Helper contributions to reproductive success in the splendid fairy-wren (Malurus splendens)

Summary In Malurus splendens, helpers were present in 65% of 226 group-years with at least one helper female in 37% of group-years. Most females helped for only one year, while many males did so for at least two years. Most were offspring of one or both present breeders, and in 53% of helper-years, helped both parents. For 159 helpers of known age and parentage, the mean coefficient of relatedness to the offspring was 0.47. Novice females with or without helpers produced fewer fledglings per season than females with one year breeding experience and the same level of help. Helpers did not affect production of fledglings per year by females with one year of experience. Females with two or more years experience and at least two helpers produced more fledglings than equivalent birds with one or no helpers. Experience and helpers have little effect on production of fledglings per nest but they lead to more females renesting after a first brood has been raised. Fewer than 20% of novices renest after fledging one brood, while for females with at least two years experience, the percent renesting after success is 40% with no help, 56% with one helper and 69% with 2 or more helpers. Experienced females begin their first clutch earlier than novices, and helpers reduce the time to renest after success from 66 days for an experienced female with no helpers to 50 days for females with at least two years experience and two or more helpers. Breeding females with helpers survive better (76%) than those with no helpers (55%), and helpers thus gain future indirect fitness. Despite their close relatedness to breeders and offspring, in only 19% of group-years did helpers increase their indirect fitness from an increase in productivity.     

A test of alternate hypotheses for helping behavior in white-fronted bee-eaters of Kenya

Summary Considerable controversy exists over the question of the importance of kin selection in the maintenance of helping behaviors among birds. We examined nine different hypotheses of how helpers might benefit from engaging in alloparental care activities. We break these into four categories: Through its activities the helper may A) improve its probability of surviving to the following breeding season, B) enhance its probability of becoming a breeder in the future, C) increase its reproductive success when it does become a breeder, and D) increase the production of non-descendent kin. The first three categories provide direct fitness gains to the helper; in the fourth, the benefit is indirect. The hypotheses are not mutually exclusive; rather their fitness effects are additive. Each hypothesis, however, makes specific and often separable predictions about both 1) the type of fitness benefits expected, and 2) the characteristics of the birds that serve as helpers. We tested these predictions using five year's data from a color marked and geneologically known population of white-fronted bee-eaters (Merops bullockoides) in Kenya. A) Survival was not related to status (breeder, helper, non-participant); nor did individuals living in large clans have better survival than those living in small ones. B) Newly formed pairs were equally likely to become future breeders irrespective of whether or not one or both individuals had helped previously. C) The mean number of young fledged by a first time breeder was unaffected by its prior helping experience. Neither were first time breeding pairs more likely to gain the services of others as their helpers than were pairs without prior helping experience. Taken together these results demonstrate that beeeaters gain very little direct benefit from alloparenting. D) Helpers did not enhance the survival of the breeders that they helped. But they did have a major effect in increasing nestling survival. Because bee-eater helpers are closely related to the nestlings they help to rear (average r=0.33), they obtain a large indirect benefit by increasing the production of non-descendent kin. We quantified the relative importance of indirect and direct benefits of helping (to the helper) using Vehrencamp's kin index, I _k (1979). I _k compares the fitness consequences of helping against an alternative strategy and calculates the proportion of the inclusive fitness gain or loss that is due to kin (indirect) benefits. Comparing the strategy sets of helping versus not helping for bee-eaters, I _k=0.89 (indicating that 89% of the benefit derived from helping is indirect). When helping was compared against the alternative of breeding, I _k=2.17. Values of I _k greater than 1.0 indicate that direct fitness gains from the alternate strategy (breeding) are greater than those from helping. The value of 2.17 indicates that the helping strategy would not be maintained except for the indirect fitness gained through the increased production of close kin. Alloparenting in white-fronted bee-eaters can thus be considered as altruistic.     

Do stress hormones suppress helper reproduction in the cooperatively breeding red-cockaded woodpecker (<Emphasis Type="Italic">Picoides borealis</Emphasis>)?

