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This thesis focuses on various factors affecting effectiveness of aposematic signals against avian predators. Adult, wild-caught as well as hand-reared juvenile great tits (Parus major) were used as predators in the experiments. The thesis consists of the following four studies. In the first study, we compared the reactions of great tits from two geographically distant populations toward aposematic firebugs (Pyrrhocoris apterus) and their non-aposematic artificially made colour variant. The birds from the Bohemian population mostly avoided aposematic firebugs and attacked non-aposematic variant. Finnish birds, which lacked experience with firebugs from their natural environment, were less hesitant to attack both firebug colour forms. Although the Bohemian birds avoided the aposematic prey variant, they were not more neophobic than Finnish birds. We conclude that the geographic differences in reactions of the birds to aposematic prey can be explained by a different population-specific experience of the birds with local aposematic prey communities. In the second study, we compared effectiveness of two chemical defence strategies in leaf beetle larvae (Chrysomela lapponica) against great tits. The birds avoided larvae devoid of external secretions after the first attack, which indicates the presence...

Social learning in animals is a frequently studied topic, including birds. Great tits belong to frequently tested passerine species in this context. For this experiment we have chosen another tit species as model birds - adult and juvenile blue tits (Cyanistes caeruleus) and coal tits (Periparus ater). We tested an effect of social information on discrimination learning between palatable and unpalatable prey. Red and green artificial prey items resembling shieldbugs with a mealworm glued underneath were offered to birds. One variant was palatable, the other was unpalatable, soaked in bitter substance. Experimental groups were allowed to observe a pretrained tutor of the same species choosing palatable prey variant and rejecting the unpalatable one. The birds were then subjected to an individual discrimination learning task with simultaneously offered palatable and unpalatable prey items. The control group was not allowed to observe tutor and learned only individually. On the following day, all birds participated in a memory test. We tested if social information has an influence on avoidance learning in juvenile and adult birds of both species. We found that both species of tits were similarly successful in discrimination learning and there was no difference in success in performance between adult...

Strikingly coloured species can form mimetic complexes. One of them is an extensive black-and-red mimetic complex in the Palearctic region. It consists of representatives across arthropod taxa with different levels of defense and investment in it. To describe individual mimetic relationships in this complex, it is necessary to know the effectiveness of these defense mechanisms against predators. Reactions of individual predators vary considerably. Differences in reactions primarily depend on the ability to perceive some components of defense, using sensory and cognitive abilities. Mechanisms and effectiveness of antipredatory defence are the subject of numerous studies, but there is still a lack of description of some mechanisms among many representatives forming this mimetic complex. For this reason, further studies are needed to allow the derivation of mimetic relationships prevailing in this mimetic complex in the future. Keywords: mimicry, aposematism, antipredatory defense, colouration, mimetic complex

The efficiency of the aposematic defenses (colouration and chemical defence) used by three rare colour forms of Harmonia axyridis were tested using in the wild caught great tits (Parus major) as predators. I predicted that visually orienting predator will pay more attention to the rare colour forms, as they are not familiar with them and need to generalize the learned aversion to some other ladybirds to these novel ones.

Warning coloration is used by prey to signal its unprofitability to potential predators. Warning colours may have different effects on various cognitive processes of predators (innate avoidance, avoidance learning, memory and generalization). Typical colours regarded as aposematic are red, orange and yellow. Red colour is considered to be the most effective signal and has a significant effect on avoidance learning, memory and generalization. Orange also represents an effective warning signal, though it has been studied less. Yellow is effective aposematic stimulus but it has frequently been found less effective compared to red and orange. Warning functions of white, blue, violet and ultraviolet colours were studied less frequently and their role in aposematism is not yet clear. Iridescent coloration might also be an effective warning signal affecting avoidance learning, memory and generalization. In this thesis I present a summary of information with regard to different warning colours and their effect on cognitive processes of predators. Key words: aposematism, warning coloration, cognitive processes, red, orange, yellow, iridescence, avoidance learning, innate avoidance, memory, generalization

Aposematic prey advertises its unprofitability to predators by means of warning signals, which can be visual, chemical, acoustic or combined with each other as multimodal signals. Most experimental studies dealing with responses of predators to aposematic prey focus on vertebrates, especially birds. This thesis focuses on the most frequently tested groups of terrestrial arthropod predators - spiders, dragonflies, and mantises - supplemented by less tested insect predators. Most of the tested arthropod predators are able to learn to avoid noxious aposematic prey based on either visual, olfactory or acoustic signals depending on their perception. Some can remember this aversion for several days. Only in case of jumping spiders, ability to generalise previous experience with a particular prey to similar prey has been confirmed. Prey chemical defenses most often consist of cardenolides or aldehydes, which proved to be aversive for tested arthropod predators.

