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.

Monitor lizards (Varanidae) are morphologically very uniform in body shape, but much diversified in body size along both phylogenetic and ontogenetic axes. A striking sexual size dimorphism exists in monitor lizards; they are capable of fast growth, metabolism and sexual maturation. I collected the data concerning body size of particular species and verified the validity of Rench's rule, which said that there is bigger difference in body size of a conspecific male and females growing with larger body size of the species. Males are markedly bigger than females. In the next step, I focused on the model species of monitor lizards, Varanus indicus. I monitored its ontogeny very carefully. I found that this monitor lizard has pronounced sexual size dimorphism, but there are only small differences in body shape. It is capable of rapid growth and sexual maturation. The sexual dimorphism in body shape is only poor, but still measurable right in those places where the selection pressures were expected. Blood sampling monitored biochemical and haematological parameters. The concentrations of the biochemical parameters revealed the economy of resources of particular sexes partially, the costs of body growth and reproduction. Although both sexes produce the same amount of biomass (the body growth of males vs. the...

The Fence Lizard (Sceloporus undulatus complex) is a wide ranging North American species complex occurring from the eastern seaboard westward through the great plains and central Rocky Mountains and into the American Southwest. A recent phylogeny suggests four species lineages occur within S. undulatus. Traits within an interbreeding species that are influenced by sexual selection are under different selection pressures and may evolve independently from the selective forces of habitat. Sceloporus lizards have several characters that are influenced by sexual selection. I investigated sexual size dimorphism and allometric relationships of body size (snout vent length), torso length, rear leg length and three measurements of head size in 12 populations from the four species in the S. undulatus complex (N=352) specifically looking for variation among the 4 species. Additionally I investigated the size of signal patches between males and females in three species (N=339 specimens of S. consobrinus, S. cowlesi, S. tristichus) of the S. undulatus complex. Sexual confusion, was recently described in a population of the Sceloporus undulatus complex occurring in White Sands, New Mexico and the behavior is correlated with variation in badge size between male and female lizards. To make inferences about sexual confusion at the species level I investigated the presence and absence of signal patches in female lizards, and compare the sizes of signal patches between males and females. My analyses suggest that torso length and head size are significant sources of sexual size dimorphism but the findings differ from earlier published investigations of sexually dimorphic characters in the species complex. I also find support for the S. undulatus complex being generally a female larger species complex. However two of the 12 populations I investigated displayed male biased sexual size dimorphism. Analysis of signal patches across three species of the S. undulatus complex suggests that sexual dimorphism in signal patch size for S. cowlesi and S. tristichus may not prevent sexual confusion. While the near total absence of signal patches in female S. consobrinus is evidence that sexual confusion is not possible with regards to signal patches.

O dimorfismo sexual nas espécies pode surgir a partir da seleção decorrente dos diferentes sistemas reprodutivos. Estudos comportamentais de espécies neotropicais são raros e pouco se sabe sobe as espécies brasileiras. Neste estudo, foram descritos o comportamento de três espécies neotropicais que ocorrem no Cerrado brasileiro: Acanthagrion truncatum, Argia reclusa (Coenagrionidae) e Heaterina rosea (Calopterygidae). Também foi evidenciado o dimorfismo sexual nestas espécies e investigou-se a partir de observações comportamentais, como o dimorfismo se desenvolve em espécies com diferentes táticas reprodutivas. Com os resultados obtidos, vemos que em espécies territoriais os machos são maiores do que as fêmeas, enquanto em espécies não-territoriais as fêmeas são maiores do que os machos. Sugere-se que, diferentemente de outros estudos, em Zygoptera o tipo de sistema reprodutivo pode determinar o dimorfismo sexual. / Sexual size dimorphism (SSD) can in some species result from the selection acting through different mating systems. Behavioral studies of neotropical species are rare, and few is known about the brazilian species. In this study, we described the behavior of three neotropical species that occur in the brazilian neotropical savannah: Acanthagrion truncatum, Argia reclusa (Coenagrionidae) and Heaterina rosea (Calopterygidae). We show the SSD in these species and investigates through behavioral observations, how SSD develops in species with different mating tactics. With our results, we can see that in territorial species the males are larger than females, while in non-territorial species the females are larger than males. We suggest that, unlike other studies, in Zygoptera the kind of mating system adopted by males may determinate the SSD in a species.

