Suppression of prolactin does not reduce infant care by parentally experienced male common marmosets (Callithrix jacchus)

High levels of prolactin have been found to correlate with the expression of paternal care in a variety of taxa. However, in mammals, there is little experimental evidence that prolactin is causally involved in the stimulation or maintenance of paternal care. Here, we suppressed prolactin production in paternally experienced common marmoset fathers in their family groups during the first 2 weeks after their infants were born. Circulating prolactin levels were suppressed using cabergoline (Dostinex: Pfizer), a long acting dopamine (D2) agonist with minimal behavioural side-effects. A within-subject design was used to compare behavioural and hormonal data on 5 paternally experienced fathers during two consecutive births. Cabergoline reduced prolactin to negligible levels in all fathers without effecting testosterone, DHT and cortisol and without adverse side-effects. However, lowering prolactin had no significant effect on the expression of majority of the behaviour patterns associated with paternal care. These included infant carrying, infant grooming and the frequency with which fathers retrieved and rejected infants. The only infant-related behaviour to be affected was the frequency with which fathers touched, licked and investigated infants. We noted a marginally significant increase in this behaviour during cabergoline treatment. Despite the lack of effect on paternal care, cabergoline did exert an effect on the affiliative/sexual behaviour of fathers as there was a significant increase in the grooming behaviour fathers directed at and received from their mates during drug treatment. This study showed that experienced male marmosets can express paternal behaviour in the absence of the high prolactin levels normally seen after infants are born.     

Carrying behaviour in captive and wild marmosets (Callithrix jacchus): A comparison between two colonies and a field site

Data on callitrichids infant carrying is highly variable, and there is some disagreement over the relative involvement of different group members. Considering that variation may result from different social and environmental conditions it is important to compare data from different environments. The purpose of this study was to compare infant carrying in 11Callithrix jacchus groups living in two colonies, in the UK and Brazil, or in a field site in the northeastern Brazil. No differences were found in the amount of time that infants were carried, in the three environmental conditions, during weeks 1 to 4, 7, and 10. Fathers and mothers carried infants for equivalent amounts of time in the three conditions, but helpers from natural groups carried infants more than their captive counterparts. In general, adult helpers carried infants more than subadults and juveniles. The results suggest that carrying is a stable feature inCallithrix jacchus groups, but there is a trend suggestive of an extended carrying period in natural groups. Mothers behaviour were remarkably stable, but fathers behaviour, although comparable, was more conspicuous in captivity than in the field, what may be attributed to paternity certainty. Finally, the greater participation of field helpers in carrying is considered as deriving mostly to age of helpers than to group size.     

Infant carrying in family groups of goeldi's monkeys (Callimico goeldii)

Goeldi's monkey (Callimico goeldii) is closely related to marmosets and tamarins. Like marmosets and tamarins, C. goeldii lives in family groups, and fathers and older offspring (helpers) participate in infant carrying. In contrast to the typical twin births in marmosets and tamarins, C. goeldii has only single offspring, and paternal carrying is delayed. We studied infant carrying following 26 births in eight groups of C. goeldii, testing hypotheses proposed in the literature on infant carrying in marmosets and tamarins. The infant was carried exclusively by the mother for the first 26.3 days after birth. Afterwards other group members participated in infant carrying. Whereas the C. goeldii mother is always the main carrier, the father does not always carry more than helpers. In contrast to other callitrichids, age and sex of sibling helpers was not found to have an effect on the participation in infant carrying. The participation in infant carrying in Callimico indicated intraindividual consistency, i.e., the amount of infant carrying performed by fathers and helpers following one birth correlated significantly with the amount of infant carrying of the same individuals following the next birth. We found a significant negative correlation between parental infant carrying and group size, indicating that helpers really do help, sharing the carrying burden with their parents. This is attributed to a clear trend for a reduction in maternal carrying in the presence of helpers, whereas fathers did not benefit from helpers. We conclude that the infant-carrying pattern in C. goeldii is different from the infant-carrying pattern in marmosets and tamarins. The main differences were that the mother instead of the father is the main carrier, and that there is a clear time delay between infant birth and when the father and helpers participate in infant carrying.     

