Stepping stones of life: natal dispersal in the group-living but noncooperative speckled warbler

In most cooperatively breeding birds the offspring of one sex, usually male, delays dispersal to remain on the natal territory and helps its parents to rear subsequent young. Thus delayed dispersal could be the first step in the evolution of cooperative breeding. We studied natal dispersal in a population of the group-living speckled warbler, Chthonicola sagittata, based on observations of a colour-banded population over 3 years. Unlike other group-living members of the Acanthizinae, all juvenile males in this population dispersed to settle on foreign territories as subordinates, which do not help rear the young. Speckled warblers showed all the life history traits that are thought to result in a saturated habitat and lead to delayed dispersal: they were sedentary, had high adult survival and had a male-biased sex ratio. However, they differed from other acanthizids in occurring at low density (0.18 birds/ha) on large breeding territories (6-12 ha), with a maximum of two males per territory. This may allow subordinates to live on foreign territories yet avoid aggression from dominants. A benefit of dispersal is that it provides an additional route to gaining a breeding vacancy. Dispersers can acquire vacancies on their new territory or on a neighbour's, but incest avoidance would be likely to constrain nondispersing males to neighbours' territories. A model of relative lifetime success showed that the survival benefits of natal philopatry are unlikely to outweigh this benefit of dispersal.     

Increased lifetime reproductive success for Siberian jay (Perisoreus infaustus) males with delayed dispersal

In a crowded environment the natal territory could serve as a haven for young and inexperienced offspring until a breeding vacancy emerges. Delayed dispersal and association with kin could then offer adaptive benefits through an individual fitness gain. Here we report that delayed dispersal is associated with a higher lifetime individual fitness in Siberian jay (Perisoreus infaustus) males. Sons bred more successfully and had more reproductive events in life when they delayed dispersal. The higher lifetime reproductive success when sons disperse later in life is sufficient to promote postponement of natal dispersal, suggesting that dispersal is delayed due to ecological constraints on access to high-quality habitats. We argue that the maintenance of this variation in the timing of dispersal and reproductive success can be reconciled with non-genetic mechanisms driving dispersal. Social dominance within broods reflecting environmental conditions during growth is such a mechanism.     

Juvenile helping in the moorhen,Gallinula chloropus

In a 3-year study of the moorhen, Gallinula chloropus, some chicks from first broods stayed on their natal territory once they had reached independence, and helped to rear their younger, second brood, siblings. Juvenile dispersal was constrained by habitat saturation, and first brood young were forced to stay on their natal territory. Juveniles that hatched early in the year were forced to stay longer, and helped more than those that hatched late. The total feeding rates to broods with and without juvenile helpers were the same, but parents with helpers reduced their feeding rates relative to parents without helpers. Pairs with helpers (=pairs attempting second broods) reared more chicks per nesting, attempt than pairs rearing chicks at the same time of year without helpers (=pairs attempting first clutch renests). This was true both for all clutches, and for hatched clutches only, even when controlling for parental quality, territory size and scasonal effects.     

NATAL DISPERSAL AND POPULATION STRUCTURE IN A MIGRATORY SONGBIRD,THE INDIGO BUNTING

Male and female indigo buntings (Passerina cyanea) were equally likely to return and breed in their natal areas. The distances and number of territories between natal and breeding sites were similar between the sexes. The breeding site was not limited by prior occupation of sites by breeding adults closer to the natal territory. Dispersal distances were independent of whether parents or siblings of either sex returned in the same year. Variation in distance within the natal area appeared to be independent of local population density, social competition, active kin recognition, and avoidance of incest, but was affected by date of birth. The local distribution of dispersal distances approximated a neutral model with a decreasing probability of settling with distance from the natal site. A population inbreeding coefficient ΔF of 0.01 was estimated from known genealogies and matings. The proportion of nestlings recaptured in a later year on their natal area was 1.65% of 1,212 nestlings at the George Reserve and 8.78% of 1,332 nestlings at Niles. Locally born birds comprised 1.6% and 13.0% of the breeding population in areas of 10 and 4 km². Evidently most buntings settle and breed more than 2 km from their natal site. The results are inconsistent with predictions of social and avoidance-of-inbreeding hypotheses that are based on the local natal demographic environment. The results are consistent with a model of neutral dispersal within a genetically open population.     

