Inter- and intra-sexual patterns of fluctuating asymmetry in the red-billed streamertail: should symmetry always increase with ornament size?

Recent work on fluctuating asymmetry has suggested that ornaments should have higher levels of fluctuating asymmetry than (1) non-ornaments and (2) homologous structures in the non-ornamented sex. In addition, as both ornament size and symmetry should increase with individual quality there should be a tendency for ornament symmetry to increase with ornament size. In non-ornaments, a U-shaped relationship between symmetry and size is expected, with the individuals at the extremes being more asymmetrical than individuals around the optimum. We tested these predictions in the red-billed streamertail (Trochilus polytmus), a sexually dimorphic endemic Jamaican hummingbird. The lengths of four bilaterally symmetrical traits (first and second outermost tail feathers, tarsi and wings) in 43 adult males and 42 females were measured. The second outermost tail feathers of adult males (which are elongated into streamers) were absolutely but not relatively more asymmetrical than non-ornaments (including the homologous feathers in females). When character size was controlled for, wings were shown to be relatively more symmetrical than other traits. Symmetry did not increase with increasing trait size in any of the morphological traits measured. There was a U-shaped relationship between asymmetry and trait size for four traits (adult male streamers, adult male wings and female outer tail feathers). These results do not support any of the predictions made by fluctuating asymmetry hypotheses and suggest that stabilising selection may act on ornaments as well as non-ornaments. These predictions have been supported in swallows and peafowl but not in sunbirds; this may be due to differences in female perception of tail ornaments. Perhaps male tails do not convey information about quality in some species, or there may be inter-specific differences in the relative costs of tail ornaments and the benefit of marginal increases in tail length and symmetry.     

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

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

Male asymmetry and postcopulatory sexual selection in the fly Dryomyza anilis

The significance of male asymmetry in postcopulatory sexual selection was studied in the fly Dryomyza anilis by examining whether male asymmetry is related to fertilization success. The traits measured were wing length, tibia length and the length of small and large claspers. The male claspers are situated at the tip of the abdomen, functional pair of claspers consists of a large and a small clasper on the same side of the body. These claspers are used to tap the female abdomen after sperm transfer, which has been shown to increase fertilization success for the mating male. Fertilization success was negatively related to the fluctuating asymmetry of wing length, suggesting either female preference for more symmetrical males or a relationship between male asymmetry and intrasexual selection which was reflected in mating performance. Fertilization success was also related to the length of small claspers, decreasing with increasing length of the claspers. In addition, males with asymmetrical small claspers enjoyed higher fertilization success than symmetrical ones. This study shows that fluctuating asymmetry in wing length is an important fitness trait in postcopulatory sexual selection. Since male tapping affects sperm distribution in the female's sperm storage organs, the higher fertilization success of males with asymmetrical small claspers could have a functional relationship with the asymmetrical position of female sperm storage organs. Received: 6 March 1997 / Accepted after revision: 8 November 1997     

Sexual selection, feather breakage and parasites: the importance of white spots in the tail of the barn swallow (Hirundo rustica)

The long outermost tail feathers of barn swallows Hirundo rustica have white spots that are larger in males than in females and in adults than in juveniles. Spot size increases with age among adults and is positively correlated with tail length. We tested the functional significance of these white spots by randomly assigning males to either of three groups during spring: (1) subjected to a considerable reduction in the size of all spots using a black permanent pen; (2) subjected to a small reduction in the size of all spots; or (3) a control group with no reduction. Experimental treatment significantly affected the total number of offspring produced per season: breeding tended to be delayed and second broods were less frequent after a considerable reduction in the size of the males' tail feather spots. White parts of feathers lacking melanin were more often found to be the site of feather breakage than the melanised parts of similar width. Furthermore, white tail spots were the preferred feeding site of feather-eating Mallophaga and, since long-tailed males have fewer Mallophaga than short-tailed males, long-tailed males may pay a lower cost for their large white tail spots than short-tailed males. Hence, the cost of increased risk of feather breakage at white tail spots and the non-random distribution of feather lice among individuals will render tail spots a reliable signal of phenotypic quality. Received: 25 May 1998 / Accepted after revision: 4 January 1998     

