Pair-bond Duration Research Articles

It is good to be average: Ecological correlates of breeding phenology in an Arctic seabird, Alle alle (Dovekie)

Abstract Recognizing the deviation in an animal’s behavior from a population mean is not only necessary to understand the evolution and stability of the whole system but also to predict the future of a population in a changing environment. Arctic seabirds are expected to exhibit high synchronization in timing of breeding at the population level, due to highly seasonal and harsh environmental conditions. Nevertheless, even in such a highly synchronized system, there are always some earlier and later breeders, and the drivers of inter-pair variation remain largely unknown. Using an 8-year dataset on Alle alle (Dovekie), a small Arctic seabird, we examined potential drivers of breeding phenology. We found that A. alle pairs were quite repeatable in their phenology, and preserved their phenological status, with their chicks hatching consistently before, during, or after the population median date for hatching, despite that the calendar position of the median shifted between years. This would suggest that timing of breeding is associated with some pair characteristics, either via properties of the nest and/or some partner’s traits. However, breeding phenology of the pair was not dependent on nest location, pair bond duration or morphological similarity of the partners. Importantly, timing of breeding was negatively associated with chick growth rate, indicating fitness consequences of phenology. A simulation in our study further suggests that the chance of fledgling survival in the context of predation may be the highest for the chicks that hatched during the peak of the hatching period. While our results suggest that the timing of the breeding is important for the reproductive outcome, further research is required to determine the drivers of the consistent breeding phenology within A. alle pairs.

Benefits of Pair-Bond Duration on Reproduction in a Lifelong Monogamous Cooperative Passerine.

AbstractLong-term social and genetic monogamy is rare in animals except birds, but even in birds it is infrequent and poorly understood. We investigated possible advantages of monogamy in a colonial, facultative cooperatively breeding bird from an arid, unpredictable environment, the sociable weaver (Philetairus socius). We documented divorce and extrapair paternity of 703 pairs over 10 years and separated effects of pair duration from breeding experience by analyzing longitudinal and cross-sectional datasets. Parts of the colonies were protected from nest predation, thereby limiting its stochastic and thus confounding effect on fitness measures. We found that 6.4% of sociable weaver pairs divorced and 2.2% of young were extrapair. Longer pair-bonds were associated with more clutches and fledglings per season and with reproducing earlier and later in the season, when snake predation is lower, but not with increased egg or fledgling mass or with nestling survival. Finally, the number of helpers at the nest increased with pair-bond duration. Results were similar for protected and unprotected nests. We suggest that long-term monogamy is associated with a better capacity for exploiting a temporally unpredictable environment and helps to form larger groups. These results can contribute to our understanding of why long-term monogamy is frequently associated with unpredictable environments and cooperation.

Breeding behavior analysis in a large captive colony of African penguins (Spheniscus demersus) and its implications for population management and conservation

The African penguin Spheniscus demersus, frequently housed in zoos, holds potential for future reintroduction efforts due to its declining wild population. This paper aims to explore various aspects of reproductive performance in African penguins within a large ex situ colony at Zoo Wrocław in Poland, covering 9 years of breeding behaviors. The analysis reveals parallels in colony growth and partner change patterns with those observed in the wild. Positive correlations were found between breeding success and pair-bond duration, with the increasing colony size influencing reproductive performance. Contrary to their wild counterparts, captive African penguins initiate breeding attempt and produce a fertilized egg at a younger age. However, successful breeding still requires gaining experience or forming pairs with more experienced partners. Our research indicates that providing captive African penguins with unlimited food resources and sufficient nesting space results in rapid colony growth. The increased colony size facilitates breeding behaviors that positively influence population dynamics, particularly through the maintenance of long-term pair bond relationships and the potential for partner changes when necessary or desirable to enhance breeding success. We present compelling case studies in pair fidelity, offering valuable insights and implications for the management of captive populations and conservation efforts.

