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Article ; Online: Individual variation explains ageing patterns in a cooperatively breeding bird, the long-tailed tit Aegithalos caudatus.

Roper, Mark / Sturrock, Nicole J / Hatchwell, Ben J / Green, Jonathan P

2022 Volume 91, Issue 7, Page(s) 1521–1534

Abstract: 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 ...

Abstract	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. 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. 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. 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. 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.
MeSH term(s)	Aging/physiology ; Animals ; Cooperative Behavior ; Longevity ; Reproduction ; Songbirds/growth & development
Language	English
Publishing date	2022-05-24
Publishing country	England
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	3024-7
ISSN	1365-2656 ; 0021-8790
ISSN (online)	1365-2656
ISSN	0021-8790
DOI	10.1111/1365-2656.13741
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Article ; Online: Inclusive fitness consequences of dispersal decisions in a cooperatively breeding bird, the long-tailed tit (

Green, Jonathan P / Hatchwell, Ben J

Proceedings of the National Academy of Sciences of the United States of America

2018 Volume 115, Issue 47, Page(s) 12011–12016

Abstract: Natal dispersal is a demographic trait with profound evolutionary, ecological, and behavioral consequences. However, our understanding of the adaptive value of dispersal patterns is severely hampered by the difficulty of measuring the relative fitness ... ...

Abstract	Natal dispersal is a demographic trait with profound evolutionary, ecological, and behavioral consequences. However, our understanding of the adaptive value of dispersal patterns is severely hampered by the difficulty of measuring the relative fitness consequences of alternative dispersal strategies in natural populations. This is especially true in social species, in which natal philopatry allows kin selection to operate, so direct and indirect components of inclusive fitness have to be considered when evaluating selection on dispersal. Here, we use lifetime reproductive success data from a long-term study of a cooperative breeder, the long-tailed tit
MeSH term(s)	Animal Migration/physiology ; Animals ; Behavior, Animal/physiology ; Biological Evolution ; Breeding ; Cooperative Behavior ; Female ; Gene Flow/physiology ; Male ; Mating Preference, Animal/physiology ; Passeriformes/physiology ; Population Dynamics ; Reproduction ; Sexual Behavior, Animal/physiology ; Songbirds/physiology
Language	English
Publishing date	2018-11-05
Publishing country	United States
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	209104-5
ISSN	1091-6490 ; 0027-8424
ISSN (online)	1091-6490
ISSN	0027-8424
DOI	10.1073/pnas.1815873115
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Article ; Online: Coordination of care is facilitated by delayed feeding and collective arrivals in the long-tailed tit

Halliwell, Chay / Beckerman, Andrew P. / Biddiscombe, Sarah J. / Germain, Marion / Hatchwell, Ben J.

Animal Behaviour.

2023

Abstract: When multiple carers invest in a shared brood, there is likely to be conflict among individuals over how much each carer invests. This conflict results in suboptimal investment to the detriment of all carers. It has been proposed that conditional ... ...

Abstract	When multiple carers invest in a shared brood, there is likely to be conflict among individuals over how much each carer invests. This conflict results in suboptimal investment to the detriment of all carers. It has been proposed that conditional cooperation, that is, 'turn taking' or 'alternation', may resolve this conflict by preventing exploitation. This contentious idea has received some empirical support, but distinguishing active alternation from that expected via passive processes has proved challenging. The aim of this study was to use detailed observations of provisioning to examine whether carers at biparental (parents only) and cooperative (parents and helpers) nests of the long-tailed tit, Aegithalos caudatus, behave in a context-dependent manner that increases the level of alternation. First, we show that carers that had been the last to feed waited near the nest (loitering) for longer before feeding when they next arrived at the nest and allowed others to feed first, thus facilitating alternation. Second, we found that the arrival of carers near the nest and their subsequent feeds were tightly synchronized, with overlapping loitering periods, allowing them to monitor the effort of other carers. Finally, we show that measures of coordination were influenced by carers arriving in a status-dependent order, with breeding females consistently arriving first and helpers last. Together, these results show how patterns of alternation and synchrony arise in long-tailed tits, and reveal the behavioural mechanisms underpinning coordination of care.
Keywords	Aegithalos ; animal behavior ; nests ; alternation ; conflict ; cooperation ; coordination ; parental care ; synchrony
Language	English
Publishing place	Elsevier Ltd
Document type	Article ; Online
Note	Pre-press version ; Use and reproduction
ZDB-ID	281-1
ISSN	0003-3472
ISSN	0003-3472
DOI	10.1016/j.anbehav.2023.04.005
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Article ; Online: Kith or Kin? Familiarity as a Cue to Kinship in Social Birds

