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ABSTRACT The Darwin–Bateman paradigm predicts that females enhance their fitness by being choosy and mating with high‐quality males, while males should compete to mate with as many females as possible. In many species, males enhance their fitness by defending females and/or resources used by females. That is, males directly defend access to mating opportunities. However, paternity analyses have repeatedly shown that females in most species mate polyandrously, which contradicts traditional expectations that male defensive behaviours lead to monandry. Here, in an extensive meta‐analysis, encompassing 109 species and 1026 effect sizes from across the animal kingdom, we tested if the occurrence of defensive behaviours modulates sexual selection on females and males. If so, we can illuminate the extent to which males really succeed in defending access to mating and fertilisation opportunities. We used four different indices of the opportunity for sexual selection that comprise pre‐mating and/or post‐mating episodes of selection. We found, for both sexes, that the occurrence of defensive behaviours does not modulate the potential strength of sexual selection. This implies that male defensive behaviours do not predict the true intensity of sexual selection. While the most extreme levels of sexual selection on males are in species with male defensive behaviours, which indicates that males do sometimes succeed in restricting females' re‐mating ability (e.g. elephant seals, Mirounga leonina), estimates of the opportunity for sexual selection vary greatly across species, regardless of whether or not defensive behaviours occur. Indeed, widespread polyandry shows that females are usually not restricted by male defensive behaviours. In addition, our results indicate that post‐mating episodes of selection, such as cryptic female choice and sperm competition, might be important factors modulating the opportunity for sexual selection. We discuss: (i) why male defensive behaviours fail to lower the opportunity for sexual selection among females or fail to elevate it for males; (ii) how post‐mating events might influence sexual selection; and (iii) the role of females as active participants in sexual selection. We also highlight that inadequate data reporting in the literature prevented us from extracting effect sizes from many studies that had presumably collected the relevant data. [ABSTRACT FROM AUTHOR]

Female spiders and praying mantises are renowned for their cannibalism of male partners before, during, or after mating. While several hypotheses have been proposed to explain species-specific examples of sexual cannibalism, much variation remains unexplained, including why the timing of cannibalism varies across taxa. Here, I outline how sexually cannibalistic behavior could evolve via sexually antagonistic selection as a type of behavioral resistance to male-imposed mating costs, and how such a generalizable interpretation provides a framework for understanding the evolution of both sexual cannibalism in females and anti-cannibalistic traits in males. I discuss how differences between mating systems that physiologically constrain males to mate only once (monogyny) or twice (bigyny) and systems where the sexes can potentially mate multiply (polygyny and polyandry) are likely to influence how sexual conflict shapes cannibalistic behavior. I review key examples from the literature that suggest how sexually cannibalistic behavior might function as a female resistance trait and provide comprehensive predictions for testing this hypothesis empirically. [ABSTRACT FROM AUTHOR]

Polygamy is a form of "one-sided" consensually non-monogamous relationship where one person has multiple committed partners, each of whom is only involved with that one person. It was likely a reoccurring feature of ancestral mating that posed adaptive problems for our ancestors. Yet polygamy, and multi-partnering more generally, is understudied in Western cultures, raising questions about the existence of polygamous interest and whether this is calibrated adaptively to personal conditions. In two studies, we examined polygamous interest in two heterosexual online samples from the UK. In Study 1 (N = 393), modest interest was found for polygamous relationships overall. Men were six times more open to polygyny than women, but there was little sex difference in openness to polyandry. Further analysis revealed that all forms of multi-partnering were undesirable relative to singlehood and monogamy; however, consensual multi-partner relationships were less undesirable than non-consensual ones. Sex differences were largest for polygyny and arrangements where men had agreed access to a casual partner alongside a committed one, yet these were two of the most acceptable forms of multi-partnering when men and women's responses were combined. Sociosexuality positively predicted interest in most forms of multi-partnering. Study 2 (N = 735) focused on polygyny and added status-linked traits as predictors. The results of Study 1 were broadly replicated, though the status-linked traits did not predict polygynous interest specifically. Instead, sociosexuality and male intrasexual competitiveness uniquely predicted general interest in multi-partner relationships. Overall, interest in polygamy appears to emerge despite social discouragement and sex differences in interest track the relative costs and benefits associated with it. However, there is no strong evidence that polygamous interest is uniquely calibrated to personal conditions when compared to other forms of multi-partnering. [ABSTRACT FROM AUTHOR]

