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Seasonal phenology and mating frequency of moths in the genus Dioryctria found sympatrically in north Okanagan Valley, British Columbia seed orchards were assessed. Female moths in the abietella, auranticella, ponderosae and schuetzeella species groups were trapped in Douglas-fir, lodgepole pine and interior spruce stands. Most species were univoltine based on one peak of flight activity per season. There is evidence that the abietella group are bivoltine in this region. Females in the abietella and auranticella groups are polyandrous; ponderosae and schuetzeella females are monandrous. The sole abietella species, D. abietivorella, recorded in British Columbia can have substantial economic impacts on seed production in commercial seed orchards. Factors influencing reproductive behaviour, longevity and fecundity of D. abietivorella were investigated. Females are synovigenic and have an income-breeding mating strategy. Reproductive behaviours are delayed post-eclosion and signalling receptivity by females coincides with egg maturation, increasing with age. Female D. abietivorella experience trade-offs between reproduction and longevity. / Ecology

Submitted by (edna.saturno@ufrpe.br) on 2016-11-22T15:55:38Z No. of bitstreams: 1 Agna Rita dos Santos Rodrigues.pdf: 1041996 bytes, checksum: 09225af0315071627118074737e4ebda (MD5) / Made available in DSpace on 2016-11-22T15:55:38Z (GMT). No. of bitstreams: 1 Agna Rita dos Santos Rodrigues.pdf: 1041996 bytes, checksum: 09225af0315071627118074737e4ebda (MD5) Previous issue date: 2008-02-01 / Conselho Nacional de Pesquisa e Desenvolvimento Científico e Tecnológico - CNPq / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / The reasons of multiple and long successive mates accomplished by Podisus nigrispinus (Dallas) (Hem.: Pentatomidae) males has been unknown. Thus, mating behavior of P. nigrispinus was investigated as function of male body size (i), previous mating status (ii), risk of predation (iii), interrupted mating (iv), and spermatogenesis (v). The female choice for mating with large or small body size males was investigated under three conditions: multiples choice within a population of large and small males; partial choice (ca. only one pair of small and large male) and no choice (only small or large male). Time to initiate a copula and its duration was observed after pairing males and females under the risk of predation imposed by the presence of the predatory wasp Polistes versicolor Oliver (Hym.: Vespidae). Furthermore, females had their reproductive success evaluated with interrupted mating at different intervals (30, 60, 120 and 240 minutes afterinitiating a mating) and when mating with males previously mated. The spermatogenesis of these mated and unmated males was investigated. The outcomes show that P. nigrispinus female mates with males disregarding their body size, previous mating status and, exhibits no mating partner preference in successive mates. In addition, duration of mating, partner choice and female fertility were not influenced by male body size. The risk of predation did not alter mating behavior and duration of mating. However, females with interrupted mating at 30, 60, 120 and 240 minutesexhibited low egg viability (0, 3.1, 7.7, and 34%, respectively) compared to females with uninterrupted mating (338 to 671 minutes) that showed egg viability of 74.2%. The dissection of males of different mating status shows that P. nigrispinus exhibits continuous spermatogenesis and absence of a storage seminal vesicle. Spermatozoa were present in the lumen of the vas deferents independent of mating status. The spermatheca does not change size according to the duration of mating, although its duct exhibited elasticity with significant deformation after mating for 30 minutes. Females under uninterrupted mating exhibited spermathecal duct fully expanded and spermatheca filled with spermatozoa. Based on the results, the reproductive success of P. nigrispinus male is related to the duration of copula and female requires multiple mates to fill spermatheca with spermatozoa. However, the reproductive success of the species does not depend on males’ body size and their mating status. / A realização de múltiplas e longas cópulas por Podisus nigrispinus (Dallas) (Hem.: Pentatomidae) tem sido uma incógnita. Assim, este trabalho teve como objetivo investigar o comportamento reprodutivo de macho de P. nigrispinus. A cópula foi estudada em função do tamanho do macho, status prévio de acasalamento do macho e risco de predação (i). Também, foram analisados a espermatogênese e o efeito do tempo de cópula na transferência de espermatozóides para fêmeas (ii), bem como foram descritas a espermateca e estruturas relacionadas quanto à morfologia interna e modificações ocorridas durante a cópula (iii). A escolha da fêmea por machos virgens ou previamente acasalados foi estudada sob chance múltipla, parcial ou sem chance de escolha por machos grandes ou pequenos. O comportamento para o acasalamento e duração da cópula sob risco de predação [presença de Polistes versicolor Oliver (Hym.: Vespidae)] foi comparado com casais sem risco de predação. Fêmeas tiveram osucesso de fertilização e inseminação avaliado através da interrupção da cópula após 30, 60, 120 e 240 minutos. Em função do status de acasalamento, machos foram dissecados e investigados quanto a produção e transferência de espermatozóides