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     

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.     

Effects of helpers on breeder survival in the red-cockaded woodpecker (Picoides borealis)

Helpers can gain future indirect fitness benefits by increasing the survival of breeders that produce offspring related to the helper. Helping augments group size through the helper's presence and, in some cases, by increasing fledging success. Breeders may then experience enhanced survivorship because of the benefits of living in large groups. Helping may also reduce the workload of the breeder, which in turn may increase the likelihood that the breeder will survive to breed again. We used Cox's proportional hazards model to examine whether breeders' survival in two populations of the red-cockaded woodpecker (Picoides borealis) was enhanced when group size was increased in the presence of (1) the helper itself, or (2) "extra" fledglings (fledglings produced by the breeder because of helping behavior). We found that in the presence of helpers, the risk of a breeder dying declined by 21–42% for males and 0–14% for females. Our results suggest reduced breeder workload as one mechanism to explain reduced breeder mortality in the presence of helpers: breeders spent less time incubating and provisioning nestlings when assisted by helpers. The risk of a breeder dying declined by 16–42% in males and 26–43% in females in the presence of "extra" fledglings. We speculate on possible mechanisms by which fledglings might affect breeder survival. Our results support the hypothesis that helpers gain future indirect benefits by reducing breeder mortality. Electronic Publication     

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     

Genetic evidence for monogamy in the cooperatively breeding red-cockaded woodpecker

We examined the genetic relationship among putative parents, offspring, and helpers in 224 red-cockaded woodpeckers (Picoides borealis) from the Sandhills of North Carolina. Comparison of DNA similarity with a pedigree constructed from 3,823 individually-marked birds observed from 1979 to 1992 provided verification of observed relatedness in the sampled population (R ² = 0.87,df = 14,P<0.001). In this population, breeding pairs may or may not have helpers, most of which are males that remain on their natal territory. Our sample included helpers assisting their parents, helpers assisting a related male and an unrelated female, and helpers assisting an unrelated pair. Band by band comparison and examination of similarity among DNA profiles indicated that all offspring from non-helped nests were sired by their putative parents (n = 28 families). Similarly, all but one offspring in helped nests were also sired by their putative parents (n = 16 families). In the exceptional case, the offspring evidently was sired by a male external to the group. Analysis of similarity values supported the conclusion that matings by helpers or extra-group males are rare or non-existent. Our results indicate that in this species advantages gained by individuals remaining on their natal territories as helpers do not generally include siring offspring. Correspondence to: S. Haig     

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.     

Replacement of breeders and within-group conflict in the cooperatively breeding acorn woodpecker

Summary Breeding males or females were removed from groups of the cooperatively breeding acorn woodpecker to study (1) competition among individuals from different groups to fill the breeding vancancies, (2) conflict between parents and offspring within groups over replacement of the removed bird, and (3) conflict between parents and offspring for breeding status. Intense fights (power struggles) among nonbreeders from other groups to fill the breeding vacancies developed after the removals. In groups containing only helpers of the opposite sex as the removed bird, replacement was rapid, but in groups containing helpers of the same sex as the removed bird replacement was delayed substantially. In these latter groups parent-offspring conflict developed: helpers could not breed because their parent of the opposite sex was present, and they attacked potential replacements, while their remaining parent advertised for a new mate by calling. Once replacement had occurred, dominance interactions between parents and offspring and among siblings appeared to prevent some individuals from breeding. We conclude that the potential for competition among close kin is high in cooperative species because breeding opportunities are limited.     

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.     

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.     

