Costs Of Mating Research Articles

Encounter with heavier females changes courtship and fighting efforts of male field crickets Gryllus bimaculatus (Orthoptera: Gryllidae)

The effects of mating experience on male mating behavior are mediated by four factors: mating cost, such as resource depletion, perception of mating opportunities, self-perception of attractiveness, and female quality. For example, encountering females might increase male expectations of prospective mating opportunities, while copulation increases self-perception of attractiveness in males. To determine the relative importance of these factors, the effect of mating on the two components of reproductive effort (courtship and fighting effort) in Gryllus bimaculatus was examined. Calling activity before and after encountering females was measured, and copulation success was recorded. Subsequently, the intensity and outcome of male–male fighting behavior was recorded. Female encounter increased calling activity irrespective of copulation, thereby indicating that the perception of mating opportunities is important factor for the males. Changes in courtship effort of males were larger and fighting success was lower when they were previously paired with relatively heavier females. These results indicate that male reproductive effort is also affected by quality of previous mating partners.

Sperm Storage Costs Determine Survival and Immunocompetence in Newly Mated Queens of the Leaf-Cutting Ant Atta colombica

In the leaf-cutting ant Atta colombica, queens receive ejaculates from multiple males during one single mating event early in their lives. A queen’s fertility and fitness therefore depend on maximizing the number of sperm cells she can store and maintain inside her spermatheca. Previous studies implied significant physiological mating costs, either originating from energetic investments maximizing sperm survival, or from resolving sexual conflicts to terminate male-driven incapacitation of rival sperm via serine proteases found in seminal fluid. Here we conducted an artificial insemination experiment, which allowed us to distinguish between the effects of sperm and seminal fluid within the queen’s sexual tract on her survival and immunocompetence. We found significantly higher mortality in queens that we had inseminated with sperm, independently of whether seminal fluid was present or not. Additionally, after receiving sperm, heavier queens had a higher probability of survival compared to lightweight queens, and immunocompetence decreased disproportionally for queens that had lost weight during the experiment. These findings indicate that queens pay significant physiological costs for maintaining and storing sperm shortly after mating. On the other hand, the presence of seminal fluid within the queens’ sexual tract neither affected their survival nor their immunocompetence. This suggests that the energetic costs that queens incur shortly after mating are primarily due to investments in sperm maintenance and not costs of terminating conflicts between competing ejaculates. This outcome is consistent with the idea that sexually selected traits in social insects with permanent castes can evolve only when they do not affect survival or life-time fitness of queens in any significant way.

Evolution of nuptial gifts and its coevolutionary dynamics with male-like persistence traits of females for multiple mating

BackgroundMany male animals donate nutritive materials during courtship or mating to their female mates. Donation of large-sized gifts, though costly to prepare, can result in increased sperm transfer during mating and delayed remating of the females, resulting in higher paternity. Nuptial gifting sometimes causes severe female-female competition for obtaining gifts (i.e., sex-role reversal in mate competition) and selection on females to increase their mating rate, changing the intensity of sperm competition and the resultant paternity gains. We built a theoretical model to simulate such coevolutionary feedbacks between nuptial gift size (male trait) and propensity for multiple mating (female trait). Donation of nuptial gifts sometimes causes development of female persistence trait for gift acquisition. We also analyzed the causes and consequences of this type of traits, taking double receptacles for nutritious seminal gifts, which are known to occur in an insect group with a “female penis” (Neotrogla spp.), as an illustrative example.ResultsOur individual-based simulations demonstrated that female-female competition for male-derived nutrients always occur when the environment is oligotrophic and mating costs are low for females. However, a positive correlation between donated gift size and the resultant paternity gain was a requisite for the co-occurrence of large gifts and females’ competitive multiple mating for the gifts. When gift donation satisfied female demands and thus resulted in monandry, exaggeration of nuptial gift size also occurred under the assumption that the last male monopolizes paternity. The evolution of double slots for gift acquisition and digestion (female persistence trait) always occurred when males could not satisfy the demands of females for gifts. However, through coevolutionary reduction in male gift size, fixation of this trait in a population drastically reduced the average female fitness.ConclusionSperm usage patterns, which have rarely been examined for animals with nuptial gifts, can be a critical factor for determining the extent of exaggeration in nuptial gifting. Sex-role reversals in mate competition, as a result of donation of nuptial gifts from males to females, can involve the evolution of male-like, persistent traits in females that reduce population productivity, as is the case with persistence traits in males.