Sexual selection and the tail ornaments of North American barn swallows

Summary In this study conducted in southeastern Ontario, Canada, we manipulated the length of outer tail feathers (streamers) of male barn swallows (Hirundo rustica) shortly after they returned to four small breeding colonies in the spring. Both streamers on most males in these colonies were experimentally either lengthened or shortened by 20 mm and as a result 10 randomly chosen males of each category had their streamer lengths manipulated before the fertile period of their mate. Males with elongated streamers had a significantly shorter pre-laying period (from arrival until first egg date) than those with shortened streamers and we interpret this as indicating that they obtained their mates more rapidly. Females mated to elongated males had significantly longer tails and laid their first egg significantly earlier than those mated to shortened males; no other indices of reproductive success differed between these two experimental groups. These results provide some support for previous work suggesting that females prefer males with longer streamers and thus that streamer length in this species is under the influence of sexual selection. We also found that males with longer natural streamers were significantly more likely to break them and that males were much more likely to break their streamers than were females. Since males participate in incubation in North America but not in Europe, we suggest that the aerodynamic disadvantages of streamer breakage that result from incubation attentiveness are at least partly responsible for the shorter streamer length of males in North America. Offprint requests to: R. Montgomerie     

Patterns of fluctuating asymmetry in beetle horns: an experimental examination of the honest signalling hypothesis

Recent theoretical arguments have claimed that negative relationships between the size and symmetry of secondary sexual traits are indicative of honest signalling of male quality. The patterns of fluctuating asymmetry in beetle horns have been proposed to support the honest signalling hypothesis. Here we examine three assumptions of the hypothesis, (1) that traits are costly to produce; (2) the levels of fluctuating asymmetry are indicative of stress imposed during development; and (3) that males with larger traits should have more symmetrical traits, using the horned beetle, Onthophagus taurus. Experimental manipulations of brood mass were used to manipulate horn size and asymmetry. The development of horns was found to be environmentally determined and costly in terms of delayed development and increased risk of pre-adult mortality. Decreasing resource availability increased relative horn asymmetry. However, horn height was positively related to absolute horn asymmetry. While the results do support the hypothesis that sexual selection on secondary sexual traits should increase levels of fluctuating asymmetry, they provide no support for the notion that the patterns of asymmetry honestly signal male quality. Horns are used in disputes between males and may be indicative of male parental investment. Thus, we conclude that while horn size may be an indication of male quality, the patterns of fluctuating asymmetry are not. Received: 16 December 1996 / Accepted after revision: 5 May 1997     

Achromatic color variation in black-capped chickadees,Poecile atricapilla: black and white signals of sex and rank

Sexual dichromatism and phenotypic variation in elaborate male traits are common products of sexual selection. The spectral properties of carotenoid and structurally-based plumage colors and the patch sizes of melanin-based plumage colors have received considerable attention as sexual signals in birds. However, the importance of variation in achromatic plumage colors (white, gray and black) remains virtually unexplored, despite their widespread occurrence. We investigated a potential signal function of the achromatic black and white plumage of black-capped chickadees (Poecile atricapilla). We captured and color-banded 178 free-living chickadees and assessed winter flock dominance hierarchies by tabulating pairwise interactions at feeders. We recaptured 73 of these birds and measured plumage coloration for six body regions using a reflectance spectrometer and the area of melanin-based plumage patches from standardized photographs. We found extensive individual variation in chickadee plumage traits and considerable sexual dichromatism. Male black-capped chickadees have significantly brighter white plumage than females, larger black patches, and greater plumage contrast between adjacent white and black plumage regions. We also found rank differences in the plumage reflectance of males; high-ranking males, who are preferred by females as both social and extra-pair partners, exhibit significantly darker black plumage and grow their feathers more rapidly than low-ranking males. This variation among individuals reveals a potential signal function for achromatic plumage coloration in birds.     