Assessing the reproductive consequences of mate retention and pair bond duration in Thorn‐tailed Rayadito (Aphrastura spinicauda), a short‐lived, socially monogamous neotropical bird

The adaptive value of mate retention has been studied in several socially monogamous birds but evidence of reproductive benefits for short‐lived species is inconclusive. Most studies come from northern latitudes but more research on tropical birds is needed, as these species typically show higher survival rates and longer pair bonds than those from temperate regions. We gathered data on the reproductive biology of a subtropical, isolated population of Thorn‐tailed Rayadito Aphrastura spinicaudaduring 2008–2017 to evaluate the reproductive consequences of mate retention. We examined data from 243 breeding attempts made by 159 breeding pairs. We found that ~30% of all breeding pairs bred together during at least two consecutive years, and some were mated for 6 years. The main cause of pair dissolution was mate loss, not divorce. Mixed‐effects models provided moderate evidence for positive effects of mate retention and successive remating on reproductive success. Newly formed pairs laid eggs later and had slightly smaller clutches than remated pairs. Furthermore, clutch size seemed to increase with successive remating. Overall, our results suggest that newly formed pairs are less efficient in reproduction and that minor yearly reproductive benefits of mate retention might accumulate for birds that are able to breed with the same partner over many years. Because breeding habitat is limited in our study population, Thorn‐tailed Rayaditos could benefit from remating if the number of individuals that can breed exceeds the number of available breeding positions. Profitable long‐term pair bonds might be more frequent in tropical birds and therefore more studies are needed to assess the prevalence of remating‐mediated effects on reproduction in relatively short‐lived monogamous species breeding in tropical regions.

Vocal recognition of distance calls in a group-living basal bird: the greylag goose, Anser anser

Vocal communication in group-living animals represents a major challenge when multiple individuals call simultaneously and generate a complex soundscape. Decoding relevant information from a vocal signal and its emitter can determine the survival of the receiver. In hierarchical social groups, information on sex, size and age can provide relevant context for vocal signals. Ultimately, individual vocal recognition is a keystone in vocal communication systems, especially in the context of sociality. Greylag geese (Aves: Anseriformes) are a basal, gregarious and highly social species, and adults produce a distance call when approaching or leaving the flock. In this study we (1) quantified the acoustic properties of distance calls to test whether they can provide information on the sex, size and age of the emitter, (2) evaluated whether distance calls have individual properties, from the sender's perspective, (3) tested whether receivers discriminate distance calls depending on the emitter's identity and (4) explored whether an individual's response strength towards its partner's distance call is associated with fitness-related traits (i.e. pair bond duration and average number of offspring per year). We recorded distance calls of individually marked geese, quantified the variation in call acoustic properties and carried out a playback experiment in the wild. Distance calls encoded information about the sex of the goose, but not its age or size. From the emitter's perspective, we found some support for individuality in distance calls. From the receiver's perspective, geese were capable of recognizing distance calls from different individuals. Finally, the response strength towards playback of the partner's call was positively correlated with pair bond duration. Vocal recognition has been proposed to be tightly related to social structure, the evolution of sociality and vocal learning. Here, we uncovered vocal recognition of individual affiliative calls in a social bird from the most basal avian group studied so far. • We studied acoustic variation in affiliative distance calls in wild greylag geese. • Geese were able to recognize distance calls from different individuals. • Fitness-related traits were correlated with vocal responses.

The macroecology of extra-pair paternity in birds.

Extra-pair paternity (EPP) is a key aspect of the mating behaviour of birds and its frequency varies widely among populations and species. Several hypotheses predict patterns of geographical variation in the occurrence and frequency of EPP, but a global-scale study on variation in this trait is still lacking. We collected data on EPP from 663 populations of 401 avian species and explored the geographical variation in the frequency of EPP among populations, species and species assemblages. We modelled the variation in the frequency of EPP within the species' breeding range accounting for the specific ecological context of each population, and used the model predictions to compute frequencies of EPP at the level of species assemblages. A global map of assemblage-level EPP rates shows clear differences between zoogeographical realms, with the highest EPP values in the Nearctic realm. Our results show that the frequency of EPP (1) decreases with latitude and increases with the distance from the breeding range boundary within the species' breeding range, (2) is negatively associated with generation length and pair-bond duration among species, and (3) decreases with latitude at assemblage level. The latitudinal decline of EPP is consistent across zoogeographical realms.

Mate fidelity improves survival and breeding propensity of a long-lived bird.

