Social environment during molt and the expression of melanin-based plumage pigmentation in male house sparrows (<Emphasis Type="Italic">Passer domesticus</Emphasis>)

Evolutionary biologists have shown much recent interest in the costliness and signal content of colorful plumage displays in birds. Although many studies suggest that both carotenoid- and structurally-based plumage colors are condition-dependent indicators of health and nutritional state at the time ornamental feathers are grown, there is little experimental evidence supporting the idea that melanin pigmentation is a reliable signal of condition during molt. Instead, melanin-based ornamental coloration often reveals the competitive ability and dominance of individuals throughout the year. However, this work does not indicate which proximate environmental factors shape the expression of melanin pigmentation at the time of feather growth. Because of the link between melanin coloration and the social environment, it is possible that the development of brightly colored plumage may be associated with aggressive social interactions during feather molt. Here, we show that melanin-based ornamental coloration in male house sparrows (Passer domesticus) is correlated with the degree to which individuals interact aggressively with conspecifics during molt. Males that were dominant (beta, but not alpha) within captive social groups during molt grew larger badges than subordinates. Groups of males that had higher rates of aggression during molt grew larger badges than less aggressive triads. To our knowledge, this is the first demonstration that melanin pigmentation and plumage-based status badges are related to the competitive history of individuals during feather development. By coupling badge size directly with aggressive experiences during molt, birds can use their status signal to honestly indicate their likelihood of winning agonistic encounters throughout the year.     

Analysis of the cyanotoxins anatoxin-a and microcystins in Lesser Flamingo feathers†

Feathers from carcasses of the Lesser Flamingo (Phoeniconaias minor), which had died after ingesting cyanobacterial toxins (cyanotoxins) contained between 0.02 and 30.0?µg microcystin-LR equivalents per gram of feather according to HPLC and ELISA analysis of feather extracts. Anatoxin-a was detected less frequently in the Lesser Flamingo feathers, up to 0.8?µg anatoxin-a per gram of feather being recorded. When feathers from different body regions were analysed and compared for microcystins and anatoxin-a, wing feathers were found to contain the highest concentrations of these cyanotoxins, the order of concentration and frequency of analytical detection being wing?>?breast?>?head. Consistent with the presence of the microcystins and anatoxin-a in gut contents and the livers of the dead birds and negligible in vitro adsorption to feathers, the cyanotoxins associated with the feathers of the dead wild flamingos are inferred to be primarily of dietary origin.     

Chernobyl as a population sink for barn swallows: tracking dispersal using stable-isotope profiles.

Stable-isotope profiles of feathers can reveal the location or habitat used by individual birds during the molting period. Heterogeneity in isotope profiles will reflect heterogeneity in molt locations, but also heterogeneity in breeding locations, because spatial heterogeneity in molt locations will be congruent with spatial heterogeneity in breeding locations in species with high connectivity between breeding and molting sites. We used information on the congruence of spatial heterogeneity in molt and breeding location to study population processes in Barn Swallows (Hirundo rustica) from a region. near Chernobyl, Ukraine, that has been radioactively contaminated since 1986; from an uncontaminated control region near Kanev, Ukraine; and from a sample of pre-1986 museum specimens used to investigate patterns prior to the nuclear disaster at Chernobyl, from both regions. Previous studies have revealed severe reductions in Barn Swallow reproductive performance and adult survival in the Chernobyl region, implying that the population is a sink and unable to sustain itself. Female Barn Swallows are known to disperse farther from their natal site than males, implying that female stable-isotope profiles should tend to be more variable than profiles of males. However, if the Barn Swallows breeding at Chernobyl are not self-sustaining, we would expect males there also to originate from a larger area than males from the control region. We found evidence that the sample of adult Barn Swallows from the Chernobyl region was more isotopically heterogeneous than the control sample, as evidenced from a significant correlation between feather sigma13C and sigma15N values in the control region, but not in the Chernobyl region. Furthermore, we found a significant difference in feather sigma15N values between regions and periods (before and after 1986). When we compared the variances in sigma13C values of feathers, we found that variances in both sexes from post-1986 samples from Chernobyl were significantly larger than variances for feather samples from the control region, and than variances for historical samples from both regions. These findings suggest that stable-isotope measurements can provide information about population processes following environmental perturbations.     