Sexual selection favours small and symmetric males in the polygynous greater sac-winged bat Saccopteryx bilineata (Emballonuridae, Chiroptera)

We investigated how morphological traits of territorial males in the polygynous bat Saccopteryx bilineata were related to their reproductive success. Because of the frequency of aerial courtship displays and defence manoeuvres, and the high energetic costs of flight, we expected small and symmetric males to be better able to court females on the wing and to monopolize copulations with females in their harems. We predicted that small and symmetric males would sire more offspring within the colony and a larger portion of the young born within their harem than large or asymmetric males. We measured size and fluctuating asymmetry of 21 territorial males and analysed their reproductive success in 6 offspring cohorts (n=209 juveniles) using 11 microsatellite loci. As predicted, small and symmetric males had, on average, a higher reproductive success in the colony than large and asymmetric males. The percentage of young sired by males within their harem increased as males decreased in size, but was not influenced by fluctuating asymmetry. As fluctuating asymmetry of males correlated with their reproductive success within the colony but not within their harems, we infer that fluctuating asymmetry is probably related to female choice, whereas male size is probably important for harem defence on the wing.Communicated by G. Wilkinson     

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.     

Sexual ornamentation,condition, and immune defence in the house sparrow Passer domesticus

Female mate preferences for extreme male ornaments may arise and be maintained because males signal their ability to raise an immune defence against parasites by the size of their ornaments. The bursa of Fabricius is an organ involved in antibody synthesis in young birds, but regresses before sexual maturity. The relationship between badge size, condition, and immune defence was investigated in male house sparrows, Passer domesticus, which have a bib of black feathers under their beak, a secondary sexual character currently subject to a directional female mate preference. The size of the bursa of Fabricius decreased during the first year of life in house sparrows. Males with large secondary sexual characters had a smaller bursa than males with small badges. Individuals with a large directional asymmetry in relative testis size had a larger bursa. The abundance of one parasite (a mallophagan) was positively related to bursa size. There was a positive relationship between relative volume of bursa of Fabricius and the number of fault bars on feathers, which is a measure of body condition. These results suggest that males with large badges have lower current levels of immune response than males with small badges, and that males in good body condition, as evidenced from the absence of fault bars, had a relatively small bursa of Fabricius. Received: 6 October 1995/Accepted after revision: 2 June 1996     

White tail markings are an indicator of quality and affect mate preference in rock sparrows

In birds, colourful and elaborate feathers are important traits in mate choice. Distinct tail white patches are present in many species of birds, but they remain little studied. Tail markings may indeed have a signal function because in many species males spread the tail offering a good view of these markings to females during courtship behaviour. Here, we investigated whether white tail spots in male rock sparrow, Petronia petronia, play a role in mate choice. In a free-living population of rock sparrows, we found a reduction in white tail spots size as the breeding season progressed due to abrasion, which was expected if tail spots act as a reliable quality indicator (i.e. a handicap). The same reduction was found under captive conditions, and males in worse condition (individuals that lost more weight) abraded a bigger part of white. This suggests that white tail markings are an indicator of male quality. In captivity, we measured female preference for males differing in white patch size in a mate choice experiment. The experimental reduction of the size of the males’ white spots resulted in a lower sexual interest by females. During courtship display, male rock sparrow shows a yellow breast patch (a carotenoid-based, sexually selected ornament) together with the white spots in the tail. The sizes of these two traits are positively correlated, but only the abraded white area in the tail correlates with a surrogate of individual quality (lost of weight). In conclusion, we can assert that the size of the white spots is preferred by female rock sparrows and it is a part of a multiple signal system.     

An experimental study of male adornment in the scarlet-tufted malachite sunbird: II. The role of the elongated tail in mate choice and experimental evidence for a handicap

Summary Scarlet-tufted malachite sunbirds (Neetarinia johnstoni) are endemic to the high altitude zones of East African mountains. On Mount Kenya the males are resident and territorial all year. The females arrive over an extended period at the beginning of the breeding season, after spending the non-breeding season in nomadic flocks at slightly lower elevations. Males are bright iridescent green with elongated central tail feathers, which are displayed during courtship. We examined the role of the tail in mate choice using natural variation and investigated the influence of tail length on male time budgets by experimental manipulation. Territorial males that paired had longer tails than territorial males which remained unpaired. Males with naturally long tails started breeding earlier, resulting in fledglings being produced earlier in the season when the weather was more clement. Birds which bred together in one year were paired together in the next year if they both survived. Thus the male's tail may only be involved as a cue in the formation of new pairs. Males which were already paired were subjected to one of three experimental treatments — having their tail elongated, shortened, or manipulated but kept at the same length. Males with experimentally shortened tails spent more time in flight and hawked for flying insects with a higher efficiency than control males. Both control and elongated tail males reduced the amount of time in flight and had a lower hawking efficiency after manipulation. These results suggest that the long tail of male scarlet-tufted malachite sunbirds is a handicap. The factors influencing the reproductive success of a pair are discussed. Offprint requests to: M.R. Evans     

