abstract: An important component of insect social structure is the number of queens that cohabitate in a colony. Queen number is highly variable between and within species. It can begin at colony initiation when often unrelated queens form cooperative social groups, a strategy known as primary polygyny. The non-kin cooperative groups formed by primary polygyny have profound effects on the social dynamics and inclusive fitness benefits within a colony. Despite this, the evolution of non-kin queen cooperation has been relatively overlooked in considerations of the evolution of cooperative sociality. To date, studies examining the costs and benefits of primary polygyny have focused primarily on the advantages of multiple queens during colony founding and early growth, but the impact of their presence extends to colony maturity and reproduction. In this dissertation, I evaluate the ecological drivers and fitness consequences of non-kin queen cooperation, by comparing the reproduction of mature single-queen versus polygynous harvester ant (Pogonomyrmex californicus) colonies in the field. I captured and quantified the total number and biomass of reproductives across multiple mating seasons, comparing between populations that vary in the proportion of single queen versus polygynous colonies, to assess the fitness outcomes of queen cooperation. Colonies in a mainly polygynous site had lower reproductive investment than those in sites with predominantly single-queen colonies. The site dominated by polygyny had higher colony density and displayed evidence of resource limitation, pressures that may drive the evolution of queen cooperation. I also used microsatellite markers to examine how polygynous queens share worker and reproductive production with nest-mate queens. The majority of queens fairly contribute to worker production and equally share reproductive output. However, there is a low frequency of queens that under-produce workers and over-produce reproductive offspring. This suggests that cheating by reproducing queens is possible, but uncommon. Competitive pressure from neighboring colonies could reduce the success of colonies that contain cheaters and maintain a low frequency of this phenotype in the population. / Dissertation/Thesis / Doctoral Dissertation Biology 2017

Polygyny is the most common mating system in mammalian species (95%), yet our understanding of polygynous systems and microevolutionary processes is still limited. Pinniped mating systems range from extreme polygyny (e.g. elephant seals) to sequential female defence by males and hence have often been used as models for mating system studies. Parentage analysis has enabled the examination of mating success, the identification of pedigrees, and the elucidation of social organisation, greatly enhancing our understanding of mating systems (Chapter 1). However, such analyses are not without pitfalls, with erroneous assignments common in open systems (i.e. when parental and offspring samplings are incomplete). We investigated the effects of the user-defined parameters on the accuracy of parental assignment using two commonly used parental allocation programme, CERVUS and PASOS (Chapter 2). We showed that inaccurate user-defined parameters in CERVUS and PASOS can lead to highly biased output e.g. the assignment rate at 95% CL of offspring with a sampled known mother to sampled males decreased from 58% to 32% when the proportion of candidate males sampled in the parameter options decreasing 4-fold. We found that the use of both CERVUS and PASOS for parentage assignment can increase the likelihood of correctly allocating offspring to sampled parents to 97% in our study system. Incorrect parental assignment can bias estimates of various biological parameters, such as lifetime reproductive success and mate choice preference, and hence bias ecological and evolutionary interpretations. Here, we propose solutions to increase the power of parentage assignment and hence decrease the bias in biological parameter estimates. In addition, we analysed the effects of the intrinsic bias in likelihood assignment approaches towards assigning higher probability of parentage on individuals with rare alleles and those with heightened offspring-parent matches, which increase with the number of homozygous loci (Chapter 3). We showed that, as a consequence of the algorithms employed in the programmes CERVUS and PASOS, heterozygote males with rare genotypes are assigned higher rates of parentage than males with common alleles. Consequently, where two males could both be biological fathers of a given offspring, parentage assignment will more often go to the male with the rarer alleles (most often in heterozygous loci). Thus, the commonly used parentage assignment methods may systematically bias the results of parentage analyses towards supporting the notion that females prefer more genetically unusual, most often heterozygous, males. Such a bias may sway investigators towards incorrectly supporting the concept that females choose genetically more unusual males for heterozygosity fitness benefits that underpin the good genes hypothesis, when in fact no such relationship may exist. In polygynous mammals, successful males mate with multiple females by competing with and limiting the access of other males to females. When the status of many males (age, size, health, genetic etc.) prevents them from achieving the primary mating tactic, theory predicts selection for a diversification of male mating tactics. Recent studies in pinnipeds have shown that observed male mating success was correlated to male paternity success in some species (elephant-seals), but not in others (grey seals). The existence of alternative mating strategies can explain those discrepancies. Chapter 4 implemented the guidelines provided in Chapter 2 and 3 and focused on the polygynous New Zealand fur seal Arctocephalus forsteri, predicting that 1) competition for females is likely to cause a diversification of male mating tactics; and 2) that alternative tactics can yield reproductive success. Our results indicated three male behavioural profiles; one corresponded to large territorial males and two illustrated a continuum of alternative tactics employed by non-territorial subordinate males. Our study highlights that holding a territory is not a necessary condition for reproductive success in a population of otariids. The degree of sexual size dimorphism in polygynous species is expected to increase with the degree of intra-sexual competition and in turn with the degree of polygyny. The life history of an individual is the pattern of resource allocations to growth, maintenance, and reproduction throughout its lifetime. Both females and males incur viability costs of mating and reproduction. However, male viability costs due to increase growth and male-male competition can be greater than female viability costs of mate choice and reproduction. Although an abundant literature on sexual dimorphism in morphology, physiology, and parasite infections is available, little is known on the intra-sexual differences in physiology and parasite infections associated to the reproductive success of different mating strategies in mammalian species. Chapter 5 examined the reproductive costs between territorial and subordinate males New Zealand fur seal related to their relative reproductive success using a multidisciplinary approach (behaviour, genetics, endocrinology, parasitology). We found that dominant New Zealand fur seal males endure higher reproductive costs due to the direct and indirect effects of high testosterone levels and parasite burdens. Our study highlights that holding a territory confers a higher reproductive success, but induces higher costs of reproduction that may impair survival. Understanding microevolutionary processes associated to polygynous systems is fundamental in light of the ongoing anthropogenic alteration of the environment through climatic variations and habitat reduction which ultimately affect opportunity for sexual selection and shape the life history trade-offs.