In the cooperatively breeding red-cockaded woodpecker (Picoides borealis), male helpers are subordinate to male breeders and do not mate with females, even when unrelated to the breeding female within their group or through extra-group matings, yet exhibit reproductive hormone profiles similar to those of breeders. We investigated whether reproduction might be suppressed in helper males via high levels of the stress hormone corticosterone. We also examined effects of group size and season on corticosterone levels by comparing baseline and maximal plasma levels of corticosterone between helper males and breeding males, and among helper males and breeders of both sexes living in groups of different sizes throughout the reproductive cycle. We also measured plasma levels of estrogen, progesterone, and testosterone to examine other potential hormonal differences between helpers and breeders. Male status did not explain variation in any hormones; therefore, our data do not support the hypothesis that helper males are reproductively suppressed via corticosterone or the other hormones investigated. However, the presence of two or more helper males in a group tended to reduce baseline corticosterone in breeding and helper males, but not breeding females, suggesting that helper males reduce parental effort of other male group members. Seasonally, maximal corticosterone peaked during the nestling provisioning phase for breeding and helper males, but not breeder females, suggesting that males show an increased response to stressors posing a potential threat to survival of offspring.     

Cooperative breeding by the Galápagos mockingbird,Nesomimus parvulus

Summary The costs and benefits of helping behavior were analyzed for 36 pairs of the Galápagos mockingbird, Nesomimus parvulus, and their associates. Helping at the nest is usually done by sons or males suspected to be offspring of the breeders. Costs and benefits to breeders were assessed by comparison of pairs with and without helpers, and costs and benefits to helpers were assessed by comparison of birds which help and those which establish themselves as novice breeders.Helping behavior benefits breeders by increasing fledging success and by reducing the adult energy load in territory defense and feeding of nestlings. Breeders assisted by helpers may also benefit by decreased nest predation. Helpers enhance their inclusive fitness by helping, and gain directly by increasing their chances of securing a territory. Helpers do not appear to gain any fitness advantage from the experience of assisting, nor do they increase their survivorship by remaining on natal territory.Ecological and demographic features such as saturated territories and low territory turn-over rates due to high adult survival may be primarily responsible for the evolution of the helping behavior, with kinselection reinforcing it. Associated features of this system are a male-biased population sex ratio, a greater energetic benefit to breeding males than to breeding females in having helpers, earlier dispersal and breeding by females than by males, and much more frequent helping by males than by females. These are interpreted as consequences of brothersister aggression that indirectly minimizes the chances of inbreeding.     

Status-dependent and strategic growth adjustments in female cooperative cichlids

Male group-living cichlids show status and strategic adjustments in growth, but females appear not to show these growth adjustments. Here, an experimental study in the Lake Tanganyika cichlid Neolamprologus pulcher elaborates on these findings. Females did not show status-dependent growth: (1) Growth in females decelerated with body size but did not depend on social status (breeder or helper), and (2) female helpers did not increase their growth rate after becoming a breeder. Females showed limited evidence for strategic growth: (3) Female helpers did not significantly adjust their growth rate depending on the treatments (comparing female helpers living in groups with a small or a large breeder female); but within the small breeder female treatment, helper growth was significantly related to their body size difference (breeder size–helper size), suggesting a strong non-linear effect of size differences on female helper growth. I conclude that these female cichlids show no status-dependent growth and only strategic growth adjustments when the size difference between the helper female and her breeder female is particularly small.     