Fear of sex: sexual conflict exposed as avoidance in a parthenogenetic invertebrate

Males and females often have divergent evolutionary interests, generating sexual conflicts. This is particularly true in organisms that exhibit facultative sexuality, whereby females are capable of reproducing without fitness costs of mating. Here, we provide the first documented evidence with quantitative tracking showing that sex interacts with social context to determine space-use of females, in a pattern resembling predator avoidance. To achieve this, we labelled Daphnia magna with fluorescent nanoparticles and utilized a 3-D tracking platform to record pairs of individuals swimming. The recordings comprised either same-sex or opposite-sex pairings. We found that females swam faster, deeper, more horizontally, and more linearly when exposed to males than when exposed to females. Simultaneously, we found that male behavior did not differ depending on swimming partner and, importantly, we observed no sexual dimorphism in swimming behaviors when swimming with the same sex. Our results suggest that the presence of males in a population has the potential to influence the distribution of individuals, similarly to known threats, such as predation. This highlights that sexual conflict has clear spatial consequences and should be considered in such ecological frameworks, like the Landscape of Fear (LOF) concept. In a broader context, the connection of the evolutionary and social concept of sexual conflict and the ecological concept of LOF may improve our understanding of population dynamics and the spatial and temporal distribution of individuals in natural ecosystems.Significance statementDespite the wealth of studies that detail how predators affect their prey’s spatial behaviors, studies on the role of sex and social context on spatial behavior are rare. Addressing this dearth of information, we studied the swimming behaviors of an organism that can reproduce with or without sex, when exposed to an individual of either the same or opposite sex. We found no difference between the sexes in swimming behaviors; however, we revealed that females avoided males by swimming deeper in the water column, reminiscent of the response to predation. Our results highlight that social conflict between the sexes strongly affects the demographics of a population and may therefore have a substantial role in the spatial ecology of organisms in the wild.

Reproductive aging and pace-of-life syndromes: more active females age faster

Abstract A common pattern of reproductive aging is that reproductive performance increases during early life and reaches a peak, followed by a decline with age. Such quadratic reproductive aging patterns can differ among individuals. Moreover, if individual differences in reproductive aging patterns reflect individual-specific life-history trade-off strategies, they are also predicted to be associated with behavior according to the pace-of-life syndrome. For example, more active, aggressive, or bolder individuals may invest more in early reproduction, resulting in more rapid reproductive aging. In this study, we estimated individual differences in quadratic reproductive aging patterns and the relationship between reproductive aging and the activity of the virgin female bean bug (Riptortus pedestris) in the absence of mating costs. We found that the egg production of virgin females followed a parabolic trajectory with age and that individuals varied significantly in their quadratic reproductive aging patterns. In addition, we found that females that were relatively more active during early life invested in egg production more heavily at a young age and suffered from a sharper decline in egg production later in life. Thus, our results indicate that individual reproductive aging patterns may be a key component in the study of pace-of-life syndromes. We suggest that within-individual plastic characteristics of life-history traits such as reproductive aging patterns may explain the mixed results from multiple studies on pace-of-life syndromes.

