Sexual size dimorphism and sex identification using morphological traits of two Aegithalidae species

The black-throated tit, Aegithalos concinnus, and long-tailed tit, A. caudatus, are two widely-distributed species of Aegithalidae. They are thought to be monomorphic and thus difficult to differentiate between sexes in the field. We determined the sex of 296 black-throated tits and 129 long-tailed tits using DNA analysis, evaluated their sexual size dimorphism, and developed discriminant models to identify sex based on morphometries, including bill length, bill depth, bill-head length, maximum tarsus length, tarsus width, wing length, tail length, total body length, and body mass. Both species were sexually dimorphic in size, with males having larger measurements than females, except for bill length in black-throated tits, and both bill length and body mass in long-tailed tits. Moreover, both species showed similar sexual size dimorphism (SSD) among the morphological features, with tail length having the highest SSD value. The multivariate discriminant models for sex identification had an accuracy of 82% for both species, which was slightly higher than the best univariate discriminatory model for each species. Because of the complicated nature of the multivariate model, we recommend univariate models for sex identification using D = 0.491 × tail length - 24.498 (accuracy 80%) for black-throated tits and D = 0.807 × wing length - 45.934 (accuracy 78%) for long-tailed tits. Females in both species showed generally higher morphological variation than did males, resulting in lower accuracies in all discriminate functions regardless of univariate or multivariate approach. This could be the result of a sex-biased selective pressure in which males have higher selective pressures for these morphological features.     

Adaptive benefits of differential post-fledging development patterns in the Lesser Flamingo (Phoenicopterus minor)

Measurements of five morphological components (mass, skull length, culmen, flattened wing and tarsus) and blood samples were taken from 154 fledged wild Lesser Flamingos Phoenicopterus minor captured during 2001 and 2002 at Lake Bogoria, Kenya (0°11'–20'N, 036°06'E). The sample included adults (>3 years old), immature birds (2–3 years old) and first-year juvenile birds of both sexes. The sex of each bird was determined by PCR amplification of the CHD-Z and CHD-W genes, using DNA extracted from blood samples. Within each gender, there were significant differences in mass and tarsus length amongst the three age groups, indicating that the skeletal size and mass of Lesser Flamingos continue to increase between fledging and attainment of adult plumage at three to four years of age. The different morphological components increased in size at different rates, although the same components appeared to increase at similar rates in both males and females. Skull and culmen lengths had reached adult size in juvenile birds, while juvenile wing length, tarsus length and mass were approximately 95%, 85% and 75% of adult size, respectively. The adaptive significance of these findings is discussed.     

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

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

Sexual size dimorphism and positive assortative mating in red-backed shrike Lanius collurio: an adaptive value?

During field studies in 1997–1999 in South Bohemia (Czech Republic), we found significant differences in size between the sexes in a local breeding population of red-backed shrike Lanius collurio. Males were significantly larger than females for wing length and tarsus length, but had smaller body mass than females. However, there was considerable overlap in the ranges of these parameters between the sexes. Interestingly, pairs were formed at random with respect to wing length and tarsus length, but assortative mating was significant for body mass/body condition. Among tested variables, only male wing length correlated significantly with nestling body mass at day 7. However, clutch size and the number of fledglings strongly depended on differences in tarsus length between mates, but not on body size of mates. Individual improvements in foraging skills and/or courtship feeding rates are proposed as possible explanations for these findings.     

Body weight and some biometrical traits of ring-necked pheasants (Phasianus colchicus) at different ages

Although the growth of pheasants is an important issue, there are few publications providing information on body weight and morphometrics of pheasants at different ages. The aim of this study was to investigate body weight, tarsus length, wing length and wingspan in farmed pheasants aged 3-16 weeks. The study was carried out on 50 pheasants (31 females and 19 males). All one-day old birds were marked and kept until the 16th week of life. In the 3rd 8 h 12th and 16th weeks of life all pheasants were weighted and tarsus length, wing length and wingspan were measured. Male chicks appeared to be significantly heavier than females (P< or = 0.01) already in the third week of life. Also their wings were longer and their wingspan attained higher values (P < or = 0.05). Absolute gains of the majority of traits also differed between sexes (being larger in males) in all studied periods. Values of body weight, tarsus length and wingspan noted in the third week of life were significantly correlated with all later measurements. Thus body weight and some biometrical traits of pheasant chicks measured already at the 3rd week of life could be applied as predictors of future body weight and size.     

