Diss. (sammanfattning) Uppsala : Uppsala universitet, 2006. / Härtill 5 uppsatser.

Diss. (sammanfattning) Uppsala : Uppsala universitet, 2006. / Härtill 5 uppsatser.

Thesis (M.S.) -- University of Montana, 2008. / Title from author supplied metadata. Description based on contents viewed on July 6, 2009. Includes bibliographical references.

Thesis (M.Sc.)--Memorial University of Newfoundland, 2003. / Includes bibliographical references. Also available online.

Thesis (M.A.)--State University of New York at New Paltz, 2007. / Also issued in electronic version. Includes bibliographical references (leaves 64-74). Online version available via the SUNY New Paltz Sojourner Truth Library : http://hdl.handle.net/1951/42568

Elaborate sexual ornaments evolve because mate choice exerts strong sexual selection favouring individuals with high levels of ornament expression. Consequently, even at evolutionary equilibrium, life history theory predicts that ornamental traits should be under directional sexual selection that opposes contrasting selection to reduce the costs associated with their maintenance. Otherwise, the resources used to maintain ornaments should be used to improve other life history functions. Elaborate female ornaments have only evolved in a few species, despite females commonly experiencing strong sexual selection. One explanation for this rarity is that male preferences for female ornaments may be self-limiting: females with higher mating success become less attractive because of the lower paternity share they provide to mates with every additional sperm competitor. The unusual species in which female ornaments do occur can provide rare insight into how selection can favour the expression of expensive characters in females despite their costs. The main goal of my thesis was to determine how sexual selection acts on exaggerated sexual ornaments, and give new insight into how these ornaments may have evolved, in spite of the self-limiting nature of selection on male preferences. To determine the strength of sexual selection acting on female ornamentation in dance flies, we developed new microsatellite markers to assess polyandry rates by genotyping stored sperm in wild female dance flies. We first used polyandry rates to determine whether ornament expression was associated with higher mating success in female Rhamphomyia longicauda, a species that has evolved two distinct and exaggerated female ornaments. Contrary to our predictions, we found no evidence that females with larger ornaments enjoy higher mating success. We then compared polyandry rates in R. longicauda to those of two other species of dance fly, one (Empis aestiva) that has i independently evolved female ornaments on its legs, and another (E. tessellata) that does not possess any discernable female ornaments. We also estimated the opportunity for sexual selection, which we found to be similar and relatively low in all three species. Moreover, the standardized sexual selection gradients for ornaments were weak and non-significant in all three species. Females with more elaborate ornaments, in both within- and cross-species comparisons, therefore did not enjoy higher mating success. Overall, these results suggested that sexual selection operates rather differently in females compared to males, potentially explaining the general rarity of female ornaments. Our amplifications of stored sperm were able to reveal more than just mate numbers. We developed new methods to study patterns of sperm storage in wild female dance flies. We investigated how the skew in sperm genotypes from mixed sperm stores changed with varying levels of polyandry. Our data suggested that sperm stores were dominated by a single male in R. longicauda, and that the proportion of sperm contributed by this dominant male was largely independent of the number of rival males’ sperm present in the spermatheca. These results were consistent with the expectation of males using sperm ‘offence strategies’ in sperm competition and that the most successful male is likely to be the female’s last partner before oviposition. As a whole, my thesis contributed new molecular resources for an understudied and fascinating group of organisms. It exploited these new resources to provide the first estimates of lifetime mating success in several related species, and suggested that the general prediction that ornament expression should covary with sexual selection intensity does not seem to hold in this group. Instead, both the unusual prevalence of ii ornaments and the inconsistent evidence for sexual selection that sustains them in dance flies may owe their existence to the confluence of two important factors. First, the conditions under which sperm competition occurs: as last male precedence is likely, males are selected to prefer the most gravid females to secure a high fraction of her offspring’s paternity as they are unlikely to mate again before oviposition. Second, potent sexually antagonistic coevolution between hungry females and discerning males: females have evolved ornaments to disguise their stage of egg maturity to receive the benefits of nuptial gifts, while males face the challenge of distinguishing between gravidity and ornamentation in females.

