Survival enhancement through food sharing: a means for parental control of natal dispersal.

The value of food sharing among relatives is analyzed for a situation where fitness equals survival. In seasonal environments the minimum food abundance may set a limit to group living. Delayed dispersal is predicted to be linked to relaxed winter competition and high parental survival. Enhanced survival for the offspring when parents share food could be a sufficient reason to delay dispersal, while early dispersal in advance of food shortage periods may be induced by a competitive relationship. At low resource abundance dominant parents do best by being competitive and retaining all resources. For food abundance higher than the expected requirements food sharing with independent offspring is possible, although it has a non-zero cost. Food sharing parents still retain most of resources to themselves, but the resource share given to subordinate offspring gradually gets larger when food abundance increases. Except for at very low food abundance, where subordinates may adopt a "suicidal" behaviour and cede their resources to the dominant, there is a conflict over how to share the resources.     

Winter resource wealth drives delayed dispersal and family-group living in western bluebirds

Delayed dispersal, where offspring remain with parents beyond the usual period of dependence, is the typical route leading to formation of kin-based cooperative societies. The prevailing explanations for why offspring stay home are variation in resource wealth, in which offspring of wealthy parents benefit disproportionately by staying home, and nepotism, where the tendency for parents to be less aggressive and share food with offspring makes home a superior place to wait to breed. These hypotheses are not strict alternatives, as only wealthy parents have sufficient resources to share. In western bluebirds, Sialia mexicana, sons usually delay dispersal until after winter, gaining feeding advantages through maternal nepotism in a familial winter group. Experimentally reducing resource wealth (mistletoe) by half on winter territories caused sons to disperse in summer, even though their parents remained on the territory during the winter. Only 8% of sons remained with their parents on mistletoe-removal territories compared to 50% of sons on control territories (t(9,10)=3.33, p<0.005). This study is the first to demonstrate that experimentally reducing wealth of a natural food resource reduces delayed dispersal, facilitating nepotism and family-group living. The results clarify the roles of year-round residency, resource limitation and relative wealth outside the breeding season in facilitating the formation of kin-based cooperative societies.     

Maternal Watchfulness in Black Howler Monkeys (Alouatta pigra)

In many vertebrates, parents protect their young by detecting predators and aggressive conspecifics before they attack. But parental investment in protection is a limited resource, whose allocation to offspring should reflect optimization strategies. Thus, we tested if maternal investment in vigilance varied with the risk faced by young black howler monkeys (Alouatta pigra). Twelve females in six groups were compared, including those with and without young and those with immatures of differing ages. The greatest increase in vigilance was seen when immatures were conspicuous, and to a lesser extent, among mothers of dependent young (neonates and infants). These findings support the idea that parents adjust their behavior to both chronic and episodic risk faced by young. We explore the possible role of predation and infanticide in shaping the vigilance of mothers.     

Low heritability in morphological characters of Tengmalm's owls: The role of cyclic food and laying date?

Summary In parent—offspring regressions, high heritability estimates of characters may simply be due to common environment: the resemblance between the living conditions of parents and their offspring in species showing restricted natal dispersal. In vole-eating Tengmalm's owls (Aegolius funereus), the natal dispersal and breeding dispersal of adult females are wide (up to > 1000 km and > 500 km, respectively), whereas adult males are resident. We found that body measurements of 183 recruits born in western Finland were independent of parental age and vole abundance in the birth year. Early-laid eggs produced longer winged recruits than late-laid eggs. The wing lengths of the daughters showed a significant positive regression on the wing length of their mothers, but the removal of the maternal effect via laying date lowered this relationship. The development time of offspring to the first autumn might also be crucial for morphological characters of females in their later life. Significant regressions were not found between the wing length of the son and his father. The mother—daughter regression was small for laying date, but positive (p = 0.08) for clutch size. These results suggest low heritability in breeding and morphological characters of owls and this low heritability may enable plastic adjustment to optimize fitness at any stage in a fluctuating environment. Nonadditive genetic variance also creates variation between offspring characters that are not genetically correlated with the parents and may explain these low heritability estimates.     

Delayed dispersal in the Zaisan mole vole (Ellobius tancrei): helping or extended parental investment?

