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This study aimed at complementing studies of sperm whale social and vocal behaviour that were restricted to the Pacific Ocean. The characteristic multi-pulsed structure of sperm whale clicks allows for estimation of whales' size from measurements of the inter-pulse intervals (IPI). I have developed two new automatic methods for IPI estimation from clicks recorded during foraging dives. When compared to other previously developed methods, the newly developed method that averages several clicks' autocorrelation function showed the best performance amongst the automatic methods. Previous studies did not support individual identity advertisement among social unit members as the function for the sperm whale communication signals called codas. I tested within coda type variation for individual specific patterns and found that, while some coda types do not allow for individual discrimination, one did so. This variation suggests that different coda types may have distinct functions. Analysis of social structure in the Azores found that, similar to the Eastern Tropical Pacific, sperm whales form long term social units of about 12 individuals. Unlike the Pacific Ocean, Azorean social units do not form temporary groups with other units, suggesting differences in the costs and benefits of group formation. I argue that these are due to differences in terms of predation pressure and intraspecific competition between the Azores and the Pacific study sites. The variation of coda repertoires in the Atlantic also showed a pattern dissimilar to that previously documented in the Eastern Tropical Pacific. In the North Atlantic, coda repertoire variation is mostly geographic, which is parsimoniously explained by random drift of culturally transmitted coda repertoires. No sympatric vocal clans with distinct dialects were found as has been noted in the Pacific. Drawing upon the differences found in social structure I argue that selection for maximization of differences between units with similar foraging strategies may have led to the Pacific vocal clans. The differences between oceans suggest that sperm whales may adaptively adjust their behaviour according to experienced ecological conditions.

A central issue in the study of primate communication is the extent to which individuals adjust their behaviour to the attention and signals of others, and manipulate others’ attention to communicate about external events. I investigated whether 13 chimpanzees (Pan troglodytes spp.), 11 bonobos (Pan paniscus), and 7 orangutans (Pongo pygmaeus pygmaeus) followed conspecific attention and led others to distal locations. Individuals were presented with a novel stimulus, to test if they would lead a conspecific to detect it in two experimental conditions. In one the conspecific faced the communicator, while another required the communicator to first attract the attention of a conspecific. All species followed conspecific attention, but only bonobos in conditions that required geometric attention following and that the communicator first attract the conspecific‘s attention. There was a clear trend for the chimpanzees to selectively produce a stimulus directional ‘hunching’ posture when viewing the stimulus in the presence of a conspecific rather than alone (the comparison was statistically non-significant, but very closely approached significance [p = 0.056]), and the behaviour consistently led conspecifics to look towards the stimulus. An observational study showed that ‘hunching’ only occurred in the context of attention following. Some chimpanzees and bonobos consistently and selectively combined functionally different behaviours (consisting of sequential auditory-stimulus-directional-behaviours), when viewing the stimulus in the presence of a non-attentive conspecific, although at species level this did not yield significant effects. While the design did not eliminate the possibility of a social referencing motive (“look and help me decide how to respond”), the coupling of auditory cues followed by directional cues towards a novel object, is consistent with a declarative and social referential interpretation of non-verbal deixis. An exploratory study, which applied the ‘Social Attention Hypothesis’ (that individuals accord and receive attention as a function of dominance) to attention following, showed that chimpanzees were more likely to follow the attention of the dominant individual. Overall, the results suggest that the paucity of observed referential behaviours in apes may owe to the inconspicuousness and multi-faceted nature of the behaviours.

Lucie Čadková: Lingvistické schopnosti nonhumánních živočichů 1 ABSTRACT The 20th century has witnessed significant advance in our knowledge of animal communication. Thanks to modern technology, ethologists have made great strides in decoding natural communication systems of non-human animals, while psychologist's attempts to teach a member of another species analogues of human language have met with first success. The unexpected findings called into question the unique status of human language capacities and gave rise to pressure to redefine human language in order to defend human uniqueness. One of the most influential definitions by which the communication systems of non-human animals are guaranteed a priori exclusion from the notion of language was developed by Charles F. Hockett in the 1950s and 1960s. Since then, his design-feature approach has been used, despite a series of paradigm changes in linguistics and key discoveries in cognitive ethology, in support of the claim that humans are the only living creatures endowed with language. The prevailing uncritical acceptance and usage of his theory in the field of animal communication was the impulse to write this thesis. The dissertation aims to shed light on the historical development of the question of animal linguistic abilities and presents the...