Multivariate selection and intersexual genetic constraints in a wild bird population

When selection differs between the sexes for traits that are genetically correlated between the sexes, there is potential for the effect of selection in one sex to be altered by indirect selection in the other sex, a situation commonly referred to as intralocus sexual conflict (ISC). While potentially common, ISC has rarely been studied in wild populations. Here, we studied ISC over a set of morphological traits (wing length, tarsus length, bill depth and bill length) in a wild population of great tits (Parus major) from Wytham Woods, UK. Specifically, we quantified the microevolutionary impacts of ISC by combining intra‐ and intersex additive genetic (co)variances and sex‐specific selection estimates in a multivariate framework. Large genetic correlations between homologous male and female traits combined with evidence for sex‐specific multivariate survival selection suggested that ISC could play an appreciable role in the evolution of this population. Together, multivariate sex‐specific selection and additive genetic (co)variance for the traits considered accounted for additive genetic variance in fitness that was uncorrelated between the sexes (cross‐sex genetic correlation = ?0.003, 95% CI = ?0.83, 0.83). Gender load, defined as the reduction in a population's rate of adaptation due to sex‐specific effects, was estimated at 50% (95% CI = 13%, 86%). This study provides novel insights into the evolution of sexual dimorphism in wild populations and illustrates how quantitative genetics and selection analyses can be combined in a multivariate framework to quantify the microevolutionary impacts of ISC.     

The evolution of sexual size dimorphism in the house finch. III. Developmental basis

Sexual size dimorphism of adults proximately results from a combination of sexually dimorphic growth patterns and selection on growing individuals. Yet, most studies of the evolution of dimorphism have focused on correlates of only adult morphologies. Here we examined the ontogeny of sexual size dimorphism in an isolated population of the house finch (Carpodacus mexicanus). Sexes differed in growth rates and growth duration; in most traits, females grew faster than males, but males grew for a longer period. Sexual dimorphism in bill traits (bill length, width, depth) and in body traits (wing, tarsus, and tail length; mass) developed during different periods of ontogeny. Growth of bill traits was most different between sexes during the juvenile period (after leaving the nest), whereas growth of body traits was most sexually dimorphic during the first few days after hatching. Postgrowth selection on juveniles strongly influenced sexual dimorphism in all traits; in some traits, this selection canceled or reversed dimorphism patterns produced by growth differences between sexes. The net result was that adult sexual dimorphism, to a large degree, was an outcome of selection for survival during juvenile stages. We suggest that previously documented fast and extensive divergence of house finch populations in sexual size dimorphism may be partially produced by distinct environmental conditions during growth in these populations.     

NATURAL SELECTION ON BEAK AND BODY SIZE IN THE SONG SPARROW

We documented temporal patterns of natural selection on beak and body traits in a song sparrow population. We looked for evidence of selection in association with reproduction and overwinter survival in order to identify the conditions under which size in beak and body traits is adaptive. We also attempted to identify the specific traits most closely associated with fitness under these conditions. Selection was observed in association with both survival and reproduction. Patterns of selection differed between the sexes. Selection on males was weak and stabilizing in association with overwinter survival. Selection on females was strong, was both stabilizing and directional, and was associated with both survival and reproduction. In females, traits that enhanced juvenile survival also reduced reproductive success; i.e., there was a trade-off between survival and reproduction. Patterns of selection in the song sparrow parallel those reported for the Galápagos finch, Geospiza fortis. However, in song sparrows, selection occurred mainly on tarsus length and beak length, and not on beak depth or width as in G. fortis. This difference may occur because most North American sparrows partition food resources by habitat, while most Galápagos ground finches partition food by seed size.     

