Elevated urinary testosterone excretion and decreased maternal caregiving effort in marmosets when conception occurs during the period of infant dependence

The proximate mechanisms that regulate transitions in mammalian female reproductive effort have not been widely studied. However, variation in circulating levels of the androgenic steroid hormone testosterone (T) appears to mediate a trade-off between investment in current and future offspring in males [Ketterson, E.D., Nolan, V., Jr., 1992. Hormones and life histories: an integrative approach. Am. Nat. 140, S33-S62; Ketterson, E.D., Nolan, V., Jr., 1994. Hormones and life histories: an integrative approach. In: L.A. Real (Ed.), Behavioral Mechanisms in Evolutionary Ecology, University of Chicago Press, Chicago, pp. 327-353; Ketterson, E. D., Nolan, V., Jr., 1999. Adaptation, exaptation, and constraint: A hormonal perspective. Am. Nat. 154S, S4-S25]. The purpose of this study was to investigate the possibility that T is also associated with transitions in the reproductive effort of females, by examining the relationship between urinary T excretion, maternal caregiving behavior, and the timing of the postpartum conception in female Wied's black tufted-ear marmosets (Callithrix kuhlii). We examined the maternal carrying effort and peripartum T profiles of six females across two conditions: (1) when they conceived during the period of infant dependence (DPID), such that gestation was coupled with lactation; and (2) when the same females conceived after the period of infant dependence (APID). We also assessed the relationship between postpartum T levels and caregiving effort. When female marmosets conceived DPID, they dramatically reduced their caregiving effort, and had higher levels of urinary T, relative to when they conceived APID. Further, the litter-to-litter changes in maternal caregiving effort that we observed were related to variation in urinary T excretion; as weekly levels of urinary T excretion increased, concurrent caregiving effort declined. Our results suggest that variation in T secretion may regulate transitions in female reproductive behavior, and that the regulation of male and female parental behavior may be mediated by homologous neuroendocrine mechanisms.     

Infant Carrying Behavior in Callitrichid Primates: Callithrix and Leontopithecus

Care of offspring by individuals other than the mother is ubiquitous in callitrichid primates. In spite of its widespread occurrence, however, there is considerable taxonomic variation in the timing and intensity of parental effort by breeding males and nonbreeding juvenile group members. These differences may be attributable to generic and specifies differences in the costs of reproduction for females or in ecological constraints on travel and foraging. We present data on patterns of infant carrying in social groups of two taxa of callitrichid primates (Callithrix and Leontopithecus) throughout the first 3 months of infant life. We evaluated patterns of care in small groups (two or fewer juvenile or subadult helpers) and in large groups (three or more helpers in addition to the breeding adult male and female). Group size had little effect on levels of maternal carrying effort in either marmosets or lion tamarins, and mothers ceased carrying infants by 3 months of age. Carrying efforts by fathers were significantly reduced in groups with many helpers relative to small groups. Helpers carried at consistent rates during the second and third months of infant life in Leontopithecus, while in Callithrix, carrying by helpers peaked during the second month. These results suggest that if helpers reduce energetic demands on lactating females, the mechanism by which helpers reduce these burdens is independent of maternal carrying effort.     

Pre- and postpartum sex steroids in female marmosets (Callithrix kuhlii): is there a link with infant survivorship and maternal behavior?

Recent studies in primates have suggested that pre- and peripartum sex steroid hormones may be important determinants of maternal behavior and motivation, since higher levels of prepartum estrogen are associated with maternal competency and infant survivorship. To test the cross-species generality of this finding, we monitored excreted profiles of estradiol (E(2)), progesterone metabolites (pregnanediol glucuronide; PdG), and E(2):PdG ratios throughout pregnancy in black tufted-ear marmosets (Callithrix kuhlii). Additionally, we wanted to determine the extent to which variability in maternal carrying effort was related to hormonal factors and relative levels of maternal experience. For six females, hormonal profiles were determined by enzyme immunoassay for two pregnancies, one in which infants survived at least 2 weeks postpartum and one in which infants did not survive. Our within-subjects analyses revealed significant differences in mean prepartum E(2) levels for females in the different infant survival conditions. In contrast to previous findings, however, females exhibited significantly higher prepartum E(2) levels when their infants did not survive a minimum of 2 weeks postpartum, relative to when their infants did survive. Maternal carrying effort was also negatively and significantly correlated with prepartum E(2) levels. Unlike previous reports in callitrichid primates, then, our data suggest that high concentrations of prepartum E(2) in callitrichid primates are not necessarily associated with competent maternal behavior and may instead be associated with poor infant survivorship and inadequate maternal care. Further, our results appear to be convergent with research focusing on human mothers and may represent a common underlying mechanism linking prepartum estrogen and postpartum affect and behavior in some primates.     

