A new interpretation of the social organization and mating system of the Callitrichidae

Tamarins and marmosets are usually described as monogamous, or as living in extended family groups. Field research on a variety of callitrichid species shows, however, that immigration and emigration by adults are common and that groups are likely to be composed largely of unrelated adults of both sexes. Despite the number of adult females present in a group, only a single female is usually reproductively active. This female may mate with more than one male. Care for the young is provided by all group members but principally by adult males. This generally is referred to as a “communal breeding system. ” However, a vertebrate social system in which more than two reproductively active adults participate in infant care is rare and has otherwise been reported only in a small number of avian and canid species. Wild callitrichid groups virtually always include helpers, but the number of adults of each sex is highly variable. We propose that the Callitrichidae tend to live in small multimalemultifemale groups, communally rear the young of one female, and exhibit a mating pattern characterized by female promiscuity but a breeding system that is functionally polyandrous. There is a significant correlation between the number of males and a group and the total number of surviving young. The possible adaptive advantages of this social organization are discussed. An erratum to this article is available at .     

TRAPPING AND RINGING OF EGYPTIAN GEESE AND AFRICAN SHELDUCK AT VOGELVLEI,CAPE

Many small plovers Charadrius spp. have sexually monomorphic plumage and cryptic sexual size dimorphism. The objective of our study was to assess the variation in body sizes between male and female plovers breeding in Madagascar. We collected blood samples and data on adult body sizes of four small plovers (Madagascar Plover Charadrius thoracicus, Kittlitz's Plover C. pecuarius, White-fronted Plover C. marginatus and Three-banded Plover C. tricollaris), and used molecular genetic markers to sex the adults. We found significant differences in body size among the four species, and between sexes. Furthermore, individuals from the southern ecoregion tended to be larger than in the western ecoregion. The Madagascar Plover's body size was significantly more dimorphic than the Kittlitz's and White-fronted Plovers. Breeding Malagasy plovers' show significant sexual size dimorphism (SSD): Madagascar Plover females were heavier and had longer wings than males, whereas the males had longer tarsi; in White-fronted Plover only wing length was different between the sexes. Taken together, our work reports SSD in small African plovers that exhibit monomorphic plumage, and we propose that SSD may be more common than currently acknowledged; we term this 'cryptic sexual size dimorphism'. Our results also suggest sexual selection and/or natural selection exert different pressures on body size in different Malagasy plover species.     

Directional selection on body size but no apparent survival cost to being large in wild New Zealand giraffe weevils

When an individual's reproductive success relies on winning fights to secure mating opportunities, bearing larger weapons is advantageous. However, sexual selection can be extremely complex, and over an animal's life the opportunity to mate is influenced by numerous factors. We studied a wild population of giraffe weevils (Lasiorhynchus barbicornis) that exhibit enormous intra and intersexual size variation. Males bear an elongated rostrum used as a weapon in fights for mating opportunities. However, small males also employ sneaking behavior as an alternative reproductive tactic. We investigated sexual selection on size by tracking individual males and females daily over two 30‐day periods to measure long‐term mating success. We also assessed how survival and recapture probabilities vary with sex and size to determine whether there might be a survival cost associated with size. We found evidence for directional selection on size through higher mating success, but no apparent survival trade‐off. Instead, larger individuals mate more often and have a higher survival probability, suggesting an accumulation of benefits to bigger individuals. Furthermore, we found evidence of size assortative mating where males appear to selectively mate with bigger females. Larger and more competitive males secure matings with larger females more frequently than smaller males, which may further increase their fitness.     

Circadian rhythmicity in cockroaches: effects of early post-embryonic development and ageing

ABSTRACT. Effects of early post-embryonic development and ageing on the circadian rhythm of locomotor activity in the cockroach, Leucophaea maderae , were investigated. Cockroach nymphs (first to fourth instars) were found to exhibit a circadian rhythm of activity generally similar to that of adults, but there appeared to be a significant change in pacemaker period (τ) early in nymphal development, and nymphs exhibited regular fluctuations in activity level which may be related to the moulting cycle. Mean τ of adults was remarkably stable with age – no significant change was found in either males or females throughout the life span of the adult. There was, however, a small but significant difference between the average period (τ) of adult males (τ= 23.72 ± 0.12h) and adult females (τ= 23.84 ± 0.13 h). Lighting conditions during post-embryonic development were found to have major effects on τ of adults. Males raised in constant darkness had a significantly shorter period (τ= 23.52 ± 0.11 h) than males raised in LD 12:12, and adult males and females which had been raised in non-24-h light cycles (T = 22 h or 26 h) exhibited major differences in τ from animals raised in LD 12:12 which persisted for several months. Animals exposed as adults to non-24-h light cycles also showed 'after-effects' on τ, but the magnitude of the effect was much less than that exhibited by animals exposed as nymphs.     