The coexistence of multiple reproductives in eusocial insects is widespread, yet the decisions leading to additional queen acceptance are not well understood. Unlike in vertebrates, acceptance decisions are likely controlled by the more numerous helper population rather than the parent reproductive. Yet there are likely to be queen-worker differences in acceptance criteria because workers and queens differ in their relatedness to a secondary queen. We develop a model that examines queen-worker conflict in two scenarios: accepting a queen's sister or daughter. We additionally ask how the mating frequency and split sex ratios affect the outcomes of these conflicts. Our results reveal that conflict over queen acceptance is highest in monandrous mating systems. We identify a "window of conflict" in which a queen is selected to accept her sister but her workers do not. Our result, that polyandry neutralizes conflict over acceptance thresholds, suggests that conflict suppression may be an additional contributor to the maintenance of polyandrous mating systems. [ABSTRACT FROM AUTHOR]

In polygynous systems, such as that exhibited by reindeer Rangifer tarandus , mate choice can be difficult to disentangle from male intrasexual competition because male behavior may constrain female choice. Multiple mating may provide an avenue for female mate choice, though it is difficult to identify using behavioral estimators alone. Molecular techniques address this issue by affording ecologists an opportunity to reassess mating systems from a genetic perspective. We assessed the frequency and possible explanations for multiple mating in reindeer using a genetic approach to determine the success of observed copulations in a semi-domesticated herd in Kaamanen, Finland. Behavioral and genetic data were synthesized with population characteristics over a 7-year period to test the hypothesis that, if present, polyandry in reindeer is driven by sexual harassment from sub-dominant males. We observed multiple mating in 42% of females, with as many as 60% exhibiting multiple mating in certain years. We found no evidence that multiple mating resulted from sexual harassment by sub-dominant males, suggesting that it is likely a deliberate strategy among females. Conversion rate of copulations into paternities varied with male size, with smaller males more likely to experience mismatch than larger males. Female preference for larger males persisted despite the occurrence of multiple mating, possibly suggesting a mechanism for cryptic post-copulatory selection. We suggest further research to delineate the possible influence of cryptic post-copulatory selection and multiple mating to defend against infertility in exhausted males. [ABSTRACT FROM AUTHOR]

Polyandry can profoundly affect the strength of pre-copulatory sexual selection acting on males because each additional mate acquired by a female means that, all else being equal, a male's paternity share declines. However, accruing additional mates could benefit male fitness if males with greater mating success also tend to have less promiscuous females as partners. If this is indeed the case, then males should experience strong sexual selection to acquire more mates. We tested these predictions by collecting detailed mating data on male and female Cook Strait giant weta (Deinacrida rugosa) in the wild via daily radio tracking. Our sexual network-based approach indeed revealed that the most polygynous male D. rugosa mated the least polyandrous females. This finding therefore suggests that the most successful males likely face lower intensities of sperm competition and so should be selected to accrue more mates. Further, our selection analysis revealed significant pre-copulatory sexual selection on males with those having relatively smaller body size, lighter body mass, and longer legs accruing more mates than otherwise. Thus, it appears that both pre- and post-copulatory episodes of sexual selection reinforce the same male phenotype. [ABSTRACT FROM AUTHOR]

