Long-tailed Manakins Chiroxiphia linearis are frugivorous birds with a lek mating system and male-male cooperation in courtship display. I studied male-male networks and correlates of male mating success in a color-banded population in Monteverde, Costa Rica, from 1981 to 1986. Males were organized in teams at scattered perch-zones (75 to 300 m apart) that were usually in aural but not visual contact. Each team consisted of 3 to 15 males (x=7.1±3.4), in an apparent linear dominance hierarchy, with an alpha and beta male who did most of the courtship display. In a study population with 50 to 60 active males per season, only 6 to 8 males were alphas. Only betas inherited alpha status (n=3). Males appear to be 8 or more years of age before attaining beta status. Alpha tenure can last 4 years. Alpha males were rarely or never seen in zones other than their 'home' zone. Lower-ranking males maintained simultaneous affiliations with males at as many as 6 different zones. Each zone, therefore, was a sort of hub at which males with different affiliations around the rim came into contact. Each of the 6 major perch-zones shared at least one constituent with each of the other zones. The mean number of males shared by zones was 3.9 ± 2.7 (range=1 to 9). Marked changes occurred in male traits with increasing age and status. These included (1) Significant declines in weight throughout the lifespan, (2) a 4-year delay in plumage maturation with well-defined stages, (3) reduction in the number of zones with which males maintained affiliations, and (4) increasing probability of copulatory success (restricted to a small subset of the oldest males, ≥ 10 years of age). Variance in copulatory success was the highest yet described for birds. Of 85 males monitored between 1983 and 1986, copulations (n=121) were distributed among only 8 males. Four of these males accounted for over 90% of the copulations, with 63% accruing to one male. The beta male of this alpha copulated twice in the absence of his partner; all the other copulaters were alphas. I examined correlates of male mating success. Female visitation correlated with the number of unison 'toledo' calls given. If a female visited, copulatory success correlated both with a residual effect of the 'toledo' output and with the duration of the 'butterfly' component of the dual-male dance performance. My correlational results suggest that females do choose, on the basis of performance cues, among the small subset of males that are well-established alpha and beta partners. Development of alliances, as much as male combat, may determine attainment of high-performance partner status. Thus, sequential male-male interactions and female choice appear to produce nested subsets of successful males leading to an extreme in variance in male mating success. Males unsuccessful in male-male interactions are not 'eligible' for female choice. By requiring partnered display, females may be implicitly narrowing the subset of potentially successful males. In other lek systems the union, rather than the intersection, of the subsets produced by intra- and intersexual selection may include successful males. In that case, intrasexual selection via disruption of copulations may enlarge the pool of potentially successful males under intersexual selection and produce lower variances in male mating success. Students of sexual selection may need to consider the extent to which intra- and intersexual selection interact as union or intersecting sets to produce variance in male mating success.

Mate choice plays an important role in sexual selection and speciation. The evolution of mate choice is intriguing in cases where choosy individuals gain little except for genetic material from the mate and where the trait used as a criterion for the choice is costly to its bearer. The sensory exploitation hypothesis is an interesting idea that applies to such cases because it suggests that sexual preferences may arise as side-effects of preferences that are under selection in other contexts. The role of mate choice in speciation is strong but is debated because the reasons for population divergence in mate preferences and sexual traits are sometimes hard to explain. Also in this context sensory exploitation offers a potential explanation in that a link between natural and sexual selection may result in divergence in sexual selection whenever populations differ in natural selection. In this thesis, I test several aspects of this hypothesis in a species of fish, the swordtail characin (Corynopoma riisei). In this species males display a flag-like ornament that grows from the operculum. Because females respond to this ornament by biting at it, it has been proposed to act as a food-mimic. By manipulating female food type and quantity, and testing the resulting female preference for the male ornament, I find support for the theory that the preference has evolved through sensory exploitation and that females indeed appear to relate the ornament to a food item. Furthermore, I show that sensory exploitation can lead to morphological divergence among natural populations in this species. Apart from the flag-ornament, other courtship signals are also investigated. The results show that the relative importance of different signals may vary depending on receiver motivation. This suggests that various aspects of both male courtship signals and the conditions during which they are being signalled should be considered to gain a full understanding of mate choice and its role in sexual selection and speciation.