An analysis of reciprocal exchange of helping behavior in the red-cockaded woodpecker (Picoides borealis)

We examine the frequency of reciprocal exchanges of helping behavior in three red-cockaded woodpecker populations to determine if such exchanges might constitute a significant fitness benefit of helping. Specifically, we determine how often helpers, once they become breeders, are assisted by young they previously helped raise. The estimated frequencies of reciprocal exchange of helping based on survival and status transition probabilities are low (2%). Observed frequencies of reciprocity are significantly higher than estimated frequencies in two of the three populations studied, suggesting that male fledglings more often remain on their natal territory as helpers if another helper is already present. High rates of retention of young males as helpers on high-quality territories or preferential helping of kin, as well as preferential helping of former care-givers, might explain this result. When the analysis is restricted to helpers unrelated to the young they help raise, which controls for preferential helping of kin and largely eliminates effects of territory quality, empirical estimates of the frequency of reciprocity do not differ from estimates calculated from population demography. We conclude that young males do not preferentially help former care-givers and that reciprocal exchange of help is not an important factor in the evolution of helping behavior in the red-cockaded woodpecker. In this species, all known benefits of helping behavior are indirect (i.e., derived through kin selection ) rather than direct. We discuss the occurrence of unrelated helpers in this context. Received: 10 August 1999 / Received in revised form: 13 February 2000 / Accepted: 19 February 2000     

Social system and reproduction of helpers in a cooperatively breeding cichlid fish (Julidochromis ornatus) in Lake Tanganyika: field observations and parentage analyses

It is suggested that some fish of the genus Julidochromis, substrate-brooding Tanganyikan cichlids with biparental care, breed cooperatively with helpers. We studied the social system of J. ornatus in the wild and analysed genetic parentage using microsatellites. Within the studied population three patterns of social system were identified: monogamous pairs (61%, 44 of 72 groups), pairs with helpers (29%, 21), and polygamous harems with helpers (controlled by either a large female or large male owner; 10%, 7). In cooperatively breeding groups, the number of helpers at each nest ranged from 1 to 6 (median 1), and male helpers were more numerous than female helpers. In both sexes, the body size was different among individuals of different social status (harem owner > breeder > helper). Helpers and harem owners of both sexes exhibited brood defence although its frequency was low. The molecular analysis revealed that (1) the helpers were mostly unrelated to dominant breeders, (2) many helpers of both sexes contributed genetically to the next generations, (3) male helpers had high siring success (41% of young in total), and (4) large young unrelated to group members were detected at 30% of observed nests, which may be due to breeder (or helper) replacements and immigration of young. We discuss the implications of these results for understanding the complex social system of this species, especially the low reproductive skew in comparison with other cooperatively breeding cichlids.     

Is helping a beneficial learning experience for red-cockaded woodpecker (Picoides borealis) helpers?

Several hypotheses have been suggested to explain the evolution and maintenance of helping behavior in cooperatively breeding birds, one of which we investigated in the red-cockaded woodpecker (Picoides borealis). Helping may provide a learning experience that improves reproductive success once the helper becomes a breeder. We used data collected from a population of red-cockaded woodpeckers in the Sandhills of North Carolina to compare the reproductive success of 2-year-old breeders that were helpers at age 1 (helping experience) to those that were floaters or solitary males at age 1 (no helping experience). Reproductive success of the two groups was similar, indicating that helping provides no experience useful in reproduction. We reject the learning experience hypothesis as an explanation for the maintenance of helping behavior in red-cockaded woodpeckers. Received: 29 July 1996 / Accepted after revision: 27 April 1997     

Do helpers increase reproductive success?