Delayed plumage maturation in the orchard oriole (Icterus spurius): tests of winter adaptation hypotheses

Summary Explanations of delayed plumage maturation (DPM) in passerines have focused on potential breeding season advantages for non-definitive (subadult) plumage. In contrast, the molt constraint hypothesis (Rohwer and Butcher 1988) proposes that subadult plumage is a winter adaptation, increasing winter survivorship by decreasing intraspecific aggression (the winter status signaling hypothesis) or predation (the winter crypsis hypothesis). Under the molt constraint hypothesis, nondefinitive breeding plumage is non-adaptive, resulting from species-specific constraints on the number of feathers replaced during the pre-alternate (spring) molt. In studies conducted on an overwintering population of orchard orioles in Panama, I tested predictions of both the winter status signaling and molt constraint hypotheses. Contrary to predictions of winter status signaling, I found no evidence that subadult plumage reduces adult male aggression toward subadults. Agonistic encounters occurred at random with respect to plumage and the intensity of adult male/subadult male encounters was not lower than the intensity of encounters occurring within age classes. Contrary to the molt constraint hypothesis, I found no evidence that the number of feathers exchanged by subadults in their pre-alternate molt is the sole constraint on the development of subadult breeding plumage. The majority of feathers grown by molting subadult orioles during January and February were of non-definitive coloration. These results, together with results of earlier breeding season experiments which tested summer communication hypotheses for DPM in this species, suggest that subadult plumage in the orchard oriole may be non-adaptive and the result of constraints on plumage development. They also indicate, however, that the extent of the pre-alternate molt is not the sole source of that constraint.     

Diet quality affects an attractive white plumage pattern in dark-eyed juncos (Junco hyemalis)

Sexually selected traits that act as signals of quality often display some degree of condition dependence. In birds, condition dependence of ornamental plumage is often mediated by production costs related to acquisition or allocation of dietary resources. White plumage ornaments, however, have often been assumed to be inexpensive because their production requires neither pigment nor specialized feather structure. In male dark-eyed juncos (Junco hyemalis), the size of a white patch on the tail contributes to attractiveness and mating success. Using captive males, we examined the effects of diet quality on the size and brightness of the tail-white patch. After removing four tail feathers to induce replacement, we maintained subjects on a subsistence (low-protein) or enriched (high-protein) diet while induced feathers grew. Birds that received an enriched diet grew their feathers more quickly and grew larger, brighter white patches. Feather growth rate was positively correlated with the increase in the size of the tail-white patch, a relationship that was stronger in the subsistence diet group. However, within diet treatments, faster-grown feathers were slightly duller. Taken together, these results suggest that variation in diet quality may lead to condition-dependent expression of tail white and that condition dependence may be stronger in more stressful environments. We suggest a mechanism by which increased feather growth rate may lead to an increase in the size of the tail-white patch and discuss potential trade-offs between signal size and brightness.     

Moulting patterns drive within-individual variations of stable isotopes and mercury in seabird body feathers: implications for monitoring of the marine environment

