Two new species of Knodus (Characidae: Stevardiinae) from the upper rio Tocantins basin, with evidence of ontogenetic meristic changes

Two new species from the upper rio Tocantins basin are described in Knodus based on the traditional definition of the genus. The new species are distinguished from other congeners by meristic and morphometric characters, such as the number of cusps in the premaxillary and dentary teeth, the number of scale series between dorsal-fin origin and lateral line, the orbital diameter and the body depth. With the two new species, the number of endemic species in the upper rio Tocantins basin upstream of the mouth of the rio Paranã, rises to 53 (89 to the confluence with rio Araguaia). The existence of a meristic character that changes through ontogeny (allomery), viz. the number of scale series between dorsal-fin origin and lateral line, was detected in some species of Knodus through a regression analysis. Additionally, this paper describes an unambiguous, more informative and precise new method for counting vertebrae, which will enhance the efficacy of this trait in species comparisons.(AU)

Duas novas espécies do alto rio Tocantins são descritas em Knodus com base na definição tradicional do gênero. As novas espécies são distinguidas das demais congêneres por caracteres merísticos e morfométricos, tais como o número de cúspides nos dentes do pré-maxilar e do dentário, o número de séries de escamas entre a origem da nadadeira dorsal e a linha lateral, o diâmetro da órbita e a altura do corpo. Com as duas novas espécies, o número de espécies endêmicas na bacia do alto rio Tocantins, acima da barra do rio Paranã, sobe para 53 (89 até a confluência com o rio Araguaia). A existência de um caráter merístico que muda ao longo da ontogenia (alomeria), ou seja, o número de séries de escamas entre a origem da nadadeira dorsal e a linha lateral, foi detectado em algumas espécies de Knodus com uma análise de regressão. Adicionalmente, este artigo descreve um novo método não-ambíguo, mas informativo e mais preciso, para a contagem de vértebras, que aumentará a eficácia deste caráter em comparações entre espécies.(AU)

Interpretation of sexual secondary characteristics (SSCs) in regulatory testing for endocrine activity in fish.

Secondary sexual characteristics (SSCs) are important features that have evolved in many fish species because of inter-individual competition for mates. SSCs are crucial not only for sexual selection, but also for other components of the reproductive process and parental care. Externally, they are especially clear in males (for instance, tubercles, fatpad, anal finnage, colouration) but are also externally present in the females (for instance, ovipositor). These characters are under hormonal control and as such there has been much interest in incorporating them as measures in fish test methods to assess the potential endocrine activity of chemicals. Here we describe the external SSCs in typical laboratory test species for endocrine testing - fathead minnow (Pimephales promelas), Japanese medaka (Oryzias latipes), zebrafish (Danio rerio) and the three-spined stickleback (Gasterosteus aculeatus L.). We also provide some examples and discuss the utility of SSC responses to the endocrine activity of chemicals in the field and the laboratory. This paper is not aimed to provide a comprehensive review of SSCs in fish but presents a view on the assessment of SSCs in regulatory testing. Due to the current regulatory importance of establishing an endocrine mode-of-action for chemicals, we also consider other, non-endocrine factors that may lead to SSC responses in fish. We conclude with recommendations for how the assessment of SSCs in fish could be usefully incorporated into the endocrine hazard and risk assessment of chemicals.

A new species of the genus Zographetus Watson, 1893 from China (Lepidoptera: Hesperiidae).

Zographetus rienki sp. nov. is described from southern China based upon two males and one female which are associated by COI barcode. The male of this new species is distinguishable by the following combination of external characters: on the ventral side of the forewing, there is a black hair tuft at basal dorsum, cubitus and basal Cu2 are swollen, and the origin of vein Cu2 is much nearer to wing base than to vein Cu1; both sides of the hindwing are unmarked. The structure of male genitalia also differs from those of all the known congeneric species. Adults and genitalia of the type specimens are illustrated. A key to species of the genus is provided based upon previous works.

Chromosomal Aberrations in Primary Amenorrhea: A Retrospective Study.