As vocalizações em anuros são comumente conspícuas e, até certo ponto, estereotipadas. Nesse grupo, o principal sinal acústico emitido por machos (canto de anúncio) basicamente codifica dois tipos de informação: a atração de fêmeas coespecíficas e o anúncio da posição do macho em atividade de vocalização, especialmente relevante na orientação espacial e delimitação de sítios de vocalização em espécies territoriais. Mais modernamente, com o devenvolvimento de métodos computacionais para incorporar a informação filogenética disponível para um dado grupo de organismos, tornou-se factível explorar tendências evolutivas e testar hipóteses adaptivas para de maneira mais apropriada, levando em conta a dependência filogenética dos táxons. Leptodactylinae é um grupo de anuros Neotropicais rico em espécies (ca. 95 spp.), cuja distribuição está essencialmente compreendida na região Neotropical. Esse táxon apresenta uma diversidade notável de vocalizações. A diversidade de padrões acústicos em Leptodactylidae por si só ressalta o potencial desse táxon representar um bom modelo para estudos focados no entendimento dos padrões e processos macroevolutivos envolvidos na evolução acústica nesse grupo de vertebrados. Nesse estudo, foi realizada uma revisão detalhada dos sinais acústicos reprodutivos, i.e. canto de anúncio, pela primeira vez analisados para esse táxon através de procedimentos e configurações padronizadas e com quantificações automatizadas. Os resultados das análises acústicas foram então associados aos padrões morfológicos na investigação dos padrões de diversidade críptica e reavaliação taxonômica de algumas populações e espécies em Leptodactylinae, além da abordagem comparativa a partir dos resultados obtidos das análises acústicas a fim de explorar as principais tendências e processos evolutivos dos sinais acústicos nesse grupo de anuros. Os resultados revelaram que existem diferentes padrões de diferenciação fenotípica nesse táxon, apontando para a necessidade de revisões taxonômicas em diferentes clados de Leptodactylinae, com resultados contundentes para o clado L. melanonotus (a reavaliação de sinônimos juniores nesse clado indicaram que duas espécies devem ser revalidadas) e Adenomera (reconhecimento de diversidade críptica em A. hylaedactyla com base em evidência acústica). A exploração de métodos comparativos filogenéticos relevou tendências evolutivas interessantes, como, por exemplo, os surgimentos e desaparecimentos recorrentes de estruturas temporais (pulsos) e mudanças acentuadas na estrutura espectral dos sinais acústicos (deslocamento da frequência fundamental em algumas espécies de Adenomera), tais fenômenos sem relação aparente com tamanho corporal ou estrutura dos habitats ocupados por essas espécies. Os dados acústicos oriundos desse estudo se mostraram eficazes nas resoluções taxonômicas em várias espécies de Leptodactylinae. Estudos futuros focando em aspectos da morfologia do aparato vocal, e biomecânica de produção de som, podem informativos acerca do entendimento dos processos envolvidos em distintos padrões acústicos reconhecidos em Leptodactylinae / In anurans, vocalizations are usually conspicuous and stereotyped to some extent. In this group, the major acoustic signal broadcast by males (advertisement signal/call) basically encodes two kinds of information: the attraction of conspecific females and the advertisement of the position of a calling male, maintaining male spacing functioning as an auditory cue in the delimitation of calling sites in territorial species. More recently, the development of computational methods as a means of incorporation of phylogenetic information available for a given group of organisms allowed scientists to explore evolutionary trends and to test for adaptative hypotheses more properly, taking into consideration phylogenetic dependence among taxa. Leptodactylinae is a species-rich taxon (ca. 95 spp.), whose distribution encompasses the Neotropical region. This frog group exhibits a remarkable diversity of vocalizations. This pattern in Leptodactylinae draws attention to the fact that this group could be a promising model organism to studies focused on macroevolutionary patterns and processes involved in the acoustic evolution in this vertebrate group. In the present study, a comprehensive acoustic review was conducted, and it was the first time for this group that the analytical procedures and standardized settings were employed in an automated fashion for the quantification of acoustic traits. The results of the acoustic analysis was then associated with morphological patterns as a means to investigate the patterns of cryptic diversity and taxonomic reappraisals for some populations and species, as well as the application of comparative methods from