Cap. 2 Com a finalidade de detectar possíveis diferenças na eficiência do processo de aprovisionamento e postura nas células de cria (POP), comparamos uma colônia monogínica e uma políginica em Melipona bicolor. Ao início das observações na colônia políginica, esta tinha três rainhas das quais uma deixou de ovipositar na metade do período de observações. Por isso separamos em duas partes a base de dados da colônia poligínica de acordo com o número de rainhas poedeiras, três ou duas. Observamos grandes diferenças entre as duas condições das colônias, a monogínica e a poligínica. Nosso estudo permitiu perceber como é a competição entre as rainhas em uma colônia poligínica. Neste tipo de colônia, as rainhas se “precipitavam" por ovipositar e como conseqüência os POPs eram muito mais curtos, diminuindo a postura de ovos tróficos. A rainha da colônia monogínica consumiu mais ovos tróficos e para conseguí-los, aumentava a duração do POP, especificamente a duração do processo de pós-aprovisionamento. O número de operárias em colônias poligínicas que participam do POP, tanto operculadoras quanto aprovisionadoras, também é maior. Isto pode refletir na eficiência do POP em colônias poligínicas: atrai um número maior de operárias que conseguem terminar o processo em menor tempo. Mas também reflete a competição entre as rainhas da colônia poligínica. Nesta, as rainhas estimulavam o POP em células que ainda não estavam com o colar pronto, dificultando seu fechamento pelas operárias. Como resultado da combinação de todos estes fatores, a produção total de ovos é maior na colônia políginica, embora cada rainha individualmente bote menos ovos na colônia poligínica do que a da colônia monogínica. Também detectamos um tipo de dominância, que se refletia no tempo que as rainhas demoravam em colocar seus ovos. Cap. 3 Para entender a poliginia, perguntou-se quais seriam as condições necessárias que levariam a uma associação estável de rainhas. A hipótese sobre “incentivos para permanecer" para as rainhas subordinadas, pode ser relevante para discutir o processo inverso testemunhado em uma colônia poligínica de Melipona bicolor que retornou à condição monogínica. Em nossa colônia poligínica, as três rainhas eram similares em vários aspectos. Não foram encontradas diferenças na competitividade na hora de ovipositar ou consumir ovos tróficos, nem discriminações por parte das operárias. Assim, cada rainha tinha igual oportunidade para pôr ovos ou se alimentar. As rainhas tinham idades semelhantes, e poder-se-ía esperar que o seu desempenho na produção de ovos fosse semelhante. Contudo, elas apresentaram uma clara desigualdade reprodutiva. Também ficou claro que a duração do processo de aprovisionamento e postura (POP) “per se", revelou diferenças como um “marca-passo" das rainhas, especialmente quando estavam sozinhas nas células de cria. A presente investigação descreve possíveis relações entre a taxa de postura das rainhas e fatores nutricionais. Discute-se sobre a provável existência de um feromônio que regularia, nas operárias, o comportamento de fechamento das células. Também se acredita que as operárias possam ter um papel importante no contexto reprodutivo entre múltiplas rainhas. Estes fatores combinados, teriam incrementado as diferenças entre as rainhas, o que levou à interrupção de postura e ao desaparecimento do favo, de duas das rainhas. Cap. 4 Ao acompanhar individualmente as operárias da Melipona bicolor, em colônias monogínica e poligínica, encontramos que existem grandes variações no comportamento de cada uma, sendo que aqueles indivíduos que tinham colocado algum tipo de ovo apareciam como os mais interessados no processo de aprovisionamento e postura (POP). Ao dividir a população de operárias em poedeiras e não poedeiras, constatamos que aquelas que colocaram ovo trófico contribuem efetivamente ao POP, mas que as operárias reprodutivas, apesar de seu pequeno número, conseguem níveis de atividade tão altos que aparentemente excluem as não poedeiras do POP. Também encontramos indivíduos “especiais" por sua extraordinária participação em um ou mais dos comportamentos básicos que compõem o POP, mas estes também parecem movidos principalmente pelo drive de colocar algum tipo de ovo. Entretanto, também encontramos diferenças entre as colônias monogínica e poligínica, mas estas foram difíceis de analisar devido ao momento em que cada colônia começou a ser filmada. Os padrões gerais encontrados tanto para a colônia monogínica, como para a poligínica são muito importantes, pois demonstram que o fato de botar ou não ovos, provoca diferenças quantitativas de comportamento, produzindo desde indivíduos “preguiçosos" até aqueles “hiperativos". Acreditamos que o desenvolvimento ovariano é a causa interna proximal que produz os diversos limiares de resposta individuais. Estes limiares produzem um efeito-cascata, já que a repetição da tarefa é reforçada no próprio indivíduo, e ao mesmo tempo, a atuação deste indivíduo afeta o repertório das outras operárias. A observação das diferenças apresentadas entre operárias poedeiras e não poedeiras, apresenta uma base muito firme para continuar na busca das causas proximais para tal diferenciação. Cap. 5 Os estudos da divisão do trabalho em insetos sociais demonstraram que embora a diferenciação de tarefas baseada na idade seria um fator importante, este mecanismo sozinho imporia uma rigidez nas sociedades que as conduziriam provavelmente à extinção. A plasticidade então é reconhecida como essencial, e a diferenciação entre indivíduos geneticamente similares deve existir. Contudo, a especialização nas tarefas de alguns indivíduos conduz a um melhor desempenho na colônia. Neste capítulo fornecemos evidências de que ocorre especialização no processo de aprovisionamento e postura (POP): poucos indivíduos “especiais" que apresentaram um repertório comportamental específico, e realizam a primeira regurgitação de alimento em Melipona bicolor. Cap. 6 A determinação de castas é um tema amplamente estudado, mas que continua deixando muitas questões a serem solucionadas. Contudo, compreendemos alguns fatos. Reconhece-se a influência de fatores externos como a quantidade e/ou a qualidade do alimento, atuando com hormônios e