Extra-pair Offspring Research Articles

Demographic drivers of reproductive failure in a threatened bird: Insights from a decade of data

Hatching failure affects up to 77% of eggs laid by threatened bird species, yet the true prevalence and drivers of egg fertilization failure versus embryo mortality as underlying mechanisms of hatching failure are unknown. Here, using ten years of data comprising 4,371 eggs laid by a population of a threatened bird, the hihi (Notiomystis cincta), we investigate the relative importance of infertility and embryo death as drivers of hatching failure and explore population-level factors associated with them. We show that of the 1,438 eggs that failed to hatch (33% of laid eggs) between 2010 and 2020, 83% failed due to embryo mortality, with the majority failing in the early stages of embryonic development. In the most comprehensive estimates of infertility rates in a wild bird population to date, we find that fertilization failure accounts for around 17% of hatching failure overall and is more prevalent in years where the population is smaller and more male biased. Male embryos are more likely to die during early development than females, but we find no overall effect of sex on the successful development of embryos. Offspring fathered by within-pair males have significantly higher inbreeding levels than extra-pair offspring; however, we find no effect of inbreeding nor extra-pair paternity on embryo mortality. Accurately distinguishing between infertility and embryo mortality in this study provides unique insight into the underlying causes of reproductive failure over a long-term scale and reveals the complex risks of small population sizes to the reproduction of threatened species.

Extra-pair paternity in two passerine birds breeding in a gradient of urbanisation.

Urbanisation has been increasing worldwide in recent decades, driving environmental change and exerting novel selective pressures on wildlife. Phenotypic differences between urban and rural individuals have been widely documented in several taxa. However, the extent to which urbanisation impacts mating strategies is less known. Here, we investigated extra-pair paternity variation in great tits (Parus major) and blue tits (Cyanistes caeruleus) breeding in nestboxes set in a gradient of urbanisation in Warsaw, Poland, over three breeding seasons. Urbanisation was quantified as the amount of light pollution, noise pollution, impervious surface area (ISA) and tree cover within a 100-m radius around each nestbox. We obtained genotypes for 1213 great tits at 7344 SNP markers and for 1299 blue tits at 9366 SNP markers with a genotyping-by-sequencing method, and inferred extra-pair paternity by computing a genomewide relatedness matrix. We report higher extra-pair paternity in blue tits breeding in more urbanised areas, for example, with higher light pollution and ISA, and lower tree cover. However, no such trend was found in great tits. Late-stage survival of individual nestlings in both species was not associated with paternity or urbanisation proxies, thus we were not able to detect fitness benefits or drawbacks of being an extra-pair offspring in relation to urbanisation. Our results contribute to the growing body of knowledge reporting on the effects of urbanisation on avian ecology and behaviour, and confirm species-specific and population-specific patterns of extra-pair paternity variation.

Extra‐pair paternity enhances the reproductive fitness of urban Chinese blackbird

Urbanization is a human‐induced process of rapid habitat change that can affect the reproductive success of animals. The occurrence of extra‐pair paternity (EPP) may be sensitive to changes in urban ecological conditions. In this study, the Chinese blackbird Turdus mandarinus, a common breeding bird in southern Chinese cities, was selected as the focus species to explore the EPP patterns of monogamous birds in developing cities and the relationship between pairs. Over four breeding seasons, 316 Chinese blackbird nestlings and 20 adults in both urban and rural areas were monitored and genotyped for 11 microsatellite loci to determine progeny affinity. We confirmed that both the brood size and hatching rate of EPP nests were significantly larger than those of within‐pair paternity (WPP) nests. The body mass, beak length, head length, and body length of 12‐day age extra‐pair offspring (EPO) were significantly greater than those of within‐pair offspring (WPO), suggesting important benefits of EPP. However, in accordance with the analyses of nest ecological variables, we concluded that EPP occurs most often in areas with low urbanization levels, indicating some constraints for this reproductive strategy in cities. Our findings provide new insights about how this growing human‐induced landscape change affects mating behaviour in birds, an important aspect in avian ecology and evolution.

