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Kin selection plays a major role in the evolution of cooperative systems. However, many social species exhibit complex within-group relatedness structures, where kin selection alone cannot explain the occurrence of cooperative behaviour. Understanding such social structures is crucial to elucidate the evolution and maintenance of multi-layered cooperative societies. In lamprologine cichlids, intragroup relatedness seems to correlate positively with reproductive skew, suggesting that in this clade dominants tend to provide reproductive concessions to unrelated subordinates to secure their participation in brood care. We investigate how patterns of within-group relatedness covary with direct and indirect fitness benefits of cooperation in a highly social vertebrate, the cooperatively breeding, polygynous lamprologine cichlid Neolamprologus savoryi. Behavioural and genetic data from 43 groups containing 578 individuals show that groups are socially and genetically structured into subgroups. About 17% of group members were unrelated immigrants, and average relatedness between breeders and brood care helpers declined with helper age due to group membership dynamics. Hence the relative importance of direct and indirect fitness benefits of cooperation depends on helper age. Our findings highlight how both direct and indirect fitness benefits of cooperation and group membership can select for cooperative behaviour in societies comprising complex social and relatedness structures. This article is protected by copyright. All rights reserved.

Biological invasions are becoming more prevalent due to the rise of global trade and expansion of urban areas. Ants are among the most prolific invaders, with many exhibiting a multi-queen colony structure, dependent colony foundation and reduced inter-nest aggression. Although these characteristics are generally associated with the invasions of exotic ants, they may also facilitate the spread of native ants into novel habitats. Native to diverse habitats across North America, the odorous house ant Tapinoma sessile has become abundant in urban environments throughout the United States. Natural colonies typically have a small workforce, inhabit a single nest, and are headed by a single queen, whereas urban colonies tend to be several orders of magnitude larger, inhabit multiple nests (i.e., polydomy) and are headed by multiple queens (i.e., polygyny). Here, we explore and compare the population genetic and breeding structure of T. sessile within and between urban and natural environments in several localities across its distribution range. We found the social structure of a colony to be a plastic trait in both habitats, although extreme polygyny was confined to urban habitats. Additionally, polydomous colonies were only present in urban habitats, suggesting T. sessile can only achieve supercoloniality within urbanized areas. Finally, we identified strong differentiation between urban and natural populations in each locality and continent-wide, indicating cities may restrict gene flow and exert intense selection pressure. Overall, our study highlights urbanization's influence in charting the evolutionary course for species.

Mating systems and patterns of reproductive success in fishes play an important role in ecology and evolution. While information on the reproductive ecology of many anadromous salmonids (Oncorhynchus spp.) is well detailed, there is less information for nonanadromous species including the Yellowstone Cutthroat Trout (O. clarkii bouvieri), a subspecies of recreational angling importance and conservation concern. Using data from a parentage‐based tagging study, we described the genetic mating system of a migratory population of Yellowstone Cutthroat Trout, tested for evidence of sexual selection, and identified predictors of mating and reproductive success. The standardized variance in mating success (i.e., opportunity for sexual selection) was significantly greater for males relative to females, and while the relationship between mating success and reproductive success (i.e., Bateman gradient) was significantly positive for both sexes, a greater proportion of reproductive success was explained by mating success for males (r 2 = 0.80) than females (r 2 = 0.59). Overall, the population displayed a polygynandrous mating system, whereby both sexes experienced variation in mating success due to multiple mating, and sexual selection was variable across sexes. Tests for evidence of sexual selection indicated the interaction between mating success and total length best‐predicted relative reproductive success. We failed to detect a signal of inbreeding avoidance among breeding adults, but the group of parents that produced progeny were on average slightly less related than adults that did not produce progeny. Lastly, we estimated the effective number of breeders (N b) and effective population size (N e) and identified while N b was lower than N e, both are sufficiently high to suggest Yellowstone Cutthroat Trout in Burns Creek represent a genetically stable and diverse population., We collected genetic samples from over 1,500 adult Yellowstone Cutthroat Trout as they entered a spawning tributary and subsequently collected ~3,000 genetic samples from outmigrating juveniles from the same creek system. Using a combination of sibship analysis and parentage‐based tagging, we identified a polygynandrous mating system, whereby both sexes experienced variation in mating success due to multiple mating, and sexual selection was variable across sexes. These data have direct implications for ongoing management and conservation efforts.

In long‐lived polygynous species, male reproductive success is often monopolized by a few mature dominant individuals. Young males are generally too small to be dominant and may employ alternative tactics; however, little is known about the determinants of reproductive success for young males. Understanding the causes and consequences of variability in early reproductive success may be crucial to assess the strength of sexual selection and possible long‐term trade‐offs among life‐history traits. Selective pressures driven by fluctuating environmental conditions may depend on age class. We evaluated the determinants of reproduction in male bighorn sheep (Ovis canadensis) aged 2–4 years using 30 years of individual‐level data. These young males cannot defend estrous ewes and use alternative mating tactics. We also investigated how the age of first detected reproduction was correlated to lifetime reproductive success and longevity. We found that reproductive success of males aged 3 years was positively correlated to body mass, to the proportion of males aged 2–4 years in the competitor pool, and to the number of females available per adult male. These results suggest that reproductive success depends on both competitive ability and population age–sex structure. None of these variables, however, had significant effects on the reproductive success of males aged 2 or 4 years. Known reproduction before the age of five increased lifetime reproductive success but decreased longevity, suggesting a long‐term survival cost of early reproduction. Our analyses reveal that both individual‐level phenotypic and population‐level demographic variables influence reproductive success by young males and provide a rare assessment of fitness trade‐offs in wild polygynous males., Little is known about the determinants of reproductive success for young polygynous male mammals, although the causes and consequences of variability in early reproductive success may be crucial to assess the strength of sexual selection and possible long‐term trade‐offs among life‐history traits. We evaluated the determinants of reproduction in male bighorn sheep (Ovis canadensis) aged 2–4 years using 30 years of individual‐level data and investigated how the age of first reproduction was correlated to lifetime reproductive success and longevity. Early reproductive success was influenced by phenotypical and demographical variables, and males that first reproduced at early ages had higher lifetime reproductive success and shorter lifespan, suggesting a long‐term survival cost of early reproduction.