Parent presence, delayed dispersal, and territory acquisition in the Seychelles warbler

The presence of parents in the natal territory may play an important,but often overlooked, role in natal dispersal and the consequentacquisition of a territory. Living with parents in a territorymay confer a fitness advantage to subordinates through, forexample, the nepotistic behavior of the parents or indirectbenefits gained by helping to raise nondescendent kin. Whena parent is replaced by a stepparent, such advantages are reducedor disappear and, as a result, subordinates may disperse. Subordinatesthat disperse after parent replacement may be constrained intheir timing of dispersal, which could have negative fitnessconsequences. In the cooperatively breeding Seychelles warbler,we show that when a parent was naturally replaced or experimentallyremoved and subsequently replaced by a stepparent from outsidethe territory, subordinates were more likely to disperse thanwhen both parents remained in the natal territory. Furthermore,subordinates dispersing from territories in which one or bothparents had been replaced were less likely to acquire a breederposition than subordinates dispersing when both parents werestill on the natal territory. Our findings suggest that thepresence of parents in the natal territory may promote delayeddispersal and facilitate the eventual acquisition of a breederposition outside the natal territory. Our results support theidea that the prolonged parental care, which long-lived speciesare able to provide, may have selected for family living.     

Habitat fragmentation shapes natal dispersal and sociality in an Afrotropical cooperative breeder

It remains poorly understood how effects of anthropogenic activity, such as large-scale habitat fragmentation, impact sociality in animals. In cooperatively breeding species, groups are mostly formed through delayed offspring dispersal, and habitat fragmentation can affect this process in two opposite directions. Increased habitat isolation may increase dispersal costs, promoting delayed dispersal. Alternatively, reduced patch size and quality may decrease benefits of philopatry, promoting dispersal. Here, we test both predictions in a cooperatively breeding bird (placid greenbul, Phyllastrephus placidus) from an Afrotropical cloud forest archipelago. Males born in fragmented forest dispersed about 1 year earlier than those born in continuous forest. Contrary to females, males also started to reproduce earlier and mostly settled within their natal patch. Females only rarely delayed their dispersal for more than 1 year, both in fragmented and continuous forests. Our results suggest that early male dispersal and reproduction is jointly driven by a decrease in the value of the natal territory and an increase in local breeding opportunities in fragmented forest. While plasticity in dispersal strategies of cooperative breeders in response to anthropogenic change is believed to optimize reproduction-survival trade-offs, to what extent it shapes the ability of species to respond to rapid environmental change remains to be studied.     

Dispersal,ranging and settling behaviour of Marsh Tits Poecile palustris in a fragmented landscape in lowland England

Capsule Natal dispersal was rapid and distances were short. Winter ranging and breeding dispersal were limited. Few birds undertook large movements.

Aims To investigate the natal and breeding dispersal of Marsh Tits, including the timing of dispersal movements.

Methods Nestlings, juveniles and adults were ringed and searched for over 4500 ha during summer, autumn–winter, and spring over six years. Dispersal distances were measured as metric distances and multiples of territory widths. Ranging distances were compared with dispersal distances.

Results Median distances of natal dispersal were 2.6 territory widths for males (704.5 m) and 3.1 territory widths for females (1065.0 m). Median distances of breeding dispersal were 0.2 territory widths for males and females (58.6 and 53.1 m respectively). Most natal dispersal was completed soon after independence, with further movement in spring. Breeding dispersal was also detected during these periods. Median ranging distances were short, and some winter floaters were identified.

Conclusion Marsh Tits had short dispersal distances, with most dispersal activity occurring in June. Results suggested that dispersal behaviour was sensitive to habitat fragmentation, resulting in poor settling success outside of the natal wood. Habitat fragmentation may, therefore, be a contributory factor in the decline of the Marsh Tit population in Britain.     