One major limitation in the use of body feathers of seabirds as a monitoring tool of the trophic structure and contamination levels of marine ecosystems is the degree of heterogeneity in feather chemical composition within individuals. Here, we tested the hypothesis that moulting patterns drive body feather heterogeneity, with synchronous moult minimizing within-individual variations, in contrast to asynchronous feather growth. Chicks of white-chinned petrels Procellaria aequinoctialis (representative of bird chicks) and adults of king penguins Aptenodytes patagonicus (representative of adult penguins) that moult their body feathers synchronously showed very low within-individual variations in their feather δ¹³C and δ¹⁵N values and mercury (Hg) concentrations. By contrast, body feathers of adults of Antarctic prions Pachyptila desolata (representative of adult seabirds with asynchronous feather growth during a protracted moult) presented much higher within-individual variances for the three parameters. These findings have three important implications for birds presenting a synchronous body moult. (1) They suggest that all body feathers from the same individual have identical δ¹³C and δ¹⁵N values and Hg content. (2) They predict negligible within-individual variations in the body feather values of other useful stable isotopes, such as δ²H and δ³⁴S, as well as in the concentrations of other compounds that are deposited in the keratin structure. (3) Analysis of one or any number of pooled body feathers is equally representative of the individual. In conclusion, we recommend that long-term routine monitoring investigations focus on birds presenting synchronous rather than asynchronous moult of body feathers both in marine and terrestrial environments. This means targeting chicks rather than adults and, for seabirds, penguins rather than adults of flying species.     

Costs of overlapping reproduction and moult in passerine birds: an experiment with the pied flycatcher

The process of moult in birds requires the investment of substantial amounts of energy and nutrients in feather production and attendant processes. Flight performance may be reduced during the moult of the wing feathers, and moulting birds may suffer from an increased predation risk. These factors may explain why the moult is usually separated in time from other energetically demanding processes, such as reproduction. In this study, we investigated the importance of the temporal separation of moult from breeding activities in the pied flycatcher Ficedula hypoleuca. We induced a moult-breeding overlap by removing the two innermost primaries on both wings, thus imitating the natural loss of these feathers during the initial stage of the moult. The experiment probably did not stimulate feather regrowth, but may have reduced flight performance. Just before fledging of the young, manipulated males and females, as well as the fledglings, had a lower body condition than control birds. The return rate of adult males in the year following the manipulation was significantly lower for males in the manipulated group than for control males. The return rate of females was lower than that of males, but there was no difference between the two female groups. The number of nestlings recruited in the year after the manipulation was significantly lower for the group with an induced moult-breeding overlap than for the control group. The reproductive performance in recruited and returning old females was independent of the manipulation, and in returning adults and recruits the feather quality in terms of number of barbs/mm was independent of the manipulation. These results suggest that a moult-breeding overlap induces fitness costs in terms of reduced recruitment rate and adult survival in a long-distance migrant. Received: 17 September 1997 / Accepted after revision: 23 February 1998     

Flightlessness and the energetic cost of wing molt in a large sea duck

Although the replacement of feathers apparently represents the major event of somatic production in the annual cycle of wild birds, knowledge about the energetics of molt has always been hampered by logistical and technical difficulties, which are exacerbated by the fact that birds are able to compensate behaviorally to buffer any variation in energy demand. During wing molt, sea ducks (Mergini) and other diving birds lose all of their wing feathers at once, leading to a period of temporary flightlessness of variable duration, a condition that considerably restricts their movements and increases the probability of predation. In the present study, we present the first results aimed at quantifying the duration of flightlessness, energy expenditure, and foraging effort during molt of a wing-propelled diving bird, the Common Eider (Somateria mollissima). Data loggers were implanted in the body cavity of 13 females to record heart rate and hydrostatic pressure (depth) every two seconds for a period of 220 days. Flight frequency and duration were assessed from elevated and constant heart rate, and the absence of flight was used to quantify the duration of flightlessness, which lasted, on average, 36 +/- 8 days (mean +/- SD). Using a period of four weeks before and four weeks after the flightless period, we found that dive depth (ranging from 1 to 2 m, on average) and daily diving time did not vary during the course of the study. Daily metabolic rate increased by 9%, and resting metabolic rate by 12% from the pre-molt period to the flightless period and remained high during the post-molt period. This study indicates that the energetic costs of replacing flight remiges in female eiders are substantial, although this is not associated with any change in foraging effort, which suggests that female Common Eiders lose mass during wing molt. Finally, estimates of energy savings associated with the total absence of flights during wing molt represent 6% of daily metabolic rate or 14% of resting metabolic rate. This finding contrasts with the classical view that little or no benefit is associated with a flightless condition. We suggest that such energy savings may have favored the evolution of temporary flightlessness in diving birds.     