Summary Although several different hypotheses have been proposed to explain the evolution of helping behavior, most are based on the assumption that helping enhances the reproductive success of recipient breeders. I tested this assumption by removal experiments in the cooperatively breeding Florida scrub jay (Aphelocoma c. coerulescens). This species lives in permanently territorial social units containing a single breeding pair and none to six nonbreeders, which are usually offspring of the breeding pair and which usually act as helpers by feeding the nestlings and fledglings produced by the breeding pair. Although experimental removals of non-breeders in 1987–1988 had no significant effect on breeder survival, egg production, or hatching success, experimental groups suffered higher rates of predation on nestlings (1987) and lower rates of fledgling survival (both years) than did unmanipulated controls. As a result, experimental groups produced an average of only 0.56 independent juveniles, compared to 1.62 young for controls. Analysis of the factors contributing to nestling and fledgling mortality indicates that helping behavior per se (i.e., the aid that nonbreeders provide to dependent young), not the mere presence of nonbreeders, was responsible for the greater reproductive success observed in control groups. Because survival rates of allofeeders (i.e., those nonbreeders that provisioned dependent young) were virtually identical to those of non-allofeeders, the costs of helping behavior in this species appear to be small. Furthermore, nonbreeders are more likely to provision dependent young within their social unit when those young are closely related. I therefore conclude that nonbreeders increase their indirect fitness by serving as helpers and that helping behavior in the Florida scrub jay is a trait that has current selective utility. It remains debatable, however, whether helping in this species is an adaptation that has been shaped by the process of natural selection.     

Plasma levels of androgens and gonadal development of breeders and helpers in the bell miner (Manorina melanophrys)

Plasma concentration of androgens and gonad development were studied in wild cooperatively breeding bell miners (Manorina melanophrys) of different age, sex and social status. Plasma levels of androgens increased with age of birds. For sexually mature (SM) bell miners, androgen levels were higher in breeders than in nonbreeding (NB) helpers (P < 0.005), and they were marginally higher in females than in males (P < 0.10). Thus female breeders showed the highest plasma levels of androgens. This pattern may be explained by an asymmetry between the sexes in the levels of competition needed to reach and defend breeding status; the degree of intra-sexual competition seems to be higher among female bell miners than among males. In general, male breeders had larger gonads than male SMNB helpers; male breeder gonads produced spermatozoa whereas most helpers' gonads did not. However, old (i.e. 34 months) SMNB male helpers had large gonads producing spermatozoa in spite of having low plasma levels of androgens. We suggest that young male bell miners may be under a voluntary mechanism of reproductive suppression perhaps favoured by inbreeding avoidance; although some aggressive interactions observed between very young helpers (about 2 months) and adult members of the breeding unit suggests that a certain degree of imposition also occurs at young ages. Older male helpers may avoid aggression with the male breeder and imposed reproductive suppression, hence the low circulating levels of androgens, but they have the potential to sneak extra-pair copulations. Gonad size decreased during the non-breeding season in both sexes.     

Reproductive enhancement by helpers and an experimental inquiry into its mechanism in the bicolored wren

Summary As part of continuing studies of sociality in the wren genus Campylorhynchus we have been studying the bicolored wren — a facultatively cooperative breeder — for the past 6 years in the central Venezuelan savanna. Reproductive groups have ranged in size from 2 to 5. In one of our study populations, only about 15% of the groups contained helpers, and nearly all these contained only a single male helper (Fig. 2). In an adjacent population, the majority of groups contained helpers, and more than half of these contained several helpers of either sex. Territory size is, on average, much smaller in the latter population. In these populations the presence of a single helper is associated with a three-fold increase in reproductive success (Table 1). Additional helpers are not associated with further reproductive enhancement. Enhancement is chiefly due to an increased proportion of nest starts that eventually produce independent juveniles. This reproductive enhancement is not merely an epiphenomenon resulting from the presence of helpers on territories which are superior for other reasons, such as greater resource availability or the quality of particular parents. It is also not a function of the mean or variance in nestling feeding rate. Predator exclusion experiments, in which certan nests were artificially protected from terrestrial predators, suggested that the mechanism of reproductive enhancement was heightened effectiveness of nest defense. Helpers are usually nondispersers from the parental territory, and have always been found to be close relative of the nestlings that they assist in rearing.     