Offspring sex ratios reflect lack of repayment by auxiliary males in a cooperatively breeding passerine

The repayment hypothesis posits that primary sex ratios in cooperative species should be biased towards the helping sex because these offspring “repay” a portion of their cost through helping behavior and therefore are less expensive to produce. However, many cooperatively breeding birds and mammals do not show the predicted bias in the primary sex ratio. Recent theoretical work has suggested that the repayment hypothesis should only hold when females gain a large fitness advantage from the presence of auxiliary adults in the group. When auxiliaries provide little or no fitness advantage, competition between relatives should lead to sex ratios biased towards the dispersing (non-helping) sex. We examined the benefits auxiliaries provide to females and corresponding offspring sex ratios in the red-backed fairy-wren (Malurus melanocephalus), a cooperatively breeding Australian bird with male auxiliary helpers. We found that auxiliaries provide little or no benefit to female reproductive success or survival. As predicted, the population primary sex ratio was biased towards daughters, the dispersing sex, and females with auxiliaries produced female-biased broods whereas females without auxiliaries produced unbiased broods. Moreover, offspring sex ratios were more strongly biased toward females in years when auxiliaries were more common in the population. These results suggest that offspring sex ratios are associated with competition among the non-dispersing sex in this species, and also that females may use cues to assess local breeding opportunities for their offspring.     

Cooperatively breeding superb fairy-wrens show no facultative manipulation of offspring sex ratio despite plausible benefits

We report a long-term study of offspring sex ratios in the cooperatively breeding superb fairy-wren Malurus cyaneus. Detailed study of this species had revealed a suite of potentially strong selection pressures on the sex ratio. First, females gain substantial fitness benefits from the presence of helpers; so females without male helpers would benefit from any strategy that increased the probability of recruiting help, such as overproduction of sons (local resource enhancement hypothesis), but large numbers of helper males compete among themselves, favouring the production of daughters (local resource competition). Second, daughters fledged early in the season have far greater chances of recruitment to the breeding population than late-fledged daughters, so mothers would benefit from production of daughters early in the breeding season (early bird hypothesis). Third, extra-group mate choice imposes strong sexual selection on males, suggesting that females mating with attractive sires could benefit from investing in sons (sexual selection hypothesis). However, the predictions from these and other sex ratio hypotheses were rejected. The only convincing evidence for manipulation of the sex ratio was a slight bias towards sons (11 sons to 10 daughters) that occurred regardless of context. This result does not support current theory.     

Family structure influences mate choice in white-fronted bee-eaters

White-fronted bee-eaters live in patrilocal, extended family groups in which several pairs may breed simultaneously and in which helping behavior by nonbreeders is common. The composition of a male's family, and his social position within it, are major predictors of his expected reproductive success. For females, the dispersing sex, a decision to pair necessitates forfeiting any potential fitness benefits available through helping in her family, in favor of the potential benefits of breeding in the male's family. Using a set of equations that incorporate the social and non-social predictors of nesting success, we develop a payoff matrix for the expected fitness (current and future) of pairing with different categories of males. Predictions generated from this payoff matrix were supported by data: (1) Females that could help close kin (and thus garner large indirect benefits) tended to remain unpaired. (2) Males with social characteristics associated with low expected reproductive success were more often rejected as potential mates by females. (3) High-quality males, those with potential helpers of their own, were nearly always paired. Simulation analyses were used to investigate female pairing decisions in more detail, and to test various hypotheses about possible decision rules. We demonstrate that pairing females incorporate an assessment of the family structure of potential partners into their mate choice decisions. Such social characters represent an aspect of female choice that has been underappreciated. We suggest that social characters will prove to be important components of mate choice decisions in many species where kinship and dominance considerations influence the likelihood, or success, of breeding.     

Experimental evidence for helping and hindering by previous offspring in the cooperative-breeding Seychelles warbler Acrocephalus sechellensis

Prebreeding Seychelles warblers (Acrocephalus sechellensis) frequently act as helpers on their natal territory, aiding in territory defence, predator mobbing, nestbuilding, incubation (only females) and feeding dependent young of their parents. In some cases helpers could attain breeding status (e.g. by joint-nesting) in their natal group and become co-breeders. Comparisons of group size and reproductive success on a given quality territory suggest that the presence of alloparents (helpers and cobreeders) significantly affects the reproductive success of their parents. The influence of alloparents on reproductive success was examined by removing alloparents from breeding units and comparing the success of natural-sized and artificially reduced groups. Removal experiments, controlled for territory quality, group size and breeder age, showed that the presence of one alloparent significantly improved the reproductive success of its parents. Analysis strongly suggests that this was entirely due to helping behaviour (i.e. providing care to offspring of their parents), thereby improving the helper's inclusive fitness benefits from staying at home. However, these experiments showed also that the presence of two or more alloparents in medium-quality territories significantly decreased reproductive success, compared with groups with one alloparent. Several lines of evidence suggest that this may have been due to the joint-nesting and reproductive competition that could occur in breeding groups, or simply to resource depression when a large number of previous offspring remained on their natal territory.     

