While antipredator strategies have been a focus of behavioral research for decades, scientists generally study the responses of prey toward overt, explicit threat. However, risk can also be significant when a threat is covert, such as when an ambush predator may be nearby or a secondary threat remains after a predator's departure. Little is known about the mechanism that prey use to assess risk in a predator's absence. Tests were conducted to determine the manner in which coyotes respond to these ambiguous threats. Specifically, I tested whether coyotes respond to prior anthropogenic activity that has occurred near their only food source, whether they investigate human activity at both profitable (feeding) and unprofitable (non-feeding) locations, and what sort of information coyotes are capable of gaining through their investigation. I explored these questions in three experiments spanning 4 years at the USDA/APHIS/WS National Wildlife Research Center's Logan field station. Test subjects were eight pairs of captive coyotes. Results showed that coyotes delayed or avoided feeding in response to prior anthropogenic activity, and that often a delay was due to investigation of human scent trails. Investigation of non-feeding areas occurred but was relatively brief. When coyotes were prevented from investigating locations of prior anthropogenic activity, foraging ceased altogether. In addition, coyotes were able to differentiate among the activity of different humans based on their association with negative, neutral, or positive threat levels, even in the presence of confounding visual and olfactory cues. They remembered these associations even after one month. This study is the first that provides evidence suggesting that canids gather and interpret complex information for cognitive inference about threat level associated with access to food.

Individual organisms face trade-offs when dealing with predation—more time spent avoiding predators often results in less time allocated to energy acquisition and reproductive-related activities. Individuals that optimize this trade-off and respond appropriately to current risk levels in their environment should be at an advantage. What is less clear is whether this tradeoff changes when individuals are repeatedly exposed to a predation threat. There may be advantages to responding consistently to every signal of threat, but it might also be advantageous to modulate individual behavior. Moreover, it is unclear how evolutionary history of a population might affect such individual responses. Our study was designed to address two questions: (1) how do B. rhabdophora respond to repeated exposures of a predation cue; and (2) do repeated responses differ based on evolutionary history? To answer these questions, we used a predation cue stimulus to repeatedly expose B. rhabdophora individuals from both high- and low-predation populations. We measured the change in total distance traveled in a 15-minute trial before and after each cue exposure, and then compared the proportional change in response to the first cue to that of each successive cue (repeated four times) to see if a decrease in response occurred. We found that fish responded consistently to each cue exposure. Both populations showed similar decreases in activity in response to each exposure and did not return to normal baseline activity until the cue was removed from the test tank. That both high- and low-predation populations respond consistently to repeated cues of predation with no reinforcement prompts questions as to the potential importance of the relative length of risk and safe periods in affecting response variation. It also provides a starting point in understanding how recent risk exposure and the evolutionary history of risk in a population both interact to influence individual response to threats over time.

Many species have evolved to live in groups. Gregarious behavior was believed to be adaptive for whole-population survival and predator evasion until selfish-herd theory was introduced. It proposed that individuals congregate not as a method of protection for the entire population, but instead to better the individual's chance of survival, thereby using the group to benefit itself (i.e., "selfishness"). Selfish behavior is a common part of mutualisms, which are complex, dynamic interactions that often change with biotic or abiotic circumstance. Here, I investigate potential selfish behavior within the mixed-species spawning aggregations hosted by bluehead chub (Nocomis leptocephalus). I hypothesized that the host benefits more directly from the aggregation it supports than previous research suggests – specifically, by using nest associates to decrease its own risk of predation (creating a selfish-herd) and simultaneously forcing associate embryos into marginal nest locations while locating its own embryo in the safer locations within the nest (creating an "embryonic selfish-herd"). In Chapter 1, I investigated the adult spawning aggregation for possible selfishness by monitoring the chub's spatial location within the aggregation and vigilance behavior under varying associate abundances. I found that the host occupied the central location within the aggregation and was less vigilant when associate abundance was high. In Chapter 2, I examined the "embryonic herd" contained within Nocomis nests for possible embryonic selfishness orchestrated by the chub host, leading to increased embryonic survival for chub young and elevated fitness for chub parents. I found that deeper nest sections support higher embryonic survival than shallower sections, and that chub embryo make up a disproportionate percentage of embryos found in those deeper sections. I also conducted a preliminary study investigating embryo-predation by host on associate embryo which produced evidence for embryo-predation by both host and two associates on each other's offspring. Cohesively, my results support the identification of the host as a selfish participant and confirm the presence of both an adult and embryonic selfish-herd in this mutualism. This is the first study to demonstrate selfish behavior on the part of the host in this system and the first to prove the existence of an embryonic selfish-herd. / Master of Science / A fish has three goals: to survive, to grow, and to reproduce. Mutualisms – when all participants experience a net benefit from interacting – arise when two or more species rely on each other to meet these objectives. In many mutualisms, however, not all participants benefit equally. Selfish-herd theory identifies those that maximize their own benefit while minimizing or negating any cost as "selfish" individuals. I examined the mutualistic mixed-species spawning aggregations hosted by a freshwater minnow, bluehead chub (Nocomis leptocephalus), for possible selfish behavior. I found that bluehead chubs disproportionately benefit from the aggregations they host and use the aggregation to improve their own survival, improve their offspring's survival, and obtain food (by eating others' offspring). Thus, I identify bluehead chub as a selfish participant and provide evidence for the existence of two selfish herds in this mutualism – one among the adult fishes and one among their young – both orchestrated by and benefiting the bluehead chub. This study is the first to identify the bluehead chub as such and rewrites our previous understanding of this mutualism, which largely denied bluehead chub as a direct beneficiary. This work contributes to the global discussion of mutualisms by attesting the complexity of these relationships and offers support for re-examination of the classification of many known interspecific interactions, such as those cursorily termed nest parasitisms.