Costs of infant‐carrying in the cotton‐top tamarin (Saguinus oedipus)

Infant‐carrying behavior among callitrichids seems to be a costly activity. Costs have been related to the physical efforts of carrying the weight of very heavy infants and to the resulting reduction in foraging efficiency. However, the costs of carrying in terms of the physical consequences for carriers have not previously been assessed. In this study, we have regarded weight loss in infant carriers as a measure of costs. We studied five families of cotton‐top tamarins (Saguinus oedipus) during the first 9 weeks following the birth of infants. Captive‐breeding conditions were required so that body weight could be measured frequently. To avoid inflicting undue stress on the subject animals, we used a noninvasive method for weighing the tamarins. Differences in carrying contribution were found amongst fathers and male and female helpers, with female helpers contributing less. We have found that carrying infants in the cotton‐top tamarins is an activity that produces a weight loss. Fathers and male helpers go through a maximal body weight loss. While carrying, the tamarins also decrease food intake. However, no relationship was found between contribution to carrying and feeding time or in energetic intake during feeding observations. Thus, it seems that a direct relation doesn't exist between the observations of feeding and weight loss. Fathers increase their contribution to carrying during mothers' periovulatory periods. In this period, male helpers and especially fathers go through a maximal body weight loss. We found body weight losses of up to 11.3% in one subadult male and 9.1percnt; in a father during the fifth week. No changes occurred in food intake in fathers or other male helpers during this period. During periovulatory periods, mothers carried less frequently but did increase their food intake. They gained weight from the second week after birth onward, especially during the periovulatory period. It seems that the infant‐carrying behavior of fathers and male helpers may contribute to the improvement of the mothers' physical condition after birth and therefore may support a consecutive pregnancy. Am. J. Primatol. 48:99–111, 1999. © 1999 Wiley‐Liss, Inc.     

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.     

Why do cotton‐top tamarin female helpers carry infants? A preliminary study

Shortly after giving birth, cotton-top tamarin mothers frequently attack the eldest female helpers [Snowdon et al., American Journal of Primatology 31:11-21, 1993]. Sometimes this aggression leads to the eviction of the eldest daughters from their natal groups 3-4 months after the birth of infants [Price & McGrew, Folia Primatologica 57:16-27, 1991]. We propose that daughters, during the act of carrying infants, may receive less aggression from mothers than when they are not carrying. On the other hand, given that mothers benefit from having others carry their infants, overall aggression received by female helpers from their mothers might be lower in those female helpers with a larger relative contribution to this activity. Four groups were observed during the first 9 weeks following the birth of infants, and aggression received was recorded for symmetrical as well as nonsymmetrical interactions. We found a positive correlation between contribution to infant carrying of female helpers and the overall aggression received from their mothers. Furthermore, the two eldest daughters with highest values of carrying contribution were evicted from their natal groups 3-4 months after the birth of infants. Although mothers do not appear to be more tolerant of female helpers that contribute the most to infant carrying, daughters do benefit from reduced aggression during the act of carrying, and remain in their natal group during the time period when infants must be carried.     

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.     

Elevated prolactin levels immediately precede decisions to babysit by male meerkat helpers

Recent studies suggest that decisions to care for the offspring of others in societies of cooperative vertebrates may have a hormonal basis. The crucial question of whether changes in hormone levels immediately precede or merely follow bouts of offspring care, however, remains largely unanswered. Here, we show that in wild groups of cooperatively breeding meerkats, male helpers that decided to babysit for the day had significantly higher levels of prolactin, coupled with lower levels of cortisol, before initiating a babysitting session compared with similarly aged individuals that decided to forage. In addition, these hormonal differences disappeared over the course of the day, suggesting that hormone levels changed in a fundamentally different way in meerkats that babysat versus those that foraged. In contrast, long-term contributions to babysitting were not significantly associated with plasma levels of prolactin, cortisol, or testosterone in individual male helpers. Our results show, for the first time, that elevated levels of prolactin may immediately precede bouts of helping behavior but differ from recent findings on the same study population in which plasma levels of cortisol, but not prolactin, were significantly and positively associated with rates of pup feeding by male helpers. Together, these results lend significant weight to the idea that decisions to help in cooperative vertebrates have a hormonal basis, although different hormones appear to be associated with different types of care.     