Tracking Natal Dispersal in a Coastal Population of a Migratory Songbird Using Feather Stable Isotope (δ2H, δ34S) Tracers

Adult birds tend to show high fidelity to their breeding territory or disperse over relatively short distances. Gene flow among avian populations is thus expected to occur primarily through natal dispersal. Although natal dispersal is a critical demographic process reflecting the area over which population dynamics take place, low recapture rates of birds breeding for the first time have limited our ability to reliably estimate dispersal rates and distances. Stable isotope approaches can elucidate origins of unmarked birds and so we generated year- and age-specific δ²H and δ³⁴S feather isoscapes (ca. 180 000 km²) of coastal-breeding Ovenbirds (Seiurus aurocapilla) and used bivariate probability density functions to assign the likely natal areas of 35 males recruited as first-year breeders into a population located in northwestern New Brunswick, Canada. Most individuals (80–94% depending on the magnitude of an age correction factor used; i.e. 28–33 out of 35) were classified as residents (i.e. fledged within our study area) and estimated minimum dispersal distances of immigrants were between 40 and 240 km. Even when considering maximum dispersal distances, the likely origin of most first-year breeders was<200 km from our study area. Our method identified recruitment into our population from large geographic areas with relatively few samples whereas previous mark-recapture based methods have required orders of magnitude more individuals to describe dispersal at such geographic scales. Natal dispersal movements revealed here suggest the spatial scale over which many population processes are taking place and we suggest that conservation plans aiming to maintain populations of Ovenbirds and ecologically-similar species should consider management units within 100 or at most 200 km of target breeding populations.     

Direct fitness benefits of group living in a complex cooperative society of carrion crows, Corvus corone corone

The social behaviour of carrion crows varies between populations. In northern Spain cooperatively breeding groups form through delayed natal dispersal and/or immigration of individuals (usually males) into the territory. In this population, carrion crows therefore breed as either unassisted pairs, pairs with nondispersing 1-2-year-old helpers (nondispersers), pairs with immigrant helpers or mixed groups (pairs with both immigrants and nondispersers). We used a microsatellite-based genotyping system to determine the parentage of 57 nestlings (19 broods). Polygamous mating was involved in 26% of the broods and reproduction was shared among group members of both sexes in at least three groups. Immigrants of both sexes can therefore gain access to mates by living in a group, while reproduction is unlikely to involve nondispersers. This implies that nondispersers and immigrants gain different sorts of benefits from group living and helping at the nest. Our genetic data confirmed that nondispersers associated with their parents on the natal territory and therefore that delayed natal dispersal leads to family formation in the carrion crow. Polygamous mating was not found in groups without immigrants, suggesting that, in this population, breeders lose parentage in their brood when sociality is extended beyond the limit of the nuclear family.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Malformed offspring,sibling matings,and selection against inbreeding in the sand lizard (Lacerta agilis)

We demonstrated that sand lizards (Lacerta agilis) are more likely to have malformed offspring when they mate with siblings. Offspring with malformations, such as deformed limbs and heads, have zero survival in a natural population. Normal-looking siblings of malformed hatchlings also had a reduced survival in the wild, compared to offspring from clutches in which all siblings appeared normal. The proportion of malformed hatchlings in the natural population was ca. 10%, in spite of differences in juvenile dispersal between males and females. Male juveniles disperse significantly further from their natal sites than do female juveniles.     

Couch potatoes do better: Delayed dispersal and territory size affect the duration of territory occupancy in a monogamous mammal

In territorial, socially monogamous species, the establishment and defense of a territory are an important strategy to maximize individual fitness, but the factors responsible for the duration of territory occupancy are rarely studied, especially in long‐lived mammals. A long‐term monitoring program in southeast Norway spanning over 18 years allowed us to follow the individual life histories of Eurasian beavers (Castor fiber) from adolescence in their natal family group to dispersal and territory establishment until the end of territory occupancy. We investigated whether territory size, resource availability, population density, and dispersal age could explain the duration of territory occupancy, which ranged from 1 to 11 years. The duration of territory occupancy was positively related to dispersal age, suggesting that individuals that delayed dispersal had a competitive advantage due to a larger body mass. This is in support with the maturation hypothesis, which states that an animal should await its physical and behavioral maturation before the acquisition of a territory. Further, we found that individuals that established in medium‐sized territories occupied them longer as compared to individuals in small or large territories. This suggests that large territories are more costly to defend due to an increased patrolling effort, and small territories might not have sufficient resources. The lifetime reproductive success ranged from zero to six kits and generally increased with an increasing duration of territory occupancy. Our findings show the importance of holding a territory and demonstrate that dispersal decisions and territory selection have important consequences for the fitness of an individual.     