Sexual size dimorphism and assortative mating for morphological traits in Passer domesticus

In this study, assortative mating for different morphological traits was studied in a captive population of house sparrows (Passer domesticus). Males were larger than females. Assortative mating was found for tail length, wing length and general body size. Males with larger badge size mated with females with longer tails. The strongest assortative mating occurred for tail length (r=0.77), and this assortative mating remained significant after controlling for wing length, mass and tarsus length, suggesting that it was not an artefact of assortative mating for body size. The possibility of sexual selection for tail length in the house sparrow is discussed.     

Genetic correlation of wing polymorphism between females and males in the oriental chinch bug,Cavelerius saccharivorus Okajima (Heteroptera: Lygaeidae)

Genetic correlations of the wing form and the relative wing length between females and males were estimated in the oriental chinch bug, Cavelerius saccharivorus, by calculating the correlation between the sexes of the proportion macropterous or the mean relative wing length in full-sib families obtained from different wing forms of parents emerged in a high density population. There was a significantly positive genetic correlation between the sexes in both the proportion macropterous and the mean relative wing length. However, the appearance rate of macropters tended to be much lower in males than in females under the rearing conditions which promote the appearance of macropters. This was evident especially in the offspring of brachypterous parents. These indicated that in C. saccharivorus the wing polymorphism of males is not a simple result of the genetic correlation of wing morphology between the two sexes. It was considered that both of the female and male fitness advantages to wing reduction, as well as the genetic correlation between the sexes, would influence the evolution of wing polymorphism in this species.     

Sex‐specific selection and sexual size dimorphism in the waterstrider Aquarius remigis

We estimated selection on adult body size for two generations in two populations of Aquarius remigis, as part of a long‐term study of the adaptive significance of sexual size dimorphism (SSD). Net adult fitness was estimated from the following components: prereproductive survival, daily reproductive success (mating frequency or fecundity), and reproductive lifespan. Standardized selection gradients were estimated for total length and for thorax, abdomen, genital and mesofemur lengths. Although selection was generally weak and showed significant temporal and spatial heterogeneity, patterns were consistent with SSD. Prereproductive survival was strongly influenced by date of eclosion, but size (thorax and genital lengths in females; total and abdomen lengths in males) played a significant secondary role. Sexual selection favoured smaller males with longer external genitalia in one population. Net adult fitness was not significantly related to body size in females, but was negatively related to size (thorax and total length) in males.     

Sexing greater flamingos by weight and linear measurements

The sex of 59 adult Greater Flamingos (Phoenicopterus ruber roseus L.) was determined by laparoscopic exam of the gonads. Concomitant body weight (kg) and linear measurements (mm) of the culmen (bill), tarsus, middle toe, and wing were taken and compared for males vs. females. Although an overlap between sexes existed in all measurements, males on average were larger than females. Student's t-test indicated significant sexual differences for all five measurements. Thus, weights and linear measurements—especially tarsus, middle toe, and wing length—appear to be a useful parameter in determining an individual's sex in lieu of laparoscopic, feather pulp, blood chromosome, or fecal steroid analysis.     

Differential Response to Selection on the Two Sexes in DROSOPHILA MELANOGASTER

Artificial selection for short wing was performed in two Drosophila melanogaster populations with partially different gene pools: the C populations were derived from a Canton stock while the H lines were derived from a cross between Canton and a b, cn, vg strain. It is shown that in both populations selection on females (CF, HF) was more effective than selection on males (CM, HM). This difference cannot be explained in terms of differences in additive genetic variability between the two sexes because: (1) both sexes contribute to the genetic variability utilized by selection applied to one sex only, and (2) switching selection pressure on females in the M lines does not result in a response comparable to that obtained in the F populations; this rules out almost completely recombination as the responsible agent for the differences between the selection limits reached by M and F selections.-These results, together with several additional observations concerning sexual dimorphism, fitness and the effect of natural selection, suggest that a complex interaction should be involved in the differential response of M and F lines, controlling the wing length phenotype.     