Evolutionary and behavioural ecologists have long been interested in factors shaping the variation in mating behaviour observed in nature. Although much of the research on this topic has focused on the consequences of mate choice and mate change on annual reproductive success, studies of a potential positive link between mate fidelity and adult demographic rates have been comparatively rare. This is particularly true for long-lived birds with multi-year, socially monogamous pair bonds. We used a 26-year capture-mark-recapture dataset of 3,330 black brent Branta bernicla nigricans to test whether breeding with a familiar mate improved future breeding propensity and survival. We predicted that experienced breeders nesting with a new partner would have rates of survival similar to familiar pairs because long-lived species avoid jeopardizing survival since their lifetime fitness is sensitive to this vital rate. In contrast, we expected that any costs of breeding with a new partner would be paid through skipping the subsequent breeding attempt. We found that unfamiliar pairs had lower subsequent breeding propensity than faithful partners. However, contrary to our expectations, individuals breeding with a new mate also suffered reduced survival. These results add to a small number of studies indicating that a positive relationship between mate retention and adult demographic rates may exist in a diverse array of avian species. Given these results, researchers should consider costs of mate change that extend beyond within-season reproductive success to fully understand the potential adaptive basis for perennial social monogamy. We caution that if mate retention enhances survival prospects, improvements in annual reproductive success with pair-bond length could be a secondary factor favouring perennial social monogamy, particularly in species with slower life-history strategies. Furthermore, some cases where annual reproductive success does not improve with pair-bond duration, yet multi-year pair bonds are common, could be explained by benefits afforded by mate fidelity to adult vital rates.

Pair bonds, reproductive success, and rise of alternate mating strategies in a social carnivore

Abstract Monogamy is commonly observed across a wide variety of species and taxa and arises when young are altricial, parental investment in young is high, and mate monopolization is generally not possible. In such species, pairs may bond for multiple breeding seasons while successfully rearing young. Individuals, however, may attempt to bypass the dominant mating strategy particularly when breeding opportunities are limited. Currently, we do not know how pair bond duration affects the efficacy of alternative mating strategies in populations with a monogamous mating system. Additionally, inferences about pair bond effects on reproductive success (i.e., both clutch size and recruitment) are largely limited to long-lived birds and little is known about effects on mammalian cooperative breeders. I used genetic sampling and pedigrees to examine the effects of pair bond duration on reproductive success (i.e., litter size, recruitment) and mating strategies in a population of gray wolves (Canis lupus) in Idaho, USA. There was a positive, marginally significant relationship between pair bond duration and apparent survival of offspring. Increased pair bond duration was also associated with a dampening in the prevalence of other alternative mating strategies such as sneaker males and polygamy. The selective advantage of alternative mating strategies is a combination of population, group (for applicable species), individual, and social influences such as pair bonds. The distribution of pair bonds in a monogamous population affects the selective advantage, and hence frequency, of various mating strategies observed.

The benefits of pair bond tenure in the cooperatively breeding pied babbler (Turdoides bicolor).

The benefits of stable pair bonds (that persist between breeding attempts) have been well described, but are relatively less well known in cooperatively breeding species. If pair bonds are beneficial, then it is possible that the bond between the behaviorally and socially dominant pair may influence factors such as reproductive success and group stability in cooperative species. Here, we used long‐term data to investigate the relationships between pair bond tenure, reproductive success, and group stability in the cooperatively breeding pied babbler (Turdoides bicolor). Pair bond tenure positively influenced both the number of offspring recruited annually per pair and total reproductive success (over entire pair bond duration), indicating that pair bond tenure has an important influence on reproductive success. The likelihood of immigration into the group was lower for groups containing a bonded pair with long tenure, indicating that the duration of pair bonds may impact group stability. These findings suggest that pair tenure, a hitherto relatively unexplored factor in cooperative species, may have an important influence on group dynamics.

Mate guarding and male body condition shape male fertilization success and female mating system in the common quail

In species with female polygamy, pair bonds are frequently established since mate guarding can determine male fertilization success. However, extending the duration of pair bonds also implies reducing the chances of finding new mates. Males face a trade-off between mate guarding and looking for new mates, which can be shaped by their body condition. Here, we investigated the effect of male body condition and mate guarding over the female mating system (genetic monogamy or polygamy) and the male fertilization success in the common quail, Coturnix coturnix, a species with sperm storage and thus the potential for postcopulatory selection, and without paternal care. We monitored 20 females and 32 males. We genotyped them, the 21 clutches laid by these females and a large proportion of the males present in the population, which could have sired the clutches, to perform paternity analyses. We tested whether it is the pairing order or the duration of the pair bond that determines the fertilization outcome in clutches with multiple fathers. We hypothesized that males with better body condition might be able to find a mate faster, reducing the cost of mate switching and increasing fertilization success by spending less time in a pair bond. We observed socially monogamous and polygamous females, and our genetic analyses revealed that broods could be sired by one and by multiple fathers. Female genetic polygamy was more frequent when first matings were with males in good body condition. We detected two or three different fathers in multiple paternity broods. The male that mate guarded for longest was the one that sired most of the clutch. Although males in better body condition seemed to establish shorter pair bonds, further data are needed to confirm this trend.