The robust behavioural courtship ritual displayed by Drosophila melanogaster males is governed by their underlying nervous system (NS). Two key genes of the Sex Determination Hierarchy, fruitless (fru) and doublesex (dsx), determine most neuronal substrates for sexual behaviour. In this study we aim to better understand the role fru plays in determining these neural substrates, as a means of better understanding the relationships between brain, behaviour and genes, and thus how the development of neuronal networks shape innate and species-specific behaviours. Fru has two major functions: control of male sexual behaviour, and viability in both sexes. Alternative splicing of fru produces transcription factors driven by four promoters: P1 transcripts are sex-specifically spliced (only viable in the male), and P2-4 transcripts are crucial to both sexes survival. The resulting proteins contain a BTB protein-protein interaction domain at the N-terminus, and one of four C-terminal zinc-finger (ZnF) DNA binding domains. Male-specific proteins (FruM) contain an additional 101 amino acid N-terminal domain, and one of three alternative C2H2 ZnF domains (FruMA, FruMB, FruMC). These male-specific isoforms are expressed in the central nervous system (CNS) beginning in the late stage larvae (L3), peaking during pupation and on into adulthood. Little is known, however, about the roles of the individual isoforms, and no clear transcriptional targets have been identified. The central aims of this thesis are to document the wild-type expression patterns of Fru isoforms throughout development in the CNS, create and characterise isoform-specific mutants, and to identify and evaluate putative transcriptional targets of Fru. Combining these findings will lead to a better understanding of the underlying molecular functions of individual FruM isoforms, as a means to understanding their roles in sexual behaviour. Expression analysis of FruM isoforms throughout development in the NS is described. To further characterise the role of individual FruM isoforms, isoform-specific mutants in fruA and fruB exons were generated using site-specific homologous recombination (HR). These novel mutants were validated by PCR and Fru isoform-specific antibody stainings. Mutants were analysed in fruM- and fru-null genetic backgrounds, to distinguish the roles of sex-specific vs. common isoforms. These analyses included: fertility, viability and morphology. FruA was found to have a role in wing extension with possible repercussions for song production. FruB was found to be developmentally lethal, in addition to having defects in male courtship behaviour. To understand the role of fru in the NS, downstream transcriptional targets of FruM isoforms were identified. DNA adenine methyltransferase identification (DamID) was used to identify putative transcription targets of FruM isoforms. The Dam protein methylates DNA in Drosophila in a sequence-specific manner allowing targets of Fru to be isolated. Candidate genes were identified using computational analysis (including gene ontology, peak analysis and motif analysis) along with a biologically significant connection with fru. The relationship between fru and six candidate genes were characterised using RNAi. The results of these studies advance our knowledge of how FruM isoforms influence the development and physiology of the NS underlying male sexual behaviour in Drosophila.

This thesis focuses on the motifs of courtship and marriage as experienced by the female characters in the novels published from the end of the 18th century till the second half of the 19th century. These novels include Evelina (1778), Pride and Prejudice (1813), Jane Eyre (1847) and The Mill on the Floss (1860). As all these novels are written by female authors, the thesis provides a unique female perspective of the issue. The chosen texts deal with the motifs of courtship and marriage, play with the conventional discourse, and at the same time challenge the established perception of the role of a woman in the process of courting. Besides discussing the novels, I explore also the way the themes of courtship and marriage are presented in the original 18th and 19th century literature. For this purpose, I scrutinized various conduct books and essays of the period. The first chapter of the thesis is introductory and explains the main ideas and terms used in the thesis. The second chapter focuses on the social background the heroine comes from - her family, character and education. Hence, the role of the family is discussed in reference to the process of courtship. Great attention is given primarily to the role of a father as a possible social guardian and the model of a future husband. The third...

