Female survival,lifetime reproductive success and mating status in a passerine bird

In facultatively polygynous birds, secondary females of polygynously mated males typically have reduced annual reproductive success, because polygynous males provide less paternal care than monogamous males. Life history theory predicts that, as a result of increased reproductive investment, secondary females should suffer from reduced survival and lifetime reproductive success, but previous studies provided only weak support for this hypothesis. We used 7 years of data to study the fitness of female collared flycatchers Ficedula albicollis in relation to mating status by estimating survival and lifetime reproductive success. Taking differences in recapture probability into account, a mark-recapture analysis revealed that females observed at least once to breed as secondary female had higher survival than other females. This relationship was not confounded by laying date, because when we assessed the impact of laying date on survival, we found similar survival patterns. Females of polygynous males had reduced breeding success in terms of number of young fledged during the current reproductive event. However, during their lifetime females found at least once in primary or secondary mating status produced significantly more eggs, and at least the same number of fledglings and recruits as monogamous females. Thus, in the collared flycatcher, females of polygynously mated males seem to suffer from mating status during the most recent reproductive event, but considering survival and lifetime reproductive success, the apparently disadvantageous mating event is not necessarily associated with reduced residual reproductive value.     

Pair bonding and "the widow effect" in female prairie voles

We conducted field and laboratory experiments with the well-studied monogamous prairie vole, Microtus ochrogaster, to distinguish among three hypotheses for the failure of females that lose their mates to bond with a new male ("the widow effect"). The reproductive value hypothesis predicts that males prefer young to older females because they potentially have a longer reproductive lifespan. The mate rejection hypothesis predicts that females will prevent repairing by aggressively deterring males that might harm their current offspring. The misdirected paternal care hypothesis assumes that females will mate during postpartum estrus and thus will be pregnant and/or nursing young throughout the breeding season; males will avoid pairing with older females to avoid providing care to unrelated offspring and/or because of a delay to the next breeding opportunity. Males associated and bred more with older than young females, allowing us to reject the reproductive value hypothesis. Our results were consistent with the male rejection hypothesis in that females were aggressive toward unfamiliar males. Our results were most consistent with the misdirected paternal care hypothesis in that once females started breeding, they continued to become pregnant and nurse young throughout the study period. Thus, our findings suggest that the potential of misdirected paternal care and delayed mating opportunity in conjunction with the aggressive behavior of females toward unfamiliar males are likely explanations for the lack of repairing for widow females.     

Age-specific mating strategies and reproductive senescence

While males gain obvious direct advantages from multiple mating, the reproductive capacity of females is more constrained. The reason why polyandry evolved in females is therefore open to many conjectures. One hypothesis postulates that females gain indirect benefits by increasing the probability of siring young from high quality males. To explore this hypothesis, we used the natural variation of the reproductive value that males and females undergo through age. The age-related variation of phenotypic performance might then induce variations in mating strategies in males and females. Using the common lizard (Lacerta vivipara) as our model system, we showed that reproductive immaturity and senescence created variability in both male and female reproductive success (including survival of offspring). Consistent with theory, males at their best-performing phenotype adopted a polygynous strategy. These males were of an intermediate age and they produced offspring of higher viability than younger and older males. In contrast, females at their best performing phenotype, also of an intermediate age, were less polyandrous than other less-performing females. Middle-aged females tended to mate with males of an intermediate age and produced litters with higher viability independently from their reproductive strategy. Males of an intermediate age enhanced their fitness by additional matings with young or old females. Young and old females increased their fitness by being more polyandrous. Polyandry therefore appears as means to seek for good males. A positive correlation between males and their partners' fitness disagree with the idea that polyandry is the result of a sexual conflict in this species.     

Cryptic female choice: frogs reduce clutch size when amplexed by undesired males

In species with internal fertilization, females can 'cryptically' choose (e.g. through sperm selection) which individuals sire their offspring, even when their overt preferences for copulatory partners are overrun by male-male competition and sexual coercion. The experiment presented here reveals that control of paternity after copulation has begun is also possible in species with external fertilization. Females of the hybridogenetic Rana essonae-Rana esculenta (LL-LR) waterfrog complex adjust their clutch size in response to mate type: they release fewer eggs when amplexed by hybrid LR males who--jeopardize successful reproduction--than when amplexed by parental LL males. This reduction in the number of eggs laid can increase a female's residual reproductive value through a second mating in the same breeding season or a larger clutch size in the next year. We argue that cryptic female choice through clutch size adjustment (i) may have evolved more often than previously assumed, and (ii) can arise even where females mate only once during a reproductive period.     

