Parental care and the transition to independence of Spanish Imperial Eagles Aquila heliaca in Doñana National Park, southwest Spain

The behaviour of five broods of radio-tagged Spanish Imperial Eagles Aquila heliaca adalberti was studied in Doñana National Park, Spain during the post-fledging period. The distance between perching sites and nest, the mean flight duration and distance, the percentage of time spent flying and the home range all increased exponentially with age. As the young got older, the parents spent less time in their vicinity. Young were not seen hunting, but depended on their parents for food. They begged and chased their parents throughout the post-fledging period, with higher intensity at the end. Nevertheless, the adults became progressively more reluctant to feed them, as reflected in the decrease in feeding frequency and in the number of approaching flights towards the young. At the end of the post-fledging period, adults often performed aerial displays and frequently chased their offspring. The age of independence of the different young studied varied between 123 and 145 days. The correlations between individual independence and the dates when the young were last fed by their parents, and when the highest intensity of parental aggressive behaviour occurred, were higher than correlations with the variables related to the maturation of flying. Therefore, it is suggested that parental 'meanness' and aggressive behaviour may be the factors determining the date of juvenile independence and dispersal from the home territory.     

The influence of mating system on the intensity of parent-offspring conflict in primates

An evolutionary conflict of interest exists between parents and their offspring over the partitioning of parental investment (PI) among siblings. When the direct fitness benefits to offspring of increased PI, outweigh the inclusive fitness costs from lost future sibling fitness, selection should favour the evolution of offspring selfishness over altruism. In theory, this conflict is heightened when females are not strictly monogamous, as current offspring should be less altruistic towards future half-siblings than they would be towards full-siblings. Using data collected on foetal growth rate (representing prenatal PI) in primates, I test the prediction from theory that the resolution of the parent-offspring conflict will be closer to the offspring's evolutionary optima in polyandrous species than in more monandrous species. Using phylogenetic comparative analysis, and controlling for allometry, I show that offspring are able to obtain more PI when the probability of future full-siblings decreases, and that this is most pronounced in taxa where there is the opportunity for direct foetal access to the maternal bloodstream. These results support the hypothesis that the resolution of prenatal PI conflict is influenced by both a species' mating system and by its placental structure.     

Is incest common in gray wolf packs?

Wolf packs generally consist of a breeding pair and their maturingoffspring that help provision and protect pack young. Becausethe reproductive tenure in wolves is often short, reproductivelymature offspring might replace their parents, resulting in siblingor parent-offspring matings. To determine the extent of incestuouspairings, we measured relatedness based on variability in 20microsatellite loci of mated pairs, parent-offspring pairs,and siblings in two populations of gray wolves. Our 16 sampledmated pairs had values of relatedness not overlapping thoseof known parent-offspring or sibling dyads, which is consistentwith their being unrelated or distantly related. These resultssuggest that full siblings or a parent and its offspring rarelymate and that incest avoidance is an important constraint ongray wolf behavioral ecology     

How costly is the honest signaling of need?

ESS models of biological signaling have shown that costly signals can provide honest information. In the context of parent-offspring conflict over the allocation of resources by parents to their young, the theory explains costly offspring solicitation behavior as an accurate signal of offspring need to parents who cannot assess offspring condition directly. In this paper, we provide a simple but general characterization of the honest signaling of need in models of parent-offspring conflict: the offspring's signaling cost is proportional to the parent's fitness loss from satisfying the offspring's resource requirement. The factor of proportionality is given by a measure of the extent of parent-offspring conflict that depends only on coefficients of relatedness. These results hold for interbrood conflict with uniparental investment even if the relationship between offspring condition and resource requirement is not monotonic, and extend to cases of biparental care, uncertainty concerning the parent's condition, and intra-brood conflict. Copyright 1999 Academic Press.     

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.     