OBJECTIVES: The aim of this study was to estimate the frequency of chromosomal abnormalities and establish the association with clinical of factors such as secondary sexual characters and gonad development in primary amenorrhea (PA). STUDY DESIGN: The study was carried out in a large cohort of PA. The chromosomal aberrations were correlated with secondary sexual characters and anatomical abnormalities. MATERIALS AND METHODS: The data of 490 cases of PA were collected retrospectively. The chromosomal preparations were done from the peripheral blood and subjected to giemsa-trypsin-giemsa banding and karyotyped according to the International System of Human Cytogenetic Nomenclature 2013. The fluorescence in situ hybridization was carried out using centromeric and whole painting probes for X and Y chromosome. STATISTICAL ANALYSIS: Statistical analysis of the data was performed using online version of social science statistics software. RESULTS: A high frequency of abnormal uterus (81.9%) and ovaries (86.7%) were detected in our study. A total of 121 (24.7%) cases were identified with abnormal karyotype. The numerical chromosomal abnormalities were identified in 53 (43.8%) cases while structural abnormalities were identified in 32 (26.4%) cases. The XY karyotype was detected in 29.8% females with PA. The PA individuals with anatomical abnormalities (84.3%) had a high frequency (24.6%) of chromosomal aberrations. CONCLUSIONS: The present study concluded that cytogenetics plays an important role in precise diagnosis which helps in the management of PA. The cytogenetic analysis should be carried out to know the genetic basis of PA.

New remarkable sexually dimorphic miniature species of Hyphessobrycon (Characiformes: Characidae) from the upper Rio Tapajós basin.

A new miniature species of Hyphessobrycon is described from an affluent of the Rio Papagaio, tributary of the Rio Juruena, upper Rio Tapajós basin, Mato Grosso State, Brazil. The new species can be distinguished from its congeners by having a remarkable secondary sexual dimorphism in its live colouration (males are red and females yellow), well-defined and relatively narrow dark midlateral stripe extending from tip of snout to tip of middle caudal-fin rays, absence of humeral blotch, 15-18 branched anal-fin rays and five or six branched pelvic-fin rays. The sexually dimorphic colouration of the new species is briefly discussed regarding its temporal variation.

Viability and expression of sexual ornaments in the barn swallow Hirundo rustica: a meta-analysis.

Sexual selection results in the evolution of exaggerated secondary sexual characters that can entail a viability cost. However, in species where sexual ornaments honestly reflect individual quality, the viability cost of secondary sexual characters may be overwhelmed by variation in individual quality, leading to expect that individuals with the largest secondary sexual characters show higher, rather than lower viability. Here, we used meta-analyses to test whether such expected positive relationship between sexual ornamentation and viability exists in the barn swallow Hirundo rustica, which is one of the most studied model species of sexual selection under field conditions. We found a mean positive effect size of viability in relation to the expression of secondary sexual characters of 0.181 (CI: 0.084-0.278), indicating that in this species the more ornamented individuals are more viable, and therefore of high quality. Analyses of moderator variables showed similar effects in males and females, the H. r. rustica subspecies rather than others and tail length rather than other secondary sexual characters. Future research emphasis on other subspecies than the European one and secondary sexual characters than tail length may help identify the sources of heterogeneity in effect sizes.

Assortative mating by colored ornaments in blue tits: space and time matter.

Assortative mating is a potential outcome of sexual selection, and estimating its level is important to better understand local adaptation and underlying trait evolution. However, assortative mating studies frequently base their conclusions on small numbers of individuals sampled over short periods of time and limited spatial scales even though spatiotemporal variation is common. Here, we characterized assortative mating patterns over 10 years in four populations of the blue tit (Cyanistes caeruleus), a passerine bird. We focused on two plumage ornaments-the blue crown and the yellow breast patch. Based on data for 1,657 pairs of birds, we found large interannual variation: assortative mating varied from positive to negative. To determine whether there was nonetheless a general trend in the data, we ran a within-study meta-analysis. It revealed that assortative mating was moderately positive for both ornaments. It also showed that mating patterns differed among populations and especially between two neighboring populations that displayed phenotypic divergence. Our results therefore underscore that long-term studies are needed to draw broad conclusions about mating patterns in natural populations. They also call for studying the potential role of assortative mating in local adaptation and evolution of ornaments in both sexes.

Clasper gland morphology and development in Potamotrygon magdalenae (Elasmobranchii: Potamotrygonidae).

