Gregarious foraging in barn swallows after the breeding season

Aerial vertebrate foragers, e.g. insectivorous bats, martins and swallows, often show gregarious behavior such as colonial breeding, communal roosting and aggregating behavior during foraging. Studies of gregariousness in aerial foragers have mostly focused on colonial breeding or communal roosting, and only a few intensive studies have dealt with gregariousness during foraging. Here, we report on large and stable aggregations of barn swallows, Hirundo rustica, observed during daytime after the breeding season in Japan. Relatively large aggregations of 20 or more birds were located around poultry, pig and cattle farms and mounds of manure. The aggregations were stable in size both within-days and between-days. Their activities consisted only of repeated cycles of foraging and resting around the farms where their prey, the large black soldier fly Hermetia illuceus (>10 mm) and other medium (5–10 mm) flies, was dense. Distributions of swallows around the farms overlapped with the distributions of prey, and the size of the aggregations significantly correlated with the abundance of prey.     

Sexually selected infanticide by adult male barn swallows

Since 1980, we have documented three definite and five suspected instances of sexually selected infanticide by unmated males in the monogamous barn swallow, Hirundo rustica. This is the first report of this type of infanticide by males in any bird species, or by either sex in any monogamous species. In our population, infanticide appears to be an adaptation serving two functions. It speeds the reproductive cycle of victimized females so that males can breed with them sooner than if they had been permitted to raise their young. In addition, it may enable unmated males to acquire females by causing them to desert their mates.     

Uninformative exaggeration of male sexual ornaments in barn swallows

Models of sexual selection suggest that mate-choice preferences are favored because differences between males in their degree of ornamental exaggeration convey useful information about the direct or indirect benefits they have to offer [1-5]. Such arguments assume that variation in male ornament size can be attributed to variation in the degree of sexually selected exaggeration. We provide the first test of this assumption by conducting tail-length experiments in male barn swallows. Over the last twenty years, a large amount of work has shown that female barn swallows are influenced by male tail length when choosing a mate [6-12]. Recent experiments have shown that a combination of natural and sexual selection results in the elongated tail streamer--a tail that is on average across the population about 12 mm (approximately 10%) longer than the aerodynamic optimum [13, 14]. We show that the aerodynamically optimal tail length varies significantly between males, whereas the extent of streamer elongation beyond the optimum does not. Similarly, the aerodynamically optimal tail length significantly predicts observed tail length and conveys information about flight performance, whereas the extent of sexually selected exaggeration of streamer length does not. Therefore, contrary to handicap models of sexual selection, the sexually selected exaggeration of this trait provides females with little information about any aspect of mate quality     

The design of complex sexual traits in male barn swallows: associations between signal attributes

Variation in the expression of sexually selected traits among individuals is widely investigated on the premise that these traits evolved to signal male quality. Significant repeatabilities of sexual signals and their associations with condition, mating success, survivorship and age may be the signatures of sexual selection. However, little is known about the relationship between these sexual attributes. Here we studied 28 acoustic and visual traits in the barn swallow, Hirundo rustica, that may potentially function in sexual selection. Based on effect sizes calculated at the between-individual level, we assessed the relationship between repeatability, condition-dependence, attractiveness, age-dependence and viability indicator value of sexual traits using sexual signals as the units of analyses. Those traits that showed high within-year repeatability also showed high between-year repeatability, indicating that between-individual variation is consistent within and among seasons. In addition, age-dependence of traits, probably causing between-year variation, was negatively related to between-year repeatability. Condition-dependence was negatively correlated with effect sizes for the extent to which traits predicted viability. Therefore, traits that are positively related to immediate condition are those that are negatively related to survival, which may be the signature of a trade-off between current and future reproductive success ultimately reflecting signal reliability. No other significant relationship was found between trait attributes. We conclude that multiple sexual signals reflect different aspects of male quality in the barn swallow.     

Intraspecies variation in avian energy expenditure: correlates and constraints

Data on energy expenditure by 553 individuals of 28 species of small bird (10–150 g) are presented. All estimates of energy expenditure were obtained using the doubly-labelled water technique. Intraspecies variation in daily energy expenditure was found to be positively correlated with brood provisioning rates, percentage of time flying and the frequency of non-resting activity. Correlations were also shown with body-mass, body-size and several environmental factors. Published data on basal metabolic rates (BMR) sometimes differed substantially from estimates either made specifically as part of the studies considered here or calculated from allometric equations. For the purpose of interspecific comparisons, specific estimates of BMR are to be preferred. When expressed as a function of BMR, energy expenditures of free-living birds ranged from 1 + to 7 + times BMR with a mode at 3 +. Values of daily energy expenditure exceeding 4 times BMR were found in up to 48% of species and 30% of individuals, so that, contrary to earlier suggestions, 4 times BMR is not a universal upper limit to the sustained work rate of small birds. Observed upper limits tended to be higher in species with energy-expensive foraging habits. Energy expended by breeding birds is likely to involve a balance between the benefits a greater expenditure has for offspring production and any fitness penalty associated with the high level of energy expediture which nest provisioning involves.     

