‘Tipping points’ in the climate system are characterised by a nonlinear response to gradual forcing, and may have severe and wide-ranging impacts. One of the best ways to identify and potentially predict threshold behaviour in the climate system is through analysis of palaeoclimate records. It has been suggested that early warning signals occur on the approach to a tipping point, generated from characteristic fluctuations in a time series as a system loses stability. Although early warning signals have been found in climate models and high-resolution marine and ice core palaeodata, studies from terrestrial records are lacking. In this study, a number of Pleistocene terrestrial records were selected to represent a range of regions strongly influenced by different climate modes which are thought to be capable of displaying threshold behaviour. These records included lake sediments from the North Atlantic, tree-rings from the South Pacific, a Chinese speleothem and were complemented by a new Greenland ice core chronology. Recently developed methods to detect signals of ‘critical slowing down’, ‘flickering’, and stability changes on the approach to a tipping point were utilised. Specific methodological issues arising from analysing palaeoclimate data were also investigated using a simple bifurcation model. A number of key criteria were found to be necessary for the reliable identification of early warning signals in palaeoclimate records, most crucially, the need for a low-noise record of sufficient data length, resolution and accuracy. Analysis of a Chinese speleothem identified the East Asian Summer Monsoon as an important climate ‘tipping element’, which may display a cascade of impacts. However, in some cases where early warning signals may fail, a deeper understanding of the underlying system dynamics is required to inform the development of more robust system-specific indicators. This was exemplified by the analysis of an abrupt, centennial-duration shutdown recorded during the Younger Dryas Chronozone in New Zealand, which demonstrated no slowing down, consistent with a freshwater pulse into the Southern Ocean. This study demonstrates that time series precursors from palaeoclimate archives provide a means of useful forewarning of many potential climate tipping points.

Many species use visual features to avoid predation by several methods, such as concealing themselves, deceiving predators and hindering capture. One of the most striking strategies is aposematism, or warning coloration, in which prey use conspicuous visual signals to advertise chemical or physical defences, and thereby deter predators from attacking. My thesis focuses on the form of these warning signals, namely which elements of visual patterns might be most effective in generating predator avoidance, as well as how these different visual features relate to defence levels and ultimately to prey survival in the wild. To address these issues, I studied the warning signals of Lepidoptera and in particular burnet moths (Zygaenidae: Zygaeninae), day-flying moths with distinctive red and black wings and the remarkable ability to both synthesise defensive compounds and sequester them from their host plants. Technological advances and a growing understanding of animal vision mean that animal signals can be studied in an increasingly precise and ecologically-relevant way. Throughout this thesis, I use sophisticated methods to quantify both the defensive chemicals and wing coloration of burnet moths, as perceived by their avian predators. I examine the key features of day-flying defended Lepidoptera, then focus on the potential for quantitative signal honesty in burnet moths. I explore the relationship between defence levels and measures of coloration, both within the six-spot burnet moth, Zygaena filipendulae, and across species in the Zygaenidae, then test the effects of variation in warning signals on predation risk for artificial burnet-like prey in the field. My work highlights some of the complicating factors that should be accounted for in the study of warning coloration, especially when investigating the potential for quantitative signal honesty. I hope my thesis will provide a basis for future research on the defensive strategies of day-flying moths and inspire others to pursue investigations into aposematism in the Zygaenidae.

