Male Barn Swallows Hirundo Rustica Research Articles

Experimental manipulation of female tail length did not cause differential allocation by males in the barn swallow

AbstractThe evolution and maintenance of female ornamentation has attracted increasing attention, because the previous explanation, that is a non‐functional copy of functional male ornamentation, seems insufficient to explain female ornamentation. A post‐mating sexual selection, differential allocation, may be more common than pre‐mating sexual selection, but few studies have investigated differential allocation by males. Here, we studied differential allocation of incubation investment by male barn swallows Hirundo rustica, a model species for the study of sexual selection, because our previous correlative study demonstrated a positive relationship between female tail length and male incubation investment. We manipulated the length of the outermost tail feathers in females after clutch completion and examined whether males adjust incubation investment according to female ornamentation. Because extra‐pair paternity is virtually absent in the study population, we were able to study differential allocation based on the tradeoff between current and future reproductive investments, rather than the tradeoff between current paternal investment and additional mating effort. The experimental treatment had no significant effect on male nest attentiveness, whereas female tail length before manipulation predicted male nest attentiveness. The observed pattern is consistent with differential access; that is, well‐ornamented individuals have greater access to mates with high reproductive (parental) ability, rather than differential allocation during incubation. Alternatively, males can directly assess eggs in their nests, and thus, as seen in other species, males might adjust their incubation investment based on the egg characteristics of long‐tailed females.

Experimental tail elongation in male Barn SwallowsHirundo rusticareduces provisioning of young, but only in second broods

At the beginning of the breeding season, the outermost tail feathers of 31 male Barn SwallowsHirundo rusticawere either shortened by 20 mm, elongated by 20 mm or left unmanipulated. In first broods, the number of feeding bouts (per nestling per hour) by males and females did not differ significantly among experimental groups. However, in second broods, males with elongated tails fed their nestlings less often than males with shortened or unmanipulated tails. Male tail elongation may have been detrimental to flight, making capture of insects more difficult. Females paired to long‐tailed males did not compensate for this reduction in feeding by males and their nestlings received less food. Neither feeding rates nor brood size differed significantly between first and second broods. Variation in abundance of large insects at different times of the day (high around noon and low in the morning and afternoon) matched variation in feeding rates, supporting the importance of large insects in the Barn Swallow diet. Different feeding rate patterns in first and second broods by males with elongated tails could have at least three explanations: (1) such males were able to adjust their parental effort to some extent, temporarily compensating for an imposed handicap; (2) they were more sensitive than males in the other experimental groups to late season food shortage (a decrease in numbers of large insects, the main Barn Swallow prey, was observed as the season progressed); and (3) they were more sensitive than males in the other experimental groups to the deterioration in their physical condition after first broods.

Dynamic Paternity Allocation as a Function of Male Plumage Color in Barn Swallows

Paternity in male animals can be influenced by their phenotypic signals of quality. Accordingly, the behavior underlying patterns of paternity should be flexible as signals of quality change. To evaluate the dynamics of paternity allocation, we analyzed paternity before and after manipulating plumage coloration, a known signal of quality, in male barn swallows Hirundo rustica. We found that, in successive breeding bouts, only males whose plumage color was experimentally enhanced received greater paternity from their social mates, demonstrating evidence for flexible and dynamic paternity allocation and the importance for males of maintaining signals of quality well after pair bond formation.

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 example of evolution of extravagant characters due to sexual selection. Such secondary sexual traits are assumed to be costly to produce and maintain, and they usually are accompanied by morphological adaptations that tend to reduce their costs. The aerodynamic costs for male barn swallows Hirundo rustica of having long tails were quantified using aerodynamics theory applied to morphological data from seven European populations. Latitudinal differences in tail length were positively correlated with differences in flight costs predicted by aerodynamics theory. A positive relationship between aerodynamic costs of long tails and the degree of sexual size dimorphism was found among populations. Latitudinal differences in foraging costs may result in tail length being relatively similar in males and females in southern populations, whereas the low foraging costs for males in northern populations may allow them to cope with higher aerodynamic costs, giving rise to large sexual size dimorphism. Enlargement of wingspan in males can alleviate but not eliminate the costs of tail exaggeration, and therefore differences in aerodynamic costs of male ornaments were maintained among populations. Sexual size dimorphism in the barn swallow arises as a consequence of latitudinal differences in the advantages of sexual selection for males and the costs of long tails for males and females.

No evidence for adjustment of sex allocation in relation to paternal ornamentation and paternity in barn swallows

Sex allocation theory predicts that parents should adjust investment in sons and daughters according to relative fitness of differently sexed offspring. In species with female preference for highly ornamented males, one advantage potentially accruing to parents from investing more in sons of the most ornamented males is that male offspring will inherit characters ensuring sexual attractiveness or high‐quality genes, if ornaments honestly reveal male genetic quality. Furthermore, in species where extra‐pair fertilizations occur, offspring sired by an extra‐pair male are expected to more frequently be male than those of the legitimate male if the latter is of lower quality than the extra‐pair male. We investigated adjustment of sex ratio of offspring in relation to ornamentation of the extra‐pair and the social mate of females by direct manipulation of tails of male barn swallowsHirundo rustica. Molecular sexing of the offspring was performed using the W chromosome‐linked avian chromo‐helicase‐DNA‐binding protein (CHD) gene while paternity assessment was conducted by typing of hypervariable microsatellite loci. Extra‐pair offspring sex ratio was not affected by ornamentation of their biological fathers relative to the experimental ornamentation of the parental male. Experimental ornamentation of the parental males did not affect the sex ratio of nestlings in their broods. Female barn swallows might be unable to bias offspring sex ratio at hatching according to the quality of the biological father. Alternatively, fitness benefits in terms of sexual attractiveness of sons might be balanced by the cost of compensating for little parental care provided by highly ornamented parental males, if sons are more costly to rear than daughters, or the advantage of producing more daughters, if males with large ornaments contribute differentially more to the viability of daughters than sons.