We investigated factors leading to variation in social complexity or 'social systems' among plateau pika family groups within a contiguous local population across 2 years. Plateau pikas are small, diurnal, nonhibernating, sexually monomorphic lagomorphs that occupy family home ranges on open alpine meadows on the Qinghai-Tibetan Plateau. Expression of the social organization, social structure, mating system and parental care system in plateau pikas did not follow expectations from traditional ecological or evolutionary explanations. Variability in plateau pika family group size and the transitions of group size between years allowed us to investigate potential advantages and disadvantages of group living. Evidence that group living served to protect pikas against predation was weak. Although social huddling could have minimized thermoregulatory costs during the extremely cold Tibetan winters, there was no correlation of overwinter survivorship among pika families of different sizes. There was no apparent group-living benefit with regard to foraging, and the occurrence of cohesive social families on the flat, continuous meadow contradicts the hypothesis that sociality is related to patchiness of critical resources. Cost of maintaining burrows appeared unrelated to group size. Most interactions between pikas occurred within family groups and were affiliative (99% of adult interactions; 97% of adult–juvenile interactions), and most interactions between adult males of different family groups were aggressive (96% of interactions). Matings were primarily within families (88% of copulations). Pikas also possess a complex vocal repertoire that enhanced interactions within social families. Demographic constraints associated with variable overwinter survivorship appeared to be the dominant precondition that produced a given family size and mating system type, coupled with selective dispersal by some pikas before the start of the breeding season. Paternal care enhanced juvenile survival, and thus led to an equalization of reproductive success among adults in families with different mating combinations. • Plateau pikas live in families occupying exclusive burrow systems and home ranges. • Pika family group size varied from one to five adults within and between years. • Pikas failed to adhere to conventional explanations for establishment of group size. • Pika families exhibited multiple social mating system types. • Pika families did not adhere to explanations for establishment of mating system types. [ABSTRACT FROM AUTHOR]

Mate choice results from the sensory or behavioural properties that cause individuals to mate with some phenotypes and not others. It can result in direct or indirect (genetic) benefits. Mate choice can also be influenced by various abiotic and biotic factors including the mating system. Here, we investigated how the mating system of the ladybird, Menochilus sexmaculatus, influences mate choice and also affects the time of commencement of mating, copulation duration, fecundity and percent egg viability. We also tested whether mate choice is dynamically affected by relatedness. We found that monandrous and polyandrous females both preferred polyandrous rather than monandrous males. In contrast, males preferred monandrous females over polyandrous females. However, females preferred unrelated monogynous males which were earlier rejected over related polygynous and males preferred unrelated polyandrous females over related monandrous females. Mating system and relatedness also impacted time to commence mating and copulation duration, fecundity and percent egg viability. This will help us to understand precisely why these shifts occur, which will be the subject of future study. [ABSTRACT FROM AUTHOR]

Despite decades of scholarly interest in the Tibetan marital practice of fraternal polyandry, very little is known about how the Tibetan family system operated in historical contexts. This study, based on a 1958 household register from Kyirong, a former district in Tibet, reconstructs nuances of family dynamics through the aid of interviews with people who were listed in the document. Kyirong's family system is shown to be very flexible. Although patrilocality was preferred, matrilocality was a viable contingency, and although polyandry was favored, monogamy and polygyny were acceptable. Despite the heterogeneity of Kyirong's family households, case studies demonstrate how people strove to achieve the monomarital stem family through polyandrous marriages in successive generations. Because polyandry created a surplus of marriageable women, joint families often arose, at least in form, when unmarried women remained with their natal families and had children, or when men discontent with their polyandrous unions moved into an adjunct house with a partner of choice. However, the offspring of these people had no rights of inheritance and thus were not integral to family continuity, so joint families in form functioned more like stem families in practice. Therefore, a discrepancy between etic definitions of form and emic understandings of process emerges when family typologies developed to facilitate cross-cultural research are incompatible with the way people actually understand rights and privileges associated with succession. The data and analysis demonstrate that Kyirong represents a unique version of a stem family society with an unambiguous stem family ideology. [ABSTRACT FROM AUTHOR]

