Divorce in cooperatively breeding long-tailed tits: a consequence of inbreeding avoidance?

The decision of whether to divorce a breeding partner between reproductive attempts can significantly affect individual fitness. In this paper, we report that 63% of surviving pairs of long-tailed tits Aegithalos caudatus divorced between years. We examine three likely explanations for the high divorce rate in this cooperative breeder. The 'better option' hypothesis predicts that divorce and re-pairing increases an individual's reproductive success. However, divorcees did not secure better partners or more helpers and there was no improvement in their reproductive success following divorce. The 'inbreeding avoidance' hypothesis predicts that females should disperse from their family group to avoid breeding with philopatric sons. The observed pattern of divorce was consistent with this hypothesis because, in contrast to the usual avian pattern, divorce was typical for successful pairs (81%) and less frequent in unsuccessful pairs (36-43%). The 'forced divorce' hypothesis predicts that divorce increases as the number of competitors increases. The pattern of divorce among failed breeders was consistent with this hypothesis, but it fails to explain the overall occurrence of divorce because divorcees rarely re-paired with their partners' closest competitors. We discuss long-tailed tits' unique association between divorce and reproductive success in the context of dispersal strategies for inbreeding avoidance

Coordinated parental provisioning is related to feeding rate and reproductive success in a songbird

In species with biparental care, there is sexual conflict over parental investment because each parent benefits when their partner bears more of the reproductive costs. Such conflict can be costly for offspring, but recent theoretical work predicts that parents can resolve sexual conflict through behavioral negotiation, specifically by alternating their trips to provision nestlings. However, this idea has received almost no empirical attention. In this study, we test the hypothesis that parents alternate their delivery of food to offspring in long-tailed tits (Aegithalos caudatus) and investigate whether this coordination of parental care is associated with greater reproductive success. We show that parents alternate provisioning trips more than would be expected by chance and that parental alternation is repeatable across multiple observation periods at a nest. More alternation is associated with increased visit synchrony and increased food delivery to nestlings. Moreover, we found that nests with more alternation were less likely to be predated, probably resulting from reduced activity around the nest when parents coordinate their provisioning behavior. Our results support the hypothesis that alternation of offspring provisioning is a behavioral adaptation to reduce the costs of sexual conflict

Incubation in a temperate passerine: do environmental conditions affect incubation period duration and hatching success?

The timing of breeding often has a profound influence on the reproductive success of birds living in seasonal environments with rapidly changing nestling food availability. Timing is typically investigated with reference to lay dates, but it is the time of hatching that determines the ambient conditions and food availability that nestlings experience. Thus, in addition to lay date, phenological studies may also have to take account of variation in the length of the incubation period, which is likely to depend on both environmental conditions and parental traits. The primary aim of this study was to use a 24-year dataset to investigate the abiotic and biotic factors influencing variation in incubation duration in long-tailed tits (Aegithalos caudatus), a species in which incubation duration varies substantially (range: 12–26 days). We found support for our predictions that drier conditions, later breeding attempts and larger clutches were associated with shorter incubation periods. Larger clutches were also more resilient to increases in incubation duration associated with wet conditions. Surprisingly, warmer ambient conditions were associated with longer incubation periods. Secondly, we assessed the consequences of variation in the length of incubation periods for the risk of nest predation and the hatching success of surviving clutches. We show that longer incubation periods are likely to be costly due to increased exposure to nest predators. In contrast, we found only marginal effects of environmental conditions or incubation duration on hatching success, implying that wet conditions cause slower embryo growth and hence longer incubation periods, rather than causing embryo fatality. We suggest that long-tailed tits’ nests and parental behavior protect eggs from mortality arising directly from adverse weather conditions

Sex Allocation Patterns across Cooperatively Breeding Birds Do Not Support Predictions of the Repayment Hypothesis