Adult and hatch-year blackpoll warblers exhibit radically different regional-scale movements during post-fledging dispersal

Using a broad-scale automated telemetry array, we explored post-fledging movements of blackpoll warblers breeding in Atlantic Canada. We sought to determine the full spatial scale of post-fledging dispersal, to assess support for three hypotheses for regional-scale post-fledging movement, and to determine whether learning influenced movement during this period. We demonstrated that both young and adults moved over distances more than 200 km prior to initiating migration. Adults moved southwest, crossing the Gulf of Maine (GOM), consistent with the commencement of migration hypothesis. Hatch-year birds exhibited less directional movements constrained geographically by the GOM. Their movements were most consistent with exploration hypotheses—that young birds develop a regional-scale map to aid in habitat selection, natal dispersal and subsequent migrations.     

Factors Affecting Vocalization in Tengmalm’s Owl (Aegolius funereus) Fledglings during Post-Fledging Dependence Period: Scramble Competition or Honest Signalling of Need?

Begging behaviour of nestlings has been intensively studied for several decades as a key component of parent-offspring conflict. There are essentially two main theories to account for intensity of food solicitation among offspring: that intensity of begging is related to some form of scramble competition between nest mates or that it offers honest signalling of need to parents. The vast majority of studies which have addressed begging behaviour have been based on observations of, and experiments on, nestlings and have not considered begging behaviour, during the post-fledging period. Begging vocalizations in this post-fledging phase of dependence have rarely been studied, despite the importance of vocalizations as a communication method between offspring and parents, particularly for nocturnal species. We radiotracked 39 fledglings of the Tengmalm’s owl (Aegolius funereus) in two years with different availability of prey: 2010 (n = 29 fledglings) and 2011 (n = 10 fledglings) and made 1320 nightly localizations in which we recorded presence or absence of begging calls. Within years, the most important measures related to the probability of vocalization were body condition at fledging, time of night, number of surviving siblings, age and weather conditions. Begging intensity increased with age in both years; however, in the year with low prey availability fledglings vocalized significantly more often. The main factor causing these differences between years was probably the different availability of prey, affecting breeding success, post-fledging behaviour, and thus also both short- and long-term needs of offspring. We believe that our results suggest honest signalling of their fledgling’s need.     

Parent-Offspring Conflict

When parent-offspring relations in sexually reproducing speciesare viewed from the standpoint of the offspring as well as theparent, conflict is seen to be an expected feature of such relations.In particular, parent and offspring are expected to disagreeover how long the period of parental investment should last,over the amount of parental investment that should be given,and over the altruistic and egoistic tendencies of the offspringas these tendencies affect other relatives. In addition, undercertain conditions parents and offspring are expected to disagreeover the preferred sex of the potential offspring. In general,parent-offspring conflict is expected to increase during theperiod of parental care, and offspring are expected to employpsychological weapons in order to compete with their parents.Detailed data on mother-offspring relations in mammals are consistentwith the arguments presented. Conflict in some species, includingthe human species, is expected to extend to the adult reproductiverole of the offspring: under certain conditions parents areexpected to attempt to mold an offspring, against its betterinterests, into a permanent nonreproductive.     

Begging in the absence of sibling competition in Wilson’s storm-petrels, Oceanites oceanicus

Begging displays of nestlings in multichick broods can signal both hunger and competitive ability. Studies of begging in species with single-chick broods exclude the influence of nestling competition and may provide especially useful models for the study of signalling during parent-offspring conflict. However, there is no evidence that chicks signal hunger by begging in the absence of sibling competition. I tested predictions of signalling models in a species with single-chick broods, the Wilson's storm-petrel. Chicks used two types of begging calls, ‘rhythmic’ calls and ‘long’ calls. I found that chicks conveyed information about their current body condition by begging. When their body condition was low, chicks increased the number and frequency of long begging calls, as well as the frequency of rhythmic calling. Parents responded to increased begging by regurgitating larger meals. The study thus demonstrates that the begging system can work in the absence of nestling competition. Chicks also called in the absence of their parents, but in this context they used only rhythmic calls and there was no correlation with current body condition. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