Female Choice of Least Costly Males; a Possible Factor in the Evolution of Leks

It is proposed that leks have evolved because females benefit by the absence of breeding males from female ranges. Non-lekking males impose several kinds of cost on female reproductive success. Under certain conditions female preference for least costly males can therefore favour males who wait, rather than search, for females.     

Sociobiology of the great apes and the hominid ancestor

This review of recent field studies of the great apes summarizes and weighs socioecological and sociobiological evidence concerning the ultimate causes of social structure. The behavioral ecology and social structures of mountain gorillas, orangutans, chimpanzees, and bonobos are reviewed and contrasted between species. Social dynamics and molecular studies indicate that, among the extant Hominoidea, the evolutionary clade of chimpanzees, bonobos, and humans probably evolved from the most recent common ancestor in the ape-human stem. The most probable phylogenetic referential model for the suite of social behaviors of the hominid ancestor consists of the behavioral traits common to all three species: female exogamy, male retention, female associations due to attraction to the same male(s), weak bonds between females, a closed, stable social group made up of a kin-group of males and containing multiple females, fusion-fission sociality in which individuals of either sex sometimes travel alone, a polygynous mating system, communal territoriality with cooperative defense by kin-related males who exhibit strong solidarity among themselves but who may kill other males in territorial disputes, low mating competition between males within communities, and moderate sexual dimorphism. It is postulated that this phylogenetic model is a useful tool for comparing goodness of fit of other referential models seeking to explain hominid evolution. It is also suggested that to construct a “strategic” or conceptual model to explain hominid evolution, the putative evolutionary processes responsible for this male-retentive system require further testing in the field by measuring individual reproductive success among the great apes and man.     

Human breasts: Unsupported hypotheses reviewed

Unlike other apes, human females’ breasts develop before first pregnancy and are permanently enlarged. Evidence suggests breasts act as signals to males but the critical data required to confirm this are lacking. These facts have led to a number of hypotheses about the evolutionary and adaptive significance of the human breast which fall into two groups. Those that address the presence of breasts in humans are (a) that they act as releasers of male sexual behaviour, (b) that they enable females to hide their reproductive condition, and (c) that they allow infants to nurse from their mother’s hip. Those that address variability in breast size are (d) that large breasts indicate lactational potential, (e) ability of mothers to invest prenatally in offspring, (f) mother’s fecundity, and (g) her longevity. Each hypothesis is reviewed and evaluated using logical or empirical arguments. Possible ways in which the adaptive significance of human breasts can be determined in contemporary populations are outlined.     

A comparative study on the evolution of reversed size dimorphism in monogamous waders

Sexual dimorphism in size is common in birds. Males are usually larger than females, although in some taxa reversed size dimorphism (RSD) predominates. Whilst direct dimorphism is attributed to sexual selection in males giving greater reproductive access to females, the evolutionary causes of RSD are still unclear. Four different hypotheses could explain the evolution of RSD in monogamous birds: (1) The ‘energy storing’ hypothesis suggests that larger females could accumulate more reserves at wintering or refuelling areas to enable an earlier start to egg laying. (2) According to the ‘incubation ability’ hypothesis, RSD has evolved because large females can incubate more efficiently than small ones. (3) The ‘parental role division’ hypothesis suggests that RSD in monogamous waders has evolved in species with parental role division and uniparental male care of the chicks. It is based on the assumption that small male size facilitates food acquisition in terrestrial habitats where chick rearing takes place and that larger females can accumulate more reserves for egg laying in coastal sites. (3) The ‘display agility’ hypothesis suggests that small males perform better in acrobatic displays presumably involved in mate choice and so RSD may have evolved due to female preference for agile males. I tested these hypotheses in monogamous waders using several comparative methods. Given the current knowledge of the phylogeny of this group, the evolutionary history of waders seems only compatible with the hypothesis that RSD has evolved as an adaptation for increasing display performance in males. In addition, the analysis of wing shape showed that males of species with acrobatic flight displays had wings with higher aspect ratio (wing span/²wing area) than non-acrobatic species, which probably increases flight manoeuvrability during acrobatic displays. In species with acrobatic displays males also had a higher aspect ratio than females although no sexual difference was found in non-acrobatic species. These results suggest that acrobatic flight displays could have produced changes in the morphology of some species and suggest the existence of selection favouring higher manoeuvrability in species with acrobatic flight displays. This supports the validity of the mechanisms proposed by the ‘display agility’ hypothesis to explain the evolution of RSD in waders.     