Made available in DSpace on 2015-03-04T18:51:18Z (GMT). No. of bitstreams: 1 TESE_PORTUGUES_JOSE_CARLOS.pdf: 4654107 bytes, checksum: 9ffae601e2a71b27032455457d38fe96 (MD5) Previous issue date: 2009-04-01 / Predation, classically described as the negative effect of the predator on the density of their prey, will be examined for their effects on the behavior of prey,in the form of antipredator responses. Antipredator responses may arise on the morphology, physiology and/or the behavior of prey, by predation or by the mere presence of the predator, in this case called non-lethal predator. In this context we mainly examine the effect of predators on foraging and change of habitat of their prey, as a antipredator response. Through the diversity of models surveyed, we exam the dynamics as often they are analised, through indirect effects mediated by density, and exam the same dynamic added of the trait-mediated indirect effects,through behavioral modeling techniques. / A predação, classicamente descrita como sendo o efeito negativo do predador sobre a densidade de suas presas, será aqui analisada a respeito de seus efeitos sobre o comportamental da presa, na forma de resposta antipredatória. Respostas antipredatórias podem surgir sobre a morfologia, a fisiologia e/ou o comportamental da presa, pela predação ou pela mera presença do predador,nesse caso denominado de predador não letal. Nesse contexto examinamos principalmente o efeito do predador sobre o forrageamento e a mudança de habitat de suas presas, como forma de resposta antipredatória. Por meio dos diversos modelos pesquisados, buscamos avaliar as dinâmicas da forma usualmente analisada, através dos efeitos indiretos mediados pela densidade, e analisar essa mesma dinâmica adicionada dos efeitos indiretos mediados por traço, através da modelagem comportamental.

Few studies have rigorously assessed the adaptive value of diel rhythms in animals. We laid the groundwork for assessing the adaptive rhythm hypothesis by assaying diel rhythms of foraging and antipredator behavior in the orb-weaving spider Cyclosa turbinata. When confronted with a predator stimulus in experimental arenas, C. turbinata exhibited thanatosis behavior more frequently and for longer durations during the day. However, assays of antipredator response within webs revealed more complex diel patterns of avoidance behaviors and no pattern of avoidance behavior duration. Assays of prey capture behavior found that the likelihood of exhibiting prey capture behavior varied significantly across times of day and test subjects, but only test subject predicted attack latencies. Although C. turbinata foraging aggression changed over the diel cycle, we found no evidence of a trade-off between foraging behavior and predator vigilance. However, overall patterns of vigilance may be masked by diel changes in antipredator strategies.

Alarm calls are one of the essential components of antipredator behaviour in birds. In this study I recorded and analysed alarm responses of the barred warbler (Sylvia nisoria) to different mounts of avian predators and nonpredators. The information about danger is encoded through graded structure in its unspecific alarm call type.

An increasing amount of attention is devoted to studying the impact of non-native animal species on native species. Among other causes of ecosystem degradation, such as climate change, polution and habitat conversion, biological invasion is considered as one of the main causes of the decrease in biological diversity all over the world. In order to detect possible predation event, tadpoles use not only visual and mechanical stimuli, but also chemical one. The tadpoles respond to the certain chemicals to be a part of predator's secretions. In aquatic systems, chemical cues are a major source of information through which animals are able to assess the current state of their environment to gain information about local predation risk. Prey use chemicals released by predators to mediate a range of behavioural, morphological and life-history antipredator defences. Tadpoles swim significatnly less and also on a less direct trajectory in the presence of chemical cues released by a turtle Trachemys scripta. This article is focused on the influence of mentioned constraints on the dynamics and the time dynamics of the larval growth of Rana temporaria. The influence was judged by the impact on the tadpole's development. We anticipated a behavioral response to the predator, which would lead to various growth reactions during larval growth, the resulting body size after metamorphosis, but also in the development time of larval stage. The results indicate that tadpoles of Rana temporaria changed behaviour in the presence of Trachemys scripta. Tadpoles in the permanent presence of the turtle grew faster, metamorphosed earlier and the resulting size after metamorphosis was smaller than of the tadpoles who developed without presence of the predator. Mentioned reactions may affect the survival and fitness of a metamorphosed individual.

