Post-reproductive Females Research Articles

Reproductive aggression and nestmate recognition in a subsocial bee

In social contexts, the ability to recognize and discriminate among individuals is advantageous, because it allows individuals to adjust their behaviour so as to enhance both individual and group fitness. Eusocial insects have finely developed mechanisms of discrimination that promote many kinds of social interactions, but discrimination may also be adaptive in noneusocial species, including solitary ones. Physical traits such as reproductive status influence rates of aggression and discrimination, permitting individuals to share common resources and nesting sites and to cooperatively care for offspring, while excluding potential aggressors or social parasites. In this study, we examined reproductive aggression and nestmate recognition in a subsocial species of small carpenter bee, Ceratina calcarata, using circle tube behavioural assays. Not only does this subsocial bee show nestmate recognition, but there is seasonal variation in aggression that correlates with seasonal variation in reproductive status, illustrating that both aggressive behaviour and the consequences of nestmate recognition are context dependent. Females that were actively reproductive (ovaries fully developed) were more aggressive than pre-reproductive (ovaries undeveloped) or post-reproductive females (ovaries resorbed). Females altered their behaviour when interacting with nestmates versus non-nestmates. As in most social Hymenoptera, agonistic behaviour was observed to be greatest between unfamiliar, reproductively active individuals. However, post-reproductive females were tolerant towards unfamiliar females. During the natural adult cohabitation phase of the nesting cycle (the mature brood phase), mothers were aggressive towards daughters, whereas same generation pairs of nestmates or non-nestmates showed no signs of aggression. These results indicate that this subsocial bee species does possess the ability to recognize nestmates but the consequences of recognition vary seasonally, sometimes resulting in greater aggression towards nestmates than towards non-nestmates.

Structural and Performance Costs of Reproduction in a Pure Capital Breeder, the Children’s PythonAntaresia childreni

Females often manage the high energy demands associated with reproduction by accumulating and storing energy in the form of fat before initiating their reproductive effort. However, fat stores cannot satisfy all reproductive resource demands, which include considerable investment of amino acids (e.g., for the production of yolk proteins or gluconeogenesis). Because capital breeders generally do not eat during reproduction, these amino acids must come from internal resources, typically muscle proteins. Although the energetic costs of reproduction have been fairly well studied, there are limited data on structural and performance costs associated with the muscle degradation required to meet amino acid demands. Thus, we examined structural changes (epaxial muscle width) and performance costs (constriction and strength) over the course of reproduction in a pure capital breeder, the children's python (Antaresia childreni). We found that both egg production (i.e., direct resource allocation) and maternal care (egg brooding) induce muscle catabolism and affect performance of the female. Although epaxial muscle loss was minimal in nonreproductive females, it reached up to 22% (in females after oviposition) and 34% (in females after brooding) of initial muscle width. Interestingly, we found that individuals with higher initial muscular condition allocated more of their muscle into reproduction. The amount of muscle loss was significantly linked to clutch mass, underscoring the role of structural protein in egg production. Egg brooding significantly increased proteolysis and epaxial loss despite no direct allocation to the offspring. Muscle loss was linked to a significant reduction in performance in postreproductive females. Overall, these results demonstrate that capital-breeding females experience dramatic costs that consume structural resources and jeopardize performance.

Utilization of prey-rich patches leads to reproductive advantages for clustered individuals of a web-building spider