Evolution of human serial pairbonding

Data on divorce taken for all available years between 1947 and 1981 from the Demographic Yearbooks of the United Nations on 58 peoples illustrate that divorce has a consistent pattern. Divorces exhibit a skewed distribution, characterized by the occurrence of the mode early in marriage (with a divorce peak on or around the fourth year) and a gradual, long-tailed decline following this peak. Divorce risk peaks in age category 25-29 for males and age categories 20-24 and 25-29 for females, the height of reproductive and parenting years, and divorce counts peak among couples with two or fewer children. These properties of divorce are unrelated to divorce rate; they occur in societies with both high and low divorce rates. Data on available horticultural and gathering/hunting societies illustrate that divorce also peaks among young couples early in marriage. Remarriage by divorced and widowed individuals of reproductive age is also common cross-culturally. It is proposed that the above four-year modal marriage duration among couples of reproductive age who divorce reflects a hominid reproductive strategy that probably evolved some time after the appearance of Homo in response to increased female "reproductive burden" and functioned to ensure the survival of the hominid infant through weaning. Serial pairbonding during the female's reproductive years had ancestral adaptive advantages, producing the modern cross-cultural pattern of serial pairbonding.     

Paternal care as a conditional strategy: distinct reproductive tactics associated with elaboration of plumage ornamentation in the house finch

When individuals in a population differ in physiological conditionand residual reproductive value, selection should favor phenotypicplasticity in reproductive investment such that individualsare able to adopt the reproductive tactic that results in thehighest fitness under given conditions. Here we examined reproductivetactics in relation to the elaboration of condition-dependentsexual ornamentation (carotenoid breast coloration) in a Montanapopulation of the house finches (Carpodacus mexicanus). Malesused distinct reproductive tactics depending on elaborationof their sexual ornamentation. Males with red pigmentation (maximum ornament elaboration) paired with females that nestedearlier, but these males did little provisioning of incubatingfemales and nestlings. In contrast, males with yellow colorationpaired with females that nested later, but these males fedfemale and nestlings more. Consequently, for red males offspringrecruitment was primarily affected by earlier nest initiation, whereas in yellow males it was affected most by male provisioning.In males with intermediate plumage coloration, all measuredcomponents, nest initiation, provisioning of incubating female,and nestling feeding, strongly contributed to offspring recruitment.The fitness consequences of alternative reproductive tacticsof males were influenced by breeding experience and fidelityof their mates. Among first-time breeders, red males achievedthe highest fecundity because of the advantage gained throughearly nesting and pairing with more experienced females andbecause of compensation by their mates for low male provisioningof nestlings. Among experienced breeders, males with intermediateplumage coloration achieved the highest fecundity because ofthe combined benefits of relatively early pairing and high parental care. High variation in sexual ornamentation in a Montana populationof house finches may favor distinct associations of sexualdisplays with a particular set of reproductive behaviors.     

The patriarch hypothesis

Menopause is puzzling because life-history theory predicts there should be no selection for outliving one’s reproductive capacity. Adaptive explanations of menopause offered thus far turn on women’s long-term investment in offspring and grandoffspring, all variations on the grandmother hypothesis. Here, I offer a very different explanation. The patriarch hypothesis proposes that once males became capable of maintaining high status and reproductive access beyond their peak physical condition, selection favored the extension of maximum life span in males. Because the relevant genes were not on the Y chromosome, life span increased in females as well. However, the female reproductive span was constrained by the depletion of viable oocytes, which resulted in menopause. Frank Marlowe, Ph.D., is Assistant Professor of Anthropology at Harvard University. He conducts research with the Hadza and his interests include the behavioral ecology of mating systems, life-history theory, and cooperation.     

Life History Dependent Behavioural Variation in Water Striders,Aquarius remigis

We investigated behavioural consequences of life history states in Aquarius (Gerris) remigis, a hemipteran surface predator and scavenger of small North American streams. A repeated-measures field study comparing reproductive and non-reproductive first summer generation females and males during the same time in summer showed that non-reproductive males and females, which were foraging to survive the winter, behaved essentially alike, often being territorial. In contrast, reproductives of both sexes were more mobile than non-reproductives, reproductive males being much more mobile than reproductive females, always in search of and attacking potential mates. This resulted in reproductive males showing litde territoriality and having lower foraging success than all other categories, while reproductive females had higher foraging success than non-reproductive females or males, thus likely increasing their fecundity. Many behavioural variables did not change with time. Exceptions were a decrease in the mating activity of reproductives of both sexes, the number of mating attempts, mobility, and the stride rate of reproductive males, and an increase in overall foraging activity of all individuals. This indicates that the behaviour of reproductive males converged towards that of non-reproductive males as the season progressed. Individuals were assigned to either state post-hoc according to whether they had been seen mating at least once during the study period (July and August). Females were further tested to ascertain if they laid eggs. These results support the hypothesis that diapause and reproductive state to some degree predetermine behaviour, thus constraining behavioural flexibility. Attributing behavioural variation to various proximate causes can be central to the interpretation of alternative strategies and tactics.     