The Role of Non-reproductive Helpers in Infant Care in Captive Callithrix jacchus

Callitrichids have been proposed as communal breeders, with non-reproductive animals acting as helpers. Reports in the literature attest to the high variability in the behaviour of helpers. The present study assessed four proximate factors involved in helping in captive Callithrix jacchus: age, gender and experience of helpers and composition of their groups. Forty-eight helpers from 15 families from Natal (Brazil) and Reading (UK) colonies were observed during the first 4 weeks of life of new-born twins. Infant carrying, transfers and retrieval, and duration of carrying bouts were registered. Results showed that the presence of helpers decreased the costs of infant carrying, with a negatively significant correlation between group size and mean percentage of carrying. Age, but not gender, was a significant factor for infant carrying, transfers and retrieval, and mean duration of carrying bouts, with older helpers contributing more than younger ones. Singletons and oldest non-reproductive offspring carried infants more, and longer, than twins and offspring with older siblings present in the group. Experience decreased motivation and actual carrying time, but not to a significant level. Age, experience and group composition appeared important factors in the display of infant care by helpers in captive Callithrix jacchus.     

The evolution of non-maternal care in anthropoid primates: a test of the hypotheses

The amount of non-maternal care (allocare) found in primates varies greatly from species to species. Our paper examines this variation and focuses on possible reasons why mothers in some anthropoid primate species are prepared to relinquish their infants to other carers whereas others are not. We use data collected from the literature to test a number of hypotheses that attempt to explain the observed variation in non-maternal care. Analyses were carried out using comparative methods that control for the influence of both body size and phylogeny on life-history and ecological variation. The results support previous studies of both primates and other mammals in finding a clear link between the amount of allocare provided and female reproductive rates. Species with high allocare levels grow rapidly post-natally and wean their infants at a younger age (but at the same relative size) than species of the same body size with lower allocare levels. This early weaning allows high allocare species to support higher birth rates than low allocare species but does not result in earlier maturation. Our results, therefore, suggest that mothers allow non-maternal care of their infants in order to increase their own reproductive output. It is not clear whether such a strategy also benefits the infant, as we could find no link between the presence of allocare and early maturation (which would increase the infant's reproductive output) or between allocare levels and infant survival (as measured by vulnerability to infanticide). This suggests that mothers and infants might come into conflict over transfer to other carers, as the benefits to the mother may not be accompanied by benefits to the infant. However, although mothers may benefit from allocare in some circumstances, they will not be expected to allow allocare if the costs are high (e.g. if there is a high risk to the infant).     

Infant-care Behavior of Non-reproductive Helpers in a Communal-care Primate,the Cotton-top Tamarin (Saguinus oedipus)

Infant-care behavior patterns of captive cotton-top tamarins were examined to assess factors defining participation by non-reproductive helpers. The time spent carrying infants and characteristics of infant transfers were examined for 47 helpers in a total of 18 groups. We predicted that age, previous experience, and carrying opportunity would all affect participation of non-reproductive helpers. Our results confirm that carrying by non-reproductive helpers was related to age, with older helpers carrying more often. However, this difference declined with increasing infant age, suggesting that body size of the carrier was not the only factor determining participation in carrying. When observations were classified relative to type of interaction with infants, older juveniles were found to both investigate and harass infants more often than subadults or younger juveniles. There was no effect of gender on carrying. The carrying behavior of subadults was not affected by their previous experience; that is, subadults with no previous exposure to infants carried as often as those with previous experience. Inter-individual variation among helpers was high; within 11 twin litters of helpers, one animal usually carried significantly more than the other.     