Genetic benefits of female from extra-pair paternity are context dependent within the socially monogamous Tree Sparrow

Females actively seek extra-pair paternity (EPP) to acquire a fitness advantage for their offspring. The “context-dependence hypothesis” posits that female extra-pair mate choice has plasticity in response to environmental conditions, and therefore magnitude of female genetic benefits from EPP depends on the environmental variation. Furthermore, chronic heavy metal pollution can cause adverse effects on fitness-related traits of wild birds. However, few studies were available on the interactions between heavy metal pollution and EPP. We selected an area that was contaminated by heavy metals for more than 60 years (Baiyin, BY), a relatively unpolluted area (Liujiaxia, LJX), and Tree Sparrows (Passer montanus) as study species to explore the response of female extra-pair mate choice and genetic benefits from EPP to heavy metal pollution in socially monogamous birds. The relatedness between social mates and extra-pair mates was investigated. Additionally, we compared the body size and heterozygosity of extra-pair offspring (EPO), within-pair offspring (WPO), social males and extra-pair males from the two Tree Sparrow populations. We found that at BY, female Tree Sparrows tended to choose extra-pair males with larger body size and lower genetic similarity, thereby producing higher heterozygosity and larger body size of EPO compared with those of WPO. However, no similar phenomenon was observed in the Tree Sparrow population from LJX. In addition, there was a significant interaction between population and paternity in the analyses of the fitness-related traits, suggesting that environmental variation could affect female genetic benefits from EPP. This study confirmed the existence of presumed interactions between environmental pollution and EPP within the natural population of socially monogamous Tree Sparrows. Our findings shed lights on the possible effects of long-term environmental stress on mating system in wild birds.

Microsatellite loci for detecting extrapair offspring in Wood Warbler (Phylloscopus sibilatrix, Aves) nests in the northeast of the species range

Microsatellite loci for detecting extrapair offspring in Wood Warbler (Phylloscopus sibilatrix, Aves) nests in the northeast of the species range

Extra-pair paternity, breeding density, and synchrony in natural cavities versus nestboxes in two passerine birds.

Most of what is known about extra-pair paternity in hole-nesting birds derives from studies using artificial nesting sites, such as nestboxes. However, it has rarely been investigated whether inference drawn from breeding events taking place in nestboxes matches what would be observed under natural conditions, that is, in natural cavities. We here report on a variation in promiscuity in blue tits and great tits nesting in natural cavities and nestboxes in an urban forest in Warsaw, Poland. Specifically, we tested whether local breeding density, local breeding synchrony, and extra-pair paternity (inferred from SNP data generated with a high-throughput genotyping by sequencing method) differed between birds nesting in natural cavities and nestboxes. In both blue tits and great tits, the frequency of extra-pair paternity was similar between the two cavity types. In blue tits, we observed shorter nearest neighbor distance, higher neighbor density, and higher synchronous neighbor density (i.e., density of fertile females) in nestboxes relative to natural cavities. No such pattern was found in great tits. Moreover, we detected a positive relationship between the proportion of extra-pair offspring in the nest and neighbor density around the nest in blue tits. Our results revealed that the provisioning of nestboxes did not change rates of extra-pair paternity, suggesting that conclusions drawn from nestbox studies might adequately represent the natural variation in extra-pair matings in some species or sites. However, the observed differences in spatiotemporal components of breeding dynamics highlight the fact that these parameters should be carefully considered when comparing mating behavior across studies and/or sites.

Extrapair offspring of the blue-footed booby show no sign of higher fitness in the first 10 years of life

Extrapair offspring of the blue-footed booby show no sign of higher fitness in the first 10 years of life

Do female bluethroats without extra-pair offspring have more MHC-compatible social mates?