Sexual conflict drives micro- and macroevolution of sexual dimorphism in immunity

BackgroundSexual dimorphism in immunity is believed to reflect sex differences in reproductive strategies and trade-offs between competing life history demands. Sexual selection can have major effects on mating rates and sex-specific costs of mating and may thereby influence sex differences in immunity as well as associated host–pathogen dynamics. Yet, experimental evidence linking the mating system to evolved sexual dimorphism in immunity are scarce and the direct effects of mating rate on immunity are not well established. Here, we use transcriptomic analyses, experimental evolution and phylogenetic comparative methods to study the association between the mating system and sexual dimorphism in immunity in seed beetles, where mating causes internal injuries in females.ResultsWe demonstrate that female phenoloxidase (PO) activity, involved in wound healing and defence against parasitic infections, is elevated relative to males. This difference is accompanied by concomitant sex differences in the expression of genes in the prophenoloxidase activating cascade. We document substantial phenotypic plasticity in female PO activity in response to mating and show that experimental evolution under enforced monogamy (resulting in low remating rates and reduced sexual conflict relative to natural polygamy) rapidly decreases female (but not male) PO activity. Moreover, monogamous females had evolved increased tolerance to bacterial infection unrelated to mating, implying that female responses to costly mating may trade off with other aspects of immune defence, an hypothesis which broadly accords with the documented sex differences in gene expression. Finally, female (but not male) PO activity shows correlated evolution with the perceived harmfulness of male genitalia across 12 species of seed beetles, suggesting that sexual conflict has a significant influence on sexual dimorphisms in immunity in this group of insects.ConclusionsOur study provides insights into the links between sexual conflict and sexual dimorphism in immunity and suggests that selection pressures moulded by mating interactions can lead to a sex-specific mosaic of immune responses with important implications for host–pathogen dynamics in sexually reproducing organisms.

Alternative reproductive tactics in male freshwater fish influence the accuracy of species recognition.

Sexual conflict can result in coercive mating. Because males bear low costs of heterospecific mating, coercive males may engage in misdirected mating attempts toward heterospecific females. In contrast, sexual selection through consensual mate choice can cause mate recognition cues among species to diverge, leading to more accurate species recognition. Some species show both coercive mating and mate choice‐associated courtship behaviors as male alternative reproductive tactics. We hypothesized that if the selection pressures on each tactic differ, then the accuracy of species recognition would also change depending on the mating tactic adopted. We tested this hypothesis in the guppy (Poecilia reticulata) and mosquitofish (Gambusia affinis) by a series of choice experiments. Poecilia reticulata and G. affinis males both showed imperfect species recognition and directed all components of mating behavior toward heterospecific females. They tended to direct courtship displays more frequently toward conspecific than heterospecific females. With male P. reticulata, however, accurate species recognition disappeared when they attempted coercive copulation: they directed coercions more frequently toward heterospecific females. We also found that heterospecific sexual interaction had little effect on the fecundity of gravid females, which suggests that prepregnancy interactions likely underpin the exclusion of G. affinis by P. reticulata in our region.

The Cost and Benefice of Multiple Mating in Desert Locust Schistocerca gregaria

The Cost and Benefice of Multiple Mating in Desert Locust Schistocerca gregaria

Reproductive interference hampers species coexistence despite conspecific sperm precedence.

Negative interspecific mating interactions, known as reproductive interference, can hamper species coexistence in a local patch and promote niche partitioning or geographical segregation of closely related species. Conspecific sperm precedence (CSP), which occurs when females that have mated with both conspecific and heterospecific males preferentially use conspecific sperm for fertilization, might contribute to species coexistence by mitigating the costs of interspecific mating and hybridization. We discussed whether two species exhibiting CSP can coexist in a local environment in the presence of reproductive interference. First, using a behaviorally explicit mathematical model, we demonstrated that two species characterized by negative mating interactions are unlikely to coexist because the costs of reproductive interference, such as loss of mating opportunity with conspecific partners, are inevitably incurred when individuals of both species are present. Second, we experimentally examined differences in mating activity and preference in two Harmonia ladybird species known to exhibit CSP. These behavioral differences may lead to local extinction of H. yedoensis because of reproductive interference by H. axyridis. This prediction is consistent with field observations that H. axyridis uses various food sources and habitats whereas H. yedoensis is confined to a less preferred prey item and a pine tree habitat. Finally, by a comparative approach, we observed that niche partitioning or parapatric distribution, but not sympatric coexistence in the same habitat, is maintained between species with CSP belonging to a wide range of taxa, including vertebrates and invertebrates living in aquatic or terrestrial environments. Taken together, it is possible that reproductive interference may destabilize local coexistence even in closely related species that exhibit CSP.