the results obtained to explore the major evolutionary trends and processes of acoustic traits in this frog group. The results revealed that there are distinctive patterns of phenotypic differentiation present across Leptodactylinae clades, which indicate that taxonomic reassessments are required, particularly in the L. melanonotus clade (revaluation of the junior synonyms showed that two species should be relavalidated) and Adenomera (cryptic diversity was uncovered under the name A. hylaedactyla based on acoustic evidence). The phylogenetic comparative methods recovered interesting evolutionary trends as, for instance, the convergent appearances and losses of structures related to the time domain (pulses) and remarkable changes in the frequency domain (doubling of the fundamental frequency in some species of Adenomera), both phenomena being apparently unrelated to differences in body size or calling habitats. The acoustic data gathered here was efficient in species discrimination and should represent a good line of evidence to address species-level taxonomy in Leptodactylinae. Future research focused on aspects of the vocal apparatus and the mechanisms of sound production might tell us more about the evolutionary processes involved in the distinctive acoustic patterns that were recognized in Leptodactylinae

Este é um estudo comparativo do comportamento e da comunicação acústica de cobaias – Cavia porcellus - e de preás – C. aperea - duas espécies muito próximas, uma com uma história recente de domesticação (há 6.000 anos nos Andes) e a outra selvagem. Inclui três etapas: (1) a elaboração de um etograma e a comparação entre cobaias e preás quanto à freqüência e duração dos comportamentos exibidos em contextos sociais particulares (encontros entre fêmeas, entre machos e entre macho e fêmea); (2) a categorização e análise sonográfica comparativa da estrutura dos sinais sonoros; e (3) a comparação entre espécies quanto ao uso dos chamados através de um estudo do comportamento do emissor e de um receptor, no momento anterior e subseqüente à emissão. Encontramos diferenças entre as espécies nas três etapas. (1) Cobaias exibiram mais comportamentos de contatos e sexuais; preás exibiram mais comportamentos de exploração. (2) Das emissões registradas, uma não foi exibida por cobaias. Analisamos 4 delas estatisticamente e todas revelaram diferenças estruturais entre espécies. (3) Houve diferença no uso do sinal de alerta e no nível de resposta eliciado por alguns sinais. Os resultados são discutidos principalmente como efeitos da domesticação. A seleção de animais maiores pode ter alterado a morfologia do trato vocal das cobaias, produzindo as mudanças na estrutura dos chamados. Ainda, com o relaxamento de seleção natural, por conta da proteção oferecida pelo cativeiro, características menos favoráveis podem ter sido mantidas e ganhado proporção na população. Como resultado, a espécie domesticada produz e reage menos aos sinais de alerta, gasta menos tempo com comportamentos de atenção ao meio e ganha tempo para os comportamentos socais e reprodutivos. / This is a comparative study of behavior and acoustic communication of wild (Cavia aperea) and domestic (C. porcellus) cavies. Animals were observed in the laboratory in three social situations (female-female, male-male and female-male pairings) and the following steps were performed: (1) comparison of the frequency and duration of behaviors, in both species; (2) description and comparison of the sonografic parameters of acoustic signals emitted by individuals of both species; and (3) identification of antecedent and subsequent behavioral contexts of such signals as an approach to the understanding of their social function. C. porcellus exhibited more contact and sexual categories of behaviors than C. aperea; C. aperea explored more than C. porcellus. Signal repertoire was almost the same (one, out of 7 calls was exclusively emitted by C. aperea). Statistical analysis revealed significant structural differences between species in four of the calls. Differences were also found in the context of use of alert vocalization and in the level of response elicited by some signals. Interspecific differences found may be partially attributed to domestication. Selection for meet production may have altered guinea pigs’ vocal tract and may have brought changes in acoustic signals structure. The absence of predatory pressure and the less demanding conditions of captivity may have favored the expression of some traits, such as the performance of long courtship bouts. The domestic cavies C. porcellus are less prone than the wild ones to emit and to respond to alert signals, spend less time with exploratory and patrolling and spend more time with social and reproductive interactions.