The benefits of helpers: Effects of group and litter size on infant care in tamarins (Saguinus oedipus)

The Callitrichidae (marmosets and tamarins) typically give birth to twins, and infant care is shared by all group members. The potential benefits to callitrichids of having helpers were investigated in a study of 21 captive cotton-top tamarin (Saguinus oedipus) infants, living in groups with two to 12 older members. Time carried and suckled and amount of food received from other family members were recorded for each infant during the first 12 weeks of life. The results showed that infants in larger groups were carried more and received more food than those in smaller groups. Twin infants in larger families were less likely to be on the same carrier. Singleton infants were carried more than twins but did not receive more food. On average, individual caretakers in larger groups carried less and shared less food with infants than those in smaller groups. When parental contributions to care were analyzed, no effects of group size were found on mothers' contributions to carrying or food sharing, but fathers in larger groups both carried infants less and shared less food with them. There appear to be several benefits of a communal rearing system to cotton-top tamarins. 1) Infants may receive more care in larger families, thus increasing their chances of survival. 2) The burden of care is spread over several animals, reducing the costs to a given individual. 3) The parents, particularly fathers, may benefit most from reducing the costs of investing in the present litter and increasing their ability to invest in future litters.     

Effect of the enclosure on carriers' body weight loss in the cotton-top tamarin (Saguinus oedipus)

Infant carrying in cotton-top tamarins is a costly activity that results in weight loss by the fathers and helpers. However, to date, measures of carrying costs have been obtained in small indoor enclosures. We studied body weight changes in adult and subadult individuals from eight large groups (n > 5) for 9 weeks after the birth of infants. Four groups were housed in large indoor/outdoor enclosures (42 m(2) x 3.3 m high), and four were housed in small indoor enclosures (12 m(2) x 2.4 m high). All of the individuals were weighed regularly at least three times a week. Reproductive males lost more weight in the big indoor/outdoor enclosures (mean=6.51%) than in the small indoor ones, as did male adult helpers (mean=5.59%) and female adult helpers (mean=4.4%). Still-growing subadult individuals also lost weight in the big indoor/outdoor enclosures (mean=3.17%), although the differences did not reach significance (P=0.07). These results support the hypothesis that cotton-top tamarins living in more natural settings experience higher weight loss than those housed under less-natural conditions.     

Allocare in a nocturnal primate: data on the spectral tarsier, Tarsius spectrum

Non-maternal infant care in many of the small-bodied New World primate species has been hypothesized by some researchers to be related to the high infant/adult weight ratio found in these species. The spectral tarsier, Tarsius spectrum, an Old World primate, has one of the highest infant/adult weight ratios of any primate, with infants weighing between 20-33% of adult weight at birth. On the basis of the hypothesized relationship between allocare and the infant/adult weight ratio, it is predicted that the spectral tarsier will also exhibit extensive allocaretaking behaviour. The results of this study indicate that although spectral tarsiers show care by male and female subadults as well as adult males, it is extremely limited compared to the extensive allocaretaking behavior observed in New World primate species such as Aotus and Callicebus. Spectral tarsier subadult females provide substantially more allocare to infants than do subadult males or adult males. Female subadults were observed sharing food, transporting, grooming, playing, alarm calling, baby-sitting and maintaining physical contact with infants more than other age/sex classes. Although the amount of allocare exhibited by adult males and subadult males was much less than that exhibited by female subadults, the data suggest that adult and subadult male spectral tarsiers do play a small part in the care and socialization of the infant. Adult males and subadult males were both observed occasionally engaging in allocaretaking behaviors such as grooming and playing, as well as frequently patrolling and defending the territory's boundaries. The results from this study suggest that although a high infant/adult weight ratio may be a prerequisite for selection to favor extensive allocare, it is not a causal factor. Additional research is needed in order to understand better the selective pressures involved and the costs and benefits of providing allocare to subadults and adult male spectral tarsiers.     