Success rate, genetic improvement and economic analysis of artificial insemination delivery models for smallholder pig production systems

Context Smallholder pig production systems in the north-eastern region of India are characterised by inefficiency due to use of unselected local pigs, and multiple breeding constraints such as paucity of superior germplasm, high mating costs, and poor access to artificial insemination (AI). Effective AI is the most viable option for genetic improvement and enhancing pig productivity. Aim The study was designed to assess the success rate and economic efficacy of four AI delivery models (Models I–IV): (I) AI at farmer’s doorstep through direct linkage to tribal farmers; (II) AI at farmer’s doorstep through trained, educated youth; (III) oestrus synchronisation and fixed-time AI; and (IV) mobile van-based AI delivery system. The study also sought to ascertain the benefits of adopting a crossbreeding program through AI over the existing use of local pigs for smallholder systems. Methods Semen collected from nine Hampshire boars was routinely utilised for AI after processing as per standard protocols. For Models I–IV, respectively, 259, 154, 183 and 284 sows from 75 villages/clusters were inseminated. Reproductive and litter performance as well as insemination cost were evaluated for each model. Growth performance of crossbred pigs obtained through AI was compared with that of local pigs in the smallholder production system. Key results Farrowing rate and mean litter size at birth were 74.90% and 8.57 ± 0.52 for Model I, 68.83% and 8.08 ± 0.37 for Model II, 80.87% and 9.31 ± 0.41 for Model III, 75.00% and 8.54 ± 0.64 for Model IV, and 81.58% and 6.81 ± 0.29 for natural service. Farrowing rate, litter size at birth and at weaning, and litter weight at weaning were significantly (P < 0.05) affected by AI delivery model in the order: Model III > Model I = Model IV > Model II. Litter weight at birth was significantly (P < 0.05) affected by AI delivery model in the order: Model III > Model I = Model II = Model IV. Lowest cost per insemination was found for Model IV (INR 319.00) followed by Model II (INR 449.00), Model III (INR 629) and Model I (INR 899). All models had lower cost than the natural service system (INR 2200). The growth performance of crossbred pigs obtained through AI was significantly (P < 0.01) higher than of the local pigs reared in the same production system. Conclusion The present study justifies the accessibility, feasibility and potential benefit of AI delivery models in smallholder/backyard pig production systems in the north-eastern region of India. Use of these AI delivery models will overcome the breeding constraints and reward farmers with more rapid genetic improvement in productivity with lower production costs. Model III (oestrus synchronisation and fixed time AI) shows best productivity and Model IV (mobile van-based AI delivery system) has lowest cost. Implications These AI delivery models will be effective for sustainable and economic piggery development and improve the socioeconomic livelihood of pig farmers in South Asian countries.

Oviposition and refractory periods of the small water strider Microvelia horvathi (Heteroptera: Veliidae) after a single mating event

AbstractMultiple mating can be costly, but also beneficial for females. When the costs outweigh the benefits, mated females can be reluctant to remate for some time. Conversely, females will be likely to remate when they need to replenish sperm and/or receive nutrients contained in the accessory ejaculate substances. Water striders have been commonly used in mating studies as many show a vigorous struggle between the sexes. Generally, water strider females frequently mate with many males and lay many eggs during mating period. However, little is known about their oviposition and refractory periods after an initial mating event. In this study, we revealed that single‐mated females of the small water strider, Microvelia horvathi, could lay eggs for 40 days, reflecting approximately half of their potential oviposition periods. It suggested that M. horvathi females do not need to mate to replenish sperm for some time once they mate. Our results also showed that no female remated after 1 or 3 days of an initial mating event. The proportion of female remating increased with time after the initial mating, however, remating levels reached 26.7% even after 14 days. In this sense, any significant cost of mating outweighs the benefit, which might be one of the factors lowering their remating levels.