Parasitic wasps are highly diverse and play a major role in suppression of herbivorous pest populations, but relatively little is known of the mechanisms driving their diversity. Molecular studies indicate that cryptic species complexes resulting from adaptations to specific hosts or host-foodplants may be common. The gregarious endoparasitoid, Cotesia congregata (Braconidae), is a model system for understanding parasitic wasp biology. It is reported to attack at least 15 species of sphingid caterpillars, most of which are plant family specialists. Molecular studies have demonstrated genetic differentiation of two host-foodplant complex sources originating from Manduca sexta on tobacco (MsT) and Ceratomia catalpae on catalpa (CcC). Response to female pheromone and elements of their courtship songs differ. Wasps from both sources mated and produced F1 hybrid offspring in the laboratory; however, 90% of hybrid females resulting from one of the reciprocal crosses failed to produce offspring. I built on this previous work by evaluating an ecological barrier, the evolution of courtship songs within the genus, and patterns of hybrid sterility among four additional host-foodplant complexes, as well as differentiation of their symbiotic bracovirus. Tests of developmental tolerance to nicotine demonstrate that MsT wasps are highly adapted to hosts feeding on tobacco, whereas CcC wasps experience high mortality. Acoustic analysis of courtship songs among host-foodplant sources of C. congregata and eleven additional species of Cotesia demonstrates that songs are species specific and appear to be correlated with genetic relatedness. Cotesia congregata from all sources mated and produced F1 hybrid offspring in the laboratory; however, hybrid females resulting from specific reciprocal crosses failed to produce progeny. Dissections of hybrid females revealed that sterile wasps lacked mature ovaries and functional bracovirus, a symbiotic virus integrated into the wasp genome and necessary to suppress the host immune system. Relative in vivo expression of wasp bracovirus genes differs between MsT and CcC host-foodplant complexes. Cumulatively, these behavioral, ecological, and genetic barriers to reproduction indicate that C. congregata is diverged into two incipient species with limited gene flow, and provides insight into the role of varied reproductive barriers in speciation of parasitic wasps.

Spotted seatrout, Cynoscion nebulosus, spawning locations as well as associated environmental variables were determined for Tampa Bay, Florida during the 2004 spawning season using a mobile hydrophone survey. Hydrophones, a type of underwater microphone, can be used to detect and record spawning sounds of soniferous fishes. During their spawning season in Tampa Bay which generally occurs between March and September, mature male spotted seatrout generate sounds associated with courtship in the crepuscular and evening periods by vibrating sonic muscles against the swim bladder. Active spawning sites can be located using hydrophones to find these calling males. Using a random stratified sampling method, 760 stations within Tampa Bay (46 % of the sampling universe) were sampled over the 2004 spawning season. Only 8% of sampled stations had large aggregations of spotted seatrout. Spawning, determined by the sound produced by large aggregations, was detected throughout the bay except for Hillsborough Bay and was most common in the lower bay and eastern region of the middle bay. Presence of submerged aquatic vegetation (SAV), proximity to shoreline, as well as high dissolved oxygen values and shallow depth were positively correlated with spawning areas. Courtship calls of sand seatrout, Cynoscion arenarius, and silver perch, Bairdiella chrysoura were also detected during the survey as they share an overlapping spawning season with spotted seatrout. Aggregations of all three species rarely occurred simultaneously. Sand seatrout and silver perch used different habitats within Tampa Bay to spawn and spawned with a much greater frequency than spotted seatrout. Courtship calls of spotted seatrout were analyzed both by ear and by received sound level to determine if signal processing could be used to assess courtship sound recordings. However, there was no clear relationship between the two methods.

Sixteenth century Elizabeth I of England has long been a figure of interest to Renaissance scholars, and their work largely focuses on how her gender impacted the power, politics, and culture of her day. Many have perceived her to be a heroine whose ingenuity and determination circumvented the limitations imposed on a female ruler in patriarchal Renaissance England. In my thesis, I examine the life and work of Elizabeth I, and the self-representations she constructed within the boundaries imposed on highborn women. In the first half of my thesis, I suggest that she embraced and utilized the female roles available to her to secure agency and a degree of safety for both herself and England. In the second half, I suggest that masculine subjects such as Sir Philip Sidney and Edmund Spenser, in turn, sought to manipulate her later self-representations to negotiate their own agency and identity which was perceived to be beset with anxieties and biases stemming from the ageing Queen's seizure and redefinition of the female gender role allotted to her. A chronological examination of the self representations evident in her personal writing, commissioned portraiture, parliamentary speeches, and sonnets, as well as the poetry of two of her foremost masculine subjects, suggests a shift in gender politics and a tension roused by an ageing Queen regnant in a rigidly patriarchal society. / Graduation date: 2012