Predation, classically described as the negative effect of the predator on the density of their prey, will be examined for their effects on the behavior of prey,in the form of antipredator responses. Antipredator responses may arise on the morphology, physiology and/or the behavior of prey, by predation or by the mere presence of the predator, in this case called non-lethal predator. In this context we mainly examine the effect of predators on foraging and change of habitat of their prey, as a antipredator response. Through the diversity of models surveyed, we exam the dynamics as often they are analised, through indirect effects mediated by density, and exam the same dynamic added of the trait-mediated indirect effects,through behavioral modeling techniques. / A predação, classicamente descrita como sendo o efeito negativo do predador sobre a densidade de suas presas, será aqui analisada a respeito de seus efeitos sobre o comportamental da presa, na forma de resposta antipredatória. Respostas antipredatórias podem surgir sobre a morfologia, a fisiologia e/ou o comportamental da presa, pela predação ou pela mera presença do predador,nesse caso denominado de predador não letal. Nesse contexto examinamos principalmente o efeito do predador sobre o forrageamento e a mudança de habitat de suas presas, como forma de resposta antipredatória. Por meio dos diversos modelos pesquisados, buscamos avaliar as dinâmicas da forma usualmente analisada, através dos efeitos indiretos mediados pela densidade, e analisar essa mesma dinâmica adicionada dos efeitos indiretos mediados por traço, através da modelagem comportamental.

Si le signalement du prédateur provoque le plus souvent la fuite des proies, il induit parfois un comportement particulier incitant la proie à s’approcher du prédateur et le harceler pour provoquer son départ plutôt que de se mettre hors de sa portée. Ce comportement de harcèlement s’accompagne de l’émission d’un signal hétérospécifique conduisant de nombreuses espèces à venir harceler le prédateur. L’objectif de mon travail de thèse est de comprendre comment est régi le transfert d’informations entre plusieurs espèces de passereaux dans le cas du comportement de harcèlement d’un prédateur. Mes travaux montrent que le comportement de harcèlement des passereaux face à un rapace nocturne dépend du risque de prédation posé par ce prédateur. Par ailleurs, les résultats indiquent un transfert d’informations entre les espèces et mettent en évidence une propension variable des différentes espèces à se rallier autour du harceleur. Les variations observées dans la réponse aux cris de harcèlement émis par des individus hétérospécifiques peuvent dépendre de la similarité acoustique, des relations interspécifiques et des variations saisonnières. Mes résultats indiquent aussi que la connaissance préalable des signaux de harcèlement n'est pas indispensable pour induire une réponse, même si un processus d’apprentissage associatif favorise sa mise en place. Mon travail suggère également une évolution convergente des cris de harcèlement, générant des signaux dont la structure permet une localisation rapide de l’émetteur, indispensable pour rameuter des proies potentielles lors du harcèlement. L’ensemble de ces avancées nous oblige désormais à considérer la communication acoustique chez les passereaux en prenant en compte le risque de prédation, les interactions hétérospécifiques et la complexité des signaux acoustiques / Signaling the presence of a predator most often causes the escape of prey, but it sometimes induces a particular behaviour prompting prey to approach and harass the predator to cause his departure. This mobbing behaviour is associated with the emission of signals leading individuals from different species to come harass the predator. The objective of this thesis is to understand how the transfer of information between several passerine species is organized in mobbing behaviour against predators. My work shows that the mobbing behaviour of passerine birds against a nocturnal raptor depends on the predation risk imposed by this predator. In addition, my results indicate a transfer of information between species and highlight a variable propensity of different species to rally around the harasser. The observed variations in the response to heterospecific calls depended on acoustic similarity, interspecific relationships and seasonality. My results also indicate that prior knowledge of harassment signals is not essential to induce a response, even if an associative learning process promotes its implementation. My work also suggests a convergent evolution in mobbing calls, generating signals with a structure that allows the emitter to be quickly located, an essential parameter to rally potential prey during harassment. My thesis consequently shows that to better understand acoustic communication in passerine birds, it is necessary to consider predation risk, heterospecific interactions and the complexity of acoustic signals