The ideal free distribution (IFD) model predicts that consumers match the distribution of resources across habitat patches and that this association should lead to equal individual fitness among individuals in a population. Here we studied to what extent predators utilize different prey groups and if the resulting spatial patterns relate to the reproductive success of individuals in a population of a web-building spider species (Phylloneta impressa Theridiidae). Web-building spiders, their prey remains, egg sacs, and spiderlings were collected in a wheat field, and coordinates of web sites were recorded for spatial point pattern analysis using pair and mark correlation functions. Spiders were aggregated in the study area, but only postreproductive females were aggregated over the full range of spatial scales. Web-owners in clusters caught a higher proportion of hemipteran prey than expected by the mean, suggesting that the availability of this prey type contributed to the clustering of individuals and the higher reproductive success of females in clusters. However, 2 abundant prey groups (Syrphidae and Hymenoptera) were not more common than expected by the mean at any distance, and another prey group (Elateridae) was even less common in webs close to each other. At least 2 prey groups (Hemiptera and Elateridae) showed deviations from the assumption of the IFD model that predators in clusters should have similar prey capture success compared to predators that are not located in clusters. Both prey groups include many cereal pest species, and our results suggest that spatially explicit information about prey utilization and predator fitness may contribute to a better understanding of the role of natural enemies in biological control.

A Papionin Multilevel Society as a Model for Hominin Social Evolution

Multilevel societies are unique in their ability to facilitate the maintenance of strong and consistent social bonds among some individuals while allowing separation among others, which may be especially important when social and sexual bonds carry significant and reliable benefits to individuals within social groups. Here we examine the importance of social and sexual bonds in the multilevel society of hamadryas baboons (Papio hamadryas) and apply these principles to social evolution in Plio-Pleistocene hominins. The behavior, adaptations, and socioecology of baboons (Papio spp.) have long been recognized as providing an important comparative sample to elucidate the processes of human evolution, and the social system of hamadryas baboons in particular shares even more similarities with humans than that of other baboons. Here we draw parallels between processes during the evolution of hamadryas social organization and those characterizing late Pliocene or early Pleistocene hominins, most likely Homo erectus. The higher costs of reproduction faced by female Homo erectus, exacerbated by an increased reliance on difficult to acquire, nutrient-dense foods, are commonly thought to have been alleviated by a strengthening of male–female bonds (via male provisioning and the evolution of monogamy) or by the assistance of older, postreproductive females (via grandmothering). We suggest that both of these social arrangements could have been present in Plio-Pleistocene hominins if we assume the development of a multilevel society such as that in hamadryas baboons. The evolution of a multilevel society thus underlies the adaptive potential for the complexity that we see in modern human social organization.

Communal nesting, activity patterns, and population characteristics in the near-threatened monito del monte, Dromiciops gliroides

The “living fossil,” Dromiciops gliroides (monito del monte), is an endemic marsupial inhabiting the temperate rain forests of South America. It is cavity-dependent and faces high energetic costs associated with thermoregulation during the austral winter. Although D. gliroides is well known for seed dispersal in temperate rain forests, its ecology, behavior, and long-term population dynamics have received little attention. We monitored a population of D. gliroides and studied variation in abundance and density and seasonal changes in body mass and body condition index (BCI). In addition, we monitored activity and communal nesting with camera traps and nest boxes, respectively. Over 4 years we documented a mean population density of 26 (95% confidence interval = 19–32) individuals/ha. We found significantly greater body mass and BCI for females than for males, suggesting different energetic strategies during the prehibernation period. Animals were nocturnal and active until dawn. Communal nesting occurred during summer and early fall, but torpor by single individuals and small groups was increasingly frequent during winter. Communal nesting could be a key behavioral strategy affecting survival. However, given the greater frequency in warm seasons and groups composed of postreproductive females and juveniles, communal nesting might be more related to parental care associated with kin selection than to thermoregulation.

Live Long and Procreate

Reproduction Speaking in generalities, once an animal leaves its reproductive phase, death is around the corner. Exceptions to this rule can be seen in humans, where postreproductive females continue to assist offspring and often subsequent generations. Another exception seems to be the nematode