Este é um estudo comparativo do comportamento e da comunicação acústica de cobaias – Cavia porcellus - e de preás – C. aperea - duas espécies muito próximas, uma com uma história recente de domesticação (há 6.000 anos nos Andes) e a outra selvagem. Inclui três etapas: (1) a elaboração de um etograma e a comparação entre cobaias e preás quanto à freqüência e duração dos comportamentos exibidos em contextos sociais particulares (encontros entre fêmeas, entre machos e entre macho e fêmea); (2) a categorização e análise sonográfica comparativa da estrutura dos sinais sonoros; e (3) a comparação entre espécies quanto ao uso dos chamados através de um estudo do comportamento do emissor e de um receptor, no momento anterior e subseqüente à emissão. Encontramos diferenças entre as espécies nas três etapas. (1) Cobaias exibiram mais comportamentos de contatos e sexuais; preás exibiram mais comportamentos de exploração. (2) Das emissões registradas, uma não foi exibida por cobaias. Analisamos 4 delas estatisticamente e todas revelaram diferenças estruturais entre espécies. (3) Houve diferença no uso do sinal de alerta e no nível de resposta eliciado por alguns sinais. Os resultados são discutidos principalmente como efeitos da domesticação. A seleção de animais maiores pode ter alterado a morfologia do trato vocal das cobaias, produzindo as mudanças na estrutura dos chamados. Ainda, com o relaxamento de seleção natural, por conta da proteção oferecida pelo cativeiro, características menos favoráveis podem ter sido mantidas e ganhado proporção na população. Como resultado, a espécie domesticada produz e reage menos aos sinais de alerta, gasta menos tempo com comportamentos de atenção ao meio e ganha tempo para os comportamentos socais e reprodutivos. / This is a comparative study of behavior and acoustic communication of wild (Cavia aperea) and domestic (C. porcellus) cavies. Animals were observed in the laboratory in three social situations (female-female, male-male and female-male pairings) and the following steps were performed: (1) comparison of the frequency and duration of behaviors, in both species; (2) description and comparison of the sonografic parameters of acoustic signals emitted by individuals of both species; and (3) identification of antecedent and subsequent behavioral contexts of such signals as an approach to the understanding of their social function. C. porcellus exhibited more contact and sexual categories of behaviors than C. aperea; C. aperea explored more than C. porcellus. Signal repertoire was almost the same (one, out of 7 calls was exclusively emitted by C. aperea). Statistical analysis revealed significant structural differences between species in four of the calls. Differences were also found in the context of use of alert vocalization and in the level of response elicited by some signals. Interspecific differences found may be partially attributed to domestication. Selection for meet production may have altered guinea pigs’ vocal tract and may have brought changes in acoustic signals structure. The absence of predatory pressure and the less demanding conditions of captivity may have favored the expression of some traits, such as the performance of long courtship bouts. The domestic cavies C. porcellus are less prone than the wild ones to emit and to respond to alert signals, spend less time with exploratory and patrolling and spend more time with social and reproductive interactions.

