Rhythmic birds show a better social integration than arrhythmic birds

The activity rhythms of Japanese quail vary from one individual to another. Performing a divergent selection, we obtained one line of quail expressing a robust circadian rhythmicity of feeding activity (R) and one line of quail expressing circadian arrhythmicity of feeding activity (A). We questioned whether the endogenous rhythmicity of an individual could predict its integration in a group. For that, we introduced either an R- or an A-line chick into stable groups of standard chicks. First, we evaluated proximity and synchronization of the introduced chicks on the activities of the other group members. R-chicks remained spatially and temporally closer to other group members than did A-chicks. Second, we evaluated interactions of the introduced chicks and the level of their acceptance by the other group members. R-chicks were more competent to gain access to food than were A-chicks, and separation from their group stressed R-chicks more than A-chicks. Last, successive introductions assessed age effects: before, around, and after dispersal time (～11th day of chicks' life). Most differences between R- and A-chicks were observed between their 7th and 15th day of life. In conclusion, individual endogenous rhythms predict social integration.     

Relationships among physical traits,age and social status in Thomson's and Grant's gazelles

Territorial male Thomson's and Grant's gazelles on a commercial game ranch near Nairobi, Kenya, were older, heavier and possessed longer horns than bachelor conspecifics. Among territorial Thomson's gazelles, males accompanied by females were older than solitary individuals but differed little in frame and horn dimensions. Small sample sizes precluded similar analyses for Grant's gazelles. These results suggest the importance of size, age and perhaps experience in determining social status.     

Nonlocal male mountain white-crowned sparrows have lower paternity and higher parasite loads than males singing local dialect

Bird song often varies geographically, and when this geographicvariation has distinct boundaries, the shared song types arereferred to as song dialects. We investigated the role of songdialect in male mating success in a wild breeding populationof mountain white-crowned sparrows (Zonotrichia leucophrysoriantha). In 2 of 3 years, males singing unusual songs ("nonlocal"males) had lower total fertilization success (measured by microsatellitepaternity analysis) than did males singing the local dialect ("local" males). Similarly, females produced disproportionately more young with local than with nonlocal males. However, dialectwas not a significant predictor of male mating success whencontrolling for other factors that might affect paternity.Instead, the low mating success of nonlocal males was apparentlydue to an interaction between song dialect and parasite load.Nonlocal males were more severely infected by bloodborne Haemoproteusthan were local males, although they did not differ in anyother measured aspect of quality. Immigrant birds may be immunologically disadvantaged, possibly due to a lack of previous experiencewith the local parasite fauna, resulting in low mating success.     

House sparrows (Passer domesticus) adjust their social status position to their physiological costs

For group-living animals, the maintenance of a position in the social hierarchy may be associated with physiological costs such as increased stress and energy expenditure or suppressed immune functions. In this study, we experimentally manipulated the social status of house sparrows so that each bird experienced two social environments in random sequence: being dominant and subordinate. For 14 males, we investigated how corticosterone concentrations, energy expenditure and immune functions were affected by these changes in social status position. We found that the cost of maintaining a social status position differed between individuals and were related to individual body size. Birds with small body size had increased costs in terms of increased stress responses and reduced cell-mediated immune responses while being experimentally kept as dominants, while birds with large body size had increased costs while they were subordinates. We also found that birds with increased energetic and immunological costs as dominants obtained a low status position in the large group, while birds with increased costs as subordinates obtained a high status position in the large group. In summary, we found that the costs associated with the maintenance of social status position differed between individuals and was related to the individuals' body size. Furthermore, in a large group, individuals maintained a social status position that minimized energetic and immunological costs.     

Comprehensive paternity assignment: genotype,spatial location and social status in song sparrows,Melospiza Melodia