Testing the functional significance of tail streamers

Studies of the evolution of elaborate ornaments have concentrated on their role in increasing attractiveness to mates. The classic examples of such sexually selected structures are the elongated tails of some bird species. Elongated tails can be divided into three categories: graduated tails, pin tails and streamers. There seems to be little debate about whether graduated and pin tails are ornaments; i.e. costly signals used in mate choice. However, in the case of streamers there is considerable discussion about their function. It has been suggested that tail streamers could be (i) entirely naturally selected, (ii) entirely sexually selected, (iii) partly naturally and partly sexually selected. The prime example of a species with tail streamers is the swallow (Hirundo rustica) in which both sexes have tail streamers. In this paper we discuss the aerodynamic consequences of different types of manipulation of the streamer and/or outer tail feather. We make qualitative predictions about the aerodynamic performance of swallows with manipulated tail streamers; these predictions differ depending on whether streamers have a naturally or sexually selected function. We demonstrate that these hypotheses can only be separated if tail streamers are shortened and changes in aerodynamic performance measured during turning flight.

Sexual selection in the barn swallow Hirundo rustica. VI. Aerodynamic adaptations

AbstractSecondary sexual characters are assumed to be costly to produce and maintain, and this will select for morphological modifications that reduce the magnitude of such costs. Here we test whether a feather ornament, the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject to a directional female mate preference, and other aspects of the morphology used for flight have been modified to increase aerodynamic performance. This was done by making comparisons among sexes within populations, among individuals varying in tail length within populations, and among populations from different parts of Europe. Male barn swallows experienced reduced drag from their elongated tail feathers by morphological modifications of the ornamental feathers as compared to females. Morphological features of the outermost tail feathers were unrelated to tail length in both males and females within populations. Wing and tail morphology (length of central tail feathers and wings, wing span, wing area, wing loading, and aspect ratio) was modified in males compared to females. Barn swallows with long tails had morphological tail and wing modifications that reduced the cost of a large ornament, and similar modifications were seen among populations. The costs of the exaggerated secondary sexual character were therefore reduced by the presence of cost‐reducing morphological modifications. The assumptions of reliable signalling theory, that signals should be costly, but more so to low than to high quality individuals, were not violated because long‐tailed male barn swallows had the largest cost‐reducing morphological characters.

Foraging costs of a tail ornament: Experimental evidence from two populations of barn swallows Hirundo rustica with different degrees of sexual size dimorphism

Secondary sexual characters are assumed to be costly to produce or maintain. A test of this assumption was performed using the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject of a directional female mate preference. A possible cost of sexual signalling in male barn swallows arises from increased flight cost during foraging in this aerially insectivorous species. A longer tail may impose a greater drag during flight and thereby affect foraging ability. This was tested by determining the relationship between experimentally modified male tail lengths and number and size of prey delivered to offspring in Spain, where sexual size dimorphism in tail length is small, compared to Denmark, where dimorphism is large. Food boluses contained significantly fewer small insects in Spain than in Denmark. Males with elongated tails captured more and smaller insects while males with shortened tails captured fewer and larger prey items at both sites. Males with naturally long tails were less affected by experimental treatment in terms of effects on the number and the size of prey delivered to their offspring, a finding consistent with a long tail being a condition-dependent viability indicator. The effect of a given degree of tail manipulation on prey size and number of prey per bolus was larger in Spain than in Denmark. These results demonstrate that (1) tail length in male barn swallows affects foraging, and (2) larger sexual size dimorphism occurs where the foraging cost of an increment in ornament size is smallest.

DIFFERENTIAL COSTS OF A SECONDARY SEXUAL CHARACTER: AN EXPERIMENTAL TEST OF THE HANDICAP PRINCIPLE.

The evolution of reliable signaling can be explained by the handicap principle, which assumes that (1) the cost of a signal guarantees its reliability, and (2) cheating is prevented because the cost of a unit of display is greater for low-quality than for high-quality individuals. A test of these two assumptions was performed using manipulations of the length of the outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject to a directional female mate preference. We found that survival decreased with tail elongation and increased with tail shortening of males, supporting the assumption that the secondary sexual character is costly. Naturally long-tailed males were better able to survive with an elongated tail, whereas naturally short-tailed males improved their survival following tail shortening. This observation supports the second assumption of a differential cost of a signal. One mechanism imposing differential costs on sexually signaling barn swallows is foraging. Males with elongated tails captured smaller, less profitable Diptera, whereas males with shortened tails captured large, profitable prey items. The conditions for reliable sexual signaling by the tail ornament of male barn swallows are thus fulfilled.