Experimental evaluation of sex differences in territory acquisition in a cooperatively breeding bird

In many species, territory ownership is a prerequisite for reproduction;consequently, factors that affect success in territory acquisitioncan have a large impact on fitness. When competing for territories,some individuals may have an advantage if, for example, theyare phenotypically superior or more familiar with the site thanothers. The relative importance of the many factors involvedin territory acquisition is, at present, unclear. We studiedpatterns of natural territory acquisition in a closed and saturatedpopulation of Seychelles warblers. Furthermore, by removingbreeders, we experimentally investigated the relative importance,to territory acquisition, of a range of factors and assessedwhether this differed between the sexes. In both sexes, themain route to natural territory acquisition was to dispersefrom the natal territory to immediately claim a vacant dominantposition. Males were older than females when acquiring a territoryfor the first time. In the removal experiment, for both sexes,the proximity of an individual's natal territory to a vacantdominant position was positively related to the individual'schance of claiming the vacancy. Older males were more likelyto gain an experimental vacant dominant position than were youngmales, whereas age did not affect territory acquisition in females.In the Seychelles warbler, the degree of intrasexual competitionfor territory ownership may be stronger for males than for femalesbecause territory ownership is a prerequisite for male reproduction,whereas females can reproduce on their natal territory. In suchcompetition, young males subsequently lose out to older ones.     

Costly reproductive competition between females in a monogamous cooperatively breeding bird

In many cooperatively breeding societies, only a few socially dominant individuals in a group breed, reproductive skew is high, and reproductive conflict is common. Surprisingly, the effects of this conflict on dominant reproductive success in vertebrate societies have rarely been investigated, especially in high-skew societies. We examine how subordinate female competition for breeding opportunities affects the reproductive success of dominant females in a monogamous cooperatively breeding bird, the Southern pied babbler (Turdoides bicolor). In this species, successful subordinate reproduction is very rare, despite the fact that groups commonly contain sexually mature female subordinates that could mate with unrelated group males. However, we show that subordinate females compete with dominant females to breed, and do so far more often than expected, based on the infrequency of their success. Attempts by subordinates to obtain a share of breeding impose significant costs on dominant females: chicks fledge from fewer nests, more nests are abandoned before incubation begins, and more eggs are lost. Dominant females appear to attempt to reduce these costs by aggressively suppressing potentially competitive subordinate females. This empirical evidence provides rare insight into the nature of the conflicts between females and the resultant costs to reproductive success in cooperatively breeding societies.     

Species‐level repertoire size predicts a correlation between individual song elaboration and reproductive success

Birdsong has long been considered a sexually selected trait that relays honest information about male quality, and laboratory studies generally suggest that female songbirds prefer larger repertoires. However, analysis of field studies across species surprisingly revealed a weak correlation between song elaboration and reproductive success, and it remains unknown why only certain species show this correlation in nature. Taken together, these studies suggest that females in numerous species can detect and prefer larger repertoires in a laboratory setting, but larger individual repertoires correlate with reproductive success only in a subset of these species. This prompts the question: Do the species that show a stronger correlation between reproductive success and larger individual repertoires in nature have anything in common? In this study, we test whether between‐species differences in two song‐related variables—species average syllable repertoire size and adult song stability over time—can be used to predict the importance of individual song elaboration in reproductive success within a species. Our cross‐species meta‐analysis of field studies revealed that species with larger average syllable repertoire sizes exhibited a stronger correlation between individual elaboration and reproductive success than species with smaller syllable repertoires. Song stability versus plasticity in adulthood provided little predictive power on its own, suggesting that the putative correlation between repertoire size and age in open‐ended learners does not explain the association between song elaboration and reproductive success.     

Male reproductive suppression in the cooperatively breeding fish Neolamprologus pulcher

In most cooperative breeders, dominants suppress the reproductionof subordinates. However, two previous studies of Neolamprologuspulcher, a cooperatively breeding cichlid fish, have suggestedthat socially subordinate helper males sneak fertilizationsfrom dominant breeding males. If such sneaking does occur, boththeoretical work and empirical studies of other fish speciessuggest that sperm competition will select for increased reproductiveinvestment by sneaker males, relative to more dominant males.To address these issues, we quantified gonadal investment andsperm characteristics of 41 N. pulcher male breeders and 62male helpers from 55 groups in Lake Tanganyika. Gonadal investmentfollowed patterns consistent with reproductive suppression,with breeders having considerably larger testes masses thanhelpers. Breeders also had faster and longer swimming spermand a higher percentage of motile sperm compared to helpers.However, sperm characteristics of large helpers were similarto those of breeders, but these same helpers had lower testesmasses. Thus, large helpers had sperm that were physiologicallyequivalent to that of breeders, but their relatively small gonadsimply that they were reproductively suppressed.     