genes. Já que são as operárias as responsáveis pela administração da quantidade ou da qualidade do alimento, a determinação de castas é socialmente controlada e talvez, entre as aprovisionadoras, existam indivíduos que prefiram criar irmãs rainhas ao invés de irmãs operárias. Para esclarecer esta pergunta, há necessidade de estabelecer a presença ou ausência de secreções glandulares nutritivas no alimento larval, além de técnicas para examinar e acompanhar as operárias aprovisionadoras de alimento larval. Cap. 7 Nos insetos sociais, demonstrou-se que o grau de desenvolvimento dos ovários está correlacionado com relações de dominância, níveis de atividade e com a produção de hormônio juvenil. No epitélio folicular dos ovários, o hormônio juvenil estimula a absorção de nutrientes, mas ao mesmo tempo sabemos que a oogênese é um processo limitado pela disponibilidade de nutrientes. Estas são as causas e efeitos inter-relacionados na reprodução de um indivíduo. Uma das diferenças maiores entre as abelhas do gênero Apis e os meliponíneos, é que as abelhas sem ferrão conseguem desenvolver os ovários e reter a capacidade de botar ovos (embora comumente de natureza trófica) em colônias com rainha. As operárias jovens normalmente desenvolvem seus ovários, principalmente aquelas atendendo ao processo de aprovisionamento e postura (POP). As diversas tarefas relacionadas ao processo são realizadas com diferentes níveis de intensidade por cada indivíduo participando dele. Como demonstramos no capítulo quatro, as operárias que colocam ovos são os indivíduos mais envolvidos no POP contribuindo eficientemente ao processo. Quando em uma colônia monogínica de Melipona bicolor dissecamos as participantes do POP, revelamos que há uma correlação significativa entre a área dos ovários de cada indivíduo com o grau de atividade por ele apresentada. Já que os fatores nutricionais tem um papel importante no desenvolvimento dos ovários, nosso método de correlacionar o peso final com a atividade das abelhas, demonstrou ser adequado para testar indiretamente a influência do alimento na oogênesis. Neste capítulo discutimos o significado evolutivo de manter a capacidade de produzir ovos, em relação com a divisão de trabalho em insetos eussociais. / Cap. 2 In order to detect possible differences in the efficiency of the Provisioning and Oviposition Process (POP), we compared a monogynous from a polygynous colony of Melipona bicolor. At the beginning of observations, the polygynous colony had three laying queens, but one of these ceased to lay when observations where just at the middle. We then decided to split in two the database of the polygynous colony depending on the number of queens laying, being these three or two. ¶ We observed great differences between monogynous and polygynous conditions. In the polygynous colony, our investigation permitted us to perceive the strong competition between queens for laying. In this type of colony, queens were in a “rush" for laying their eggs. As a consequence, POP were very short and this diminished the number of trophic eggs laid. The queen of the monogynous colony consumed many more trophic eggs and in order to achieve this, she would increase the duration of the POP, specifically, the post-provisioning phase. In the polygynous colony, the number of workers, both dischargers and sealers were also greater than in the monogynous colony. This fact could be reflecting a more efficient process in polygynous conditions: more workers are attracted and complete the process in less time. However, again we may perceive the competition between the queens of this colony. The queens stimulated POP in cells that were still not ready, increasing the difficulty for its sealing by the workers. The total outcome of the combination of all these facts is that there is significant major production of queen eggs in the polygynous colony, although the number of eggs laid by each queen is smaller. We also detected a type of dominance reflected on the time queens spent laying their eggs. Cap. 3 There are hypotheses on which could be the necessary conditions that lead to stable multiple queen associations. Some hypothesis state mechanisms such as “staying incentives" for subordinated queens, and may be relevant for discussing the reversal process witnessed in a polygynous colony of Melipona bicolor that shifted to a monogynous condition. In a polygynous colony, three queens were similar in various aspects. Egg laying or consumption of trophic eggs, presented no differences or discriminations and each queen had an equal chance to lay or eat. The queens were of the same age, so it could be expected that they would have a similar performance in egg laying. However, a clear reproductive skew developed. It was also clear that the duration of the provisioning and oviposition process (POP), revealed differences in the pace-marking for the queens principally when alone. ¶ The present paper describes the possible relations between egg laying rate and nutritional factors. We discuss the possible existence of a queen pheromone, regulating the sealing behavior of the workers. Furthermore, workers might play a role in the contest for reproduction among multiple queens. In combination, these factors give rise to increasing differences among the queens resulting in egg laying cessation and disappearance of two of the queens from the comb. Cap. 4 When we followed individual workers of Melipona bicolor in monogynic and polygynic colonies, we found that there are great variations between the behaviors of each bee. Individuals that had laid some type of egg seemed to be the most interested in the provisioning and oviposition process (POP). Therefore, we then analyzed the population dividing them into layers and non layers. We proved that workers that lay trophic eggs contribute effectively to the POP, but reproductive workers, in spite of being so few, achieve such high levels of activity that it seems they exclude the non-laying bees from the process. We also found “special" individuals with an extraordinary participation in one or more of the basic behaviors of the POP, but these also seem to be motivated by the same drive of laying some type of egg. However, we also