Some effects of energy costs on foraging strategies

We consider the effect of including energy costs on the optimal strategy for animals exploiting a depleting food resource. In the context of central place foraging this leads to the problem of what load size should be brought back to the central place. Two strategies are discussed: (i) maximize gross rate of energy delivery and (ii) maximize net rate of energy delivery. The optimal load size (or optimal patch time) for net maximizers is not always larger than for gross maximizers, as has been claimed. Instead, the difference in optimal load size has the same sign as the difference between metabolic rates of travelling and foraging. We point out that the influence of costs has not always been correctly incorporated in experimental tests of the theory.     

Experimental manipulation of tail ornament size affects the hematocrit of male barn swallows (Hirundo rustica)

Ornamental tail feathers of male barn swallows (Hirundo rustica) confer an advantage in sexual selection because long-tailed males are preferred by females. However, the size of tail ornaments exceeds the natural selection optimum and males are predicted to pay an energetic cost for flying, directly related to tail length. An increase in hematocrit is an adaptive response to enhance oxygen uptake, for example during periods of intense locomotory activity. In this study, we analyzed the effect of experimental manipulation of tail length on the hematocrit of male barn swallows from an Italian and a Spanish population. We predicted that the natural decrease in hematocrit during the breeding season would be reduced by experimental elongation and enhanced by experimental shortening of tail ornaments. The results showed that the decrease in hematocrit was significantly different among tail treatments, and tail-elongated males had the smallest hematocrit reduction. In Italy, the hematocrit of tail-elongated males did not change after tail manipulation, while that of two control groups and tail-shortened males decreased. A comparatively high hematocrit in males with experimentally enlarged tail ornaments may be a response to increased energetic requirements and, hence, to oxygen demands for flying imposed by their tail morphology. Received: 22 June 1996 / Accepted: 23 October 1996     

Daily energy expenditure of singing great reed warblers Acrocephalus arundinaceus

According to honest signalling theory, signals must be costly to produce to retain information about the signaller's quality. The song produced by male birds during breeding is a vocal "ornament" used for intra- and inter-sexual purposes. The energetic cost of this vocal signal remains a contentious issue. We used the doubly labelled water method to measure field metabolic rate by estimating CO₂ production and then convert this to daily energy expenditure (DEE) in great reed warbler males singing under natural conditions (10 at low to moderate intensity and 7 at very high intensity from dawn to dusk). There was a significant positive relationship between singing intensity and DEE. From this relationship we extrapolated the average DEE for intensely singing males (i.e., males producing song sounds 50% of the time and hence sitting at their elevated song post in the top of a reed stem more or less continuously throughout the ∼20 h of daylight) to 3.3×BMR (basal metabolic rate) and for non-singing males to 2.2×BMR. The mean DEE measured for the seven males singing with very high intensity was 3.1×BMR. The maximum measured DEE for a single male was 3.9×BMR, i.e. close to the maximum sustainable DEE (4×BMR), and the minimum DEE was 2.1×BMR for a male singing at very low intensity. These results imply that producing intensive advertising song in birds may incur a substantial cost in terms of increased energy expenditure.     

Behavioural and energetic responses to body state in male and female barn swallows (Hirundo rustica)

Females and males often have different roles when attending young. The factors responsible for shifts in the balance of effort when both sexes provision their young are not clear. This study asked if sex-specific behaviour and provisioning rules in barn swallows (Hirundo rustica) were dependent on individual body-state or affected by the state of their provisioning partner. To assess this male and female barn swallows underwent overnight warming manipulations whilst provisioning 10-14 day old nestlings, a time when energetic demands are maximal. The overnight warming treatment reduced thermoregulatory costs and provided birds with extra energy reserves at dawn. Only one member of a pair was manipulated in this way, whilst their partners were left un-manipulated to assess their response to their partner's elevated body-state. The energetic and behavioural responses to these manipulations were followed over the subsequent 24-h. I found that warmed male and female barn swallows increased their energy expenditure and nest visitation rates to the same extent after manipulation, implying a trade-off in resource allocation, which was biased towards reproductive effort. There was no effect of gender. Males paired with manipulated females showed no energetic or behavioural adjustments, however, females paired with manipulated males tended to increase both energy expenditure and nest visitation. This study provides evidence that energy reserves constrain behaviour, and that male and female swallows normally follow the same state-dependent rules when provisioning young.     

