Formosan macaques were observed to collect data on intergroup encounters, the strategies of male and female and the factors which influence inter-group dominance relationships. Four hypotheses exist to explain the strategies of male and female in inter-group encounter, including female resource defense, male mate defense, male resource defense and male mate attraction via infanticide. Behavioral observations were recorded from March 2008 to March 2009. Data collections included scan sampling and focal sampling of troop members which participated in encounter, containing their sex, age, rank and behavior. Duration of inter-group interaction and distances between two troops were also recorded. There is a significant positive relationship between chance of winning and troop number (AM+AF+SAM), and this chance of winning is higher than 50% when troop size exceeded 30. There is also a significant linear regression between weight win¡]main group and branch group¡^and troop number , but chance of weight win in some troops is higher than 90% when troop size is only between 20 to 30. Approach, line-up, displace and be-displaced have significant correlation with troop size type. Large troops displayed higher displace (65%), line-up (6%) and approach (28%) than other type of troops, and small troop were easily displaced by other type of troops (60%). Aggressive behavior also has significant correlation with mating season significantly; troops of Formosan macaques display more aggressive behavior in mating season (70%) than in non-mating season (41%). Troops of Formosan macaques display higher approach¡]33%¡^ and displace (93%) when troops encounter with all male troop than with bisexual troop. Inter-group dominance relationships exist among main group and branch group which newly split of Formosan macaques. But there is uncertain inter-group dominance relationships between main group and branch group which split early for the difference of maternal dominance hierarchies, variation of troop numbers, and the character of central males. Adult male and female Formosan macaques adopt different strategy in inter-group encounter. Adult female Formosan macaques participated (8.62%, n=8) and displayed aggressive behavior (3.66%, n=8) in more food-related encounter than in non-food-related encounter, so female resource defense hypothesis is supported. Adult male Formosan macaques display higher aggressive behavior (7.92%, n=23) significantly than adult female and sub-adult male in inter-group encounter, and the target of aggression is higher for males than for females. Adult and sub-adult male Formosan macaques display higher aggressive behavior in mating season significantly than in non-mating season. So male mate defense hypothesis is also supported. This study provides support for the male resource defense hypothesis. Adult male Formosan macaques would defend food resource directly and indirectly. Besides, frequencies of participation and aggression in low ranking adult male Formosan macaques are higher than high ranking ones to exchange mating opportunity.

No description available.

Cavidades naturais constituem um recurso reprodutivo monopolizável, cuja disponibilidade determina a intensidade da competição macho-macho que, por sua vez, pode influenciar os custos do cuidado paternal, pois quanto maior o risco de perda do recurso, menor deve ser a freqüência de forrageio dos machos. Fêmeas do opilião Magnispina neptunus utilizam cavidades naturais como sítios de nidificação, que são monopolizados por machos em um sistema de poliginia por defesa de recursos. Após a oviposição, as fêmeas abandonam os ninhos e todo o cuidado parental é exercido pelos machos. O objetivo deste trabalho foi avaliar o efeito da disponibilidade de ninhos sobre a competição intrassexual e os custos do cuidado paternal em M. neptunus. Dois grupos experimentais foram estabelecidos em laboratório: alta (8 ninhos) e baixa (4 ninhos) disponibilidade de sítios reprodutivos. Cada grupo era composto por sete terrários contendo 12 machos e 12 fêmeas. Apesar da redução dos sítios de nidificação ter reduzido o número de machos detentores de ninhos, o tamanho dos indivíduos não exerceu nenhum efeito sobre o sucesso de monopolização de ninhos. Aparentemente, o pequeno diâmetro de entrada dos ninhos experimentais reduz a importância do tamanho dos machos sobre sua probabilidade de monopolizar um sítio de nidificação. Apenas a residência prévia parece determinar o resultado das disputas entre machos e, portanto, os custos de abandonar os ninhos devem ser altos. De fato, machos guardiões permanecem quase todo o tempo dentro dos seus ninhos em ambos os grupos experimentais, o que aumenta os custos do cuidado. Como conseqüência, o canibalismo filial é freqüente, pois os custos em termos de redução no tamanho da desova são menores do que os benefícios de manter a posse do ninho e dos ovos, que sabidamente aumentam a atratividade dos machos e suas chances de obterem novas desovas / Natural cavities are a reproductive resource that can be monopolized, and their availability determines the intensity of male-male competition, which in turn may influence the costs of paternal care since the higher the risk of loosing the resource, the low should be male foraging frequency. Females of the harvestman Magnispina neptunus use natural cavities as nest site, which are monopolized by males in typical resource