found differences between the monogynous and polygynous colonies, but these differences were difficult to analyze due to the filming moments for each colony. The general patterns found for both monogynous and polygynous colonies are very important because they demonstrate that the fact of laying an egg provokes quantitative differences in the behaviors, producing from “lazy" individuals up to those “hyperactive". Ovary development is then thought to be the internal proximal cause producing the diverse response thresholds that in a cascade effect, produces an auto-reinforcement when repeatedly performed the task, that at the same time affects the performance of others. The observation on the differences presented by layers and non layers, represents a strudy base to continue exploring the proximal causes for behavioral differentiation. Cap.5 Studies on division of labor have demonstrated that despite being an important factor, etharism alone would impose rigidity to societies and thus, would make them less efficient probably leading them to extinction. Plasticity of behavior is then recognized as essential being expressed as individual differentiation among genetically similar sisters. Furthermore, task specialization of some individuals leads to a better performance of the colony as a whole. In this chapter we give evidence for specialization in the provisioning and oviposition process: a task such as the first discharge of food in Melipona bicolor, is carried out mainly by few, specialized individuals that exhibit a particular behavioral repertory. Cap. 6 Caste determination is a complex topic that has received much attention but remains elusive, leaving many questions to be solved. However, there are some facts that have been comprehended. It is recognized the influence of external factors such as the amount and/or the quality of the food ingested by the larvae, interacting with hormones and genes. Since the workers administrate the larval food, caste determination is socially controlled. Could it happen that among the nurse bees there are individuals that would prefer to raise sister queens than sister workers? To answer this question, it would be necessary to establish the presence or absence of glandular secretions of high nutritional value in the larval food, together with techniques to follow and assess the food dischargers. Through video recordings made on the Provisioning and Oviposition Process (POP) in Melipona bicolor, we recognized two types of food discharges: the first fluid and shiny and another discharge dense and opaque. From each cell filmed we compiled its historical plus the caste in formation. We did not find any relation between the resulting caste and the number of solid or liquid discharges it had received. We discuss why our method may not be an accurate technique to study this difficult and complex topic, but may support future investigations. Cap. 7 Ovary development in social insects has been found to be correlated with dominance relationships, levels of activity and production of juvenile hormone. In the follicular epithelium of ovaries, juvenile hormone stimulates the absorption of nutrients; however, oogenesis is a nutrient limited process. The above roughly summarizes the interrelated causes and effects involved in the reproduction of an individual. One of the biggest differences between Apis bees and stingless bees, is that in queenright conditions, stingless bee workers develop their ovaries retaining their capacity to lay eggs (although most are of a trophic nature). Ovary development is common among young bees attending the provisioning and oviposition process (POP). The several tasks that are involved in the POP are carried on with different levels of intensity by the individuals participating in the process. As we demonstrated in the previous chapter, the individuals that are egg layers are most involved in the POP contributing efficiently to the process. Dissections revealed that there is a significant correlation between the area of the ovaries and the levels of activity presented by the workers of a monogynouos colony of Melipona bicolor. Since nutritional factors also play an important role, our method of correlating the final weight of the bees with activity, proved to be adequate to indirectly test the influence of food in oogenesis. In this chapter we discuss the meaning of retaining the capacity to produce eggs in an evolutionary scenario related to division of labor in eusocial insects.

An important question in the analysis of fertility is does the type of marital union matter? Is women's fertility affected by their marital union type? Does being in a monogamous or polygynous union have any effect on the number of children women will have? What is the nature of this difference? Does the social setting play a role? And, what kind of role does it have? Most studies, especially those based on African societies, where polygyny is most prevalent, note that female fertility is negatively associated with polygyny. This dissertation examines the polygyny-fertility relationship among married women in a different social setting, namely in an oil-rich country, that is, the United Arab Emirates. Using data from the 1999 National Family Survey for the Characteristics of the Native Households in the UAE, I was able to examine the effect of fertility using two statistical methods to model my dependent variables: Logistic Regression and Negative Binomial Regression. My findings show that when controlling for numerous biological and social variables, polygyny has a positive effect on women?s current fertility, which is the likelihood of having a baby in the past 12 months, but a negative effect on women?s cumulative fertility, which is the children ever born to them. However, when the number of wives in the union was introduced into the models, polygyny effect on women?s current and cumulative fertility has disappeared, i.e., polygyny no longer has a significant effect on women?s fertility. I discuss the implications of these findings for UAE national policy to increase the fertility of UAE women, an important consideration given that UAE nationals comprise only 20 percent of the UAE population in 2005.