Resolving the conundrum of inbreeding depression but no inbreeding avoidance: Estimating sex‐specific selection on inbreeding by song sparrows (Melospiza melodia)

Inbreeding avoidance among interacting females and males is not always observed despite inbreeding depression in offspring fitness, creating an apparent “inbreeding paradox.” This paradox could be resolved if selection against inbreeding was in fact weak, despite inbreeding depression. However, the net magnitude and direction of selection on the degree to which females and males inbreed by pairing with relatives has not been explicitly estimated. We used long‐term pedigree data to estimate phenotypic selection gradients on the degree of inbreeding that female and male song sparrows (Melospiza melodia) expressed by forming socially persistent breeding pairs with relatives. Fitness was measured as the total numbers of offspring and grand offspring contributed to the population, and as corresponding expected numbers of identical‐by‐descent allele copies, thereby accounting for variation in offspring survival, reproduction, and relatedness associated with variation in parental inbreeding. Estimated selection gradients on the degree to which individuals paired with relatives were weakly positive in females, but negative in males that formed at least one socially persistent pairing. However, males that paired had higher mean fitness than males that remained socially unpaired. These analyses suggest that net selection against inbreeding may be weak in both sexes despite strong inbreeding depression, thereby resolving the “inbreeding paradox.”     

Sex differences in survival selection in the serin,Serinus serinus

Natural selection is demonstrated in most natural populations which suggests that populations are dispatched from their adaptive peaks as a result of selection on correlated characters, or conflicting selection between the sexes. We analysed patterns of survival selection in a population of serins (Serinus serinus) outside Barcelona over a period of 13 years. There was directional selection for increased wing length in males and females accompanied by strong disruptive selection on both tail and wing length in males and a selection against a positive correlation between the two characters in males. In females there was directional selection for increased bill width but decreased bill depth, which should be contrasted to the stabilizing selection acting on bill depth in males. There were conflicting selection on the characters within a sex and conflicting selection of the same characters between sexes, which constrain the rate of access to the nearest adaptive peak.     

Sex differences in body size and body condition in breeding Temminck’s Stints Calidris temminckii

There have been very few reports of body size measurements of live Temminck’s Stints, but earlier studies have shown sex differences in body mass and tarsus length. Here we use molecular techniques to determine the sex of Temminck’s Stints from a Norwegian breeding population. In total, we report measurements of body weight, wing length, tarsus length, bill length, skull length and keel length from 17 males and 30 females. We found significant sex differences in all of these variables, with the exception of tarsus length. The differences in skull length disappeared after the bill lengths had been subtracted from the measurements. A discriminant function analysis based on wing length and bill length correctly classified 86% of the cases (12/16 males, 25/27 females). Female Temminck’s Stints are known to regularly lay more than one clutch of eggs per season and could therefore be expected to be physiologically deprived of bodily energy stores. Nevertheless, we found females to be in better body condition than males.     

SEXUAL SELECTION AND THE EVOLUTION OF SEXUAL SIZE DIMORPHISM IN THE WATER STRIDER,AQUARIUS REMIGIS

Sexual size dimorphism (SSD) is often attributed to sexual selection, particularly when males are the larger sex. However, sexual selection favoring large males is common even in taxa where females are the larger sex, and is therefore not a sufficient explanation of patterns of SSD. As part of a more extensive study of the evolution of SSD in water striders (Heteroptera, Gerridae), we examine patterns of sexual selection and SSD in 12 populations of Aquarius remigis. We calculate univariate and multivariate selection gradients from samples of mating and single males, for two sexually dimorphic traits (total length and profemoral width) and two sexually monomorphic traits (mesofemoral length and wing form). The multivariate analyses reveal strong selection favoring larger males, in spite of the female-biased SSD for this trait, and weaker selection favoring aptery and reduced mesofemoral length. Selection is weakest on the most dimorphic trait, profemoral width, and is stabilizing rather than directional. The pattern of sexual selection on morphological traits is therefore not concordant with the pattern of SSD. The univariate selection gradients reveal little net selection (direct + indirect) on any of the traits, and suggest that evolution away from the plesiomorphic pattern of SSD is constrained by antagonistic patterns of selection acting on this suite of positively correlated morphological traits. We hypothesize that SSD in A. remigis is not in equilibrium, a hypothesis that is consistent with both theoretical models of the evolution of SSD and our previous studies of allometry for SSD. A negative interpopulation correlation between the intensity of sexual selection and the operational sex ratio supports the hypothesis that, as in several other water strider species, sexual selection in A. remigis occurs through generalized female reluctance rather than active female choice. The implications of this for patterns of sexual selection are discussed.     

