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>Diss. (sammanfattning) Umeå : Umeå universitet, 1992, härtill 8 uppsatser</p> / digitalisering@umu

Male mate choice occurs in species in which males face high mating costs, leading to few opportunities to copulate. Additionally, if male-male competition is strong, male mate choice should be influenced by male fighting ability. Good competitors should choose high quality females because they may be able to fight off contenders, while poor competitors should choose lower quality females. In Nephila clavipes spiders, males have limited sperm supply and fight for access to females. Here, using field experiments and observational data, we tested whether female quality and male size (a proxy of fighting ability) affect the pairing likelihood of males. In our experiments, we found that males did not express mate choice initially, but, after a few hours, chose their mates based on female size and female recent pairing status. Importantly, male mate choice direction and intensity varied with male size, as large males guarded larger females that were not recently paired and small males guarded smaller females that were recently paired. The observational data provided little evidence of assortative pairing regarding body size. With our field experiments, we show that crucial information can be collected by assessing mate choice in multiple moments. Taken together, our findings reveal the importance of considering male traits and the social context in understanding variation in male mate choice, which is often neglected in empirical studies / A escolha de parceiras por machos ocorre em espécies em que machos sofrem altos custos associados ao acasalamento, o que gera poucas oportunidades para copular. Adicionalmente, se a competição entre machos é intensa, a escolha de parceiras por machos deveria ser afetada pela habilidade de luta dos machos. Bons competidores deveriam escolher fêmeas de alta qualidade porque eles seriam capazes de afastar rivais, enquanto maus competidores deveriam escolher fêmeas de má qualidade. Em aranhas da espécie Nephila clavipes, machos possuem um estoque limitado de esperma e lutam pelo acesso a fêmeas. Usando experimentos de campo e dados observacionais, testamos se a qualidade das fêmeas e o tamanho dos machos (variável operacional de habilidade de luta) afetam a probabilidade de pareamento dos machos. Nos nossos experimentos, vimos que machos não expressam escolha de parceiras inicialmente, mas, depois de algumas horas, escolhem parceiras baseado no tamanho e no estado recente de pareamento delas. A direção e intensidade da escolha de parceiras por machos variou com o tamanho dos machos, pois machos grandes guardaram fêmeas maiores que não estavam pareadas recentemente, enquanto machos pequenos guardaram fêmeas menores que estavam pareadas recentemente. Nossos dados observacionais forneceram pouca evidência para pareamento assortativo referente a tamanho corporal. Com nossos experimentos de campo, mostramos que informação crucial pode ser coletada avaliando a escolha de parceiras em diferentes momentos. Em conjunto, nossos resultados revelam a importância de considerar características dos machos e contexto social no entendimento da variação na escolha de parceiras por machos, que é comumente negligenciada em estudos empíricos

Sexual selection plays an important role in the evolution of animals. Today we already know that it takes place not only before copulation (precopulatory sexual selection), but also after copulation. This type of sexual selection is called postcopulatory sexual selection, and occurs mainly in promiscuous species, where females mate with multiple males. In this case, sperm competition occurs in the female reproductive tract. To increase the likelihood of their reproductive success, males began to develop surprisingly diverse sperm adaptations at the morphological, physiological, or behavioural levels. These adaptations often affect sperm velocity (motility), which is a key factor for successful egg fertilization. However, the result of reproductive success can also be influenced by females, who may prefer sperm with a certain phenotype in the process of cryptic female choice, and thus, for example, obtain better genes for offspring. In some species, females even can have the ability to sort and store sperm in specialized organs in which the sperms are nourished for some time, and then used to fertilize the egg. Even in this case, the storage of sperm is often affected by sperm morphology. Due to these mechanisms of postcopulatory sexual selection, sperm are under strong selection pressure, which can...