Hormonal changes related to paternal and alloparental care in common marmosets (Callithrix jacchus)

The physiological mechanisms of parental and alloparental care in cooperatively breeding nonhuman primate species such as the common marmoset (Callithrix jacchus) are poorly known. In this study, we examined prolactin and cortisol plasma levels of fathers and older offspring of both sexes, with and without previous experience in infant carrying, around parturition and during infant carrying. Blood samples were collected from fathers and older offspring and prolactin and cortisol were measured by RIA and EIA, respectively. Prolactin levels of both caretakers were not influenced by infant's birth, previous experience or proximity to parturition. However, prolactin levels increased in both caretakers while in physical contact with infants and also with the number of infants being carried in older offspring. These findings suggest that increased prolactin seems to be mainly due to physical effort rather than a physiological trigger of paternal and alloparental care in common marmosets. Cortisol levels were higher for experienced fathers shortly before parturition which could act to reinforce affiliative bonds between breeding males and females at this time or in the ability of males to detect the proximity of the parturition or both.     

Nighttime Wakefulness Associated with Infant Rearing in Callithrix kuhlii

Parent-infant cosleeping occurs in human and nonhuman primates, yet studies on the impact of cosleeping on parental sleep patterns have been limited to human mothers. We examined the effects of cosleeping on the nighttime wakefulness of a biparental New World primate, Wied's black tufted-ear marmoset (Callithrix kuhlii). We compared the sleep patterns of marmoset parents caring for young infants to those without infants, using an 8 mm videocamera and timelapse VCR under infrared illumination. The presence of young infants significantly impacted the sleep of mothers but not fathers. In fact, mothers rearing young infants were awake >3 times as often as mothers without infants. We also examined the nighttime wakefulness of marmoset parents across the first 9 weeks of infant life (birth through weaning). Although callitrichid mothers tend to reduce their daytime investment in offspring very early in infant life by relinquishing the care of infants to fathers and alloparents, increased nighttime wakefulness was not limited to the early postpartum period for the mothers. Instead, mothers exhibited more nighttime wakefulness than fathers did across the first 9 weeks of infant life. Our results indicate that the presence of infants has a greater impact on the sleep patterns of Callithrix kuhlii mothers than fathers, suggesting that mothers are more involved in infant care than previously realized and that fathers are not nearly as involved in nighttime care as their behavior during the day would suggest.     

Alloparents in the Mongolian gerbil: impact on long-term reproductive performance of breeders and opportunities for independent reproduction

The presence of alloparents in cooperatively breeding groupsmay enhance the reproductive performance of the breeding pairbut tests of this hypothesis in mammalian cooperative breedershave typically examined short periods in the reproductive lifespan of breeders. The present experiment was designed to evaluatethe long-term effects of the presence of alloparents on reproductiveperformance under laboratory conditions in Mongolian gerbils(Mertones unguiculalus), and to evaluate the possibility thatalloparents may attempt independent reproduction during theirtenure as alloparents. Pairs were established with either 0or 4 alloparents and breeding performance was monitored overa 13.5-month period. The presence of alloparents did not enhancereproductive rates in breeding pairs, and neither enhanced nordelayed developmental rates in offspring. Variation in littersize, offspring survival, pup growth, and interbirth intervaldid not vary significantly as a function of the breeding females'reproductive tenure. Seven of 49 female alloparents of breedingage in family groups produced offspring. Alloparent breedingwas associated with long interbirth intervals in the breedingadult female. Neonatal survival in litters born to alloparentswas higher in litters that were temporally synchronized withlitters produced by the older breeding female than in asynchronouslyproduced litters. Under the conditions in this experiment, then,few indirect benefits accrue to helpers as a consequence oftheir alloparental effort, and immediate (the opportunity forindependent reproductive attempts) and delayed (parental skillsacquisition) direct benefits may account for helping behaviorin this species.     