Altruistic Colony Defense by Menopausal Female Insects

Recent studies have suggested that an extended postreproductive life span, such as life after menopause in human females, will evolve when the indirect (kin-selected) fitness benefits from altruistic behavior are greater than the direct fitness benefits from continuing reproduction. Under some conditions in which postreproductive altruism is more beneficial and/or continuing reproduction is more costly, the postreproductive life span can be shaped by natural selection. However, indirect fitness benefits during postreproductive survival have been documented mainly in intelligent mammals such as humans and cetaceans, in which elder females possess enhanced social knowledge through learning. Here we show that postreproductive females of the gall-forming aphid Quadrartus yoshinomiyai (Nipponaphidini) can gain indirect fitness benefits through their altruistic colony defense. These females cease reproduction around the time of gall opening and defend the colony by sticking themselves to intruding predators with a waxy secretion that is accumulated in their body with aging. Our results suggest that the presence of an age-related trait for altruistic behavior promotes the evolution of postreproductive altruism in this social insect via kin selection under natural selection imposed by predators.

THE IMPACT OF MATERNAL BIRTH MONTH ON REPRODUCTIVE PERFORMANCE: CONTROLLING FOR SOCIO-DEMOGRAPHIC CONFOUNDERS

Based on a 1900 census sample of 34,166 post-reproductive females (> or =45 years), the birth month effect was put to the test, for both lifetime fertility as well as child survival, controlling for maternal birth cohort (1826-1835, 1836-1845, 1846-1855), Duncan's SEI, urbanity, nativity, literacy and marital duration. Testing for potential cohort effects did not indicate a temporal trend in fertility by maternal birth month (seasonal Mann-Kendall test, p=0.578), while a minute increase in offspring survival was detected (p<0.001, Sen's estimator of slope=0.02, 95% CI=0.02 to 0.03). Further analyses of the maternal birth month effect on child survival were therefore seized. For lifetime fertility, ANOVA results indicated that maternal birth month was a major predictor for total offspring count (F11, 33606=1809.0, p<0.001), accounting for 37.2% of the total variability. In addition to main effects, a statistically significant interaction effect was observed (F538, 33606=2.2, p<0.001), with a corresponding effect size of eta2=0.40. Planned contrasts revealed that birth-month-specific differences in fertility achieved statistical significance (F11, 31798=1712.9, p<0.001), while post-hoc multiple comparisons for literacy and nativity displayed an inverse relationship with fertility, which meets demographic expectations. Controlling for all factors of interest, models of cohort-specific offspring counts (independent ANOVAs for 1826-1835: F157, 3467=26.3, p<0.001; 1836-1845: F182, 10299=75.5, p<0.001; 1846-1855: F199, 19859=137.9, p<0.001) indicated that women born in the first half of the year (particularly, January, February, April and May) achieved above-average parity, while those born in the latter half (namely, July, October, November and December) displayed markedly lower fertility averages. These monthly disparities are in line with previous observations and appear to be linked to seasonal optimal ripening of the oocyte or seasonal preovulatory over-ripeness ovopathy (Jongbloet, 1992).

Cooperative breeding in South American hunter-gatherers.

Evolutionary researchers have recently suggested that pre-modern human societies habitually practised cooperative breeding and that this feature helps explain human prosocial tendencies. Despite circumstantial evidence that post-reproductive females and extra-pair males both provide resources required for successful reproduction by mated pairs, no study has yet provided details about the flow of food resources by different age and sex categories to breeders and offspring, nor documented the ratio of helpers to breeders. Here, we show in two hunter-gatherer societies of South America that each breeding pair with dependent offspring on average obtained help from approximately 1.3 non-reproductive adults. Young married males and unmarried males of all ages were the main food providers, accounting for 93-100% of all excess food production available to breeding pairs and their offspring. Thus, each breeding pair with dependants was provisioned on average by 0.8 adult male helpers. The data provide no support for the hypothesis that post-reproductive females are the main provisioners of younger reproductive-aged kin in hunter-gatherer societies. Demographic and food acquisition data show that most breeding pairs can expect food deficits owing to foraging luck, health disabilities and accumulating dependency ratio of offspring in middle age, and that extra-pair provisioning may be essential to the evolved human life history.