Experimental manipulation of tail length in female barn swallows (Hirundo rustica) affects their future reproductive success

Previous studies have shown no significant effect of experimentaltail length manipulation in female barn swallows (Hirundo rustica)at the beginning of a breeding season on reproductive successor behavior during that breeding season. In the present study,we investigate if tail length manipulation had any effect onreproductive performance the following year, the so-called long-termeffect, in contrast to the short-term effects already studied.We found that females with experimentally elongated externaltail feathers at the beginning of a breeding season producedless offspring during the breeding season the following yearthan did females with shortened or unmanipulated tails. Theseresults suggest that tail elongation caused flight deficienciesthat deteriorated the condition of females and eventually reducedreproductive success. The finding of long-term effects but nosignificant short-term effects for female tail elongation suggeststhat female barn swallows have the ability to adjust immediateparental investment. Detrimental effects of long tails in femalesin terms of decreased reproductive success might explain whyfemale tails are not as long as those of males. Finally, femalesmated to long-tailed (sexually attractive) males decreased theirreproductive success the following year more than did femalesmated to short-tailed males, possibly owing to differentialparental effort causing a deterioration of their condition.     

Aerodynamic costs of long tails in male barn swallows Hirundo rustica and the evolution of sexual size dimorphism

Exaggerated tail feathers of birds constitute a standard exampleof evolution of extravagant characters due to sexual selection.Such secondary sexual traits are assumed to be costly to produceand maintain, and they usually are accompanied by morphologicaladaptations that tend to reduce their costs. The aerodynamiccosts for male barn swallows Hirundo rustica of having longtails were quantified using aerodynamics theory applied to morphologicaldata from seven European populations. Latitudinal differencesin tail length were positively correlated with differences inflight costs predicted by aerodynamics theory. A positive relationshipbetween aerodynamic costs of long tails and the degree of sexualsize dimorphism was found among populations. Latitudinal differencesin foraging costs may result in tail length being relativelysimilar in males and females in southern populations, whereasthe low foraging costs for males in northern populations mayallow them to cope with higher aerodynamic costs, giving riseto large sexual size dimorphism. Enlargement of wingspan inmales can alleviate but not eliminate the costs of tail exaggeration,and therefore differences in aerodynamic costs of male ornamentswere maintained among populations. Sexual size dimorphism in thebarn swallow arises as a consequence of latitudinal differencesin the advantages of sexual selection for males and the costsof long tails for males and females.     

Daily energy expenditure in active and inactive persons with spinal cord injury.

Daily energy expenditure (DEE) was evaluated seventeen male subjects with spinal cord injury (SSCI), who had active (N = 9) and inactive (N = 8) lifestyles, and in seven normal males. DEE was estimated from the mean 24-hr heart rate and heart rate-energy expenditure relationship determined in an arm cranking exercise. The DEE of SSCI on active days did not differ from those of normal subjects. On inactive days, SSCI had significantly lower DEE than on active days and than normal subjects. In contrast, the mean 24-hr heart rates of SSCI on active days and inactive SSCI were significantly greater than those of normal subjects, suggesting that SSCI, particularly inactive SSCI, exhibited a slight degree of tachycardia when compared to normal subjects. On inactive days, the DEE was fairly independent of the level of spinal cord injury. However, on active days, there was a clear tendency for SSCI with a low lesion level to have a higher DEE. Even if a SSCI with a high lesion level engaged in active sports, his DEE was relatively small. This lower DEE was largely attributable to the smaller functional muscle mass due to the limitation of physical activity.     

Daily energy expenditure and cell-mediated immunity in pied flycatchers while feeding nestlings: interaction with moult

Ecological immunology posits a trade-off between parental effort and immunocompetence underlying the cost of reproduction. The moult-breeding overlap observed in several bird species represents a conflict in resource allocation between two energy-demanding processes. Moult processes have been associated with enlargements of immune system organs. In the present study. we measured simultaneously daily energy expenditure (DEE) and the T-cell-dependent immune response of pied flycatchers, Ficedula hypoleuca, caring for grown nestlings. We used the doubly labelled water technique and the phytohaemagglutinin (PHA) injection assay on both males and females, while recording provisioning rates and moult scores. DEE and the PHA response were negatively correlated for females, but not for males. A significantly higher proportion of males than females initiated moult. Provisioning rates were strongly correlated with DEE for females but only for non-moulting males. The DEE of moulting males was marginally correlated with moult score. For moulting males, there was a marginally significant positive correlation between moult score and immune response. The trade-off between DEE and immunity for females could underlie the cost of reproduction. However, the moult-breeding overlap found in males may offset this trade-off, thereby reducing the implications of immunosuppression for parental survival.     