Who teaches children to forage? Exploring the primacy of child-to-child teaching among Hadza and BaYaka Hunter-Gatherers of Tanzania and Congo

Teaching is cross-culturally widespread but few studies have considered children as teachers as well as learners. This is surprising, since forager children spend much of their time playing and foraging in child-only groups, and thus, have access to many potential child teachers. Using the Social Relations Model, we examined the prevalence of child-to-child teaching using focal follow data from 35 Hadza and 38 BaYaka 3- to 18-year-olds. We investigated the effect of age, sex and kinship on the teaching of subsistence skills. We found that child-to-child teaching was more frequent than adult-child teaching. Additionally, children taught more with age, teaching was more likely to occur within same-sex versus opposite-sex dyads, and close kin were more likely to teach than non-kin. The Hadza and BaYaka also showed distinct learning patterns; teaching was more likely to occur between sibling dyads among the Hadza than among the BaYaka, and a multistage learning model where younger children learn from peers, and older children from adults, was evident for the BaYaka, but not for the Hadza. We attribute these differences to subsistence and settlement patterns. These findings highlight the role of children in the intergenerational transmission of subsistence skills.     

Leptin Manipulation Reduces Appetite and Causes a Switch in Mating Preference in the Plains Spadefoot Toad (Spea bombifrons)

Condition- or context-dependent mate choice occurs when females modify their mate preferences depending on their internal or external environment. While the ecological and evolutionary factors that favor the evolution of such plasticity are emerging, relatively little is known of the mechanisms underlying such choice. Here we evaluated whether leptin, a protein hormone involved in the regulation of appetite, might affect the expression of condition-dependent mate choice decisions. To do so, we administered leptin to spadefoot toads, Spea bombifrons, which exhibit condition-dependent mate choice for males of their own species versus congeneric males of S. multiplicata. In particular, poor-condition S. bombifrons are more likely than are good-condition S. bombifrons to prefer S. multiplicata males, but only in environments where hybridization between the two species is beneficial. We found that our leptin treatment reduced appetite in S. bombifrons adults, as was expected from leptin''s known effects on appetite. However, although we predicted that leptin would reduce female preferences for heterospecific males, we found the opposite. In particular, our leptin treatment generated a consistent, repeatable preference for heterospecifics in an environment where females generally prefer conspecifics regardless of condition. These results indicate that leptin has the potential to affect female mate choice, but that it might do so in non-intuitive ways.     

Child fostering as a reproductive strategy among Southern African pastoralists

Information on child fosterage was obtained on 1902 individuals in the course of collecting 611 reproductive histories from Herero and Mbanderu pastoralists in northwestern Botswana in southern Africa. The hypothesis that women foster out their children as a strategy for increasing the number of children they produce is tested. Analysis of these data indicate that sex and birth year of children and parents' marital status significantly influence a child's risk of being fostered, but the relationship of fostering to fertility is unclear. More recently born individuals, females, and children born to parents who were not married to each other are more likely to be fostered than older individuals, males, and children born to parents who were married to each other. A child's risk of being fostered does not appear to be influenced by mother's age if the parents were married to each other, while children born to parents who were not married to each other are more likely to be fostered if the mother is very young, but this pattern is not clear. No association between fostering and mortality was found.     

Courting strategies and conflicts in a monogamous,biparental termite

In species where parental investment is extensive for both sexes, both males and females are predicted to exhibit choosiness when seeking their lifetime partners. Evidence is presented that both courting males and females are choosy in the wood-dwelling, biparental termite Zootermopsis nevadensis. There are, however, sex differences both in the method of mate rejection and in the mate-choice criteria. In particular, females are more likely than males to invite an extra suitor into the nest, whereas males are more likely than females to leave their nest and partner and seek a replacement elsewhere. Correlational analyses of adults show that head width is a mate-choice criterion used by females, whereas body mass and fat mass are used by males. Among males and females that sought replacement mates, those that had been rejected previously were more likely to end up paired with an adult that also had been rejected, compared with adults that had not been rejected. Additional findings reveal a rich repertoire of courting behaviours characterized by extensive intra- and intersexual conflict, including mate rejection, intrasexual combat, and even intrapair aggression.     