Genes of the major histocompatibility complex (MHC) are crucial for adaptive immunity in jawed vertebrates, and theory predicts that there should be mate choice for optimizing MHC constitution in the offspring. In a previous study, we demonstrated a non-random female choice of extra-pair males in the bluethroat (Luscinia svecica), yielding offspring that was closer to an intermediate MHC class II (MHCII) allele count than their within-pair halfsiblings. The present study tests whether social pairs with only within-pair young (WPY) in their brood, in the same study population, had a combined MHC-constitution closer to a presumed intermediate optimum, than social pairs with extra-pair young (EPY), with a corresponding pattern in their offspring. As expected, we found that WPY from pure WPY-broods were more MHC-optimal than WPY from mixed broods, but only in broods of young (second year) males. Correspondingly, there was a tendency for social pairs with only WPY in their brood to be more MHC-compatible than social pairs with EPY in their brood, when the male was young. Older bluethroat males have considerably larger testes than young males, and their higher sperm competitiveness could help them secure paternity in their own brood, also when they are not MHC-compatible. In other words, in the sexual conflict over paternity, females may be more likely to realise their preference for a MHC-compatible mate when paired to a young male. As a possible fitness indicator, immune responsiveness to an injected antigen (PHA) was elevated for offspring closer to “the golden mean” in MHCII allele count.Significance statementThis study contributes to our understanding of MHC-based mate choice in extra-pair mating systems, by showing that female bluethroats (Luscinia svecica) with an MHCII-compatible social mate tend to have no extra-pair young in their brood, but only when the social male is young. This elucidates a possible sexual conflict, in which older social males are able to override female preferences and prevent other males from gaining paternity in their brood through higher sperm production. Studying systems in which extra-pair paternity occurs offers an insight into the genetic benefits of mate choice, as extra-pair males, in contrast to social males, generally contribute only sperm. Further, the strict and thorough genotyping scheme applied in this study enabled us to demonstrate a preference for “the golden mean” in MHC-diversity in a species with one of the highest MHC class II-diversity known to date.

Experimentally advancing morning emergence time does not increase extra-pair siring success in blue tit males.

Extra-pair paternity occurs frequently in socially monogamous birds, but there is substantial variation in extra-pair siring success among males. Several studies have shown that siring success relates to the timing of morning activity, with the earliest active males being more successful, suggesting that early activity is important for acquiring extra-pair copulations. However, these studies are correlational, and it, therefore, remains unclear whether the relationship between timing and extra-pair siring success is causal. An alternative explanation is that successful extra-pair sires tend to be active earlier (e.g., because they are of high quality or in good condition), but that early activity in itself does not increase siring success. We experimentally advanced the emergence time of male blue tits by exposing them to light about half an hour before their natural emergence time. Although males that were exposed to the light treatment emerged from their roost substantially earlier than males that were exposed to a control treatment, light-treated males were not more likely to sire extra-pair offspring. Furthermore, whereas control males showed the expected relation between emergence time and siring success (although not statistically significant), there was no relation between emergence time and extra-pair siring success among light-treated males. Our results suggest that the timing of emergence from the roost is not an important factor underlying extra-pair siring success.

Jealousy: self-inflicted agony and ruin

This chapter analyses the concept of jealousy (as different from envy) in romantic and sexual relationships. It demonstrates that jealousy is both logically and empirically untenable, as it is: (1) a self-contradictory emotion; (2) a self-destructive activity. As to (1) feelings of jealousy are incompatible with true care for the well-being and satisfaction of the loved partner. Logically, the notion of jealousy is self-defeating: it professes to be an expression of love while at the same time forbidding the loved one the freedom to act autonomously, thus killing the very notion of love. As to (2) nearly all empirical data point to the destructive effects to relationships of the feeling of jealousy, Shakespeare’s Othello being its archetypal expression and its scathing analysis. The astounding fact, however, is that in many (most?) cultures jealousy is seen as an expression of love, while in reality it is an emanation of possessive feeling void of affection for the person ‘loved’. A thorough cultural analysis, coupled to recently available DNA analyses, however, reveal a wholly different picture of extra-pair offspring, fully undermining the foundations upon which the notion of jealousy if based. Recent developments like ‘open relationships’ and ‘polyamory’ may be signs of overthrowing the contradictory and destructive aspects of jealousy. They presuppose, however, to undo deeply ingrained social attitudes accompanying love relationships.

A high level of extra-pair paternity in the Chestnut Thrush (Turdus rubrocanus)