Sexual selection is considered a powerful evolutionary force responsible for the enormous diversity found in reproductive morphology, physiology, and behavior. I addressed questions related to selection in the Cape ground squirrel (Xerus inauris), a species characterized as highly social and promiscuous. These attributes often are responsible for variance in male reproductive success and as such, sexual selection theory predicts increased opportunity for sexual selection. I confirm that the predominant mechanism underlying genital evolution and competition for paternity in X. inauris is sperm competition. I find evidence that investment in sperm competition is costly and may reflect immunocompetence. I quantify reproductive success as it relates to alternative male tactics and female resource distribution. I find that male X. inauris alternative reproductive tactics differ within and across populations most likely due to differences in female resource distribution. In areas where females are evenly distributed, dispersed males encounter more estrous females, and therefore have increased breeding opportunities. However, the decision to remain natal does not preclude reproduction. I determine that these tactics are most likely conditional with equal fitness payoffs. Males, regardless of tactic, invest more in post-copulatory competition (e.g. sperm competition, copulatory plugs) than males within a population with a clustered distribution of breeding females. In the latter area, males form dominance hierarchies that affect copulatory success and lead to greater skews in reproduction among males. Both sites have evidence of a highly skewed variance in reproduction and intense sexual selective pressure. My results suggest these populations have increased opportunities for selection but that different mechanisms of intrasexual competition may result in rapid evolutionary change within this species.

Sperm competition is a widely-recognized and powerful selective force. Male accessory glands are organs found across animal taxa that can influence sperm performance, and thus may be selected for in competitive contexts. In fishes, these organs are in fact rare, but display great diversity in form and function across species. Although the accessory gland is known to play a role in mate attraction, parental care, fertilization, or post-copulatory competition in a few select species, the role of this organ remains a mystery in most species. Many fishes with accessory glands also exhibit alternative reproductive tactics (ARTs), which add an extra layer of complexity to how species respond to sperm competition. Because males of different ARTs typically experience different levels of sperm competition risk, it’s possible they may differentially invest in accessory glands to overcome this competition. In this thesis, I used the plainfin midshipman fish (Porichthys notatus), a species with both ARTs and an accessory gland, to experimentally investigate the role of the accessory gland in sperm competition and uncover how this organ may differ between ARTs. Over a two-year period, I studied tactic-specific investment in the accessory gland in fish from the beaches of British Columbia. I also examined the effects of seminal fluid, produced in part by the accessory gland, on sperm performance and morphology. I found that males adopting the ‘guarder’ male tactic invested more in one region (the lobules) of this organ, while males adopting the smaller ‘sneaker’ male tactic invested more the other region of the gland (the nodes). Using data collected over five years, I found that guarder males also invested more in their whole accessory glands. Additionally, I report that sperm swam faster in the presence of seminal fluid, and seminal fluid increased sperm head size in both male tactics and increased midpiece size in guarder males. These results suggest that the plainfin midshipman accessory gland may have dual functions, one of which may be to aid sperm competitive ability through enhancements in swimming speed and potentially more successful sperm morphology. Taken together, the results of my thesis improve our knowledge of the role of non-sperm components like seminal fluid and the accessory gland in sperm competition, and demonstrate how species with ARTs can have varying physiological responses to such competition. Only a handful of studies have considered the effects of seminal fluid on sperm performance. By examining sperm competition in a more biologically relevant way and incorporating the effects of a little-studied organ that impacts sperm competition, we should be able to more generally and accurately appreciate the dynamics of post-copulatory competition and fertilization. / Thesis / Master of Science (MSc)

Postcopulatory sexual selection—sperm competition and cryptic female choice—has become a major area of research over the past 40 years. Within this field there are many outstanding questions at every level of analysis, from proximate to ultimate. The fitness consequences for both sexes in the period after copulation and before fertilization are considerable, but are obscured within the female reproductive tract. Our understanding of postcopulatory mechanisms is especially sparse in taxa other than birds and insects. Nearly nothing is known in reptiles except that multiple paternity is common and widespread, and often results from long-term sperm storage across breeding seasons. We present some of the very first data on the determinants of fertilization success in the context of sperm competition in reptiles, a group that accounts for 30% of terrestrial vertebrates. In the first chapter, "Asymmetric