A full understanding of any biological trait requires investigation of its evolutionary origin. Primates inspire great curiosity amongst researchers due to the remarkable diversity across species in both anatomical and behavioural traits, including sociality, sexual behaviour, life histories, neuro-anatomy, cognitive abilities and behavioural repertoires. The study of primates has involved comparative approaches since its inception, however, the necessary tools for statistically investigating the macro-evolutionary processes responsible for current diversity in biological traits have been developed only in the last 30 years or so, namely phylogenetic reconstruction and phylogenetic comparative methods. Amongst a multitude of evolutionary questions that can be addressed by phylogenetic comparative analyses, this thesis attempts to address two in particular, concerning primates. First, chapters 3 and 4 use meta-analysis and phylogenetic comparative analyses to investigate the evolution of large, brightly coloured ‘exaggerated sexual swellings' in female Catarrhine (‘Old World') primates. Together, chapters 3 and 4 show that such swellings are signals of temporal fertility, and present evidence to suggest that swellings co-evolved with conditions favouring male mate choice and cryptic female choice, therefore shedding light on the general conditions under which female signals of temporal fertility should evolve. Second, chapters 5 and 6 use phylogenetic comparative analyses investigate the evolution of enlarged brain size in the primate order. Together, chapters 5 and 6 suggest that multiple selection pressures have contributed to diversity in brain size and cognitive traits across primates, including sociality, intra-sexual competition and extended life history. Further, analyses presented in chapter 6 suggest that reliance on learned behaviour is a self-reinforcing evolutionary process, favouring ‘runaway' increases in cognitive abilities and reliance on culture in some primate lineages, which parallels increases in brain size, cognitive ability and reliance on culture in human evolution.