Title from PDF of title page. / Document formatted into pages; contains 69 pages. / Thesis (M.S.)--University of South Florida, 2003. / Includes bibliographical references. / Text (Electronic thesis) in PDF format. / ABSTRACT: I studied the reproductive characteristics and mating systems of a central Florida population of gopher tortoises (Gopherus polyphemus). Using x-radiography, females were monitored for stage in egg-shelling and clutch size. Eggs began to appear on x-ray photographs in the first week of May in both 2001 and 2002; however, fully shelled eggs were not found before the end of May. In total 55% of the females x-rayed were gravid. Clutch sizes ranged from 4-12 with a mean of 7.29, with a mean clutch mass of 40.9 g. Clutch size increased with an increase in mean carapace length and mean plastron length. Mean clutch mass also increased with mean carapace length of females. Hatchlings began to emerge in late August, with incubation times ranging from 83 to 96 days. 50% of the eggs hatched, with 16.2% of the eggs showing no signs of development when opened. Hatchling mass averaged 30.7 g and was positively correlated with egg mass. / ABSTRACT: DNA was extracted from blood samples obtained from females and their offspring, and from the sexually mature males in the population. Nine microsatellite loci were amplified and genotypes constructed for each individual. There is evidence for promiscuous mating in gopher tortoises. Multiple paternity was detected in two of the seven clutches (28.6 %). In the clutches with multiple fathers, fertilization was highly skewed to one male, with primary male fertilizing over 70% of the clutch. Females with multiple-sired clutches were significantly smaller than females with single-sired clutches. Among the clutches assayed only one male fertilized more than one clutch, indicating that insemination of females is evenly spread among males of similar sizes. However, males assigned as fathers were significantly larger than other sampled males which may mean that larger males have an advantage in fertilization of clutches. / ABSTRACT: Conservation efforts should consider the impact of the mating system on reproduction in a population, and the possible impact of the relocation of larger males on recipient populations. / System requirements: World Wide Web browser and PDF reader. / Mode of access: World Wide Web.

Polygyny is the most common mating system in mammalian species (95%), yet our understanding of polygynous systems and microevolutionary processes is still limited. Pinniped mating systems range from extreme polygyny (e.g. elephant seals) to sequential female defence by males and hence have often been used as models for mating system studies. Parentage analysis has enabled the examination of mating success, the identification of pedigrees, and the elucidation of social organisation, greatly enhancing our understanding of mating systems (Chapter 1). However, such analyses are not without pitfalls, with erroneous assignments common in open systems (i.e. when parental and offspring samplings are incomplete). We investigated the effects of the user-defined parameters on the accuracy of parental assignment using two commonly used parental allocation programme, CERVUS and PASOS (Chapter 2). We showed that inaccurate user-defined parameters in CERVUS and PASOS can lead to highly biased output e.g. the assignment rate at 95% CL of offspring with a sampled known mother to sampled males decreased from 58% to 32% when the proportion of candidate males sampled in the parameter options decreasing 4-fold. We found that the use of both CERVUS and PASOS for parentage assignment can increase the likelihood of correctly allocating offspring to sampled parents to 97% in our study system. Incorrect parental assignment can bias estimates of various biological parameters, such as lifetime reproductive success and mate choice preference, and hence bias ecological and evolutionary interpretations. Here, we propose solutions to increase the power of parentage assignment and hence decrease the bias in biological parameter estimates. In addition, we analysed the effects of the intrinsic bias in likelihood assignment approaches towards assigning higher probability of parentage on individuals with rare alleles and those with heightened offspring-parent matches, which increase with the number of homozygous loci (Chapter 3). We showed that, as a consequence of the algorithms employed in the programmes CERVUS and PASOS, heterozygote males with rare genotypes are assigned higher rates of parentage than males with common alleles. Consequently, where two males could both be biological fathers of a given offspring, parentage assignment will more often go to the male with the rarer alleles (most often in heterozygous loci). Thus, the commonly used parentage assignment methods may systematically bias the results of parentage analyses towards supporting the notion that females prefer more genetically unusual, most often heterozygous, males. Such a bias may sway investigators towards incorrectly supporting the concept that females choose genetically more unusual males for heterozygosity fitness benefits that underpin the good genes hypothesis, when in fact no such relationship may exist. In polygynous mammals, successful males mate with multiple females by competing with and limiting the access of other males to females. When the status of many males (age, size, health, genetic etc.) prevents them from achieving the primary mating tactic, theory predicts selection for a diversification of male mating tactics. Recent studies in pinnipeds have shown that observed male mating success was correlated to male paternity success in some species (elephant-seals), but not in others (grey seals). The existence of alternative mating strategies can explain those discrepancies. Chapter 4 implemented the guidelines provided in Chapter 2 and 3 and focused on the polygynous New Zealand fur seal Arctocephalus forsteri, predicting that 1) competition for females is likely to cause a diversification of male mating tactics; and 2) that alternative tactics can yield reproductive success. Our results indicated three male behavioural profiles; one corresponded to large territorial males and two illustrated a continuum of alternative tactics employed by non-territorial subordinate males. Our study highlights that holding a territory is not a necessary condition for reproductive success in a population of otariids. The degree of sexual size dimorphism in polygynous species is expected to increase with the degree of intra-sexual competition and in turn with the degree of polygyny. The life history of an individual is the pattern of resource allocations to growth, maintenance, and reproduction throughout its lifetime. Both females and males incur viability costs of mating and reproduction. However, male viability costs due to increase growth and male-male competition can be greater than female viability costs of mate choice and reproduction. Although an abundant literature on sexual dimorphism in morphology, physiology, and parasite infections is available, little is known on the intra-sexual differences in physiology and parasite infections associated to the reproductive success of different mating strategies in mammalian species. Chapter 5 examined the reproductive costs between territorial and subordinate males New Zealand fur seal related to their relative reproductive success using a multidisciplinary approach (behaviour, genetics, endocrinology, parasitology). We found that dominant New Zealand fur seal males endure higher reproductive costs due to the direct and indirect effects of high testosterone levels and parasite burdens. Our study highlights that holding a territory confers a higher reproductive success, but induces higher costs of reproduction that may impair survival. Understanding microevolutionary processes associated to polygynous systems is fundamental in light of the ongoing anthropogenic alteration of the environment through climatic variations and habitat reduction which ultimately affect opportunity for sexual selection and shape the life history trade-offs.

