremos a dinˆamica energ´etica do processo de busca unidimensional com extremos absorventes. Vamos discutir a transic¸ ˜ao de fase que o buscador experimenta de um estado ativo (“vivo”), t´ıpico de ambientes com abundˆancia de recursos, para um estado est´atico absorvente (“morto”), onde a busca ´e encerrada pela falta de energia oriunda do encontro de recursos. Obteremos os expoentes cr´ıticos relativos a essa transic¸ ˜ao atrav´es de abordagens te ´ oricas, tais como o m´etodo de primeira passagem para o estado de energia nula, e num´ericas, baseadas na hip´otese de escala. Mostraremos a independˆencia destes expoentes com a forma funcional da func¸ ˜ao gasto de energia. Por fim, faremos uma breve revis˜ao da literatura sobre a equac¸ ˜ao de Fokker-Planck canˆonica e tamb´em sobre as suas vers˜oes utilizando derivadas fracion´arias, numa prepararac¸ ˜ao para uma futura abordagem, durante o programa de Doutorado, do problema da busca aleat´oria envolvendo difus˜oes anˆomalas (por exemplo, superdifus˜ao) via equac¸ ˜oes diferenciais. / In this work we study the energy dynamics of random searches applied to the foraging problem, in which animals search for food or mates in scarce environments. Firstly, we discuss a statistical model of random search walks using the L´evy distribution of step lengths, which has been reported in the literature as an optimal solution to the problem. In the sequence we include in the model energy gains and losses during the search walk, and discuss the energy dynamics of the search process in a one dimensional space with absorbing boundaries. We discuss the phase transition that the searcher experiences from an active (“alive”) state, typical of environments abundant in resources, to a static absorbed (“dead”) one, in which the search is terminated due to the lack of energy obtained from the encounters.We obtain the critical exponents for this transition through both theoretical (such as the first-passage method to the state of zero energy) and numerical approaches, based on the scale hypothesis.We show the independence of the exponents with the functional form of the energy cost. Finally, we provide a brief review of the literature on the canonical Fokker-Planck equation and also on its version using fractional derivatives, in a preparation for a future approach of the random search problem involving anomalous diffusion (e.g., superdiffusion) through differential equations during the Ph.D. program.

Most albatrosses are critically endangered, endangered or vulnerable due to the deleterious impact of fisheries, pollution, introduced species, habitat alteration, and climate change. Foraging behaviour influences many aspects of seabird biology, and a detailed understanding of foraging ecology is required to better predict the impacts of significant changes to the marine environment. Campbell Albatross (Thalassarche impavida) is a threatened endemic, confined to a small number of locations on Campbell Island, New Zealand and was recently split from the closely related Black-browed Albatross (T. melanophrys). We currently lack much basic information on the foraging behaviour of this species, hindering our ability to understand how change may have occurred in the past and make predictions about it’s long-term future. First, I used GPS loggers and stable isotope analysis of blood to investigate how distribution and foraging effort (distance travelled and duration) varied with sex and breeding stage. I found that Campbell Albatrosses are sexually dimorphic and showed sex-specific foraging behaviour and habitat use – although this varied by stage of reproduction. Because males and females may be vulnerable to different threats, such as interactions with fisheries, I compared the spatial overlap and high resolution spatio-temporal overlaps between fisheries vessels and albatrosses within New Zealand’s Exclusive Economic Zone (EEZ). Albatrosses utilised 32% of the EEZ, however they overlapped with fisheries vessels in only 0.20% of the area. Previous research has demonstrated that the influence of fisheries vessels goes beyond the immediate location of the boat itself. Campbell Albatross have low levels of spatio-temporal overlap with fisheries – with males overlapping more than females. More generally, my results indicate that adding data on fine scale interactions will improve fisheries risk assessments, and provide information needed for the conservation and management of the Campbell Albatross. A key development in recent ecological research has been a greater appreciation that inter-individual variation in foraging behaviour can have profound population-level consequences. Accordingly I tested for individual differences in foraging behaviour in Campbell Albatrosses. The majority of individuals demonstrated both annual and inter-annual individual consistency in foraging locations, and the degree of specialisation was influenced by both sex and year. Consistent terminal latitude and longitude of foraging trips indicated high foraging area fidelity with a degree of flexibility in the fine-scale location. During brooding, females used the Campbell Plateau and showed more consistent behaviours than males, which tended to forage in the Southern Ocean. This adds to a growing body of evidence of individual foraging specialisation among seabirds in general and albatrosses in particular and reveals marked inter-individual differences in vulnerability to threats. In light of the evidence of individual foraging specialisations in the Campbell albatross, I also preformed a literature review of individual foraging specialisations across all seabirds. I found studies examining foraging specialisation for 35 species, with 28 (80%) providing evidence of consistent inter-individual differences (i.e. specialisation). Current studies suggest that specialisation is influenced by environmental variability and resource predictability, however, with limited data in tropical regions, more studies are needed to test these links. In summary, my thesis has provided new information on Campbell Albatross foraging ecology. Sex specific variations in behaviour and habitat use may influence conservation and management strategies. I have been able to contextualise the consistent individual differences in foraging distribution described for this species in light of global patterns of individual foraging specialisation in seabirds and highlight future areas of research.

