Outermost Tail Feathers Research Articles

Revisiting an Old Issue: Sex Identification of Short-eared Owls (Asio flammeus) at an Asian Wintering Site

Abstract At wintering sites of the migratory Short-eared Owl (Asio flammeus) in subtropical Asia, behavioral interactions between males and females are limited, and the body mass and plumage features between the sexes overlap significantly. Thus, sexing the species morphometrically for research and conservation activities is difficult. We aimed to develop a quantitative method for sexing Short-eared Owls, and we here present a new formula to do so using plumage features. We used a total of 198 Short-eared Owls (163 from bird rescue efforts at 15 airports and 35 preserved specimens from a museum in Taiwan) and carefully examined their morphological data and plumage images, then determined their sex using polymerase chain reactions of chromodomain helicase DNA-binding protein genes on sex chromosomes or gonad inspection. Eight of the ten traits we evaluated differed significantly between males and females. Females had a greater number of cross bars on primaries, secondaries, and outermost tail feathers; a higher proportion of yellowish-brown underwing coverage; and larger measurements in head length, bill length, tarsus length, and body mass. The best-fit model suggested simplifying the formula to just the proportion of the yellowish-brown underwing coverage, which provided sexing accuracy exceeding 95.9% for the 49 live individuals in the test data set and 94.3% for the 35 preserved specimens. This formula addresses the challenges posed by ambiguous individuals in the wintering region and offers an efficient and accurate means for sexing Short-eared Owls when DNA or gonad inspection is unavailable.

Feather growth rate and hormone deposition vary with elevation but not reproductive costs in resident Mountain Chickadees

Abstract Many organisms engage in metabolic tradeoffs to manage costs associated with reproductive output which often leads to these costs carrying over into the future. Compensatory mechanisms vary across life history strategies and are expected to result in near-optimal fitness gains for the investor. Here we investigated whether environmental differences associated with increasing montane elevation and variation in reproductive output of a resident passerine songbird, the Mountain Chickadee (Poecile gambeli), were related to physiological conditions during annual molt. Higher elevations are associated with harsher environmental conditions during the winter, which results in later and shorter breeding seasons than at lower elevations. We sampled the outermost tail feathers from adult birds in the fall after their prebasic molt, which initiates closely after reproduction (e.g., after parental care has ceased, ~1–3 weeks). We measured the hormone corticosterone deposited in feathers (fCORT) and feather growth rates for evidence of physiological effort predicted to be driven by several units of reproductive output (e.g., breeding timing, clutch and brood size, and offspring mass). There were no relationships between any measure of reproductive output and feather characteristics between elevations or across years, despite substantial variation in reproductive output in the wider population across this same time. However, birds at the high-elevation site grew their tail feathers significantly faster and had higher fCORT deposition compared to low-elevation birds. These results suggest that although differences in reproductive output and any related signals of associated physiological effort (e.g., fCORT and feather growth rate) may not extend into individual conditions during annual molt, shorter breeding seasons associated with harsher environmental conditions may favor faster feather growth as required by earlier onset of winter.

Compensatory traits can explain the concave cost function of purely sexual traits.

The cost of ornamentation is often measured experimentally to study the relative importance of sexual and viability selection for ornamentation, but these experiments can lead to a misleading conclusion when compensatory trait is ignored. For example, a classic experiment on the outermost tail feathers in the barn swallow Hirundo rustica explains that the concave (or U-shaped) aerodynamic performance cost of the outermost tail feathers would be the evolutionary outcome through viability selection for optimal tail length, but this conclusion depends on the assumption that compensatory traits do not cause reduced performance. Using a simple "toy model" experiment, I demonstrated that ornamentation evolved purely though sexual selection can produce a concave cost function under the presence of compensatory traits, which was further reinforced by a simple mathematical model. Therefore, concave cost function (and the low performance of individuals with reduced ornaments) cannot be used to infer the evolutionary force favoring ornamentation, due to a previously overlooked concept, "overcompensation," which can worsen the whole body performance.

When Older Males Sire More Offspring—Increased Attractiveness or Higher Fertility?