abstract: Persistent cooperation between unrelated conspecifics rarely occurs in mature eusocial insect societies. In this dissertation, I present evidence of non-kin cooperation in the Nearctic honey ant Myrmecocystus mendax. Using microsatellite markers, I show that mature colonies in the Sierra Ancha Mountain of central Arizona contain multiple unrelated matrilines, an observation that is consistent with primary polygyny. In contrast, similar analyses suggest that colonies in the Chiricahua Mountains of southeastern Arizona are primarily monogynous. These interpretations are consistent with field and laboratory observations. Whereas cooperative colony founding was observed frequently among groups of Sierra Ancha foundresses, founding in the Chiricahua population was restricted to individual foundresses. Furthermore, Sierra Ancha foundresses successfully established incipient laboratory colonies without undergoing queen culling following emergence of the first workers. Multi-queen laboratory Sierra Ancha colonies also produced more workers and repletes than haplometrotic colonies, and when brood raiding was induced between colonies, queens of those with more workers had a higher survival probability. Microsatellite analyses of additional locations within the M. mendax range suggest that polygyny is also present in some other populations, especially in central-northern Arizona, albeit at lower frequencies than that in the Sierra Anchas. In addition, analyses of multiple types of genetic data, including microsatellites, the mitochondrial barcoding region, and over 2000 nuclear ultra-conserved elements indicate that M. mendax populations within the southwestern U.S. and northwestern Mexico are geographically structured, with strong support for the existence of two or more divergent clades as well as isolation-by-distance within clades. This structure is further shown to correlate with variation in queen number and hair length, a diagnostic taxonomic feature used to distinguish honey ant species. Together, these findings suggest that regional ecological pressures (e.g. colony density , climate) may have acted on colony founding and social strategy to select for increasing workforce size and, along with genetic drift, have driven geographically isolated M. mendax populations to differentiate genetically and morphologically. The presence of colony fusion in the laboratory and life history traits in honey ant that are influenced by colony size, including repletism, brood raiding, and tournament, support this evolutionary scenario. / Dissertation/Thesis / Doctoral Dissertation Biology 2018

Cap. 2 Com a finalidade de detectar possíveis diferenças na eficiência do processo de aprovisionamento e postura nas células de cria (POP), comparamos uma colônia monogínica e uma políginica em Melipona bicolor. Ao início das observações na colônia políginica, esta tinha três rainhas das quais uma deixou de ovipositar na metade do período de observações. Por isso separamos em duas partes a base de dados da colônia poligínica de acordo com o número de rainhas poedeiras, três ou duas. Observamos grandes diferenças entre as duas condições das colônias, a monogínica e a poligínica. Nosso estudo permitiu perceber como é a competição entre as rainhas em uma colônia poligínica. Neste tipo de colônia, as rainhas se “precipitavam” por ovipositar e como conseqüência os POPs eram muito mais curtos, diminuindo a postura de ovos tróficos. A rainha da colônia monogínica consumiu mais ovos tróficos e para conseguí-los, aumentava a duração do POP, especificamente a duração do processo de pós-aprovisionamento. O número de operárias em colônias poligínicas que participam do POP, tanto operculadoras quanto aprovisionadoras, também é maior. Isto pode refletir na eficiência do POP em colônias poligínicas: atrai um número maior de operárias que conseguem terminar o processo em menor tempo. Mas também reflete a competição entre as rainhas da colônia poligínica. Nesta, as rainhas estimulavam o POP em células que ainda não estavam com o colar pronto, dificultando seu fechamento pelas operárias. Como resultado da combinação de todos estes fatores, a produção total de ovos é maior na colônia políginica, embora cada rainha individualmente bote menos ovos na colônia poligínica do que a da colônia monogínica. Também detectamos um tipo de dominância, que se refletia no tempo que as rainhas demoravam em colocar seus ovos. Cap. 3 Para entender a poliginia, perguntou-se quais seriam as condições necessárias que levariam a uma associação estável de rainhas. A hipótese sobre “incentivos para permanecer” para as rainhas subordinadas, pode ser relevante para discutir o processo inverso testemunhado em uma colônia poligínica de Melipona bicolor que retornou à condição monogínica. Em nossa colônia poligínica, as três rainhas eram similares em vários aspectos. Não foram encontradas diferenças na competitividade na hora de ovipositar ou consumir ovos tróficos, nem discriminações por parte das operárias. Assim, cada rainha tinha igual oportunidade para pôr ovos ou se alimentar. As rainhas tinham idades semelhantes, e poder-se-ía esperar que o seu desempenho na produção de ovos fosse semelhante. Contudo, elas apresentaram uma clara desigualdade reprodutiva. Também ficou claro que a duração do processo de aprovisionamento e postura (POP) “per se”, revelou diferenças como um “marca-passo” das rainhas, especialmente quando estavam sozinhas nas células de cria. A presente investigação descreve possíveis relações entre a taxa de postura das rainhas e fatores nutricionais. Discute-se sobre a provável existência de um feromônio que regularia, nas operárias, o comportamento de fechamento das células. Também se acredita que as operárias possam ter um papel importante no contexto reprodutivo entre múltiplas rainhas. Estes fatores combinados, teriam incrementado as diferenças entre as rainhas, o que levou à interrupção de postura e ao desaparecimento do favo, de duas das rainhas. Cap. 4 Ao acompanhar individualmente as operárias da Melipona bicolor, em colônias monogínica e poligínica, encontramos que existem grandes variações no comportamento de cada uma, sendo que aqueles indivíduos que tinham colocado algum tipo de ovo apareciam como os mais interessados no processo de aprovisionamento e postura (POP). Ao dividir a população de operárias em poedeiras e não poedeiras, constatamos que aquelas que colocaram ovo trófico contribuem efetivamente ao POP, mas que as operárias reprodutivas, apesar de seu pequeno número, conseguem níveis de atividade tão altos que aparentemente