<p>Researchers have long assumed that black widow coloration functions as a warning signal to avian predators. However adult female black widow coloration does not resemble typical warning coloration in two distinct ways. First, black widows are less colorful than most other documented aposematic species. Second, the hourglass shape of an adult female varies both between species and within a site. Here I examine the ultimate causes and consequences of North American black widow coloration. </p><p>In chapter two I present data that suggest that black widow coloration not only functions as an aposematic signal to avian predators, but has also been selected to be inconspicuous to insect prey. In choice experiments with wild birds, I found that the red-and-black coloration of black widows deters potential predators: wild birds were ~3 times less likely to attack a black widow model with a red hourglass than one without. Using visual-system appropriate models, I also found that a black widow's red-and-black color combo is more apparent to a typical bird than typical insect (Euclidean color distance ~2.2 times greater for birds than insects). Additionally, an ancestral reconstruction revealed that red dorsal coloration is ancestral in black widows and that at some point some North American black widows lost their red dorsal coloration (while maintaining the ventral hourglass). Behaviorally, differences in red dorsal coloration between two North American species are accompanied by differences in microhabitat that affects how often a bird will view a black widow's dorsal region. All observations are consistent with a cost-benefit tradeoff of being conspicuous to potential predators while being inconspicuous to prey. I suggest that avoiding detection by prey --- combined with Müllerian mimicry --- may help explain why red-and-black aposematic signals occur frequently in nature.</p><p>In chapter three, I examine the variation in hourglass shape. Classical aposematic theory predicts near uniformity in warning signal appearance because a uniform signal is easier to learn to avoid than a variable signal. However the shape of the hourglass of North American black widows appears to vary both within and between sites in ways that are inconsistent with classical aposematic theory. Using 133 black widows of three different species from nine sites across the United States, I quantified the variation in hourglass shape and examined how Müllerian mimicry, species type, and condition each influenced hourglass shape. A principle components analysis revealed that 84.5% of the variation in hourglass shape can be explained by principle components 1, 2, and 3, which corresponded to hourglass size (PC1), the separation between hourglass parts (PC2), and the slenderness of the hourglass (PC3). Both a black widow's condition and species significantly predicted hourglass shape; however I found no support for localized Müllerian mimicry within different geographical regions. My results suggest a relaxed role for selection on hourglass shape. I discuss several hypotheses that could explain the variation in hourglass morphology including that potential predators may avoid any red markings rather than an exact shape (categorical rather than continuous perception).</p><p>In chapter four I expand on my work from chapter two to examine the eavesdropper's perspective on private communication channels. Private communication may benefit signalers by reducing the costs imposed by potential eavesdroppers such as parasites, predators, prey, or rivals. It is likely that private communication channels are influenced by the evolution of signalers, intended receivers, and potential eavesdroppers, but most studies only examine how private communication benefits signalers. Here, I address this shortcoming by examining visual private communication from a potential eavesdropper's perspective. Specifically, I ask if a signaler would face fitness consequences if a potential eavesdropper could detect its signal more clearly. By integrating studies on private communication with those on the evolution of vision, I suggest that published studies find few taxon-based constraints that could keep potential eavesdroppers from detecting most hypothesized forms of visual private communication. However, I find that private signals may persist over evolutionary time if the benefits of detecting a particular signal do not outweigh the functional costs a potential eavesdropper would suffer from evolving the ability to detect it.</p> / Dissertation

Aposematic species of true bugs (Heteroptera) have multimodal signalization, which warns potential predators. This signalization consists of optical (coloration), chemical (unpalatable or repugnant substance) and acoustic (stridulation) warning signals. The aim of this thesis was to test whether the selected chemical substances have antipredatory function towards avian predators. Antipredatory function is anticipated in the chemical substances that form the majority in secretion in many taxa of true bugs (aldehydes and tridecane). In experiments with wild-caught great tits (Parus major) and blue tits (Cyanistes caeruleus) we tested if chemical substances and age of birds have influence on the latency related to the first manipulation with the prey. It was found that both species of tits reacted aversively to the mixture of aldehydes (2-decenal, 2-octenal, 2-hexenal) and to the total secretion of metathoracic glands of Graphosoma lineatum, whereas the mixture of the aldehydes with tridecane did not have any effect. The effect of age was not significant. We also tested the influence of immediate experience with striated shieldbug Graphosoma lineatum on naive great tits and their reactions to the prey with olfactoric signal of the shieldbug. Additionally, we investigated whether tested chemicals cause...

In lane departure warning systems (LDWS) it is important that the auditory warning triggers a fast and appropriate reaction from the driver. The rumble strip noise is a suitable warning to alert the driver of an imminent lane departure. A short reaction time is important in lane departure scenarios, where a late response may have fatal consequences. For abstract sounds an increase in intensity can influence the perceived urgency level of the warning, which may also trigger a faster reaction from the listener. In this thesis, the effect of a rumble strip warning with looming (increasing) intensity was analyzed by letting test persons drive a driving simulator and measuring how quickly they reacted to the auditory warning. These results were compared with those for a rumble strip warning with a constant intensity, and two versions of an abstract warning; constant intensity and looming intensity. A survey regarding the perceived urgency, annoyance and acceptance of the warnings was also carried out. The results show no differences in reaction time between the four warning signals. This may be because the test persons expected the warnings, or because of their limited experience. The survey suggests that adding a looming intensity to the rumble strip warning results in a higher urgency, while keeping the annoyance low, which could be of importance to avoid unwanted reactions from the driver. / I varningssystem för personbilar används ofta ett system som signalerar ett stundande ofrivilligt lämnande av körfältet, s.k. lane departure warning systems (LDWS), genom att en varningssignal ljuder. Det är viktigt att en sådan akustisk varningssignal frammanar en snabb och lämplig reaktion från föraren. Ljudet av en bullerräffla är en lämplig varningssignal för detta ändamål. En kort reaktionstid är viktig när fordon är på väg att ofrivilligt lämna körfältet, då en långsam reaktion kan ha förödande konsekvenser. Studier på abstrakta akustiska varningssignaler har visat att en ökande intensitet kan få en varning att verka mer brådskande, vilket i sin tur kan leda till att lyssnaren reagerar snabbare. I denna rapport analyseras hur ett bullerräffleljuds ökande intensitet påverkar förarens reaktionstid. Analysen gjordes genom att mäta reaktionstiden hos testpersoner som körde en bilsimulator med fyra olika varningssignaler; en bullerräffleljudsvarning och en abstrakt varning, båda med konstant intensitet och ökande intensitet. Reaktionstiderna för de olika signalerna jämfördes, varpå en enkät utfärdades där testpersonerna uppgav hur brådskande och irriterande de uppfattade varningarna, samt till vilken grad de skulle acceptera varningarna i ett verkligt körscenario. Resultaten visar inga skillnader i reaktionstid mellan varningarna, vilket kan bero på att testpersonerna förutsåg när varningarna skulle komma, eller på grund av deras begränsade erfarenhet av bullerräffleljud. Enkätens utfall antyder att bullerräffleljudsvarningen med ökande intensitet är mer brådskande än versionen med konstant intensitet, men att irritationsnivån inte påverkas när intensiteten ökar, vilket kan vara viktigt för att inte framkalla oönskade reaktioner hos föraren.