Extra-pair paternity and antiparasitic defence

BackgroundExtra-pair paternity (EPP) in birds provides benefits in terms of more offspring, and characteristics for maintenance of this behaviour have been the subject of investigation. Microorganisms are known to be transmitted during mating, especially when mating with multiple partners, and factors reducing this cost of multiple mating are expected. Further, plumage brightness and colour intensity have been shown to be important traits to benefits from multiple mating as predicted by sexual selection. The aim of this study was to investigate the relationship between the rate of extra-pair paternity and the relative size of the uropygial gland at the interspecific level, as the uropygial gland is an exocrine gland hypothesized to produce antiparasitic substances and further identified to affect plumage brightness. Because of the expected benefits of large uropygial gland in scenarios of sexual selection, we predicted a positive correlation with EPP.MethodsWe collected information from the literature of uropygial gland size and frequency of extra-pair paternity of 60 avian species of different families and explored the predicted positive correlation between them. We did so with means of comparative analyses that considered phylogenetic relationship as random factor and included body mass as covariate. We used Markov chain Monte Carlo generalized linear mixed models that were weighted by number of nests used to estimate extra-pair paternity.ResultsWe detected a positive relationship between level of extra-pair paternity and uropygial gland size at an interspecific level. This finding is consistent with the prediction.ConclusionsWe discuss the importance of this result in scenarios of sexual selection and argue that the detected relationship may have arisen by utilizing antiparasitic secretions through secondary sexual characters indicating parasite resistance.

Cost of mating in male Menochilus sexmaculatus (Fabricius) (Coleoptera: Coccinellidae)

AbstractMating and reproduction not only provide benefits but can also incur substantial costs. An increasing number of recent studies have indicated that mate searching, mating, and ejaculate production requires large amount of energy which can result in high cost to males. Here, we examine the effect of number of matings of males and mating duration on mating and reproductive success in Menochilus sexmaculatus (Coleoptera: Coccinellidae) by subjecting them to 3 consecutive matings of 30, 60, 90 min, and complete mating (a) with no interval and (b) with interval of 24 hr between the matings. The results reveal a reduction in reproductive performance for both the setup and reduction is more when mating is consecutive without gap. This shows that between subsequent matings males require time to recuperate the sperm/ejaculate. As a male engages in a greater number of matings, the time it takes to begin mating when paired with a female increases and the overall duration of mating decreases. However, frequent mating attempts made by males result in reduction in their body weight. Thus, in this ladybird beetle mating and related physiological process consume lots of energy and the body weight of males tend to decrease which further modifies their reproductive performance. The study helps us to identify the cost of mating from male's perspective and indicates that males need to rejuvenate between the matings.

Polyandry provides reproductive and genetic benefits in colonising populations

Polyandry, when females mate with more than one male, is theorised to play an important role in successful colonisation of new habitats. In addition to possible benefits from sexual selection, even mild polyandry could facilitate colonisation by protecting against inbreeding and reducing the costs of mating with incompatible or infertile males. Here, we measure the importance of mild polyandry for population viability and reproductive fitness following experimental founder events into a higher‐temperature regime. Using colonisation experiments with the model beetle Tribolium castaneum, in which females can produce offspring for up to 140 days following a single mating, we founded more than 100 replicate populations using single females that had been given the opportunity to mate with either one or two males and then tracked their subsequent population dynamics. Following population viability and fitness across 10 generations, we found that extinction rates were significantly lower in populations founded by females given polyandrous opportunities to mate with two males (9%) compared to populations founded by monogamous females (34%). In addition, populations founded by females that had been provided with opportunities to store sperm from two different males showed double the median productivity following colonisation compared to monogamous‐founded populations. Notably, we identified short‐term and longer‐term benefits to post‐colonisation populations from double‐mating, with results suggesting that polyandry acts to both protect against mating with incompatible males through the founder event, and reduce inbreeding depression as the colonisation proceeds for 10 generations. Our results therefore show that even mild polyandry provides both reproductive and genetic benefits for colonising populations.

