José Javier Cuervo

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.

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

Abstract 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.

Sexually Selected Egg Coloration in Spotless Starlings

It has been recently proposed that the blue‐green coloration in eggs of many avian species may constitute a sexually selected female signal. Blue‐green color intensity would reflect the physiological condition of females, and hence it might also affect the allocation of male parental care. In this study, we use three different experimental approaches to explore the importance of sexual selection on blue‐green egg coloration of spotless starling (Sturnus unicolor) eggs. First, experimental deterioration of female body condition (by means of wing feather removal) negatively affected the intensity of blue‐green egg coloration. Second, blue‐green color intensity of artificial model eggs had a significant positive influence on paternal feeding effort. Finally, we found a negative relationship between the effect of experimental food supply on nestling immunocompetence and the intensity of blue‐green coloration of eggs, suggesting that egg color predicts nutritional conditions that nestlings will experience during development. All these results taken together strongly support a role of sexual selection in the blue‐green coloration of spotless starling eggs.

Concentration of trace elements in feathers of three Antarctic penguins: Geographical and interspecific differences

Antarctica is often considered as one of the last pristine regions, but it could be affected by pollution at global and local scale. Concentrations of Al, Cr, Mn, Fe, Ni, Cu, Zn, As, Se, Cd and Pb were determinated by ICP-MS in feathers (n = 207 individuals) of gentoo, chinstrap and Adélie penguin collected in 8 locations throughout the Antarctic Peninsula (2006-2007). The highest levels of several elements were found in samples from King George Island (8.08, 20.29 and 1.76 μg g(-1) dw for Cr, Cu and Pb, respectively) and Deception Island (203.13, 3.26 and 164.26 μg g(-1) dw for Al, Mn and Fe, respectively), where probably human activities and large-scale transport of pollutants contribute to increase metal levels. Concentrations of Cr, Mn, Cu, Se or Pb, which are similar to others found in different regions of the world, show that some areas in Antarctica are not utterly pristine.

Components of phenotypic variation in avian ornamental and non-ornamental feathers

Phenotypic variation, measured as the coefficient of variation (CV), is usually larger in secondary sexual characters than in ordinary morphological traits. We tested if intraspecific differences in the CV between ornamental and non-ornamental feather traits in 67 evolutionary events of feather ornamentation in birds were due to differences in (1) the allometric pattern (slope of the regression line when regressing trait size on an indicator of body size), or (2) the dispersion of observations around the regression line. We found that only dispersion of observations around the regression line contributed significantly to total variation. A large dispersion of observations around the regression line for ornamental feathers is consistent with these characters showing condition-dependence, supporting indicator models of sexual selection more strongly than a pure Fisher process. Ornamental feathers in males demonstrated negative allometry when regressed on tarsus length, which is a measure of skeletal body size. This finding is consistent with ornamental feather traits being subject to directional selection due to female mate preferences, where large body size is less important than in male–male competition. This pattern of phenotypic variation for avian secondary sexual characters contrasts with patterns of variation for insect genitalia, supposedly subject to sexual selection, since the latter traits only differ from ordinary morphology traits in allometry coefficient. The prevailing regime of selection (directional or stabilizing) and the effects of environmental factors are proposed to account for these differences among traits.

An analysis of continent-wide patterns of sexual selection in a passerine bird

Patterns of selection are widely believed to differ geographically, causing adaptation to local environmental conditions. However, few studies have investigated patterns of phenotypic selection across large spatial scales. We quantified the intensity of selection on morphology in a monogamous passerine bird, the barn swallow Hirundo rustica, using 6495 adults from 22 populations distributed across Europe and North Africa. According to the classical Darwin-Fisher mechanism of sexual selection in monogamous species, two important components of fitness due to sexual selection are the advantages that the most attractive males acquire by starting to breed early and their high annual fecundity. We estimated directional selection differentials on tail length (a secondary sexual character) and directional selection gradients after controlling for correlated selection on wing length and tarsus length with respect to these two fitness components. Phenotype and fitness components differed significantly among populations for which estimates were available for more than a single year. Likewise, selection differentials and selection gradients differed significantly among populations for tail length, but not for the other two characters. Sexual selection differentials differed significantly from zero across populations for tail length, particularly in males. Controlling statistically for the effects of age reduced the intensity of selection by 60 to 81%, although corrected and uncorrected estimates were strongly positively correlated. Selection differentials and gradients for tail length were positively correlated between the sexes among populations for selection acting on breeding date, but not for fecundity selection. The intensity of selection with respect to breeding date and fecundity were significantly correlated for tail length across populations. Sexual size dimorphism in tail length was significantly correlated with selection differentials with respect to breeding date for tail length in male barn swallows across populations. These findings suggest that patterns of sexual selection are consistent across large geographical scales, but also that they vary among populations. In addition, geographical patterns of phenotypic selection predict current patterns of phenotypic variation among populations, suggesting that consistent patterns of selection have been present for considerable amounts of time.