Helping at the nest,allofeeding and social status in immature arabian babblers

Summary Arabian babblers, Turdoides sqamiceps, are cooperatively breeding, group territorial birds, occurring in desert habitats. Non-breeders participate in several types of cooperative behaviour, including care of eggs, nestlings and fledglings, and bringing food to one another (allofeeding). This paper reports on observations of helping with eggs and young, and peer allofeeding, performed by immature babblers, fledged the previous season or earlier in the same season. The birds were from a babbler population studied since 1971 in the Arava valley of eastern Israel. They belonged to groups in which all individuals had been colourringed and were accustomed to human observers. Individuals and nests in the territories of these groups could be watched from distances of 1–2 m without causing alarm, so fine details of behaviour could be observed in the field. In most allofeeding interactions observed, a more dominant individual brought food to a subordinate. Sometimes the subordinate bird avoided accepting the proferred food, and was then hit or chased by the bird that had attempted to feed it. Occasionally a subordinate bird attempted to feed a more dominant individual. Dominant birds always refused to take food offered by a subordinate, and hit or chased any subordinate attempting to feed them. Frequencies of visits to nests were correlated with helper rank for those visits where the incoming bird displaced a previous visiter from the nest. Frequences of visits to unattended nests, however, were not correlated with helper rank. Similarly, frequencies of feeding visits to fledglings were correlated with helper rank until incubation of the next brood began. After this, the correlation disappeared. Numerous instances of aborted nest visits were observed in which a helper, often carrying food, arrived at the nest-tree but left again without visiting the nest. In many other cases, a helper arrived at the nest-tree but delayed visiting the nest for several minutes. Aborted and delayed visits usually occurred when a more dominant bird was in the vicinity of the nest. This suggests interference between helpers. Direct interference between helpers visiting fledglings was occasionally observed. In such cases, a more dominant helper snatched food from a subordinate approaching fledglings, and then fed this food to the fledglings itself. Interference between helpers, and conflict between proferers and recipients of food during allofeeding, are not easily explained by kin selection or reciprocity. On the other hand, such behaviour is readily explained by a hypothesis which suggests that individuals may increase their social status in the group by performing cooperative behaviour. Babblers that establish status by demonstrating ability to bear the short-term costs of cooperative behaviour, rather than through direct aggression towards rivals, are likely to forge collaborative relationships with other group members. Since babblers must collaborate to establish and defend a territory, such relationships are essential to reproductive success.     

Does the presence of non-breeders enhance the fitness of breeders? An experimental analysis in the clown anemonefish<Emphasis Type="Italic"> Amphiprion percula</Emphasis>

The stability of animal societies depends on individuals decisions about whether to tolerate or evict others and about whether to stay or leave. These decisions, in turn, depend on individuals costs and benefits of living in the group. The clown anemonefish, Amphiprion percula, lives in groups composed of a breeding pair and zero to four non-breeders. To determine why breeders accept the presence of non-breeders in this species I investigated the effect of non-breeders on multiple components of the breeders fitness. Non-breeders did not assist breeders in any obvious way. Experimental removal of non-breeders had no significant effect on the survival, growth, or reproductive success of breeders. Experimental removal of one of the breeding pair showed that non-breeders had little effect on the time taken for a widowed breeder to recommence breeding. The results indicate that the presence of non-breeders neither enhances, nor reduces, the fitness of breeders in A. percula. I suggest that non-breeders might modulate their effect on the fitness of breeders, either by reducing the costs they inflict or by increasing the benefits they provide, such that it just pays breeders to tolerate, rather than to evict, them. This study illustrates that animal societies can be stable even when some individuals gain nothing from the association.Communicated by M. Abrahams     

Cooperative breeding in purple gallinules: the role of helpers in feeding chicks

Summary The participation of breeders and helpers in the feeding of 21 broods of chicks was studied in a population of cooperatively breeding purple gallinules (Porphyrula martinica). In the breeding group, all birds over the age of 2 months fed chicks. Female breeders fed chicks at the highest rate, followed by male breeders and adult helpers, old juvenile helpers, and young juvenile helpers. The amount of food breeders fed chicks was independent of the number of helpers in the breeding group. However, breeders made fewer feeding visits when they had helpers. Male and female breeders spent similar amounts of time feeding chicks. Helpers spent significantly less time feeding chicks than did breeders. As helpers grew older they fed chicks at a faster rate, made more feeding visits and spent more time feeding chicks. Analysis of variance was used to determine which variables explained the variation in the brood feeding rate (amount of food delivered to an entire brood during one observation). Age of chicks had a significant nonlinear effect, and size of brood and number of helpers had significant linear effects on the brood feeding rate. Chicks in groups with helpers received more food and were accompanied for longer periods of time than chicks in groups without helpers; either or both of these factors may have led to increased chick survival.     