Introduction of agriculture and its effects on women's oral health

This study explores the dynamic relationship between the introduction of agriculture and its effects on women's oral health by testing the hypothesis that female reproductive physiology contributes to an oral environment more susceptible to chronic oral disease and that, in a population undergoing the foraging to farming transition, females will exhibit a higher prevalence of oral pathology than males. This is tested by comparing the presence, location, and severity of caries lesions and antemortem tooth loss across groups of reproductive aged and postreproductive females (n = 71) against corresponding groups of males (n = 71) in an Early Agricultural period (1600 B.C.-A.D. 200) skeletal sample from northwest Mexico. Caries rates did not differ by sex across age groups in the sample; however, females were found to exhibit significantly more antemortem tooth loss than males (P > 0.01). Differences were initially minimal but increased by age cohort until postreproductive females experienced a considerable amount of tooth loss, during a life stage when the accumulation of bodily insults likely contributed to dental exfoliation. Higher caries rates in females are often cited as the result of gender differences and dietary disparities in agricultural communities. In an early farming community, with diets being relatively equal, women were found to experience similar caries expression but greater tooth loss. We believe this differential pattern of oral pathology provides new evidence in support of the interpretation that women's oral health is impacted by effects relating to reproductive biology.

The role of menopause and reproductive senescence in a long-lived social mammal

BackgroundMenopause is a seemingly maladaptive life-history trait that is found in many long-lived mammals. There are two competing evolutionary hypotheses for this phenomenon; in the adaptive view of menopause, the cessation of reproduction may increase the fitness of older females; in the non-adaptive view, menopause may be explained by physiological deterioration with age. The decline and eventual cessation of reproduction has been documented in a number of mammalian species, however the evolutionary cause of this trait is unknown.ResultsWe examined a unique 30-year time series of killer whales, tracking the reproductive performance of individuals through time. Killer whales are extremely long-lived, and may have the longest documented post-reproductive lifespan of any mammal, including humans. We found no strong support for either of the adaptive hypotheses of menopause; there was little support for the presence of post-reproductive females benefitting their daughter's reproductive performance (interbirth interval and reproductive lifespan of daughters), or the number of mature recruits to the population. Oldest mothers (> 35) did appear to have a small positive impact on calf survival, suggesting that females may gain experience with age. There was mixed support for the grandmother hypothesis – grandoffspring survival probabilities were not influenced by living grandmothers, but grandmothers may positively influence survival of juveniles at a critical life stage.ConclusionAlthough existing data do not allow us to examine evolutionary tradeoffs between survival and reproduction for this species, we were able to examine the effect of maternal age on offspring survival. Our results are consistent with similar studies of other mammals – oldest mothers appear to be better mothers, producing calves with higher survival rates. Studies of juvenile survival in humans have reported positive benefits of grandmothers on newly weaned infants; our results indicate that 3-year old killer whales may experience a positive benefit from helpful grandmothers. While our research provides little support for menopause evolving to provide fitness benefits to mothers or grandmothers, our work supports previous research showing that menopause and long post-reproductive lifespans are not a human phenomenon.

Estradiol induces sexual behavior in female túngara frogs

Steroid hormones play an important role in regulating vertebrate sexual behavior. In frogs and toads, injections of exogenous gonadotropins, which stimulate steroid hormone production, are often used to induce reproductive behavior, but steroid hormones alone are not always sufficient. To determine which hormonal conditions promote sexual behavior in female túngara frogs, we assessed the effect of hormone manipulation on the probability of phonotaxis behavior toward conspecific calls in post-reproductive females. We injected females with human chorionic gonadotropin (HCG), estradiol, estradiol plus progesterone, saline, or HCG plus fadrozole (an aromatase blocker) and tested their responses to mating calls. We found that injections of HCG, estradiol, and estradiol plus progesterone all increased phonotaxis behavior, whereas injections of saline or HCG plus fadrozole did not. Since injections of estradiol alone were effective at increasing phonotaxis behavior, we concluded that estradiol is sufficient for the expression of phonotaxis behavior. Next, to determine if estradiol-injected females display the same behavioral preferences as naturally breeding females, we compared mating call preferences of naturally breeding females to those of post-reproductive females injected with estradiol. We found that, when injected with estradiol, females show similar call preferences as naturally breeding females, although they were less likely to respond across multiple phonotaxis tests. Overall, our results suggest that estradiol is sufficient for the expression of sexual responses to mating calls in túngara frogs. To our knowledge, ours is the only study to find that estradiol alone is capable of promoting phonotaxis behavior in a frog.