Kin selection, not group augmentation, predicts helping in an obligate cooperatively breeding bird

Kin selection theory has been the central model for understanding the evolution of cooperative breeding, where non-breeders help bear the cost of rearing young. Recently, the dominance of this idea has been questioned; particularly in obligate cooperative breeders where breeding without help is uncommon and seldom successful. In such systems, the direct benefits gained through augmenting current group size have been hypothesized to provide a tractable alternative (or addition) to kin selection. However, clear empirical tests of the opposing predictions are lacking. Here, we provide convincing evidence to suggest that kin selection and not group augmentation accounts for decisions of whether, where and how often to help in an obligate cooperative breeder, the chestnut-crowned babbler (Pomatostomus ruficeps). We found no evidence that group members base helping decisions on the size of breeding units available in their social group, despite both correlational and experimental data showing substantial variation in the degree to which helpers affect productivity in units of different size. By contrast, 98 per cent of group members with kin present helped, 100 per cent directed their care towards the most related brood in the social group, and those rearing half/full-sibs helped approximately three times harder than those rearing less/non-related broods. We conclude that kin selection plays a central role in the maintenance of cooperative breeding in this species, despite the apparent importance of living in large groups.     

Negative effects of individual heterozygosity on reproductive success in a wild bird population

The evolutionary consequences of individual genetic diversity are frequently studied by assessing heterozygosity–fitness correlations (HFCs). The prevalence of positive and negative HFCs and the predominance of general versus local effects in wild populations are far from understood, partly because comprehensive studies testing for both inbreeding and outbreeding depression are lacking. We studied a genetically diverse population of blue tits in southern Germany using a genome‐wide set of 87 microsatellites to investigate the relationship between proxies of reproductive success and measures of multilocus and single‐locus individual heterozygosity (MLH and SLH). We used complimentary measures of MLH and partitioned markers into functional categories according to their position in the blue tit genome. HFCs based on MLH were consistently negative for functional loci, whereas correlations were rather inconsistent for loci found in nonfunctional areas of the genome. Clutch size was the only reproductive variable showing a general effect. We found evidence for local effects for three measures of reproductive success: arrival date at the breeding site, the probability of breeding at the study site and male reproductive success. For these, we observed consistent, and relatively strong, negative effects at one functional locus. Remarkably, this marker had a similar effect in another blue tit population from Austria (~400 km to the east). We suggest that a genetic local effect on timing of arrival might be responsible for most negative HFCs detected, with carry‐over effects on other reproductive traits. This effect could reflect individual differences in the distance between overwintering areas and breeding sites.     

Variable fledging age according to group size: trade-offs in a cooperatively breeding bird

Group living can provide individuals with several benefits, including cooperative vigilance and lower predation rates. Individuals in larger groups may be less vulnerable to predation due to dilution effects, efficient detection or greater ability to repel predators. Individuals in smaller groups may consequently employ alternative behavioural tactics to compensate for their greater vulnerability to predators. Here, we describe how pied babbler (Turdoides bicolor) fledging age varies with group size and the associated risk of nestling predation. Nestling predation is highest in smaller groups, but there is no effect of group size on fledgling predation. Consequently, small groups fledge young earlier, thereby reducing the risk of predation. However, there is a cost to this behaviour as younger fledglings are less mobile than older fledglings: they move shorter distances and are less likely to successfully reach the communal roost tree. The optimal age to fledge young appears to depend on the trade-off between reduced nestling predation and increased fledgling mobility. We suggest that such trade-offs may be common in species where group size critically affects individual survival and reproductive success.     

Fine‐scale genetic structure and helping decisions in a cooperatively breeding bird

In animal societies, characteristic demographic and dispersal patterns may lead to genetic structuring of populations, generating the potential for kin selection to operate. However, even in genetically structured populations, social interactions may still require kin discrimination for cooperative behaviour to be directed towards relatives. Here, we use molecular genetics and long‐term field data to investigate genetic structure in an adult population of long‐tailed tits Aegithalos caudatus, a cooperative breeder in which helping occurs within extended kin networks, and relate this to patterns of helping with respect to kinship. Spatial autocorrelation analyses reveal fine‐scale genetic structure within our population, such that related adults of either sex are spatially clustered following natal dispersal, with relatedness among nearby males higher than that among nearby females, as predicted by observations of male‐biased philopatry. This kin structure creates opportunities for failed breeders to gain indirect fitness benefits via redirected helping, but crucially, most close neighbours of failed breeders are unrelated and help is directed towards relatives more often than expected by indiscriminate helping. These findings are consistent with the effective kin discrimination mechanism known to exist in long‐tailed tits and support models identifying kin selection as the driver of cooperation.     

Bateman's principle is reversed in a cooperatively breeding bird

Bateman''s principle is not only used to explain sex differences in mating behaviour, but also to determine which sex has the greater opportunity for sexual selection. It predicts that the relationship between the number of mates and the number of offspring produced should be stronger for males than for females. Yet, it is unclear whether Bateman''s principle holds in cooperatively breeding systems where the strength of selection on traits used in intrasexual competition is high in both sexes. We tested Bateman''s principle in the cooperatively breeding superb starling (Lamprotornis superbus), finding that only females showed a significant, positive Bateman gradient. We also found that the opportunity for selection was on average higher in females, but that its strength and direction oscillated through time. These data are consistent with the hypothesis that sexual selection underlies the female trait elaboration observed in superb starlings and other cooperative breeders. Even though the Bateman gradient was steeper for females than for males, the year-to-year oscillation in the strength and direction of the opportunity for selection likely explains why cooperative breeders do not exhibit sexual role reversal. Thus, Bateman''s principle may not hold in cooperative breeders where both sexes appear to be under mutually strong sexual selection.     