Impact of helpers on colony productivity in a primitively eusocial bee

Small societies of totipotent individuals are good systems in which to study the costs and benefits of group living that are central to the origin and maintenance of eusociality. For instance, in eusocial halictid bees, some females remain in their natal nest to help rear the next brood. Why do helpers stay in the nest? Do they really help, and if yes, is their contribution large enough to voluntarily forfeit direct reproduction? Here, we estimate the impact of helpers on colony survival and productivity in the sweat bee Halictus scabiosae. The number of helpers was positively associated with colony survival and productivity. Colonies from which we experimentally removed one helper produced significantly fewer offspring. However, the effect of helper removal was very small, on average. From the removal experiment, we estimated that one helper increased colony productivity by 0.72 additional offspring in colonies with one to three helpers, while the increase was smaller and not statistically significant in larger colonies. We conclude that helpers do actually help in this primitively eusocial bee, particularly in small colonies. However, the resulting increase in colony productivity is low, which suggests that helpers may be constrained in their role or may attempt to reproduce.     

Helpers and egg investment in the cooperatively breeding acorn woodpecker: testing the concealed helper effects hypothesis

In cooperatively breeding acorn woodpeckers (Melanerpes formicivorus), helper males have a large positive effect on fledging success in good acorn crop years but only a small positive effect in poor acorn crop years, while helper females exhibit the opposite pattern. Based on these findings, we tested the “concealed helper effects” hypothesis, which proposes that laying females reduce investment in eggs (with respect to their size, number, or quality) in a way that confounds helper effects and results in an absence of a relationship between helpers and breeding success. Results generally failed to support the hypothesis. Mean egg size was positively related to temperatures during the 10 days prior to egg-laying and negatively related to the food supply as indexed by the prior fall’s acorn crop, but there were no significant differences vis-à-vis helpers except for interactions with the acorn crop that only partly corresponded to those predicted. With respect to clutch size, females laid larger clutches when assisted by female helpers, opposite the pattern predicted. Although our results suggest that egg size is adjusted to particular ecological circumstances, we conclude that neither egg nor clutch size is adjusted in a way that confounds the apparent effects of helpers, as proposed by the concealed helper effects hypothesis.     

Morphological variation in a population of grey-crowned babblers: Correlations with variables affecting social behavior

Summary Variation in weight, wing length and bill length in a population of grey-crowned babblers is influenced primarily by sex and age, but correlations with size of social unit, with reproductive success, and with vegetation are also detectable (Table 1). The latter correlations vary with sex, age, and status as helper or breeder. Differential wear according to behavioral role, competition for status, incubation, and inheritance are discussed as possible causal mechanisms. Helpers were not detectably smaller in any dimension than breeders of the same age and sex. Male and female non-breeding helpers differ in patterns of morphological correlation, suggesting that they have different behavioral roles. Breeding males have a unique pattern of morphological correlation, suggesting that their foraging behavior differs from breeding females and non-breeders.     