波兰东南部乌灰鹞在雏鸟出羽后期的同窝雏间冲突

通过对波兰东南部的13巢乌灰鹞(Circus pygargus) ,37只雏鸟观察,对同窝雏鸟间的竞争进行了研究。无论是在食物的数量或者生物量上,早孵出的雏鸟占有空中喂养与巢内喂养的食物源的绝对优势。空中学习捕食的食物量占喂养食物量的比例随着雏鸟年龄的增长而增加。雏鸟在空中喂养中的攻击行为通过对空中食物传递进行研究。在370次的食物传递中,由于雏间竞争而引起的落地食物的比例57次(7·8 %)。由于雏间竞争而不能获得亲鸟喂养的幼鸟有时捕食周围的雀形目小鸟。亲鸟很少喂养较小的雏鸟。窝雏数越大,雏鸟间对食物的争斗持续时间越长。空中食物传递中捕食的成功率与雏鸟的空间分布相关。这种建立在雏鸟早期的等级制一直持续到雏鸟的出羽后期,从而保证早出雏具备开始迁徙的良好身体条件[动物学报51 (5) : 790 -796 ,2005]。     

The effect of parent sex on prey deliveries to fledgling Eurasian Sparrowhawks Accipiter nisus

The relative contribution of each parent when providing for the fledglings has been recorded in only a few raptor species. We studied prey deliveries by Eurasian Sparrowhawk Accipiter nisus parents to fledglings at seven nests in southern Norway. Parents and young were fitted with radio-transmitters. Males delivered a larger number of prey to the young than did females throughout the post-fledging period (on average c.  80% of the deliveries). Two females were never observed to deliver food to the offspring, and their mates apparently raised the young to independence alone. The duration of the post-fledging period was positively related to per-capita delivery rate in the late stage.     

Games among cannibals: competition to cannibalize and parent-offspring conflict lead to increased sibling cannibalism

Sibling cannibalism occurs in many species, yet understanding of sibling cannibalism as an adaptation currently lags behind understanding of other antagonistic interactions among siblings. Observed sibling cannibalism phenotypes likely reflect the interaction between competitive games among siblings and parent-offspring conflict. Using a game-theoretic approach, we derive optimal offspring cannibalism behaviour and parental modifiers that limit or facilitate cannibalism. The results are compared to contemporary frequency-independent analysis. With the addition of game interactions among siblings or parent-offspring co-evolution, our model predicts increased cannibalism (compared to the frequency-independent prediction), as offspring compete to eat siblings. When infertile eggs are present--strengthening competition--offspring risk eating viable siblings in order to gain access to infertile eggs, intensifying parent-offspring conflict. We use the results to make new predictions about the occurrence of sibling cannibalism. Additionally, we demonstrate the utility of trophic egg laying as a maternal mechanism to promote egg eating.     

Post-fledging interactions between the Common Cuckoo Cuculus canorus and its cavity-nesting Common Redstart Phoenicurus phoenicurus host

Brood parasite–host interactions during the incubation and nestling stages have been well studied, but the post-fledging period remains virtually unknown. Using radiotracking, we provide the first detailed data on post-fledging interactions between the Common Cuckoo Cuculus canorus and its only regular cavity-nesting host, the Common Redstart Phoenicurus phoenicurus. Cuckoos raised alone (‘solitary’) fledged at higher mass, with higher wing and tarsus length and started to fly at a younger age than Cuckoos raised alongside young Redstarts (‘mixed’). However, a further 23 fledging and post-fledging parameters measured at five pre-determined times (fledging, first-flight, predation, starvation, independence) did not differ between solitary and mixed Cuckoos. In addition, none of the parameters measured during the post-fledging period (growth, dispersal distances, number of flights) differed between solitary and mixed Cuckoos. Redstart fledglings from non-parasitized broods (‘solitary’) showed generally similar fledging and post-fledging parameters to fledglings reared alongside a Cuckoo (‘mixed’). Surprisingly, there were no significant differences in post-fledging predation rate, starvation or overall survival rates between mixed and solitary Cuckoos or mixed and solitary Redstarts. Thus, during the post-fledging period, mixed Cuckoo fledglings successfully compensated for the poorer performance experienced during the nestling stage whereas mixed and solitary Redstarts did not differ in any measured parameters. This suggests that the regular occurrence of mixed broods in this host–parasite system – which is unique among the many Cuckoo hosts – is evolutionarily stable for both hosts and parasites.     

Parentally biased favouritism: why should parents specialize in caring for different offspring?