Diminished maternal responsiveness during pregnancy in multiparous female common marmosets

Common marmoset monkeys (Callithrix jacchus) live in small groups in which, usually, only a single female breeds and all group members provide infant care. When two females breed concurrently, however, they may commonly kill one another's infants, especially during the peripartum period. To investigate the mechanisms underlying infanticide by breeding females, we characterized responses of multiparous females to infants and determined circulating hormone levels in adult females during early pregnancy, late pregnancy, and the early postpartum period. Additionally, we compared the responses of postpartum females to their own infants and infants of other females (unfamiliar infants). Postpartum females were highly maternal toward both their own and unfamiliar infants, and showed no differences in their behavioral or hormonal responses to the two. During both early and late pregnancy, however, these females exhibited longer latencies to initially approach unfamiliar infants and spent less time carrying unfamiliar infants. Moreover, females spent less time carrying unfamiliar infants during late pregnancy than early pregnancy. Most late pregnant females never carried infants, and those that did rejected them quickly. Prolactin concentrations were higher and progesterone concentrations lower postpartum than in early or late pregnancy, while estradiol concentrations, the estradiol-to-progesterone ratio, and cortisol levels were higher during late pregnancy. Within reproductive conditions, however, maternal behaviors were not correlated with hormone levels. These results suggest that maternal responsiveness in marmosets may be attenuated during pregnancy, especially late pregnancy, and this may contribute to infanticide by breeding females.     

What is cooperative breeding in mammals and birds? Removing definitional barriers for comparative research

Cooperative breeding (i.e. when alloparents care for the offspring of other group members) has been studied for nearly a century. Yet, inconsistent definitions of this breeding system still hamper comparative research. Here, we identify two major inconsistencies, discuss their consequences and propose a way forward. First, some researchers restrict the term ‘cooperative breeding’ to species with non-breeding alloparents. We show that such restrictive definitions lack distinct quantitative criteria to define non-breeding alloparents. This ambiguity, we argue, reflects the reproductive-sharing continuum among cooperatively breeding species. We therefore suggest that cooperative breeding should not be restricted to the few species with extreme reproductive skew and should be defined independent of the reproductive status of alloparents. Second, definitions rarely specify the type, extent and prevalence of alloparental care required to classify species as cooperative breeders. We thus analysed published data to propose qualitative and quantitative criteria for alloparental care. We conclude by proposing the following operational definition: cooperative breeding is a reproductive system where >5% of broods/litters in at least one population receive species-typical parental care and conspecifics provide proactive alloparental care that fulfils >5% of at least one type of the offspring's needs. This operational definition is designed to increase comparability across species and disciplines while allowing to study the intriguing phenomenon of cooperative breeding as a behaviour with multiple dimensions.     

A comparative perspective on the evolution of tamarin and marmoset social systems

Tamarins and marmosets (callitrichids) present an unusual opportunity for study of the determinants of primate social systems, because both the mating and infant care patterns of callitrichids are variable, even within individual populations. In this paper, I briefly describe three characteristics of callitrichid social systems that distinguish them from most other primates: extensive male parental care, helping by nonreproductive individuals, and variable mating patterns. I then discuss the evolution of these characteristics and of the frequent twinning exhibited by callitrichids. I suggest that an ancestor of modern callitrichids gave birth to a single offspring at a time, mated monogamously, and had significant paternal care. The idea that males of this ancestral form must have provided paternal care, even though only single infants were born, derives from a comparison of litter/mother weight ratios in modern primate species. Twinning perhaps then evolved because of a combination of dwarfing in the callitrichid lineage, leading to higher litter/mother weight ratios, and a high infant mortality rate, and because the extensive paternal care already present facilitated the raising of twins. I propose that the helping behavior of older offspring may have coevolved with twinning, because helpers would have increased the chances of survival of twins, and the presence of twins would have increased the benefits of helping. Finally, the high costs of raising twins and the variability of group compositions, especially the fact that some groups would not have had older offspring to serve as helpers, may have selected for facultative polyandry in saddle-back tamarins (Saguinus fuscicollis) and perhaps in other callitrichid species. Both helping and cooperative polyandry have been extensively studied in bird species, and I apply some of the conclusions of these studies to the discussion of the evolution of callitrichid social systems.     