Use of trace element analysis of feathers as a tool to track fine-scale dispersal in birds

Tracking dispersal and migratory movements of animals over small and large spatial scales is a challenge. In birds, a promising yet underutilized tool is the trace elemental composition of feathers. The elemental profile of a feather may reveal information about the geographic origin of a bird provided that molting occurs on the breeding grounds and that elemental differences exist between breeding areas. Here, we explore the use of trace elemental composition in body feathers of the Puget Sound white-crowned sparrow Zonotrichia leucophrys pugetensis as a tool (1) to discriminate among birds collected in four different song dialect populations along a 400-km stretch of the Pacific Northwest coast and (2) to assign males singing nonlocal dialects in one population to potential natal populations. Inductively coupled plasma mass spectrometry detected 34 trace elements in sampled feathers and in a discriminant function analysis seven of these elements differed among the four source populations. Half of the six nonlocal dialect singers, who were likely to have immigrated into the focal population, were assigned to a population that matched their song dialect. Our study suggests that feather microchemistry is a promising tool for identifying geographic origins of dispersing birds over small geographic scales and in combination with other markers, such as song, may give insight into ecological and evolutionary processes.     

Differences in stable isotopes in blood and feathers of seabirds are consistent across species, age and latitude: implications for food web studies

Stable isotopes of growing feathers and blood both represent assimilated diet, and both tissues are used to study the diet and foraging distribution of marine and terrestrial birds. Although most studies have assumed that both tissues represent a difference of one trophic level to diet, the enrichment factors of blood and feathers may differ, especially where endogenous reserves are used as precursors during feather synthesis. In this study, we compare carbon and nitrogen stable isotopes of blood and simultaneously growing feathers of five species of Procellariiformes, representing five genera, different geographical regions and different life stages (chicks and adults). In all species, feathers were enriched in ¹⁵N and ¹³C compared with blood. Isotopic values of carbon and nitrogen were correlated in different tissues growing simultaneously for most species analyzed, suggesting that mathematical corrections could be used to compare different tissues. Our results imply that more care needs to be taken when comparing stable isotope signatures across studies assuming different tissues are equivalent indicators of trophic ecology.     

Differences in δ13C and δ15N values between feathers and blood of seabird chicks: implications for non-invasive isotopic investigations

Blood and feathers are the most targeted tissues for isotopic investigations in avian ecology, primarily because they can be easily and non-destructively sampled on live individuals. Comparing blood and feather isotopic ratios can provide valuable information on dietary shifts, trophic specialization and migration patterns, but it requires a good knowledge of the isotopic differences between the two tissues. Here, δ¹³C and δ¹⁵N values of whole blood (in blood cells of a few species) and simultaneously grown body feathers were measured in seabird chicks to quantify the tissue-related isotopic differences. Seabirds include 27 populations of 22 wild species that were sampled in 2000–2008, and a review of the literature added 8 groups (including adult birds) to the analysis. The use of a large data set that overall encompasses wide δ¹³C and δ¹⁵N ranges allowed us to depict for the first time accurate relationships between blood and feather isotopic ratios across avian taxa. Blood was impoverished in ¹³C and generally in ¹⁵N compared with feathers. Both mean δ¹³C and δ¹⁵N values of feathers and blood were highly positively and linearly related [feather δ¹³C = 0.972 (±0.020) blood δ¹³C + 0.962 (±0.414), and feather δ¹⁵N = 1.014 (±0.056) blood + 0.447 (±0.665), respectively; both P < 0.0001]. The regressions should be applied to mathematically correct feather or whole blood δ¹³C and δ¹⁵N values when comparing isotopic ratios within and between ecological studies on birds.     