In birds with extrapair mating, older males usually have higher fertilization success than younger males. Two hypotheses can potentially explain this pattern: 1) females prefer older, and often more ornamented males, or 2) older males invest more in reproduction and fertility than younger males. Here we studied factors associated with age-related male fertilization success in a population of barn swallows Hirundo rustica in Canada. We document that male fertilization success increased gradually up to a minimum age of four-year old. The age effect was especially strong for the number of extrapair offspring obtained and the occurrence of a second brood. The higher fertilization success of older males was also associated with an early start of breeding in spring. The length of the elongated outermost tail feathers, a postulated male ornament preferred by females, also increased with age (in both sexes), but it was not a significant predictor of male fertilization success within age classes. Male fertility traits, especially testis size, but also sperm motility and sperm velocity, increased significantly across age groups. Our results suggest that the higher fertilization success by older males is due to their higher reproductive investments and that their longer tails are an adaptation to early arrival on the breeding grounds.Significance statementThe barn swallow is a socially monogamous passerine with extensive extrapair mating. We found that males become more successful in siring both withinpair and extrapair offspring as they become older. Their increased fertilization success was associated with a higher reproductive effort as indicated by larger testes, more motile sperm, and an earlier start of breeding in spring. The length of the outer tail feathers increased with age in both sexes, but long tails did not enhance male fertilization success among males of the same age. Long tails are probably an adaptation to rapid migration and earlier arrival on the breeding grounds. Our findings suggest that the commonly observed age-related increase in male fertilization success in passerine birds is better explained by life history theory than by sexual selection theory.

Telomere length is highly repeatable and shorter in individuals with more elaborate sexual ornamentation in a short-lived passerine.

Quantifying an individual's state as a fitness proxy has proven challenging, but accumulating evidence suggests that telomere length and attrition may indicate individual somatic state and success at self-maintenance, respectively. Sexual ornamentation is also thought to signal phenotypic quality, but links between telomeres and sexual ornamentation have been little explored. To address this issue, we examined whether telomere length and dynamics are predicted by the expression of a sexually selected ornament, the length of the outermost tail feathers (streamers), using longitudinal data from a population of European barn swallows (Hirundo rustica). In 139 adult individuals, each measured twice, we further assessed associations of telomere length with age, sex, breeding status and survival. Telomere length showed high individual repeatability (R=.97) across years while shortening with age in both sexes. Telomere length and dynamics were not significantly associated with survival to the next year, remaining lifespan or reproduction status (comparing breeding and nonbreeding yearlings). Tail streamer length, a sexually selected trait in barn swallows, was negatively associated with telomere length, independent of sex. Thus, telomere length may reflect the costs of carrying an elaborated sexual ornament, although ornament size did not significantly predict telomere shortening. In conclusion, telomere length in adult barn swallows is a highly consistent trait that shows a negative relationship with sexual ornamentation, suggesting a trade-off between sexual ornamentation and telomere length.

Experimental tail shortening affects feeding rate depending on original tail length in female barn swallows Hirundo rustica gutturalis

Long tail feathers of the barn swallow Hirundo rustica are a classic example of an intersexually selected trait, but previous aerodynamic analyses indicate that the tail feather is only 10–12 mm longer than the aerodynamic optimum even in the nominate subspecies with long tails. Here, by experimentally shortening female tail length, we studied the feeding cost of long tail feathers in Japanese barn swallows, Hirundo rustica gutturalis, which have ca. 10 mm shorter tails than the nominate subspecies. Female feeding rate was explained by the interaction between treatment and original female tail length: feeding rate decreased with decreasing original female tail length in control, but not in tail-shortened females. Because the interaction term was far from significant in the analysis of female incubation investment, the observed pattern would be specific to feeding rate, which is greatly affected by the aerodynamic properties associated with tail length. Differential allocation of paternal feeding investment was not observed in the current data set. Long tails would be costly at least in short-tailed females, supporting differential costs of ornamentation as predicted by sexual selection theory. Female outermost tail feathers are costly ornamentation in short-tailed Japanese barn swallows.

