Great expectations: dominance,breeding success and offspring sex ratios in red deer

Differences in dominance rank among red deer (Cervus elaphus) females (hinds) on Rhum were related to their breeding success as well as to the comparative success of male and female offspring. Males (stags) born to mothers above median rank were more successful than hinds, while hinds born to subordinate mothers were more successful than stags. The ratio of male to female calves produced by dominant mothers was significantly higher than that produced by subordinates. Since dominance rank among hinds is related to their body weight as adults and to their birth weight, these results suggest that the birth sex ratio may be affected by environmental factors operating during a female's early development.     

Female macaques compete for ‘power’ and ‘commitment’ in their male partners

The formation of male-female social bonds and the resulting competition among females for male partners is a core element of human societies. While female competition for a male partner outside the mating context is well studied in humans, evidence from non-human primates is scarce, and its evolutionary roots remain to be explored. We studied two multi male – multi female groups of wild Assamese macaques (Macaca assamensis), a species where females gain benefits from selectively affiliating with particular males. Using a behavioral data set collected over several years, we tested whether females competed over access to male social partners, whether success in competition was driven by female dominance rank, and which male traits were most attractive for females. We found assortative bonding by dominance rank between females and males, which together with females initiating and maintaining contact suggests direct female competition over males. Two male traits independently predicted male attractiveness to females: (1) current dominance rank, a measure of 'power' or a male's ability to provide access to resources, and (2) prior male affiliation with immatures, a measure of a male's potential paternal proclivity or 'commitment' to infant care. Both traits have been consistently identified as drivers of female partner choice in humans. Our study adds to the evidence that female competition for valuable male partners is not unique to humans, suggesting deep evolutionary origins of women's mate choice tendencies for ‘power’ and ‘commitment’.     

Maternal effects are long‐lasting and influence female offspring's reproductive strategy in the swordtail fish Xiphophorus multilineatus

The adaptive benefits of maternal investment into individual offspring (inherited environmental effects) will be shaped by selection on mothers as well as their offspring, often across variable environments. We examined how a mother's nutritional environment interacted with her offspring's nutritional and social environment in Xiphophorus multilineatus, a live‐bearing fish. Fry from mothers reared on two different nutritional diets (HQ = high quality and LQ = low quality) were all reared on a LQ diet in addition to being split between two social treatments: exposed to a large adult male during development and not exposed. Mothers raised on a HQ diet produce offspring that were not only initially larger (at 14 days of age), but grew faster, and were larger at sexual maturity. Male offspring from mothers raised on both diets responded to the exposure to courter males by growing faster; however, the response of their sisters varied with mother's diet; females from HQ diet mothers reduced growth if exposed to a courter male, whereas females from LQ diet mothers increased growth. Therefore, we detected variation in maternal investment depending on female size and diet, and the effects of this variation on offspring were long‐lasting and sex specific. Our results support the maternal stress hypothesis, with selection on mothers to reduce investment in low‐quality environments. In addition, the interaction we detected between the mother's nutritional environment and the female offspring's social environment suggests that female offspring adopted different reproductive strategies depending on maternal investment.     

Female Remating Behavior in a Lekking Moth

In polyandrous species, male reproductive success will at least partly be determined by males' success in sperm competition. To understand the potential for post‐mating sexual selection, it is therefore important to assess the extent of female remating. In the lekking moth Achroia grisella, male mating success is strongly determined by female choice based on the attractiveness of male ultrasonic songs. Although observations have indicated that some females will remate, only little is known about the level of sperm competition. In many species, females are more likely to remate if their first mating involved an already mated male than if the first male was virgin. Potentially, this is because mated males are less well able to provide an adequate sperm supply, nutrients, or substances inhibiting female remating. This phenomenon will effectively reduce the strength of pre‐copulatory sexual selection because attractive males with high mating success will be more susceptible to sperm competition. We therefore performed an experiment designed both to provide a more precise estimate of female remating probability and simultaneously to test the hypothesis that female remating is influenced by male mating history. Overall, approximately one of five females remated with a second male. Yet, although females mated to non‐virgin males were somewhat more prone to remate, the effect of male mating history was not significant. The results revealed, however, that heavier females were more likely to remate. Furthermore, we found that females' second copulations were longer, suggesting that, in accordance with theory, males may invest more sperm in situations with an elevated risk of sperm competition.     