defense polygyny mating system. After oviposition, females leave the nests and all parental activities are in charge of males. The goal of this study was to evaluate the effect of nest site availability on the intrasexual competition and the costs of paternal care in M. neptunus. Two experimental groups were established in the laboratory: high (8 nests) and low (4 nests) nest site availability. Each group was composed of seven terraria containing 12 males and 12 females. Although the reduction of in the number of nests sites has indeed reduced the number of males holding nests, male size did not influence the probability of acquiring and maintaining a nest. Apparently, the small diameter of entrance in the experimental nests reduces the importance of male size on their resource holding power. Only previous residence seems to influence the output of agonistic interactions between two males and, therefore, the costs of leaving the nest to forage are likely to be high. In fact, guarding males remain almost all the time inside their nests, which increases the costs of paternal care. As a consequence, filial cannibalism is frequent in both experimental groups because the costs in terms of reduction in clutch size are likely to lower than the benefits of holding a nest containing eggs, which is known to increase male attractiveness and his chance of acquiring additional eggs

Zahlreiche Forschungsarbeiten haben gezeigt, dass nicht nur Menschen, sondern auch Tiere konstante individuelle Unterschiede im Verhalten aufweisen. Zu verstehen warum sich diese Verhaltensunterschiede im Laufe der Evolution entwickelt haben, ist ein Ziel dieses Forschungsbereiches. In dieser Arbeit wurde untersucht wie verschiedene Modulatoren das Nahrungssuchverhalten von Blütenfledermäusen (Glossophaginae) beeinflussen um individuelle Verhaltensunterschiede zu quantifizieren und theoretische Vorhersagen zu testen. Alle Experimente wurden in naturnaher Umgebung mit programmierbaren, künstlichen Blüten durchgeführt. Es wird angenommen, dass die Plastizität von Verhalten ein generelles Merkmal ist in dem sich Tiere unterscheiden, da manche Individuen allgemein stärker auf Reize aus der Umwelt reagieren könnten als andere. Um diese Vorhersage zu testen, wurde die Nahrungsverfügbarkeit experimentell manipuliert und zwei Arten von Verhaltensplastizität in denselben Individuen gemessen. Die Ergebnisse unterstützen diese Annahme jedoch nicht, da die beiden Arten von Verhaltensplastizität nicht korrelieren. Neben Umwelteinflüssen können auch innere Merkmale wie die Stoffwechselrate das individuelle Nahrungssuchverhalten beeinflussen. Die Ergebnisse zeigen, dass sich die Individuen in ihrem täglichen Energieverbrauch unterscheiden und dass diese Unterschiede mit dem Explorationsverhalten während der Nahrungssuche korrelieren. Zusätzlich kann das individuelle Nahrungssuchverhalten auch von sozialen Faktoren beeinflusst werden. Bei limitierter Nahrungsverfügbarkeit verteidigen einzelne Männchen Blüten gegen andere Männchen, jedoch nicht gegenüber Weibchen. Individuelle Unterschiede in der Aggression und Aktivität werden dagegen nicht von der sozialen Gruppenzusammensetzung beeinflusst. In dieser Arbeit wurden nicht nur individuelle Unterschiede im Nahrungssuchverhalten von Blütenfledermäusen bestimmt, sondern auch Vorhersagen aus dem Bereich der Persönlichkeitsforschung bei Tieren überprüft. / Animal personality research has shown that animals express individual differences in their behavior that are consistent over time and/or across situations. Furthermore, animal personality research aims to understand how these individual differences in behavior evolve and how they are maintained within populations. This thesis focuses on how different modulators influence the foraging behavior of nectar-feeding bats (Glossophaginae) in order to investigate consistent individual differences in their behavior and to test predictions proposed in the field of animal personality. All experiments were conducted in a semi-natural environment by using a setup of computer-controlled artificial flowers. One prediction of animal personality research is that behavioral plasticity is a single trait in which individuals differ because some individuals might be generally more responsive to changes in the environment than others. In order to test this prediction, resource availability was manipulated and two types of behavioral plasticity were quantified within the same individual. However, the two types of plasticity did not correlate and therefore this hypothesis was not supported. Individual foraging behavior can not only be modulated by changes in the environment but also by individual differences in internal traits like metabolic rates. Experiments confirmed that individual differences in daily energy expenditure correlated with consistent individual differences in exploration. Additionally, foraging behavior can also be modulated by social factors. When resources are limited some males started to defend flowers against other males but not against females. Furthermore, it could be shown that individual differences in aggression and activity are internally driven and not influenced by changes in the social group composition. This thesis not only assessed individual differences in the foraging behavior of nectar-feeding bats but also investigated predictions proposed in the field of animal personality research.