Eusocial insects tend to present low genetic diversity (GD) within colonies, which can increase with the co-occurrence of multiple queens (polygyny) or with multiple mating by a single queen (polyandry). Therefore, it is important to elucidate how these strategies influence GD, which in turn mediate population ecology and how organisms respond to their environment. We studied two carpenter ant species from the Brazilian savanna, Camponotus renggeri and C. rufipes. Using microsatellites, we evaluated the number of breeders, the genetic relatedness and the contribution of polygyny and polyandry to GD within colonies. Both species exhibited facultative polygyny. In C. renggeri , low related queens formed colonies jointly and present low mating frequency. In this species, colony GD increased with the number of queens. Contrastingly, closely related queens of C. rufipes formed polygynous colonies, exhibiting high mating frequency. In C. rufipes , both queens and males contributed to colony GD. Despite the differences, the two species have similar GD at the colony scale. Under low mating frequency, our data support that polygyny has evolutionary importance for increasing GD in ant colonies, a mechanism mainly conferred to polyandry. Although the impact of GD in variable ecological and adaptive contexts remains uncertain, this study highlights how distinct reproductive strategies may generate similar patterns of GD in ants. [ABSTRACT FROM AUTHOR]

Knowledge on the reproductive biology of target insect pest is essential for the effective implementation of pheromone-based pest management tactics. In mealybugs, the second largest family of scale insects, the existence of female multiple mating was recently suggested. In this study, we aimed at testing how general is this behavior in mealybugs, by investigating polygyny and polyandry in two cosmopolitan pest mealybugs, Planococcus citri and Pseudococcus calceolariae. Males of these species were able to mate an average of 11.9 and 13.3 females, respectively, during their lifespan. The number of fertilized females per male decreased with male age/mating history for both mealybugs. We found no differences in female fecundity and fertility, when fertilized by males with different mating history. When we used male age as a proxy of mating history, we observed a significant negative effect on female fecundity. The females of both species remained receptive after first copula and eventually mated multiple times. The percentage of remated females of P. citri decreased linearly with time since first copula, with some maintaining receptivity up to 24 h. Males showed no preference between virgin and mated females, in static-air olfactometer tests. We found no benefit of female multiple mating in relation to fecundity. In biparental mealybugs, the mating system of males is possibly scramble competition polygyny; and that of females is possibly polyandry, with female receptivity restricted to a relatively short period. We discuss the practical implications of the results for pest management. [ABSTRACT FROM AUTHOR]

Human marriage systems, characterized by long-term partnerships and extended windows of parental care, differ from the mating systems of pulsed or seasonally breeding non-human animals in which Bateman's principles were originally tested. These features, paradigmatic of but not unique to humans, complicate the accurate measurement of mating success in evaluating Bateman's three principles. Here, we unpack the concept of mating success into distinct components: number of partners, number of years partnered, the timing of partnerships, and the quality of partners. Drawing on longitudinal records of marriage and reproduction collected in a natural-fertility East African population over a 20-year period, we test and compare various models of the relationship between mating success and reproductive success (RS), and show that an accurate assessment of male and female reproductive behaviour requires consideration of all major components of mating success. Furthermore, we demonstrate that while Bateman's third principle holds when mating success is defined in terms of years married, women's fitness increases whereas men's fitness decreases from an increase in the number of marriage partners, holding constant the total effective duration of marriages. We discuss these findings in terms of the distinct, sex-specific pathways through which RS can be optimized, and comment on the contribution of this approach to the broader study of sexual selection. [ABSTRACT FROM AUTHOR]

Zorocrates guerrerensis Gertsch & Davis, 1940 is a Mexican cribellate spider with almost no information about its biology. As the species could potentially be of medical interest, it was decided to study basic aspects of its life history under laboratory conditions. The life cycle lasts a year, with spiders undergoing up to 12 instars to reach maturity. The courtship behaviour includes tactile communication. Copulation consists of two palpal insertions, each with a single haematodochal expansion. Both males and females can mate more than once; components of the first and second copulation do not differ. On average, the total copulation duration lasts for more than five minutes. Some details of the copulation process are discussed and compared with those of other lycosoids. [ABSTRACT FROM AUTHOR]