Determinants of fecundity and reproductive success in captive vervet monkeys

Between 1975 and 1983, adult female vervet monkeys (Cercopithecus aethiops sabaeus) over 3.5 years of age, living in two undisturbed social groups in a captive colony in Sepulveda, California, have averaged 1.0 births per female year with a mean interbirth interval of 10.7 months. Increased fecundity did not result in decreased survival rates of offspring in this population. Fecundity was influenced by the mother's age and dominance rank. The primary factor in the age-fecundity relationship was the age at first birth, which varied from three to five years. High-ranking females contributed the most to the high rate of fecundity, with significantly shorter interbirth intervals, more births per female year, and more surviving infants compared to low-ranking females.     

Mating Group Size and Stability in Reindeer Rangifer tarandus: The Effects of Male Characteristics,Sex Ratio and Male Age Structure

In polygynous mating systems, males compete intensely for mates and may mate several females during a single reproductive season. Accordingly, factors influencing the ability of males to control a larger number of females during the breeding season can provide information on the processes underlying sexual selection. In ungulates, age, body mass and social rank are considered good predictors of the reproductive success of males, but how male age structure and sex ratio in the population influence mating group (MG) dynamics has received little empirical testing. Between 1996 and 2005, we manipulated male age‐ and sex structure and monitored MG dynamics in a reindeer (Rangifer tarandus) population. We investigated the influence of male characteristics, percentage of males and male age structure on MG size and stability. We found that males with higher social rank (that were also older and heavier) controlled larger MGs (therefore had greater mating opportunities) and had more stable MGs (corresponding to a higher ability to maintain and control females) than males of lower social rank. Moreover, MG size and MG stability decreased as the percentage of males in the population increased, most likely resulting from greater male–male competition and increased female movements. Male age structure did not influence MG stability. Given the positive relationship between mating success and MG size (and likely MG stability), frequent female movements and intense competition among males to control females seem to be the principal components of reindeer MGs dynamic.     

Domestic Calves (Bos taurus) Recognize their Own Mothers by Auditory Cues

The goal of this study was to determine if auditory cues are important in maternal recognition by domestic cattle calves, Bos taurus. Cows and their calves were separated and the vocalizations of the mothers were recorded. During experimental playbacks in a test enclosure, each calf (n = 9) was given a choice between a tape-recorded vocalization of its mother and that of a strange mother. Calves significantly preferred their own mother's vocalization as compared to the vocalization of the unfamiliar mother. Calves spent significantly more time near the speaker that played their own mother's call, and approached significantly closer to their mother's speaker. These results demonstrate that 3–5-wk-old calves can recognize their mothers by auditory cues alone. Visual inspection of audiospectrograms of the cows' vocalizations suggests that there are individual differences among cows.     

Contrasting the Effects of Maternal and Behavioral Characteristics on Fawn Birth Mass in White-Tailed Deer

Maternal care influences offspring quality and can improve a mother’s inclusive fitness. However, improved fitness may only occur when offspring quality (i.e., offspring birth mass) persists throughout life and enhances survival and/or reproductive success. Although maternal body mass, age, and social rank have been shown to influence offspring birth mass, the inter-dependence among these variables makes identifying causation problematic. We established that fawn birth mass was related to adult body mass for captive male and female white-tailed deer (Odocoileus virginianus), thus maternal care should improve offspring fitness. We then used path analysis to identify which maternal characteristic(s) most influenced fawn birth mass of captive female white-tailed deer. Maternal age, body mass and social rank had varying effects on fawn birth mass. Maternal body mass displayed the strongest direct effect on fawn birth mass, followed by maternal age and social rank. Maternal body mass had a greater effect on social rank than age. The direct path between social rank and fawn birth mass may indicate dominance as an underlying mechanism. Our results suggest that heavier mothers could use dominance to improve access to resources, resulting in increased fitness through production of heavier offspring.     

Social rank,feeding and winter weight loss in red deer: any evidence of interference competition?

During winter at northern latitudes, large herbivores often exploit patches of concentrated, relatively high quality forage, which may lead to interference competition. The factors affecting success in contests and subsequent dominance rank, such as age and body weight, remain key issues in ungulate behavioural ecology. Maternal effects on offspring body weight are well known, but few studies have investigated if mothers social rank influence offspring rank. Moreover, no study has related dominance rank in ungulates to weight loss during winter. Outcomes of social interactions (n=7,609), feeding time and spatial position in red deer (Cervus elaphus) hinds and calves, and weight loss of calves, were registered from 1981 to 1996 at six winter-feeding sites within the county of Sør-Trøndelag in Norway. The level of aggressiveness was higher among calves than among adult hinds, and the factors determining the outcome of contests also differed. The initiator won the majority of interactions (more than 90% in both hinds and calves). Social rank was related to both age and body weight in adult hinds, and related to body weight and mother rank in calves. The relationship between feeding time and rank was non-linear. Feeding time was correlated with rank only among high ranked hinds, while there was no such relationship among low ranked hinds or calves. There was no correlation between winter weight loss and social rank in calves. Our study therefore underlines that, although frequent aggression is observed at artificial feeding sites of northern herbivores, this is not necessarily sufficient to give rise to interference competition.     