The fire ant Solenopsis invicta exists in two alternate social forms: monogyne nests contain a single reproductive queen and polygyne nests contain multiple reproductive queens. This colony-level social polymorphism corresponds with individual differences in queen physiology, queen dispersal patterns and worker discrimination behaviours, all evidently regulated by an inversion-based supergene that spans more than 13 Mb of a "social chromosome," contains over 400 protein-coding genes and rarely undergoes recombination. The specific mechanisms by which this supergene influences expression of the many distinctive features that characterize the alternate forms remain almost wholly unknown. To advance our understanding of these mechanisms, we explore the effects of social chromosome genotype and natal colony social form on gene expression in queens sampled as they embarked on nuptial flights, using RNA-sequencing of brains and ovaries. We observe a large effect of natal social form, that is, of the social/developmental environment, on gene expression profiles, with similarly substantial effects of genotype, including: (a) supergene-associated gene upregulation, (b) allele-specific expression and (c) pronounced extra-supergene trans-regulatory effects. These findings, along with observed spatial variation in differential and allele-specific expression within the supergene region, highlight the complex gene regulatory landscape that emerged following divergence of the inversion-mediated Sb haplotype from its homologue, which presumably largely retained the ancestral gene order. The distinctive supergene-associated gene expression trajectories we document at the onset of a queen's reproductive life expand the known record of relevant molecular correlates of a complex social polymorphism and point to putative genetic factors underpinning the alternate social syndromes.

Ungulates exhibit diverse mating systems that range from monogamous pair territories to highly polygynous leks. We review mating systems and behaviors across ungulates and offer a new approach synthesizing how interacting factors may shape those mating systems. Variability exists in mating systems among and within species of ungulates and likely is affected by predation risk, availability of resources (food and mates), habitat structure, and sociality. Ungulate mating systems may be labile as a consequence of the varying strength of those interacting factors. In addition, degree of polygyny and sexual dimorphism in size are associated with the evolution of mating systems. Neither male–male combat nor paternal care, however, can completely explain differences in sexual size dimorphism for ungulates, a necessary component in understanding the development of some mating systems. Whatever the evolutionary pathway, sexual segregation limits paternal care allowing more intense male–male competition. Selection of habitat structure, because it modifies risk of predation, is a major determinant of sociality for ungulates. Likewise, ruggedness and steepness of terrain limit the types of mating systems that can occur because of limitations in group size and cohesiveness, as well as the ability of males to herd even small groups of females effectively. The quality and defensibility of resources affect mating systems, as does the defensibility of females. Population density of females also may be a critical determinant of the types of mating systems that develop. Size of groups likewise constrains the types of mating tactics that males can employ. Our aim was to use those relationships to create a broad conceptual model that predicts how various environmental and social factors interact to structure mating systems in ungulates. This model provides a useful framework for future tests of the roles of both ecological and social conditions in influencing the social systems of ungulates., We provide a new approach to understanding mating systems in ungulates. We integrate both social and environmental factors in doing so. We develop a predictive model to help understand the evolution of these unique behaviors, which should help to direct future studies.

In most animals, females are larger than males. Paradoxically, sexual size dimorphism is biased towards males in most mammalian species. An accepted explanation is that sexual dimorphism in mammals evolved by intramale sexual selection. I tested this hypothesis in primates, by relating sexual size dimorphism to seven proxies of sexual selection intensity: operational sex ratio, mating system, intermale competition, group sex ratio, group size, maximum mating percentage (percentage of observed copulations involving the most successful male), and total paternity (a genetic estimate of the percentage of young sired by the most successful male). I fitted phylogenetic generalised least squares models using sexual size dimorphism as the dependent variable and each of the seven measures of intensity of sexual selection as independent variables. I conducted this comparative analysis with data from 50 extant species of primates, including Homo sapiens, Pan troglodytes, and Gorilla spp. Sexual dimorphism was positively related to the four measures of female monopolisation (operational sex ratio, mating system, intermale competition, and group sex ratio) and in some cases to group size, but was not associated with maximum mating percentage or total paternity. Additional regression analyses indicated that maximum mating percentage and total paternity were negatively associated with group size. These results are predicted by reproductive skew theory: in large groups, males can lose control of the sexual behaviour of the other members of the group or can concede reproductive opportunities to others. The results are also consistent with the evolution of sexual size dimorphism before polygyny, due to the effects of natural, rather than sexual, selection. In birds, the study of molecular paternity showed that variance in male reproductive success is much higher than expected by behaviour. In mammals, recent studies have begun to show the opposite trend, i.e. that intensity of sexual selection is lower than expected by polygyny. Results of this comparative analysis of sexual size dimorphism and sexual selection intensity in primates suggest that the use of intramale sexual selection theory to explain the evolution of polygyny and sexual dimorphism in mammals should be reviewed, and that natural selection should be considered alongside sexual selection as an evolutionary driver of sexual size dimorphism and polygyny in mammals.

Multiple mating by females is common and often driven by social constraints on female mate choice. However, females mate with multiple males even in systems without these social constraints and rates of multiple mating tend to be highly variable within and between populations. In lek mating systems, females are able to assess multiple males and their choice is unrestricted by pair bonds or the need for biparental care, yet some females mate with multiple males. To better understand the factors affecting variation in multiple mating, we investigated the occurrence of multiple paternity within clutches in a highly polygynous lek mating system. Using long-term data on genetic paternity, survival, social status and individual age from a population of lance-tailed manakins Chiroxiphia lanceolata, a species where males lek in cooperative alpha-beta pairs, we tested five non-exclusive hypotheses about the causes of variation in multiple mating and its benefits in females. We found that inexperienced males, including new alphas and rare beta sires, were disproportionately likely to share paternity when they sired any chicks. In contrast, female age (experience) was unrelated to multiple paternity. Multiple mating did not result in higher reproductive success or reduced variance in success for females, and there were neither consistently promiscuous females nor males that consistently shared paternity. The occurrence of multiple paternity in this lek mating system was best explained by female choice related to male characteristics that change with male experience. Our results support the hypothesis that there is a developmental component to the occurrence of multiple mating, and suggest females choose to mate multiply when their choices are not optimal.