gametic isolation between two populations of red-sided garter snakes", we discuss the use of between-population crosses to reveal gametic isolation. The effect of population density and operational sex ratios on mating systems and the speciation process has fueled theoretical debate. We attempted to address these issues using two populations of red-sided garter snakes (Thamnophis sirtalis parietalis) from Manitoba, Canada. Our study populations differ markedly in their density mating aggregations, with a 10-fold difference between them. Using microsatellite markers for paternity analysis of litters produced from within and between population crosses. We found that the population with highest aggregation density, and presumably with the highest level of sexual conflict (i.e., when the evolutionary interests of the sexes differ) over mating, was also the population that exhibited homotypic sperm precedence. The less dense population showed a distinct postcopulatory male-size advantage. We also demonstrated that sperm stored within the female over hibernation can father 20-30% of offspring in a litter. In the second chapter, "Sperm competition and mate-order effects in red-sided garter snakes", we test whether females use mate-order effects to ensure that a larger (fitter) male will sire her offspring. Does that second male should have precedence in sperm competition? We tested for second-male precedence using singly-mated females that mated with a second male. Average proportion of paternity was shared equally among the first (P₁, i.e., proportion of offspring from a litter fathered by the first male to mate) and second males (P₂) to mate, and stored sperm (P[subscript ss]). This may be a case where last male precedence breaks down with more than two males. All females were spring virgins (they had not mated that spring, but may have stored sperm from fall matings); thus sperm stored presumably from fall matings is important in this system. As the interval between matings increased P₁ increased at the expense of P[subscript ss]. As the second male to mate's copulation duration increased, P₁ also increased at the expense of P₂. This last result may indicate female influence over sperm transfer during coerced matings. Copulatory plugs (CPs) are found in many taxa, but the functional significance is debated. Male garter snakes produce a gelatinous copulatory plug during mating that occludes the opening of the female reproductive tract for approximately two days. In chapter three, "Not just a chastity belt: the role of mating plugs in red-sided garter snakes revisited", we experimentally tested the role of the CPs. In snakes, sperm are produced in the testes and delivered through the ductus deferens, and the copulatory plug is thought to be produced by the sexual segment of the kidney and conveyed through the ureter. We manipulated the delivery of the two fluids separately by ligating the ducts. We confirmed that the CP is not formed in ureter-ligated males and that sperm leaks out immediately after copulation. The CP is analogous to a spermatophore. The protein matrix contains most of the sperm which are liberated as the plug dissolves within the female's vaginal pouch. One of the fundamental principles in sperm competition is that increased sperm numbers increase the odds of winning in competitions for fertilization success and males will adjust their ejaculate relative to competition and the quality of his mate. In chapter four, "Sperm depleted males and the unfortunate females who mate with them", we detect significant among-male variation in the number of sperm ejaculated, and that male mate-order reduces sperm numbers. Male sperm numbers drop significantly from one mating to the next, and this has implications for sperm competiveness, as Thamnophis sirtalis exhibits a disassociated reproductive tactic, in that sperm stores are produced outside the breeding season, and thus cannot be replenished after mating. Interestingly, however, the on average the mobility of the sperm increased for a male's second mating. Therefore, increased sperm quality may compensate for reduced numbers in a competitive context. Further, females increase their remating rate when mating with males that are unable to deliver sperm. In chapter five, "Sexual conflict during mating in red-sided garter snakes as evidenced by genital manipulation", we revisited the CP in the context of sexual conflict. Sex-differences in optimal copulation duration can be a source of conflict, as increased copulation duration may be advantageous for males as it delays female remating. Males of many species actively guard females to prevent them from remating, and in some cases males produce copulatory plugs to prevent remating. If precopulatory choice is limited at the time of her first mating, conflict may be especially onerous to a female. The size of the plug is influenced by the copulation duration. We experimentally tested the contribution of male and female control over copulation duration. We ablated the largest basal spine on the male's hemipene and found a reduction in copulation duration and an increase in the variation of plug mass. Further, we anesthetized the female's cloaca and found copulation duration increased, which suggests that males benefit from increased copulation duration while females actively try to reduce copulation duration. Therefore, sexual conflict is manifest in divergent copulation duration optima for males and females. / Graduation date: 2013