The highly social subterranean voles of genus Ellobius (mole vole) represent a unique model to compare with both social bathyergids and surface-dwelling voles. Unlike most arvicolines, mole voles display the prolonged obligatory delay of dispersal. In subterranean rodents, this delay may be associated with benefits of cooperation (weaned offspring help their parents), an extended parental investment (parents care for weaned offspring), or both. To identify the role offspring aged <3 months play in mole vole families, we estimated their contribution to important group activities. We found that juveniles contributed very little to daily living activities up to the age of approximately 2 months. The older offspring carried objects at least as frequently as breeders, but they engaged in gnawing obstacles less frequently than their fathers. Male breeders and non-breeders engaged in gnawing more than the respective categories of females. Although young mole voles clearly did not surpass their parents in performing the more energetically costly activities, they gained body mass by 26 g (130 %) between days 30 and 90, often surpassing their parents in weight. Based on our results, in E. tancrei an extremely prolonged infancy is followed by intensive building of body reserves. This ontogenetic trajectory appears to be distinct from the patterns described for social bathyergids and for most voles.     

Delayed dispersal and prolonged brood care in a family‐living beetle

Delayed juvenile dispersal is an important prerequisite for the evolution of family‐based social systems, such as cooperative breeding and eusociality. In general, young adults forego dispersal if there are substantial benefits to remaining in the natal nest and/or the likelihood of dispersing and breeding successfully is low. We investigate some general factors thought to drive delayed juvenile dispersal in the horned passalus beetle, a family‐living beetle in which young adults remain with their families in their natal nest for several months before dispersing. Fine‐scale population genetic structure indicated high gene flow between nest sites, suggesting that constraints on mobility are unlikely to explain philopatry. Young adults do not breed in their natal log and likely disperse before reaching breeding age, suggesting that they do not gain direct reproductive benefits from delayed dispersal. We also examined several ways in which parents might incentivize delayed dispersal by providing prolonged care to adult offspring. Although adult beetles inhibit fungal growth in the colony by manipulating both the nest site and deceased conspecifics, this is unlikely to be a major explanation for family living as both parents and adult offspring seem capable of controlling fungal growth. Adult offspring that stayed with their family groups also neither gained more mass nor experienced faster exoskeleton development than those experimentally removed from their families. The results of these experiments suggest that our current understanding of the factors underlying prolonged family living may be insufficient to explain delayed dispersal in at least some taxa, particularly insects.     

Patterns of pollen flow in a dense population of the insect-pollinated canopy tree species Castanopsis sieboldii

Insect pollinations of tree species with high-density populations have rarely been studied. Since the density of adults can affect effective pollen dispersal, short-distance pollination, even by insects, may frequently occur in high-density populations. To test this prediction, we investigated pollination patterns in a high-density population of the insect-pollinated canopy tree species Castanopsis sieboldii by paternity analysis using genotypes at 8 microsatellite loci of 145 adult trees and 439 seeds from 11 seed parents in a 4-ha plot. We then explored their genetic effects on the population by calculating other population genetics parameters. Although C. sieboldii has high potential for long-distance dispersal of pollen (as indicated by a fat-tailed dispersal kernel), the cumulative pollination at the local scale was spatially limited and strongly dependent on the distance between parents due to the high density of adults. Genetic diversity estimates for pollen pools accepted by each seed parent converged on a maximum as the effective number of pollen parents increased. The genetic diversity of pollen pool bulked over all the seed parents from inside the plot did not differ from that of the total pollen pools. Therefore, although pollen flow from distant pollen parents may help to maintain the genetic diversity of offspring, pollen parents neighboring seed parents may be the main contributors to the genetic diversity of the offspring at the seed stage.     

Natal dispersal and personalities in great tits (Parus major)

Dispersal is a major determinant of the dynamics and genetic structure of populations, and its consequences depend not only on average dispersal rates and distances, but also on the characteristics of dispersing and philopatric individuals. We investigated whether natal dispersal correlated with a predisposed behavioural trait: exploratory behaviour in novel environments. Wild great tits were caught in their natural habitat, tested the following morning in the laboratory using an open field test and released at the capture site. Natal dispersal correlated positively with parental and individual exploratory behaviour, using three independent datasets. First, fast-exploring parents had offspring that dispersed furthest. Second, immigrants were faster explorers than locally born birds. Third, post-fledging movements, comprising a major proportion of the variation in natal dispersal distances, were greater for fast females than for slow females. These findings suggest that parental behaviour influenced offspring natal dispersal either via parental behaviour per se (e.g. via post-fledging care) or by affecting the phenotype of their offspring (e.g. via their genes). Because this personality trait has a genetic basis, our results imply that genotypes differ in their dispersal distances. Therefore, the described patterns have profound consequences for the genetic composition of populations.     