广州市白云山雀形目鸟体内汞、砷与硒残留分析

测定了广州市白云山白头鹎(Pycnonotus sinensis)、暗绿绣眼鸟(Zosterops japonicus)、大山雀(Parus major)和白眉鵐(Emberiza tristrami )飞羽、胸肌和肝脏中的汞、砷和硒残留量。汞和砷在鸟体内残留量的分布模式都是,羽毛>肝脏>肌肉;而硒在鸟体内残留量的分布模式是肝脏>肌肉>羽毛,与汞和砷的分布模式不同。鸟组织中的汞质量分数存在种间差异,但不同组织间差异程度不同,白眉鵐肝脏、肌肉中的汞质量分数极显著(或显著)高于暗绿绣眼鸟肝脏、肌肉中的汞质量分数,而2种鸟羽毛中的汞质量分数没有显著差异。鸟组织中砷和硒质量分数的种间差异不明显。羽毛对汞有高的富集作用,用羽毛监测环境中的汞污染比用肝脏和肌肉监测环境中的汞污染更灵敏。在鸟组织中,汞质量分数、砷质量分数和硒质量分数间的相关性较差,只有白眉鵐羽毛中的汞质量分数与硒质量分数存在显著相关性,其回归方程为Y汞=0.657X硒-0.237(p=0.025)。几个地区鸟组织中汞、砷和硒的质量分数的比较表明,广州市白云山受重金属和硒污染程度较低。     

Exposing males to female scent increases the cost of controlling <Emphasis Type="Italic">Salmonella</Emphasis> infection in wild house mice

Secondary sexual characters often provide indicators of a male’s resistance to infectious diseases to rivals and potential mates, but it is unclear why. It is often suggested that males honestly signal their health due to energetic and other physiological trade-offs between investing into secondary sexual traits vs resistance to infectious diseases. Our aim was to determine whether such a trade-off exists using wild-derived male house mice (Mus domesticus). We exposed male mice to female scent, a manipulation that induces elevations in testosterone concentration and the expression of a variety of testosterone-mediated secondary sexual traits, and tested whether this sexual stimulation impaired the males’ ability to resolve or cope with an experimental infection (Salmonella enterica). We kept the males on a controlled diet to prevent them from compensating by eating more food. We found that sexually stimulated males were able to control bacterial growth as effectively as sham-stimulated controls; however, to do so, they lost more body mass during infection compared to the controls. In contrast, we found no evidence that sexual stimulation reduced the body mass of uninfected male mice. These results indicate that males’ responses to female odor are not immunosuppressive per se, yet they increase the energetic costs of controlling infection. Our findings support the idea that there is a physiological trade-off between secondary sexual signaling vs resistance to infectious diseases and suggest that studies using only immunocompetence assays might fail to detect such energetic trade-offs. We dedicate this paper to the late Professor Chris Barnard who conducted pioneering research on this topic.     

The effects of preen oils and soiling on the UV–visible reflectance of carotenoid-pigmented feathers

Plumage coloration, particularly when carotenoid-based, is important in social signaling in birds. Although feather color is a relatively stable trait, individuals may modify it with “cosmetic” substances such as preen oils. In addition, dirt accumulation may influence plumage coloration and further affect signal perception by receivers. Here, we analyze the separate potential effects of preen oils and soil accumulation on the reflectance properties of carotenoid-pigmented feathers across the visual range of most bird species, which includes the ultraviolet (UV). Using the yellow portion of tail feathers of Bohemian waxwings (Bombycilla garrulus), we performed two separate experiments where: (a) preen oils and/or soil were removed, or (b) preen oils (from black-billed magpies Pica pica or eagle owls Bubo bubo) were added. Preen oil addition reduced brightness but increased UV hue and yellow chroma. UV chroma was reduced by the addition of magpie (but not owl) preen oil. Soil accumulation had little effect on plumage reflectance in the UV range but significantly reduced yellow chroma. According to models of avian vision, both of these effects are detectable by birds and biologically meaningful when compared with natural variation between the sexes and age classes. We conclude that preen oil and soil accumulation can significantly affect the UV–visible reflectance of carotenoid-based plumages. As such traits typically advertise individual quality, preening and soiling have the potential to modify the information content of carotenoid-based plumage traits and how these signals are perceived by receivers.     