Contrairement à d’autres groupes animaux, chez les primates, la hiérarchie de dominance ne détermine pas systématiquement le succès reproductif des mâles. Afin de comprendre pourquoi, j’ai étudié les stratégies de reproduction des mâles et des femelles dans un groupe de macaques rhésus de la population semi-libre de Cayo Santiago (Porto Rico), collectant des données comportementales, hormonales et génétiques pendant deux saisons de reproduction. Les résultats se résument en cinq points. 1. Les nouveaux mâles qui ont immigré dans le groupe d’étude occupaient tous les rangs les plus subordonnés de la hiérarchie de dominance et ont monté en rang suite au départ de mâles plus dominants. Ainsi, l’acquisition d’un rang supérieur s’est faite passivement, en absence de conflits. Par conséquent, les mâles dominants étaient généralement d’âge mature et avaient résidé plus longtemps dans le groupe que les mâles subordonnés. 2. L’accès des mâles aux femelles est en accord avec le « modèle de la priorité d’accès » selon lequel le nombre de femelles simultanément en œstrus détermine le rang de dominance du mâle le plus subordonné qui peut avoir accès à une femelle (p. ex. le mâle de rang 4 s’il y a quatre femelles en œstrus). Bien que les mâles dominants aient eu plus de partenaires et aient monopolisé les femelles de qualité supérieure (dominance, parité, âge) pendant leur période ovulatoire (identifiée grâce au profil hormonal de la progestérone), le rang de dominance n’a pas déterminé le succès reproductif, les mâles intermédiaires ayant engendré significativement plus de rejetons que prédit. Il est possible que ces jeunes adultes aient produit un éjaculat de meilleure qualité que les mâles dominants d’âge mature, leur donnant un avantage au niveau de la compétition spermatique. 3. Les mâles dominants préféraient les femelles dominantes, mais cette préférence n’était pas réciproque, ces femelles coopérant plutôt avec les mâles intermédiaires, plus jeunes et moins familiers (c.-à-d. courte durée de résidence). Au contraire, les femelles subordonnées ont coopéré avec les mâles dominants. La préférence des femelles pour les mâles non familiers pourrait être liée à l’attrait pour un nouveau bagage génétique. 4. L’intensité de la couleur de la peau du visage des femelles pendant le cycle ovarien était corrélée au moment de la phase ovulatoire, une information susceptible d’être utilisée par les mâles pour maximiser leur probabilité de fécondation. 5. Les femelles retiraient des bénéfices directs de leurs liaisons sexuelles. En effet, les femelles en liaison sexuelle bénéficiaient d’un niveau de tolérance plus élevé de la part de leur partenaire mâle lorsqu’elles étaient à proximité d’une source de nourriture défendable, comparativement aux autres femelles. En somme, bien que les mâles dominants aient bénéficié d’une priorité d’accès aux femelles fertiles, cela s’est avéré insuffisant pour leur garantir la fécondation de ces femelles parce que celles-ci avaient plusieurs partenaires sexuels. Il semble que l’âge et la durée de résidence des mâles, corrélats de leur mode d’acquisition du rang, aient confondu l’effet du rang de dominance. / In contrast to most animal groups, dominance hierarchy does not systematically determine male reproductive success in primates. In order to investigate why, I studied male and female reproductive strategies in a group of free-ranging rhesus macaques on Cayo Santiago, Puerto Rico. I collected behavioural, genetic, and hormonal data during two consecutive mating seasons. My results are summarized below. 1. All new males who immigrated into the study group occupied the lowest-ranking position in the dominance hierarchy and rose in rank as the higher-ranking males left the group. Achieving a higher dominance rank occurred passively, without physical conflict. Thus, dominant males were mature individuals who resided longest in the group. 2. Male access to oestrus females followed the predictions of the ‘priority of access’ model, in which the number of females in oestrus determines the rank of the lowest-ranking male who can access a female (e.g. the fourth ranking male if four females are in oestrus). Even though dominant males obtained more mating partners and monopolised higher quality females (dominance, parity, age) during the ovulation window (as identified using progesterone profiles), dominance rank did not determine reproductive success, as intermediate-ranking males sired significantly more infants than predicted. It is likely that those young, intermediate-ranking adult males produced high quality ejaculate, giving them an advantage in sperm competition. 3. Dominant males preferred high-ranking females, but this preference was not reciprocal; high-ranking females cooperated with younger and less familiar intermediate-ranking males. Conversely, subordinate females cooperated with dominant males. Female preference for non-familiar males (i.e. short residency in the group) may be explained by an attraction to a novel genetic pool. 4. Female facial color intensity during the ovarian cycle was correlated with the timing of the ovulation window. This information may be used by males in order to maximize their fertilisation probability. 5. Consort females enjoyed a higher level of tolerance from their male partner when they were in proximity to a monopolisable food source, compared to other, non-consort females. This suggests that females obtained direct benefits from their sexual consorts. In conclusion, even though dominant males had priority access to ovulating females in the group, this was insufficient to guarantee fertilisation when females had several sexual partners. It appears that males’ age and length of residency, both correlates of their rank acquisition mode, may have been confounding factors in dominance rank.