I describe the natural history and reproductive behaviours of the Emei Moustache Toad (Leptobrachium boringii), testing the hypotheses that the species exhibits resource defense polygyny, and that combat, and paternal care lead to the evolution of male-biased sexual size dimorphism. In this study I document combat behaviour and paternal care for the first time in this species. Between February and March of 2011 and 2012, 26 female and 55 male L. boringii from Mount Emei UNESCO World Heritage Site, Sichuan, China, were observed throughout the breeding season. Prior to the breeding season, males grow 10-16 keratinized maxillary spines, which fall off once the season has ended. Throughout this time, males construct and defend aquatic nests where they produce advertisement calls to attract females. In a natural setting, I documented 14 cases involving a total of 22 males where males used their moustaches for aggressive interaction, and nest take over was observed on seven occasions. Despite my predictions, neither male body size nor body condition significantly affects the outcome of an aggressive interaction, though this may be representative of a low sample size. Males were also observed to possess injuries resulting from combat. Combat trials conducted in artificial nests demonstrated heightened aggression from resident males towards intruders. Genetic analysis using microsatellite markers revealed several cases of multiple paternity, both within nest and within clutch, indicating that some alternative male reproductive strategy, such as satellite behaviour is occurring. Larger males were observed to mate more frequently, and in multiple nests, suggesting that females are selecting for larger males, or that larger males are more capable of defending high quality territories. Males showed evidence of paternal care behaviours by remaining with the nests once females had left, moving throughout the nest cleaning, touching the eggs, and blowing bubbles into the centre of the doughnut-shaped egg masses. From this study I conclude that the male biased sexual size dimorphism in L. boringii is likely the result of both combat and paternal care behaviours creating a selection pressure on male body size.

Cavidades naturais constituem um recurso reprodutivo monopolizável, cuja disponibilidade determina a intensidade da competição macho-macho que, por sua vez, pode influenciar os custos do cuidado paternal, pois quanto maior o risco de perda do recurso, menor deve ser a freqüência de forrageio dos machos. Fêmeas do opilião Magnispina neptunus utilizam cavidades naturais como sítios de nidificação, que são monopolizados por machos em um sistema de poliginia por defesa de recursos. Após a oviposição, as fêmeas abandonam os ninhos e todo o cuidado parental é exercido pelos machos. O objetivo deste trabalho foi avaliar o efeito da disponibilidade de ninhos sobre a competição intrassexual e os custos do cuidado paternal em M. neptunus. Dois grupos experimentais foram estabelecidos em laboratório: alta (8 ninhos) e baixa (4 ninhos) disponibilidade de sítios reprodutivos. Cada grupo era composto por sete terrários contendo 12 machos e 12 fêmeas. Apesar da redução dos sítios de nidificação ter reduzido o número de machos detentores de ninhos, o tamanho dos indivíduos não exerceu nenhum efeito sobre o sucesso de monopolização de ninhos. Aparentemente, o pequeno diâmetro de entrada dos ninhos experimentais reduz a importância do tamanho dos machos sobre sua probabilidade de monopolizar um sítio de nidificação. Apenas a residência prévia parece determinar o resultado das disputas entre machos e, portanto, os custos de abandonar os ninhos devem ser altos. De fato, machos guardiões permanecem quase todo o tempo dentro dos seus ninhos em ambos os grupos experimentais, o que aumenta os custos do cuidado. Como conseqüência, o canibalismo filial é freqüente, pois os custos em termos de redução no tamanho da desova são menores do que os benefícios de manter a posse do ninho e dos ovos, que sabidamente aumentam a atratividade dos machos e suas chances de obterem novas desovas / Natural cavities are a reproductive resource that can be monopolized, and their availability determines the intensity of male-male competition, which in turn may influence the costs of paternal care since the higher the risk of loosing the resource, the low should be male foraging frequency. Females of the harvestman Magnispina neptunus use natural cavities as nest site, which are monopolized by males in typical resource defense polygyny mating system. After oviposition, females leave the nests and all parental activities are in charge of males. The goal of this study was to evaluate the effect of nest site availability on the intrasexual competition and the costs of paternal care in M. neptunus. Two experimental groups were established in the laboratory: high (8 nests) and low (4 nests) nest site availability. Each group was composed of seven terraria containing 12 males and 12 females. Although the reduction of in the number of nests sites has indeed reduced the number of males holding nests, male size did not influence the probability of acquiring and maintaining a nest. Apparently, the small diameter of entrance in the experimental nests reduces the importance of male size on their resource holding power. Only previous residence seems to influence the output of agonistic interactions between two males and, therefore, the costs of leaving the nest to forage are likely to be high. In fact, guarding males remain almost all the time inside their nests, which increases the costs of paternal care. As a consequence, filial cannibalism is frequent in both experimental groups because the costs in terms of reduction in clutch size are likely to lower than the benefits of holding a nest containing eggs, which is known to increase male attractiveness and his chance of acquiring additional eggs