Exploring the origin of egg protein in an oviparous water snake (Natrix maura)

Reptiles are typical capital breeders that fuel reproduction by the use of lipids stored in fat bodies. However, little is known about the origin (exogenous or endogenous) of egg protein. We have examined the origin of egg protein by means of the analysis of protein content of liver and carcass (skeletal muscle) of the oviparous snake Natrix maura throughout an annual cycle. We have also measured monthly variation of the digestive-content mass and the ovarian mass. Results showed that protein in liver peaked during vitellogenesis according to the role of the liver in the synthesis of vitellogenin. Partial correlations showed the path of protein from the prey (digestive-content) to the liver, and finally to the ovaries, as well as an inverse relation between carcass protein and ovarian mass. Carcass muscle is the only body part that may act as a potential reserve for endogenous protein, although we did not find significant variation in carcass protein during female reproduction. As females with large follicles did not stop foraging activity, we assumed that egg protein was derived from the diet as partial correlations indicated. Our results suggest that N. maura is a capital breeder for lipids and tend to be income breeder for protein. This conclusion contrasts with that observed in capital breeders for which egg protein was derived from muscle. We discuss the idea that flexibility in the origin of egg protein could affect the body condition in post-reproductive females.

The Evolution of Senescence and Post-Reproductive Lifespan in Guppies (Poecilia reticulata)

The study of post-reproductive lifespan has been of interest primarily with regard to the extended post-menopausal lifespan seen in humans. This unusual feature of human demography has been hypothesized to have evolved because of the “grandmother” effect, or the contributions that post-reproductive females make to the fitness of their children and grandchildren. While some correlative analyses of human populations support this hypothesis, few formal, experimental studies have addressed the evolution of post-reproductive lifespan. As part of an ongoing study of life history evolution in guppies, we compared lifespans of individual guppies derived from populations that differ in their extrinsic mortality rates. Some of these populations co-occur with predators that increase mortality rate, whereas other nearby populations above barrier waterfalls are relatively free from predation. Theory predicts that such differences in extrinsic mortality will select for differences in the age at maturity, allocation of resources to reproduction, and patterns of senescence, including reproductive declines. As part of our evaluation of these predictions, we quantified differences among populations in post-reproductive lifespan. We present here the first formal, comparative study of the evolution of post-reproductive lifespan as a component of the evolution of the entire life history.Guppies that evolved with predators and that experienced high extrinsic mortality mature at an earlier age but also have longer lifespans. We divided the lifespan into three non-overlapping components: birth to age at first reproduction, age at first reproduction to age at last reproduction (reproductive lifespan), and age at last reproduction to age at death (post-reproductive lifespan). Guppies from high-predation environments live longer because they have a longer reproductive lifespan, which is the component of the life history that can make a direct contribution to individual fitness. We found no differences among populations in post-reproductive lifespan, which is as predicted since there can be no contribution of this segment of the life history to an individual's fitness.Prior work on the evolution of post-reproductive lifespan has been dominated by speculation and correlative analyses. We show here that this component of the life history is accessible to formal study as part of experiments that quantify the different segments of an individual's life history. Populations of guppies subject to different mortality pressures from predation evolved differences in total lifespan, but not in post-reproductive lifespan. Rather than showing the direct effects of selection characterizing other life-history traits, post-reproductive lifespan in these fish appears to be a random add-on at the end of the life history. These findings support the hypothesis that differences in lifespan evolving in response to selection are confined to the reproductive lifespan, or those segments of the life history that make a direct contribution to fitness. We also show, for the first time, that fish can have reproductive senescence and extended post-reproductive lifespans despite the general observation that they are capable of producing new primary oocytes throughout their lives.