The effects of residential locality on parental and alloparental investment among the Aka foragers of the central African Republic

In this paper I examine the intracultural variability of parental and alloparental caregiving among the Aka foragers of the Central African Republic. It has been suggested that maternal kin offer higher frequencies of allocare than paternal kin and that maternal investment in infants will decrease when alloparental assistance is provided. Behavioral observations were conducted on 15 eight- to twelve-monthold infants. The practice of brideservice and the flexibility of Aka residence patterns offered a means to test the effect of maternal residence on parental and alloparental investment. There was significant variation in the frequency of investment and who supplied care to infants depending on whether mothers resided with their kin or their husbands’ kin. However, in spite of the variation in allocare, when all categories of caregivers were examined collectively, infants received similar overall levels of care. Courtney L. Meehan is a Ph.D. student in cultural anthropology at Washington State University. Her research interests include parenting, alloparenting, female social networks, and female-female cooperation and competition.     

Direct Male Care and Hominin Evolution: Why Male–Child Interaction Is More Than a Nice Social Idea

ABSTRACT  Early members of the genus Homo experienced heightened absolute metabolic costs, partially owing to increases in body size. However, as is characteristic of modern humans, they also likely began reproducing with shortened interbirth intervals. Male investment in offspring may help explain how this life history shift occurred. Evolutionary models of hominin male investment in offspring have traditionally focused on provisioning of females and young, yet the extent to which direct male care of offspring was evolutionarily important, from an energetic perspective, is largely unaddressed. I propose an evolutionary model of direct male care, demonstrating that males could have helped reduce the energetic burden of caregiving placed on mothers by carrying young. In doing so, males would have assisted females in achieving and maintaining an energetic condition sufficient for reproduction, thereby hastening the advent of shortened interbirth intervals that played a formative role in the success of our genus.     

Different ovarian responses to potential mates underlie species-specific breeding strategies in common marmoset and Goeldi's monkey

Callitrichids are cooperative breeders, characterized by obligate twinning, extensive paternal care, and monopolization of reproduction by the dominant female. This is the case in the common marmoset, and in common marmoset groups of more than one adult female, subordinate females are typically acyclic consistent with infertility. However, one callitrichid, Goeldi's monkey, gives birth to singletons and exhibits low paternal care. Given these reproductive traits of Goeldi's monkey, we hypothesized that there would not be suppression of ovarian activity. To test this hypothesis, we applied non-invasive endocrine methods in a step-wise experiment with laboratory groups of both species. In each species, six pairs of sisters were studied alone, in visual contact with an unrelated male and in a polygynous trio with the male, and urine samples were collected for determination of oestrogen titres reflecting ovarian activity. Common marmoset sister pairs exhibited a marked difference in social status: during the study 5 of 6 dominant females conceived but only 1 of 6 subordinate females; the remaining 5 subordinates were acyclic at the end of the study, and instances of ovulation typically resulted in aggression. Goeldi's monkey sister pairs showed no status differences: in all pairs, however, both sisters exhibited a temporary cessation of ovarian cyclicity on trio formation, followed by ovulation and conception. We conclude that these marked differences in ovarian responses reflect the differences in inter-female competition for paternal caregiving resources. In common marmosets with high inter-female competition, suppression of ovulation functions to reduce aggression received by subordinate females; in Goeldi's monkey with low competition, temporary cessation of ovulation could facilitate female choice.     

Do maternal food deprivation and offspring predator cues interactively affect maternal effort in fish?

The state of the environment parents are exposed to during reproduction can either facilitate or impair their ability to take care of their young. Thus, the environmental conditions experienced by parents can have a transgenerational impact on offspring phenotype and survival. Parental energetic needs and the variance in offspring predation risk have both been recognized as important factors influencing the quality and amount of parental care, but surprisingly, they are rarely manipulated simultaneously to investigate how parents adjust care to these potentially conflicting demands. In the maternally mouthbrooding cichlid Simochromis pleurospilus, we manipulated female body condition before spawning and exposure to offspring predator cues during brood care in a two‐by‐two factorial experiment. Subsequently, we measured the duration of brood care and the number and size of the released young. Furthermore, we stimulated females to take up their young by staged predator attacks and recorded the time before the young were released again. We found that food‐deprived females produced smaller young and engaged less in brood care behaviour than well‐nourished females. Final brood size and, related to this, female protective behaviour were interactively determined by nutritional state and predator exposure: well‐nourished females without a predator encounter had smaller broods than all other females and at the same time were least likely to take up their young after a simulated predator attack. We discuss several mechanisms by which predator exposure and maternal nutrition might have influenced brood and offspring size. Our results highlight the importance to investigate the selective forces on parents and offspring in combination, if we aim to understand reproductive strategies.     