Racketed tail of the male and female turquoise-browed motmot: male but not female tail length correlates with pairing success,performance, and reproductive success

Both males and females of many avian species maintain elaborate plumage traits, and elaborate monomorphic plumage may convey adaptive benefits to one or both sexes as inter- or intraspecific signals. Both sexes of the turquoise-browed motmot (Eumomota superciliosa) are elaborately plumed with long racket-tipped tail. I investigated whether the racketed tail functions as a sexually selected signal in one or both sexes by testing the predictions that males and/or females with the largest tails have: (1) greater pairing success, (2) greater reproductive performance (clutch-initiation date, clutch size, and hatching success), and (3) greater reproductive success. Yearling males with longer denuded rachises (wires) on the central tail feathers had greater pairing success. In addition, adult males with longer wires paired with females who laid larger clutches, had greater hatching success independent of clutch size, and fledged more young. There was no relationship between female tail plumage and pairing success, reproductive performance, or fledgling success. These results are consistent with the hypothesis that male tail plumage functions as a mate choice or status signal, but that the tail of the female does not function in a sexually selected context. I discuss alternative hypotheses for the evolutionary maintenance of the elaborate female tail plumage.     

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.     

Male tail streamer length predicts fertilization success in the North American barn swallow (Hirundo rustica erythrogaster)

In socially monogamous species, extra-pair paternity has the potential to increase the variance in male reproductive success, thereby affecting the opportunity for sexual selection on male extravagant ornamentation. In the European barn swallow (Hirundo rustica rustica), the tail streamer length is a sexually selected male ornament and an honest indicator of viability. The North American barn swallow (Hirundo rustica erythrogaster) also shows sexual dimorphism in tail streamer length, but whether this trait holds the same signalling function in this subspecies is a controversial issue, and the available literature is presently scarce. Here, we present data on paternity in the North American barn swallow, including a complete sampling of extra-pair sires in four colonies. We analysed how extra-pair paternity affected the variance in male fertilization success and examined whether male tail streamer (i.e. the outermost tail feather) length correlated with fertilization success (n=86 males). Extra-pair paternity constituted 31% of all offspring and significantly increased the variance in male fertilization success. The number of offspring sired by extra-pair males accounted for almost half of the total variance in male fertilization success. Males with naturally long tail streamers had a higher fertilization success than males with shorter tail streamers, and this pattern was mainly caused by a higher extra-pair success for males with long tail streamers. Males with long tail streamers also paired with early breeding females in prime body condition. These results are consistent with the idea that there is directional sexual selection on male tail streamer length, possibly mediated through male extra-pair mating success or the timing of breeding onset.     

Testosterone and sexual signalling in male house sparrows (Passer domesticus)

Female preference for mates with elaborated ornaments has often been explained on the basis that exaggerated secondary sexual traits might reflect individual quality and females might gather direct and indirect benefits in mating with such males. Sexual signals must however also entail costs to be reliable indicators of male quality. Androgens have been suggested as a physiological link between sexual signals and individual quality for several reasons, including their immunosuppressive effect. In this study, we tested two hypotheses linked to the hormonal basis of sexual signal expression. First, we investigated whether testosterone is correlated with the size of the black feather bib on the throat of male house sparrows (Passer domesticus) - a trait involved in intra- and inter-sexual selection. Second, we tested whether testosterone affects the seasonal exposure of the trait. Observational work conducted in 1998 showed that the testosterone level was positively correlated with badge size both in spring and in the subsequent fall, after molt. In 1999, we experimentally reduced spring testosterone levels using silastic implants filled with cyproterone acetate, an antiandrogen. Male house sparrows implanted with cyproterone acetate showed reduced exposure of the badge, because the white tips of the black feathers of the badge wore off later than in control males implanted with empty silastic tubes. This result suggests that testosterone can be causally involved in the expression of a secondary sexual trait in house sparrows, at least in terms of its seasonal exposure.     