'Parentally biased favouritism' occurs when the two parents differentially care for individual offspring or kinds of offspring. Examples in birds include brood division and differential investment by the two parents in relation to the size or sex of the offspring. This paper uses mathematical models to investigate which ideas can, in theory, explain parentally biased favouritism. One previous explanation is that the parents differ in their cost of reproduction and that the parent who consequently invests least concentrates its care on the more valuable offspring. However, a mathematical model predicts the total care given by each parent and received by each offspring, not how much each parent cares for each offspring, and hence does not explain parentally biased favouritism. Parentally biased favouritism towards particular types of offspring can be explained by a difference between the parents in the benefits of caring for a given type of offspring or in the effort incurred in providing care to a given type of offspring, but then it is extreme, with at least one of the parents providing care to only one type of offspring. Parentally biased favouritism towards particular individual offspring (brood division) can be explained by parent-offspring conflict or sexual conflict.     

Parental aggression in black-capped chickadees

Black-capped chickadee (Parus atricapillus) adults are aggressivetoward their young after the young have fledged. We examinetwo hypotheses to explain the function of this aggression. Eightfamilies were observed throughout the period after fledgingto see if aggressionfunctions in discouraging the youngs' dependenceon their parents for feedings or in encouraging dispersal ofyoung from the natal territory. Aggression consisted mostlyof chases, supplantings, and attacks. Most aggressive interactionsoccurred when young were following a parent closely or begging.After an aggressive act, fledglings tended to move away fromthe parent and forage on their own. The frequency of aggressiveacts peaked at 13 days after fledging, then decreased. Thispeak occurred when fledglings were making the transition fromdependence on parental feedings to independent foraging. Mostfledglings had not dispersed by 22 days after fledging. Ourresults support the hypothesis that parental aggression encouragesfledglings to forage independently, rather than to dispersefrom the natal territory.     

Cost of parasitism and host immune defence in the sand martin Riparia riparia: a role for parent-offspring conflict?

Parent-offspring conflict may arise because the lifetime reproductive success of the parent is more influenced by its life span than by reproductive success during a particular reproductive event, while the fitness of an offspring depends firstly on its own survival as a juvenile and only subsequently on its own reproductive success. The naive immune system of young animals may allow offspring to be much more affected by parasites than their parents, and thus cause an initial asymmetry in a potential parent-offspring conflict. We investigated this type of conflict by assessing the health status and the immune response of parent and offspring sand martins Riparia riparia infested with manipulated loads of ticks Ixodes lividus (nests either treated with pyrethrin, water, or just visited). The prevalence and the intensity of tick infestations differed among treatments, with low tick loads in nests with the pyrethrin treatment. Ticks reduced the reproductive success of the host and increased offspring wing length. Broods with ticks had higher leukocyte concentrations and concentrations of immunoglobulins. The concentration of immunoglobulins in nestlings was negatively related to brood size and nestling tarsus length. Nestlings receiving the control treatments had a positive association between wing length and the concentration of immunoglobulins and a negative association between tarsus length and immunoglobulins. In contrast, adult sand martins did not respond to the parasite treatment in terms of immune response. Hence, the naive immune system of nestlings may be the crucial factor causing the parent-offspring conflict over costs of parasitism to be resolved to the advantage of parents that may sacrifice nestlings in heavily parasitized nests. Received: 30 March 1998 / Accepted: 5 December 1998     

Post-fledging body mass increase in Common Terns Sterna hirundo: influence of age, sex and year

We studied the inter-year and inter-sex variation of the post-fledging body mass development of Common Terns Sterna hirundo in 2000 and 2001 at the Banter See colony, northern Germany. Here, post-fledglings can be identified and weighed remotely and automatically by a transponder system that makes use of automated balances installed at the colony. Individuals were sexed with PCR amplification methods. After fledging, young generally continued to increase their mass. However, in 2000, the young did not significantly increase their mass during the post-fledging period. In 2001, conditions were more favourable and body mass increased continuously. Further, in 2001, male post-fledglings were significantly heavier than female post-fledglings. Once having left the colony area (on average 18–23 days after fledging in 2000, and 14–16 days after fledging in 2001), post-fledgling body mass had still not reached adult body mass. The longer a juvenile stayed at the colony, the higher was its final body mass, which if acting as a threshold level may control departure time. Neither brood size nor hatching order affected post-fledging mass or period. In the unfavourable year 2000, when many individuals were found dead after fledging, fledging age but not fledging mass was found to be a predictor of post-fledging survival before departure: individuals fledging when older had a lower survival probability. Our results stress the importance of the post-fledging period for body mass increase and survival prior to departure. The variation in post-fledging mass growth between years and between the sexes is discussed with respect to parental effort and a possible selective provisioning of sons over daughters.     