A Prisoner's Dilemma model of the evolution of paternal care

The heavy energetic demands of gestation, lactation and rearing of offspring mean that studies of paternal care in primates usually focus on female reproductive effort. Here it is shown that both male and female reproductive effort must be considered in order to understand how paternal care evolved. This is done using the Prisoner's Dilemma, best known as a model of reciprocal altruism. It is found that the relative cost of reproduction for males and females is crucially important in determining co-operative and competitive strategies. In particular, when male reproductive costs are less than female reproductive costs, males co-operate with females even when females do not reciprocate. This surprising behaviour, termed non-reciprocal altruism, is comparable with male investment in a female and her offspring.     

A theoretical study on the evolution of male parental care and female multiple mating: effects of female mate choice and male care bias

Male parental care and female multiple mating are seen in many species in spite of the cost they entail. Moreover, they even coexist in some species though polyandry, by reducing paternity confidence of caregiving males, seems to hinder the evolution of paternal care. Previous studies have investigated the coevolutionary process of paternal care and polyandry under various simplifying assumptions, including random mating and random provision of male care. We extend these models to examine possible effects of female mate choice and male care bias, assuming that (a) monandrous females mate preferentially with caregiving males while polyandrous females compromise their preference in order to mate with multiple males and (b) caregiving males tend to direct their care to offspring of monandrous females. Our models suggest that both the female preference and the male bias always favor caregiving males while they may not always facilitate the evolution of monandry.     

Male burying beetles extend,not reduce,parental care duration when reproductive competition is high

Male parents spend less time caring than females in many species with biparental care. The traditional explanation for this pattern is that males have lower confidence of parentage, so they desert earlier in favour of pursuing other mating opportunities. However, one recent alternative hypothesis is that prolonged male parental care might also evolve if staying to care actively improves paternity. If this is the case, an increase in reproductive competition should be associated with increased paternal care. To test this prediction, we manipulated the level of reproductive competition experienced by burying beetles, Nicrophorus vespilloides (Herbst, 1783). We found that caregiving males stayed for longer and mated more frequently with their partner when reproductive competition was greater. Reproductive productivity did not increase when males extended care. Our findings provide support for the increased paternity hypothesis. Extended duration of parental care may be a male tactic both protecting investment (in the current brood) and maximizing paternity (in subsequent brood(s) via female stored sperm) even if this fails to maximize current reproductive productivity and creates conflict of interest with their mate via costs associated with increased mating frequency.     

Alloparental Care in the African Striped Mouse Rhabdomys pumilio is Age‐Dependent and Influences the Development of Paternal Care

Alloparents contribute to offspring care and alleviate the workload of breeders. The help provided varies with the age and/or experience of helpers, but it is not known whether breeders vary their investment based on the age of helpers by adjusting the parental care they provide. We studied the alloparental care provided by juvenile and subadult philopatric daughters in biparental African striped mice Rhabdomys pumilio with and without the mother as a measure of alleviation of maternal workload. We showed in a previous study that alleviation of maternal workload directly affects the development of paternal care in their sons, so we studied the expression of paternal care in young males raised by helpers as a proxy of the long‐term consequences of helping. Both juvenile and subadult daughters provided care but the level of alloparental care and concomitant alleviation of maternal care was age‐dependent. In the absence of the mother, juvenile daughters provided just 6% of care compared with 24% of subadult daughters. Sons raised by mothers and juvenile helpers displayed the expected exaggerated levels of care also observed when mothers raise litters on their own. While our results show the direct value of subadult daughters, juvenile daughters could contribute indirectly (e.g. nest maintenance) to alleviating maternal workload. The development of paternal care indicates that mothers do distinguish between the care provided by different aged helpers. Overall, the type of alloparental care provided by female striped mice is expected to change over their lifetimes, resulting in increased inclusive fitness through caring for siblings and acquisition of parenting skills.     