The superseded female's dilemma: ultimate and proximate factors that influence guarding behaviour of the carpenter bee Xylocopa pubescens

Both solitary and primitively social nests of the facultatively social carpenter bee Xylocopa pubescens can be found throughout most of the breeding season. In social nests there is reproductive division of labour between a dominant forager and a guarding female. Two types of guarding females can be discerned: the young pre-reproductive guards, and older, formerly reproductive guards. The latter type of guard is found when, after a take-over of reproductive dominance either by a nestmate (mostly a daughter) or an intruder, the defeated female stays in the nest instead of leaving to try and found or usurp another nest. She is then manipulated into the role of a guard. The dominant female profits from the presence of the guard since she protects the nest against pollen robbery by conspecifics (Hogendoorn and Velthuis 1993). We have studied why superseded females might prefer to remain as a guard, rather than try their luck somewhere else. The hypotheses investigated pertain to (1) the difficulty for the defeated female of finding a new nest and of restarting reproductive activities due to (a) ecological constraints (nest and pollen shortage) and (b) the effect of age and wear on the defeated female; (2) the effects of guarding in terms of inclusive fitness. We found that superseded females remained as guards significantly more often when a nestmate (not necessarily close kin) took over reproductive dominance than when an intruder did so. Other factors associated with the decision of the defeated female to stay or leave were her age and the number of her own young still present after the supersedure. The probability of finding or constructing a new nest was lower for old than for young females. After finding a nest, old females produced less brood than young foundresses. As a result of these two factors old superseded females gained, in terms of inclusive fitness, by staying as guards, whereas young females profited from leaving the nest. We interpret these results as an indication that guarding behaviour has evolved due to kin selection. However, kin discrimination apparently did not occur. Therefore we conclude that in this species kin selection is not, in the proximate frame of reference, based on kin recognition and preference for helping kin. Correspondence to: K. Hogendoorn     

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.     

Cooperatively breeding carrion crows adjust offspring sex ratio according to group composition

In cooperatively breeding species, parents should bias offspring sex ratio towards the philopatric sex to obtain new helpers (helper repayment hypothesis). However, philopatric offspring might increase within-group competition for resources (local resource competition hypothesis), diluting the benefits of helper acquisition. Furthermore, benefits of offspring sex bias on parents’ fitness may depend on different costs of production and/or different breeding opportunities of sons and daughters. Because of these counteracting factors, strategies of offspring sex allocation in cooperative species are often difficult to investigate. In carrion crows Corvus corone corone in northern Spain, sons are more philopatric and more helpful at the nest than daughters, which disperse earlier and have higher chances to find a breeding vacancy. Consistent with the helper repayment hypothesis, we found that crows fledged more sons in groups short of subordinate males than in groups with sufficient helper contingent, where daughters were preferred. Crow females also proved able to bias primary sex ratio, allocating offspring sex along the hatching sequence in a way that provided the highest fledging probability to sons in the first breeding attempt and to daughters in the following ones. The higher cost of producing male offspring may explain this pattern, with breeding females shifting to the cheapest sex (female) as a response to the costs generated by previous reproductive attempts. Our results suggest complex adjustments of offspring sex ratio that allowed crows to maximize the value of daughters and sons.     

Flexible helper structure as an ecological adaptation in the pied kingfisher (Ceryle rudis rudis L.)

Summary The pied kingfisher has two types of helpers at the nest: primary ones, helping their own parents, and secondary ones, helping birds other than parents. Primary helpers are always accepted by breeding pairs, secondary helpers only in poor environmental conditions where the time and energy budget of parents is not sufficient for rearing the offspring alone. Under these conditions, helpers increase the breeding success of pairs (Tables 2 and 3) by providing additional food for the young (Table 4). Thus the flexible helper structure can be seen as an ecological adaptation. It is argued that-originating from a skewed sex ratio and breeding in colonies-this adaptation evolved through the combined effects of individual and kin selection.     