Life history and the evolution of family living in birds

The reason why some bird species live in family groups is an important question of evolutionary biology that remains unanswered. Families arise when young delay the onset of independent reproduction and remain with their parents beyond independence. Explanations for why individuals forgo independent reproduction have hitherto focused on dispersal constraints, such as the absence of high-quality breeding openings. However, while constraints successfully explain within-population dispersal decisions, they fail as an ultimate explanation for variation in family formation across species. Most family-living species are long-lived and recent life-history studies demonstrated that a delayed onset of reproduction can be adaptive in long-lived species. Hence, delayed dispersal and reproduction might be an adaptive life-history decision rather than 'the best of a bad job'. Here, we attempt to provide a predictive framework for the evolution of families by integrating life-history theory into family formation theory. We suggest that longevity favours a delayed onset of reproduction and gives parents the opportunity of a prolonged investment in offspring, an option which is not available for short-lived species. Yet, parents should only prolong their investment in offspring if this increases offspring survival and outweighs the fitness cost that parents incur, which is only possible under ecological conditions, such as a predictable access to resources. We therefore propose that both life-history and ecological factors play a role in determining the evolution of family living across species, yet we suggest different mechanisms than those proposed by previous models.     

NM+: software implementing parentage-based models for estimating gene dispersal and mating patterns in plants

NM+ is computer software designed for making inferences on plant gene dispersal and mating patterns by modelling parentage probabilities of offspring based on the spatially explicit neighbourhood model. NM+ requires a sample of mapped and genotyped candidate parents and offspring; however, offspring may optionally be assigned to single maternal parents (forming so-called half-sib progeny arrays). Using maximum likelihood, NM+ estimates a number of parameters, including proportions of offspring due to self-fertilization, pollen immigration from outside of a defined study site, parameters of pollen (and/or seed) dispersal kernels (exponential-power, Weibull, geometric or 2Dt) and selection gradients relating covariates (phenotypic traits) with male (and/or female) reproductive success. NM+ allows for missing data both in parents and in offspring. It accounts for null alleles and their frequencies can optionally be considered as estimable parameters. Data files are formatted in a table-like structure so they can be easily prepared in a spreadsheet software. By default NM+ is for studying plant populations, however, it can be used for any organism as long as data requirements and model assumptions are met. NM+ runs under Windows, but it can be launched under Linux using WINE emulator. NM+ can be downloaded free of charge from http://www.genetyka.ukw.edu.pl/index_pliki/software.htm.     

Parental nepotism enhances survival of retained offspring in the Siberian jay

The evolutionary payoff accruing to parents from breeding offspringcould be an incentive for prolonged investments in the offspring.Enhanced survival for offspring as a result of such a prolongedparental investment would increase the value of remaining inthe natal territory for the offspring. Here we show that first-yearsurvival in Siberian jays is higher in the company of theirparents. Two observations point to that the enhanced survivalof retained offspring is due to nepotistic parents rather thanto the quality of a shared habitat. First, winter survivalis higher only for those retained offspring whose parents havesurvived too ; this precludes the possibility that the linkbetween timing of dispersal and survival should reflect a higher phenotypic quality of retained offspring in general. Second,there is no support for the more parsimonious explanation thatthis link between the survival of parents and retained offspringreflects habitat quality of a shared territory. We could, withhigh statistical power, reject the possibility of a correlationbetween the survival of parental birds and unrelated immigrantsto the territory. Such a correlation would have been expectedif survival reflected habitat quality and not kinship. Our data instead suggest a direct fitness gain to retained offspringin enhanced survival through parental nepotism (parental facilitation).The behavior of parents in allowing retained offspring accessto food that is denied to immigrants is one proximate mechanismmediating a benefit of delayed dispersal.     