Movement patterns of the White-tailed Sea Eagle (Haliaeetus albicilla): post-fledging behaviour,natal dispersal onset and the role of the natal environment

Exploratory movements and natal dispersal form essential processes during early life history stages of raptors, but identifying the factors shaping individual movement decisions is challenging. Global positioning system (GPS) telemetry thereby provides a promising technique to study movement patterns on adequate spatio-temporal scales. We analysed data of juvenile White-tailed Sea Eagles Haliaeetus albicilla (WTSE) in north-east Germany (n = 24) derived from GPS tracking to extensively analyse movements between fledging and emigration from the natal territory. Our goal was to determine the time point of fledging, characterize pre-emigration movements and the onset of natal dispersal while investigating the influence of the natal environment. WTSE fledged at an average age of 72 days and showed strong excursive behaviour during the post-fledging period regarding the number, distance and duration of excursions, yet with high individual variability. Excursive behaviour did not differ between sexes. On average, WTSE left the parental territory 93 days after fledging. The quantity of excursive behaviour delayed the timing of emigration and WTSE tended to postpone their emigration when foraging water was accessible within the boundaries of their parental territory. The overall results suggest that young WTSE assess the quality of the natal environment via pre-emigration movements and stay in their territory of origin for as long as internal and external conditions allow for it. Our study is one of the first to characterize post-fledging and natal dispersal movements of young WTSE to such an extent and applies modern techniques to understand related movements in relation to the natal environment. The results emphasize the urgent necessity for the extension of currently existing nest protection periods and guaranteeing sustainable management of potential breeding and foraging grounds for WTSE. Ultimately, the results are relevant for all large raptor species sensitive to human-related disturbance, as they support the increasing importance of regulations with spatio-temporal specifications for breeding populations of large raptors in densely human-inhabited areas with increasing alteration of land.     

Mating system and sex allocation in the gregarious parasitoid Cotesia glomerata

The gregarious parasitoid Cotesia glomerata (L.) is often presumed to possess the characteristic attributes of a species that manifests local mate competition (LMC), as it commonly produces female-biased broods. However, our field surveys of sex ratio and laboratory observations of adult behaviour showed that this species is subject to partial local mate competition caused by natal dispersal. On average, 30% of males left their natal patch before mating, with the proportion of dispersing males increasing with an increase in the patch's sex ratio (i.e. proportion of males). Over 50% of females left their natal patch before mating, and only 27.5% of females mated with males emerging from the same natal patch. Although females showed no preference between males that were and were not their siblings, broods from females that mated with siblings had a significantly higher mean brood sex ratio (0.56) than broods from females that mated with nonsiblings (0.39). Furthermore, brood sex ratios increased as inbreeding was intensified over four generations. A field population of this wasp had a mean brood sex ratio of 0.35 over 3 years, which conformed well to the evolutionarily stable strategy sex ratio (r=0.34) predicted by Taylor's partial sibmating model for haplodiploid species. These results suggest that the sex allocation strategy of C. glomerata is based on both partial local mate competition in males and inbreeding avoidance in females. In turn, this mating system plays a role in the evolution of natal dispersal behaviour in this species.Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Sibling resemblance in natal dispersal distance and direction in the Ortolan Bunting Emberiza hortulana

Natal dispersal distance and direction determine the likelihood that siblings will settle close together and hence the risk of inbreeding. Several studies have shown a sibling resemblance in dispersal distance, but few studies have analysed sibling resemblance in dispersal direction or the distance between siblings after dispersal at the landscape level. I studied the entire Norwegian population of Ortolan Buntings Emberiza hortulana, which is patchily distributed in an area covering c. 500 km². Males and females did not differ in dispersal distance (overall median 3.7 km), but directions were different. Natal dispersal distances and directions were positively related within sibling pairs, but comparisons with control individuals suggested that any effects were due to spatial effects of configuration of habitat patches in the study area. Brother–sister pairs (n = 16) were at least as similar as brother–brother pairs (n = 18). Distance between siblings after natal dispersal increased with dispersal distance, but even so, five of 35 sibling pairs settled < 1 km apart, despite dispersal distances of 8.3–9.9 km for two of these pairs. Including movements later in life, eight sibling pairs were < 1 km apart at some time (four pairs of brothers and four brother–sister pairs), and in one case a brother mated with its sister. Another case of mating between close relatives (father and daughter) involved short female natal dispersal. These data indicate that female‐biased natal dispersal and long‐distance dispersal may reduce, but do not exclude, the possibility of inbreeding.     