Intermittent breeding is associated with breeding group turnover in a cooperatively breeding bird

Oecologia - Intermittent breeding, in which an adult skips a breeding opportunity, can represent a non-adaptive constraint or an adaptive response to the tradeoff between current and future...     

Brood sex ratio variation in a cooperatively breeding bird

In cooperatively breeding species, the fitness consequences of producing sons or daughters depend upon the fitness impacts of positive (repayment hypothesis) and negative (local competition hypothesis) social interactions among relatives. In this study, we examine brood sex allocation in relation to the predictions of both the repayment and the local competition hypotheses in the cooperatively breeding long-tailed tit Aegithalos caudatus. At the population level, we found that annual brood sex ratio was negatively related to the number of male survivors across years, as predicted by the local competition hypothesis. At an individual level, in contrast to predictions of the repayment hypothesis, there was no evidence for facultative control of brood sex ratio. However, immigrant females produced a greater proportion of sons than resident females, a result consistent with both hypotheses. We conclude that female long-tailed tits make adaptive decisions about brood sex allocation.     

Inbreeding,inbreeding depression,and infidelity in a cooperatively breeding bird*

Inbreeding depression plays a major role in shaping mating systems: in particular, inbreeding avoidance is often proposed as a mechanism explaining extra‐pair reproduction in socially monogamous species. This suggestion relies on assumptions that are rarely comprehensively tested: that inbreeding depression is present, that higher kinship between social partners increases infidelity, and that infidelity reduces the frequency of inbreeding. Here, we test these assumptions using 26 years of data for a cooperatively breeding, socially monogamous bird with high female infidelity, the superb fairy‐wren (Malurus cyaneus). Although inbred individuals were rare (～6% of offspring), we found evidence of inbreeding depression in nestling mass (but not in fledgling survival). Mother–son social pairings resulted in 100% infidelity, but kinship between a social pair did not otherwise predict female infidelity. Nevertheless, extra‐pair offspring were less likely to be inbred than within‐pair offspring. Finally, the social environment (the number of helpers in a group) did not affect offspring inbreeding coefficients or inbreeding depression levels. In conclusion, despite some agreement with the assumptions that are necessary for inbreeding avoidance to drive infidelity, the apparent scarcity of inbreeding events and the observed levels of inbreeding depression seem insufficient to explain the ubiquitous infidelity in this system, beyond the mother–son mating avoidance.     

Kinship affects investment by helpers in a cooperatively breeding bird

Helping behaviour in cooperative breeding systems has been attributed to kin selection, but the relative roles of direct and indirect fitness benefits in the evolution of such systems remain a matter of debate. In theory, helpers could maximize the indirect fitness benefits of cooperation by investing more in broods with whom they are more closely related, but there is little evidence for such fine-scale adjustment in helper effort among cooperative vertebrates. In this study, we used the unusual cooperative breeding system of the long-tailed tit Aegithalos caudatus to test the hypothesis that the provisioning effort of helpers was positively correlated with their kinship to broods. We first use pedigrees and microsatellite genotypes to characterize the relatedness between helpers and breeders from a 14 year field study. We used both pedigree and genetic approaches because long-tailed tits have access to pedigree information acquired through social relationships, but any fitness consequences will be determined by genetic relatedness. We then show using both pedigrees and genetic relatedness estimates that alloparental investment by helpers increases as their relatedness to the recipients of their care increases. We conclude that kin selection has played a critical role in moulding the investment decisions of helpers in this cooperatively breeding species.     

The deterrent effect of bird song in territory defense

Using the responses of territory owners to playback to inferthe territorial function of acoustic signals is common practice.However, difficulties with interpreting the results of suchexperiments have obscured our understanding of territorial signalling.For instance, a stronger response to playback is often interpretedas more aggressive, but there is no consensus as to whetherthis should be in response to the least or most threateningsimulated intruder. Rather than following a gradual increaseor decrease, the relationship between signal intensity and responsestrength may instead describe a peaked curve. We manipulatedbanded wren (Thryophilus pleurostictus) songs to simulate low-,median-, and high-performance singers and used these songs asstimuli in playback experiments. Banded wrens were less likelyto approach the high-performance stimulus compared with thelow- and median-performance stimuli. However, the birds thatdid approach the high-performance stimulus sang more than thosethat approached the low-performance stimulus. In addition, birdswere more likely to match the songs when exposed to the median-and high-performance stimuli compared with the low-performancestimuli, and song matching predicted approach behavior. Theseresults are in accordance with theoretical models of aggressiveencounters in which low-performance opponents are challengedwithout further assessment. Median- and high-performance opponents,however, may require further assessment, and the latter maybe perceived as too intimidating for approach.