A retractable larynx and adaptations of the vocal folds in the males of several polygynous ruminants serve for the production of rutting calls that acoustically announce larger than actual body size to both rival males and potential female mates. Here, such features of the vocal tract and of the sound source are documented in another species. We investigated the vocal anatomy and laryngeal mobility including its acoustical effects during the rutting vocal display of free‐ranging male impala (Aepyceros melampus melampus) in Namibia. Male impala produced bouts of rutting calls (consisting of oral roars and interspersed explosive nasal snorts) in a low‐stretch posture while guarding a rutting territory or harem. For the duration of the roars, male impala retracted the larynx from its high resting position to a low mid‐neck position involving an extensible pharynx and a resilient connection between the hyoid apparatus and the larynx. Maximal larynx retraction was 108 mm based on estimates in video single frames. This was in good concordance with 91‐mm vocal tract elongation calculated on the basis of differences in formant dispersion between roar portions produced with the larynx still ascended and those produced with maximally retracted larynx. Judged by their morphological traits, the larynx‐retracting muscles of male impala are homologous to those of other larynx‐retracting ruminants. In contrast, the large and massive vocal keels are evolutionary novelties arising by fusion and linear arrangement of the arytenoid cartilage and the canonical vocal fold. These bulky and histologically complex vocal keels produced a low fundamental frequency of 50 Hz. Impala is another ruminant species in which the males are capable of larynx retraction. In addition, male impala vocal folds are spectacularly specialized compared with domestic bovids, allowing the production of impressive, low‐frequency roaring vocalizations as a significant part of their rutting behaviour. Our study expands knowledge on the evolutionary variation of vocal fold morphology in mammals, suggesting that the structure of the mammalian sound source is not always human‐like and should be considered in acoustic analysis and modelling., Male impala retract the larynx while producing their impressive rutting roars. The low fundamental frequency of these roars is produced by a highly modified vocal fold, a ‘vocal keel’. The anatomical and acoustical correlates of male impala's mobile larynx are discussed.

Fil: Cassini, Marcelo Hernan. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Instituto de Biologia y Medicina Experimental. Fundacion de Instituto de Biologia y Medicina Experimental. Instituto de Biologia y Medicina Experimental; Argentina

Women make important contributions to household food production in sub‐Saharan Africa. Women's agricultural productivity is often reduced, however, by inefficient intrahousehold allocation of agricultural resources. Complex marital structures found in polygynous households may complicate resource allocation. Using three waves of the Tanzania Living Standards Measurement Survey–Integrated Survey on Agriculture, we measure the effect of the marital structure, wife position, and plot management on agricultural productivity and input allocations. We find evidence of cooperation within polygynous households. Plots managed by husbands and wives in polygynous households produce more valuable crops, have higher yields, and are more likely to use fertilizer than their monogamous counterparts. Within polygynous households, we observe that plots jointly managed by husbands, first wives, and second wives (together) have significantly more family labor than plots managed by husbands and first wives. This result may provide evidence of different production technologies across plot managers within the same households.

Dispersal is ubiquitous throughout the tree of life: factors selecting for dispersal include kin competition, inbreeding avoidance and spatiotemporal variation in resources or habitat suitability. These factors differ in whether they promote male and female dispersal equally strongly, and often selection on dispersal of one sex depends on how much the other disperses. For example, for inbreeding avoidance it can be sufficient that one sex disperses away from the natal site. Attempts to understand sex‐specific dispersal evolution have created a rich body of theoretical literature, which we review here. We highlight an interesting gap between empirical and theoretical literature. The former associates different patterns of sex‐biased dispersal with mating systems, such as female‐biased dispersal in monogamous birds and male‐biased dispersal in polygynous mammals. The predominant explanation is traceable back to Greenwood's (1980) ideas of how successful philopatric or dispersing individuals are at gaining mates or the resources required to attract them. Theory, however, has developed surprisingly independently of these ideas: models typically track how immigration and emigration change relatedness patterns and alter competition for limiting resources. The limiting resources are often considered sexually distinct, with breeding sites and fertilizable females limiting reproductive success for females and males, respectively. We show that the link between mating system and sex‐biased dispersal is far from resolved: there are studies showing that mating systems matter, but the oft‐stated association between polygyny and male‐biased dispersal is not a straightforward theoretical expectation. Here, an important understudied factor is the extent to which movement is interpretable as an extension of mate‐searching (e.g. are matings possible en route or do they only happen after settling in new habitat – or can females perhaps move with stored sperm). We also point out other new directions for bridging the gap between empirical and theoretical studies: there is a need to build Greenwood's influential yet verbal explanation into formal models, which also includes the possibility that an individual benefits from mobility as it leads to fitness gains in more than one final breeding location (a possibility not present in models with a very rigid deme structure). The order of life‐cycle events is likewise important, as this impacts whether a departing individual leaves behind important resources for its female or male kin, or perhaps both, in the case of partially overlapping resource use.

The recent advances of new genomic technologies have enabled the identification and characterization of sex chromosomes in an increasing number of nonmodel species, revealing that many plants and animals undergo frequent sex chromosome turnovers. What evolutionary forces drive these turnovers remains poorly understood, but it was recently proposed that drift might play a more important role than generally assumed. We analysed the dynamics of different types of turnovers using individual-based simulations and show that when mediated by genetic drift, turnovers are usually easier to achieve than substitutions at neutral markers, but that their dynamics and relative likelihoods vary with the type of the resident and emergent sex chromosome system (XY and/or ZW) and the dominance relationships among the sex-determining factors. Focusing on turnovers driven by epistatically dominant mutations, we find that drift-mediated turnovers that preserve the heterogamety pattern are 2-4× more likely than those along which the heterogametic sex changes. This ratio nevertheless decreases along with effective population size and can even reverse in case of extreme polygyny. This can be attributed to a 'drift-induced' selective force, known to influence transitions between male and female heterogamety, but which according to our study does not affect turnovers that preserve the heterogametic sex.