Oxycarenus lavaterae is species of Heteroptera and possessing pink blots on corium and distinctive, gleaming membranaceous wings. This appearance could correspond to the definition of a warning color. The design is similar to other coreoid Heteroptera, which live in the Czech Republic; though Oxycarenus lavaterae is a species originating from Mediterranean and quickly spreading north. In Czech Republic it is a new species and predators thus have only short experience with it.The information about chemical protection of Oxycarenus lavaterae is weak and the effect on predators has never been tested. The characteristic behaviour for this species is creating large aggregations, primarily at the time of hibernation. At that time, several thousands of individuals are gathered in apertures or at the surface of tree bark. The aggregation may enhance the warning appearance of this species. The aim of this study was to test the response of bird predators to the adult individuals of Oxycarenus lavaterae. Three species of insectivorous birds differing in the level of foraging and diet specialization was chosen as predator (namely - Great tit (Parus major), Blue tit (Cyanistes caeruleus), Eurasian blackcap (Sylvia atricapilla)). Unmodified individuals of Oxycarenus lavaterae were served to these predators. In the case of the Parus major, the aggregation of Oxycarenus lavaterae (about 100 pieces) was also presented to test the effect of gregarious behavior on the warning signal. This study showed that the Parus major and the Sylvia atricapilla attacked the individual Oxycarenus more often than the Cyanistes caeruleus showing high level of restraint. In experiments with aggregatedOxycarenus lavaterae Parus major attacked them less often than when presented solitarily. However, after eating some Oxycarenus lavaterae, the Parus major showed symptoms of disgust and loss of appetite.

Social learning is a topic of many studies. We tested the effect of social learning on the acquisition of avoidance against aposematic prey. We have chosen wild-caught adult and naive hand-reared juvenile great tits (Parus major) as a model predator species, because their individual avoidance learnig of aposematic prey is well-studied. We used red and green paper dummies of bugs with a mealworm (Tenebrio molitor) stuck underneath as an artificial prey. Mealworms were soaked in water or in bitter-tasting solution of quinine. We used two types of conspecific demonstrators - naive birds that showed aversive reactions while tasting an unpalatable prey and experienced birds that were trained not to handle the unpalatable pray at all. We compared effects of both demonstrators on discrimination learning and we tested whether these effects differ in adult and juvenile birds. Observing an experienced demonstrator had an effect on the performance of observers at the beginning of learning process. The observers did not reject the unpalatable prey completely, but the number of trials in which they correctly chose the palatable prey was higher in comparison with birds that observed naive demonstrators and birds from the control group with no demonstrator. Latencies to handle the unpalatable prey in the first...

By warning signalling prey advertises its unpalatability to the predators. Typical examples are true bugs (Heteroptera) combining optical and chemical signals. The aim of this study was to find out the effectiveness of chemical defence against wild-caught adults of great tits (Parus major) and blue tits (Cyanistes caeruleus) and towards hand-reared juveniles of great tits. The tested prey were adults and larvae of two invasive species of genus Oxycarenus (Heteroptera: Oxycarenidae) (aposematic O. lavaterae, non-aposematic O. hyalinipennis), adults of Horvathiolus superbus (Heteroptera: Lygaeidae) and crickets (Gryllus assimilis) as a control prey. We were focusing on the influence of seed bugs to the initial reaction, the learning process and to the displays of discomfort behaviour in tits. Reactions affected by the tit species, age and sex were compared in adults. Juveniles were divided into two independent experimental groups, one group was offered adults of genus Oxycarenus, the second was offered sunflower (Helianthus sp.) or Digitalis sp. seed fed adults of H. superbus. The first bug offered did not elicit an initial aversive reaction in tits, but birds reacted aversively to the bugs after having further handling experience. All juveniles, contrary to the adult tits, attacked at least one bug...

The effect of the search image on the ability of hand reared skinks (Chalcides sexlineatus) to reveal a "fake" Batesian mimic was tested with respect to their previous experience with palatable experimental prey (Guyana spotted cockroach Blaptica dubia) which served as a motivational prey as well as midsized mealworm beetle larvae (Tenebrio molitor). The red firebug (Pyrrhocoris apterus) was used as an aposematic model.