Clasper gland morphology and development in Potamotrygon magdalenae and its relation with the acquisition of reproductive maturity is described in males of different developmental stages (embryos, neonates, juveniles, and reproductively active and resting adults). The glands are subcutaneous masses in the proximal base of each clasper. They are partially bilobate organs with a ventral groove that bears a row of papillae. Glands tend to be asymmetric, the left gland has a larger size, a trend that has been observed in other organs of elasmobranchs. Glands are formed by radially organized tubular secretory units lined with a simple columnar epithelium with basal nuclei and granular eosinophilic cytoplasm; vascularized loose connective tissue surrounds the gland units. The gland is covered by two layers of striated muscle tissue in circular and longitudinal arrangement. The clasper glands begin to develop in neonates and their secretory activity begins in juveniles. The active secretion of the clasper gland is observed in mature males, it includes glycoproteins and sulfated mucopolysaccharides. The size of the glands has a positive and direct relationship with body size, measured as disc width. Significant differences in clasper gland size were found between mature (active and resting) and immature (neonates and juveniles) males, suggesting that the acquisition of the sexual maturity involves the increase in the size of the gland due to a highly augmented secretory activity. Therefore, clasper glands are clearly associated with the reproductive activity of males and their secretion should have an endocrine control as other sexual secondary organs. J. Morphol. 278:369-379, 2017. © 2017 Wiley Periodicals, Inc.

Parabolic variation in sexual selection intensity across the range of a cold-water pipefish: implications for susceptibility to climate change.

While an understanding of evolutionary processes in shifting environments is vital in the context of rapid ecological change, one of the most potent selective forces, sexual selection, remains curiously unexplored. Variation in sexual selection across a species range, especially across a gradient of temperature regimes, has the potential to provide a window into the possible impacts of climate change on the evolution of mating patterns. Here, we investigated some of the links between temperature and indicators of sexual selection, using a cold-water pipefish as model. We found that populations differed with respect to body size, length of the breeding season, fecundity, and sexual dimorphism across a wide latitudinal gradient. We encountered two types of latitudinal patterns, either linear, when related to body size, or parabolic in shape when considering variables related to sexual selection intensity, such as sexual dimorphism and reproductive investment. Our results suggest that sexual selection intensity increases toward both edges of the distribution and that the large differences in temperature likely play a significant role. Shorter breeding seasons in the north and reduced periods for gamete production in the south certainly have the potential to alter mating systems, breeding synchrony, and mate monopolization rates. As latitude and water temperature are tightly coupled across the European coasts, the observed patterns in traits related to sexual selection can lead to predictions regarding how sexual selection should change in response to climate change. Based on data from extant populations, we can predict that as the worm pipefish moves northward, a wave of decreasing selection intensity will likely replace the strong sexual selection at the northern range margin. In contrast, the southern populations will be followed by heightened sexual selection, which may exacerbate the problem of local extinction at this retreating boundary.

A Case of Primary Hypogonadism with Features of Albright's Syndrome.

BACKGROUND: McCune Albright syndrome is rare with an estimated prevalence of 1 in 100,000 to 1 in 1,000,000 persons. The classical clinical triad consists of fibrous dysplasia of the bone, café-au-lait skin spots and precocious puberty. However, in rare cases, there may be primary hypogonadism and amenorrhea. CASE PRESENTATION: An eighteen-year-old female presented with amenorrhea. She had a short stature, round face, thick neck, and short fourth metacarpals and metatarsals. The secondary sexual characters were absent. Serum calcium, phosphorus and parathyroid concentrations were normal, but gonadotropin hormones were very low. X-ray examination revealed short fourth and fifth metacarpals, short left metatarsal, and short fibula. CONCLUSION: These local bony abnormalities along with the biochemical findings helped us to diagnose this case as an unusual presentation of primary hypogonadism with features of McCune Albright's syndrome where there was amenorrhea rather than preocious puberty.

Do male secondary sexual characters correlate with testis size and sperm length in the small hairy maggot blowfly?