O dimorfismo sexual nas espécies pode surgir a partir da seleção decorrente dos diferentes sistemas reprodutivos. Estudos comportamentais de espécies neotropicais são raros e pouco se sabe sobe as espécies brasileiras. Neste estudo, foram descritos o comportamento de três espécies neotropicais que ocorrem no Cerrado brasileiro: Acanthagrion truncatum, Argia reclusa (Coenagrionidae) e Heaterina rosea (Calopterygidae). Também foi evidenciado o dimorfismo sexual nestas espécies e investigou-se a partir de observações comportamentais, como o dimorfismo se desenvolve em espécies com diferentes táticas reprodutivas. Com os resultados obtidos, vemos que em espécies territoriais os machos são maiores do que as fêmeas, enquanto em espécies não-territoriais as fêmeas são maiores do que os machos. Sugere-se que, diferentemente de outros estudos, em Zygoptera o tipo de sistema reprodutivo pode determinar o dimorfismo sexual. / Sexual size dimorphism (SSD) can in some species result from the selection acting through different mating systems. Behavioral studies of neotropical species are rare, and few is known about the brazilian species. In this study, we described the behavior of three neotropical species that occur in the brazilian neotropical savannah: Acanthagrion truncatum, Argia reclusa (Coenagrionidae) and Heaterina rosea (Calopterygidae). We show the SSD in these species and investigates through behavioral observations, how SSD develops in species with different mating tactics. With our results, we can see that in territorial species the males are larger than females, while in non-territorial species the females are larger than males. We suggest that, unlike other studies, in Zygoptera the kind of mating system adopted by males may determinate the SSD in a species.

Sexual selection is one of main selective pressure affecting body size, and subsequently leads to the evolution of sexual size dimorphism (SSD). The eyelid geckoes, family Eublepharidae, are a monophyletic group with considerable variability in SSD, including both male-larger and female-larger species. In general, it was supposed that eyelid geckos are highly variable in presence of male combats and in complexity of male pre-copulatory behaviour, and that this variability in this conspicuous male behaviour may lead to differences in SSD. The aim of this study was to reveal relationships between the direction of SSD and presence/absence of tail vibration during precopulatory phase and male combat behaviour. Using behavioural testing, it was revealed that male combats are present in all tested species, even in species, where the absence of such behaviour was supposed so far. In several species, the strong effect of seasonality to male aggression was observed, which may play a role in the evolution of SSD. The evolutionary changes in the presence/absence of tail vibration during precopulatory phase were independent on changes in the direction of SSD, the presence of tail vibration seems to be ancestral state for these lizards. During the evolution of this group, the tail vibrations disappeared four...

6 Abstract Due to its effect on fitness and many biological processes is body size one of the most important attribute of organisms. Body size is positively correlated with fecundity in insects and other ectotherms. Growth, which determine body size, is therefore crucial feature of animals. Study of growth can elucidate some aspects of body size evolution. Unfortunatelly, many insects life-history studies do not consider its complexity, especially the existence of distinct larval instars. Inaccurate record of growth trajectory may result also in biased differences in growth between sexually dimorphic sexes. Aim of this thesis is to record growth trajectories of two flower beetle species (Coleoptera: Scarabaeidae: Cetoniinae). These growth trajectories enable to define proximate mechanisms of growth with regard to individual instars. Determination of developmental mechanisms of sexual size dimorphism is another goal of this study. The growth is clearly divided in three distinct periods. In each individual period (i.e. instar) is described by asymptotic curve. The instars are not independent on each other - the growth in following instar is influenced by growth in previous. There are no differences in growth characteristics between sexes. Sexual size dimorphism is caused by differences in growth rate between...

This project describes various aspects of the breeding ecology and behaviour of the brown falcon Falco berigora, a common but poorly study Australian raptor. In particular it examines (a) the main influences on reproductive success; (b) tests predictions of theories proposed to explain the evolution and maintenance of sexual size dimorphism (RSD; females the larger sex) in raptors; and (c) investigates sex allocation patterns in the light of current sex ratio and parental investment theory. The study was conducted between July 1999 and June 2002 approximately 35 km southwest of Melbourne, at the Western Treatment Plant (WTP), Werribee (38°0’S 144°34’E) and surrounds, a total area of approximately 150 km2.¶ · In all plumage and bare part colouration of 160 free-flying falcons was described. The majority of variation in these characters could be attributed to distinct age and/or sex differences as opposed to previously described colour ‘morphs’.¶ · Nestling chronology and development is described and formulae based on wing length derived for determining nestling age. An accurate field-based test for determining nestling sex at banding age is also presented.¶ · Strong sex role differentiation was apparent during breeding; typical of falcons females performed most parental duties whilst males predominantly hunted for their brood and partner. Based on observations of marked individuals, both sexes of brown falcons aggressively defended mutual territories throughout the year, with just 10% of each sex changing territories during the entire study period. Males performed territorial displays more frequently than females, the latter rarely displaying alone.¶ · The diet of the population as a whole was very broad, but within pairs both sexes predominantly specialised on either lagomorphs, small ground prey (e.g. house mice Mus musculus), small birds, large birds or reptiles, according to availability.¶ · Reproductive parameters such as clutch size and the duration of parental care were constant across all years, however marked annual differences in brood size and the proportion of pairs breeding were evident.¶ · Age was an important influence upon reproductive success and survival, with immature birds inferior to adults in both areas. However, interannual differences were by far the most influential factor on breeding success and female survival. Heavy rain downpours were implicated as the main determinant of reproductive success and adult female mortality in a population largely devoid of predation or human interference.¶ · Female-female competition for territorial vacancies was intense; larger adult females were more likely to be recruited and once breeding fledged more offspring. In contrast, male recruitment and breeding success was unrelated to either body size or condition indices, although smaller immature males were more likely to survive to the next breeding season. This directional selection is consistent only with the predictions of the intrasexual competition hypothesis.¶ · Despite marked RSD (males c. 75% of female body mass), throughout the nestling phase female nestlings did not require greater quantities of food than their male siblings. However, female parents fed their last-hatched sons but not daughters, resulting in the complete mortality of all last-hatched female offspring in focal nests. Given last-hatched nestlings suffered markedly reduced growth rates and female, but not male, body size is important in determining recruitment patterns, the biased allocation amongst last-hatched offspring is likely to reflect differing benefits associated with investing in small members of each sex, consistent with broad-scale Trivers-Willard effects. Recruitment patterns support this, with surviving last-hatched females, in contrast to males, unable to gain recruitment into the breeding population upon their return to the study site.¶ Thus selection appears to act at the nestling, immature and adult stages to maintain RSD in the focal population. Larger females were favoured in the nestling phase, at recruitment and once breeding had greater reproductive success. In contrast, selection favoured a reduction or maintenance of immature male size as smaller birds had a greater chance of survival in the year following recruitment than their larger counterparts; thereafter male size was unimportant. Together, this directional selection favouring increased female competitive ability is consistent only with the predictions of the intrasexual competition hypothesis, which appears the most probable in explaining the maintenance and perhaps evolution of RSD in raptors.