Extra-pair copulation (EPC) can potentially maximize individual reproductive fitness, and this process may involve sexual selection of male and female traits that reflect individual quality. Previous studies have implied that adult characteristics are associated with the probability of extra-pair paternity (EPP), but it differs between species. Moreover, there are relatively few examples of the adaptive rationale for females’ engagement in EPCs based on an examination of these more traditionally recorded traits, in combination with female flight-mediated traits such as wing length. We investigated whether EPP existed in the wild Chestnut Thrush (Turdus rubrocanus) population during three breeding seasons (2019–2021), and whether paternity was related to morphological traits of males and females. Eight highly variable microsatellite loci were used to identify paternity, and generalized linear mixed models were used to analyze the relationship between paternity and morphological traits. We found that EPP existed in the Chestnut Thrush. 53.3% (N ​= ​41/77) of the broods contained at least one extra-pair offspring (EPO), and 34.6% (N ​= ​72/208) of the nestlings were EPO. We also found that male wing length was negatively associated with the probability of EPP and the proportion of EPO. Female body length was positively related to the probability of EPP. Both female body condition and mass were negatively associated with the proportion of EPO. While other traits of male and female did not relate to the probability of EPP or the proportion of EPO. Extra-pair males had better body condition compared to the males they cuckolded. EPO did not differ from their half-siblings in terms of body size or body condition. The results suggest that body size and body condition were associated with EPP in the Chestnut Thrush. This study provides fundamental information for further studies on the evolution and maintenance of EPP in the Chestnut Thrush, and it is also useful for the comparison of EPP among Turdus species.

Patterns of extra-territorial nest-box visits in a songbird suggest a role in extrapair mating.

Many animals make visits outside of their territory during the breeding period, but these are typically infrequent and difficult to observe. As a consequence, comprehensive data on extra-territorial movements at the population-level are scarce and the function of this behavior remains poorly understood. Using an automated nest-box visit tracking system in a wild blue tit population over six breeding seasons, we recorded all extra-territorial nest-box visits (n = 22 137) related to 1195 individual breeding attempts (761 unique individuals). Sixty-two percent of breeders made at least one extra-territorial visit between the onset of nest building and the day of fledging of their offspring, and individuals visited another nest-box on average on 11% of the days during this period. Visit behavior differed clearly between the sexes, with males making over three times as many extra-territorial forays as females. There was a strong overall seasonal decline in visit behavior, but this was sex dependent, with females showing a strong reduction in the number of extra-territorial visits before the onset of egg laying and males showing a strong and sudden reduction on the day their offspring hatched. The likelihood of visiting a particular nest-box declined sharply with the distance to that box, and blue tits almost exclusively visited direct neighbors. Individuals were more likely to have extrapair offspring with an individual whose box they visited, but they were not more likely to disperse to a box they had visited. Thus, our results are inconsistent with the hypothesis that extra-territorial nest-box visits serve to inform dispersal decisions, but suggest that such visits are linked to extrapair mating opportunities.

When Older Males Sire More Offspring—Increased Attractiveness or Higher Fertility?

In birds with extrapair mating, older males usually have higher fertilization success than younger males. Two hypotheses can potentially explain this pattern: 1) females prefer older, and often more ornamented males, or 2) older males invest more in reproduction and fertility than younger males. Here we studied factors associated with age-related male fertilization success in a population of barn swallows Hirundo rustica in Canada. We document that male fertilization success increased gradually up to a minimum age of four-year old. The age effect was especially strong for the number of extrapair offspring obtained and the occurrence of a second brood. The higher fertilization success of older males was also associated with an early start of breeding in spring. The length of the elongated outermost tail feathers, a postulated male ornament preferred by females, also increased with age (in both sexes), but it was not a significant predictor of male fertilization success within age classes. Male fertility traits, especially testis size, but also sperm motility and sperm velocity, increased significantly across age groups. Our results suggest that the higher fertilization success by older males is due to their higher reproductive investments and that their longer tails are an adaptation to early arrival on the breeding grounds.Significance statementThe barn swallow is a socially monogamous passerine with extensive extrapair mating. We found that males become more successful in siring both withinpair and extrapair offspring as they become older. Their increased fertilization success was associated with a higher reproductive effort as indicated by larger testes, more motile sperm, and an earlier start of breeding in spring. The length of the outer tail feathers increased with age in both sexes, but long tails did not enhance male fertilization success among males of the same age. Long tails are probably an adaptation to rapid migration and earlier arrival on the breeding grounds. Our findings suggest that the commonly observed age-related increase in male fertilization success in passerine birds is better explained by life history theory than by sexual selection theory.