The social transmission of social behaviours in nonhuman primates has been understudied, experimentally, relative to instrumental, food-related behaviours. This is disproportional in relation to the comparatively high percentage of potential social traditions reported in wild primates. I report a systematic survey of the social learning literature and provide quantitative evidence of the discrepancy (Watson and Caldwell, 2009). Addressing the identified deficit in experimental work on social behaviours, I also report three empirical studies investigating the contagious nature of affective states in captive, socially housed marmosets. I carried out an observational study, to determine whether marmosets are influenced by spontaneously produced neighbour calls to perform a range of behaviours associated with similar affect. My results supported a neighbour effect for anxiety in marmosets. Consistent with previous findings for chimpanzees (Baker and Aureli, 1996; Videan et al., 2005), I also found evidence for neighbour effects for aggression and affiliation (Watson and Caldwell, 2010). Through experimental playback, I investigated contingent social contagion in the auditory and visual modalities. The playback of pre-recorded affiliative (chirp) calls was found to be associated with marmosets spending increased time in a range of affiliative behaviours. Playback of video showing conspecifics engaged in a positive affiliative behaviour (allogrooming) also appeared to cause marmosets to spend longer performing various affiliative behaviours. My results indicate that social contagion of affiliation is a multi-modal phenomenon in marmosets and also represent the first evidence that allogrooming is visually contagious in primates. Sapolsky (2006) conceptualised culture as the performance of species-typical behaviours to an unusual extent, termed ‘social culture’. Researchers have yet to directly investigate a transmission mechanism. I investigated whether a social culture of increased affiliation could be initiated in marmosets through the long-term playback, of positive calls, or of video of positive behaviour. The results were consistent with a relatively long-lasting influence of the playback of affiliative calls across several affiliative behaviours. The effect appeared to last substantially beyond the specific hours of playback, between playbacks, and after playback had ceased, potentially indicating a temporary shift in social culture. These results are preliminary but provide some support for the proposal that auditory social contagion may be a transmission mechanism for social culture. The long-term video playback of allogrooming appeared to result in a transitory shift in performance of the identical behaviour (increased allogrooming) after playbacks had ceased. In addition to theoretical implications for social cognition and social culture, my findings have potential practical application for the enhancement of welfare in captive marmosets through sensory, and non-contact social, enrichment.

Flamboyant colors are widespread throughout the animal kingdom. While many of these traits arise through sexual selection, bright coloration can also evolve through natural selection. Many aposematic species, for example, use conspicuous warning coloration to communicate their noxiousness to predators. Recent research suggests these signals can also function in the context of mate choice. Studies of warning color evolution can therefore provide new insights into how the interplay of natural and sexual selection impact the trajectory of conspicuous signal evolution. For my dissertation, I investigated the potential for male-male competition to impact the warning color evolution of a species of poison frog. I focused my work on an exceptionally bright and toxic population of the strawberry poison frog (Oophaga pumilio) where males are brighter than females, a classic signature of sexual selection. In Chapter 1, I used theoretical models of predator and frog visual systems to determine which can see the variation in bright warning coloration within this population. I found that birds, the presumed major predator, likely cannot see this variation, indicating that sexual selection can work under the radar of predators in this species. In Chapter 2, I tested the aggressive responses of males using a two-way choice paradigm that manipulated the perceived brightness of stimulus males. I found that males directed more of their behaviors to bright stimulus frogs, and brighter focal frogs more readily approached stimuli and directed more of their attention to the brighter rival. In Chapter 3, I tested the outcomes of dyadic interactions between males of varying brightness and observed male reactions to simulated intruders in their territories. I found that brighter males initiated aggressive interactions with rivals more readily, and brightness asymmetries between males settled interactions in a way that is consistent with classic hypotheses about male sexual signals. In Chapter 4 I sought to describe physiological correlates of male warning color brightness. While male brightness did not co-vary with classic measures of body condition (circulating testosterone and skin carotenoids), it did correlate with toxins sequestered from the diet and thus appears to be a reliable signal of toxicity in this population. / text