Amy E. Leedale / Jianqiang Li / Ben J. Hatchwell

Frontiers in Ecology and Evolution, Vol

2020 Volume 8

Abstract: Interacting with relatives provides opportunities for fitness benefits via kin-selected cooperation, but also creates potential costs through kin competition and inbreeding. Therefore, a mechanism for the discrimination of kin from non-kin is likely to ... ...

Abstract	Interacting with relatives provides opportunities for fitness benefits via kin-selected cooperation, but also creates potential costs through kin competition and inbreeding. Therefore, a mechanism for the discrimination of kin from non-kin is likely to be critical for individuals of many social species to maximize their inclusive fitness. Evidence suggests that genetic cues to kinship are rare and that learned or environmental cues offer a more parsimonious explanation for kin recognition in most contexts. This is particularly true among cooperatively breeding birds, where recognition of familiar individuals is usually regarded as the most plausible mechanism for kin discrimination. In this article, we first review the evidence that familiarity provides an effective decision rule for discrimination of kin from non-kin in social birds. We then consider some of the complexities of familiarity as a cue to kinship, especially the problems of how individuals become familiar, and how familiar individuals are recognized. We conclude that while familiarity as a mechanism for kin recognition may be more parsimonious and widespread than genetic mechanisms, its apparent simplicity as a decision rule governing social interactions may be deceptive. Finally, we identify directions for future research on familiarity as a kin recognition mechanism in social birds and other taxa.
Keywords	kin discrimination ; kin recognition ; cooperation ; familiarity ; social birds ; Evolution ; QH359-425 ; Ecology ; QH540-549.5
Subject code	590
Language	English
Publishing date	2020-04-01T00:00:00Z
Publisher	Frontiers Media S.A.
Document type	Article ; Online
Database	BASE - Bielefeld Academic Search Engine (life sciences selection)

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Article ; Online: Fine-scale genetic structure reflects limited and coordinated dispersal in the colonial monk parakeet, Myiopsitta monachus.

Dawson Pell, Francesca S E / Senar, Juan Carlos / Franks, Daniel W / Hatchwell, Ben J

Molecular ecology

2021 Volume 30, Issue 6, Page(s) 1531–1544

Abstract: The genetic structure of animal populations has considerable behavioural, ecological and evolutionary implications and may arise from various demographic traits. Here, we use observational field data and molecular genetics to determine the genetic ... ...

Abstract	The genetic structure of animal populations has considerable behavioural, ecological and evolutionary implications and may arise from various demographic traits. Here, we use observational field data and molecular genetics to determine the genetic structure of an invasive population of monk parakeets, Myiopsitta monachus, at a range of spatial scales, and investigate the demographic processes that generate the observed structure. Monk parakeets construct large nests that can house several pairs occupying separate chambers; these nests are often aggregated within nesting trees. We determined patterns of relatedness within compound nests, within nesting trees and between trees. Spatial autocorrelation analyses of pairwise genetic relatedness revealed fine-scale genetic structure with relatives of both sexes spatially clustered within, but not beyond, nesting trees. In addition, males were more related to males sharing their compound nests than to other males occupying the same nesting tree. By contrast, males and females within compound nests were not significantly more closely related than elsewhere in the same tree, and we found no evidence for inbreeding. Adults showed high breeding site fidelity between years despite considerable disturbance of nest sites. Natal dispersal was female-biased, but dispersal distances were relatively short with some natal philopatry observed in both sexes. Sibling coalitions, typically of males, were observed amongst both philopatric and dispersing birds. Our results show significant clustering of kin within compound nests and nesting trees resulting from limited and coordinated natal dispersal, with subsequent breeding site fidelity. The resulting genetic structure has implications for social behaviour in this unusual parrot species.
MeSH term(s)	Animals ; Biological Evolution ; Female ; Male ; Microsatellite Repeats/genetics ; Parakeets/genetics ; Parrots ; Plant Breeding
Language	English
Publishing date	2021-02-09
Publishing country	England
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	1126687-9
ISSN	1365-294X ; 0962-1083
ISSN (online)	1365-294X
ISSN	0962-1083
DOI	10.1111/mec.15818
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Article: Individual variation explains ageing patterns in a cooperatively breeding bird, the long‐tailed tit Aegithalos caudatus