Effects of age and sex on intra-group spacing behaviour in juvenile savannah baboons,Papio cynocephalus cynocephalus

Twenty juvenile members of known genealogies in two baboon groups were studied over a 16-month period to evaluate a number of predictions about juvenile spacing behaviour based on the natural history of savannah baboons. Young juveniles (1–2·5 years old) approached more frequently and spent more time in proximity to other group members than did old juveniles (3–5·5 years old). In particular, young juveniles associated more closely with their mothers, particular adult males (possible fathers) and age-peers than did old juveniles. Approaches of young juveniles towards unrelated, high-ranking adults were more likely to occur during feeding than were those of old juveniles. Also, following such an approach, young juveniles were more likely than old juveniles to begin feeding immediately. The overall rates of feeding of old juveniles were depressed when they were in proximity to unrelated, high-ranking adults, whereas the feeding rates of young juveniles were not. Juvenile males approached adult males more often than did juvenile females. Juvenile females approached unrelated adult females more often than did juvenile males. Sex differences also existed in juveniles' choices of unrelated adult female neighbours. Juvenile females associated most often with lactating females, whereas juvenile males associated primarily with cycling females. During group resting, juvenile females approached adult females from higher-ranking matrilines more often than they approached adult females from lower-ranking matrilines. Juvenile males did not exhibit this attraction. Also, among old juveniles, females associated closely with their mothers, whereas males did not. Taken together, the results support the hypotheses that juvenile baboons associate with group members in ways that (1) enhance the probability of surviving an early period of high mortality, (2) create opportunities for social learning of sex-typical behaviours/skills, and, for females, (3) facilitate acquisition of familial dominance status.     

Benefits of size dimorphism and copulatory silk wrapping in the sexually cannibalistic nursery web spider,Pisaurina mira

In sexually cannibalistic animals, male fitness is influenced not only by successful mate acquisition and egg fertilization, but also by avoiding being eaten. In the cannibalistic nursery web spider, Pisaurina mira, the legs of mature males are longer in relation to their body size than those of females, and males use these legs to aid in wrapping a female''s legs with silk prior to and during copulation. We hypothesized that elongated male legs and silk wrapping provide benefits to males, in part through a reduced likelihood of sexual cannibalism. To test this, we paired females of random size with males from one of two treatment groups—those capable of silk wrapping versus those incapable of silk wrapping. We found that males with relatively longer legs and larger body size were more likely to mate and were less likely to be cannibalized prior to copulation. Regardless of relative size, males capable of silk wrapping were less likely to be cannibalized during or following copulation and had more opportunities for sperm transfer (i.e. pedipalpal insertions). Our results suggest that male size and copulatory silk wrapping are sexually selected traits benefiting male reproductive success.     

The evolutionary ecology of early weaning in Kilimanjaro,Tanzania

Public health recommendations promote prolonged breastfeeding of all children; however, parental investment (PI) theory predicts that breastfeeding will be allocated among a mothers'' offspring to maximize her reproductive success. We evaluated PI in terms of risk for weaning before age two among 283 children in Kilimanjaro, Tanzania. Results demonstrate: (i) a Trivers–Willard effect—high socioeconomic status (SES) females and low SES males were more likely to be weaned early; (ii) later-born children were less likely to be weaned early; (iii) higher birthweight children were less likely to be weaned early, and (iv) no effect of cattle (a source of supplementary milk) ownership. These associations were largely independent and remained significant in models controlling for potential confounders; however, the inverse association between early weaning and birth order lost significance in the model containing birthweight. These patterns were observed despite public health recommendations encouraging breastfeeding for at least two years.     

Female Remating Behavior in a Lekking Moth

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

Behavioural allometry and interdemic variation in sexual behaviour of the sailfin molly,Poecilia latipinna (Pisces: Poeciliidae)