Female rank and reproductive success among arashiyama B Japanese macaques (Macaca fuscata)

Five hypotheses that related female rank and reproductive success were tested in an intact troop of free-ranging, provisioned, Japanese macaques. The hypotheses stated that high-ranking females (1) begin parturition earlier in life than low-ranking females; (2) produce more offspring than low-ranking females; (3) give birth during some optimal time during the birth season to a greater extent than low-ranking females; (4) experience less infant mortality than low-ranking females;and (5) more frequently produce male offspring, while low-ranking females more frequently produce female offspring. A statistical analysis of the data which included three birth seasons and 55 adult females and 34 pubescent females, all of known age, rank, and matrifocal membership in the Arashiyama B troop, revealed few significant results. An association was found between the rank of the matrifocal unit and the age of first birth. However, the relationship was the reverse of hypothesis 1, i.e., females of the lower-ranking matrifocal units began parturition earlier than females of higher-ranking matrifocal units. Therefore, in this troop of Japanese monkeys— where alternative feeding strategies existed— there was little association between female rank and reproductive success.     

Recognition of Individual Males' Songs by Female Dunnocks: a Mechanism Increasing the Number of Copulatory Partners and Reproductive Success

Previous studies have shown that a female dunnock Prunella modularis increases her reproductive success on average by copulating with more than one male resident on her territory and thereby obtaining extra help in raising offspring. Here we document behavior by females that affects which males copulate with them. During her period of receptivity to copulation, a female in a territory shared by two males often left the dominant (or alpha) male, which guarded her most of the time, and approached the subordinate (or beta) male when he sang. A female's responses to individual males thus tend to increase her own reproductive success by increasing her chances for copulation with both males sharing her territory. Playbacks of tape-recorded songs in the field showed that females approached only songs of resident males, not neighbors. They can therefore discriminate individual males by their songs alone, a capability not previously established for female songbirds. Despite intensive guarding of females by males, mating success among male dunnocks depends in part on female choice.     

Longitudinal stability in adult status hierarchies among vervet monkeys (Cercopithecus aethiops)

Status hierarchies were examined in a captive group over a 12-year period (1968–1979) to establish the stability and structure of vervet monkey dominance relationships. Dyadic records of spontaneous avoid, bite, chase, and supplant behaviors are summarized into a win/lose matrix to produce a rank order of status at the end of each calendar year. The 13,717 agonistic acts tabulated are drawn from both focal and ad lib data. Relative rank is computed by dividing the rank position of each individual by the number of individuals ranked for that year. Relative rank was found to be quite stable for adults of both sexes, although males were more variable than females. There was a strong matinrline effect on all captive born animals' rank position. No gender difference in rank was demonstrable. Infants were very subordinate at the end of their first year, but rose to positions near those of their mothers by the end of the fourth year. Presence or absence of canine teeth had no demonstrable effect on status. The results emphasize the relative importance of social learning in the establishment and maintenance of dominance relationships. The apparent gender difference in rank among wild living vervets is hypothesized to be a consequence of the male intergroup mobility. If a male is restrained from emigration, he is strongly affected by his mother's rank as he establishes his own dominance position in the troop.     

Mother — Offspring Conflict in the Saharan Arrui,Ammotragus lervia sahariensis: Relation to Weaning and Mother's Sexual Activity

Behavioural conflict between mother and offspring was investigated in a captive population of the Saharan arrui (Ammotragus lervia sahariensis, Artiodactyla: Bovidae). The frequency of agonistic behaviours (mother's aggressiveness and calf's retreats) and unsuccessful suckling attempts, Hinde's proximity index and the distances between mother and calf have been analysed in relation to weaning and sexual activity of the mother. Mothers that conceive in the following mating season accelerate the weaning process of their current offspring; furthermore, high-ranking females wean their male calves earlier. Behavioural conflict occurs during the resumption of the mother's sexual activity; however, no conflict appears to happen during weaning.     