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.     

Sexual segregation in western grey kangaroos: testing alternative evolutionary hypotheses

In sexually dimorphic ungulates, sexual segregation is hypothesized to have evolved because of sex-specific differences in body size and/or reproductive strategies. We tested these alternative hypotheses in kangaroos, which are ecological analogues of ungulates. Kangaroos exhibit a wide range of body sizes, particularly among mature males, and so the effects of body size and sex can be distinguished. We tested predictions derived from these hypotheses by comparing the distribution of three sex–sex size classes of western grey kangaroos Macropus fuliginosus , in different habitats, and the composition of groups of kangaroos, across seasons. In accordance with the predation risk-reproductive strategy hypothesis, during the non-breeding season, females, which were more susceptible to predation than larger males, and were accompanied by vulnerable young-at-foot, were over-represented in secure habitats. Large males, which were essentially immune to predation, occurred more often than expected in nutrient-rich habitat, and small males, which faced competing demands of predator avoidance and feeding, were intermediate between females and large males in their distribution across habitats. During the breeding season, females continued to be over-represented in secure habitats when their newly emerged pouch young were most vulnerable to predation. All males occupied these same habitats to maximize their chances of securing mates. Consistent with the social hypotheses, groups composed of individuals of the same sex, irrespective of body size, were over-represented in the population during the non-breeding season, while during the breeding season all males sought females so that mixed-sex groups predominated. These results indicate that body size and reproductive strategies are both important, yet independent, factors influencing segregation in western grey kangaroos.     

Sex-specific costs of reproduction in Eastern Bluebirds Sialia sialis

In species with bi-parental care, individuals must partition energy between parental effort and mating effort. Typically, female songbirds invest more than males in reproductive activities such as egg-laying and incubation, but males invest more in secondary sexual traits used in attracting mates. Animals that breed more than once within a season must also allocate time and energy between first and subsequent breeding attempts and between current and future breeding seasons. To investigate strategies of reproductive investment by males and females and the consequences of such strategies, we manipulated the size of broods of Eastern Bluebirds Sialia sialis . Pairs with enlarged first broods were less likely to produce a second clutch or took longer to initiate one than pairs with reduced broods. After rearing enlarged broods, females were less likely than males to survive to the following year. Although plumage coloration is a sexually selected trait in Eastern Bluebirds that is influenced by nutritional stress, we did not detect an effect of brood-size manipulation on female coloration. Past research, however, demonstrates that, in males, plumage colour is negatively affected by increasing brood size. We suggest that there are sex-specific strategies of reproductive investment in Eastern Bluebirds, and that researchers should incorporate measures of residual reproductive value in studies of life-history evolution.     

Similar gender dimorphism in the costs of reproduction across the geographic range of Fraxinus ornus

BACKGROUND AND AIMS: The reproductive costs for individuals with the female function have been hypothesized to be greater than for those with the male function because the allocation unit per female flower is very high due to the necessity to nurture the embryos until seed dispersal occurs, while the male reproductive allocation per flower is lower because it finishes once pollen is shed. Consequently, males may invest more resources in growth than females. This prediction was tested across a wide geographical range in a tree with a dimorphic breeding system (Fraxinus ornus) consisting of males and hermaphrodites functioning as females. The contrasting ecological conditions found across the geographical range allowed the evaluation of the hypothesis that the reproductive costs of sexual dimorphism varies with environmental stressors. METHODS: By using random-effects meta-analysis, the differences in the reproductive and vegetative investment of male and hermaphrodite trees of F. ornus were analysed in 10 populations from the northern (Slovakia), south-eastern (Greece) and south-western (Spain) limits of its European distribution. The variation in gender-dimorphism with environmental stress was analysed by running a meta-regression between these effect sizes and the two environmental stress indicators: one related to temperature (the frost-free period) and another related to water availability (moisture deficit). KEY RESULTS: Most of the effect sizes showed that males produced more flowers and grew more quickly than hermaphrodites. Gender differences in reproduction and growth were not minimized or maximized under adverse climatic conditions such as short frost-free periods or severe aridity. CONCLUSIONS: The lower costs of reproduction for F. ornus males allow them to grow more quickly than hermaphrodites, although such differences in sex-specific reproductive costs are not magnified under stressful conditions.     