The availability of reproductive sites is a major factor shaping the behavior of males and females in species with resource-based mating systems. Using a meta-analytic approach, we tested five predictions directly or indirectly derived from the mating system theory. We expected that reproductive site limitation would lead to: (1) intense male-male competition for resource possession; (2) high variance in male reproductive success, generating high values of opportunity for sexual selection; (3) high intensity of selection on male traits related to resource possession; (4) high sperm competition risk; and (5) low quality of paternal care. We compiled information from observational and experimental studies that compared the reproductive behavior of individuals of the same species under low and high reproductive site limitation. We found that, when reproductive sites are limited, there is a slight increase in male-male competition, with higher rates of nest takeover and agonistic interactions, and a slight increase in the selection gradient on male traits, with successful males tending to be larger than unsuccessful males. Reproductive site limitation has no consistent effect on the opportunity for sexual selection and on the sperm competition risk. However, territorial males invest more in gonads and lose less paternity when reproductive sites are limited. There is also no clear effect of reproductive site limitation on the quality of paternal care, but few studies have addressed this subject. Taken together, our findings indicate that predictions on how reproductive site limitation affects several aspects of resource-based mating systems have weak empirical support. These predictions do not consider the plasticity in the mating tactics of males and females, which make them too simplistic. Moreover, since the original proposition of the mating system theory, our understanding of sperm competition increased a lot. We now know that accurate predictions on the intensity and direction of sexual selection should take into account both pre- and post-copulatory processes. Finally, the interplay between sexual selection and parental care is complex, and the original framework of mating system theory does not provide sufficient elements to derive clear and taxonomically broad predictions / A disponibilidade de sítios reprodutivos influencia tanto o comportamento de machos quanto de fêmeas em espécies cujo sistema de acasalamento depende da defesa de recursos. Usando uma abordagem meta-analítica, testamos cinco previsões direta ou indiretamente relacionadas à teoria de sistemas de acasalamento. Esperávamos que a limitação de sítios reprodutivos promoveria: (1) aumento na competição masculina pela posse de recursos; (2) aumento na variância do sucesso reprodutivo dos machos, gerando valores alto de oportunidade para a seleção sexual; (3) aumento da intensidade da seleção sobre características masculinas relacionadas à posse de recursos; (4) aumento no risco de competição espermática e (5) diminuição da qualidade do cuidado paternal. Compilamos informações de estudos observacionais e experimentais que compararam o comportamento reprodutivo de indivíduos da mesma espécie em situação de alta e baixa disponibilidade de sítios reprodutivos. Encontramos que, quando os sítios reprodutivos são escassos, há um ligeiro aumento na competição masculina, com maiores taxas de roubo de ninhos e interações agonísticas, e um ligeiro aumento no gradiente de seleção sobre características masculinas, com machos bem sucedidos tendendo a ser maiores do que machos mal sucedidos em monopolizar recursos. A disponibilidade de sítios reprodutivos não teve nenhum efeito consistente sobre a oportunidade para a seleção sexual e o risco de competição espermática. Entretanto, machos territoriais investiram mais em gônadas e perderam menos paternidade quando os sítios reprodutivos eram escassos. Não encontramos nenhum efeito da disponibilidade de sítios reprodutivos sobre a qualidade do cuidado paternal. Em conjunto, nossos resultados indicam que as previsões sobre como a disponibilidade de sítios reprodutivos influencia diversos aspectos de sistemas de acasalamento baseados na defesa de recursos têm fraco respaldo empírico. Tais previsões não consideram a plasticidade nas táticas de acasalamento de machos e fêmeas, tornando-as demasiadamente simplistas. Adicionalmente, desde a formulação original da teoria de sistemas de acasalamento, nossa compreensão sobre competição espermática aumentou. Sabemos atualmente que previsões acuradas sobre a intensidade e direção da seleção sexual devem levar em consideração processos pré- e pós-copulatórios. Finalmente, a interconexão entre seleção sexual e cuidado parental é complexa e a teoria original de sistemas de acasalamento não provê elementos suficientes para a construção de previsões claras e de amplo escopo taxonômico