Gonopore sexing technique allows determination of sex ratios and helper composition in eusocial shrimps

An evaluation of the social organization and sexual system of eusocial species of Synalpheus has been hindered because it has not been possible to determine the sexual composition of colony helpers (workers). The external sexual characters typically used to sex caridean shrimps are lacking in Synalpheus. We used SEM sexing technique to determine the sexual composition of helpers in colonies of Synalpheus regalis, S. rathbunae, S. chacei, S. rathbunae A (see Morrison et al. Mol Phylogen Evol 30:563–568, 2004), and S. filidigitus. Colonies consisted of both sexes and sex ratios of helpers generally conformed to 50:50 female to male. Females were characterized by gonopores with U-shaped slits on the coxae of the third pereopods (first walking legs) while males had oval gonopore openings on the coxae of the fifth pereopods (third or last walking legs). In S. chacei, S. filidigitus, and S. rathbunae A, a few helpers were found that had both male and female gonopores (intersexes). All three reproductive females (queens) of S. filidigitus examined were intersexes. Sexing of helpers allowed us to test some hypotheses about sexual differences in helper morphology that might indicate task specialization (division of labor). Male helpers were not different from female ones in body size (except in S. regalis: males somewhat larger) and in fighting chela size. The lack of sexual dimorphism in these characters suggests no male–female specialization in colony tasks such as defense. The presence of male and female helpers similar in size suggests that the sexual system of these eusocial species is gonochoristic, although protandry of some sort in S. filidigitus can not be ruled out. The intersexuality observed in a few individuals may be due to developmental anomalies, protandry, or even simultaneous hermaphroditism. Finally, the sexing technique allowed us to establish that new colonizers of unoccupied sponges in S. rathbunae are a single male and female of helper size.     

A critical analysis of ‘false-feeding’ behavior in a cooperatively breeding bird: disturbance effects,satiated nestlings or deception?

‘False feeding,’ where helpers arrive at nests with food but fail to provision the young, has been reported in several cooperative species. This and other potentially ‘deceptive’ behavior has been interpreted as indicating that helping may operate as a signal within such social groups. We critically examine these phenomena in the provisioning behavior of the bell miner Manorina melanophrys. Excessively close observation distances can artificially elevate the rate of false feeding in this (and other) species, but once this had been accounted for, there was little evidence for any ‘deceptive’ behavior by helpers or breeders. Natural and experimentally induced variation in the presence of a potential conspecific audience at the nest did not have any consistent influence upon the rate of false feeds, which was low at 7.94% of 6,880 nest visits. Instead, encountering unexpectedly low levels of brood demand provided a more parsimonious explanation for those visits where helpers failed to feed nestlings or ate the food themselves. Failure to completely transfer a load to nestlings was more likely when the load contained a high proportion of sticky lerp, indicating a simple prey-transfer problem. Finally, individuals that arrived at nests without prey were often members of neighboring breeding pairs, suggesting that these few non-feeding visits may instead involve an information-gathering function. We, therefore, suggest that future studies explicitly exclude the possibility of observer disturbance and all aspects of normal provisioning behavior before applying the terms ‘false feeding’ or ‘deceptive’ and inferring anything more than straightforward helping at the nest.     

Testosterone and helping behavior in the azure-winged magpie (Cyanopica cyanus): natural covariation and an experimental test

In this study, we investigated patterns of natural covariation between testosterone and reproductive status in a cooperatively breeding bird species, the azure-winged magpie (Cyanopica cyanus). To assess the relationship between testosterone and breeding behavior, we also manipulated testosterone (T) levels early in the season (before breeding started) using testosterone-filled or empty implants. Our results do not support the hypothesis that circulating testosterone levels affect the occurrence of helping behavior in the azure-winged magpie. Helping males had similar T levels to breeding males. Furthermore, experimentally augmented plasma T did not affect the likelihood of becoming either a helper or a breeder. Overall, these results are consistent with previous findings in other bird species and only give some support to the behavioral suppression hypothesis, suggesting that helping in the azure-winged magpie is a flexible behavioral option moderated in the short-term by social and ecological factors. Experimentally elevated testosterone levels, however, reduced the reproductive success of male breeders because of markedly lower levels of paternal care, had similar effects on their mates, but had the contrary effect on helpers, which raised levels of parental effort. We suggest increases in the share of paternity and in social prestige as possible explanations for these results.Communicated by: A. Cockburn