excluem as não poedeiras do POP. Também encontramos indivíduos “especiais” por sua extraordinária participação em um ou mais dos comportamentos básicos que compõem o POP, mas estes também parecem movidos principalmente pelo drive de colocar algum tipo de ovo. Entretanto, também encontramos diferenças entre as colônias monogínica e poligínica, mas estas foram difíceis de analisar devido ao momento em que cada colônia começou a ser filmada. Os padrões gerais encontrados tanto para a colônia monogínica, como para a poligínica são muito importantes, pois demonstram que o fato de botar ou não ovos, provoca diferenças quantitativas de comportamento, produzindo desde indivíduos “preguiçosos” até aqueles “hiperativos”. Acreditamos que o desenvolvimento ovariano é a causa interna proximal que produz os diversos limiares de resposta individuais. Estes limiares produzem um efeito-cascata, já que a repetição da tarefa é reforçada no próprio indivíduo, e ao mesmo tempo, a atuação deste indivíduo afeta o repertório das outras operárias. A observação das diferenças apresentadas entre operárias poedeiras e não poedeiras, apresenta uma base muito firme para continuar na busca das causas proximais para tal diferenciação. Cap. 5 Os estudos da divisão do trabalho em insetos sociais demonstraram que embora a diferenciação de tarefas baseada na idade seria um fator importante, este mecanismo sozinho imporia uma rigidez nas sociedades que as conduziriam provavelmente à extinção. A plasticidade então é reconhecida como essencial, e a diferenciação entre indivíduos geneticamente similares deve existir. Contudo, a especialização nas tarefas de alguns indivíduos conduz a um melhor desempenho na colônia. Neste capítulo fornecemos evidências de que ocorre especialização no processo de aprovisionamento e postura (POP): poucos indivíduos “especiais” que apresentaram um repertório comportamental específico, e realizam a primeira regurgitação de alimento em Melipona bicolor. Cap. 6 A determinação de castas é um tema amplamente estudado, mas que continua deixando muitas questões a serem solucionadas. Contudo, compreendemos alguns fatos. Reconhece-se a influência de fatores externos como a quantidade e/ou a qualidade do alimento, atuando com hormônios e genes. Já que são as operárias as responsáveis pela administração da quantidade ou da qualidade do alimento, a determinação de castas é socialmente controlada e talvez, entre as aprovisionadoras, existam indivíduos que prefiram criar irmãs rainhas ao invés de irmãs operárias. Para esclarecer esta pergunta, há necessidade de estabelecer a presença ou ausência de secreções glandulares nutritivas no alimento larval, além de técnicas para examinar e acompanhar as operárias aprovisionadoras de alimento larval. Cap. 7 Nos insetos sociais, demonstrou-se que o grau de desenvolvimento dos ovários está correlacionado com relações de dominância, níveis de atividade e com a produção de hormônio juvenil. No epitélio folicular dos ovários, o hormônio juvenil estimula a absorção de nutrientes, mas ao mesmo tempo sabemos que a oogênese é um processo limitado pela disponibilidade de nutrientes. Estas são as causas e efeitos inter-relacionados na reprodução de um indivíduo. Uma das diferenças maiores entre as abelhas do gênero Apis e os meliponíneos, é que as abelhas sem ferrão conseguem desenvolver os ovários e reter a capacidade de botar ovos (embora comumente de natureza trófica) em colônias com rainha. As operárias jovens normalmente desenvolvem seus ovários, principalmente aquelas atendendo ao processo de aprovisionamento e postura (POP). As diversas tarefas relacionadas ao processo são realizadas com diferentes níveis de intensidade por cada indivíduo participando dele. Como demonstramos no capítulo quatro, as operárias que colocam ovos são os indivíduos mais envolvidos no POP contribuindo eficientemente ao processo. Quando em uma colônia monogínica de Melipona bicolor dissecamos as participantes do POP, revelamos que há uma correlação significativa entre a área dos ovários de cada indivíduo com o grau de atividade por ele apresentada. Já que os fatores nutricionais tem um papel importante no desenvolvimento dos ovários, nosso método de correlacionar o peso final com a atividade das abelhas, demonstrou ser adequado para testar indiretamente a influência do alimento na oogênesis. Neste capítulo discutimos o significado evolutivo de manter a capacidade de produzir ovos, em relação com a divisão de trabalho em insetos eussociais. / Cap. 2 In order to detect possible differences in the efficiency of the Provisioning and Oviposition Process (POP), we compared a monogynous from a polygynous colony of Melipona bicolor. At the beginning of observations, the polygynous colony had three laying queens, but one of these ceased to lay when observations where just at the middle. We then decided to split in two the database of the polygynous colony depending on the number of queens laying, being these three or two. ¶ We observed great differences between monogynous and polygynous conditions. In the polygynous colony, our investigation permitted us to perceive the strong competition between queens for laying. In this type of colony, queens were in a “rush” for laying their eggs. As a consequence, POP were very short and this diminished the number of trophic eggs laid. The queen of the monogynous colony consumed many more trophic eggs and in order to achieve this, she would increase the duration of the POP, specifically, the post-provisioning phase. In the polygynous colony, the number of workers, both dischargers and sealers were also greater than in the monogynous colony. This fact could be reflecting a more efficient process in polygynous conditions: more workers are attracted and complete the process in less time. However, again we may perceive the competition between the queens of this colony. The queens stimulated POP in cells that were still not ready, increasing the difficulty for its sealing by the workers. The total outcome of the combination of all these facts is that there is significant major production of queen eggs in the polygynous colony, although the number of eggs laid by each queen is smaller. We also detected a type of dominance reflected on the time queens spent laying their eggs. Cap. 3 There are hypotheses on which could be the necessary conditions that lead to stable multiple queen associations. Some hypothesis state mechanisms such as “staying incentives” for subordinated queens, and may be relevant for discussing the reversal process witnessed in a polygynous colony of Melipona