Migration pattern of Yellow-throated buntings revealed by isotope-based geographic assignment

ABSTRACTMigratory birds have evolved diverse migration strategies in response to a variety of factors, but information about the detailed migration patterns of Asian songbirds is not yet available. To understand the short-distance migration pattern of declining Yellow-throated Buntings (Emberiza elegans) in East Asia, we analyzed stable isotopes from the outermost tail feathers (rectrices) of individual buntings collected in Korea and Japan. Temporal changes in feather hydrogen (δ2Hf) and oxygen (δ18Of) isotopic values at stopover islands suggested that northern populations start migration earlier than southern populations, especially in autumn. Latitudinal gradient in δ2Hf values of three wintering populations implied that northern breeders wintered farther north than southern breeders. The migration pattern of this bunting, known as Type II chain migration, was also inferred from hydrogen isotope-based geographic assignments of feather growth origins. Our data demonstrate that stable isotope analysis may help to bridge current knowledge gaps in songbird migration despite coarsely mapped isoscapes and as-of-yet undetermined isotope calibration functions in Asia.

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.

Variation in melanin pigmentation of a sexually selected plumage trait and its adaptive value in the Common SnipeGallinago gallinago

There is increasing evidence that melanin‐based plumage coloration correlates with different components of fitness and that it may act as a social or sexual signal of individual quality. We analysed variation in melanin pigmentation in the outermost tail feathers of the Common SnipeGallinago gallinago. During courtship flights, male Snipe use their outermost tail feathers to generate a drumming sound, which plays a role in territory establishment and mate choice. As the outermost tail feathers are displayed to females during these flights, we predicted that conspicuous variation in their rusty‐brown (pheomelanin‐based) coloration may act as an honest signal of individual quality. To test this prediction, we spectrophotometrically measured brightness (an indicator of total melanin content) and red chroma (an indicator of pheomelanin content) of the outermost tail feathers in 180 juvenile and adult Common Snipe. An age‐related decline in feather brightness was found exclusively in females, suggesting that melanization could have evolved by natural selection to camouflage incubating birds. In both sexes, brightness of the tail feathers was inversely correlated with their structural quality (as measured with mass–length residuals), suggesting that melanization could increase mechanical properties of feathers and, in males, enhance the quality of courtship sonation. Red chroma positively correlated with total plasma protein concentration, supporting our prediction that pheomelanin pigmentation of tail feathers may act as an honest signal of condition. Our study indicated that variation in the melanin‐based coloration of the outermost tail feathers in the Common Snipe could have evolved as a result of several different selection pressures and it emphasizes the complexity of the processes that underlie the evolution of melanin‐based plumage coloration in birds.

The maintenance of phenotypic divergence through sexual selection: An experimental study in barn swallows Hirundo rustica.

Previous studies have shown that sexual signals can rapidly diverge among closely related species. However, we lack experimental studies to demonstrate that differences in trait-associated reproductive performance maintain sexual trait differences between closely related populations, in support for a role of sexual selection in speciation. Populations of Northern Hemisphere distributed barn swallows Hirundo rustica are closely related, yet differ in two plumage-based traits: ventral color and length of the outermost tail feathers (streamers). Here we provide experimental evidence that manipulations of these traits result in different reproductive consequences in two subspecies of barn swallow: (H. r. erythrogaster in North America and H. r. transitiva in the East Mediterranean). Experimental results in Colorado, USA, demonstrate that males with (1) darkened ventral coloration and (2) shortened streamers gained paternity between two successive reproductive bouts. In contrast, exaggeration of both traits improved reproductive performance within H. r. transitiva in Israel: males with a combination treatment of darkened ventral coloration and elongated streamers gained paternity between two successive reproductive bouts. Collectively, these experimental results fill an important gap in our understanding for how divergent sexual selection maintains phenotype differentiation in closely related populations, an important aspect of the speciation process.

Sexual Dimorphism and Population Differences in Structural Properties of Barn Swallow (Hirundo rustica) Wing and Tail Feathers.