The vigilance components of begging and sibling competition

The conflict between siblings over how parental resources are divided has promoted the evolution of specific behaviour to outcompete each other. Young animals look out for parents’ arrival in order to start begging as quickly as possible, since a rapid begging reaction increases the likelihood of being fed before nestmates. If the young can physically intercept the parents, selection might be operating on the offspring ability to monitor parent arrival (vigilance towards parents) and any sudden modifications in siblings’ behaviour (vigilance towards siblings). To investigate the adaptive value of nestling vigilance in the context of family interactions, we recorded which direction barn owl Tyto alba siblings were facing in 89 two‐chick broods before the first parental feeding visit of the night. Nestlings were more vigilant towards nest entrance than expected by chance suggesting that vigilance towards parents is an important component of sibling competition. When positioned near the nest‐box entrance where parents predictably deliver food, the younger individual (i.e. junior) looked more towards the entrance than its older sibling. Thus, when the likelihood of obtaining a food item is relatively high, juniors are more vigilant than seniors to detect the incoming parent. When positioned at the back of the nest, the senior looked relatively more frequently towards its sibling than the junior did in the same situation. This suggests that when the likelihood of obtaining a food item is relatively low, seniors are more vigilant than juniors to observe their sibling. Because vigilance was not related to hunger level and prey obtaining, we propose the hypothesis that vigilance towards parents and siblings only indirectly influences the outcome of sibling competition.     

Family movements before independence influence natal dispersal in a territorial songbird

Dispersal behaviour in territorial species is typically assumed to be independent of parental behaviour except for the possible role of parental eviction from the natal territory. Great tits defend exclusive territories at the onset of breeding but after fledging undertake substantial excursions with dependent offspring, sometimes covering distances equivalent to ten or more breeding territories and even moving across open spaces into neighbouring woodlots. We show that postfledging family movements are significantly associated with subsequent dispersal directions of recruits by comparing observed angles of movement with a simulated distribution taking into account the patchy nature of the landscape. However, the extent of family movements did not predict dispersal distances. Our findings provide an explanation for previously observed similarities in dispersal direction between siblings in the same study population, as well as for effects of nest site location on dispersal. More generally we show the existence of a novel mechanism for parents to influence dispersal and fine-scale kinship structure among their offspring.     

Dispersal distance is influenced by parental and grand-parental density

Non-genetic transmission of information across generations, so-called parental effects, can have significant impacts on offspring morphology, physiology, behaviour and life-history traits. In previous experimental work using the two-spotted spider mite Tetranychus urticae Koch, we demonstrated that dispersal distances increase with local density and levels of genetic relatedness. We here show that manipulation of parental and grand-parental density has a significant effect on offspring dispersal distance, of the same order of magnitude as manipulation of offspring density. We demonstrate that offspring exposed to the same density disperse further if they were born to parents exposed to higher density compared with parents exposed to low density. Offspring dispersal distance also increases when grand-parents were exposed to higher density, except for offspring exposed to low densities, which disperse at shorter distances whatever the grand-parental density. We also show that offspring from mothers exposed to higher densities were overall larger, which suggests that parents in high densities invest more in individual offspring, enabling them to disperse further. We propose that our findings should be included in models investigating the spread rate of invasive species or when predicting the success of conservation measures of species attempting to track changing climates.     

Reduced mortality selects for family cohesion in a social species

Delayed dispersal is the key to family formation in most kin-societies. Previous explanations for the evolution of families have focused on dispersal constraints. Recently, an alternative explanation was proposed, emphasizing the benefits gained through philopatry. Empirical data have confirmed that parents provide their philopatric offspring with preferential treatment through enhanced access to food and predator protection. Yet it remains unclear to what extent such benefits translate into fitness benefits such as reduced mortality, which ultimately can select for the evolution of families. Here, we demonstrate that philopatric Siberian jay (Perisoreus infaustus) offspring have an odds ratio of being killed by predators 62% lower than offspring that dispersed promptly after independence to join groups of unrelated individuals (20.6% versus 33.3% winter mortality). Predation was the sole cause of mortality, killing 20 out of 73 juveniles fitted with radio tags. The higher survival rate among philopatric offspring was associated with parents providing nepotistic predator protection that was withheld from unrelated group members. Natal philopatry usually involves the suppression of personal reproduction. However, a lower mortality of philopatric offspring can overcome this cost and may thus select for the formation of families and set the scene for cooperative kin-societies.     