De acordo com o paradigma Darwin-Bateman, fêmeas não ganham benefícios em copular com múltiplos machos (i.e., poliandria), dado que seu sucesso reprodutivo depende principalmente da qualidade dos seus parceiros sexuais. Porém, a poliandria é um comportamento amplamente encontrado nas espécies com reprodução sexuada. O objetivo central desta tese foi explorar as possíveis causas e consequências da poliandria. No Capítulo 1, utilizamos experimentos em laboratório para investigar como a experiência social antes e depois da maturação influencia o grau de poliandria. A espécie estudada foi o grilo australiano Teleogryllus commodus, para a qual o ambiente acústico antes da maturação determina o comportamento reprodutivo das fêmeas após a maturação. Nossos resultados mostram que o grau de poliandria não varia entre fêmeas criadas em dois ambientes acústicos distintos, um composto por machos de qualidade variada (QV) e outro composto por machos de alta qualidade (AQ). As fêmeas do grupo AQ foram menos responsivas aos machos de alta qualidade e aceitaram machos independentemente de sua qualidade, um padrão diferente daquele encontrado para as fêmeas do grupo QV. Quando a qualidade média dos machos encontrados pelas fêmeas do grupo QV foi baixa, o número total de machos aceitos foi baixo, porém quando a qualidade média dos machos foi alta, o número de machos aceitos foi alto. Portanto, há uma interação entre a experiência social antes e após a maturação na determinação do grau de poliandria. No Capítulo 2, utilizamos métodos comparativos para testar se o risco de competição espermática promovido pela poliandria determina a evolução de características masculinas relacionadas à monopolização de fêmeas. Nosso modelo de estudo foram rãs da subfamília Leptodactylinae, cujas espécies podem depositar ovos na água (onde o risco de competição espermática é alto) ou dentro de tocas na terra (onde o risco de competição espermática é baixo). Encontramos que há correlação entre o tipo de local de reprodução (aquático x terrestre) e quatro características masculinas relacionadas à monopolização de fêmeas. Nossos resultados provêem uma relação causal entre características dos machos e a reprodução terrestre, mas os padrões encontrados nem sempre seguiram as previsões propostas pela hipótese de que o risco de competição espermática é menor em espécies com reprodução terrestre. Tais resultados sugerem que as pressões seletivas que agem nas quatro características masculinas exploradas aqui não são as mesmas, provavelmente devido a diferenças nas funções que cada característica tem durante as interações inter- e intra-sexuais. Ao fim desta tese, fornecemos evidências de uma nova causa da poliandria, além de mostrarmos as consequências da poliandria sobre a evolução de características dos machos / According to the Darwin-Bateman paradigm, females derive no benefit from multiple mating (i.e., polyandry) and their reproductive success is mainly influenced by the quality of their partners. However, there is increasing evidence that polyandry is ubiquitous across many animal groups. The main goal of this thesis was to explore possible causes and consequences of polyandry. In Chapter 1, we conducted an experiment to investigate how the social experience pre- and post-maturity influences the degree of polyandry. We studied the Australian cricket Teleogryllus commodus, for which we know that the acoustic environment prior to maturity alters female reproductive behavior after maturity. We found that the degree of polyandry did not differ between females reared in two acoustic environments, one composed of varied-quality (VQ) males and other composed of only high-quality (HQ) males. However, whereas females from the HQ group were less responsive to high quality male calls, accepting males regardless of their quality, females from the VQ group showed a different response. When the mean male quality experienced by a female from the VQ group was low, the total number of accepted males was low, and when the mean male quality was high, the total number of accepted males was high. Thus, the social experience pre- and post-maturation interact to determine the degree of polyandry in females of the Australian cricket. In Chapter 2, we used comparative methods to test if the risk of sperm competition promoted by polyandry drives the evolution of male traits related to female monopolization. We studied frogs of the subfamily Leptodactylinae, which lay eggs either on the water (where sperm competition risk is high) or inside terrestrial burrows (where sperm competition risk is low). We found evidence of an evolutionary correlation between the type breeding site (aquatic x terrestrial) and four male traits related to female monopolization. Our study creates a causal link between male traits and terrestrial reproduction, but the patterns not always followed the hypothesis that sperm competition risk is lower in terrestrial breeders. This finding suggests that the selective pressure acting on male traits is not the same, probably because they have different functions during inter- and intrasexual interactions. In conclusion, we provide evidence of a new cause of polyandry and show that polyandry may have consequences for the evolution of weaponry traits in males