Roper, Mark / Sturrock, Nicole J. / Hatchwell, Ben J. / Green, Jonathan P.

journal of animal ecology. 2022 July, v. 91, no. 7

2022

Abstract: 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 ...

Abstract	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. 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. 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. 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. 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.
Keywords	Aegithalos ; adults ; alloparental behavior ; birds ; body weight ; decline ; longevity ; probability ; progeny ; reproductive performance ; reproductive success
Language	English
Dates of publication	2022-07
Size	p. 1521-1534.
Publishing place	John Wiley & Sons, Ltd
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	3024-7
ISSN	1365-2656 ; 0021-8790
ISSN (online)	1365-2656
ISSN	0021-8790
DOI	10.1111/1365-2656.13741
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Article: Who to help? Helping decisions in a cooperatively breeding bird with redirected care

Sturrock, Nicole J. / Hatchwell, Ben J. / Firth, Josh A. / Green, Jonathan P.

Behavioral ecology and sociobiology. 2022 June, v. 76, no. 6

2022

Abstract: 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 ... ...

Abstract	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.
Keywords	Aegithalos ; birds ; decision making ; kinship ; nests ; predation ; progeny ; sociobiology
Language	English
Dates of publication	2022-06
Size	p. 83.
Publishing place	Springer Berlin Heidelberg
Document type	Article
ZDB-ID	194510-5
ISSN	1432-0762 ; 0340-5443
ISSN (online)	1432-0762
ISSN	0340-5443
DOI	10.1007/s00265-022-03190-w
Database	NAL-Catalogue (AGRICOLA)

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Article ; Online: Helping decisions and kin recognition in long-tailed tits: is call similarity used to direct help towards kin?

Leedale, Amy E / Lachlan, Robert F / Robinson, Elva J H / Hatchwell, Ben J

Philosophical transactions of the Royal Society of London. Series B, Biological sciences

2020 Volume 375, Issue 1802, Page(s) 20190565

Abstract: Most cooperative breeders live in discrete family groups, but in a minority, breeding populations comprise extended social networks of conspecifics that vary in relatedness. Selection for effective kin recognition may be expected for more related ... ...

Abstract	Most cooperative breeders live in discrete family groups, but in a minority, breeding populations comprise extended social networks of conspecifics that vary in relatedness. Selection for effective kin recognition may be expected for more related individuals in such kin neighbourhoods to maximize indirect fitness. Using a long-term social pedigree, molecular genetics, field observations and acoustic analyses, we examine how vocal similarity affects helping decisions in the long-tailed tit
MeSH term(s)	Animal Communication ; Animals ; Auditory Perception ; Cooperative Behavior ; Cues ; Decision Making ; Helping Behavior ; Male ; Nesting Behavior ; Songbirds/physiology
Language	English
Publishing date	2020-05-18
Publishing country	England
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	208382-6
ISSN	1471-2970 ; 0080-4622 ; 0264-3839 ; 0962-8436
ISSN (online)	1471-2970
ISSN	0080-4622 ; 0264-3839 ; 0962-8436
DOI	10.1098/rstb.2019.0565
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Article ; Online: Mechanistic home range analysis reveals drivers of space use patterns for a non-territorial passerine.