Abstract: Objective: The socioecological model (SEM) is a popular collection of controversial models purporting to explain mating systems in terms of ecological and social parameters. Despite its guise of objectivity, several of its hypotheses assume Victorian gender stereotypes of active, competing males heedlessly sowing their seeds, and cautious, passive females, imprisoned by greater costs of reproduction and their consequent resourceߚdependence. Methods: We enter this debate by taking a previously neglected explanatory approach borrowed from species theory. According to the Recognition Concept of sexual species, the unit of reproductive success/fitness is irreducible to fewer than two integrated subparts (minimally a male and a female). Phyletic changes in mating systems logically effect changes in fertilization systems, leading to reproductive isolation. We take our primary assumption of the average equivalence of female and male contributions to successful reproduction from the writings of the natural philosopher, Antoinette Blackwell. Results: We revisit the SEM with its contradictions and extrapolations, and develop a genderߚneutral alternative hypothesis termed SpecificߚMate Contact (SMC), centered on two fundamental mating strategies: sexual animals may behave as synchronous mateߚattractors or asynchronous mateߚseekers, generating four possible mating system combinations (monogamy: two attractors; promiscuity: two seekers; polygyny: male attractor and female seeker; polyandry: female attractor and male seeker). Conclusions: Our approach predicts all known primate mating systems using a neutral (nonߚsexist) principle. The approach is also neutral in the sense that it does not invoke either competition or cooperation: fertilization success is considered a posteriori and males and females are coߚadapted to this end rather than cognitively cooperative. [ABSTRACT FROM AUTHOR]

The longhorn beetle Cerambyx welensii is an emerging pest involved in oak decline episodes, whose damage is increasingly reported in dehesa open woodlands. Knowledge of the reproductive biology of C. welensii is a crucial goal due to its new pest status. In this study, we assess the reproductive traits of both sexes in the laboratory (25°C and 60% relative humidity ). In females, body length was 44.9 ± 0.9 mm (mean ± SE), fecundity 132 ± 12 eggs, fertility 70 ± 1 %, longevity 70 ± 3 days, preoviposition period 2 ± 0.2 days, oviposition period 44 ± 3 days and postoviposition period 19 ± 3 days. Fecundity was positively correlated with female size, longevity and oviposition period. Daily fecundity was 3.0 ± 0.2 eggs/day and showed a fluctuating synovigenic pattern with a slight decreasing trend over time. Egg length was 4.24 ± 0.01 mm and egg volume 8.14 ± 0.04 mm3. Egg size was correlated with female size but the relative size of eggs was larger in smaller females. Incubation time was 13.9 ± 0.1 days and hatching did not depend on egg size. Neonate size was positively correlated with egg length. Females were polyandrous (more than 20 lifetime matings) but multiple mating did not increase fecundity, fertility or longevity. In males, body length was 43.7 ± 0.6 mm and longevity 52 ± 3 days. Unlike with females, longevity was positively correlated with male size. Males were polygynous (up to 30 lifetime matings) but mating history did not affect male longevity. Rather to the contrary, long-lived males mated more times because they had more mating chances. Lastly, C. welensii reproductive traits were compared with those other Cerambycidae species and discussed from an adaptive perspective. Our data will be useful to improve management of C. welensii in order to prevent or mitigate its impact in dehesa woodlands and other oak forests. [ABSTRACT FROM AUTHOR]

Polyandry is more widespread than anticipated from Bateman’s principle but its ultimate (evolutionary) causes and proximate (mechanistic) correlates are more difficult to pinpoint than those of polygyny. Here, we combined mating experiments, quantification of reproductive traits and microsatellite genotyping to determine the fitness implications of polyandry in two predatory mite species, where males are highly polygynous (up to 45 fertilized females during life), whereas females range from monandry to various polyandry levels. The medium-level polyandrous (up to eight male mates possible) Neoseiulus californicus received clear direct and indirect benefits: multiply mated females produced more offspring with higher survival chances over longer times than singly mated females. In contrast, singly and multiply mated females of the low-level polyandrous (commonly two male mates at maximum) Phytoseiulus persimilis produced similar numbers of offspring having similar survival chances. In both species, multiple mating resulted in mixed offspring paternities, opening the chance for indirect fitness benefits such as enhanced genetic compatibility, complementarity and/or variability. However, the female re-mating likelihood and the paternity chance of non-first male mates were lower in P. persimilis than in N. californicus. Regarding proximate factors, in both species first mating duration and female re-mating likelihood were negatively correlated. Based on occasional fertilization failure of first male mates in P. persimilis, and mixed offspring paternities in both species, we argue that fertilization assurance and the chance to gain indirect fitness benefits are the ultimate drivers of polyandry in P. persimilis, whereas those of N. californicus are higher offspring numbers coupled with enhanced offspring viability and possibly other indirect fitness benefits. Overall, the adaptive significance and proximate events well reflected the polyandry levels. Our study provides a key example for linking behavioral experiments, quantification of reproductive traits and paternity analysis via offspring genotyping to explain the evolution of differing levels of polyandry. [ABSTRACT FROM AUTHOR]