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The increasing number of electric and hybrid vehicles in urban areas has shown to be beneficial in reducing both air and noise pollution. However, the lack of sound when driving at low speed has negatively affected the pedestrian safety since many rely on the vehicle sound for orientation. Regulatory bodies have therefore introduced minimum sound requirements for all silent vehicles, which has resulted in a key challenge for car manufacturers to develop external warning signature sounds. The objective of this project has been to study how these signals should sound in order to fit the image of electric and hybrid vehicles and minimize noise and annoyance. To complete the study, five sound concepts with different characteristics and rhythms were developed using the concept generation process. In order to gain subjective impressions of the sounds, three measurement methods were utilized. The first listening test was performed to eliminate the possibility that the sound samples would be perceived variously loud in the upcoming tests. The second listening test was performed to gain information on how suitable the signals are for electric and hybrid vehicles, and the third listening test was conducted to obtain information on how much annoyance the signals contributed with. A reference signal representing an internal combustion engine was included in the tests as well. The results of the measurement methods showed that the sound samples with long tone sequences were more preferred than those with short tone sequences, and that the artificial approach was more popular than the sound of an internal combustion engine vehicle. It was also established that additional tests need to be performed in order to confirm that these results are valid. It was suggested that field tests should be conducted and that new sound concepts should be developed based on the results of these tests.

The ability to memorize and recognize edible prey from inedible prey is essential for an individuals survival. Many species use aposematic signals for their defense. These are most often represented by distinctive colors or contrast patterns. The aim of this study was to find out in which time the memory consolidation for aposematic pray is achieved and if color or pattern increase its memorability. The choosen model species was the Great tit (Parus major). The comparision was done between handreared naïve birds and wild-caught adults of different age and sex. During the discriminatory task of consolidation experiment, birds were simultaneously presented with palatable and unpalatable prey in the form of paper dummies of bugs, differing in color (red versus green). The birds were divided into three groups with a different interval (0, 1 or 3 hours) for consolidation. The results of consolidation test show that adult birds were more successfull in solving the task than juvenile birds. The only difference between the experimental groups was that the group with one-hour interval achieved better results than other groups. The effect of color of palatable and unpalatable prey on discrimination learning was also found only in adults. Memorability of warning signals was tested using paper bugs of...

Aposematic animals advertise their defensive mechanisms to potential predators using warning signals. Signalling through more than one sensory pathway is called multimodal warning display. Most experimental studies of aposematism have been focused on the effect of a particular warning signal rather than on importance of multimodal signalling. Focusing on the multimodal signalling of real prey is the best way how to understand its effect in nature. The present study is focused on comparing the effect of multimodal warning display of insect prey with its particular warning signals on two species of bird predators. Multimodal warning signalisation of the burrowing bug, Tritomegas sexmaculatus consists of visual (black and white coloration), chemical (odour, possibly taste) and acoustic (stridulation) signals. We compared reactions of wild-caught great tit (Parus major) and blue tit (Cyanistes caeruleus) to three types of bugs with different warning displays. The non-manipulated bugs displayed multimodally, the brown painted bugs had their warning coloration manipulated and the dealatized bugs couldn't emit acoustic signals. The wild-caught birds of both species avoided all types of bugs. In an experiment with naive hand reared great tits we compared their reactions to non-manipulated and dealatized bugs. Naive...

In this thesis we empirically examine whether the Turkish stock market is driven by rational bubbles over the period between March 1990 and February 2012. The bubble periods are estimated using a recently developed right-tailed unit root test, the generalized sup augmented Dickey-Fuller test of Phillips, Shi and Yu (2011a). Applying their bubble detection and location strategies to weekly price dividend ratio series, we find strong evidence for the existence of rational bubbles in the Turkish stock market benchmark indices as well as sector indices. Our located bubble periods may give early warning signals of the subsequent Turkish financial crisis.