Hatching asynchrony, sibling hierarchies and brood reduction in the Chinstrap penguin Pygoscelis antarctica

We studied patterns of chick growth and mortality in relation to egg size and hatching asynchrony during two breeding seasons (1991 and 1992) in a colony of chinstrap penguins sited in the Vapour Col rookery, Deception Island, South Shetlands. Intraclutch variability in egg size was slight and not related to chick asymmetry at hatching. Hatching was asynchronous in 78% (1991) and 69% (1992) of the clutches, asynchrony ranging from 1 to 4 days (on average 0.9 in 1991 and 1.0 days in 1992). Chicks resulting from oneegg clutches grew better than chicks in families of two in 1991. In 1992, single chicks grew to the same size and mass at 46 days of age as chicks of broods of two, suggesting food limitation in 1991 but not in 1992. In 1991, asymmetry between siblings in mass and flipper length was significantly greater in asynchronous than in synchronous families during the initial guard stage, but these differences disappeared during the later créche phase. In 1992, asymmetry in body mass increased with hatching asynchrony and decreased with age. Only the effect of age was significant for flipper length and culmen. Asymmetries at 15 days were similar in both years, but significantly lower in 1992 than in 1991 at 46 days of age. There were relatively frequent reversals of size hierarchies during both phases of chick growth in the two years, reversals being more common in 1991 than in 1992 for small chicks. In 1991, survivors of brood reduction grew significantly worse than chicks in nonreduced broods. In both years, chicks of synchronous broods attained similarly large sizes before fledging as both A and B chicks of asynchronous broods. In 1991, chick mortality rate increased during the guard stage due to parental desertions, decreased during the transition to crèches (occurs at a mean age of 29 days) and returned to high constant levels during the crèche stage, when it is mostly due to starvation (in total 66% of hatched chicks survived to fledging). In contrast, in 1992, mortality was relatively high immediately after hatching and almost absent for chicks older than 3 weeks (87% of chicks survived to fledging). Mortality affected similarly one- and two-chick families. In 1991, asynchronous families suffered a significantly greater probability of brood reduction than synchronous families, but this probability was not significantly related to degree of asymmetry between siblings. No association between asynchrony and mortality was found in 1992. These results show that there is food limitation in this population during the crèche phase in some years, that asynchronous hatching does not facilitate early brood reduction and that it does not ensure stable size hierarchies between siblings. Brood reduction due to starvation is not associated to prior asymmetry and does not facilitate the survival or improve the growth of the surviving chick. Asynchronous hatching may be a consequence of thermal constraints on embryo development inducing incubation of eggs as soon as they are laid.

Is the relation between colour and immune response mediated by nutritional condition in spotless starling nestlings

The hypothesis that nestling coloration is important for parenteoffspring communication, because it influences parental feeding decisions, has received strong experimental support. In European starlings, Sturnus vulgaris, and Alpine swifts, Apus melba, manipulation of ultraviolet reflectance of nestlings’ mouth and skin affected the amount of food parents provided, and skin brightness of starling nestlings predicted their T-cell-mediated immune response. Therefore, a link between nestling coloration and immunity, mediated by parental effort, was suggested. We explored this hypothesis by experimentally feeding some spotless starling, Sturnus unicolor, nestlings while leaving others in the same nest as a control. First, we found a significant effect of food supplementation on nestlings’ immune response, which is a requirement for the hypothesis. Second, we confirmed in spotless starlings the association between skin brightness and ability to raise an immune response. However, this correlation disappeared when we controlled for between-nest variation. These results suggest that parental feeding preference is not the only factor explaining nestling immunity, and that covariation between mean brood nestling coloration and parental quality, and/or intrinsic (i.e. genetic) quality of nestlings, may explain the association between immunity and coloration of nestlings. Finally, within-nest variation in nestling coloration partially explained immune responses because food supplementation had more effect on nestlings with brighter skin. We discuss these results as possible evidence of nestling coloration partially reflecting intrinsic characteristics that affect both ability to produce efficient immune responses and parental feeding preferences.

The function of feeding chases in the chinstrap penguin, Pygoscelis antarctica

Crrching chinstrap penguin chicks chase their parents on the run before being fed. This characteristic behaviour of Pygoscelid penguins has been interpreted in several ways. In an observational study of several colonies in a rookery on Deception Island, South Shetlands, the frequency and duration of feeding chases in families with one and two chicks were compared. Significantly more feedings occurred outside the crrche in two-chick than in one-chick families. Chases were significantly more frequent and prolonged in families with two than in those with single chicks. This difference was independent of the number of chicks present in the interactions (one or two siblings in two-chick families). Chases during feedings by single chicks were significantly less frequent and prolonged than those by one chick when its sibling was absent. Siblings chasing more intensively obtained more feedings. There was no increase in chasing effort with chick age. These results suggest that feeding chases allow parents to regulate food distribution between siblings according to their needs or hunger but they could also allow brood reduction in times of food crises.

Energetic cost of tail streamers in the barn swallow (Hirundo rustica)

Different hypotheses stress the importance of natural or sexual selection to explain the evolution and maintenance of long outermost tail feathers in the barn swallow (Hirundo rustica). Since energy costs are predicted to arise from tail length manipulation, we measured the daily energy expenditure in three experimental groups (tail-shortened, tail-elongated, and control birds) with the doubly labelled water technique. Though we did not directly measure flight cost, we assumed this to be positively related to daily energy expenditure. Mass independent daily energy expenditure (kJ/mass0.67 day) average daily metabolic rate (ml CO2/g h), and water flux (ml H2O/g day) did not show any significant difference among treatments in either sex. Males had higher values than females for the three parameters. Males with short original tail length experienced a higher water flux than originally long-tailed males. Females that laid more eggs during the breeding season or had heavier broods also showed higher levels of water flux which could imply a higher food intake. Our expectation of finding energetic costs of manipulated tail length in barn swallows with an integrated measure of metabolism was not fulfilled, and we did not find evidence for behavioural changes in the birds involved in the experiment.