Long-term study of infant-carrying behavior in captive common marmosets (Callithrix jacchus): Effect of nonreproductive helpers on the parents' carrying performance

Infant-carrying behavior in four families ofCallithrix jacchus was investigated over a period covering six to eight litters. We evaluated the effect of the total number of helpers and the number of adult helpers on the parents' carrying performance for the total carrying period and for three age stages of the infants. The carrying performance of the parents reached an asymptote beginning with litters 3 to 5. In two groups, the carrying performance of the parents was significantly negatively correlated with group size. However, considerable differences existed within each group and between the groups. The mother was the primary caregiver in the early weeks of the infants' lives. In this phase the helpers' contribution to carrying was less than might be expected. The relief of the parents from infant-carrying was greatest when the infants were 4 to 7 weeks old. Breeding males benefited most and from every helper, while breeding females benefited most from adult helpers, i.e., the carrying performance of the breeding female was considerably reduced until the respective group had grown to 8 to 10 members (infants not included). This group size is considered to be ideal for maximal relief of the parents from infant carrying. An increasing number of adult nonreproductive helpers (>4 or 5) does not induce a further reduction of the parents' carrying performance. Infant-rearing experience of nonreproductive helpers seems to be more important for the parents' and, especially, the breeding female's relief from infant-carrying than the overall number of helpers is. The data also strongly evidence that a group member's participation in infant-carrying is influenced by housing conditions and the demographic history of the group.     

Testosterone and prolactin are associated with emotional responses to infant cries in new fathers

To determine the responsiveness of new fathers and non-fathers toward infant cues, we exposed fathers and non-fathers to infant cries and to control stimuli and we measured affective, heart-rate, and endocrine responses, including salivary testosterone and cortisol and plasma prolactin concentrations prior to and after cry presentations. We found that (1) fathers hearing the cry stimuli felt more sympathetic and more alert compared to groups who did not hear the cries or to non-fathers who heard the cries; (2) fathers and non-fathers with lower testosterone levels had higher sympathy and/or need to respond to the infant cries than fathers with higher testosterone levels; (3) fathers with higher, as opposed to lower, prolactin levels were also more alert and more positive in response to the cries; (4) fathers hearing the cry stimuli showed greater percentage increase in testosterone than fathers not hearing the cry stimuli; (5) experienced fathers hearing the cries showed a greater percentage increase in prolactin levels compared to first-time fathers or to any group of fathers hearing control stimuli; finally, (6) partial correlations with parity and experience entered as a covariates indicated that both experience and testosterone contributed to the variance in fathers' affective responses to infant cries. Taken together, these results indicate that, as with a number of other biparental species, human fathers are more responsive to infant cues than are non-fathers and fathers' responses to infant cues are related to both hormones and to caregiving experience.     

Kin and social relationships in splendid fairy-wrens: recognition by song in a cooperative bird

Cooperatively breeding splendid fairy-wren,s Malurus splendens, were tested with songs recorded from individuals of known social and kin relationships. Both males and females sang and responded aggressively to songs of wrens from other social groups. Wrens responded similarly to songs of non-kin and songs of close kin in the absence of social familiarity with them. Breeding females responded more intensely to songs of helpers from other groups than to songs of helpers in their own group. The songs of male and female helpers elicited similar responses by breeding females. The response to other females may be associated with competition for breeding status and helpers. Two females sometimes breed in social groups with two older females; no interference was observed. Song may allow individuals to recognize other wrens in their group and to direct their behaviour towards non-dispersing relatives by location and social familiarity rather than by kinship identifiers.     