Sibship effects on dispersal behaviour in a pre‐industrial human population

Understanding dispersal behaviour and its determinants is critical for studies on life‐history maximizing strategies. Although many studies have investigated the causes of dispersal, few have focused on the importance of sibship, despite that sibling interactions are predicted to lead to intrafamilial differences in dispersal patterns. Using a large demographic data set from pre‐industrial Finland (n = 9000), we tested whether the sex‐specific probability of dispersal depended on the presence of same‐sex or opposite‐sex elder siblings who can both compete and cooperate in the family. Overall, following our predictions, the presence of same‐sex elder siblings increased the probability of dispersal from natal population for both sexes, whereas the number of opposite‐sex siblings had less influence. Among males, dispersal was strongly linked to access to land resources. Female dispersal was mainly associated with competition over availability of mates but likely mediated by competition over access to wealthy mates rather mate availability per se. Besides ecological constraints, sibling interactions are strongly linked with dispersal decisions and need to be better considered in the studies on the evolution of family dynamics and fitness maximizing strategies in humans and other species.     

Dispersal, migration, and offspring retention in saturated habitats

We examine the evolutionary stability of year-round residency in territorial populations, where breeding sites are a limiting resource. The model links individual life histories to the population-wide competition for territories and includes spatial variation in habitat quality as well as a potential parent-offspring conflict over territory ownership. The general form of the model makes it applicable to the evolution of dispersal, migration, partial migration, and delayed dispersal (offspring retention). We show that migration can be evolutionarily stable only if year-round residency in a given area would produce a sink population, where mortality exceeds reproduction. If this applies to a fraction of the breeding habitat only, partial migration is expected to evolve. In the context of delayed dispersal, habitat saturation has been argued to form an ecological constraint on independent breeding, which favors offspring retention and cooperative breeding. We show that habitat saturation must be considered as a dynamic outcome of birth, death, and dispersal rates in the population, rather than an externally determined constraint. Although delayed dispersal often associates with intense competition for territories, life-history traits have direct effects on stable dispersal strategies, which can often override the effect of habitat saturation. As an example, high survival of floaters selects against delayed dispersal, even though it increases the number of competitors for each breeding vacancy (the "habitat saturation factor"). High survival of territory owners, by contrast, generally favors natal philopatry. We also conclude that spatial variation in habitat quality only rarely selects for delayed dispersal. Within a population, however, offspring retention is more likely in high-quality territories.     

Sex difference in natal dispersal distances of Golden Eagles Aquila chrysaetos in Scotland

Natal dispersal distance (NDD) is critical in understanding and defining populations and their conservation. It is defined as the linear distance between the natal location and first reproductive (‘effective NDD’) or potential reproductive (‘gross NDD’) location. It is a measure of gene flow and the functional connectivity across generations between individuals breeding in the same or different geographies. NDD is difficult to record in large raptors. GPS-satellite telemetry has facilitated its recording. Previous Scottish studies showed that gross and effective NDD were apparently equivalent, and an algorithm based on telemetric data could identify the territory settlement timing and location of birds originally tagged as nestlings. We analysed natal dispersal data from 39 Golden Eagles GPS-tagged in Scotland to estimate NDD. Raw median estimates were 29.8 km for males (n = 22) and 58.6 km for females (n = 17), 38.1 km averaged across sexes. Males had significantly shorter NDD, as theoretically predicted. Our NDD estimates were shorter but not grossly dissimilar to those from the USA, where sex differences in NDD had not been confirmed. Respective sample sizes may underly the latter contrast in confirmation. We also showed that in the absence of data from sexed birds, NDD estimates can be different. Natal dispersal duration was not related to NDD, suggesting that time to prospect a territory opportunity was not associated with the selected territory's distance from the natal site. The previous status of the subsequent settled territory (occupied or vacant) was also not related to NDD. We conclude that sex differences in NDD are important in application to population demography and conservation. Although we found no support for two other potential drivers of NDD (natal dispersal duration and previous territory status), identifying additional influences on NDD is in its infancy in large raptors and deserves more study.     