para as fêmeas. Os resultados mostram que o acasalamento em P. nigrispinus independe do tamanho e status prévio de acasalamento dosmachos, além de não apresentar preferência de parceiro. A duração da cópula, escolha do parceiro e a fertilidade das fêmeas não sofrem influência do tamanho do macho. Também, o risco de predação não alterou o comportamento de acasalamento e duração de cópula. Fêmeas com cópulas interrompidas após 30, 60, 120 e 240 minutos apresentaram baixa viabilidade de ovos (0; 3,1; 7,7; e 34%, respectivamente), enquanto aquelas sem interrupção de cópula (338 a 671 minutos) obtiveram 74,2% de viabilidade de ovos. Também, foi observado que machos possuem espermatogênese contínua e ausência de vesícula seminal. A análise do ducto deferente mostra espermatozóides presentes no seu lúmen independente do status de acasalamento (machos virgens, 0h, 12h ou 24h após término da cópula). Em fêmeas, o volume da espermateca não sofre alteração com a cópula. Contudo, ocorre mudança de formato do ducto elástico da espermateca a partir de 30 minutos do início da cópula. Fêmeas que tiveram cópula sem interrupção apresentaram espermatecas completamente preenchidas com espermatozóides. Para P. nigrispinus o sucesso de inseminação e fertilização de fêmeas é dependente da duração da cópula e de múltiplas cópulas. Por outro lado, o sucesso reprodutivo não depende do tamanho do macho e de seu status de acasalamento.

As abelhas são insetos da ordem Hymenoptera e taxonomicamente estão reunidas na superfamília Apoidea. Os Meliponini, da subfamília Apinae, são popularmente chamados abelhas indígenas sem ferrão e algumas espécies são essenciais para a polinização de plantas selvagens e lavouras. As abelhas Nannotrigona testaceicornis, utilizadas neste trabalho, são pequenas, possuem o tórax marrom escuro e opaco e não são agressivas. No Brasil, elas são encontradas na Bahia, Espírito Santo, Goiás, Minas Gerais, Rio de Janeiro e São Paulo. Assim como muitas espécies de Hymenoptera, elas também podem nidificar em agregações e analisar a variabilidade genética dentro e entre os agregados foi um dos objetivos deste trabalho, além de avaliar os prováveis múltiplos acasalamentos que parecem ocorrer dentro dessa espécie de meliponíneos. Para tanto, foram coletados indivíduos de ninhos de N. testaceicornis no Campus da USP de Ribeirão Preto (SP), Campinas (SP), Bonfim Paulista (SP), Uberlândia (MG) e Caratinga (MG). 302 indivíduos (operárias) de um total de 32 ninhos foram macerados e tiveram seu DNA extraído. Foram utilizados oito loci microssatélites específicos para o estudo da variabilidade genética e múltiplos acasalamentos. Um total de 38 alelos foi observado. A diversidade genética média entre as agregações foi de 35,4%, a heterozigose média esperada foi de 41,4% e a observada foi de 28,3%. O coeficiente de endogamia (Fis) foi estatisticamente significativo para todas as agregações. Também foram observados desvios no equilíbrio de Hardy-Weinberg para todos os loci analisados. A análise do Fst mostrou diferenciação entre as agregações, porém, apesar da indicação pelo AMOVA de uma discreta estruturação entre as agregações, não foi possível agrupá-las. A partir dos dados obtidos foi possível observar que a diversidade genética nestas agregações está baixa, além disso, os desvios do equilíbrio, o valor significativo do Fis, e a heterozigose observada menor do que a esperada em todos os loci demonstrou a ocorrência de cruzamentos endogâmicos resultando em alto nível de homozigose nesta população. Com relação aos múltiplos acasalamentos, dos 32 ninhos estudados, somente 11 (34,4%) apresentaram uma patrilínea. A presença de alelos nulos, a inclusão acidental de machos na amostra, mudanças recentes de rainhas, operárias de outros ninhos e a contínua conexão entre ninho mãe-filho podem ser possibilidades para estes resultados, entretanto, a frequência de mais de uma patrilínea é muito alta para que a poliandria não seja considerada. / Bees are insects belonging to the Hymenoptera order and to the Apoidea superfamily. The Meliponini, from the subfamily Apinae, are popularly known as stingless bees and some species are essential to the pollination of wild and farming plants. Nannotrigona testaceicornis, studied in this work, are small bees presenting a dark brown and opaque thorax, and non-aggressive behavior. In Brazil, they are found at Bahia, Espírito Santo, Goiás, Minas Gerais, Rio de Janeiro and São Paulo states. As well as many other Hymenoptera species, they can nest on aggregations. To analyze the genetic variability within and among the aggregates was one of the objectives of this work, besides to evaluate the possibility of multiple matings which seem to occur in this species. For this purpose, samples of N. testaceicornis were collected from nests found at University of São Paulo Campus at Ribeirão Preto (SP), Campinas (SP), Bonfim Paulista (SP), Uberlândia (MG) and Caratinga (MG). Three hundred and two individuals (workers) from thirty-two sampled nests had their DNA extracted. Eight specific microsatellite loci were used to study the genetic variability and multiple matings. A total of thirty-eight alleles were observed. The mean genetic diversity among the aggregations was 35.4%, the mean expected heterozygosity was 41.4% and the observed was 28.3%. The inbreeding coefficient (Fis) was statistically significant for all the aggregations. Deviations from Hardy-Weinberg Equilibrium