Sexual selection and aerodynamic forces affecting structural properties of the flight feathers of birds are poorly understood. Here, we compared the structural features of the innermost primary wing feather (P1) and the sexually dimorphic outermost (Ta6) and monomorphic second outermost (Ta5) tail feathers of barn swallows (Hirundo rustica) from a Romanian population to investigate how sexual selection and resistance to aerodynamic forces affect structural differences among these feathers. Furthermore, we compared structural properties of Ta6 of barn swallows from six European populations. Finally, we determined the relationship between feather growth bars width (GBW) and the structural properties of tail feathers. The structure of P1 indicates strong resistance against aerodynamic forces, while the narrow rachis, low vane density and low bending stiffness of tail feathers suggest reduced resistance against airflow. The highly elongated Ta6 is characterized by structural modifications such as large rachis width and increased barbule density in relation to the less elongated Ta5, which can be explained by increased length and/or high aerodynamic forces acting at the leading tail edge. However, these changes in Ta6 structure do not allow for full compensation of elongation, as reflected by the reduced bending stiffness of Ta6. Ta6 elongation in males resulted in feathers with reduced resistance, as shown by the low barb density and reduced bending stiffness compared to females. The inconsistency in sexual dimorphism and in change in quality traits of Ta6 among six European populations shows that multiple factors may contribute to shaping population differences. In general, the difference in quality traits between tail feathers cannot be explained by the GBW of feathers. Our results show that the material and structural properties of wing and tail feathers of barn swallows change as a result of aerodynamic forces and sexual selection, although the result of these changes can be contrasting.

Intensity of Melanin-Based Color and Risk of Predation in the Barn SwallowHirundo rustica

Predators are supposed to exert strong selection pressures on their prey, especially when phenotypic traits such as secondary sexual characters promote mating success at the expense of costs in terms of natural selection. Signaling theory predicts that individuals of superior phenotypic quality will enjoy an advantage in term of mating success, but also in term of natural selection, if such individuals are in prime condition both before and after development of exaggerated secondary sexual characters. We tested this prediction in the Barn Swallow Hirundo rustica being preyed upon by the Eurasian Sparrowhawk Accipiter nisus, using extensive samples of feathers from prey and non-prey. We measured tail length and coloration of outermost tail feathers in the black area of the proximal and distal part of tail feathers, but also the white spot of the tail feathers. Prey had significantly less dark distal, but not proximal parts of their tails, while there was no difference in coloration of the white spot between prey and non-prey. Prey had significantly paler tail feathers than non-prey, especially among long-tailed individuals. These results suggest that Barn Swallows with long tails that fail to deposit large amounts of melanin in their tail feathers run an elevated risk of predation.

Effects of experimental tail shortening on the phenotypic condition of barn swallowsHirundo rustica: implications for tail‐length evolution

Some studies have suggested that tail streamers in the barn swallowHirundo rusticamay have been elongated 10–12 mm by sexual selection, but according to other studies, the length of these feathers is at the aerodynamic optimum or very close to it. To shed light on this issue, outermost tail feathers were experimentally shortened in male and female barn swallows by 1, 11 or 21 mm. Changes in four physiological parameters commonly used to estimate phenotypic condition in birds (weight, erythrocyte sedimentation rate, blood leukocyte concentration and heterophil/lymphocyte ratio) were checked one month later. Health improved (blood leukocyte concentration decreased) in the group of birds with tails shortened by 11 mm (both males and females), but body condition deteriorated (weight decreased) compared to the other two experimental groups. There was no significant effect of tail‐length manipulation on the other two physiological parameters. These contradictory results suggest trade‐offs between components of phenotypic condition. Possible negative relationships between condition‐related traits imply that using one or very few physiological parameters to estimate phenotypic condition might not be appropriate. The most plausible explanation for the turning point in phenotypic condition when streamers were shortened by 11 mm is that these feathers are 7–15 mm longer than the aerodynamic optimum in both sexes. Therefore, our results are consistent with the hypothesis that tail streamers have been elongated 10–12 mm by sexual selection. This conclusion disagrees with a previous study on the effect of experimental tail shortening on haematocrit, but the complexity of interpreting changes in haematocrit might account for this discrepancy.