The repayment hypothesis predicts that reproductive females in cooperative breeding systems overproduce the helping sex. Thanks to well-documented examples of this predicted sex ratio bias, repayment has been considered an important driver of variation in sex allocation patterns. Here we test this hypothesis using data on population brood sex ratios and facultative sex allocation from 28 cooperatively breeding bird species. We find that biased sex ratios of helpers do not correlate with production biases in brood sex ratios, contrary to predictions. We also test whether females facultatively produce the helping sex in response to a deficiency of help (i.e., when they have fewer or no helpers). Although this is observed in a few species, it is not a significant trend overall, with a mean effect size close to zero. We conclude that, surprisingly, repayment does not appear to be a widespread influence on sex ratios in cooperatively breeding birds. We discuss possible explanations for our results and encourage further examination of the repayment model

Context-dependent strategies of food allocation among offspring in a facultative cooperative breeder

Natural selection should favor adoption of parental strategies that maximize fitness when allocating investment among offspring. In birds, begging displays often convey information of nestling need and quality, allowing parents to make adaptive food allocation decisions. We investigated how adults utilized cues likely to represent nestling competitive ability (begging position) and need (begging intensity) and a cue independent of nestling control (nestling sex) to distribute food among nestlings in a facultative cooperative breeder, the black-throated tit (Aegithalos concinnus). We found that parents reduced their efforts when helped, suggesting that parents of helped broods would have the potential to satisfy nestling needs more than unhelped parents. This suggestion was supported by the fact that nestling mass increased faster in helped than in unhelped nests. We found no effect of nestling sex on food allocation, but, as predicted, we found that adults responded differently to begging signals in relation to the presence of helpers and brood size. First, helped parents were more responsive to nestling begging intensity than parents without helpers. Second, female parents and helpers had a stronger preference for nestling begging position in large than in small broods. Third, the preference for nestling begging position was greater for unhelped than for helped female parents. These results provide evidence that carers adjust their preference for different offspring begging signals based on availability of food resources

Who to help? Helping decisions in a cooperatively breeding bird with redirected care

Cooperative breeding sometimes occurs when adult breeders form groups following natal dispersal and mating. In such cases, individuals typically face a choice of social partner with whom to cooperate. Selecting appropriate social partners is crucial to maximising the fitness payoffs from cooperation, but our understanding of the criteria guiding partner choice is limited. Here, we analyse helping decisions by long-tailed tits (Aegithalos caudatus), which may redirect their care to assist breeders in raising offspring following the failure of their own nests. In this species, helpers prefer to help relatives at nearby nests, but it is unclear whether other criteria that may affect helper fitness also influence helping decisions. When choosing among broods of equivalent kinship, we found that helpers did not prefer those broods that offered the greatest indirect fitness returns. Further analyses revealed that helpers did not choose nests on the basis of brood size or age, but were more likely to help broods that were closer to their own failed nests and that were already being cared for by other helpers. Both effects likely reflect the limited choice available to helpers: although individuals breed close to relatives within kin neighbourhoods, a high rate of nest predation constrains helpers’ choice of broods. In other species where cooperatively breeding groups form after natal dispersal, a greater range of options may be available and here detailed analysis of group formation will be helpful for determining the decision rules that underpin partner choice and permit stable cooperation in the face of alternative options. Significance statement Cooperative breeding occurs most frequently when offspring delay dispersal from their natal site and help to care for their younger siblings. In some species, however, individuals first disperse and then come together as adults to cooperate in rearing young. In the latter case, multiple social partners may be available — what then determines which partner is helped? We studied partner choice in long-tailed tits, which may help to feed other broods if their own brood fails. When multiple related broods were available, individuals were more likely to help those close by but showed no preference for broods offering the greatest indirect fitness returns. One explanation for this result is that helping options for most individuals are limited by high levels of nest predation, favouring a simpler decision-making process based on identifying close relatives breeding in close proximity