Comprehensive, accurate paternity assignment is critical to answering numerous questions in evolutionary ecology. Yet, most studies of species with extra‐pair paternity (EPP) fail to assign sires to all offspring. Common limitations include incomplete and biased sampling of offspring and males, particularly with respect to male location and social status, potentially biasing estimated patterns of paternity. Studies that achieve comprehensive sampling and paternity assignment are therefore required. Accordingly, we genotyped virtually all males and >99% of 6‐day‐old offspring over 16 years in a song sparrow (Melospiza melodia) population and used three complementary statistical methodologies to attempt complete paternity assignment for all 2207 offspring. Assignments were highly consistent across maximum likelihood methods that used solely genotype data, and heuristic and integrated Bayesian analyses that included data describing individual locations. Sires were assigned to >99% of all genotyped offspring with ≥95% confidence, revealing an EPP rate of c. 28%. Extra‐pair sires primarily occupied territories neighbouring their extra‐pair offspring; spatial location was therefore highly informative for paternity assignment. EPP was biased towards paired territorial males, although unpaired territorial and floater males sired c. 13% of extra‐pair offspring. Failing to sample and include unpaired males as candidate sires would therefore substantially reduce assignment rates. These analyses demonstrate the integration of genetic and ecological information to achieve comprehensive paternity assignment and direct biological insight, illustrate the potential biases that common forms of incomplete sampling could have on estimated patterns of EPP, and provide an essential basis for understanding the evolutionary causes and consequences of EPP.     

Young women with major depression live on higher homeostatic sleep pressure than healthy controls

There is mounting evidence for the involvement of the sleep-wake cycle and the circadian system in the pathogenesis of major depression. However, only a few studies so far focused on sleep and circadian rhythms under controlled experimental conditions. Thus, it remains unclear whether homeostatic sleep pressure or circadian rhythms, or both, are altered in depression. Here, the authors aimed at quantifying homeostatic and circadian sleep-wake regulatory mechanisms in young women suffering from major depressive disorder and healthy controls during a multiple nap paradigm under constant routine conditions. After an 8-h baseline night, 9 depressed women, 8 healthy young women, and 8 healthy older women underwent a 40-h multiple nap protocol (10 short sleep-wake cycles) followed by an 8-h recovery night. Polysomnographic recordings were done continuously, and subjective sleepiness was assessed. In order to measure circadian output, salivary melatonin samples were collected during scheduled wakefulness, and the circadian modulation of sleep spindles was analyzed with reference to the timing of melatonin secretion. Sleep parameters as well as non-rapid eye movement (NREM) sleep electroencephalographic (EEG) spectra were determined for collapsed left, central, and right frontal, central, parietal, and occipital derivations for the night and nap-sleep episodes in the frequency range .75-25 Hz. Young depressed women showed higher frontal EEG delta activity, as a marker of homeostatic sleep pressure, compared to healthy young and older women across both night sleep episodes together with significantly higher subjective sleepiness. Higher delta sleep EEG activity in the naps during the biological day were observed in young depressed women along with reduced nighttime melatonin secretion as compared to healthy young volunteers. The circadian modulation of sleep spindles between the biological night and day was virtually absent in healthy older women and partially impaired in young depressed women. These data provide strong evidence for higher homeostatic sleep pressure in young moderately depressed women, along with some indications for impairment of the strength of the endogenous circadian output signal involved in sleep-wake regulation. This finding may have important repercussions on the treatment of the illness as such that a selective suppression of EEG slow-wave activity could promote acute mood improvement.     

Changes in social environment induce higher emotional disturbances than changes in physical environment in quail

In intensive rearing, birds are exposed to frequent changes in both their social and physical environments, and this can have an impact on animal welfare under commercial conditions. The aim of the present study was to compare the emotional responses induced by social and non-social changes and to study the influence of the familiarity on these responses.Twenty-two female quail were first reared with a ball in 15–20-individual groups for 3 weeks from hatching. Then, each experimental subject was allocated either a familiar congener (CONG) or a familiar object (BALL). At 6 weeks of age the birds were fitted with a telemetric device to collect motor and cardiac activities. After that, each subject was exposed first to a separation from its cage-mate/familiar object, and secondly to the (re-)introduction of either the cage-mate/familiar object or of an unknown conspecific/another ball. Emotional responses were assessed through behavioural reactions and heart rate variability.Before being separated, both BALL and CONG quail showed stereotyped pacing that was more pronounced in BALL quail. BALL quail were not affected by the ball withdrawal, unlike CONG quail which reacted to the separation from their cage-mate by reducing activity and exploratory behaviour (P < 0.05). After the re-introduction, BALL quail remained closer to their ball and CONG quail spent more time with “stretched necks” than before the separation (P < 0.05). Surprisingly, CONG quail showed less stereotyped pacing when their partner was removed and recovered the initial level of stereotyped behaviour when their cage-mate was re-introduced (P < 0.01). Likewise, CONG quail also recovered the initial values of heart rate after the re-introduction of their cage-mate, indicating an increase in sympathetic activity compared to the isolation period. When the unfamiliar congener or ball was introduced in their cage, BALL quail pecked the unknown ball more than the familiar ball and had more contact with it (P < 0.05) and CONG quail had more contact with the unknown congener than with the cage-mate (P < 0.05).In conclusion, the quail appeared to experience a negative affective state before being separated, and there was no clear evidence of negative emotion in quail in response to the separation from either a social partner or an inanimate object. Nevertheless, the emotional responses of the quail in reaction to the (re-)introduction was influenced by the familiarity of the congener or the ball. Finally, changes in the quail's social environment induced more behavioural and cardiac modifications than changes in its non-social environment.     