Sailfin mollies, Poecilia latipinna, exhibit remarkable levels of intraspecific variation in reproductive behaviour. Larger males exhibit higher rates of courtship and lowered rates of gonoporal nibbling and gonopodial thrusting (forced copulation attempts). Larger males have relatively longer and higher dorsal fins than smaller males. The dorsal fin is a prominent component of the courtship display. Variation in fin measurements, behaviour patterns, and body size of mature males is continuous, and fin shape and behaviour patterns are allometrically related to body size. The allometric pattern is displayed by individual traits as well as by the morphological or behavioural traits in ensemble. Eight populations of mollies from markedly distinct habitats exhibited similar consistent levels of intrademic variation in the size of mature males. Detailed studies on three populations showed that dorsal fin shape could be described by the same regression relationship in all populations, and indicted that a male's shape was determined by his absolute body size. By contrast, there was some variation among populations in the relation of behaviour patterns to male body size. The pattern of this interdemic variation indicated that a male's behaviour patterns were influenced by his relative size in a population. Successful inseminations following forced copulations were rare. The average size of successful males was smaller than the average size of unsuccessful males in all three populations. These results indicated that successful insemination through forced copulation was, like behaviour patterns generally, more a function of the relative size of the male, than his absolute size.     

Patterns of interventions and the effect of coalitions and sociality on male fitness

In group living animals, especially among primates, there is consistent evidence that high-ranking males gain a higher reproductive output than low-ranking males. Primate studies have shown that male coalitions and sociality can impact male fitness; however, it remains unclear whether males could potentially increase their fitness by preferentially supporting and socializing with females. Here we investigate patterns of male interventions and the effect of coalitions and sociality on male fitness in rhesus macaques (Macaca mulatta) with particular focus on male-female interactions. We combined behavioural observations collected on Cayo Santiago with genetic data analysed for male reproductive output and relatedness. Our results revealed that the ten top-ranking males provided the majority of all male support observed. In contrast to other primates, male rhesus macaques mainly formed all-down coalitions suggesting that coalitions are less likely used to enhance male dominance. Males supporting females during and before their likely conception were not more likely to fertilize those females. We also found no evidence that males preferably support their offspring or other close kin. Interestingly, the most important predictor of male support was sociality, since opponents sharing a higher sociality index with a given male were more likely to be supported. Furthermore, a high sociality index of a given male-female dyad resulted in a higher probability of paternity. Overall, our results strengthen the evidence that sociality affects fitness in male primates, but also suggest that in species in which males queue for dominance, it is less likely that males derive fitness benefits from coalitions.     

Temporal patterns of territory settlement and detectability in mated and unmated Nightingales Luscinia megarhynchos

In many bird species the sex ratio of adults is male-biased, which is likely to have consequences for the ecology as well as for the conservation of a species. For example, when some males remain unpaired in a population, there should be strong selection on behavioural traits that enhance pairing success. A surplus of males is also likely to have important implications for the interpretation of breeding bird survey data. In our study population of Nightingales Luscinia megarhynchos , about half of the males stayed unpaired, suggesting that the number of males encountered singing was greater than the number of breeding pairs. Furthermore, the detectability (the probability of encountering a male singing) of mated males was only two-thirds that of unmated males when censused in the morning or late in the breeding season. The relative detectability was more similar early in the season and during the twilight periods before sunrise and after sunset. Males that arrived earlier on the breeding grounds were more successful in attracting a mate than males arriving later. Some of the unmated males deserted their territories and prospected areas up to 4000 m distant, whereas others settled on the study site only late in the season and may actually have changed territories. We suggest that adult sex ratios and the time of the census should be taken into account when interpreting the results of breeding bird surveys.     

Opportunities for female choice in the bed bug Cimex lectularius

Bed bugs are cited as exemplars of sexual conflict because mating can only occur via traumatic insemination. However, past antagonistic coevolution between the sexes does not necessarily preclude current female choice. Here, we investigate opportunities for precopulatory female choice in bed bugs. We examined whether females seek out mating opportunities when they gain the most benefit: when females are virgin and/or have recently fed. But, we found that female mating and feeding status had little effect on female attraction to males and male odor. To determine whether females approach male harborages (home crevices) to seek matings in nature, we investigated where matings occurred among unfamiliar pairs of bed bugs. We found that, despite female attraction to male odor, matings were most likely to take place in the female's harborage rather than the male's harborage. We also examined the effect of feeding on male and female ability to mate. Whereas previous research reported that engorgement impaired female ability to refuse matings, we found that male feeding status had a larger effect on the success of mating encounters than female feeding status. Fed males had poor mating success, suggesting that males may be faced with a trade‐off between mating and feeding.     