were observed in all studied loci. The Fst analysis showed differentiation among the aggregations but, despite the AMOVA indication of structuring among them, it was not possible to group them. From the obtained data we observed that the genetic diversity of the aggregations is low. The significant Fis value and the observed heterozygosity lower than the expected one in all the studied loci, demonstrated inbreeding events resulting in a high level of homozygosis in this population. In relation to the possibility of multiple mating, from the 32 studied nests, only 11 (34.4%) presented one patriline. The presence of null alleles, the accidental inclusion of males in the sample, recent queen changes, workers drifting among nests, continuing connections between mother and daughter nests or multiple matings might be possibilities for these results. However, the frequency of more than one patriline is very high to not consider the polyandry.

<p>Multiple mating by females is common in nature. Yet, the evolution and maintenance of polyandry remains a bit of an evolutionary puzzle. It was my aim in this thesis to reach a greater understanding of this phenomenon as well as to investigate the consequences of polyandry on the evolution of reproductive divergence in insects. In an extensive meta analysis addressing the direct effects of multiple mating on female fitness in insects, I found that insects gain from multiple matings in terms of increased lifetime offspring production. In species without nuptial feeding, increased mating rate leads to decreased female lifespan and my results strongly support the existence of an intermediate optimal female mating rate. However, results from an experimental study where I examined the relationship between female fitness and mating rate in the bean weevil (Callosobruchus maculatus) showed that female fitness was maximized at two alternative mating rates, indicating that some species may exhibit a more complex relationship between the costs and benefits of mating. In the meta analysis on species with nuptial feeding, I found only positive effects of increased mating rate and the puzzle is rather what constrains the actual mating rates of females in these groups.</p><p>Sexual selection is a very potent driver of rapid evolutionary change in reproductive characters. Most research has focussed on precopulatory sexual selection, but in promiscuous species sexual selection continues after copulation and variance in male fertilization success gives rise to postcopulatory sexual selection. In this thesis I found that three allopatric populations of the red flour beetle (Tribolium castaneum) have diverged in traits related to reproduction. Male genotype affected all aspects of female reproduction, but more interestingly, males and females interacted in their effect on offspring production and reproductive rate, showing that the divergence was due to the evolution of both male and female reproductive traits.</p><p>When studying postcopulatory sexual selection, sperm competition has been put forward as the main source of variance in fertilization success. The results from a set of double-mating experiments, using the same populations of flour beetles, provided strong evidence that cryptic female choice is also important in generating variance in male fertilization success. I found not only main effects of female genotype on male fertilization success but also male-female interactions which provide more unambiguous evidence for cryptic female choice. Finally, I attempted to uncover which male signals-female receptors are involved in the reproductive divergence observed in the Tribolium populations. In a double-mating experiment I manipulated female perception of two male reproductive signals, copulatory courtship and cuticular hydrocarbons, and the results indicate that, within populations, both signals are sexually selected. However, only male cuticular hydrocarbons seem to be involved in the reproductive divergence between the populations. </p><p>In conclusion, multiple mating by female insects can be understood solely in terms of direct fitness benefits resulting from increased offspring production. I have shown that postcopulatory sexual selection can lead to rapid divergence in reproductive traits related to mating and that cryptic female choice plays an important role in this divergence.</p>

<p>Harmony and cooperation was for long believed to dominate sexual interactions. This view slowly started to change 25 years ago and is today replaced with a view where males and females act based on what is best from a costs-benefits perspective. When sex specific costs and benefits differ, concerning reproductive decision influenced by both sexes, sexual conflict will occur. The basis for discordant reproductive interests between the sexes is that males produce many small gametes, while females’ produce few and large gametes. One result of this difference is that the optimal mating rate differs between the sexes. Males, with their many small sperm, maximize their reproductive output by mating with many females, while females often do best by not mating more frequently than to fertilize their eggs, since mating often entails a cost. Sexual conflict over mating is thus an important factor shaping the interactions between the sexes. In this thesis I study this and related conflicts between the sexes, using mathematical models, fruit flies and comparative methods. Mathematical