Multiple Mating Increases Fecundity, Fertility and Relative Clutch Mass in the Female Leopard Gecko (Eublepharis macularius)

Females across many taxa may mate with several males or mate more than once with the same male within one reproductive event. Although many researchers have discussed the effects of multiple mating on reproductive success of females, few studies have attempted to disentangle whether the reproductive success of females differs with respect to whether females mate with multiple males or mate more than once with one male. In this study, we hypothesized that female leopard geckos (Eublepharis macularius) increase aspects of their reproductive success, such as fecundity, fertility and relative clutch mass, by mating more than once within one reproductive event, either by mating repeatedly with the same male or multiply mating with different males. We controlled for the potentially confounding variables of mating frequency and mate number by allowing females to mate once with one male, twice with the same male, or twice with two different males. We found that females that mated with more than one male laid more clutches, exhibited increased egg fertility and invested more in clutches relative to females that mated only once with one male, whereas females that mated twice to the same male were intermediate for these variables. Thus, reproductive success is higher among female leopard geckos that mated with more than one male compared to female leopard geckos that mated only once.     

Female weight and reproductive condition in a population of olive baboons (Papio anubis)

The study reported here tested Altmann's prediction [Baboon Mothers and Infants. Cambridge, Harvard University Press, 1980] that lactating female baboons endure a weight loss. Data from 64 adult female olive baboons (Papio anubis) residing in six troops in Kenya revealed that reproductive condition was related to weight. Lactating females weighed less and pregnant females weighed more than cycling females. There was a negative correlation between the weight of cycling females and the number of months postweighing to their next conception. These results indicate that lactation in wild baboons imposes energy costs that result in lost weight. It is suggested that female baboons may have to surpass a minimum weight threshold prior to resumption of postlactational cycling and that nutritional status is more influential than rank in affecting female reproductive success.     

Reproductive success of cat fleas,Ctenocephalides felis,on calves as unusual hosts

Abstract. Cat fleas, Ctenocephalides felis, were released onto calves as unusual hosts, and sampled at intervals for histological examination. Egg output from fleas on age-matched male and female calves was monitored. Using indicators of reproductive maturation and regression together with egg output data, the reproductive success and fertility of cat fleas on male and female calves were estimated. Comparisons were made with fleas taken from cats. The mean egg output of fleas on the bull calf was highly significantly different from that on the age-matched female calf: 28.14 ± 2.96 (SE) eggs/h compared with 16.21 ± 1.96 (SE) eggs/h. A higher proportion of sampled fleas (83.0%) was reproductively mature on the feline hosts compared with the calves (45.4-62.5%). Blue bodies resulting from oocyte resorption were present in the ovarioles of 10.4-19.0% of fleas sampled from the calves. No blue bodies were present in fleas removed from cats. Eggs laid by fleas on calves were viable and larvae were reared to adulthood. The mean percentage hatching success on the age-matched male and female calves was 46.7% and 51.7%. This represents a reduction in viability of 28-33% compared with eggs laid by fleas on cats. Factors which may account for reduced reproductive maturation of fleas on calves, including protein content of the host's blood, are discussed.     

Female competition and its evolutionary consequences in mammals

Following Darwin's original insights regarding sexual selection, studies of intrasexual competition have mainly focused on male competition for mates; by contrast, female reproductive competition has received less attention. Here, we review evidence that female mammals compete for both resources and mates in order to secure reproductive benefits. We describe how females compete for resources such as food, nest sites, and protection by means of dominance relationships, territoriality and inter‐group aggression, and by inhibiting the reproduction of other females. We also describe evidence that female mammals compete for mates and consider the ultimate causes of such behaviour, including competition for access to resources provided by mates, sperm limitation and prevention of future resource competition. Our review reveals female competition to be a potentially widespread and significant evolutionary selection pressure among mammals, particularly competition for resources among social species for which most evidence is currently available. We report that female competition is associated with many diverse adaptations, from overtly aggressive behaviour, weaponry, and conspicuous sexual signals to subtle and often complex social behaviour involving olfactory signalling, alliance formation, altruism and spite, and even cases where individuals appear to inhibit their own reproduction. Overall, despite some obvious parallels with male phenotypic traits favoured under sexual selection, it appears that fundamental differences in the reproductive strategies of the sexes (ultimately related to parental investment) commonly lead to contrasting competitive goals and adaptations. Because female adaptations for intrasexual competition are often less conspicuous than those of males, they are generally more challenging to study. In particular, since females often employ competitive strategies that directly influence not only the number but also the quality (survival and reproductive success) of their own offspring, as well as the relative reproductive success of others, a multigenerational view ideally is required to quantify the full extent of variation in female fitness resulting from intrasexual competition. Nonetheless, current evidence indicates that the reproductive success of female mammals can also be highly variable over shorter time scales, with significant reproductive skew related to competitive ability. Whether we choose to describe the outcome of female reproductive competition (competition for mates, for mates controlling resources, or for resources per se) as sexual selection depends on how sexual selection is defined. Considering sexual selection strictly as resulting from differential mating or fertilisation success, the role of female competition for the sperm of preferred (or competitively successful) males appears particularly worthy of more detailed investigation. Broader definitions of sexual selection have recently been proposed to encompass the impact on reproduction of competition for resources other than mates. Although the merits of such definitions are a matter of ongoing debate, our review highlights that understanding the evolutionary causes and consequences of female reproductive competition indeed requires a broader perspective than has traditionally been assumed. We conclude that future research in this field offers much exciting potential to address new and fundamentally important questions relating to social and mating‐system evolution.     