Breeding strategies and parental effects play a major role in genetic selection programmes for quality trait improvement. The black-lip pearl oyster, Pinctada margaritifera, is a protandrous hermaphrodite species in which the number of mature females is low, and these only appear after 4 years of culture. This difference underlines the importance of studying the contributions of dams and sires in the production and selection of donor pearl oyster families to choose the most appropriate breeding strategies for cultured pearl quality trait improvement. Laboratory crosses were made using wild dams and sires to produce half-sib families from polyandry and polygyny mating designs. The resulting half-sib families were used as donor oysters in a large grafting experiment of 900 grafts with random recipient oysters, using nuclei of the same size and brand, the same operating technician and the same grow-out site. After 18 months of culture, 497 cultured pearls were harvested. Comparison between half-sib pairs revealed that for nacre thickness, weight and shape, a polygyny effect exists, whereas no significant effect was observed for the polyandry mating. By contrast, no evidence of dam- or sire-related benefits was found for the presence/absence of circles, darkness level and colour categories or grade and its components: lustre and surface defects. These results should have important implications for the selective breeding of donor pearl oysters for hatchery management. [ABSTRACT FROM AUTHOR]

The evolutionary trajectories of reproductive systems, including both male and female multiple mating and hence polygyny and polyandry, are expected to depend on the additive genetic variances and covariances in and among components of male reproductive success achieved through different reproductive tactics. However, genetic covariances among key components of male reproductive success have not been estimated in wild populations. We used comprehensive paternity data from socially monogamous but genetically polygynandrous song sparrows ( Melospiza melodia) to estimate additive genetic variance and covariance in the total number of offspring a male sired per year outside his social pairings (i.e. his total extra-pair reproductive success achieved through multiple mating) and his liability to sire offspring produced by his socially paired female (i.e. his success in defending within-pair paternity). Both components of male fitness showed nonzero additive genetic variance, and the estimated genetic covariance was positive, implying that males with high additive genetic value for extra-pair reproduction also have high additive genetic propensity to sire their socially paired female's offspring. There was consequently no evidence of a genetic or phenotypic trade-off between male within-pair paternity success and extra-pair reproductive success. Such positive genetic covariance might be expected to facilitate ongoing evolution of polygyny and could also shape the ongoing evolution of polyandry through indirect selection. [ABSTRACT FROM AUTHOR]

Pair bonding is a psychological construct that we attempt to operationalize via behavioral and physiological measurements. Yet, pair bonding has been both defined differently in various taxonomic groups as well as used loosely to describe not just a psychological and affective phenomenon, but also a social structure or mating system (either social monogamy or just pair living). In this review, we ask the questions: What has been the historical definition of a pair bond? Has this definition differed across taxonomic groups? What behavioral evidence do we see of pair bonding in these groups? Does this observed evidence alter the definition of pair bonding? Does the observed neurobiology underlying these behaviors affect this definition as well? And finally, what are the upcoming directions in which the study of pair bonding needs to head? [Display omitted] • Pair bonding has been defined differently across animal taxa. • We review these definitions as well as the occurrence of probable pair bonding behavior. • We revise the definition of pair bonding, with an emphasis on affective components. [ABSTRACT FROM AUTHOR]