modelling was used to explore how males and females may coevolve under sexual conflict over mating. This model shows that sexual conflict over mating results in the evolution of costly female mate choice, in terms high resistance to matings, and costly exaggerated male sexual traits, aimed to manipulate females into mating. A key assumption in this model is that males which females find attractive also are more harmful to females. This assumption was tested by housing fruit fly females with either attractive or unattractive males. Females kept with attractive males were courted and mated more, and suffered a 16 percent reduction in lifetime offspring production. In another study I measured genetic variation in two antagonistic male traits used to compete over females; offence - a male’s ability to acquire new mates and supplant stored sperm, and defence - a male’s ability to induce fidelity in his mates and prevent sperm displacement when remating occurs. Independent additive genetic variation and positive selection gradients were found for both these traits, indicating an ongoing arms race between these male antagonistic traits. This arms race also had a negative impact on females, since high values of offence compromised female fitness. Genetic variation in female ability to withstand male harm was also tested for and found, indicating that females evolve counter adaptations to reduce the effect of harmful male traits. Finally, the proposed link between sexual conflict and speciation was tested. Theory suggests that perpetual sexual arms races will cause allopatric populations to evolve along different evolutionary trajectories, resulting in speciation. This theory was tested using comparative methods by contrasting the number of extant species in taxa with high and low opportunity for sexual conflict. The study showed that taxa with high opportunity for sexual conflict, on average, has four times as many species as those with low opportunity, supporting that sexual conflict is a key process in speciation.</p>

Multiple mating by females is common in nature. Yet, the evolution and maintenance of polyandry remains a bit of an evolutionary puzzle. It was my aim in this thesis to reach a greater understanding of this phenomenon as well as to investigate the consequences of polyandry on the evolution of reproductive divergence in insects. In an extensive meta analysis addressing the direct effects of multiple mating on female fitness in insects, I found that insects gain from multiple matings in terms of increased lifetime offspring production. In species without nuptial feeding, increased mating rate leads to decreased female lifespan and my results strongly support the existence of an intermediate optimal female mating rate. However, results from an experimental study where I examined the relationship between female fitness and mating rate in the bean weevil (Callosobruchus maculatus) showed that female fitness was maximized at two alternative mating rates, indicating that some species may exhibit a more complex relationship between the costs and benefits of mating. In the meta analysis on species with nuptial feeding, I found only positive effects of increased mating rate and the puzzle is rather what constrains the actual mating rates of females in these groups. Sexual selection is a very potent driver of rapid evolutionary change in reproductive characters. Most research has focussed on precopulatory sexual selection, but in promiscuous species sexual selection continues after copulation and variance in male fertilization success gives rise to postcopulatory sexual selection. In this thesis I found that three allopatric populations of the red flour beetle (Tribolium castaneum) have diverged in traits related to reproduction. Male genotype affected all aspects of female reproduction, but more interestingly, males and females interacted in their effect on offspring production and reproductive rate, showing that the divergence was due to the evolution of both male and female reproductive traits. When studying postcopulatory sexual selection, sperm competition has been put forward as the main source of variance in fertilization success. The results from a set of double-mating experiments, using the same populations of flour beetles, provided strong evidence that cryptic female choice is also important in generating variance in male fertilization success. I found not only main effects of female genotype on male fertilization success but also male-female interactions which provide more unambiguous evidence for cryptic female choice. Finally, I attempted to uncover which male signals-female receptors are involved in the reproductive divergence observed in the Tribolium populations. In a double-mating experiment I manipulated female perception of two male reproductive signals, copulatory courtship and cuticular hydrocarbons, and the results indicate that, within populations, both signals are sexually selected. However, only male cuticular hydrocarbons seem to be involved in the reproductive divergence between the populations. In conclusion, multiple mating by female insects can be understood solely in terms of direct fitness benefits resulting from increased offspring production. I have shown that postcopulatory sexual selection can lead to rapid divergence in reproductive traits related to mating and that cryptic female choice plays an important role in this divergence.