Inter-specific and intra-specific variation in reactions towards novel and aposematic prey was found in several species of tits (Paridae). This Ph.D. thesis is focusing on various factors influencing reactions towards novel and aposematic prey in three European species of tits. We tested differences in exploration behaviour, neophobia, dietary conservatism, personality, age and experience as well as ability of avoidance learning and generalisation. We found no difference in exploration behaviour and in reaction towards novel prey in two different populations of great tits (Parus major). But the birds from the Finnish population were more neophobic than Czech birds, but they attacked aposematic firebug (Pyrrhocoris apterus) more often and faster than Czech birds. The difference can be explained by a different experience with local aposematic prey communities. Than we studied initial wariness in naive juveniles of great tits (P. major), coal tits (Periparus ater) and blue tits (Cyanistes caeruleus), and we tested how the initial wariness towards novel and aposematic prey can be deactivated by experience with palatable prey. Great tits and coal tits from experienced groups significantly decreased their neophobia towards both types of prey while blue tits did not change their strongly neophobic...

An aposematism is a type of defence mechanism, by which a prey demonstrates towards predator that it is unpalatable and toxic. The reactions of predators towards chemical warning signals of prey differ due to the quantity and a combination of signals showed by prey, the predator species and its primary orientation. Most of works tested reactions of birds towards gustatory and olfactory stimuli, often in combination with signals from other modalities. Although the birds are predators which orientate primarily visually, olfaction, gustation and chemesthesis significantly helped them with determining unpalatable food. Among other tested predators were fish, amphibians, reptiles, mammals and invertebrates and their reactions towards the same chemicals of prey are noted for comparison. The main chemicals used in the experiments regarding birds were denatonium benzoate (Bitrex), quinin and pyrazines. Predators often showed an aversive behaviour, neophobic reactions, accelerated learning and more effective memorizing of the unpalatable prey if it was presented together with chemical signals. Some of the predators were able to control their toxin intake relative to the amount of toxin ingested or to discriminate between palatable and unpalatable food according to the chemical concentration in it. Key...

1 Abstract: Multimodal signaling is quite common in nature. Even if we focus only on signaling among prey and its predator, we can still find many examples of communication combining more than one modality (vision, hearing, smell, taste). This thesis is about multimodal warning visual and olfactory signaling between true bugs (Heteroptera) and their avian predators. Aldehydes are an important compound of repellent secretion of most true bug species. They are considered to have defence effects against predators. In our experiments, we tested the effect of aldehydes mixture on behaviour of predators - naive hand-reared and experienced wild-caught adult great tits (Parus major) - towards several kinds of prey - aposematic true bugs Pyrrhocoris apterus, nonaposematic true bugs Pyrrhocoris tibialis and red painted mealworms (Tenebrio molitor larvae). We were focusing on the effect of aldehydes on initial reaction, learning process and predator's memory. Aldehydes elicited innate wariness in naive hand- reared great tits towards aposematic P. apterus, but not towards P. tibialis. Therefore it seems possible, that aldehydes act as a signal of unpalatability for naive predators, but only in multimodal combination with visual warning signal. Similar effect of aldehydes was not found in experiments with wild-caught...

Ţádný organismus v přírodě není izolován od organismů okolních. Pokud se ţivočich neměl stát zcela vyhubenou kořistí, v průběhu biologické evoluce se u něj musely vyvinout mechanismy, které mu umoţnily co nejlépe se vyhnout ulovení. Vzniká tak evoluční vztah mezi kořistí a lovcem, při kterém zdokonalení lovu lovcem vyvolá u kořisti zdokonalení mechanismů, jak predaci uniknout. Tato tzv. koevoluce je jedním z hnacích motorů evoluce. Výsledkem je víceméně ustálená rovnováha mezi lovcem a kořistí. Tento material vznikl z finanční podpory Evropského sociálního fondu a státního rozpočtu České republiky v rámci projektu „Popularizace vědy a badatelsky orientované výuky“, registrační číslo CZ.1.07/2.3.00/45.0007.

The problem of the supposedly mimetic relationship between some species of king snakes and coral snakes is rather scattered in literature and opinions on it vary greatly. In this work, I focus on summarizing and generalizing the existing pieces of knowledge about the colouration of these snakes and its consequences for evolutionary biology - neodarwinistic and especially portmannistic. In other words, natural selection does not solve all the questions and in this case in particular, it is obvious that mimesis interests not only biologists, but also mathematicians, chemists, physicists and philosophers. All of these are trying to find the answer to whether mimesis in these snakes is actually occurring and what would be the purpose of such colouration if it is not. Adolf Portmann is very clear on this subject: organisms show their inner properties on the outside as a means to self-present. The specific appearance is not determined and it does not have to have any adaptive value at all.