Mate searching is assumed to be performed mostly by males, but when females benefit from multiple mating or are under risk of failing to mate, they may also perform mate searching. This is especially important in scramble competition polygynies, in which mate searching is the main mechanism of mate competition. Typically, more mobile individuals are expected to achieve higher mating success because mobility increases their probability of finding mates. If we assume individual movements are mainly explained by mate searching in scramble competition polygynies, we can investigate searching strategies by asking when individuals should leave their location and where they should go. We hypothesize that individuals will leave their locations when mating opportunities are scarce and will seek spatially close sites with better mating opportunities. We tested these hypotheses for males and females of Leptinotarsa undecimlineata, a leaf beetle with scramble competition polygyny in which both sexes are promiscuous. Individuals mate and feed exclusively on Solanum plants, and thus, individual movements can be described as switches between plants. Females were less likely than males to leave isolated plants, and both males and females moved preferentially to neighboring plants. Males were more likely to leave when the local number of females was low, and the number of males was high. They moved to plants with more females, a behavior consistent with a mate searching strategy. Females were more likely to move to plants with fewer males and many females, a behavior consistent with male harassment avoidance. Strategic movement is widely considered in foraging context, but seldom in a mate searching context. Considering that selection to minimize searching costs, maximize mating success, and minimize harassment may be ubiquitous in nature, we argue that strategic movements by mate searching individuals are likely to occur in many species.

Effective population size, a central concept in conservation biology, is now routinely estimated from genetic surveys and can also be theoretically predicted from demographic, life-history, and mating-system data. By evaluating the consistency of theoretical predictions with empirically estimated effective size, insights can be gained regarding life-history characteristics and the relative impact of different life-history traits on genetic drift. These insights can be used to design and inform management strategies aimed at increasing effective population size. We demonstrated this approach by addressing the conservation of a reintroduced population of Asiatic wild ass (Equus hemionus). We estimated the variance effective size (Nev ) from genetic data (N ev =24.3) and formulated predictions for the impacts on Nev of demography, polygyny, female variance in lifetime reproductive success (RS), and heritability of female RS. By contrasting the genetic estimation with theoretical predictions, we found that polygyny was the strongest factor affecting genetic drift because only when accounting for polygyny were predictions consistent with the genetically measured Nev . The comparison of effective-size estimation and predictions indicated that 10.6% of the males mated per generation when heritability of female RS was unaccounted for (polygyny responsible for 81% decrease in Nev ) and 19.5% mated when female RS was accounted for (polygyny responsible for 67% decrease in Nev ). Heritability of female RS also affected Nev ; hf2=0.91 (heritability responsible for 41% decrease in Nev ). The low effective size is of concern, and we suggest that management actions focus on factors identified as strongly affecting Nev, namely, increasing the availability of artificial water sources to increase number of dominant males contributing to the gene pool. This approach, evaluating life-history hypotheses in light of their impact on effective population size, and contrasting predictions with genetic measurements, is a general, applicable strategy that can be used to inform conservation practice.

The Acacia drepanolobium (also known as Vachellia drepanolobium) ant‐plant symbiosis is considered a classic case of species coexistence, in which four species of tree‐defending ants compete for nesting space in a single host tree species. Coexistence in this system has been explained by trade‐offs in the ability of the ant associates to compete with each other for occupied trees versus the ability to colonize unoccupied trees. We seek to understand the proximal reasons for how and why the ant species vary in competitive or colonizing abilities, which are largely unknown. In this study, we use RADseq‐derived SNPs to identify relatedness of workers in colonies to test the hypothesis that competitively dominant ants reach large colony sizes due to polygyny, that is, the presence of multiple egg‐laying queens in a single colony. We find that variation in polygyny is not associated with competitive ability; in fact, the most dominant species, unexpectedly, showed little evidence of polygyny. We also use these markers to investigate variation in mating behavior among the ant species and find that different species vary in the number of males fathering the offspring of each colony. Finally, we show that the nature of polygyny varies between the two commonly polygynous species, Crematogaster mimosae and Tetraponera penzigi: in C. mimosae, queens in the same colony are often related, while this is not the case for T. penzigi. These results shed light on factors influencing the evolution of species coexistence in an ant‐plant mutualism, as well as demonstrating the effectiveness of RADseq‐derived SNPs for parentage analysis.

The polygyny threshold (PT) is a critical transition point in the sexual selection process for many organisms in natural populations, characterizing when females choose to mate with an already mated male over an unmated one to improve fitness. Understanding its causes and consequences is therefore of high interest. While both theoretical and empirical work suggest that the degree of polygyny within a species is plastic and a function of male inequality, the functional relationship between underlying availability of resources occupied by breeding males under variable climatic conditions and the dynamics of PTs across space and time has received less attention. Here, we use a standardized measure of male mating inequality as the culmination of female mate choices to analyse how spatially dynamic PTs in a naturally regulated feral horse (Equus ferus caballus) population emerge along a geographic gradient in a known, limiting resource (freshwater) each year from variable climatic conditions. Polygyny threshold distance from permanent freshwater increased with increasing precipitation during the breeding season of each year, suggesting a relationship between annual resource availability and female mate choice. The mechanism by which climatic conditions underpin the spatial dynamics of PTs was likely through precipitation providing ephemeral freshwater sources across the study area that effectively weakened the gradient in availability of permanent freshwater, thereby providing mating males that occupied home ranges far from permanent water with access to this limiting resource and enabling them to attract and retain females. Increased precipitation also coincided with a decreased proportion of males in the population that experienced sexual selection pressure attributed to female mate choice in relation to the acquisition and/or defence of freshwater sources. Climatic conditions caused spatial shifts in PTs annually along the geographic gradient in resource availability. Our findings reveal that such environmental gradients may either buffer or amplify impacts of climatic variation on selection pressure operating in natural populations, and emphasize the importance of integrating spatially explicit PTs with atmospheric fluctuations when predicting the effect of climatic change on selection processes within populations that occupy environmental gradients.