Food production and food consumption have been shown to have a great impact on our ecosystem. Human beings have been exploiting the planet in order to feed themselves. This will have negative consequences for future life on the planet. Modern food production and consumption are among the main causes of natural resource exploitation and the problem is very likely to increase. Indeed, during the past thirty years, the global population has grown exponentially by almost one billion every decade, and it is still growing at the same pace. This demographic explosion means that dramatic shifts in the production and consumption of food will be required. Working with food is a great chance to achieve or at least lead towards a condition of recovering, understanding the world around us and managing our natural resourcesÅ. Increasing control and efficiency in food production and consumption cannot solve the problem. There is a much broader spectrum of causes contributing to the ecological decline. It is necessary to look beyond the technological and economic aspects. It is, therefore, necessary to focus on cultural and behavioural causes, promoting the involvement of local peopleÇ. With this in mind, this research explores the potential of urban foraging for generating social consciousness about ecological sustainability using design as method of intervention, and involving food consumers in the process of production and consumption of food in a more sustainable way. In order to do that, this research focuses on a small scale urban foraging project. In this case, by food, I refer to spontaneous food that grow in the natural urban environment of Växjö, and that can be used as a resource for citizens. In summary, this research aims to promote the involvement of local people and to support knowledge exchange in order to pursue socio-ecological sustainability. Engaging with more participants, the research gains the capacity of addressing complexity in a more coherent manner, and use its outcome as a usable resource for the local community that aims to promote its self-sustenance.

No description available.

Master of Science / Department of Entomology / David C. Margolies / James R. Nechols / Many crops grown in greenhouses are damaged by the twospotted spider mite, Tetranychus urticae. The predatory mite, Phytoseiulus persimilis, is a commercially-available predator that is commonly used to control twospotted spider mites on greenhouse crops; but its efficacy varies among crops, and it is generally ineffective at low prey densities. In general, predator foraging efficiency depends on how well predators find prey patches, the length of stay in prey patches, and consumption of prey while in prey patches. With respect to P. persimilis, I asked how this predator responds to different prey distributions, as might be encountered at different stages of spider mite infestations. I also asked how components of foraging, namely consumption rate and dispersal tendency, affected predator efficiency. To examine the former, I established T. urticae eggs on 6-leafed cucumber plants in two distributions. To examine the latter, I imposed artificial selection on a population of P. persimilis to create a line that exhibited extremely high consumption and one that demonstrated a greater tendency for dispersal. Subsequently, foraging efficiency was assessed by observing predator oviposition and consumption of twospotted mite eggs on individual leaves of 6-leafed cucumber plants. The number of eggs laid by predators corresponded to the number of prey consumed regardless of predator line. In addition, predators from both lines distributed their eggs proportional to where they fed. However, prey consumption differed between selected lines in response to prey distribution. Predators selected for high consumption fed more on the basal leaf where they were released; whereas prey consumption by the high dispersal and control lines were more evenly distributed throughout the plant. These results contribute to a better understanding of how foraging behavior is modified in plant landscapes under different levels of expression of foraging traits. They also indicate that predator release strategies likely would need to modified in accordance with the kind of foraging trait(s) used in artificial selection programs. In general, my research, when combined with future studies at a broader landscape level, will facilitate decisions by biological control practitioners about whether changes in foraging efficiency resulting from artificial selection justify the cost investment of producing selected lines of P. persimilis