bicolor that shifted to a monogynous condition. In a polygynous colony, three queens were similar in various aspects. Egg laying or consumption of trophic eggs, presented no differences or discriminations and each queen had an equal chance to lay or eat. The queens were of the same age, so it could be expected that they would have a similar performance in egg laying. However, a clear reproductive skew developed. It was also clear that the duration of the provisioning and oviposition process (POP), revealed differences in the pace-marking for the queens principally when alone. ¶ The present paper describes the possible relations between egg laying rate and nutritional factors. We discuss the possible existence of a queen pheromone, regulating the sealing behavior of the workers. Furthermore, workers might play a role in the contest for reproduction among multiple queens. In combination, these factors give rise to increasing differences among the queens resulting in egg laying cessation and disappearance of two of the queens from the comb. Cap. 4 When we followed individual workers of Melipona bicolor in monogynic and polygynic colonies, we found that there are great variations between the behaviors of each bee. Individuals that had laid some type of egg seemed to be the most interested in the provisioning and oviposition process (POP). Therefore, we then analyzed the population dividing them into layers and non layers. We proved that workers that lay trophic eggs contribute effectively to the POP, but reproductive workers, in spite of being so few, achieve such high levels of activity that it seems they exclude the non-laying bees from the process. We also found “special” individuals with an extraordinary participation in one or more of the basic behaviors of the POP, but these also seem to be motivated by the same drive of laying some type of egg. However, we also found differences between the monogynous and polygynous colonies, but these differences were difficult to analyze due to the filming moments for each colony. The general patterns found for both monogynous and polygynous colonies are very important because they demonstrate that the fact of laying an egg provokes quantitative differences in the behaviors, producing from “lazy” individuals up to those “hyperactive”. Ovary development is then thought to be the internal proximal cause producing the diverse response thresholds that in a cascade effect, produces an auto-reinforcement when repeatedly performed the task, that at the same time affects the performance of others. The observation on the differences presented by layers and non layers, represents a strudy base to continue exploring the proximal causes for behavioral differentiation. Cap.5 Studies on division of labor have demonstrated that despite being an important factor, etharism alone would impose rigidity to societies and thus, would make them less efficient probably leading them to extinction. Plasticity of behavior is then recognized as essential being expressed as individual differentiation among genetically similar sisters. Furthermore, task specialization of some individuals leads to a better performance of the colony as a whole. In this chapter we give evidence for specialization in the provisioning and oviposition process: a task such as the first discharge of food in Melipona bicolor, is carried out mainly by few, specialized individuals that exhibit a particular behavioral repertory. Cap. 6 Caste determination is a complex topic that has received much attention but remains elusive, leaving many questions to be solved. However, there are some facts that have been comprehended. It is recognized the influence of external factors such as the amount and/or the quality of the food ingested by the larvae, interacting with hormones and genes. Since the workers administrate the larval food, caste determination is socially controlled. Could it happen that among the nurse bees there are individuals that would prefer to raise sister queens than sister workers? To answer this question, it would be necessary to establish the presence or absence of glandular secretions of high nutritional value in the larval food, together with techniques to follow and assess the food dischargers. Through video recordings made on the Provisioning and Oviposition Process (POP) in Melipona bicolor, we recognized two types of food discharges: the first fluid and shiny and another discharge dense and opaque. From each cell filmed we compiled its historical plus the caste in formation. We did not find any relation between the resulting caste and the number of solid or liquid discharges it had received. We discuss why our method may not be an accurate technique to study this difficult and complex topic, but may support future investigations. Cap. 7 Ovary development in social insects has been found to be correlated with dominance relationships, levels of activity and production of juvenile hormone. In the follicular epithelium of ovaries, juvenile hormone stimulates the absorption of nutrients; however, oogenesis is a nutrient limited process. The above roughly summarizes the interrelated causes and effects involved in the reproduction of an individual. One of the biggest differences between Apis bees and stingless bees, is that in queenright conditions, stingless bee workers develop their ovaries retaining their capacity to lay eggs (although most are of a trophic nature). Ovary development is common among young bees attending the provisioning and oviposition process (POP). The several tasks that are involved in the POP are carried on with different levels of intensity by the individuals participating in the process. As we demonstrated in the previous chapter, the individuals that are egg layers are most involved in the POP contributing efficiently to the process. Dissections revealed that there is a significant correlation between the area of the ovaries and the levels of activity presented by the workers of a monogynouos colony of Melipona bicolor. Since nutritional factors also play an important role, our method of correlating the final weight of the bees with activity, proved to be adequate to indirectly test the influence of food in oogenesis. In this chapter we discuss the meaning of retaining the capacity to produce eggs in an evolutionary scenario related to division of labor in eusocial insects.

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Thesis (Ph. D.)--Ohio State University, 2006. / Available online via OhioLINK's ETD Center; full text release delayed at author's request until 2009 May 24