Frequent mating reduces male mating rate but not offspring quality or quantity in a neriid fly

Frequent mating can deplete sperm and seminal fluid, limiting male ability to sire offspring. Frequent mating could also deplete non-genetic ejaculate components that affect offspring quality. These effects of frequent mating on male reproductive success may be mediated by male condition, or by modification of subsequent male mating behaviour. Using the neriid fly Telostylinus angusticollis, we conducted two experiments to examine whether a history of frequent mating affects males’ subsequent mating rate and offspring traits. The first experiment tested whether male condition (manipulated by varying larval diet quality) and mating history affects male performance in a subsequent mating with a single novel female whereby we predicted effects of previous mating may be more prevalent in low-condition males. Prior mating resulted in a reduction in mating rate with the novel female, but we did not detect an effect of mating history or male condition on offspring quality or quantity. The second experiment tested whether costs of mating become more evident when males encounter multiple novel females. Surprisingly, while prior mating once again resulted in a reduction in mating rate with the novel females, we still did not detect an effect of condition or mating history on any offspring traits. Therefore, male neriid flies appear to be able to mate many times without suffering a reduction in offspring quality or quantity. The apparent lack of an effect of frequent mating on such traits could be mediated by a reduction in mating rate, reflecting male prudence with ejaculate expenditure.

Disentangling the costs of mating and harassment across different environments

Why females mate multiply has been a long-standing question in evolutionary ecology. In attempts to answer this question, many studies on diverse taxa have highlighted various costs and benefits associated with reproduction. However, how the costs of mating differ from the costs of harassment and whether they vary with environmental conditions are unknown. To explore this, we compared various fitness traits of singly mated Callosobruchus maculatus females, with or without access to water, subjected to different levels of male exposure: no male (i.e. no mating or harassment), ablated male present (i.e. harassment only), intact male present (i.e. both mating and harassment), exposure to males for a short time (i.e. repeated mating but with minimal harassment). We found that, overall, females lived longer in wet than in dry environments, and that the effects of male exposure on female life span differed between environments: wet environments showed cumulative costs of harassment and mating, while differing levels of male exposure had no effect in dry environments. Further, while females laid more eggs in dry environments, females mating repeatedly were more fecund than females housed with ablated males in both environments, suggesting both benefits of repeated mating and costs of harassment, which are environmentally independent. Finally, offspring survival was dependent on a complex interaction between environment and male exposure, potentially reflecting environmental differences in resource allocation trade-offs.

Females of a solitary bee reject males to collect food for offspring

AbstractThe time dedicated to courtship and copulation is the most general cost of mating for females. However, quantitative estimates of this cost and the consequences for female mating behavior have been investigated for only a few model organisms and mostly under laboratory conditions. We determined the costs of copulations and persistent courtship by males in terms of time for females of the solitary bee Anthrenoides micans. We estimated the rate and duration of male mating behaviors and the consequences for sexual interactions for females with respect to the loss of foraging opportunity in the wild. Males invested most of their time searching for mates and intercepted foraging females every 3 min. Copulas lasted, on average, 10 times longer than the time females took to resist male mating attempts. Despite the high frequency of these rejections (82%), females spent 3-fold more time copulating than rejecting males. Considering the rate of encounters with males and the mean duration of flower visits by females, we estimated that females would perform 64% fewer flower visits per hour if they accepted all copulation attempts. The loss of time is especially significant in the natural habitat of the species, where host cacti blossom for extraordinary short periods of time and females compete with other cacti-specialized bees and conspecifics. Because the offspring production of a female solitary bee depends on its pollen collection capacity, reduced foraging performance directly influences female reproductive success.

Is selfing a reproductive assurance promoting polyploid establishment? Reduced fitness, leaky self-incompatibility and lower inbreeding depression in neotetraploids.