SEXUAL SELECTION IN THE BARN SWALLOW (HIRUNDO RUSTICA). IV. PATTERNS OF FLUCTUATING ASYMMETRY AND SELECTION AGAINST ASYMMETRY

The patterns of variation in fluctuating asymmetry were studied in four morphological characters of the barn swallow Hirundo rustica. The level of absolute and relative asymmetry was larger in the secondary sexual character “outer tail length” than in three nonsexual morphological traits (wing, central tail, and tarsus length). The extent of individual asymmetry in outer tail length was negatively correlated with tail-ornament size, whereas the relationship between asymmetry of all other morphological characters and their size was flat or U-shaped. Asymmetry in outer tail length was unrelated to asymmetry in other morphological characters, whereas asymmetries in the length of wing, central tail, and tarsus were positively correlated. Male bam swallows exhibited larger asymmetry in outer tail length than females. Asymmetry of most morphological traits exhibited intermediate repeatabilities between years, with the exception of male and female outer tail length, which were highly repeatable. Tail asymmetry of offspring weakly, though significantly, resembled that of their parents. Asymmetry in wing and outer tail length was also significantly related to several fitness components. Male barn swallows that acquired a mate were less asymmetric in wing and outer tail length than unmated males. Females with more asymmetrical tails laid eggs significantly later. Annual reproductive success was unrelated to fluctuating asymmetry. Male barn swallows that survived were less asymmetric in wing and outer tail length than nonsurvivors, whereas female survivors were less asymmetric in outer tail length than nonsurvivors. These results suggest that levels of fluctuating asymmetry in barn swallows are associated with differences in fitness.     

NATURAL SELECTION ON BODY SIZE AND LAYING DATE IN THE TREE SWALLOW

I measured natural selection on body size and laying date in a population of tree swallows (Tachycineta bicolor) from 1986 to 1988. There was little evidence of selection on body size associated with overwinter survival. Disruptive selection on tarsus length, associated with female reproductive success, was detected in one of three years. Both repeatability and mother-daughter regression suggested that laying date was heritable. I found weak evidence of selection on laying date, associated with both overwinter survival and reproduction in females. The ecological implications of both tarsus length and laying date variation in this population could not be identified. Consequently, although I was able to identify the targets of natural selection, the ecological link between trait variation and selection remains unknown.     

Mate choice by marsh wrens: the influence of male and territory quality

If females choose breeding situations to maximize their fitness, then those features of the male and/or territory that are important in mate selection by females should affect female fitness, be assessable prior to mating, and vary among males. The purpose of this study was to identify characteristics of male quality and territory quality that fit these criteria for marsh wrens Cistothorus palustris. The male's contribution to nest defence appeared not to affect female success or choice. However, females that received male assistance with feeding young produced more and heavier fledglings than females without assistance. Males that fed young did not attract more mates. Few males in this population fed young, and no relationship was found between feeding effort and physical or behavioural features of the male. Results of multivariate analyses suggest that, within sites, the measures of territory quality used here cannot explain the variance in the number of young fledged per female or male harem size. The examination of arbitrary territories showed that male pairing success, female settlement order and predation patterns were not significantly correlated between years. The results suggest that territory quality does not influence female success or choice. These results are discussed in the light of earlier attempts to relate features of males and territories to mate selection by female passerines.