<p>Males and females differ in morphology and behaviour, so that selection acts differently on the two sexes. This changes the relative reproductive success of males and females, and it is beneficial for parents to bias the sex ratio of their broods in favour of the sex with the best survival and breeding prospects. Differences between the sexes and brood sex ratio in Tengmalm’s owl (Aegolius funereus) in northern Sweden were investigated, using a molecular sexing technique based on PCRamplification of sex-linked CHD1 genes. Among owls caught during autumn migration, females were commoner than males, especially within juveniles. However, in contrast to earlier studies, it was shown that adult males sometimes undertake migratory movements indicatory of nomadism. Measurements of these owls revealed that sexual size dimorphism in Tengmalm’s owl is not as great as previously reported from studies carried out during the breeding season. Females were slightly larger (4% by mass) than males, probably owing to the different roles of males and females during breeding, when this dimorphism is greater. The size difference between male and female nestlings was found to be similar to that for adults in autumn, and to investigate whether this led to differential mortality, the effect of supplementary feeding on mortality of male and female nestlings was studied. Supplementary feeding reduced male mortality when vole abundance was low, and it was concluded that larger female nestlings out-competed their smaller brothers, who then suffered increased mortality when food was scarce. Recruitment of male nestlings into the breeding population declined with decreasing food supply at the time of fledging, a pattern not observed in females. Juvenile males were therefore more vulnerable to food shortage than females, both in the nest and after fledging. Mean brood sex ratio varied significantly among years characterized by different phases of the vole cycle and associated vole abundance. Broods were male-biased (63% males) in a year when the food supply was favourable during spring and summer, neutral (50%) in a year with an intermediate food supply, and female-biased (35% males) in a year when food was in short supply. Parents appeared to adaptively adjust the sex ratio of their broods according to the relative mortality risk and reproductive potential of sons and daughters.</p>

Males and females differ in morphology and behaviour, so that selection acts differently on the two sexes. This changes the relative reproductive success of males and females, and it is beneficial for parents to bias the sex ratio of their broods in favour of the sex with the best survival and breeding prospects. Differences between the sexes and brood sex ratio in Tengmalm’s owl (Aegolius funereus) in northern Sweden were investigated, using a molecular sexing technique based on PCRamplification of sex-linked CHD1 genes. Among owls caught during autumn migration, females were commoner than males, especially within juveniles. However, in contrast to earlier studies, it was shown that adult males sometimes undertake migratory movements indicatory of nomadism. Measurements of these owls revealed that sexual size dimorphism in Tengmalm’s owl is not as great as previously reported from studies carried out during the breeding season. Females were slightly larger (4% by mass) than males, probably owing to the different roles of males and females during breeding, when this dimorphism is greater. The size difference between male and female nestlings was found to be similar to that for adults in autumn, and to investigate whether this led to differential mortality, the effect of supplementary feeding on mortality of male and female nestlings was studied. Supplementary feeding reduced male mortality when vole abundance was low, and it was concluded that larger female nestlings out-competed their smaller brothers, who then suffered increased mortality when food was scarce. Recruitment of male nestlings into the breeding population declined with decreasing food supply at the time of fledging, a pattern not observed in females. Juvenile males were therefore more vulnerable to food shortage than females, both in the nest and after fledging. Mean brood sex ratio varied significantly among years characterized by different phases of the vole cycle and associated vole abundance. Broods were male-biased (63% males) in a year when the food supply was favourable during spring and summer, neutral (50%) in a year with an intermediate food supply, and female-biased (35% males) in a year when food was in short supply. Parents appeared to adaptively adjust the sex ratio of their broods according to the relative mortality risk and reproductive potential of sons and daughters.