Natal dispersal and parental escorting predict relatedness between mates in a passerine bird

Although relatedness between mates is of considerable evolutionary and ecological significance, the way in which the level of relatedness is determined by different behavioural processes remains largely unknown. We investigated the role of behaviour in predicting mate relatedness in great tits using genotypic markers and detailed observations. We studied how mate relatedness is influenced by natal dispersal, inbreeding/outbreeding avoidance after natal dispersal and a behaviour not previously considered that influences membership to social aggregations, namely family escorting behaviour by parents. Among locally born individuals, the level of mate relatedness decreased with natal dispersal distance for females, but not for males. In contrast, mate relatedness was negatively related to the extent of family movements for males, but not for females. However, family movements did not predict dispersal distance for either sex. Local recruits were more related to their mates than immigrants, but this was only significant for females. No evidence was found for inbreeding/outbreeding avoidance after dispersal. Our results suggest that, in highly mobile species, mating options are spatially and/or socially limited, and that parents influence mating options of their offspring before dispersal.     

Effects of parental and own early developmental conditions on the phenotype in zebra finches (Taeniopygia guttata)

The performance of an individual can be critically influenced by its experience early in life as well as trans-generationally by the conditions experienced by its parents. However, it remains unclear whether or not the early experience of parents and offspring interact with each other and adapt offspring when the parental and own early environmental conditions match. Here, zebra finches (Taeniopygia guttata) that had experienced either early low or high nutritional conditions raised their offspring under either matched or mismatched nutritional conditions. Parental and offspring early conditions both separately affected the offspring’s adult phenotype, but early conditions experienced by parents and offspring did not interact as predicted. Offspring that grew up under conditions matching those their parents had experienced did not do better than those that grew up in a mismatched environment. Thus, transgenerational effects remain a lifelong burden to the offspring acting in addition to the offspring’s own early life experience. The lack of evidence for adaptive programming to matching environmental conditions may result from non-predictive environments under natural conditions in such opportunistic breeders.     

Sex-biased preferential care in the cooperatively breeding Arabian babbler

In many social birds there are sex differences in dispersal patterns, with males commonly remaining in their natal group whereas females typically disperse at adolescence. Group members may therefore increase their fitness by preferentially caring for offspring of a particular sex according to social circumstances. Although previous studies have focussed on intragroup social factors that may affect preferential care, we propose that the relative size of neighbouring groups is of primary importance. Here we show that in the cooperatively breeding Arabian babbler (Turdoides squamiceps), parents preferentially feed male offspring when relative group size is small, and female offspring when group size is large. Unlike parents, helpers consistently favour young of the opposite sex to themselves, suggesting the risk of competition with members of the same sex for future breeding opportunities may override other considerations. These results emphasize the complexity of investment strategies in relation to social circumstances and the variable benefits of raising males vs. females in a species with sex-biased philopatry.     

Experimentally increased food resources in the natal territory promote offspring philopatry and helping in cooperatively breeding carrion crows

Kin-based societies, where families represent the basic social unit, occur in a relatively small number of vertebrate species. In the majority of avian kin societies, families form when offspring prolong their association with the parents on the natal territory. Therefore, the key to understanding the evolution of families in birds is to understand natal philopatry (i.e. the tendency to remain on the natal territory). It has been shown that, within populations, the strength of the association between parents and offspring (i.e. family stability) increases when offspring dispersal is constrained by external environmental factors, but it is unclear whether and how family wealth influences juvenile dispersal decisions. Here, we show that young carrion crows (Corvus corone corone) from territories that were food-supplemented year-round were more philopatric and more likely to help at their family's nest than the unfed ones. The results suggest that offspring philopatry and helping behaviour are influenced by the quality of 'home' and that the availability of food resources positively affects the cohesion of the family.     

Effect of parental flowering and dispersal times on offspring fate in Daucus carota (Apiaceae)

Seeds collected from parents that flowered at different times were dispersed onto experimental plots at different times during the normal dispersal season. Parental flowering and dispersal times, which are correlated with each other, independently affected offspring germination, growth, and time of reproduction. Estimated population growth rates were highest for offspring that were dispersed early in the dispersal season and that came from early flowering parents. The data provide evidence that 1) an individual's fate is determined by the environment of the previous generation, and that 2) an individual's fitness should be calculated from life history data that span more than one generation.