Brood defense and age of young: a test of the vulnerability hypothesis

Summary In many altricial species including the great tit (Parus major) the intensity of brood defense against predators has been shown to increase with the age of the offspring. This effect has been ascribed amongst others to the young becoming more vulnerable as they age (vulnerability hypothesis). In a great tit population suffering heavy losses from brood depredation by the great spotted woodpecker (Dendrocopus major), we rendered first and second broods more vulnerable by artificially enlarging the entrance of the nest hole. Contrary to the vulnerability hypothesis, 16 experimental pairs defended their brood against a dummy great spotted woodpecker less vigorously than did 16 control pairs. Nest concealment behavior potentially compromising active defense was minimized by simultaneous playback of nestling distress calls, thus simulating the act of nest predation. This leaves the brood value hypothesis as an alternative functional explanation of the defense level — age effect. It predicts that parents should defend their brood in proportion to the reproductive value (or some more suitable cohortal equivalent measure) of their offspring. At present, this explanation pertains to one predator species. In first broods, but not in second broods, males defended them more vigorously than did their females. While this parallels previous experiments on brood defense against predators posing a much greater risk to the parents, two functional explanations previously put forward can hardly apply.     

Facultative polyandry and the role of infant-carrying in wild saddle-back tamarins (Saguinus fuscicollis)

Summary Wild saddle-back tamarins (Saguinus fuscicollis) in southeastern Peru have a variable mating system that can differ both between territories at any one time and within territories over time. Groups are usually monogamous or cooperatively polyandrous, but are occasionally even polygynous. This study addressed the following questions: why does this population contain both monogamous and polyandrous groups simultaneously? What factors determine whether specific groups are monogamous or polyandrous? The data from this study population tentatively support the hypothesis that adults should mate monogramously only if they have nonreproductive helpers (usually older offspring) to help rear infants. Without helpers, the reproductive success of both males and females is hypothesized to be higher, on average, if they mate polyandrously than if they mate monogamously. The proposed benefits of polyandry to males and females differ quantitatively, but in both cases benefits stem from the help that males provide in rearing young. The following results support this hypothesis. (1) Lone pairs were never seen to attempt breeding, and calculations suggest that the costs of lactation and infant-carrying are too great for lone pairs to have a high probability of being able to raise twin offspring (the normal litter size). (2) Polyandrous males and nonreproductive offspring contributed substantially to infant care, particularly infant-carrying (fig. 2). (3) Adult males carried infants approximately twice as often as did lactating females, presumably because of the combined costs of (a) lactation (Fig. 3) and (b) infant-carrying (Fig. 4). The proximate causes of cooperative polyandry inS. fuscicollis appear to be different from those responsible in several bird species, showing that cooperative polyandry is a complex phenomenon.     

Genetic evidence for monogamy in the cooperatively breeding red-cockaded woodpecker

We examined the genetic relationship among putative parents, offspring, and helpers in 224 red-cock-aded woodpeckers (Picoides borealis) from the Sandhills of North Carolina. Comparison of DNA similarity with a pedigree constructed from 3,823 individually-marked birds observed from 1979 to 1992 provided verification of observed relatedness in the sampled population (R ² = 0.87, df = 14, P < 0.001). In this population, breeding pairs may or may not have helpers, most of which are males that remain on their natal territory. Our sample included helpers assisting their parents, helpers assisting a related male and an unrelated female, and helpers assisting an unrelated pair. Band by band comparison and examination of similarity among DNA profiles indicated that all offspring from non-helped nests were sired by their putative parents (n = 28 families). Similarly, all but one offspring in helped nests were also sired by their putative parents (n = 16 families). In the exceptional case, the offspring evidently was sired by a male external to the group. Analysis of similarity values supported the conclusion that matings by helpers or extra-group males are rare or non-existent. Our results indicate that in this species advantages gained by individuals remaining on their natal territories as helpers do not generally include siring offspring. Correspondence to: S. Haig