Relationship between sexual signals and louse (Insecta: Phthiraptera) infestation of breeding and migrating Barn swallows (Hirundo rustica) in Hungary

Abstract Lice (Phthiraptera) chew characteristic holes on the remiges and rectrices of Barn swallows (Hirundo rustica). The number of these holes correlate positively with the intensity of louse infestation, hence hole counts are useful to quantify lousiness. Several papers showed that lice affect both life expectancy and reproductive success of hosts. In male Barn swallows, the length of the outermost tail feathers act as a sexual signal. Females prefer long-tailed males, which have significantly fewer feather holes. In this study we sampled breeding and migrating Barn swallows and compared their louse burden, and the relationship between tail length and the number of feather holes. We found significant negative correlation between feather holes and tail length in breeding males; however, we found non-significant correlation in migrating males. We suggest that attractive males have more physical interactions (e.g. extra-pair copulation) during the breeding season, than less attractive males, hence they are more exposed to louse transmission, and therefore the difference in the infestation declines towards the end of the breeding season. However, given that migrating swallow groups include colonial and solitary breeding birds, it cannot be excluded that a potentially different louse distribution on solitary breeding birds may contribute to the results.

Population differences in density and resource allocation of ornamental tail feathers in the barn swallow

Many organisms show well-defined latitudinal clines in morphology, which appear to be caused by spatially varying natural selection, resulting in different optimal phenotypes in each location. Such spatial variability raises an interesting question, with different prospects for the action of sexual selection on characters that have a dual purpose, such as locomotion and sexual attraction. The outermost tail feathers of barn swallows (Hirundo rustica) represent one such character, and their evolution has been a classic model subject to intense debate. In the present study, we examined individuals from four European populations to analyze geographical variation in the length and mass of tail feathers in relation to body size and wing size. Tail feather length differed between sexes and populations, and such variation was a result of the effects of natural selection, acting through differences in body size and wing size, as well as the effects of sexual selection that favours longer tails. The extra enlargement of the tail promoted by sexual selection (i.e. beyond the natural selection optimum) could be achieved by increasing investment in ornaments, and by modifying feather structure to produce longer feathers of lower density. These two separate processes accounting for the production of longer and more costly tail feathers and less dense feathers, respectively, are consistent with the hypothesis that both Zahavian and Fisherian mechanisms may be involved in the evolution of the long tails of male barn swallows. We hypothesize that the strength of sexual selection increases with latitude because of the need for rapid mating as a result of the short duration of the breeding season at high latitudes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105, 925–936.

Effects of immune activation and glucocorticoid administration on feather growth in greenfinches

Elevation of glucocorticoid (GC) hormone levels is an integral part of stress response (as well as its termination) and immunomodulation. These hormones are also responsible for mobilizing energy stores by stimulation of gluconeogenesis and inhibition of protein synthesis. Elevation of GCs is thus incompatible with other protein-demanding processes, such as moult. Previous studies have shown that chronic elevation of GC hormones suppresses feather growth. Here, we asked whether similar effect would also occur in the case of acute GC elevation and induction of an inflammatory response by foreign antigen. We performed an experiment on captive wild-caught greenfinches (Carduelis chloris) injecting birds with phytohaemagglutinin (PHA) and dexamethasone (DEX) in a factorial design. To assess the possible somatic impacts of these manipulations, we removed one of the outermost tail feathers before the experiment and measured mass and rachis diameter and length of the replacement feathers grown in captivity. Immunostimulation by PHA reduced rachis length, but did not affect feather mass or rachis diameter. Single injection of a synthetic GC hormone DEX significantly reduced all three parameters of feather size. Altogether, these findings demonstrate the sensitivity of feather growth to manipulation of immune and adrenal functions. Our results corroborate the somatic costs of immune activation and suggest that even a short-term elevation of GC hormones may induce long-term somatic costs with a potential impact on fitness. Our findings also imply that a single injection of DEX, frequently used as a diagnostic tool, can have lasting effects and researchers must consider this when designing experiments.