Reduced Feather Growth Rates of Two Common Birds Inhabiting Central Amazonian Forest Fragments

Abstract: Forest fragmentation may negatively affect populations typically found within continuous forest tracts. Some effects, such as absence from small fragments, are obvious, but other effects may be subtle and easily overlooked. We evaluated the hypothesis that forest birds dwelling in fragments, where microclimatic conditions have been shown to be hotter and drier than in continuous forest, may be in poorer physiological condition than those in the forest interior. We studied two bird species, the Wedge-billed Woodcreeper (  Glyphorynchus spirurus ) and the White-crowned Manakin (    Pipra pipra ), common to the fragmented landscape north of Manaus, Brazil. We analyzed feather growth rates in Pipra and Glyphorynchus captured in 1-, 10-, and 100-ha forest fragments and continuous forest. Mean daily feather growth rates of the outer right rectrix of birds captured in fragments were significantly slower than feather growth rates of birds captured in continuous forest. Based on recapture data, Wedge-billed Woodcreepers probably grew their feathers in sites where they were first captured. White-crowned Manakins, however, were highly mobile and were recaptured rarely. Although we cannot conclusively show that fragmentation caused birds to be in poorer physiological condition, the data indicate that birds in poorer physiological condition were more likely to be captured in fragments than in continuous forest. Thus, our data suggest that forest fragmentation may have subtle but important effects on species that are relatively common after landscape alteration.     

The effects of parasites on male ornaments and female choice in the lek-breeding black grouse (Tetrao tetrix)

Summary We describe the results of two studies of parasitic infection in the black grouse (Tetrao tetrix). The first deals with our own observations of lekking black grouse in which the parasite levels of two blood parasites, the protozoan Leucocytozoon lovati and microfilaria, probably produced by a nematode worm Splendidofilaria tuvensis, were scored. We also obtained measures of age, survival, number of copulations, body mass and length of the ornamental tail feathers (the lyre) of the lekking males. The second study analysed the data from Lund (1954) which involved eight gut parasites obtained from birds which were killed. In the first study we found higher levels of infection of Leucocytozoon in adults relative to young birds. Parasites had no effect on male survival and there was no correlation in infection between the two types of parasites. Birds infected with microfilaria had shorter tail ornaments. There was no relationship between parasitic infection and mating success. However, the data indicated that such a trend indeed may exist for Leucocytozoon and the most successful males on the leks were less often infected by Leucocytozoon than other males. Results of the second study showed a negative relationship between parasite load (a combined measure of all parasites) and both ornamental tail feather length and body mass. These observations are compatible with, but not conclusive evidence for, the hypothesis of Hamilton and Zuk (1982) on the evolution of secondary sexual characters, where females choose to mate with genetically resistant males which show their resistance by expressing larger and more showy secondary sexual characters. Alternative explanations for the observed patterns are: females avoid infected males for some immediate benefit; and/or parasite loads are indicators of general stress rather than genetical resistance. Under the latter hypothesis females could mate with more vigorous males for reasons unrelated to parasite resistance.     

Schwermetalle in Federn

Results from the literature concerning feathers as a matrix for heavy metal accumulation were presented. Especially problems like the differentiation between endo and exogenous contributions, cleaning procedures or the right choice of feather parts were described. The question of the use of feathers for monitoring air, food or bird contamination is also discussed, as well as the advantages or disadvantages of this matrix in comparison to other methods.     