Development of chicks and predispersal behaviour of young in the Eagle Owl Bubo bubo

Little quantitative information on the development and behaviour of chicks and young is available for many species, despite the crucial importance of such data and the sensitivity of this stage in a bird's life. For Eagle Owls Bubo bubo , despite the large amount of scientific literature on this species, much basic information is lacking. This study provides a photographic and morphometric guide for age estimation of nestlings and fledglings, as well as data on the call behaviour of young, and patterns of movements during the post-fledging dependence period. The most remarkable event in chick development is the rapid increase in mass, and size gain, during the first 30 and 40–45 days, respectively. Because after this time morphometric differences become less evident, young-feather development is more useful for ageing. Patterns of chick call behaviour showed that the time spent calling increased with age and, from 110 days of age, chick vocalizations were usually uniformly distributed through the whole night and most synchronized at sunset and sunrise (the maximum recorded number of vocalizations per chick and per night was 1106 calls). During the post-fledging dependence period, radiotagged Owls moved widely, up to 1500 m from the nest after the age of 80–90 days. During such movements, the mean distance among siblings increased with age, from 168 m on average for juveniles less than 100 days old, to 489 m for those older than 100 days. Definitive dispersal started when young were about 150–160 days old. Information on chick call behaviour and movements is crucial for unbiased census and nest checking, as well as for the definition of young post-fledging areas. Knowledge of the latter is very important in terms of conservation and management (especially for those species that move largely around their nest before dispersal) owing to the high mortality that can occur during this period.     

Parent-offspring conflict and co-adaptation: behavioural ecology meets quantitative genetics

The evolution of the complex and dynamic behavioural interactions between caring parents and their dependent offspring is a major area of research in behavioural ecology and quantitative genetics. While behavioural ecologists examine the evolution of interactions between parents and offspring in the light of parent-offspring conflict and its resolution, quantitative geneticists explore the evolution of such interactions in the light of parent-offspring co-adaptation due to combined effects of parental and offspring behaviours on fitness. To date, there is little interaction or integration between these two fields. Here, we first review the merits and limitations of each of these two approaches and show that they provide important complementary insights into the evolution of strategies for offspring begging and parental resource provisioning. We then outline how central ideas from behavioural ecology and quantitative genetics can be combined within a framework based on the concept of behavioural reaction norms, which provides a common basis for behavioural ecologists and quantitative geneticists to study the evolution of parent-offspring interactions. Finally, we discuss how the behavioural reaction norm approach can be used to advance our understanding of parent-offspring conflict by combining information about the genetic basis of traits from quantitative genetics with key insights regarding the adaptive function and dynamic nature of parental and offspring behaviours from behavioural ecology.     

Begging and bleating: the evolution of parent-offspring signalling

The evolution of biological signalling in the face of evolutionary conflicts of interest is an active area of evolutionary ecology, and one to which Maynard Smith has made important contributions. We explore the major theoretical challenges in the field, concentrating largely on how offspring signal to their parents when there is the potential for parent-offspring conflict. Costly offspring solicitation (begging etc.) has been interpreted in terms of a Zahavi Grafen honest handicap signal, but this has been challenged on the grounds of' the costs of signalling. We review this controversy and also explore the issue of pooling versus separating signalling equilibrium. An alternative explanation for costly begging is that it is due to sibling competition, and we discuss the relationship between these ideas and signalling models in families with more than one offspring. Finally we consider signal uncertainty, how signalling models can be made dynamic, and briefly how they may be tested experimentally.