The availability of reproductive sites is a major factor shaping the behavior of males and females in species with resource-based mating systems. Using a meta-analytic approach, we tested five predictions directly or indirectly derived from the mating system theory. We expected that reproductive site limitation would lead to: (1) intense male-male competition for resource possession; (2) high variance in male reproductive success, generating high values of opportunity for sexual selection; (3) high intensity of selection on male traits related to resource possession; (4) high sperm competition risk; and (5) low quality of paternal care. We compiled information from observational and experimental studies that compared the reproductive behavior of individuals of the same species under low and high reproductive site limitation. We found that, when reproductive sites are limited, there is a slight increase in male-male competition, with higher rates of nest takeover and agonistic interactions, and a slight increase in the selection gradient on male traits, with successful males tending to be larger than unsuccessful males. Reproductive site limitation has no consistent effect on the opportunity for sexual selection and on the sperm competition risk. However, territorial males invest more in gonads and lose less paternity when reproductive sites are limited. There is also no clear effect of reproductive site limitation on the quality of paternal care, but few studies have addressed this subject. Taken together, our findings indicate that predictions on how reproductive site limitation affects several aspects of resource-based mating systems have weak empirical support. These predictions do not consider the plasticity in the mating tactics of males and females, which make them too simplistic. Moreover, since the original proposition of the mating system theory, our understanding of sperm competition increased a lot. We now know that accurate predictions on the intensity and direction of sexual selection should take into account both pre- and post-copulatory processes. Finally, the interplay between sexual selection and parental care is complex, and the original framework of mating system theory does not provide sufficient elements to derive clear and taxonomically broad predictions / A disponibilidade de sítios reprodutivos influencia tanto o comportamento de machos quanto de fêmeas em espécies cujo sistema de acasalamento depende da defesa de recursos. Usando uma abordagem meta-analítica, testamos cinco previsões direta ou indiretamente relacionadas à teoria de sistemas de acasalamento. Esperávamos que a limitação de sítios reprodutivos promoveria: (1) aumento na competição masculina pela posse de recursos; (2) aumento na variância do sucesso reprodutivo dos machos, gerando valores alto de oportunidade para a seleção sexual; (3) aumento da intensidade da seleção sobre características masculinas relacionadas à posse de recursos; (4) aumento no risco de competição espermática e (5) diminuição da qualidade do cuidado paternal. Compilamos informações de estudos observacionais e experimentais que compararam o comportamento reprodutivo de indivíduos da mesma espécie em situação de alta e baixa disponibilidade de sítios reprodutivos. Encontramos que, quando os sítios reprodutivos são escassos, há um ligeiro aumento na competição masculina, com maiores taxas de roubo de ninhos e interações agonísticas, e um ligeiro aumento no gradiente de seleção sobre características masculinas, com machos bem sucedidos tendendo a ser maiores do que machos mal sucedidos em monopolizar recursos. A disponibilidade de sítios reprodutivos não teve nenhum efeito consistente sobre a oportunidade para a seleção sexual e o risco de competição espermática. Entretanto, machos territoriais investiram mais em gônadas e perderam menos paternidade quando os sítios reprodutivos eram escassos. Não encontramos nenhum efeito da disponibilidade de sítios reprodutivos sobre a qualidade do cuidado paternal. Em conjunto, nossos resultados indicam que as previsões sobre como a disponibilidade de sítios reprodutivos influencia diversos aspectos de sistemas de acasalamento baseados na defesa de recursos têm fraco respaldo empírico. Tais previsões não consideram a plasticidade nas táticas de acasalamento de machos e fêmeas, tornando-as demasiadamente simplistas. Adicionalmente, desde a formulação original da teoria de sistemas de acasalamento, nossa compreensão sobre competição espermática aumentou. Sabemos atualmente que previsões acuradas sobre a intensidade e direção da seleção sexual devem levar em consideração processos pré- e pós-copulatórios. Finalmente, a interconexão entre seleção sexual e cuidado parental é complexa e a teoria original de sistemas de acasalamento não provê elementos suficientes para a construção de previsões claras e de amplo escopo taxonômico