Animals experience variation in their environment because of natural changes. However, due to anthropogenic disturbance, the speed and severity of these changes have recently increased. This thesis investigates how reproductive behaviours may be affected by human induced environmental change. In specific, I investigate how visual and chemical changes in the aquatic environment, caused by eutrophication, affect mating systems and sexual selection in fish. Broad-nosed- and straight-nosed pipefish, which both have been studied in detail for a long period, were used as model organisms. These two species are particularly suitable model organisms since they perform complex courtship behaviours, including the advertisement of ornaments and a nuptial dance. Further, two distinct populations were studied, one on the Swedish west coast and one in the Baltic Sea, as these two locations vary in the degree and extent of environmental disturbance, in particular turbidity. I found that changes in the visual environment had no impact on the development of female sexual ornaments in these sex-role reversed pipefishes, but it hampered adaptive mate choice. Turbidity also had a negative effect on reproductive success in the Baltic Sea population. Changes in the chemical environment in the form of increased pH reduced the probability to mate, while hypoxia did not alter mating propensity. However, hypoxic water delayed the onset of both courting and mating. Hence, human induced change in aquatic environments may alter the processes of sexual selection and population dynamics.

Recent advances in courtship theory emphasize day-today interaction and the environment in which interaction takes place as critically important in the understanding of relationship development. The purpose of this study was to determine the influence of college course assignments on time spent in relationship activities and, similarly, the influence of time spent in relationship activities on relationship satisfaction. As college couples are often the subjects of relationship studies, consideration of the college environment seemed both appropriate and overdue. The sample consisted of 35 serious dating couples in which both partners were full-time students. A telephone survey methodology was developed so that couples could report coded relationship behaviors both conveniently and confidentially. Data were collected twice a week for 8 1/2 weeks during winter term 1987 at a large northwestern university. Results indicated: (1) previous, current, and up-coming course assignments were influential regarding reported time in selected relationship activities, and, in general, tended to increase time in activities; (2) when assignments decreased relationship activity, men's assignments were more influential. Women's assignments, particularly previous assignments, were found likely to increase relationship activity. Regarding the influence time spent in activities had on relationships satisfaction, the data indicated that time spent eating together and in affectionate behavior were activities that increased relationship satisfaction. Discussion centered on the "interpersonal process" framework of relationships development and on the timing of course workload on relationships. Conclusions suggested the academic environment does have an effect on dating relationships and that this effect may be similar to work and family issues that society as a whole is facing. / Graduation date: 1990

Behavior is a complex process resulting from the integration of genetic and environmental information. Thus, the genetically tractable Drosophila melanogaster was utilized to better understand the interplay between these factors since Drosophila males and females exhibit sex-specific courtship behaviors that are innate yet modifiable. These sex-specific behaviors, as well as sexually dimorphic development, are regulated, in part, by the somatic sex-determination hierarchy. Since reproductive behaviors rely on the rapid integration of multiple sensory cues, it is likely that the perception and integration of such cues and mating-induced physiological changes are mediated in part by changes in gene expression. Therefore, it was hypothesized that assaying gene expression changes in response to courtship or mating in Drosophila males would uncover new targets of the sex-determination hierarchy and other behaviorally important loci. We took a novel approach to find these behaviorally-responsive loci by utilizing microarray technology to assess courtship- or mating-induced gene expression changes in Drosophila male whole bodies or heads. Mutations in candidate loci were tested for effects on reproductive behaviors and present the first data showing that egghead (egh) and female-specific independent of transformer (fit) affect male reproductive behavior. egh is up regulated in male heads 20 min after courting and is required post-developmentally in a subset of neurons for robust male courtship behavior. fit, a fat body-expressed sex-determination hierarchy target gene, is up regulated in male whole bodies after 5 min of courtship. fit is also up regulated in male heads after 20 min of courtship or 2 hrs after mating. Mutations in fit result in male-male courtship; more specifically, fit mutants direct courtship towards males and also elicit courtship from wild-type males. By analyzing fit's role in courtship behavior, we also shed light on the role the fat body plays in modulating behavior. These studies provide the first pieces of evidence that gene expression changes occur in Drosophila males performing reproductive behaviors. This novel approach identified behaviorally important loci that are expressed in the nervous system and the fat body, indicating that both tissues modulate behavior. Also identified were sex-determination hierarchy target genes and it is likely that further analysis of the remaining candidates will reveal more members of this genetic cascade.