F. de Lope

Female reproductive effort depends on the degree of ornamentation of their mates

Sexual selection theory assumes that secondary sexual characters do not influence female reproductive effort. Female animals may invest relatively more in reproduction if they acquire mates of high phenotypic quality, because offspring sired by preferred males may be relatively more viable than offspring sired by less preferred males. Here we report for the first time in a field study that females of the monogamous barn swallow Hirundo rustica adjust their reproductive effort to the attractiveness of their mates. Experimental manipulation of male tail length, which is a trait currently subject to a directional female mating preference, affected the reproductive effort by females in single broods as well as their decision on the seasonal number of clutches. These results, and those of previous experiments, demonstrate that female barn swallows assess the quality of their mates throughout the reproductive season and adjust their reproductive decisions accordingly. This result has important implications for the theory of sexual selection and for the possibility of testing current models of female mate preferences, because the viability of offspring will be confounded by differential reproductive effort.

Effects of malaria double infection in birds: one plus one is not two

Avian malaria parasites are supposed to exert negative effects on host fitness because these intracellular parasites affect host metabolism. Recent advances in molecular genotyping and microscopy have revealed that coinfections with multiple parasites are frequent in bird–malaria parasite systems. However, studies of the fitness consequences of such double infections are scarce and inconclusive. We tested if the infection with two malaria parasite lineages has more negative effects than single infection using 6 years of data from a natural population of house martins. Survival was negatively affected by both types of infections. We found an additive cost from single to double infection in body condition, but not in reproductive parameters (double‐infected had higher reproductive success). These results demonstrate that malaria infections decrease survival, but also have different consequences on the breeding performance of single‐ and double‐infected wild birds.

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.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.

Increased detrimental effects of ectoparasites on their bird hosts during adverse environmental conditions

Parasites are usually assumed to negatively affect fitness components of their hosts, particularly during adverse environmental conditions. We experimentally tested whether the ectoparasitic house martin bug (Oeciacus hirundinis) had larger negative effects on its house martin (Delichon urbica) host during second clutches. Conditions for reproduction decreased during the season, being less favourable for the second clutch than the first clutch. This was demonstrated by smaller clutch sizes, lower breeding success and smaller and lighter nestlings during the second clutch, as compared to the first, in the absence of parasites. Experimental manipulation of the intensity of parasitism resulted in the reproductive success of the host being reduced with increasing numbers of house martin bugs. This effect was much stronger during the second, as compared to the first, clutch. The house martin bug therefore had a greater detrimental effect on the reproduction of its host when conditions for reproduction were poor.

Sexual selection and tail streamers in the barn swallow

The functional significance of elongated, narrow tips of the tail feathers of certain birds, so–called tail streamers, has recently been discussed from an aerodynamic point of view, and the effects of sexual selection on such traits have been questioned. We review our long–term field studies using observational and experimental approaches to investigate natural and sexual selection in the barn swallow, Hirundo rustica, which has sexually size–dimorphic outermost tail feathers. Experimental manipulation of the length of the outermost tail feathers has demonstrated sexual selection advantages of tail elongation and disadvantages of tail shortening, with opposite effects for natural selection in terms of foraging efficiency, haematocrit and survival. These findings are contrary to the prediction of a general deterioration from both shortening and elongation, if the tail trait was determined solely by its effects on aerodynamic efficiency and flight manoeuvrability. Patterns of sexual selection in manipulated birds conform with patterns in unmanipulated birds, and selection differentials for different components of sexual selection in manipulated birds are strongly positively correlated with differentials in unmanipulated birds. Age and sex differences in tail length, and geographical patterns of sexual size dimorphism, are also consistent with sexual selection theory, but inconsistent with a purely natural selection advantage of long outermost tail feathers in male barn swallows.

Parasitism, immune response and reproductive success in the house martin Delichonurbica

Abstract Parasites often exert strong selection pressures on their hosts that have evolved anti-parasite defences to counter the negative effects of parasites. We studied the relationship between intensity of parasitism, one aspect of host immune response, and host reproductive success, using the house martin bug Oeciacushirundinis and its house martin Delichonurbica host as a model system. Experimental manipulation of parasite load of nests during laying of the first clutch altered the intensity of parasitism. Parasites reduced the reproductive success of their hosts measured in terms of body condition and survival of nestlings. Host immune response, measured as the concentration of gammaglobulins and total plasma proteins, was positively associated with parasite reproduction, estimated as the number of juvenile parasites, but was only weakly related to the intensity of adult parasites. The concentration of gammaglobulins was negatively related to nestling body mass, implying a trade-off between immune function and body condition. Parasite reproduction thus exerts a cost on hosts by increasing the immune response.

Yolk androgens in the barn swallow (Hirundo rustica): a test of some adaptive hypotheses.

Maternal effects such as androgen in avian eggs can mediate evolutionary responses to selection, allowing manipulation of offspring phenotype and promoting trans‐generational adaptive effects. We tested the predictions of two adaptive hypotheses that have been proposed to explain female variation in yolk androgen allocation in birds, using the barn swallow Hirundo rustica as a model. We found no support for the first hypothesis proposing that yolk androgen varies as a function of breeding density in order to prepare offspring for different breeding densities. However, we found experimental support for the hypothesis that female yolk androgen allocation depends on mate attractiveness and that it constitutes an example of differential allocation. Females increased the concentration of androgens in their eggs when mated to males with experimentally elongated tails. Female phenotypic quality as measured by arrival date and clutch size was positively related to egg androgen concentration, consistent with the hypothesis that this is a costly investment, constrained by female condition. We found correlative evidence of a direct relationship between egg androgen concentration and performance of offspring as measured by mass increase.

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

Secondary 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.

Nest and egg crypsis in the ground-nesting stone curlew burhinus oedicnemus

We studied the selective advantage of nest and egg crypsis in the Stone Curlew Burhinus oedicnemus. Experimental removal of camouflaging nest material resulted in replacement of a similar amount and size of nest material for the specific nest site. The match between the colour of the eggs and the ground was usually perfect and mismatches in coloration increased the predation rate by visually searching avian nest predators. Both nest building and choice of nest substrate appeared to increase crypsis and improved hatching success.

Ectoparasites and host energetics: house martin bugs and house martin nestlings

We measured the energy cost of ectoparasitism in an experimental study of the house martin bug Oeciacus hirundinis and its main host, nestlings of the house martin Delichon urbica. Nests were randomly assigned to inoculation with 0 (control) 10, or 100 bugs during egg laying, and this resulted in statistically significant differences in parasite loads following fledging of nestlings. Parasite loads negatively affected house martin nestlings as estimated from their body mass at age 16 days and from mass loss estimated over 1 day late in the nestling period. Daily energy expenditure (KJ/d), average daily metabolic rate (ml CO2/g h), and mass independent daily energy expenditure (kJ/mass0.67d) did not differ significantly between experimental treatments. However, average daily metabolic rate increased with increasing intensities of ectoparasitism. Mass independent daily energy expenditure also increased with higher levels of parasite infestation. These results demonstrate that the bug imposes an energy cost on its host by elevating the level of metabolism.