Effect of captivity in semi-natural enclosures on the reproductive endocrinology of female lizards

Long-term captivity can result in abnormal behavior and physiology in many vertebrates. Here, we examine whether semi-natural, outdoor captive environments can offer a compromise, allowing much of the experimental control afforded by captivity, while providing the environmental conditions essential for normal behavior and physiology. We first determined plasma concentration of the sex steroid hormones progesterone, testosterone, and estradiol of free-ranging female striped plateau lizards ( Sceloporus virgatus) throughout the reproductive season, and second, compared the results to those from conspecific females maintained in semi-natural outdoor enclosures. Among free-ranging females, levels of progesterone and estradiol were elevated during vitellogenesis, with an apparent surge in progesterone and testosterone occurring in association with ovulation. Following ovulation, estradiol fell to non-reproductive levels while progesterone remained elevated during the month-long period of gravidity. Long-term captivity in outdoor enclosures did not significantly affect progesterone or estradiol levels at any stage of the reproductive cycle, and did not affect testosterone levels during normal ovarian development and gravidity. However, (1) females with delayed oviposition in captivity had lower testosterone levels than free-ranging females with normal oviposition, and (2) when all females sampled were post-reproductive, captive females continued to have lower testosterone levels than free-ranging females. Thus, the endocrine response to captivity may be greater among post-reproductive females than among reproductive females. Our results are in marked contrast to many studies that show captivity can dramatically impair reproduction of female vertebrates.

Availability and adaptive value of reproductive and postreproductive Japanese macaque mothers and grandmothers

Prior tests of the grandmother hypothesis have suggested that postreproductive female Japanese macaques,Macaca fuscata , do not significantly improve the survivorship of their descendents. However, not all postreproductive females are grandmothers, and not all grandmothers are postreproductive. In this study we looked at the daughters and grandchildren of 70 female Japanese macaques to assess the availability and adaptive value of reproductive and postreproductive mothers and grandmothers. We found that postreproductive Japanese macaque grandmothers are very rare. Only 2.8% of the sample of daughters had a postreproductive grandmother available when they began to reproduce, and only 4.2% of the reproductive life span of the daughter (less than 6 months for the average female) was spent with a postreproductive grandmother available to help the daughter. Grandchildren who survived to age 5 had a postreproductive grandmother available to them for only 4% of the first 5 years of life (i.e. a couple of months on average). The presence of a living mother, irrespective of her reproductive status, was associated with improved reproduction in her adult daughters, and the presence of a postreproductive grandmother was associated with significantly improved survivorship of grandchildren to age 1. While improved maternal investment does not appear to be the primary explanation for reproductive termination in Japanese monkeys, the few postreproductive females that have unweaned grandchildren available appear to have a positive influence on their survival. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.

Hematology and clinical chemistry in dystocic and healthy post-reproductive female chameleons.

Prevalence of egg-laying disorders and hematology and blood chemical parameters were analyzed in free-ranging common chameleons (Chamaeleo chamaeleon) from southern Spain. During four reproductive seasons oviposition failures occurred only during dry years. Egg binding appeared to be an important cause of mortality. Oviductal eggs of dystocic females were longer and wider than those of females with normal oviposition. Blood cell counts, white blood cell differentials, hematocrit, and concentration of hemoglobin, total plasma protein, glucose, uric acid, aspartate aminotransferase, and creatinine phosphokinase in blood were determined for eight apparently healthy post-reproductive females and considered as reference values. These chameleons differed from other reptiles in high concentrations of glucose in blood, averaging 362 mg/dl, and in high numbers of heterophils. Females with dystocia differed from reference values in an increase in monocytes and in the high concentrations of aspartate aminotransferase probably associated with tissue trauma.