The use of incubation behavior to adjust avian reproductive costs after egg laying

Reproduction in birds requires the input of time and energy during discrete breeding phases leading to investment trade-offs between laying date, clutch size, body mass, and incubation constancy. We investigated costs during incubation by experimentally enlarging 25 clutches of white-tailed ptarmigan Lagopus leucurus. The experiment was conducted in 2 years, one with harsh weather that forced a natural delay in reproduction. When forced to delay egg-laying, females began incubation with poorer body condition and foraged more during incubation. Rates of mass loss during incubation were not affected by clutch enlargement, and did not differ between harsh and benign years; however, females that were heavier at the start of incubation lost more mass than lighter females. Clutch-enlarged females had reduced nest attendance compared to control birds in both years and incubation periods increased by up to 2 days relative to controls. In the harsh year, there was a trend for clutch-enlarged females to have lower nest success, but there was no effect on overwinter survival. Different behavioral responses by females in the 2 years showed that incubation costs may depend on other factors such as female quality, food supply, or weather conditions. Incubation is a dynamic period during which birds may adjust energy balances by varying body condition and food intake. Received: 28 March 2000 / Received in revised form: 18 August 2000 / Accepted: 2 July 2000     

Using carbon and nitrogen isotopic values of body feathers to infer inter- and intra-individual variations of seabird feeding ecology during moult

To determine whether stable isotope measurements of body feathers can be used to investigate the isotopic niche of moulting (inter-nesting) adult seabirds, we examined the stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopic composition of body feathers of breeding wandering albatrosses (Diomedea exulans) from Crozet Islands, southern Indian Ocean. First we showed that the isotopic composition of body feathers was not significantly different from that of wing feathers, being thus a safe alternative to flight feathers whose collection impairs the birds’ flying ability. Second, we looked at the variances in δ¹³C and δ¹⁵N values resulting from the isotopic measurement of a single feather, four different feathers, and a pool of four feathers per bird, to delineate the best isotopic analytical procedure. A two-step protocol is proposed that allows investigating both the intra- and inter-individual components of the niche width of the species. In a first step, isotopic measurements on a single feather per bird are used to define isotopic specialist from isotopic generalist populations. In a second step and for generalist populations only, measurements on additional (three) feathers per bird are used to delineate type A from type B isotopic generalists (Bearhop et al. in J Anim Ecol 73:1007–1012, 2004). Third, from a biological point of view, our data showed different moulting isotopic niches for adult males and females, and also within female wandering albatrosses. Since the isotopic composition of body feathers in this species reflects that of wing feathers, our results suggest that, after validation, body feathers have the potential for investigating the foraging ecology of other Procellariiforms and seabirds during the poorly known inter-nesting period.     

Assessing the aerodynamic effects of tail elongations in the house martin (Delichon urbica): implications for the initial selection pressures in hirundines

Of the three species of hirundine that breed sympatrically across the U.K., one, the barn swallow, has outer tail feathers elongated into streamers, whereas the other two species, the house martin and the sand martin, do not. The tail streamer of the barn swallow is regarded as a classic example of a sexually selected trait. Recent evidence, however, has suggested that streamers may have evolved largely through natural selection for enhanced flight performance and increased maneuverability. We tested the hypotheses that small streamers (1) increase performance in turning flight, but (2) decrease performance in flight variables related to velocity. We manipulated the lengths of house martin outer tail feathers and measured changes in their free-flight performance, using stereo-video to reconstruct the birds" three-dimensional flight paths. Five flight variables were found to best describe individual variation in flight performance. Of these five, the three variables determining maneuverability predicted that flight performance would be optimized by a 6- to 10-mm increase in the length of the outer tail feathers. In contrast, for mean velocity and mean acceleration, extension of the outer tail feathers appears to have a detrimental effect on flight performance. We suggest that the initial selection pressure for streamers in ancestral short-tailed "barn swallows" was via natural selection for increased maneuverability. In addition, we propose that the benefits of increased maneuverability have differed between hirundines in the past, such that the cost of increasing the length of the outer tail feather has, to date, outweighed the benefits of doing so in streamerless hirundines. Received: 14 February 2000 / Revised: 2 July 2000 / Accepted 18 July 2000