Spatial movements and social networks in juvenile male song sparrows

The time between fledging and breeding is a critical period in songbird ontogeny, but the behavior of young songbirds in the wild is relatively unstudied. The types of social relationships juveniles form with other individuals can provide insight into the process through which they learn complex behaviors crucial for survival, territory establishment, and mate attraction. We used radio telemetry to observe social associations of young male song sparrows (Melospiza melodia) from May to November. Juvenile song sparrows were frequently observed in social flocks and generally associated with more birds in the summer than in the autumn months. Most juvenile subjects formed stable social relationships with other birds and were seen with the same individual on up to 60% of the days observed. The strongest associations occurred with other juvenile males, and these individuals were often seen <1 m from the subject, even when the subject moved large distances between tracking observations. Associations also had long-term behavioral consequences as subjects were more likely to establish territories near their associates and learn shared song types. Our results indicate that male song sparrows spend a large percentage of the juvenile life stage forming social relationships and suggest that these associations may be important for the ecology of young birds and the ontogeny of their behaviors.     

Cognitive adaptations of social bonding in birds

The 'social intelligence hypothesis' was originally conceived to explain how primates may have evolved their superior intellect and large brains when compared with other animals. Although some birds such as corvids may be intellectually comparable to apes, the same relationship between sociality and brain size seen in primates has not been found for birds, possibly suggesting a role for other non-social factors. But bird sociality is different from primate sociality. Most monkeys and apes form stable groups, whereas most birds are monogamous, and only form large flocks outside of the breeding season. Some birds form lifelong pair bonds and these species tend to have the largest brains relative to body size. Some of these species are known for their intellectual abilities (e.g. corvids and parrots), while others are not (e.g. geese and albatrosses). Although socio-ecological factors may explain some of the differences in brain size and intelligence between corvids/parrots and geese/albatrosses, we predict that the type and quality of the bonded relationship is also critical. Indeed, we present empirical evidence that rook and jackdaw partnerships resemble primate and dolphin alliances. Although social interactions within a pair may seem simple on the surface, we argue that cognition may play an important role in the maintenance of long-term relationships, something we name as 'relationship intelligence'.     

Visual perception and social foraging in birds

Birds gather information about their environment mainly through vision by scanning their surroundings. Many prevalent models of social foraging assume that foraging and scanning are mutually exclusive. Although this assumption is valid for birds with narrow visual fields, these models have also been applied to species with wide fields. In fact, available models do not make precise predictions for birds with large visual fields, in which the head-up, head-down dichotomy is not accurate and, moreover, do not consider the effects of detection distance and limited attention. Studies of how different types of visual information are acquired as a function of body posture and of how information flows within flocks offer new insights into the costs and benefits of living in groups.     

Longer telomeres associated with higher survival in birds

Differences in individual quality and survival within species are a major focus in evolutionary ecology, but we know very little about the underlying physiological mechanisms that determine these differences. Telomere shortening associated with cellular senescence and ageing may be one such mechanism. To date, however, there is little evidence linking telomere length and survival. Here, we show that tree swallows (Tachycineta bicolor) with relatively short telomeres at the age of 1 year have lower survival than tree swallows of the same age with relatively long telomeres. The survival advantage in the long telomere group continues for at least three breeding seasons. It will be important to identify mechanisms that link telomere length with survival early in life.     

Automated mapping of social networks in wild birds

Growing interest in the structure and dynamics of animal social networks has stimulated major advances [1-3], but recording reliable association data for wild populations has remained challenging. While animal-borne 'proximity' tags have been available for some time [4], earlier devices were comparatively heavy, had limited detection ranges and/or necessitated recovery for data retrieval. We have developed wireless digital transceiver technology ('Encounternet') that enables automated mapping of social networks in wild birds, yielding datasets of unprecedented size, quality and spatio-temporal resolution. Miniature, animal-borne tags record the proximity and duration of bird encounters, and periodically transfer logs to a grid of fixed receiver stations, from which datasets can be downloaded remotely for real-time analysis. We used our system to chart social associations in New Caledonian crows Corvus moneduloides[5,6]. Analysis of ca. 28,000 encounter logs for 34 crows over a 7-day period reveals a substantial degree of close-range association between non-family birds, demonstrating the potential for horizontal and oblique information exchange.     