Summary 1.In mammals, maternal expenditure on offspring is a complex mix of several factors including the species’ mating system, offspring sex and the condition and age of the mother. While theory suggests that in polygynous species mothers should wean larger male offspring than females when resources and maternal conditions allow, the evidence for this remains equivocal. 2.Southern elephant seals are highly dimorphic, polygynous capital breeders existing in an environment with highly variable resources and should therefore provide clear evidence to support the theoretical expectations of differential maternal expenditure in male and female pups. 3.We quantified maternal size (mass and length) and pup size at birth and weaning for 342 elephant seal mothers at Macquarie Island. The study was conducted over 11 years of contrasting sea-ice and Southern Annular Mode values, both indices of maternal prey resources. 4.Overall, large females weaned male pups that weighed 17 kg (15.5%) more than female pups. Maternal condition varied by as much as 59 kg among years, and was positively related to SAM, and negatively to maximum sea-ice extent. Smaller mothers weaned relatively larger male pups under favourable conditions, this effect was less apparent for larger mothers. 5.We developed a simple model linking environmental variation to maternal masses post-partum, followed by maternal masses post-partum to weaning masses and then weaning masses to pup survival and demonstrated that environmental conditions affected predicted survival so that the pups of small mothers had an estimated 7% increase in first year survival in “good” vs “bad” years compared to 1% for female pups of large mothers. 6.Co-occurrence of environmental quality and conservative reproductive tactics suggests that mothers retain substantial plasticity in maternal care, enhancing their lifetime reproductive success by adjusting reproductive expenditure relative to both prevailing environmental conditions and their own capabilities. This article is protected by copyright. All rights reserved.

The ponerine ant Brachyponera chinensis was introduced to the USA, where it has become invasive. Although various ecological data have been collected for B. chinensis populations in the USA, most aspects concerning the biology and ecology of native populations in Japan, a presumed origin, remain unknown. Here we investigated the social structure and nestmate discrimination in native populations of B. chinensis and a closely related species, B. nakasujii. Both species showed functional polygyny over seasons. Only in B. nakasujii was there a seasonal change in the numbers of queens and workers per nest. In arena tests, workers of neither species showed aggressive behaviors to conspecific non-nestmates from the same population, and the mean aggression score did not increase with the distance between nests. However, some differences in non-aggressive responses were detected between nestmate and non-nestmate pairs in both species. In an experiment to introduce a single worker into a nest, B. chinensis accepted non-nestmates with a high probability just like nestmates, whereas in B. nakasujii non-nestmates were less accepted than nestmates. These findings suggest that native populations of B. chinensis already possess some of the key characteristics shared by many invasive exotic ants in introduced ranges, such as stable polygyny, weak internest aggression and acceptance of non-nestmates. These tendencies are remarkable in comparison to the closely related B. nakasujii.

According to the Trivers-Willard hypothesis, females of polygynous mammal species can adaptively modify offspring sex ratio, which would provide them evolutionary benefits in terms of a higher number of grand-offspring. In red deer (Cervus elaphus), a species where the reproductive success of males is significantly more variable than that of females, mothers that are in good condition are expected to have a male-biased offspring sex ratio but only in a population below carrying capacity. We aimed at testing the Trivers-Willard hypothesis on a low-density population of red deer in large woodlands of north-eastern Poland. The overall foetal sex ratio did not differ from parity. The mother's probability of having male offspring increased with her body mass independently of age and population density. Such condition-based adjustment of the offspring sex ratio in female red deer from a low-density population conforms to the Trivers–Willard hypothesis. However, full support of Trivers–Willard predictions would require further investigations comparing survival and reproductive output of male and female offspring.

The use of alternative reproductive tactics (ARTs) is widespread in animals. Males of some species may change tactics depending on age, body condition and social environment. Many bat species are polygynous where a fraction of males only have access to fertile females. For polygynous bats, knowledge of the reproductive success of males using different ARTs is scarce, and it remains unclear how age of males is related to switching decisions between social statuses. We studied a large captive population of Carollia perspicillata, where males are either harem holders, bachelors or peripheral males. Using a multistate procedure, we modelled the age-related switches in reproductive tactics and in survival probability. From the model, we calculated the reproductive success and the frequencies of males displaying different reproductive tactics. As in mammals, the switch between social statuses is often related to age, we predicted that the transition probability of bachelor and peripheral males to harem status would increase with age. We show, however, that social status transition towards a harem holding position was not related to age. Reproductive success changed with age and social status. Harem males had a significantly higher reproductive success than bachelor males except between a short period from 3.8 to 4.4 years of age where success was similar, and a significantly higher reproductive success than peripheral males between 2.6 and 4.4 years of age. Harem males showed a clear decrease in the probability of maintaining social status with age, which suggests that senescence reduces resource holding potential.

Despite being regularly bred in zoos, giraffes remain a challenge, especially in terms of feeding. Assessment of factors influencing growth and weight changes during ontogeny, as well as analysis of weight fluctuations in adult individuals, may become a critical point in captive diet evaluation. Knowledge about weight is a crucial husbandry tool; however, such data are rarely acquired. Using a unique dataset from regularly weighed Rothschild giraffes (Giraffa camelopardalis rothschildi) from Prague zoo, we determined the growth functions of male and female giraffes and calculated weight gains during giraffe ontogeny. The mean weights of adult males and females were 1307 ± 52 and 835 ± 45 kg, respectively confirming the large overall dimensions of G. c. rothschildi in comparison with other giraffe subspecies. As the giraffe is a polygynous species showing considerable sexual dimorphism, we expected male calves to have larger first weights and faster growth during the most intensive period of maternal care. Growth rates and daily weight gains were higher in males than in females during the whole postnatal period. Males grew faster and longer than females. However, differences in weight between males and females appeared as late as after 1 year of age. The weight of adult males and non-pregnant adult females fluctuated significantly across seasons, being the highest during the autumn and winter months, respectively which may reflect the different effects of sexual activity and feeding ratios in males and females. Zoo Biol. 35:423-431, 2016. © Wiley Periodicals, Inc.