Tail size and female choice in the guppy (Poecilia reticulata)

Summary Under laboratory conditions, female guppies demonstrate a clear preference for males with larger tails, and this preference translates into enhanced reproductive fitness for these males. Females also prefer males with higher display rates, a behavior which appears to be linked to tail size, but which can be experimentally disassociated. This appears to be a case of female-choice sexual selection.     

Testosterone stress does not increase asymmetry of a hormonally mediated sexual ornament in a lizard

The theory that fluctuating asymmetry is sensitive to both environmental and genetic stress is gaining acceptance among evolutionary biologists. Most empirical work has focused on ornamental traits on the assumption that they are more susceptible to stress. In tegumentary coloration is a common ornament in nature, and frequently has a hormonal basis. Earlier studies in the lizard Psammodromus algirus indicate that testosterone induces the development of head nuptial coloration in large males and, at the same time, produces an increase of the ectoparasite load and higher mortality. Hence, the manipulation of testosterone levels may be a way to increase ornament expression and simultaneously create conditions that may make symmetrical development difficult. This positive covariation between character size and symmetry is opposite to that expected in theory for sexually selected traits, so the predicted elevation of asymmetry due to the treatment cannot be confounded by any intrinsic association between symmetry and character size. We firstly consider the effect that testosterone supplementation has on two variables that reflect the symmetry of bilateral throat nuptial coloration in large P. algirus males. Also, we examine whether ornament symmetry is positively associated with reproductive success, a prediction of theory of sexually selected symmetry. Testosterone treatment did not increase the fluctuating asymmetry of throat coloration. Size asymmetry increased with character size in individuals with a fragmented colour pattern, but changed suddenly to a highly symmetric pattern in individuals with non-fragmented coloration. Mirror asymmetry decreased steadily with character size. These results suggest that the development of coloration on both sides of the throat midline follows a random pattern. Asymmetry did not correlate with variables that estimated reproductive success, suggesting that asymmetry is not affected by physiological stress and that this trait is not a sexually selected signal in P. algirus. Received: 1 July 1996 / Accepted after revision: 4 May 1997     

Sexual selection in a monomorphic lek-breeding bird: correlates of male mating success in the great snipe Gallinago media

Summary In the lek-breeding great snipe, male morphology, behaviour, and territory features were recorded for individually marked birds on two adjacent leks. Partial correlation sshowed that male mating success, expressed as the number of female solicitations and copulations, was negatively correlated with the distance of a display territory to the lek center and positively correlated with the number of displays per unit time given by a male. No other variables were directly correlated with male matin success. Thus, central males obtain more matings than peripheral males and successful males display more per unit time than do less successful males, independently of position on the lek. Central males were found to be older than peripheral ones and were present more often on the lek. Furthermore, central males had a larger number of white tail feathers, which are usea as visual signals in the displays, but this may be explained by the fact that these males were older. It is suggested that male great snipe are subject to sexual selection mainly in behavioural and vocal cues and that this may explain the absence of size and plumage dimorphism in this species.     

Age before beauty? Relationships between fertilization success and age-dependent ornaments in barn swallows

When males become more ornamented and reproduce more successfully as they grow older, phenotypic correlations between ornament exaggeration and reproductive success can be confounded with age effects in cross-sectional studies, and thus say relatively little about sexual selection on these traits. This is exemplified here in a correlative study of male fertilization success in a large colony of American barn swallows (Hirundo rustica erythrogaster). Previous studies of this species have indicated that two sexually dimorphic traits, tail length and ventral plumage coloration, are positively correlated with male fertilization success, and a mechanism of sexual selection by female choice has been invoked. However, these studies did not control for potential age-related variation in trait expression. Here, we show that male fertilization success was positively correlated with male tail length but not with plumage coloration. We also show that 1-year-old males had shorter tails and lower fertilization success than older males. This age effect accounted for much of the covariance between tail length and fertilization success. Still, there was a positive relationship between tail length and fertilization success among older males. But as this group consisted of males from different age classes, an age effect may be hidden in this relationship as well. Our data also revealed a longitudinal increase in both tail length and fertilization success for individual males. We argue that age-dependent ornament expression and reproductive performance in males complicate inferences about female preferences and sexual selection.