Delayed dispersal and elevated monoaminergic activity in free-ranging rhesus monkeys

Male rhesus monkeys typically disperse from their groups of birth when they are between 3 and 5 years of age. Some males, however, delay dispersal from their natal groups until after they are 5 years old. The current study evaluated central monoaminergic neurotransmitter activity as a potential correlate of such “delayed” dispersal among 54 randomly selected adolescent and adult male rhesus monkeys (Macaca mulatta) captured on Cayo Santiago during an annual trapping season. Specifically, cerebrospinal fluid (CSF) concentrations of 5-hydroxyindoleacetic acid (5-HIAA, a serotonin metabolite), 3-methoxy-4-hydroxyphenylglycol (MHPG, a norepinephrine metabolite), and homovanillic acid (HVA, a dopamine metabolite) were compared in monkeys 60 months of age or more that had either dispersed (n = 33) or were still in their natal groups (n = 5). The monkeys still in their natal groups had higher CSF concentrations of both 5-HIAA and HVA (but not MHPG) than did the animals that had emigrated (Ps < 0.05). Subsequent analysis indicated that only 5-HIAA independently differentiated dispersing monkeys from delayed dispensers. Of monkeys less than 60 months of age (n = 16), only two had dispersed from their natal groups; in this age class, there were no significant differences between dispersing and natal individuals in any CSF monoaminergic metabolite (all Ps = NS). Finally, there was no difference in the CSF 5-HIAA concentrations of the five delayed dispersers and those of younger animals (P = NS), suggesting a failure to experience the frequently reported adolescent decline in serotonergic activity. In contrast, the CSF 5-HIAA concentrations of the dispersing animals were lower than those of the younger animals (P < 0.05), consistent with either an agerelated decline or an effect of dispersal per se. © 1995 Wiley-Liss, Inc.     

Mating Strategies in Solitary Aphid Parasitoids: Effect of Patch Residence Time and Ant Attendance

Mate finding and dispersal from the natal patch in parasitoid Hymenoptera are influenced by the availability of host resources and interactions with other organisms. We compared the mating behavior of three solitary aphid parasitoids, Aphidius ervi Haliday, Lysiphlebus hirticornis Mackauer and Pauesia pini (Haliday) (Hymenoptera: Braconidae: Aphidiinae) that differ in host resource exploitation and ant mutualism. In L. hirticornis, which is obligately ant-attended, the residence time on the natal patch was approximately 4 h compared with less than 2 h in the facultatively ant-attended P. pini; the sexes did not differ in residence time. Females of A. ervi, which is not attended by ants, stayed for slightly more than 2 h on the natal patch while their male siblings remained for only 1 h. In L. hirticornis, 90% of all siblings in a clutch mated on the natal patch but only 13% in A. ervi and 42% in P. pini did so. Off-patch matings (23%) were observed only in A. ervi. Males and females of L. hirticornis were 12-times more likely to mate on the natal patch when aphids and ants were present than when either of the latter species was removed; and patch residence time declined from approximately 4 h to approximately 2.5 h in the absence of either aphids or ants. We propose that, in aphidiine wasps and perhaps other quasigregarious parasitoids, mating behavior is influenced by the availability of resources on the natal patch and the presence or absence of trophobiotic ants. Partial sib mating is expected in species producing large clutches and having a long patch residence time.     

Strategic reduction of help before dispersal in a cooperative breeder

In cooperative breeders, sexually mature subordinates can either queue for chances to inherit the breeding position in their natal group, or disperse to reproduce independently. The choice of one or the other option may be flexible, as when individuals respond to attractive dispersal options, or they may reflect fixed life-history trajectories. Here, we show in a permanently marked, natural population of the cooperatively breeding cichlid fish Neolamprologus pulcher that subordinate helpers reduce investment in territory defence shortly before dispersing. Such reduction of effort is not shown by subordinates who stay and inherit the breeding position. This difference suggests that subordinates ready to leave reduce their investment in the natal territory strategically in favour of future life-history perspectives. It seems to be part of a conditional choice of the dispersal tactic, as this reduction in effort appears only shortly before dispersal, whereas philopatric and dispersing helpers do not differ in defence effort earlier in life. Hence, cooperative territory defence is state-dependent and plastic rather than a consistent part of a fixed life-history trajectory.