In large multimale-multifemale primate groups, individual adult males and females may form close social relationships that extend beyond the mating context, a surprising finding for polygynandrous mammals. The patterns of these associations can be relatively stable across time. Here we investigate whether dyadic social relationships between the sexes transcend mere association in wild Assamese macaques and may be characterized as strong, equitable, and stable affiliative relationships or social bonds. We collected >9,000 hr of focal animal data on adult males and females from two groups of wild Assamese macaques (Macaca assamensis) at Phu Khieo Wildlife Sanctuary, Thailand. Using dyadic composite sociality indices, we found male-female affiliative relationships to be highly differentiated. The stronger the relationships were, the more likely partners were to reciprocate grooming and the more stable were the relationships. In addition, the strongest dyadic relationships remained stable over multiple years as long as both partners remained in the group. These results indicate that in a polygynous species particular males and females form strong, equitable, and enduring affiliative relationships qualitatively similar to the same-sex bonds described for female baboons and male chimpanzees. Am. J. Primatol. 78:872-882, 2016. © 2016 Wiley Periodicals, Inc.

Lifetime reproductive success of males is often dependent upon the ability to physically compete for mates. However, species variation in social structure leads to differences in the relative importance of intraspecific aggression. Here, we present a large comparative dataset on sexual dimorphism in skeletal shape in Carnivora to test the hypotheses that carnivorans exhibit sexual dimorphism in skeletal anatomy that is reflective of greater specialization for physical aggression in males relative to females and that this dimorphism is associated with the intensity of sexual selection. We tested these hypotheses using a set of functional indices predicted to improve aggressive performance. Our results indicate that skeletal shape dimorphism is widespread within our sample. Functional traits thought to enhance aggressive performance are more pronounced in males. Phylogenetic model selection suggests that the evolution of this dimorphism is driven by sexual selection, with the best-fitting model indicating greater dimorphism in polygynous versus nonpolygynous species. Skeletal shape dimorphism is correlated with body size dimorphism, a common indicator of the intensity of male-male competition, but not with mean body size. These results represent the first evidence of sexual dimorphism in the primary locomotor system of a large sample of mammals.

Parents should bias sex allocation toward offspring of the sex most likely to provide higher fitness returns. Trivers and Willard proposed that for polygynous mammals, females should adjust sex-ratio at conception or bias allocation of resources toward the most profitable sex, according to their own body condition. However, the possibility that mammalian fathers may influence sex allocation has seldom been considered. Here, we show that the probability of having a son increased from 0.31 to 0.60 with sire reproductive success in wild bighorn sheep (Ovis canadensis). Furthermore, our results suggest that females fertilized by relatively unsuccessful sires allocated more energy during lactation to daughters than to sons, while the opposite occurred for females fertilized by successful sires. The pattern of sex-biased offspring production appears adaptive because paternal reproductive success reduced the fitness of daughters and increased the average annual weaning success of sons, independently of maternal allocation to the offspring. Our results illustrate that sex allocation can be driven by paternal phenotype, with profound influences on the strength of sexual selection and on conflicts of interest between parents.

The partitioning of reproduction among individuals in communally breeding animals varies greatly among species, from the monopolization of reproduction (high reproductive skew) to similar contribution to the offspring in others (low skew). Reproductive skew models explain how relatedness or ecological constraints affect the magnitude of reproductive skew. They typically assume that individuals are capable of flexibly reacting to social and environmental changes. Most models predict a decrease of skew when benefits of staying in the group are reduced. In the ant Leptothorax acervorum, queens in colonies from marginal habitats form dominance hierarchies and only the top‐ranking queen lays eggs (“functional monogyny”). In contrast, queens in colonies from extended coniferous forests throughout the Palaearctic rarely interact aggressively and all lay eggs (“polygyny”). An experimental increase of queen:worker ratios in colonies from low‐skew populations elicits queen–queen aggression similar to that in functionally monogynous populations. Here, we show that this manipulation also results in increased reproductive inequalities among queens. Queens from natural overwintering colonies differed in the number of developing oocytes in their ovaries. These differences were greatly augmented in queens from colonies with increased queen:worker ratios relative to colonies with a low queen:worker ratio. As assumed by models of reproductive skew, L. acervorum colonies thus appear to be capable of flexibly adjusting reproductive skew to social conditions, yet in the opposite way than predicted by most models.

In most mammals, both sexes display different survival patterns, often involving faster senescence in males. Being under intense sexual competition to secure mating opportunities, males of polygynous species allocate resources to costly behaviors and conspicuous sexual traits, which might explain these observed differences in longevity and senescence patterns. However, comparative studies performed to date have led to conflicting results. We aimed to resolve this problem by first reviewing case studies of the relationship between the strength of sexual selection and age-specific survival metrics. Then, we performed a comprehensive comparative analysis to test whether such relationships exist among species of captive ruminants. We found that the strength of sexual selection negatively influenced the onset of actuarial senescence in males, with males senescing earlier in polygynous than in monogamous species, which led to reduced male longevity in polygynous species. Moreover, males of territorial species senesced earlier but slower, and have a shorter longevity than males of species displaying other mating tactics. We detected little influence of the strength of sexual selection on the rate of actuarial senescence. Our findings demonstrate that the onset of actuarial senescence, rather than its rate, is a side effect of physiological mechanisms linked to sexual selection, and potentially accounts for observed differences in longevity.

Objectives Genealogies contain information on the prevalence of different sibling types that result from past reproductive behavior. Full sibling sets stem from stable monogamy, paternal half siblings primarily indicate male reproductive skew, and maternal half siblings reflect unstable pair bonds. Methods Full and half sibling types are calculated for a total of 61,181 siblings from published genealogies for 80 small-scale societies, including foragers, horticulturalists, agriculturalists, and pastoralists from around the world. Results Most siblings are full (61%) followed by paternal half siblings (27%) and maternal half siblings (13%). Paternal half siblings are positively correlated with more polygynous marriages, higher at low latitudes, and slightly higher in nonforagers, Maternal half sibling fractions are slightly higher at low latitudes but do not vary with subsistence. Partible paternity societies in Amazonia have more paternal half siblings indicating higher male reproductive skew. Conclusions Sibling counts from genealogies provide a convenient method to simultaneously investigate the reproductive skew and pair-bond stability dimensions of human mating systems cross-culturally. Am. J. Hum. Biol., 2015. © 2015 Wiley Periodicals, Inc.