Orientador: Woodruff Whitman Benson / Tese (doutorado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-12T18:41:50Z (GMT). No. of bitstreams: 1 Peixoto_PauloEnriqueCardoso_D.pdf: 865304 bytes, checksum: 19540c783ae14daa3ba0b5f8dffc1d86 (MD5) Previous issue date: 2009 / Resumo: Brigas territoriais entre machos de insetos voadores são resolvidas das mais variadas formas. Em borboletas, existem evidencias de que tamanho, idade e motivação são importantes determinantes das chances de vitória. No entanto, as regras utilizadas nas brigas, o contexto biológico que afeta a intensidade dos comportamentos agonisticos e o papel funcional das características dos machos na geração de custos durante os confrontos ainda são pouco conhecidos. Neste trabalho investiguei uma serie de características que podem influenciar os custos e benefícios dos confrontos territoriais utilizando as espécies de satirineos Hermeuptychia hermes (Fabricius, 1775), Moneuptychia soter (Buttler, 1877) e Paryphthimoides phronius (Buttler, 1867) como modelos de estudo. No capitulo 1 investiguei se as disputas intra-especificas pela posse de territórios em machos de H. hermes e de M. soter ocorrem com contato físico e quais podem ser as características funcionalmente relevantes para a distinção entre machos territoriais e não territoriais. Para tanto, documentei as brigas entre machos utilizando filmagens de alta velocidade e realizei comparações morfológicas e fisiológicas entre machos possuidores de territórios (residentes) e machos que ocuparam territórios nos quais os residentes originais foram removidos (intrusos). Machos residentes de H. hermes apresentaram massa corporal e muscular similares as dos seus pares intrusos, porem, possuíram menor desgaste alar e maior quantidade de lipídeos. Machos residentes de M. soter, por outro lado, foram mais pesados que os machos intrusos que ocuparam seus territórios. Machos de H. hermes podem dividir a interação em vôos circulares, espirais ascendentes e perseguições lineares, enquanto machos de M. soter usam vôos circulares mais lentos, os quais são seguidos por perseguições lineares e emissões de som (estalos). Contatos físicos não foram registrados para nenhuma espécie. No capitulo 2 investiguei o papel da residência previa na resolução de conflitos territoriais entre machos de H. hermes. Ao induzir disputas entre dois machos que se comportaram como residentes (um residente original e um residente induzido) e compara-las com disputas naturais entre machos residentes e intrusos, foi possível demonstrar que a residência previa aumenta as chances de vitória em um combate. Massa corporal afetou positivamente as chances de vitória nas brigas induzidas, mas não nas brigas naturais. Finalmente no capitulo 3, investiguei a influencia da presença de recursos alimentares no estabelecimento de territórios de acasalamento por machos de P. phronius. Machos desta espécie defendem sítios de acasalamento localizados em manchas de sol sem recursos evidentes. No entanto, manchas de sol previamente desocupadas passaram a ser defendidas depois de receberem frutas fermentadas. Apesar de machos que se estabeleceram nestes locais aparentemente se alimentarem das frutas, experimentos de escolha de territórios sugerem que eles preferem defender sítios sem recursos e utilizam a defesa de áreas com alimento como tática alternativa de acasalamento. Este estudo representa a primeira investigação deste tipo de variação nas táticas de acasalamento em borboletas. Alem de auxiliar a compreensão da evolução de determinados sistemas de localização de parceiros, esse sistema permite testar qual a implicação da existência de mais de uma tática reprodutiva na intensidade dos confrontos territoriais entre machos / Abstract: Territorial conflicts in flying insects may be resolved through many different ways. In male butterflies, size, age and motivation are often reported as important determinants of the winning chances. However, the rules used when fighting, the biological context that affects the intensity of agonistic behaviors, and the functional role of male traits in the contests costs generation are poorly known. In this study I investigated a series of characteristics that may influence the costs and benefits of territorial conflicts using the satyrine species Hermeuptychia hermes (Fabricius, 1775), Moneuptychia soter (Buttler, 1877), and Paryphthimoides phronius (Buttler, 1867) as study organisms. In chapter 1, I used high-speed video imagery to investigate whether intraspecific disputes between male H. hermes and male M. soter occur with physical contact. Additionally, I removed resident males from their defended sites and the subsequent intruders that established on those areas after the resident removal to assess if wing wear, body mass, fat content and flight muscle ratio are important determinants of male residency status. Resident males of H. hermes were similar in body weight and thoracic muscle mass to their intruder rivals, but had fewer wing wear and higher fat content. Resident males of M. soter, on the other hand, were heavier than intruder males. Male H. hermes divided their interaction in up to three phases consisting of circular flights, ascending spirals and back and forth persecutions, whereas male M. soter used much slower circular flights that were often followed by linear persecutions and clicking sound emissions. Physical contacts were not observed for both species. In chapter 2, I investigated the role of previous residence on contest resolution in the butterfly H. hermes. By inducing territorial interactions between males that behaved as residents (one original resident and another resident-induced one) and comparing them with natural contests between resident and intruder males, it was possible to show that previous residence increases the chances of victory. Body mass positively affected the winning chances in the induced fights, but was unimportant among the natural ones. Finally in chapter 3, I investigated the role of feeding resources in the territory establishment by males of the butterfly P. phronius. Male of this species typically defend sunny clearings on the forest edge which do not contain any evident resource. However, previously undefended sunny clearings were occupied by territorial males after receiving soft fermenting fruit. Although males which established on sites containing feeding resources seem to forage on the fermenting fruit, territory selection experiments shows that males prefer to defend sites without resources. This type of variation in the mate locating tactics has never been previously investigated for butterflies. In addition of helping the understanding of the evolutionary processes leading to different mate-locating strategies, this system allows the testing of the influence of different reproductive behaviors on the intensity of territorial conflicts between males / Doutorado / Doutor em Ecologia