Ellison, Natasha / Hatchwell, Ben J / Biddiscombe, Sarah J / Napper, Clare J / Potts, Jonathan R

The Journal of animal ecology

2020 Volume 89, Issue 12, Page(s) 2763–2776

Abstract: Home ranging is a near-ubiquitous phenomenon in the animal kingdom. Understanding the behavioural mechanisms that give rise to observed home range patterns is thus an important general question, and mechanistic home range analysis (MHRA) provides the ... ...

Abstract	Home ranging is a near-ubiquitous phenomenon in the animal kingdom. Understanding the behavioural mechanisms that give rise to observed home range patterns is thus an important general question, and mechanistic home range analysis (MHRA) provides the tools to address it. However, such analysis has hitherto been principally restricted to scent-marking territorial animals, so its potential breadth of application has not been tested. Here, we apply MHRA to a population of long-tailed tits Aegithalos caudatus, a non-territorial passerine, in the non-breeding season where there is no clear 'central place' near which birds need to remain. The aim is to uncover the principal movement mechanisms underlying observed home range formation. Our foundational models consist of memory-mediated conspecific avoidance between flocks, combined with attraction to woodland. These are then modified to incorporate the effects of flock size and relatedness (i.e. kinship), to uncover the effect of these on the mechanisms of home range formation. We found that a simple model of spatial avoidance, together with attraction to the central parts of woodland areas, accurately captures long-tailed tit home range patterns. Refining these models further, we show that the magnitude of spatial avoidance by a flock is negatively correlated to both the relative size of the flock (compared to its neighbour) and the relatedness of the flock with its neighbour. Our study applies MHRA beyond the confines of scent-marking, territorial animals, so paves the way for much broader taxonomic application. These could potentially help uncover general properties underlying the emergence of animal space use patterns. This is also the first study to apply MHRA to questions of relatedness and flock size, thus broadening the potential possible applications of this suite of analytic techniques.
MeSH term(s)	Animals ; Homing Behavior ; Movement ; Passeriformes ; Pheromones ; Territoriality
Chemical Substances	Pheromones
Language	English
Publishing date	2020-08-10
Publishing country	England
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	3024-7
ISSN	1365-2656 ; 0021-8790
ISSN (online)	1365-2656
ISSN	0021-8790
DOI	10.1111/1365-2656.13292
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Article ; Online: The evolution of cooperative breeding in birds: kinship, dispersal and life history.

Hatchwell, Ben J

Philosophical transactions of the Royal Society of London. Series B, Biological sciences

2009 Volume 364, Issue 1533, Page(s) 3217–3227

Abstract: The evolution of cooperation among animals has posed a major problem for evolutionary biologists, and despite decades of research into avian cooperative breeding systems, many questions about the evolution of their societies remain unresolved. A review ... ...

Abstract	The evolution of cooperation among animals has posed a major problem for evolutionary biologists, and despite decades of research into avian cooperative breeding systems, many questions about the evolution of their societies remain unresolved. A review of the kin structure of avian societies shows that a large majority live in kin-based groups. This is consistent with the proposed evolutionary routes to cooperative breeding via delayed dispersal leading to family formation, or limited dispersal leading to kin neighbourhoods. Hypotheses proposed to explain the evolution of cooperative breeding systems have focused on the role of population viscosity, induced by ecological/demographic constraints or benefits of philopatry, in generating this kin structure. However, comparative analyses have failed to generate robust predictions about the nature of those constraints, nor differentiated between the viscosity of social and non-social populations, except at a coarse level. I consider deficiencies in our understanding of how avian dispersal strategies differ between social and non-social species, and suggest that research has focused too narrowly on population viscosity and that a broader perspective that encompasses life history and demographic processes may provide fresh insights into the evolution of avian societies.
MeSH term(s)	Animals ; Biological Evolution ; Birds/physiology ; Cooperative Behavior ; Family ; Female ; Male ; Phylogeny ; Sexual Behavior, Animal
Language	English
Publishing date	2009-10-05
Publishing country	England
Document type	Journal Article
ZDB-ID	208382-6
ISSN	1471-2970 ; 0080-4622 ; 0264-3839 ; 0962-8436
ISSN (online)	1471-2970
ISSN	0080-4622 ; 0264-3839 ; 0962-8436
DOI	10.1098/rstb.2009.0109
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