Male dominance rank and reproductive success in primate groups

The relationship between male dominance rank and reproductive success has been a topic of interest since the beginning of primatology. From a theoretical point of view the existence or absence of this relationship has great implications with respect to the meaning of dominance rank and more general of the social relationships between individuals within social groups. Until fairly recently mating behaviour has been used as an indicator of reproductive success, but these two variables need not be correlated. The relations between mating success, reproductive success, and dominance rank indicate whether selective mating is involved (different mating partners at different phases of fertility, for example through male contest or female choice). With the development of genetic techniques to determine reproductive success directly, it has now become possible to investigate these relations. In our study on wild long-tailed macaques (Macaca fascicularis) we find a relatively strong correlation between rank and reproductive success which is attributable to selective mating by the alpha male during fertile periods of the females. In most previous studies no such clear results have been obtained, and we discuss the differences in outcome in relation to the study conditions, group sizes, and possible differences between species in terms of reproductive strategies.     

Female behavioral strategies during consortship in Tibetan macaques (Macaca thibetana)

Consortship has been defined as a temporary association between an adult male and an estrous/receptive female. It has been considered as male mating strategies to improve male mating success and potential reproductive success. However, the female roles have been more or less neglected, and thus, less is known about female behavioral strategies during the consortship periods. In this study, during the two consecutive mating seasons, we collected behavioral data of free‐ranging Tibetan macaques (Macaca thibetana) habituated in Mt. Huangshan, China, to investigate female behaviors when she was consorted by an adult male. The results showed that (a) females were more likely to approach and exhibit sexual solicitation to their consorting males during the consorted period, and females also exhibited less approach to their nonconsorting males; (b) females exhibited strong responses (either departed distantly or formed affiliative relationships with their consorting male partner) when their consorting males mated with rival females or showed sexual motivation toward rival females; (c) female preferences were positively correlated to the duration of consortships and the frequencies of ejaculation copulations, independent of the social ranks of their consorting male partners. Our results suggested that female strategies played much more important roles in forming and maintaining consortship than previously assumed. It provides new insight into understanding female adaptive strategies to male strategies by forming consortships in multimale–multifemale primate species when males could not identify female''s fertile phase accurately.     

Measuring male-female relationships during the mating season in wild Japanese macaques (Macaca fuscata yakui)

Heterosexual relationships during one mating season were examined in a wild troop of Japanese macaques (Macaca fuscata yakui) on Yakushima Island, Japan. Validation tests of putative mate choice behaviors demonstrated that female initiation and maintenance of proximity, female lookback at the male, and sexual presents to the male, were associated with increased mating. Male grooming the female was also associated with increased mating. Ten dyadic social behaviors were subject to principal components analysis to empirically define behavioral dimensions of male-female relationships. The analysis yielded four relationship dimensions: ‘Mutual Choice and Male Coercion,’ ‘Female Choice’ (two types), and ‘Mutual Choice’ Dyads tended to be characterized by more than one dimension. The results suggested that females sought matings with multiple males of various dominance ranks. Female relationships with high ranking males contained elements of male coercion and mate guarding, however, because these males attempted to inhibit females from mating with lower ranking males. The correlation between each relationship dimension and mating success depended, in part, on the dominance rank of males. Relationships involving high ranking males, which were most likely to contain elements of male coercion and mate guarding, were associated with mating success. Relationships involving low ranking males, which usually lacked such coercive elements. were less strongly correlated with mating success. These results, obtained from a wild troop, are compared to those previously obtained in captive and provisioned groups of Japanese macaques.