Anfíbios anuros representam um grupo de vertebrados cujo plano corpóreo apresenta uma série de modificações associadas ao desempenho locomotor através de saltos, sendo estas especializações muito antigas e conservadas filogeneticamente. Embora estudos comparativos venham demonstrando associações entre desempenho de salto e diversidade de habitat e história de vida para esse grupo filogenético, as relações entre o desempenho locomotor e o comportamento antipredatório dentro do contexto da diversificação do uso do micro-habitat permanecem inexploradas. Primeiramente, nós testamos modelos adaptativos de evolução morfológica associados com a diversidade do uso do micro-habitat (aquático, arborícola, fossorial, reofílico e terrestre) em espécies de anuros e foi examinada a relação da distância máxima do salto como uma função dos componentes das variáveis morfológicas e do uso do micro-habitat. Nós também investigamos, tanto em nível intra quanto interespecífico a influência da complexidade do microambiente (arena vazia, arena com folhiços ou arena com folhiço e arbustos) e do tipo de estímulo (aproximação versus toque) na manifestação do comportamento antipredatório em anuros. Adicionalmente, nós investigamos o efeito do uso de diferentes tipos de refúgio pelos anuros quando sujeitos aos testes de simulação predatória no laboratório. Nossos resultados demostram a existência de múltiplos ótimos adaptativos para os comprimentos dos membros associados aos diferentes usos do micro-habitat, com uma tendência de aumento dos membros posteriores em espécies reofílicas, arborícolas e aquáticas quando comparadas com espécies terrestres e fossoriais, as quais evoluíram em direção ao ótimo adaptativo com membros posteriores mais curtos. Além disso, espécies reofílicas, arborícolas e aquáticas apresentaram maior desempenho para o salto e membros posteriores mais longos quando comparadas com espécies terrestres e fossoriais. Em seguida foi abordada a influencia da complexidade do ambiente no comportamento antipredatório e nossas análises intraspecíficas mostraram que o número de respostas ativas (salto) e passivas é fortemente dependente do tipo de estímulo e varia de acordo com a complexidade estrutural do ambiente. Simultaneamente, nossas análises comparativas interespecíficas mostraram que anuros modulam a distância saltada em resposta ao toque do predador de acordo com a complexidade ambiental, e nós ressaltamos uma variação interespecífica associada ao uso de micro-habitat. Por fim, foi investigado o uso de refúgios pelos anuros expostos a testes de simulação predatória. Nossos resultados mostram que os anuros usam a vegetação arbustiva, o folhiço e a água como areas de refúgio quando sujeitadas a simulação predatória. Além disso, ocorreu variação interespecífica na escolha de refúgios potencialmente associados à diversificação do uso de microhábitat / Anuran amphibians represent a group of vertebrates whose body plan presents a series of changes associated with jumping locomotor performance; these specializations are very old and phylogenetically conserved. Although comparative studies have shown associations among jumping performance, habitat diversity and life history for this phylogenetic group, the relationship between locomotor performance and antipredator behavior remains unexplored within the context of diversification of microhabitat use and habitat. First, we tested adaptive models of morphological evolution associated with the diversity of microhabitat use (aquatic, arboreal, fossorial, torrent and terrestrial) in species of anurans and examined the relation of the maximum distance jumped as a function of components of morphological variables and microhabitat use. We also investigated, both at intra and interspecific levels, the influence of the complexity of microenvironment (empty arena, with leaf litter or bushes) and stimulus type (approach versus touch) on the manifestation of antipredator behavior in anurans. In addition, we investigated the effect of the use of different types of refuges by anurans when subjected to simulated predator tests in the laboratory. Our results demonstrate the existence of multiple optima of limb lengths associated to different microhabitats, with a trend of increasing hindlimbs in torrent, arboreal, aquatic species whereas fossorial and terrestrial species evolve toward optima with shorter hindlimbs. Moreover, arboreal, aquatic and torrent anurans have higher jumping performance and longer hindlimbs, when compared to terrestrial and fossorial species. Then, was addressed the influence of the complexity of environment on antipredator behavior and our intraspecific analyzes showed that the number of active responses (jump) and passive responses is strongly dependent on stimulus type and varies according to the structural complexity of the environment. At the same time, our interspecific comparative analyzes showed that anurans modulate the distance jumped in response to the predator\'s touch according to environmental complexity, and we highlight an interspecific variation associated with the use of microhabitat. Lastly, was investigated the use of refuges by anurans exposed to simulated predation events. Our results show that anurans use bushes, leaf litter and water as refuge areas when subjected to simulated predator. Moreover, there is interspecific variation in the choice of refuges potentially associated to diversification of microhabitat use