Better the devil you know: common terns stay with a previous partner although pair bond duration does not affect breeding output.

In a monogamous species two partners contribute to the breeding process. We study pair formation as well as the effect of pair bond length and age on breeding performance, incorporating individual heterogeneity, based on a high-quality dataset of a long-lived seabird, the common tern (Sterna hirundo). To handle missing information and model the complicated processes driving reproduction, we use a hierarchical Bayesian model of the steps that lead to the number of fledglings, including processes at the individual and the pair level. The results show that the age of both partners is important for reproductive performance, with similar patterns for both sexes and individual heterogeneity in reproductive performance, but pair bond length is not. The terns are more likely to choose a former partner independent of the previous breeding outcome with that partner, which suggests a tendency to retain the partner chosen at the beginning of the breeding career.

Social and environmental factors modulate leucocyte profiles in free-living Greylag geese (Anser anser)

BackgroundBlood parameters such as haematocrit or leucocyte counts are indicators of immune status and health, which can be affected, in a complex way, by exogenous as well as endogenous factors. Additionally, social context is known to be among the most potent stressors in group living individuals, therefore potentially influencing haematological parameters. However, with few exceptions, this potential causal relationship received only moderate scientific attention.MethodsIn a free-living and individually marked population of the highly social and long-lived Greylag goose, Anser anser, we relate variation in haematocrit (HCT), heterophils to lymphocytes ratio (H/L) and blood leucocyte counts to the following factors: intrinsic (sex, age, raising condition, i.e. goose- or hand-raised), social (pair-bond status, pair-bond duration and parental experience) and environmental (biologically relevant periods, ambient temperature) factors. Blood samples were collected repeatedly from a total of 105 focal birds during three biologically relevant seasons (winter flock, mating season, summer).ResultsWe found significant relationships between haematological parameters and social as well as environmental factors. During the mating season, unpaired individuals had higher HCT compared to paired and family individuals and this pattern reversed in fall. Similarly, H/L ratio was positively related to pair-bond status in a seasonally dependent way, with highest values during mating and successful pairs had higher H/L ratio than unsuccessful ones. Also, absolute number of leucocytes tended to vary depending on raising condition in a seasonally dependent way.DiscussionHaematology bears a great potential in ecological and behavioural studies on wild vertebrates. In sum, we found that HTC, H/L ratio and absolute number of leucocytes are modulated by social factors and conclude that they may be considered valid indicators of individual stress load.

Neotropical wrens learn new duet rules as adults.

Although song development in songbirds has been much studied as an analogue of language development in humans, the development of vocal interaction rules has been relatively neglected in both groups. Duetting avian species provide an ideal model to address the acquisition of interaction rules as duet structure involves time and pattern-specific relationships among the vocalizations from different individuals. In this study, we address the development of the most striking properties of duets: the specific answering rules that individuals use to link their own phrase types to those of their partners (duet codes) and precise temporal coordination. By performing two removal experiments in canebrake wrens (Cantorchilus zeledoni), we show that individuals use a fixed phrase repertoire to create new phrase pairings when they acquire a new partner. Furthermore, immediately after pairing, individuals perform duets with poor coordination and poor duet code adherence, but both aspects improve with time. These results indicate that individuals need a learning period to be able to perform well-coordinated duets that follow a consistent duet code. We conclude that both duet coordination and duet code adherence are honest indicators of pair-bond duration.

Let's stay together? Intrinsic and extrinsic factors involved in pair bond dissolution in a recolonizing wolf population.