Sex allocation and sexual selection have been heavily studied, but rarely linked. In this thesis I investigated the interface between them in the gregarious parasitoid wasp Nasonia vitripennis, both directly and through their interactions with the mating system and sexual conflict. Chapter 2 investigated sexual selection and mating at the natal site: earlier eclosing males mated more females independently of body size. Nasonia follows Local Mate Competition, which describes how a female laying eggs alone on a patch of resources (a so-called single-foundress) should lay an extremely female-biased brood to minimise competition between her sons, yet ensure all her daughters are fertilised. Based on this I predicted that males with with fewer brothers would be better inseminators. Despite finding significant among-strain variation in (1) single-foundress sex ratio, (2) mate competitiveness when alone and (3) when in competition, (4) sperm resources, but not (5) sperm-depletion (Chapters 3 & 4), I did not find the predicted relationship. Conversely males from strains with more brothers had a higher mating success under competition (Chapter 3) leading to the question: does mating success select on sex ratio or vice versa? Either way it is a result of an interaction between sexual selection and sex allocation. Chapter 5 investigated the role of male post-copulatory courtship on female re-mating, and found that among- strain variation in female re-mating was not associated with variation in the duration of the post-copulatory courtship. Chapter 6 reviewed sexual conflict in the Hymenoptera: their haplodiploid genetics, newly sequenced genomes and varied life- histories provides a base for future research to build on. Finally I highlight the novel links between sexual selection, sex allocation, sexual conflict and the mating system found during my studies that will hopefully prompt future research on this topic.

[Truncated abstract] Despite the costs associated with mating, females of many taxa solicit multiple mates during a single reproductive event (polyandry). Polyandry is clearly adaptive when females gain direct benefits from males at mating. However, polyandry has also been shown to increase female fitness in the absence of direct benefits. Thus, a number of genetic benefit hypotheses have been developed to account for the origin of this behaviour. Although not mutually exclusive, a distinction lays between genetic benefits that propose defense against reproductive failure (nonadditive genetic effects), and those that propose benefits from intrinsic sire effects (additive genetic effects). Nonadditive genetic benefits of polyandry have been documented in a number of species; by soliciting multiple mates females can avoid inbreeding and other forms of incompatibility between parental genotypes. Polyandry may also increase female reproductive success when genetically superior males have greater success in sperm competition, and produce better quality offspring. An inevitable consequence of polyandry is that sperm from rival males will overlap in the female reproductive tract and compete to fertilise the ova. The outcome of sperm competition is typically determined by bias in sperm use by the females, interactions between parental genotypes, and ejaculate characteristics that provide a fertilisation advantage. Thus, sperm competition is recognised as a persuasive force in the evolution of male reproductive traits. Comparative analyses across species, and competitive mating trials within species have suggested that sperm competition can influence the evolution of testis size and sperm production, and both sperm form and sperm function. ... After six generations of selection I observed phenotypic divergence in litter size - litter size increased in the polyandrous lines but not in the monandrous lines. This result was not attributable to inbreeding depression, or environmental/maternal effects associated with mating regime. Genetic benefits associated with polyandry could account for this result if increased litter size were attributable to increased embryo survival. However, males from the polyandrous lineages were subject to sperm competition, and evolved ejaculates with more sperm, suggesting that evolutionary increases in litter size may in part be due to improved male fertility. Finally, Chapter Five is an investigation of the natural variation in levels of polyandry in the wild, and the potential for sperm competition to drive macroevolutionary changes in male reproductive traits among geographically isolated island populations of house mice. I sampled seven island populations of house mice along the coast of Western Australia and, by genotyping pregnant females and their offspring, determined the frequency of multiply sired litters within each population. I applied the frequency of multiple paternity as an index of the risk of sperm competition, and looked for selective responses in testis size and ejaculate traits. I found that the risk of sperm competition predicted testis size across the seven island populations. However, variation in sperm traits was not explained by the risk of sperm competition. I discuss these results in relation to sperm competition theory, and extrinsic factors that influence ejaculate quality.