The phenotype-linked fertility hypothesis proposes that secondary sexual characters (SSCs) advertise a male's fertility to prospective mates. However, findings from empirical studies attempting to test this hypothesis are often ambivalent or even contradictory, and few studies have simultaneously evaluated how both morphological and behavioural SSCs relate to ejaculate characteristics. Males of the small hairy maggot blowfly, Chrysomya varipes, possess conspicuous foreleg ornaments and display highly stereotyped courtship behaviour. These traits are favoured by females during pre-copulatory mate choice, but it remains unknown whether they correlate with post-copulatory traits expected to influence male fertility. The aim of this study was to investigate whether male courtship and ornamentation correlate with testis size and sperm length in C. varipes. We found that males investing more in courtship had bigger testes, and males with more extensive foreleg ornamentation released sperm with longer tails. Based on the assumption that larger testes enable males to produce more sperm, and that sperm with longer tails have greater propulsive force, our findings suggest that more vigorous and more ornamented males may be more fertile. These findings lend support to the phenotype-linked fertility hypothesis. However, a complete test of this hypothesis will require evaluating whether testis size and sperm length influence male fertilisation ability, as well as female fecundity and/or fertility.

Sex allocation according to multiple sexually dimorphic traits of both parents in the barn swallow (Hirundo rustica).

Parents should differentially invest in sons or daughters depending on the sex-specific fitness returns from male and female offspring. In species with sexually selected heritable male characters, highly ornamented fathers should overproduce sons, which will be more sexually attractive than sons of less ornamented fathers. Because of genetic correlations between the sexes, females that express traits which are under selection in males should also overproduce sons. However, sex allocation strategies may consist in reaction norms leading to spatiotemporal variation in the association between offspring sex ratio (SR) and parental phenotype. We analysed offspring SR in barn swallows (Hirundo rustica) over 8 years in relation to two sexually dimorphic traits: tail length and melanin-based ventral plumage coloration. The proportion of sons increased with maternal plumage darkness and paternal tail length, consistently with sexual dimorphism in these traits. The size of the effect of these parental traits on SR was large compared to other studies of offspring SR in birds. Barn swallows thus manipulate offspring SR to overproduce 'sexy sons' and potentially to mitigate the costs of intralocus sexually antagonistic selection. Interannual variation in the relationships between offspring SR and parental traits was observed which may suggest phenotypic plasticity in sex allocation and provides a proximate explanation for inconsistent results of studies of sex allocation in relation to sexual ornamentation in birds.

The effects of sexual selection on life-history traits: an experimental study on guppies.

Sexual selection is often prevented during captive breeding in order to maximize effective population size and retain genetic diversity. However, enforcing monogamy and thereby preventing sexual selection may affect population fitness either negatively by preventing the purging of deleterious mutations or positively by reducing sexual conflicts. To better understand the effect of sexual selection on the fitness of small populations, we compared components of female fitness and the expression of male secondary sexual characters in 19 experimental populations of guppies (Poecilia reticulata) maintained under polygamous or monogamous mating regimes over nine generations. In order to generate treatments that solely differed by their level of sexual selection, the middle-class neighbourhood breeding design was enforced in the monogamous populations, while in the polygamous populations, all females contributed similarly to the next generation with one male and one female offspring. This experimental design allowed potential sexual conflicts to increase in the polygamous populations because selection could not operate on adult-female traits. Clutch size and offspring survival showed a weak decline from generation to generation but did not differ among treatments. Offspring size, however, declined across generations, but more in monogamous than polygamous populations. By generation eight, orange- and black-spot areas were larger in males from the polygamous treatment, but these differences were not statistically significant. Overall, these results suggest that neither sexual conflict nor the purging of deleterious mutation had important effects on the fitness of our experimental populations. However, only few generations of enforced monogamy in a benign environment were sufficient to negatively affect offspring size, a trait potentially crucial for survival in the wild. Sexual selection may therefore, under certain circumstances, be beneficial over enforced monogamy during captive breeding.

Phylogenetic systematics of Schacontia Dyar with descriptions of eight new species (Lepidoptera, Crambidae).