Summary For socially monogamous species, breeder bond dissolution has important consequences for population dynamics, but the extent to which extrinsic or intrinsic population factors causes pair dissolution remain poorly understood, especially among carnivores.Using an extensive life‐history data set, a survival analysis and competing risks framework, we examined the fate of 153 different wolf (Canis lupus) pairs in the recolonizing Scandinavian wolf population, during 14 winters of snow tracking and DNA monitoring.Wolf pair dissolution was generally linked to a mortality event and was strongly affected by extrinsic (i.e. anthropogenic) causes. No divorce was observed, and among the pair dissolution where causes have been identified, death of one or both wolves was always involved. Median time from pair formation to pair dissolution was three consecutive winters (i.e. approximately 2 years). Pair dissolution was mostly human‐related, primarily caused by legal control actions (36·7%), verified poaching (9·2%) and traffic‐related causes (2·1%). Intrinsic factors, such as disease and age, accounted for only 7·7% of pair dissolutions. The remaining 44·3% of dissolution events were from unknown causes, but we argue that a large portion could be explained by an additional source of human‐caused mortality, cryptic poaching.Extrinsic population factors, such as variables describing the geographical location of the pair, had a stronger effect on risk of pair dissolution compared to anthropogenic landscape characteristics. Population intrinsic factors, such as the inbreeding coefficient of the male pair member, had a negative effect on pair bond duration. The mechanism behind this result remains unknown, but might be explained by lower survival of inbred males or more complex inbreeding effects mediated by behaviour.Our study provides quantitative estimates of breeder bond duration in a social carnivore and highlights the effect of extrinsic (i.e. anthropogenic) and intrinsic factors (i.e. inbreeding) involved in wolf pair bond duration. Unlike the effects of intrinsic and extrinsic factors that are commonly reported on individual survival or population growth, here we provide quantitative estimates of their potential effect on the social unit of the population, the wolf pair.

Breeding experience, but not mate retention, determines the breeding performance in a passerine bird

Individuals that retain their former partners often perform better than conspecifics that switch partners. This may be due to high-quality individuals being more faithful to their partners and more productive. Investigations of the fitness benefits of mate retention that also control for potential confounding effects of individuals are scarce. We studied the influence of mate retention and breeding experience on breeding performance of the hair-crested drongo (Dicrurus hottentottus) by carrying out both cross-sectional and longitudinal analyses. Pairs with longer pair-bond duration did not fledge more young or fledglings of better body condition nor did they produce more or better fledglings than newly formed pairs consisting of at least 1 experienced breeder, that is, individuals that had bred before. Individuals produced fewer fledglings when they were paired with an inexperienced breeder, especially when females were paired with inexperienced males. Although clutch size was not affected by mate retention or breeding experience, pairs consisting of inexperienced breeder(s) had a relatively higher predation rate of eggs and/or nestlings because they may be less effective in nest defense. The onset of breeding was advanced in the year following mate retention, but not in the second year thereafter, when pairs still remained together. Furthermore, only the breeding experience of the male determined the onset of breeding: Pairs consisting of inexperienced males bred later in the season. Our results suggest that breeding experience, and particularly the breeding experience of the male, but not mate retention, is important in determining the breeding performance of hair-crested drongos.

Do individual breeding experience and parental effort affect breeding season length in blackbirds?

Weather is a crucial factor affecting the onset and end of breeding in birds. However, a few studies have clearly indicated that other factors—breeding parameters and/or individual traits—can also exert an influence. Our study aimed to determine how breeding experience and parental investment are related to the start and the end of the breeding season. We used age and pair-bond duration as predictors of breeding experience. We presumed, firstly, that older birds with a longer pair-bond duration would start breeding earlier, and secondly, that greater parental investment (expressed as breeding success in the previous season) favored later breeding. We tested the above hypotheses on an urban population of blackbirds (Turdus merula) in 2 parks in the city of Szczecin (western Poland) over 19 years. Despite the importance of weather variables, breeding experience expressed as age and pair-bond was shown to be a significant factor, too. Older birds with more bond experience started breeding sooner. Toward the end of the breeding season older, more experienced birds prolonged this period. We also observed senescence, which was manifested as the shortening of the breeding season. Those birds with more success in the previous season prolonged their breeding period in the following season; this contradicted our hypothesis. Temperature and precipitation were negatively related to the initiation date of the first clutch but positively to the initiation date of the last one. Our survey clearly showed the importance of breeding experience with the same partner and underlined the advantages of monogamy.

Perennial Pair Bonds in an Accipiter: A Behavioral Response to an Urbanized Landscape?