Effects of allocare-givers on fitness of infants and parents in callitrichid primates

The effects of callitrichid primate helpers (allocare-givers other than an infant's father) on the survival, reproduction or behavior of infants and parents are reviewed, using both published studies and data from free-ranging golden lion tamarins (Leontopithecus rosalia). Three lines of evidence suggest that helpers may increase their own inclusive fitness: (1) The number of adult males acting as helpers in free-ranging groups is correlated with the number of surviving infants in 3 callitrichid species. However, the lack of a negative correlation with number of infants dying suggests that activities other than direct infant care (e.g. territory defense) may be more important, especially in newly formed groups. (2) In 2 species, captive groups with helpers carry infants for longer periods of time than do groups without helpers. Whether such differences would translate into meaningful survival differences in free-ranging groups is unclear. (3) Helpers reduce the energetic burden of parents by reducing the amount of time they spend transporting or provisioning infants in at least 4 species. Reproductive males are more likely than reproductive females to benefit from the presence of helpers, reducing their investment in infant care activities as the number of helpers in the group increases. In free-ranging golden lion tamarins, the reproductive tenure of males, but not females, increases with the number of helpers in the group, suggesting that a reduction in energetic investment may translate into increased survival. 'Decisions' made by helpers to participate in infant transport are weighed against competing needs for foraging, vigilance, territory defense and, in some cases, prospecting for breeding opportunities. Given this complexity, a sophisticated model may be required to answer the question of how helpers 'decide' to participate in infant care versus other activities.     

The relationship between social status, behaviour, growth and steroids in male helpers and breeders of a cooperatively breeding cichlid

We tested whether subordinate helper males of the Lake Tanganyika cooperatively breeding cichlid Neolamprologus pulcher show elevated excretion levels of the stress hormone cortisol and reduced levels of 11-ketotestosterone and testosterone when living in groups with a small breeder male, compared to similar helper males living in groups with a large breeder male, in a full-factorial repeated measures experimental design. We also measured the same hormones in breeder males with and without helper males. Previous research showed that the size difference between large male helpers and male breeders in groups of this species influences behaviour and growth decisions. Contrary to our expectation, no effect of the size-difference between helper males and breeder males on helper hormone levels was detected. Furthermore, helper males had similar hormone excretion levels to those of size-matched breeder males without helpers, and to small breeder males. There was no influence of egg laying on breeder male and helper hormone levels during the experiment. Interestingly, all three hormone levels were significantly lower in helpers showing elevated levels of submissive behaviour towards the breeders, independently of the size of the breeder males. The low cortisol levels suggest that helper males can successfully reduce stress by appeasing breeder males through submission. Furthermore, helper males showing a high level of submissive behaviour had lower levels of androgens than less submissive helpers, suggesting a lower reproductive potential in submissive helpers. We propose that helper submission may be used as an honest signal of reduced interest in reproduction towards the breeder male in this species.     

Positive effects of helpers on reproductive success in the brown treecreeper and the general importance of future benefits

1. Numerous studies of cooperatively breeding species have tested for effects of helpers on reproductive success to evaluate hypotheses for the evolution of cooperation, but relatively few have used experimental or statistical approaches that control for the confounding effects of breeder and territory quality. 2. In the brown treecreeper Climacteris picumnus, most helpers are male offspring of the breeding pair that have delayed dispersal. We analysed 5 years of data (97 territory-years) using hierarchical linear modelling to test for effects of helpers on reproductive success while controlling for confounding factors. 3. The number of helpers was related positively to reproductive success even after controlling for differences between territories and breeders. A threshold effect was observed, with success increasing most with the presence of a second helper (i.e. at group size of four). 4. Feeding at the nest was one mechanism responsible for this effect, as larger groups had higher total feeding rates at all nesting stages. Higher total feeding rates, as well as higher feeding rates by helpers, were correlated in turn with greater reproductive success. 5. An analysis of the effects of helper feeding rate on reproductive success in groups with just one helper produced only weak support for a positive effect of helpers. Controlled comparisons of this kind utilize only a small fraction of the total data available and thus have limited statistical power compared to hierarchical or mixed-modelling. 6. A number of hypotheses to explain the evolution and maintenance of helping behaviour are consistent with our results for brown treecreepers including kin selection and hypotheses based on future direct benefits. 7. A previous synthesis of studies of helper effects that controlled for confounding factors suggested a pattern in which male helpers rarely have positive effects on reproductive success. However, revising that synthesis to include recent hierarchical or mixed-modelling studies suggests that helpers of both sexes usually have positive effects, and that the relative importance of future direct benefits may have been underestimated.