Understanding Extra-Pair Mating Behaviour: A Case Study of Socially Monogamous European Pied Flycatcher (Ficedula hypoleuca) in Western Siberia

Extra-pair copulation (EPC) occurred in most socially monogamous bird species. The mechanisms leading to the frequent occurrence of extra-pair offspring (EPO, EPY) in socially monogamous couples, as well as the ‘function’ of EPC, are the subjects of strong debates and raise many unanswered questions. We studied the relationship between extra-pair paternity (EPP) and the different characteristics of males and females in the European pied flycatcher in Western Siberia (Russia). The analysis was based on the genotyping of 232 males, 250 females, 1485 nestlings (250 nests). The European pied flycatchers were predominantly socially and genetically monogamous, but about 20% of birds could be involved in EPP. Loss of paternity tended to be more frequent in one-year-old males. EPCs could be multiple: one individual may have up to three extra-pair partners. The EPP rate was independent of the breeding time. The extra-pair mates of an individual were mainly its near neighbours. The EPC status of an individual was unrelated to most of its morpho-physiological traits. The occurrence of EPP was almost twice as high in females nesting in good quality territories. The fitness of within-pair offspring, EPO, paternal half-sibs of EPO and maternal half-sibs of EPO did not differ statistically significantly. Assuming very low heritability of extra-pair mating, we argued that EPCs could be incidental side effects (by-product) of selection. We believe that the evolution and maintenance of extra-pair mating are the episelective processes in the case of the European pied flycatcher.

Extra-pair paternity explains cooperation in a bird species

In many social animals, females mate with multiple males, but the adaptive value of female extra-pair mating is not fully understood. Here, we tested whether male pied flycatchers (Ficedula hypoleuca) engaging in extra-pair copulations with neighboring females were more likely to assist their neighbors in antipredator defense. We found that extra-pair sires joined predator-mobbing more often, approached predators more closely, and attacked predators more aggressively than males without extra-pair offspring in the neighboring nest. Extra-pair mating may incentivize males to assist in nest defense because of the benefits that this cooperative behavior has on their total offspring production. For females, this mating strategy may help recruit more males to join in antipredator defense, offering better protection and ultimately improving reproductive success. Our results suggest a simple mechanism by which extra-pair mating can improve reproductive success in breeding birds. In summary, males siring extra-pair offspring in neighboring nests assist neighbors in antipredator defense more often than males without extra-pair offspring.

High level of extra-pair paternity in the socially monogamous Marsh Tits (Poecile palustris)

BackgroundExtra-pair copulation behavior has been widely studied among socially monogamous birds. Many species revealed high rates of extra-pair paternity. But few of the studies have been carried out in the Asian population.MethodsFrom 2012 to 2019, we explored the extra-pair paternity of Marsh Tits (Poecile palustris) in Xianrendong National Nature Reserve, Liaoning Province, China. During the study, adult Marsh Tits were captured with mist nets and parental birds, with nest-box traps. Blood samples were taken from the brachial vein. Parentage analyses were carried out using nine highly variable microsatellite loci through Cervus 3.0 software and maximum likelihood approach.ResultsForty-nine offspring (15.08%) from 20 nests (45.45%) were the results of extra-pair fertilization out of a total of 325 offspring in 44 nests. The average extra-pair offspring ratio was 33.54%, with a set varying from 11.11 to 71.43%. Nine extra-pair fathers had been successfully identified, four of whom were the close neighbors of the focus nest while the nests of the remaining five were relatively far. No significant difference was found in the genetic similarity between the social and extra-pair mates of the female, nor in the heterozygosis among the maternal half-siblings.ConclusionsIn general, our study proved that the extra-pair paternity in Marsh Tits and its extra-pair mating is independent of the genetic compatibility hypothesis. This complements the understudied bird’s extra-pair paternity in Asian area and contributes to the comprehensive insight of birds’ extra-pair paternity behaviors.

Improvement of sperm traits related to the high level of extra-pair fertilization in tree sparrow population under long-term environmental heavy metal pollution

Environmental stress can affect sperm traits whose changes have been reported to be associated with extra-pair fertilization (EPF) level in natural animal populations. However, little is known regarding how exposure to environmental heavy metals influences sperm traits and EPF level in free-living bird populations. In a previous study, we found that a tree sparrow (Passer montanus) population that has been exposed to heavy metal pollution over 60 years (Baiyin, BY) exhibits increased sperm quality compared with a population from a relatively unpolluted area (Liujiaxia, LJX). The high sperm quality could be related to extra-pair mating rates. Therefore, the present study investigated EPF level (the ratio of extra-pair offspring) in tree sparrow populations from BY and LJX, and analyzed the relationship between sperm traits (morphology, velocity and quantity) and EPF success. EPF success of tree sparrows was significantly correlated with their sperm velocity (p = 0.048) and total sperm length (p = 0.045), indicating that these sperm traits were important for EPF success. Tree sparrows from the BY population produced longer sperm with lower head/flagellum ratio and faster swimming sperm and showed a significantly higher EPF level than conspecifics from LJX. Thus, adaptive variation of sperm characteristics was related to the high EPF level in tree sparrows under long-term environmental heavy metal pollution. The findings are of scientific significance for exploring the evolution of mating tactics in wild bird populations under environmental stress.