The availability of reproductive sites is a major factor shaping the behavior of males and females in species with resource-based mating systems. Using a meta-analytic approach, we tested five predictions directly or indirectly derived from the mating system theory. We expected that reproductive site limitation would lead to: (1) intense male-male competition for resource possession; (2) high variance in male reproductive success, generating high values of opportunity for sexual selection; (3) high intensity of selection on male traits related to resource possession; (4) high sperm competition risk; and (5) low quality of paternal care. We compiled information from observational and experimental studies that compared the reproductive behavior of individuals of the same species under low and high reproductive site limitation. We found that, when reproductive sites are limited, there is a slight increase in male-male competition, with higher rates of nest takeover and agonistic interactions, and a slight increase in the selection gradient on male traits, with successful males tending to be larger than unsuccessful males. Reproductive site limitation has no consistent effect on the opportunity for sexual selection and on the sperm competition risk. However, territorial males invest more in gonads and lose less paternity when reproductive sites are limited. There is also no clear effect of reproductive site limitation on the quality of paternal care, but few studies have addressed this subject. Taken together, our findings indicate that predictions on how reproductive site limitation affects several aspects of resource-based mating systems have weak empirical support. These predictions do not consider the plasticity in the mating tactics of males and females, which make them too simplistic. Moreover, since the original proposition of the mating system theory, our understanding of sperm competition increased a lot. We now know that accurate predictions on the intensity and direction of sexual selection should take into account both pre- and post-copulatory processes. Finally, the interplay between sexual selection and parental care is complex, and the original framework of mating system theory does not provide sufficient elements to derive clear and taxonomically broad predictions / A disponibilidade de sítios reprodutivos influencia tanto o comportamento de machos quanto de fêmeas em espécies cujo sistema de acasalamento depende da defesa de recursos. Usando uma abordagem meta-analítica, testamos cinco previsões direta ou indiretamente relacionadas à teoria de sistemas de acasalamento. Esperávamos que a limitação de sítios reprodutivos promoveria: (1) aumento na competição masculina pela posse de recursos; (2) aumento na variância do sucesso reprodutivo dos machos, gerando valores alto de oportunidade para a seleção sexual; (3) aumento da intensidade da seleção sobre características masculinas relacionadas à posse de recursos; (4) aumento no risco de competição espermática e (5) diminuição da qualidade do cuidado paternal. Compilamos informações de estudos observacionais e experimentais que compararam o comportamento reprodutivo de indivíduos da mesma espécie em situação de alta e baixa disponibilidade de sítios reprodutivos. Encontramos que, quando os sítios reprodutivos são escassos, há um ligeiro aumento na competição masculina, com maiores taxas de roubo de ninhos e interações agonísticas, e um ligeiro aumento no gradiente de seleção sobre características masculinas, com machos bem sucedidos tendendo a ser maiores do que machos mal sucedidos em monopolizar recursos. A disponibilidade de sítios reprodutivos não teve nenhum efeito consistente sobre a oportunidade para a seleção sexual e o risco de competição espermática. Entretanto, machos territoriais investiram mais em gônadas e perderam menos paternidade quando os sítios reprodutivos eram escassos. Não encontramos nenhum efeito da disponibilidade de sítios reprodutivos sobre a qualidade do cuidado paternal. Em conjunto, nossos resultados indicam que as previsões sobre como a disponibilidade de sítios reprodutivos influencia diversos aspectos de sistemas de acasalamento baseados na defesa de recursos têm fraco respaldo empírico. Tais previsões não consideram a plasticidade nas táticas de acasalamento de machos e fêmeas, tornando-as demasiadamente simplistas. Adicionalmente, desde a formulação original da teoria de sistemas de acasalamento, nossa compreensão sobre competição espermática aumentou. Sabemos atualmente que previsões acuradas sobre a intensidade e direção da seleção sexual devem levar em consideração processos pré- e pós-copulatórios. Finalmente, a interconexão entre seleção sexual e cuidado parental é complexa e a teoria original de sistemas de acasalamento não provê elementos suficientes para a construção de previsões claras e de amplo escopo taxonômico