Cere Colour as a Basis for Extra-pair Preferences of Paired Male Budgerigars (Melopsittacus undulatus: Psittacidae: Aves)

Male budgerigars may be limited in the time they have available to pursue extra-pair copulations because of conflicts with other reproductive behaviour. If males could selectively court only immediately fertilizable extra-pair females they should do so. The cere colour of a female budgerigar may be used as a cue to her reproductive readiness. We experimentally tested the hypothesis that males prefer females with dark ceres, which are indicative of developed ovaries, over females with light-coloured ceres, characteristic of birds with undeveloped ovaries. In a pairwise choice experiment between females that had their cere colour artificially manipulated, pair-bonded males were more likely to choose the female with the dark-coloured cere.     

Sexual Cannibalism in the Brown Widow Spider (Latrodectus geometricus)

Sexual cannibalism may represent an extreme form of male monogamy. According to this view, males gain reproductive success by sacrificing themselves to females. We studied the occurrence and timing of sexual cannibalism in the brown widow spider Latrodectus geometricus and compared male courtship and mating behavior with virgin and with previously mated females. We found that events of sexual cannibalism are frequent, that they occur during copulation and that males initiate cannibalism by placing the abdomen in front of the female’s mouth‐parts during copulation (somersault behavior). Both the somersaults and mating occurred more frequently with virgins than with previously mated females. Our results support the hypothesis that sexual cannibalism is a male strategy in this species. The somersault behavior was previously known only from the redback spider, Latrodectus hasselti. It is as yet unknown whether self‐sacrifice has evolved more than once in this genus.     

Sex differences in biparental care as offspring develop: a field study of convict cichlids (<Emphasis Type="Italic">Amatitlania siquia</Emphasis>)

Parental investment theory states that parents should contribute more to older offspring. Differences between the sexes also influence how each parent contributes to offspring in biparental species. Here, we examined a naturally occurring population of biparental convict cichlids in Costa Rica to determine how each parent cared for offspring during two distinct offspring development stages. Consistent with the predictions of the reproductive value hypothesis, we hypothesized that the levels of parental contribution would be relative to the value that each parent places on a brood. We predicted that female parents would contribute more than male parents because female convict cichlids have lower future reproductive success than males. Additionally, we predicted that both parents should contribute more to older offspring, either due to the young’s increased susceptibility to predation (i.e., the vulnerability hypothesis) or because of the longer period of time parents have been interacting with older offspring (i.e., feedback hypotheses). This increase in investment by males should coincide with a change in the coordination of care between parents. Detailed observations of parental pairs in their natural habitat supported these predictions. Females contributed more to broods than males and were relatively unaffected by offspring age while males spent significantly more time with older, free-swimming fry. Additionally, males tended to leave younger offspring more than females did, and were more likely to do so consecutively with younger offspring. This suggests that the coordination of duties between parents changes as parental investment changes. Overall, these data support both the reproductive value and the vulnerability hypotheses, but not necessarily the feedback hypothesis.     

Sex-specific predation on a monogamous rat, Hypogeomys antimena (Muridae: Nesomyinae)

Sex-specific predation on adult individuals is often predicted by different behaviour in males and females resulting from different reproductive strategies and social systems. High predation pressure and the need for biparental care are considered a possible ecological basis for the evolution of monogamy, the most puzzling social system in mammals. In species where adults and offspring are vulnerable to the same predators, males and females may protect their offspring to different extents because of conflicting demands of investment in current and future offspring. I present the first empirical data on age- and sex-specific predation pressure by top predators on a monogamous rodent and the sex-specific behavioural responses of the prey species to different rates of predation. During an annual predation peak, only offspring and adult males were killed but no females. Whereas males and females travelled similar distances at night before the period of high predation on offspring, males moved further during this period. At the same time, males and females increased their distance from their offspring but males stayed closer to them than females. As a consequence, the distance between the members of the pair increased during the predation peak. The males' behaviour could lead to their encountering predators more frequently which would reduce survival prospects. The different behaviour of males and females provides empirical evidence that males invest in the welfare of current offspring at the cost of higher predation risk whereas females protect their residual reproductive value. Copyright 2000 The Association for the Study of Animal Behaviour.