Marriage as a reproductive contract: Patterns of marriage,divorce, and remarriage

Patterns of marriage, divorce, remarriage, and redivorce were examined in several representative Western cultures through survey questions and archival data to test the hypothesis that marriage and divorce can be understood as expressions of underlying gender-specific, fitness maximization strategies. Differences between males and females were found for the relationship between age and patterns of both marriage and divorce, with females being far more likely at almost all ages to initiate divorce proceedings than males. Once divorced, however, formerly married females were less likely to remarry than formerly married males. The presence of children from a prior marriage had the effect of further decreasing the likelihood of remarriage for females, but not for males. Formerly married males without children tended to remarry females who had never been married, whereas just the opposite was true for divorced males with children. Consistent with our view of marriage as a reproductive contract, the absence of children was not only conducive to divorce and remarriage, but appeared to increase the likelihood of redivorce as well.     

Tourism and Infant-Directed Aggression in Tibetan Macaques (Macaca thibetana) at Mt. Huangshan,China

ABSTRACT

Previous studies of habituated Tibetan macaques (Macaca thibetana) at Mt. Huangshan, China demonstrated the negative impact that some tourism management strategies had on monkeys' annual infant mortality, but data on precise causes and perpetrators of infant injuries were not available. We worked at the same site and recorded rates of aggression that older monkeys directed toward group infants as a function of tourist numbers and proximity. We found a negative correlation between the number of people on the viewing platform and the frequency of monkeys' aggression toward the group's infants, but the effect is slight. We noted that infant-directed aggression (IDA) was more likely to occur in the provisioning zone where tourists bounded the monkeys on two sides, despite the fact that infants spent very little time in that location. The alpha and beta males engaged in more IDA than expected, while the other three adult males, adult females, and juveniles exhibited less IDA than expected. Males' IDA included biting, chasing, grabbing, and pushing. Adult females bit and grabbed mostly their own infants in the contexts of punishment and weaning. Juveniles' IDA was uncommon and consisted mostly of open-mouth threats. To minimize the severity and frequency of male-toinfant IDA, we recommend that sections of the tourist viewing platform be closed off so that tourists do not surround the monkeys, particularly when infants are near the viewing platforms and tourist numbers are few. The rule against tourists feeding monkeys should also be strictly enforced, so that altercations between the highest-ranked males and other group members, particularly infants, are less likely to occur.     

Mate choice and competition in the barklouseLepinotus patruelis (Psocoptera: Trogiidae): The effect of diet quality and sex ratio

Courtship varies among individuals, partly because individuals differ in quality. To explore proximate factors affecting courtship behavior, I investigated the effect of diet quality on mate choice and competition in the barklouseLepinotus patruelis Pearman (Psocoptera: Trogiidae) in the laboratory. The effect of sex ratio on mate choice was also addressed. Some males were found to exhibit active mate choice, and rejected females in both male- and female-biased sex ratio groups, although they were more likely to do so in a female-biased sex ratio group. Diet quality affected male mate choice: males on high-quality diets were significantly more likely to reject females than males on low-quality diets. Males exhibited choice significantly more often than females, who showed no overt signs of choosiness. Both males and females competed for, access to mates: both sexes attempted to interfere with mounted pairs and females grappled. The choosiness of the male may have directly affected the incidence of female competition. The results also suggest that the patterns of mate choice inL. patruelis differ from those expected by conventional sex role theory.     

Temporal variation of juvenile survival in a long-lived species: the role of parasites and body condition

Studies of population dynamics of long-lived species have generally focused on adult survival because population growth should be most sensitive to this parameter. However, actual variations in population size can often be driven by other demographic parameters, such as juvenile survival, when they show high temporal variability. We used capture–recapture data from a long-term study of a hunted, migratory species, the greater snow goose (Chen caerulescens atlantica), to assess temporal variability in first-year survival and the relative importance of natural and hunting mortality. We also conducted a parasite-removal experiment to determine the effect of internal parasites and body condition on temporal variation in juvenile survival. We found that juvenile survival showed a higher temporal variability than adult survival and that natural mortality was more important than hunting mortality, unlike in adults. Parasite removal increased first-year survival and reduced its annual variability in females only. Body condition at fledging was also positively correlated with first-year survival in treated females. With reduced parasite load, females, which are thought to invest more in their immune system than males according to Bateman’s principle, could probably reallocate more energy to growth than males, leading to a higher survival. Treated birds also had a higher survival than control ones during their second year, suggesting a developmental effect that manifested later in life. Our study shows that natural factors such as internal parasites may be a major source of variation in juvenile survival of a long-lived, migratory bird, which has implications for its population dynamics.