Shiny Cowbirds Molothrus bonariensis and Screaming Cowbirds Molothrus rufoaxillaris are closely related brood parasites but the former is socially polygynous or promiscuous and an extreme host generalist, whereas the latter is socially monogamous and parasitizes almost exclusively one host. Females of both species lay in relative darkness, before dawn, relying for host nest location on previous days’ prospecting activity, or possibly on following better-informed roost associates. We studied the temporal and spatial patterns of roosting behaviour in these species to test the hypothesis that roosting behaviour of cowbirds is related to their breeding strategy (brood parasitism) and reflects differences in strategies between species. We recorded fidelity to a roost, location fidelity within a roost, inter-individual spatial associations and timing of roost departures and parasitic events, using tagged individuals. Female Shiny Cowbirds and both sexes of Screaming Cowbirds showed marked fidelity in roosting location, and roost departures occurred both during and after the known time window for parasitism, with earlier departures probably corresponding to laying days. Screaming Cowbird females and males that were trapped together and showed high levels of association during the day, also showed high levels of association in the roost. We describe the spatial and temporal patterns of a relatively poorly known aspect of avian ecology in general and the behaviour of brood parasites in particular. Fil: Scardamaglia, Romina Clara. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Ecología, Genética y Evolución de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Ecología, Genética y Evolución de Buenos Aires; Argentina Fil: Kacelnik, Alex. University of Oxford; Reino Unido Fil: Reboreda, Juan Carlos. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Ecología, Genética y Evolución de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Ecología, Genética y Evolución de Buenos Aires; Argentina

Polygyny in social insects can greatly reduce within‐nest genetic relatedness. In polygynous ant species, potential rival queens in colonies with multiple queens are often executed by other queens, workers, or both. The Argentine ant, Linepithema humile, native to South America, forms a “supercolony” that is composed of a large number of nests and is considered to contribute to the ant's invasion success. Currently, four mutually antagonistic supercolonies are contiguously distributed within a small area of Japan. Here, we analyzed the genetic structure and relatedness within and among the four supercolonies using microsatellite markers to clarify how L. humile maintains its supercoloniality. The results of AMOVA and BASP, the FST values, and the existence of several private alleles indicated that the L. humile population in the Kobe area had a characteristic genetic structure. Within a given supercolony, there was significant genetic differentiation (FST) among workers collected in May and those collected in September. The significant deviation from Hardy–Weinberg equilibrium increased, and the relatedness among workers significantly increased from May to September in all supercolonies. This result suggested that the supercolonies replaced old queens with new ones during the reproductive season, thus supporting the plausibility of queen execution. From the perspective of kin selection, workers collectively eliminate queens, thereby increasing their own inclusive fitness. Restricted gene flow among supercolonies, together with mating with sib and queen execution, could help to maintain the unique social structure of L. humile, the distribution of which is expanding worldwide.

Objectives One-male social systems are usually characterized by polygyny and reproductive exclusion by a single resident male. Sometimes, however, secondary males join these groups, and this may carry fitness costs and/or benefits to the resident male. In hamadryas baboons (Papio hamadryas hamadryas), which live in one-male units (OMUs) with female defense polygyny within a multi-level social system, secondary “follower” males often reside in OMUs. Our aim here is to examine possible benefits of these secondary males to hamadryas resident males. Materials and Methods Using 6 years of data from 65 OMUs in a band of wild hamadryas baboons in Ethiopia, we compared demographic and reproductive parameters of OMUs with and without secondary “follower” males to assess whether their presence conferred any reproductive benefits to resident “leader” males. Results Leaders with followers had tenure lengths almost twice as long, acquired more than twice as many females, retained females longer, and had three times as many infants during their tenure compared to leaders without followers. Discussion Hamadryas follower males enabled leaders to retain females for longer periods of time—likely through unit defense, social relationships with OMU members, and/or infant protection. Hamadryas leaders appear to be able to monopolize access to females despite the presence of followers, and as such any enhanced reproduction derived from the presence of followers likely increases the fitness of the leader rather than the follower. Thus the relationship between leaders and followers in hamadryas society appears to be a mutually beneficial one and tolerance of secondary males may be an adaptive reproductive strategy characterizing hamadryas leader males. Am J Phys Anthropol 158:501–513, 2015. © 2015 Wiley Periodicals, Inc.

In polygynous primates, a greater reproductive variance in males have been linked to their reduced life expectancy relative to females. The mortality patterns of monogamous pair-bonded primates, however, are less clear. We analyzed the sex differences in mortality within wild (NMales = 70, NFemales = 73) and captive (NMales = 25, NFemales = 29) populations of Azara's owl monkeys (Aotus azarae), a socially and genetically monogamous primate exhibiting biparental care. We used Bayesian Survival Trajectory Analysis (BaSTA) to test age-dependent models of mortality. The wild and captive populations were best fit by the logistic and Gompertz models, respectively, implying greater heterogeneity in the wild environment likely due to harsher conditions. We found that age patterns of mortality were similar between the sexes in both populations. We calculated life expectancy and disparity, the latter a measure of the steepness of senescence, for both sexes in each population. Males and females had similar life expectancies in both populations; the wild population overall having a shorter life expectancy than the captive one. Furthermore, captive females had a reduced life disparity relative to captive males and to both sexes in the wild. We interpret this pattern in light of the hazards associated with reproduction. In captivity, where reproduction is intensely managed, the risks associated with gestation and birth are tempered so that there is a reduction in the likelihood of captive females dying prematurely, decreasing their overall life disparity. Am. J. Primatol. © 2015 Wiley Periodicals, Inc.