Harmony and cooperation was for long believed to dominate sexual interactions. This view slowly started to change 25 years ago and is today replaced with a view where males and females act based on what is best from a costs-benefits perspective. When sex specific costs and benefits differ, concerning reproductive decision influenced by both sexes, sexual conflict will occur. The basis for discordant reproductive interests between the sexes is that males produce many small gametes, while females’ produce few and large gametes. One result of this difference is that the optimal mating rate differs between the sexes. Males, with their many small sperm, maximize their reproductive output by mating with many females, while females often do best by not mating more frequently than to fertilize their eggs, since mating often entails a cost. Sexual conflict over mating is thus an important factor shaping the interactions between the sexes. In this thesis I study this and related conflicts between the sexes, using mathematical models, fruit flies and comparative methods. Mathematical modelling was used to explore how males and females may coevolve under sexual conflict over mating. This model shows that sexual conflict over mating results in the evolution of costly female mate choice, in terms high resistance to matings, and costly exaggerated male sexual traits, aimed to manipulate females into mating. A key assumption in this model is that males which females find attractive also are more harmful to females. This assumption was tested by housing fruit fly females with either attractive or unattractive males. Females kept with attractive males were courted and mated more, and suffered a 16 percent reduction in lifetime offspring production. In another study I measured genetic variation in two antagonistic male traits used to compete over females; offence - a male’s ability to acquire new mates and supplant stored sperm, and defence - a male’s ability to induce fidelity in his mates and prevent sperm displacement when remating occurs. Independent additive genetic variation and positive selection gradients were found for both these traits, indicating an ongoing arms race between these male antagonistic traits. This arms race also had a negative impact on females, since high values of offence compromised female fitness. Genetic variation in female ability to withstand male harm was also tested for and found, indicating that females evolve counter adaptations to reduce the effect of harmful male traits. Finally, the proposed link between sexual conflict and speciation was tested. Theory suggests that perpetual sexual arms races will cause allopatric populations to evolve along different evolutionary trajectories, resulting in speciation. This theory was tested using comparative methods by contrasting the number of extant species in taxa with high and low opportunity for sexual conflict. The study showed that taxa with high opportunity for sexual conflict, on average, has four times as many species as those with low opportunity, supporting that sexual conflict is a key process in speciation.