The relationship between foraging behaviour and energy expenditure in Antarctic fur seals

By using time-depth recorders to measure diving activity and the doubly-labelled water method to determine energy expenditure, the relationship between foraging behaviour and energy expenditure was investigated in nine Antarctic fur seal females rearing pups. At-sea metabolic rate (MR) (mean of 6.34 ± 0.4 W. kg^-1; 4.6 times predicted BMR) was positively correlated to foraging trip duration (mean of 4.21 ± 0.54 days; r²= 0.5, P < 0.04). There were no relationships between MR and the total number of dives, the total time spent diving or the total vertical distance travelled during the foraging trip. There was, however, a close negative sigmoidal relationship (r²= 0.93) between at-sea MR and the proportion of time at sea spent diving. This measure of diving behaviour may provide a useful, inexpensive means of estimating foraging energy expenditure in this species and possibly in other otariids. The rate of diving (m.h^-1) was also negatively related to at-sea MR (r²= 0.69, P < 0.005). Body mass gain during a foraging trip had a positive relationship to the time spent at sea (r²= 0.58, P < 0.02) and the total amount of energy expended while at sea (r²= 0.72, P < 0.004) such that, while females undertaking long trips have higher metabolic rates, the energetic efficiency with which females gain mass is independent of the time spent at sea. Therefore, within the range of conditions observed, there is no apparent energetic advantage for females in undertaking foraging trips of any particular duration.     

Immune response of male barn swallows in relation to parental effort, corticosterone plasma levels, and sexual ornamentation

Life-history theory posits trade-offs between fitness components. Reproduction negatively affects physiology and immune systemfunctioning, and the effect of this form of stress may be mediatedby glucocorticosteroids. We manipulated brood size of barnswallows (Hirundo rustica) to study the effect of stress arisingfrom reproductive effort on corticosterone levels of males.We also measured T-cell—mediated immunocompetence by intradermally injecting birds with phytohemagglutinin, whichis mitogenic to T-lymphocytes. The results confirmed the predictionof a negative effect of parental effort on lymphoproliferativeresponse. We found no covariation between immune response andcorticosterone levels. Males with long tails, an ornament currentlyunder directional sexual selection, had a relatively large T-cell response to the mitogen, consistent with models of parasite-mediated sexual selection predicting higher levels of immune defensein highly ornamented males. In addition, males with large sexualornaments had relatively low corticosterone levels at the endof the parental period. These results can be reconciled withthe hypothesis proposing a trade-off between parental activitiesand adaptive immunity and suggest that highly ornamented malesare less exposed or less susceptible to stress arising fromparental effort.     

Daily energy expenditure across the course of lactation among urban Bangladeshi women

Measures of energy intake of lactating women in developing countries show that intakes are often lower than those recommended by international bodies, while fat-mass losses are often substantially less than the 3-4 kg used in the calculations of recommendations, suggesting that physiological adaptation must be commonplace among such women. The cost of lactation may be met by reduction in energy expenditure, including reduced physical activity, as well as by mobilization of bodily soft tissue. However, daily energy expenditure of lactating women has been shown to increase across the course of lactation among women in a rural population in the Philippines and an urban population in India, with a decline in body weight across the course of lactation in both studies. In the present study, total daily energy expenditure and anthropometric body composition were measured longitudinally in 68 mothers from a poor urban area of Dhaka, Bangladesh, at 0, 1, 2, 4, and 8 months of lactation, to determine whether the increasing energy expenditure across lactation observed elsewhere also occurs in Bangladeshi women. In addition, the extent to which an extended period of lactation was accompanied by weight and body fat change in these women was determined. Energy expenditure by heart-rate monitoring and activity report, and body composition from anthropometry was carried out four times across the 8-month period of lactation. A small decline in body fat mass and a significant increase in total energy expenditure across this period were observed, confirming similar observations elsewhere in the developing world.     

Plumage coloration, not length or symmetry of tail-streamers, is a sexually selected trait in North American barn swallows

Sexual adornments often vary markedly across a species' range,which presumably is owing to differences in local environmentalconditions and the associated selection pressures, such as naturalversus sexual selection or the relative signaling value of differentornamental traits. However, there are only a few reported examplesin which the information content of mating signals varies geographically,and even fewer in which a set of secondary sexual traits servesdifferent signaling functions in different populations. Classicstudies of sexual selection in the European barn swallow (Hirundorustica rustica) demonstrate that elongate tail-streamers provideseveral reproductive advantages to males and females and areused as reliable signals of mate quality. Here, we show thattail-streamers do not appear to confer these same benefits ina population of barn swallows from North America (Hirundo rusticaerythrogaster). Instead, ventral plumage coloration, which ismore exaggerated in North American swallows compared with theirEuropean counterparts, predicts patterns of assortative matingand annual reproductive success in H. r. erythrogaster. Theseobservations support the idea that ornamental traits can servedifferent functions among animal populations and suggest thatgeographic variation in different sexual signals may facilitatepopulation divergence, which may ultimately lead to speciation.