An additional field method to sex adult Barn Swallows during the non-breeding season in Zambia: white spot length in the outer tail feather

Adult Barn Swallows Hirundo rustica exhibit strong sexual size dimorphism in the length of the outermost tail feathers, which are longer in males compared with females. This trait is traditionally used to sex adult Barn Swallows in the field. However, due to the wear and breakage of the tips of the outer tail feather and tail moult during the non-breeding season, sexing becomes unreliable or even impossible. We therefore tested whether the length of the white spot on the outer tail feather is sexually dimorphic, and whether it can be used as an additional sexing method for adult Barn Swallows. The white spot length was sexually dimorphic, based on DNA analysis of 101 adult individuals caught at their roost during the non-breeding season in Zambia. Accuracy in sex determination of 95% could be obtained by classifying individuals with a white spot length <17.5 mm as females and those with a white spot length >29.5 mm as males. When applying the length of the white spot as an additional method to sex adult Barn Swallows on all birds caught in Zambia during the study period (N = 759), the percentage of birds that could successfully be sexed increased to more than 55%. Therefore we emphasise the importance of measuring the white spot length in addition to the tail fork depth and tail length to sex adult Barn Swallows in the non-breeding season.

Quantitative genetics of migration syndromes: a study of two barn swallow populations

Migration is a complex trait although little is known about genetic correlations between traits involved in such migration syndromes. To assess the migratory responses to climate change, we need information on genetic constraints on evolutionary potential of arrival dates in migratory birds. Using two long-term data sets on barn swallows Hirundo rustica (from Spain and Denmark), we show for the first time in wild populations that spring arrival dates are phenotypically and genetically correlated with morphological and life history traits. In the Danish population, length of outermost tail feathers and wing length were negatively genetically correlated with arrival date. In the Spanish population, we found a negative genetic correlation between arrival date and time elapsed between arrival date and laying date, constraining response to selection that favours both early arrival and shorter delays. This results in a decreased rate of adaptation, not because of constraints on arrival date, but constraints on delay before breeding, that is, a trait that can be equally important in the context of climate change.

Fluctuating asymmetry in great tit nestlings in relation to diet quality, calcium availability and pollution exposure

Stress during development may cause fluctuating asymmetry (FA), i.e. non-directional and random deviations from perfect symmetry in otherwise symmetrical morphological traits. These deviations affect the phenotypic quality of an individual. We manipulated the diet of nestling great tits, Parus major, to investigate how food quality and quantity affect FA in the length and mass of the outermost tail feathers of great tit nestlings in a polluted and an unpolluted area. High carotenoid diet groups and the control group had higher FA in tail feather length compared to a mealworm-supplemented (low carotenoid) group. This suggests that high carotenoid content in the diet may either directly or indirectly induce higher FA in tail feather length. Calcium is an essential element for birds and important component of feathers. The less calcium there was in the diet, the higher was the FA in tail feather length, which suggests that calcium availability may be an important determinant of the developmental stability of tail feather length. In the control group, in which nestlings were fully dependent upon natural food resources provided by their parents, FA in feather mass was higher in polluted than in unpolluted sites. Diet quality and quantity seemed to differentially affect FA in tail feather length and mass between the polluted and the unpolluted areas. FA in tail feather length in the control group was unaltered by pollution, while FA in tail feather mass was lower in the control group in the unpolluted than in the polluted area. Our study also demonstrates for the first time that the developmental stability of tail feather length and mass are affected by different factors.

Why Do Calypte Hummingbirds “Sing” with Both Their Tail and Their Syrinx? An Apparent Example of Sexual Sensory Bias

Courtship displays frequently include complex signals that females use to pick a mate. Male Costa's hummingbirds (Calypte costae) generate two acoustic signals during courtship: a vocal song produced close to a female and a dive-sound produced during a courtship dive. The song and dive-sound sound similar, and both were assumed to be produced vocally by the syrinx. Here, we show that they are not; whereas the song is produced by the syrinx, the dive-sound is produced by high-frequency fluttering of the outermost tail feathers. The Anna's hummingbird (Calypte anna), sister to the Costa's, also sings a vocal song and produces a dive-sound with the wings and outermost tail feathers that sounds similar to a portion of the song. The interspecific match in signal form between the two species is not as strong as the intraspecific match. Phylogenetic reconstruction indicates that the dive-sounds may have evolved first, suggesting that the song may have evolved to mimic the dive-sound. We propose the "sexual sensory bias" hypothesis as an explanation for the match in form between the song and the dive-sound within each species, in which we suggest that new sexual signals can arise in response to preexisting female preferences for older sexual signals.