The freshwater prawn, Macrobrachium rosenbergii (de Man, 1879) is an important aquaculture species but one that has the disadvantage of heterogeneous individual growth (HIG) according to different morphotypes. Chemical cues, especially, pheromones, are one of the most important communication types between individual prawns, along with visual and tactile methods. Testing pheromones, whilst restricting other cues, may therefore lead to a better understanding of the influence of these communicatory compounds on the prawn reproductive process. The three principle objectives of this study were therefore: 1) To examine the effect of moult stage and morphotype on pheromone-induced sexual behaviour 2) To examine the role of pheromone / urine concentrations on sexual attraction behaviour 3) To describe the functional morphology of the antennal gland and examine its possible role in pheromone production and release. Identical bioassay tanks were designed and constructed to study the reproductive behaviour of prawns. Experiments were set up to examine responses to pheromone release by live prawns over 30 minutes and behavioural response observations were made with the aid of a Closed-Circuit Videotape System (CCVS). Results were statistically analysed using a repeated measures general linear model (GLM). Three trials were designed to test the effect of moult stage of both males and females and male morphotypes on sexual attraction behavioural responses. Twelve prawns were used for each trial and each prawn was used five times (1 no-pheromone control and 4 for experimental tests). The first trial studied the effect of female moult stages (pre-, inter and newly-moulted) on sexual attraction behaviour of blue claw (BC) male. Results of this trial showed that newly-moulted females spent significantly (p<0.05) less time approaching the BC male than the pre- and inter-moult females. The second trial studied the effect of male moult stage (pre-, inter and newly-moulted) on sexual attraction to receptive females. Results showed that the time taken by the inter-moult males was (p<0.05) less than the pre- and newly-moulted males in approaching the newly-moulted female. The third trial tested the effect of male morphotypes (small male, SM, orange claw, OC and dominant blue claw, BC) on sexual attraction behaviour towards newly-moulted females. Results showed that the BC male was significantly more attractive (p<0.05) than other morphotypes to newly-moulted females and that the OC male was the least attractive. The role of moulting stage for both male and female prawns on reproductive response behaviour was investigated. Because BC males responded significantly faster towards newly-moulted female more than to either pre-or inter-moult females, results of the first trial suggest that BC males are able to use different chemical cues to gather information about a conspecific’s gender and can differentiate female’s moult stages. Since BC males responded significantly faster towards newly-moulted females more than to either pre-or inter-moult females, this suggests that females at this particular stage released a distinct sexual pheromone or concentration of pheromone that differed from those pheromones released by both pre- and inter-moult females. In contrast, newly-moulted females prefer the inter-moult BC males which indicate that females have an ability to distinguish the moult status of BC males. Furthermore, it indicates that pheromone characteristics change with the moult status of BC males. Also, newly-moulted females are most likely to be avoiding the potential costs of mate guarding with soft shell BC males. Results obtained from the third trial suggested that a newly-moulted female can discriminate male morphotypes (SM, OC and BC) from their pheromone cues. This indicates that male morphotypes release pheromones which differ from each other in some way. Newly-moulted females responded positively to both SM and BC males with different levels of attraction with the greatest attraction to BC males to BC males suggesting that pheromone released from the BC male may carry information relating to dominance status. Urine is believed to be one of the main carriers of pheromone and is usually released from the antennal gland. Different urine concentrations (0.1, 1.0, 2.0, 3.0, 5.0 and 10µl l-1) of collected urine from BC males were used to test the sexual attraction behaviour of receptive newly-moulted females. Also, the attractant capability of fresh urine following exposure to different temperature regimes (cooled at 4ºC, frozen at -70ºC and heated at 70ºC) was tested. Since newly-moulted female M. rosenbergii were attracted to BC male urine, this indicates the existence of sex pheromone in the fresh urine. Also, it was found that the sexual response of females to fresh urine of BC males was directly proportional to urine concentration with faster responses observed with increasing urine concentrations. At the three fresh urine concentrations 0.1 µl l-1, 1.0 µl l-1 and 2.0 µl l-1, statistical analysis indicated no significant difference (p>0.05) between these three concentrations while a significant (P<0.05) response was to concentrations more than 3.0 µl l-1. This may indicate that these three concentrations were not sufficient to elicit attraction behaviour in newly-moulted females. A concentration of 3.0 µl l-1 of fresh urine is suggested to be a sufficient concentration to elicit a significant sexual attraction under laboratory conditions. Response of newly-moulted female prawns to the various temperature treatments tested declined in response to nominally increasingly degradative treatments. Also, statistical analysis showed that temperature treatment and concentration added both had a significant effect on the response of females. The greatest degradation of urine attractiveness was found with the 70ºC heat treatment. It can be concluded that the pheromone components of prawn urine are friable when exposed to high temperatures. Using light and transmission electron microscopes, ultrastructural observation of the antennal gland (AG) of M. rosenbergii suggests that it has four distinct regions, the coelomosac, the nephridial tubules, the labyrinth and the bladder. Morphological and functional descriptions of each of these regions were compared with those of other aquatic Crustacea.