Is There Adaptive Value to Reproductive Termination in Japanese Macaques? A Test of Maternal Investment Hypotheses

Evolutionary biologists often argue that menopause evolved in the human female as the result of selection for a postreproductive phase of life, during which increased maternal investment in existing progeny could lead to enhanced survivorship of descendents. Adaptive theories relating menopause to enhanced maternal investment are known as the mother (first-generation) and grandmother (second-generation-offspring) hypotheses. Although menopause—universal midlife termination of reproduction—has not been documented in primates other than humans, some researchers have argued that postreproductive alloprimates also have a positive impact on the survivorship of first and second generation progeny. We tested the maternal investment hypotheses in Japanese macaques by comparing the survivorship of offspring, final infants, and great-offspring of females that terminated reproduction before death with females that continued to reproduce until death. SURVIVAL analyses revealed no significant difference in the survivorship of descendents of postreproductive and reproductive females, though final infants of postreproductive females were 13% more likely to survive than final infants of females that reproduced until death were. We also explored possible differences between these two groups of females, other than survivorship of progeny. We found no difference in dominance rank, matrilineal affiliation, body weight, infant sex ratio, age at first birth, fecundity rate or lifetime reproductive success. However, postreproductive females are significantly longer-lived than reproductive females and as a result experienced more years of reproduction and produced more infants in total. Apart from final infants, offspring survival is marginally lower in postreproductive females. Since offspring survival is not significantly enhanced in postreproductive females, the greater number of infants produced did not translate into greater lifetime reproductive success. Our findings fail to support the maternal investment hypotheses and instead suggest that reproductive termination in this population of Japanese macaques is most closely associated with enhanced longevity and its repercussions.

Reproductive cessation in female mammals.

In female mammals, fertility declines abruptly at an advanced age. The human menopause is one example, but reproductive cessation has also been documented in non-human primates, rodents, whales, dogs, rabbits, elephants and domestic livestock. The human menopause has been considered an evolutionary adaptation, assuming that elderly women avoid the increasing complications of continued childbirth to better nurture their current children and grandchildren. But an abrupt reproductive decline might be only a non-adaptive by-product of life-history patterns. Because so many individuals die from starvation, disease and predation, detrimental genetic traits can persist (or even be favoured) as long as their deleterious effects are delayed until an advanced age is reached, and, for a given pattern of mortality, there should be an age by which selection would be too weak to prevent the onset of reproductive senescence. We provide a systematic test of these alternatives using field data from two species in which grandmothers frequently engage in kin-directed behaviour. Both species show abrupt age-specific changes in reproductive performance that are characteristic of menopause. But elderly females do not suffer increased mortality costs of reproduction, nor do post-reproductive females enhance the fitness of grandchildren or older children. Instead, reproductive cessation appears to result from senescence.

The behavior of postreproductive females in a wild population of toque macaques (Macaca sinica) in Sri Lanka

I compared the behavior of three old postreproductive females in a wild population of toque macaques (Macaca sinica)in Polonnaruwa with those of reproductive females via focal-animal sampling techniques. Postreproductives foraged less, slept more, and were less active overall than reproductive females were. They also had significantly lower rates of agonistic behavior, were more peripheral, and had lower frequencies of overall affiliative contact. Although postreproductives initiated contact with others as frequently as reproductives did, group members initiated contact with them significantly less than they did with reproductive females. Postreproductives associated more with adult females than reproductives did and less with adult and subadult males than high-ranking reproductives did. Juvenile and infant females associated more frequently with reproductive females of high or low rank than with postreproductives. Postreproductives resembled low-ranking reproductive females in giving less grooming to others than they received. This contrasts with high-ranking females, which gave more grooming to others than they received. The results suggest that old age and cessation of reproduction are evident through the manifestation of distinct behavioral characteristics in toque macaque females.