Corticosterone Levels Correlate With Alloparental Care in a Sex‐Dependent Manner in African Striped Mice,Rhabdomys pumilio

Alloparental care of non‐breeders is the main characteristic of cooperatively breeding species. While many studies have contributed to the understanding of the evolutionary reasons why individuals provide care to young that are not their own offspring, the variables influencing and causing alloparental care are less understood. We tested in African striped mice (Rhabdomys pumilio) whether age, sex, testosterone and corticosterone were correlated with alloparental care of non‐breeding helpers. We studied 11 family groups under controlled conditions in the laboratory, each with two juvenile and two adult helpers, one being male and one being female in each age category. We predicted male helpers to show more alloparental care than female helpers, as males are the dispersing sex and might thus have to pay for staying. We also expected adult helpers to show more alloparental care than juvenile helpers and both corticosterone and testosterone to correlate negatively with alloparental care. We found high levels of alloparental care in non‐breeding striped mice, which spent a significant amount of time in the nest, huddling and licking pups. There was neither a difference between the sexes nor between age categories (although both factors were significant in interaction terms), indicating either low costs and/or high benefits of alloparental care. Mothers showed significantly more care than helpers, and fathers showed similar levels of parental care as mothers but not significantly more than helpers. Although testosterone levels differed significantly between helpers of different age and sex, with adult male helpers showing the highest levels, we did not find any relationships between testosterone and the amount of alloparental care. Corticosterone levels were negatively correlated with alloparental care, and these effects were modulated by the sex and the age of helpers. In females, less alloparental care was shown with increasing corticosterone levels, while in males, the relationship was positive. Also, younger individuals with lower corticosterone levels showed more alloparental care than older individuals with low corticosterone levels. In sum, alloparental care is well developed in male and female non‐breeding helpers of striped mice, both in adult and juvenile helpers, but independently of testosterone, with corticosterone showing an age‐ and sex‐specific relationship with alloparental care.     

Irreversible transfer of brood care duties and insights into the burden of caregiving in incipient subterranean termite colonies

1. In incipient termite colonies, biparental brood care rapidly shifts towards alloparental brood care. This transition was suggested to recapitulate the evolutionary trajectory from subsocial wood roach ancestors to eusociality in termites. 2. Incipient colonies of the subterranean termite Coptotermes gestroi (Wasmann) were investigated to determine if this transition was irreversible and if the burden of brood care on the first alloparents could be measured. To do so, the initial size of the work force necessary for an incipient colony to survive once the brood care became alloparental was determined. 3. The results of the study show that within 5 months after foundation, brood care duties were fully transferred to workers and the primary reproductives became irreversibly dependent on these workers for survival, reproduction, and colony growth. 4. Once the brood care became strictly alloparental, the presence of a single worker was enough to maintain the survival of the king and queen, confirming that ‘reversed parental care’ was also achieved. However, major brood loss and suppressed egg‐laying activity from the queen was observed, suggesting that the burden of brood care was too high for a single worker to absorb. Therefore, once brood care has shifted to alloparents, a critical number of workers is necessary to prevent brood loss and initiate colony growth. 5. As the initial cost of brood care is rapidly absorbed with colony growth in termites, the performance in brood care of the first few alloparents in a subsocial wood roach ancestor may have contributed to the emergence of eusociality in this clade.     

What makes a nest-building male successful? Male behavior and female care in penduline tits

Why do females increase parental effort when caring for theoffspring of attractive males? First, attractive males may bepoor fathers so that their females are compelled to increasetheir own contribution in order to fledge some young (the partner-compensationhypothesis). Second, females mated to attractive males may bewilling to increase their parental effort to reap high indirectbenefits for their offspring, and in turn males can decreasetheir own contribution (the differential allocation hypothesis[DAH]). We investigated these hypotheses in the penduline titRemiz pendulinus, a small passerine bird that has sequentialpolygamy by both sexes and strict uniparental care either bythe male or the female. We focused on two sexually selectedmale traits: nest size and nest-building behavior. We show thatmale care is unrelated to nest-building behavior, whereas femalesare more likely to care for the offspring of those males thatspend more time nest building. Females also more likely carefor the offspring of males that build large nests. Consequently,the reproductive success of males increases with nest size andnest-building behavior. Our results are consistent with theDAH and suggest that nest-building behavior and nest size areunder postmating sexual selection in penduline tits.