Female differential allocation in response to extrapair offspring and social mate attractiveness.

Renewed debate over what benefits females might gain from producing extra‐pair offspring emphasizes the possibility that apparent differences in quality between within‐pair and extra‐pair offspring are confounded by greater maternal investment in extra‐pair offspring. Moreover, the attractiveness of a female's social mate can also influence contributions of both partners to a reproductive attempt. Here, we explore the complexities involved in parental investment decisions in response to extra‐pair offspring and mate attractiveness with a focus on the female point of view. Adult zebra finches paired and reproduced in a colony setting. A male's early‐life diet quality and his extra‐pair reproductive success were used as metrics of his mating attractiveness. Females paired with males that achieved extra‐pair success laid heavier eggs than other females and spent less time attending their nests than their mates or other females. Extra‐pair nestlings were fed more protein‐rich hen's egg than within‐pair nestlings. Females producing extra‐pair offspring had more surviving sons than females producing only within‐pair offspring. Collectively, results show that females differentially allocate resources in response to offspring extra‐pair status and their social mate's attractiveness. Females may also obtain fitness benefits through the production of extra‐pair offspring.

Female and male plumage color is linked to parental quality, pairing, and extrapair mating in a tropical passerine

AbstractSexual selection has been proposed to drive the evolution of elaborate phenotypic traits in males, which often confer success in competition or mating. However, in many species, both males and females display such traits. Studies examining how selection acts on both sexes are scarce. In this study, we investigated whether plumage ornamentation is sexually selected in females and males lovely fairy-wren Malurus amabilis, a cooperatively breeding songbird. We found that female and male plumage color was correlated with parental quality but not with individual quality or survival. We also found evidence of positive assortative mating based on plumage color. Microsatellite analyses of paternity indicated that the lovely fairy-wren has high levels of extrapair paternity (EPP), with 53% of offspring (in 58% of broods of 57% of females) resulting from extrapair (EP) mating. Female and male plumage color did not predict reproductive success or the proportion of EP offspring in their own nest, but less colorful males obtained higher EPP when paired with more colorful females and gained overall higher total paternity (own nest and other nests). We argue that plumage color may be under sex-specific selection, highlighting the importance of looking at both sexes in studies of sexual selection and ornament evolution. The current findings together with the previous study suggest that plumage color in female and male lovely fairy-wrens appears to be an honest signal relevant in both intrasexual and intersexual competition contexts.

Drivers of alloparental provisioning of fledglings in a colonially breeding bird

Abstract Reproduction is costly. Despite this, evidence suggests that parents sometimes feed unrelated offspring. Several hypotheses could explain this puzzling phenomenon. Adults could feed unrelated offspring that are 1) of their close social associates to facilitate these juveniles’ integration into their social network (the social inheritance hypothesis), 2) potential extrapair offspring, 3) at a similar developmental stage as their own, 4) coercing feeding by begging, or 5) less-developed (to enhance their survival, which could benefit the adult or its offspring; the group augmentation hypothesis). Colonial breeders are ideal for investigating the relative importance of these hypotheses because offspring are often kept in crèches where adults can exhibit allofeeding. Using automated monitoring of replicated captive zebra finch (Taeniopygia guttata) colonies, we found that while parents selectively fed their own offspring, they also consistently fed unrelated offspring (32.48% of feeding events). Social relationships among adults prior to breeding did not predict allofeeding, nor was allofeeding directed toward potential genetic offspring. Instead, adults with more-developed offspring preferentially fed less-developed non-offspring over non-offspring at a similar developmental stage as their own offspring, and this tendency was not explained by differences in begging behavior. Our study suggests that allofeeding is consistent with group augmentation, potentially benefiting adults through colony maintenance or increased offspring survival.