Primary controls on species richness in higher taxa

The disparity in species richness across the tree of life is one of the most striking and pervasive features of biological diversity. Some groups are exceptionally diverse, whereas many other groups are species poor. Differences in diversity among groups are frequently assumed to result from primary control by differential rates of net diversification. However, a major alternative explanation is that ecological and other factors exert primary control on clade diversity, such that apparent variation in net diversification rates is a secondary consequence of ecological limits on clade growth. Here, I consider a likelihood framework for distinguishing between these competing hypotheses. I incorporate hierarchical modeling to explicitly relax assumptions about the constancy of diversification rates across clades, and I propose several statistics for a posteriori evaluation of model adequacy. I apply the framework to a recent dated phylogeny of ants. My results reject the hypothesis that net diversification rates exert primary control on species richness in this group and demonstrate that clade diversity is better explained by total time-integrated speciation. These results further suggest that it may not possible to estimate meaningful speciation and extinction rates from higher-level phylogenies of extant taxa only.     

Wintering forest birds roost in areas of higher sun radiation

We analyze environmental determinants of roost site selection by tree gleaning passerines wintering in a Mediterranean montane oakwood at a craggy area of high variation in altitude and hill-shading pattern. We hypothesize that in temperate latitudes of cold winter climate, birds should spend the night in areas of low altitudes, higher temperatures, and higher solar radiation in order to minimize thermoregulation costs during resting time and to improve foraging conditions just before and after roosting. We study night occupation of woodland locations by the presence of feces in 159 wooden nest boxes (i.e., under identical controlled roosting situations). We employ GIS methods to quantify solar radiation at each location surrounding the nest boxes and data loggers to measure air temperature in the field. Birds prefer to roost in forest patches with higher solar radiation, where the period of light available for foraging is extended and thermoregulation costs during daytime are minimized. They also selected woodland patches with taller trees, a pattern consistent with their foraging preferences for trunks and branches. Other environmental variables played a negligible role in determining the selection of roost sites. Here, we show, for the first time, the importance of sun radiation determining where to spend the night in wintering birds and call attention on considering the thermal space in forest management. Forest management should preserve woodland patches with taller trees more exposed to solar radiation to enhance winter habitat suitability for birds in these Mediterranean oakwoods.     

Lean birds in the city: body size and condition of house sparrows along the urbanization gradient

1. Urbanized habitats differ from natural ones in several ecological features, including climate, food availability, strength of predation and competition. Although the effects of urbanization on avian community composition are well known, there is much less information about how individual birds are affected by these human-generated habitat differences. 2. In this study we investigated the relationships between the morphological characteristics and the degree of habitat urbanization in house sparrows, Passer domesticus (Linne 1758) . We collected data for more than 1000 non-breeding adult birds in Hungary between 1997 and 2006, from seven sites including farmlands, suburban areas and city centres. 3. We found that the body mass, tarsus length and body condition of free-living sparrows differed among the sites: birds in more urbanized habitats were consistently smaller and in worse condition than birds in more rural habitats. A composite measure of habitat urbanization (based on building density, road density and vegetation cover) explained over 75% of variance between sites in the studied traits, after we controlled for the effects of sex, year, season and time of capture. 4. The difference in body mass between rural and urban sparrows was significant when birds were kept in aviaries under identical conditions, with constant ad libitum food availability. It is therefore unlikely that the reduced body size and condition of urban sparrows are a consequence of reduced access to food for adults (e.g. due to strong competition), or their short-term responses to high food predictability (e.g. by strategic mass regulation). 5. We suggest that habitat differences in nestling development or adaptive divergence of sparrow populations due to distinct environmental conditions (such as differing predation pressure) may account for the differences along the urbanization gradient.     