Abstract In some urban environments, human activities enhance resources for avian species, providing habitat that can support year-round occupancy. If both members of a mated pair stay on their breeding territories year-round, close proximity of pair members throughout the year may increase the potential for interactions outside the breeding season. Under these circumstances, avian species that would otherwise terminate their bonds following the breeding season may form perennial pair bonds. We examined behavior of mated pairs of adult Cooper’s Hawks (Accipiter cooperii) during the nonbreeding season in an urban environment to determine whether pairs retained their breeding territories outside the breeding season and if year-round maintenance of territories influenced the duration of pair bonds. Home ranges and core areas of pair members largely overlapped. Pair members remained close to the nest site they used during the previous breeding season, avoided neighboring conspecifics of the same sex, and sele...

Mate Familiarity Affects Pairing Behaviour in a Long‐Term Monogamous Lizard: Evidence from Detailed Bio‐Logging and a 31‐Year Field Study

AbstractLong‐term monogamy is most prevalent in birds but is also found in lizards. We combined a 31‐year field study of the long‐lived, monogamous Australian sleepy lizard, Tiliqua rugosa, with continuous behavioural observations through GPS data logging, in 1 yr, to investigate the duration of pair bonds, rates of partner change and whether either the reproductive performance hypothesis or the mate familiarity hypothesis could explain this remarkable long‐term monogamy. The reproductive performance hypothesis predicts higher reproductive success in more experienced parents, whereas the mate familiarity hypothesis suggests that effects of partner familiarity select for partner retention and long‐term monogamy. Rates of partner change were below 34% over a 5‐yr period and most sleepy lizards formed long‐term pair bonds: 31 partnerships lasted for more than 15 yr, 110 for more than 10 yr, and the recorded maximum was 27 yr (ongoing). In the year when we conducted detailed observations, familiar pairs mated significantly earlier than unfamiliar pairs. Previous pairing experience (total number of years paired with previous partners) had no significant effect. Early mating often equates to higher reproductive success, and we infer that is the case in sleepy lizards. Early mating of familiar pairs was not due to better body condition. We propose two suggestions about the proximate mechanisms that may allow familiar pair partners to mate earlier than unfamiliar partners. First, they may have improved coordination of their reproductive sexual cycles to reach receptivity earlier and thereby maximise fertilisation success. Second, they may forage more efficiently, benefiting from effective information transfer and/or cooperative predator detection. Those ideas need empirical testing in the future. Regardless of the mechanism, our observations of sleepy lizard pairing behaviour support the mate familiarity hypothesis, but not the reproductive performance hypothesis, as an explanation for its long‐term monogamous mating system.

Better stay together: pair bond duration increases individual fitness independent of age-related variation.

Prolonged pair bonds have the potential to improve reproductive performance of socially monogamous animals by increasing pair familiarity and enhancing coordination and cooperation between pair members. However, this has proved very difficult to test robustly because of important confounds such as age and reproductive experience. Here, we address limitations of previous studies and provide a rigorous test of the mate familiarity effect in the socially monogamous blue-footed booby, Sula nebouxii, a long-lived marine bird with a high divorce rate. Taking advantage of a natural disassociation between age and pair bond duration in this species, and applying a novel analytical approach to a 24 year database, we found that those pairs which have been together for longer establish their clutches five weeks earlier in the season, hatch more of their eggs and produce 35% more fledglings, regardless of age and reproductive experience. Our results demonstrate that pair bond duration increases individual fitness and further suggest that synergistic effects between a male and female's behaviour are likely to be involved in generating a mate familiarity effect. These findings help to explain the age- and experience-independent benefits of remating and their role in life-history evolution.

Long-life partners or sex friends? Impact of parental pair bond on offspring personality.

Previous investigations reported that some traits of parental relationships, including pair-bond duration or mate behavioural compatibility, influence subsequent offspring fitness by acting on their behaviour and growth and thus their early survival. We hypothesized that the development of a pair bond between sexual partners would have a prenatal influence. This study investigated the impact of two pairing managements on the egg characteristics and development of offspring of Japanese quail (Coturnix c. japonica). Thirty males and 30 females were paired either continuously (C; mates together all the time) or non-continuously (NC; pairs met only three times a week for 5 min). Separation-reunion tests evaluated parental pair bond. Egg yolk testosterone and androstenedione levels were evaluated, and the somatic and behavioural development of C and NC chicks was assessed. Our results revealed that members of C pairs were attached to their mates and, although no significant differences in androgen levels could be evidenced between egg sets, a higher proportion of C pairs' eggs were fertilized and their chicks appeared less emotive and more social. Our results revealed that the parental relationship can modulate the behavioural development of their offspring, probably via non-genetic effects, and this could play a major role in the emergence of inter-individual variability.