Newly formed polyploids face significant obstacles to persistence and population establishment because of fitness costs of intercytotype mating. Selfing provides the opportunity to escape mate limitation, enabling production of new individuals and increasing the likelihood of fixation of new polyploid lineages. Still, association between self-compatibility and polyploidy is not always clear. We compared self-incompatibility and inbreeding depression in neotetraploids and their diploid progenitor to explore the direct effects of whole genome duplications on self-incompatibility and the implications of ploidy-driven changes for polyploid establishment. Outcross and self-pollinations were performed in diploids and synthetic neotetraploids of Jasione maritima var. maritima, and reproductive success was measured through fruit and seed production and seed germination. Self- and outcross offspring were grown under controlled conditions, and plant performance was measured through several fitness parameters. Neotetraploids showed an overall lower performance than diploids. Reproductive success was negatively affected by selfing in both cytotypes. However, greater variation in the expression of self-incompatibility was observed in neotetraploids; additionally, developmental and physiological parameters were not affected by selfing on neotetraploids, with an overall similar fitness of outcrossed and selfed individuals, resulting in lower inbreeding depression indexes. Neotetraploids might have benefited from selfing at initial stages after their formation. Genome duplications resulted in leaky self-incompatibility, enabling the production of offspring under minority cytotype disadvantage with similar fitness as outcrossed offspring. Our results support theoretical assumptions that selfing might be important for neopolyploid establishment, although changes in self-incompatibility might not be abrupt.

Costs and benefits of multiple mating in a species with first-male sperm precedence.

Different patterns of sperm precedence are expected to entail different costs and benefits of mating for each sex that translate into distinct predictions regarding mating system evolution. Still, most studies addressing these costs and benefits have focused on species with mixed paternity or last male precedence, neglecting first-male sperm precedence. We attempted to understand whether this latter pattern of sperm precedence translates into different costs and benefits for each sex in the haplodiploid spider mite Tetranychus urticae, a species in which female multiple mating is prevalent but most offspring aresired by first males. First, we assessed the stability of the sperm precedence pattern. To do so, we measured offspring paternity after exposing females to a different number of matings and mating intervals. Next, to determine the potential costs or benefits of multiple mating for females under different contexts, we measured the fecundity and survival of females that re-mated at different time points. To measure the potential costs of multiple mating for males, we analysed male survival in the presence of different numbers of virgin or mated females. We also tested whether males can reduce offspring production of their competitors, by reducing the production of fertilized offspring of mated females. We found no change in the pattern of sperm precedence, independently of the mating interval between matings and the number of matings. Females paid a cost of mating, as multiply-mated females laid fewer eggs than once-mated females. However, while males had reduced survival when exposed to an intermediate number of virgin females, they paid no additional costs of mating with mated females. Moreover, females that mated multiple times produced fewer fertilized offspring than females that mated once. Thus, males that copulated with mated females reduced the fitness of other males, potentially leading to a relative fitness benefit for themselves. Our results show that complex costs and benefits may arise in males in species with first-male sperm precedence. How these costs and benefits affect the maintenance of selection for polyandry remains an open question.

Individual Quality and Extra-Pair Paternity in the Blue Tit: Sexy Males Bear the Costs.

Adaptive explanations for the evolution of extra-pair paternity (EPP) suggest that females seek extra-pair copulations with high quality males. Still, the link between ornamentation, individual quality, and paternity remains unclear. Moreover, honest signaling is essential when explaining EPP because it is needed for sexual selection to occur; yet, it is understudied in multiple ornaments. Because blue tits (Cyanistes caeruleus) show variable color expression in several plumage patches, we tested: (i) over two seasons, whether males in better condition, more ornamented and less infected by blood parasites gain EPP and have higher reproductive success, and (ii) over three seasons, whether mating patterns affect color change. Males with more saturated yellow feathers, brighter tails, and in better condition had higher reproductive success in one of the seasons. Contrary to expectation, in another season, males that gained EPP were parasitized by blood parasites, suggesting increased vector exposure during extra-pair copulations. Our results for two seasons show that males siring more extra-pair young were older and grew brighter cheek or tail feathers for the following season. Despite the increased mating costs, in socially monogamous avian systems, high quality males incur in EPP without compromising traits that may be under sexual selection.