Microsatellite characterisation and sex-typing in two invasive parakeet species, the monk parakeet Myiopsitta monachus and ring-necked parakeet Psittacula krameri

Invasive species can have wide-ranging negative impacts, and an understanding of the process and success of invasions can be vital to determine management strategies, mitigate impacts and predict range expansions of such species. Monk parakeets (Myiopsitta monachus) and ring-necked parakeets (Psittacula krameri) are both widespread invasive species, but there has been little research into the genetic and social structure of these two species despite the potential links with invasion success. The aim of this study was to isolate novel microsatellite loci from the monk parakeet and characterise them in both monk and ring-necked parakeets in order to facilitate future investigations into their behaviour and population ecology. Sex-typing markers were also tested in both species. Of the 20 microsatellite loci assessed in 24 unrelated monk parakeets, 16 successfully amplified and were polymorphic displaying between 2 and 14 alleles (mean = 8.06). Expected heterozygosity ranged from 0.43 to 0.93 and observed heterozygosity ranged from 0.23 to 0.96. Nine of the 20 loci also successfully amplified and were polymorphic in the ring-necked parakeet, displaying between 2 and 10 alleles. Suitable markers to sex both species and a Z-linked microsatellite locus were identified. A multiplex marker set was validated for monk parakeets. These novel microsatellite loci will facilitate fine and broad-scale population genetic analyses of these two widespread invasive species

Individual variation explains ageing patterns in a cooperatively breeding bird, the long-tailed tit Aegithalos caudatus

1. Alloparental care in cooperatively breeding species may alter breeder age-specific survival and reproduction and subsequently senescence. The helping behaviour itself might also undergo age-related change, and decisions to help in facultative cooperative breeders are likely to be affected by individual condition. 2. Helpers in long-tailed tits Aegithalos caudatus assist relatives after failing to raise their own brood, with offspring from helped nests being more likely to recruit into the breeding population. 3. Using data collected over 25 years, we examined the age trajectories of survival and reproduction in adult long-tailed tits to determine how these were affected by the presence or absence of helpers and how helper behaviour changed with age. 4. There was evidence for increased reproductive performance with breeder age, but no effect of age on the probability of survival. We found no evidence of significant senescent decline in survival or reproductive performance, although individuals accrued less inclusive fitness in their last year of life. Lifetime reproductive success was positively related to both reproductive life span and body mass. Within a season, breeders that were assisted by helpers enjoyed greater reproductive success through enhanced offspring recruitment in the following year. We found no evidence that age affected an individual's propensity to help, or the amount of indirect fitness accrued through helping. 5. We found a positive correlation between life span and multiple components of reproductive success, suggesting that individual variation in quality underpins age-related variation in fitness in this species. Helping decisions are driven by condition, and lifetime inclusive fitness of immigrants was predicted by body mass. These findings further support individual heterogeneity in quality being a major driver for fitness gains across the life course of long-tailed tits

Sex and age effects on monk parakeet home-range variation in the urban habitat

Home-range size is a key aspect of space-use, and variation in home-range size and structure may have profound consequences for the potential impact of damage and control strategies for invasive species. However, knowledge on home-range structure of naturalized parrot species is very limited. The aim of this study was to quantify patterns of home-range variation according to sex and age of the monk parakeet Myiopsitta monachus, an invasive parakeet in Europe. Mean kernel home-range size was 12.4 ± 1.22 ha (range 1.7–74.1 ha; N = 73 birds). Juveniles had a larger home-range size than adults, but sexes did not differ in kernel home-range size. The mean maximum distance moved by monk parakeets was 727 ± 37.0 m (range: 150–1581 m), and it was not dependent on either the sex or age of the birds. Having a small home range is one of the conditions for the feasible eradication of an invasive species; hence, the small home range of urban monk parakeets that we report here is good news for pest managers. However, this small home-range size can limit the effectiveness of culling operations with traps or feeders with contraceptives or poison, and other alternatives, such as funnel nets or traps, should be used