Mimicry is one of strategies which provides protection for the prey from predator attack. There are several kinds of mimetic relationships, one of them is Müllerian mimicry, i.e. mutualistic relationship between two unpalatable prey species. If there are more animal species in this relationship, it is called Müllerian mimetic complex. Relationships in mimetic complexes are affected by many factors like defence level and aposematic signals resemblance of particular species, their different frequencies, alternative prey, prey community structure or predator species and their experience level. These factors determine whether the relationship of the species involved is mutualistic or parasitic. They also affect predators behaviour and their learning rate, memory and generalisation between mimetic species. Powered by TCPDF (www.tcpdf.org)

The dissertation thesis deals with human aesthetic preference to other species and its anthropological and conservation aspect. The aesthetic preference to animal species has rarely been systematically studied before and quantitative analyses, especially on a fine taxonomic scale did not exist. On the other hand, it was known that attractive species often receive more support for their conservation. From these simple facts rose the idea to test human aesthetic preference to snake species (and consequently to species across major vertebrate taxa) and use this quantified preference to explain the conservation effort devoted to captive breeding worldwide (measured as size of zoo populations). We confirmed that the perceived attractiveness (preference and/or species' body size) succesfully predicts the size of zoo populations across mammal, bird and reptile taxa. On contrary, we found no effect of the IUCN listing of the species. To find out whether we work with Czech students' preference only or we may generalize to other populations, we carried out the same experiment to determine human preference to boas and pythons in eight cultures of five continents. Despite profound differences of the studied ethnics, we revealed a considerable agreement. Moreover, we found an agreement between pre-school...

Neophobia and exploratory behaviour are two processes applied in great measure in birds' reactions towards novel stimuli. The specific demonstration of these two processes may differ between different species of birds as well as within them. We compared neophobia and exploratory behaviour in two species of tits (Paridae). Adult Great tits (Parus major) and Blue tits (Cyanistes caeruleus) did not differ neither in exploring novel environment and novel object nor in neophobia towards novel object near food. We have found great differences between juveniles of these two species. Juvenile Blue tits were more neophobic towards both novel objects. Age, sex and personality affected some differences within tested species. We tested whether there exist any differences in reactions towards novel food between and within species. We used red painted mealworm (larvae of Tenebrio molitor), house cricket (Acheta domestica) and aposematic bug (Pyrrhocoris apterus) as novel food. Adult wild-caught tits with plentiful experience with various feeds did not differ in reaction towards these types of novel food. In contrast hand-reared juvenile tits differed quite a lot. Juvenile Blue tits were more neophobic than juvenile Great tits towards all three types of novel food. We found dieraty conservatism, phenomenon...

So far, nobody examined a human reaction on aposematic signals. I used red firebug's coloration and its modifications in color and pattern for testing the human reactions. People showed refusing reaction on white color and on some types of eye spots. There is a big difference between reactions on red color and black color in different human groups. This needs further research.

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...

Jan Raška: Reactions of the jumping spider Evarcha arcuata to aposematic true bugs Abstract: This paper studies the effect of qualities of a prey on predatory behaviour of Evarcha arcuata (Araneae: Salticidae). Naturally coloured aposematic larveae of Pyrrhocoris apterus have been compared to two types of prey: 1) to identically coloured but in defensive secretion different larvae of Scantius aegyptius, and 2) differently coloured larvae of white mutants of P. apterus. The defensive secretion of S. aegyptius has been found less effective in the first interaction with predator than that of P. apterus, however, after several trials the difference of reactions of the spider has not been noticeable. When changed the species of prey, E. arcuata has symmetrically and considerably generalised between the two species. Significant difference has been established in memory test - while P. apterus has been avoided even after one day, S. aegyptius has been attacked as thought the spiders were naive. The learning sequence has been similar in both colour forms of P. apterus; the generalisation of the two colours has been symetrical as well. Both colour forms succeeded in memory test (e.g. spiders have proven difference in comparison with the naive ones), however, white mutants have been successfully attacked...

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...

Warning meaning of several artificial as well as natural colour modifications of the Harlequin ladybird (Harmonia axyridis) has been tested in experiments with avian predators (Great tit ? Parus major). I was able to show a certain effect of black spots as their presence provided partial protection to the brown form, while the red form was protected even when unspotted.

The present PhD thesis comprises four published research papers and two manuscripts in preparation dealing with importace of particular parts of the warning signal in protection of insect prey against avian predators.

The present RNDr. thesis comprises an original research paper testing the efficacy of signal of the Batesian mimic under various familiarity of the predator with the ancestral species from which the mimic evolved.

Personality and individual differences in reactions of the Great tit to aposematic prey were investigated. The aim of this study was to assess differences in personality of forty Great tits and find out possible correlation to the reactions of these birds to aposematic prey, the fifth larval instar of firebug Pyrrhocoris apterus.