Sexual dimorphism in mammals typically is reduced in monogamous species relative to polygynous species, with promiscuous species being intermediate. This pattern of dimorphism characterizes adult behavior and body mass of prairie voles, a monogamous species, when compared with meadow voles, a closely related polygynous or promiscuous species. We examined whether the pattern also applies to young of the two species by observing individual pups living in family groups in seminatural environments. Observations during the second week of life revealed no sex differences in pup behavior or body mass. However, we detected species differences in suckling behavior, jockeying for position within the huddle (especially among males), and body mass that replicate and extend our previous observations. These data indicate that patterns of sexual dimorphism associated with different mating systems may not be evident in juvenile mammals, but that species differences in behavior and body mass can be obvious at this stage.

Social and reproductive systems remain among the main predictors affecting mammalian birth sex ratio. The two extant hippopotamus species differ in their social and reproductive systems. While common hippopotamus (Hippopotamus amphibius) form herds and tend to be polygynous, solitary living pygmy hippopotamus (Choeropsis liberiensis) are promiscuous. Although it is one of the most studied topics, only few empirical studies using large sample sizes have reported distorted birth sex ratio. We examined the birth sex ratio in both hippopotamus species using international studbooks including large data sets exceeding a thousand individuals (1138 for common hippopotamus and 1161 for pygmy ones). In both species, the birth sex ratio differed from 1:1. Whereas more males than females were recorded in common hippopotamus (53.9% males), the opposite was found in pygmy hippopotamus (41.5% males). We also found that the birth sex ratio was affected by individual dams in common hippopotamus, and by individual sires in pygmy hippopotamus. The most plausible explanation for differentially skewed birth sex ratios in both species may be related to differences in social and reproductive systems. Whereas the polygynous, sexually dimorphic common hippopotamus biased the birth sex ratio towards males, the promiscuous and sexually monomorphic pygmy hippopotamus skewed the sex ratio in favour of females. Our results are in line with recent studies showing that not only manipulation by the mother (in common hippopotamus), but also by the father (in pygmy hippopotamus), may be responsible for the birth sex ratio in different species.

Tamarins are reported to live in small multimale-multifemale groups characterized by a single breeding female. Here we present information on the composition and genetic relatedness of individuals in 12 wild-trapped groups of Weddell's saddleback tamarins (Saguinus weddelli) from northern Bolivia to determine if groups are best described as nuclear or extended families suggesting social monogamy or whether groups contain several unrelated same sex adults indicative of social polyandry/polygyny. Mean group size was 6.25 including an average of 2.16 adult males (range 1-4) and 2.08 adult females (1-3). No group contained only one adult male and one adult female and 25% of groups contained two parous females. We estimated the genetic relatedness among individuals using 13 polymorphic microsatellite markers. Across the population, mean relatedness was low and not significantly different among adult males versus among adult females, suggesting that both sexes disperse from their natal groups. Adults of both sexes also tended to have close same-sex adult relatives within their groups; relatedness among adult females of the same group averaged 0.31 and among adult males was 0.26. This suggests that tamarins of one or both sexes sometimes delay dispersal and remain as adults in their natal group or that emigration of same-sexed relatives into the same group may be common. Finally, parentage analyses indicated that, whereas the parents of juveniles generally were present in the group, this was not always the case. Based on these data, published reports of the presence of multiple breeding males and occasionally multiple breeding females in the same group, and the fact that less than 10% of groups in the wild contain a single adult male-adult female pair, we argue that social polyandry best characterizes the composition of tamarin groups and that monogamy is not a common mating pattern in Saguinus weddelli or other tamarin species.

Academics frequently serve as expert witnesses in legal cases yet their role as transmitters of social scientific knowledge remains under-examined. The present study analyzes the deployment of social science within British Columbias polygamy reference trial where research is used to support the assertion that polygamy is inherently harmful to society. Within the trial record and the written decision the protection of monogamy as an institution is performed in part through the marginalization of qualitative methodology and the concurrent privileging of quantitative studies that purportedly demonstrate widespread social harms associated with the practice of polygyny.

Actuarial senescence is widespread in age-structured populations. In growing populations, the progressive decline of Hamiltonian forces of selection with age leads to decreasing survival. As actuarial senescence is overcompensated by a high fertility, actuarial senescence should be more intense in species with high reproductive effort, a theoretical prediction that has not been yet explicitly tested across species. Wild boar (Sus scrofa) females have an unusual life-history strategy among large mammals by associating both early and high reproductive effort with potentially long lifespan. Therefore, wild boar females should show stronger actuarial senescence than similar-sized related mammals. Moreover, being polygynous and much larger than females, males should display higher senescence rates than females. Using a long-term monitoring (18 years) of a wild boar population, we tested these predictions. We provided clear evidence of actuarial senescence in both sexes. Wild boar females had earlier but not stronger actuarial senescence than similar-sized ungulates. Both sexes displayed similar senescence rates. Our study indicates that the timing of senescence, not the rate, is associated with the magnitude of fertility in ungulates. This demonstrates the importance of including the timing of senescence in addition to its rate to understand variation in senescence patterns in wild populations.

Although dispersal requires context-dependent decision-making in three distinct stages (emigration, transit, immigration), these decisions are commonly ignored in simple models of dispersal. For sexually reproducing organisms, mate availability is an important factor in dispersal decisions. Difficulty finding mates can lead to an Allee effect where population growth decreases at low densities. Surprisingly, theoretical studies on mate finding and on sex-biased dispersal produce opposing predictions: in the former, one sex is predicted to move less if the other sex evolves to search more, whereas in the latter, mate-finding difficulties can select for less sex bias in dispersal when mate finding occurs after dispersal. Here, we develop a pair of models to examine the joint evolution of dispersal and settlement behaviour. Our first model resolves the apparent contradiction from the mate search and dispersal literatures. Our second model demonstrates that the relationship between mating system and sex-biased dispersal is more complex than a simple contrast between resource defence monogamy and female defence polygyny. Our results highlight that a key factor is the timing of mating relative to dispersal (before, during, or after). We also show that although movement has the potential to alleviate a mate-finding Allee effect, in some cases, it can actually exacerbate the effect.