As abelhas são insetos da ordem Hymenoptera e taxonomicamente estão reunidas na superfamília Apoidea. Os Meliponini, da subfamília Apinae, são popularmente chamados abelhas indígenas sem ferrão e algumas espécies são essenciais para a polinização de plantas selvagens e lavouras. As abelhas Nannotrigona testaceicornis, utilizadas neste trabalho, são pequenas, possuem o tórax marrom escuro e opaco e não são agressivas. No Brasil, elas são encontradas na Bahia, Espírito Santo, Goiás, Minas Gerais, Rio de Janeiro e São Paulo. Assim como muitas espécies de Hymenoptera, elas também podem nidificar em agregações e analisar a variabilidade genética dentro e entre os agregados foi um dos objetivos deste trabalho, além de avaliar os prováveis múltiplos acasalamentos que parecem ocorrer dentro dessa espécie de meliponíneos. Para tanto, foram coletados indivíduos de ninhos de N. testaceicornis no Campus da USP de Ribeirão Preto (SP), Campinas (SP), Bonfim Paulista (SP), Uberlândia (MG) e Caratinga (MG). 302 indivíduos (operárias) de um total de 32 ninhos foram macerados e tiveram seu DNA extraído. Foram utilizados oito loci microssatélites específicos para o estudo da variabilidade genética e múltiplos acasalamentos. Um total de 38 alelos foi observado. A diversidade genética média entre as agregações foi de 35,4%, a heterozigose média esperada foi de 41,4% e a observada foi de 28,3%. O coeficiente de endogamia (Fis) foi estatisticamente significativo para todas as agregações. Também foram observados desvios no equilíbrio de Hardy-Weinberg para todos os loci analisados. A análise do Fst mostrou diferenciação entre as agregações, porém, apesar da indicação pelo AMOVA de uma discreta estruturação entre as agregações, não foi possível agrupá-las. A partir dos dados obtidos foi possível observar que a diversidade genética nestas agregações está baixa, além disso, os desvios do equilíbrio, o valor significativo do Fis, e a heterozigose observada menor do que a esperada em todos os loci demonstrou a ocorrência de cruzamentos endogâmicos resultando em alto nível de homozigose nesta população. Com relação aos múltiplos acasalamentos, dos 32 ninhos estudados, somente 11 (34,4%) apresentaram uma patrilínea. A presença de alelos nulos, a inclusão acidental de machos na amostra, mudanças recentes de rainhas, operárias de outros ninhos e a contínua conexão entre ninho mãe-filho podem ser possibilidades para estes resultados, entretanto, a frequência de mais de uma patrilínea é muito alta para que a poliandria não seja considerada. / Bees are insects belonging to the Hymenoptera order and to the Apoidea superfamily. The Meliponini, from the subfamily Apinae, are popularly known as stingless bees and some species are essential to the pollination of wild and farming plants. Nannotrigona testaceicornis, studied in this work, are small bees presenting a dark brown and opaque thorax, and non-aggressive behavior. In Brazil, they are found at Bahia, Espírito Santo, Goiás, Minas Gerais, Rio de Janeiro and São Paulo states. As well as many other Hymenoptera species, they can nest on aggregations. To analyze the genetic variability within and among the aggregates was one of the objectives of this work, besides to evaluate the possibility of multiple matings which seem to occur in this species. For this purpose, samples of N. testaceicornis were collected from nests found at University of São Paulo Campus at Ribeirão Preto (SP), Campinas (SP), Bonfim Paulista (SP), Uberlândia (MG) and Caratinga (MG). Three hundred and two individuals (workers) from thirty-two sampled nests had their DNA extracted. Eight specific microsatellite loci were used to study the genetic variability and multiple matings. A total of thirty-eight alleles were observed. The mean genetic diversity among the aggregations was 35.4%, the mean expected heterozygosity was 41.4% and the observed was 28.3%. The inbreeding coefficient (Fis) was statistically significant for all the aggregations. Deviations from Hardy-Weinberg Equilibrium were observed in all studied loci. The Fst analysis showed differentiation among the aggregations but, despite the AMOVA indication of structuring among them, it was not possible to group them. From the obtained data we observed that the genetic diversity of the aggregations is low. The significant Fis value and the observed heterozygosity lower than the expected one in all the studied loci, demonstrated inbreeding events resulting in a high level of homozygosis in this population. In relation to the possibility of multiple mating, from the 32 studied nests, only 11 (34.4%) presented one patriline. The presence of null alleles, the accidental inclusion of males in the sample, recent queen changes, workers drifting among nests, continuing connections between mother and daughter nests or multiple matings might be possibilities for these results. However, the frequency of more than one patriline is very high to not consider the polyandry.

The aim of the project was to test female advantages of promiscuous and/or polyandrous mating. Although, previous studies reported improved reproductive success of females copulating with multiple males in many model species, including Eublepharis macularius, this study failed to demonstrate such a benefit of polyandry. Traits reflecting female reproductive success, i.e. the number of produced eggs, egg mass, hatchability, hatchling mass, growth and survival were not influenced by copulation with multiple partners, but some of them were dependent on female condition. Only initial stages of juvenile growth were influenced by female identity and/or investment.