Perceptual consequences of early social experience in precocial birds

This article reviews recent work with precocial avian hatchlings demonstrating the important contribution that social experience with conspecifics can make to the development of species-typical perceptual preferences. In particular, experiments on the role of sibling social interaction in the development of early auditory and visually directed maternal preferences are surveyed. Results reveal that young hatchlings denied the opportunity for direct social experience with siblings consistently display auditory and visual preferences different from those shown by hatchlings allowed ongoing experience with their broodmates during the period immediately following hatching. Taken together, the studies reviewed here demonstrate that the perceptual preferences underlying the process of filial imprinting, long thought to be simply ‘innate’ or ‘instinctive’, are sensitive to an array of social factors present in the young bird's posthatching environment. The findings also provide support for the view that the minimum unit for the developmental analysis of species-typical behavior must be the developmental system, comprised of both the developing organism and its specific stimulative environment.     

Sex of opponent influences response to a potential status signal in house sparrows

We evaluated the possible use of a sexually dimorphic plumage trait as a status signal in inter- and intrasexual social interactions in free-living house sparrows, Passer domesticus. Males with larger black throat patches (bibs) tended to be dominant regardless of the opponent's sex. This relationship was maintained after controlling for body size and age in multivariate analyses. However, the nature of social interactions varied according to the sex of the opponent and a male's bib size. Males with larger bibs tended to be less aggressive to male opponents, and received significantly less aggression from those birds, a key predicted benefit of a status signal. However, when the opponent was a female, males with large bibs displayed significantly more aggression than males with small bibs, and this relationship persisted even after controlling for the dominance status of the participants. Females were significantly more aggressive towards male opponents than towards female opponents, and females interacting with two males of known bib size were more aggressive to the one with the larger bib. These results suggest that the bib of male house sparrows may be a signal of status, but that in interactions with females, the bib may also have other functions. Frequent testing by females of the underlying quality being signalled by the bib may be linked to later mate choice, and might be adaptive if the information value of the bib varies among years. Such testing may also contribute to mechanisms for maintaining the bib as a reliable signal of status. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Signals of status in wintering white-crowned sparrows, Zonotrichia leucophrys gambelii

The possibility of plumage status signalling within the social systems of wintering birds has been a controversial issue. Our results are the first to demonstrate conclusively the reality of such signalling. Data from eight groups of captive white-crowned sparrows (Zonotrichia leucophrys gambelii), each with 8 to 11 different individuals, show that immature and adult females with crowns painted to resemble more brightly coloured, dominant adult males consistently win encounters with control birds of their own age and sex. These experiments demonstrate that signals that correlate with age (adult versus immature) and sex (adult male versus adult female) are used by the birds as reliable indicators of relative dominance position. Our demonstration of status signalling draws attention to the need to explain how such a system can be evolutionarily stable and we discuss some suitable models.     

Offspring fitness varies with parental extra-pair status in song sparrows, Melospiza melodia

Numerous studies have tested for indirect selection on female extra-pair reproduction (EPR) by quantifying whether extra-pair young (EPY) are fitter than their within-pair young (WPY) maternal half-siblings. In contrast, the hypothesis that offspring of EPY and WPY (rather than the EPY and WPY themselves) differ in fitness has not been tested, even though inter-generational effects of parental extra-pair status on offspring fitness could alter the magnitude and direction of indirect selection on EPR. We tested whether offspring of EPY song sparrows, Melospiza melodia, were more likely to recruit or produce hatched or recruited offspring over their lifetimes than offspring of WPY. Hatchlings with one or two EPY parents were more likely to recruit and produce hatched offspring than hatchlings with two WPY parents. Furthermore, these relationships differed between maternal versus paternal extra-pair status. Hatchlings with EPY fathers were more likely to recruit and produce offspring than hatchlings with WPY fathers. In contrast, hatchlings with EPY mothers were as likely to recruit as hatchlings with WPY mothers and tended to be less likely to produce recruited offspring. Depending on the causal genetic and environmental mechanisms, such conflicting inter-generational relationships between parental extra-pair status and offspring fitness could substantially influence the evolutionary dynamics of EPR.     

Female house sparrows "count on" male genes: experimental evidence for MHC-dependent mate preference in birds

Background  

Females can potentially assess the quality of potential mates using their secondary sexual traits, and obtain "good genes" that increase offspring fitness. Another potential indirect benefit from mating preferences is genetic compatibility, which does not require extravagant or viability indicator traits. Several studies with mammals and fish indicate that the genes of the major histocompatibility complex (MHC) influence olfactory cues and mating preferences, and such preferences confer genetic benefits to offspring. We investigated whether individual MHC diversity (class I) influences mating preferences in house sparrows (Passer domesticus).