The Neotropical genus SchacontiaDyar (1914) is reviewed and revised to include eleven species. Schacontia replica Dyar, 1914, syn. n. and Schacontia pfeifferi Amsel, 1956, syn. n. are synonymized with Schacontia chanesalis (Druce, 1899) and eight new species are described: Schacontia umbra,sp. n., Schacontia speciosa,sp. n., Schacontia themis, sp. n., Schacontia rasa, sp. n., Schacontia nyx,sp. n., Schacontia clotho, sp. n., Schacontia lachesis, sp. n., and Schacontia atropos, sp. n. Three species, Schacontia medalba, Schacontia chanesalis, and Schacontia ysticalis, are re-described. An analysis of 64 characters (56 binary, 8 multistate; 5 head, 13 thoracic, 13 abdominal, 25 male genitalic, and 8 female genitalic) scored for all Schacontia and three outgroup species (Eustixia pupula Hübner, 1823, Glaphyria sesquistrialis Hübner, 1823, and Hellula undalis (Fabricius, 1781)) retrieved 8 equally most parsimonious trees (L=102, CI=71, RI=84) of which the strict consensus is: [[[[medalba + umbra] + chanesalis] + speciosa] + [ysticalis + [rasa + themis + [atropos + lachesis + nyx + clotho]]]]. The relevance of male secondary sexual characters to the diagnosis of Schacontia species is discussed.

Diferenciación sexual en la langosta de agua dulce Cherax quadricarinatus (Decapoda: Parastacidae) / Sexual differentiation in the freshwater crayfish Cherax quadricarinatus (Decapoda: Parastacidae)

Determinamos el momento del desarrollo postembrionario en que se produce la diferenciación sexual primaria en la langosta Cherax quadricarinatus. Esta es evidenciada por la presencia de las gónadas y sus respectivos conductos. También determinamos la diferenciación sexual definida por la aparición de los caracteres sexualessecundarios. Se observaron 797 machos, 506 hembras y 456 individuos intersexos de 0.02 a 89.96 g (de criadero y laboratorio). Disecamos una submuestra de 106 machos, 69 hembras y 59 individuos intersexos para la caracterización macroscópica de la estructura gonadal. La diferenciación de los gonoporos se inicia aproximadamente a los 0.10 g, en sincronía con la diferenciación del sistema reproductor en machos, hembras e intersexos. La adquisición de la forma definitiva de ovario, oviducto, testículo y vaso deferente son posteriores. El appendix masculina iniciasu diferenciación a los 0.12-0.2 g y adquiere los rasgos característicos del appendix de los adultos a partir de 1-2 g.La diferenciación de la mancha roja (red patch) ocurre a partir de los 2.3 g.

We determined the earliest stage of postembryonic development at which primary and secondary sexual differentiations occur in the freshwater (“red claw”) crayfish Cherax quadricarinatus. For this purpose,797 males, 506 females and 456 intersex specimens within a weight range of 0.02-89.96 g were observed under stereoscopic microscope to determine the presence of the genital openings at the basis of the third (females) or fifth (males) pair of pereiopods. Animals presenting both pairs of genital openings were considered as intersex. A subsample of 106 males, 69 females and 59 intersex were dissected for the macroscopic characterization of gonad morphology. The development of the genital openings began approximately at 0.10 g, simultaneously with the differentiation of the reproductive system in females, males and intersex. Although the differentiation of the reproductive system started very early in the postembryonic development, the definitive form and colour of ovaries, oviducts, testes and vasa deferentia was acquired later. The differentiation of the appendix masculina began at 0.12-0.2 g and acquired the elongated shape of the adult at 1-2 g. The soft red patch characteristic of adult males started at 2.3 g of body weight in both chelipeds.

POPULATION DIVERGENCE IN SEXUAL ORNAMENTS: THE WHITE FOREHEAD PATCH OF NORWEGIAN PIED FLYCATCHERS IS SMALL AND UNSEXY.

Models of sexual selection suggest that populations may easily diverge in male secondary sexual characters. Studies of a Spanish population of the pied flycatcher, Ficedula hypoleuca, and a Swedish population of the closely related collared flycatcher, F. albicollis, have indicated that the white forehead patch of males is a sexually selected trait. We studied the white forehead patch of male pied flycatchers (n = 487) in a Norwegian population over seven years. Males with large forehead patches were in general more brightly colored, but patch height was not correlated to body mass, body size, or parasite loads. Conditions during the nestling period did not seem to influence patch height as an adult. Patch height increased slightly from the first to the second year as adults, but then remained relatively constant at higher ages. Patch height was not related to survival. Year-to-year changes showed that males who increased in patch height also increased in body mass, suggesting that expression of the forehead patch may be partly condition dependent. However, changes in body mass explained only a small proportion of the variance in patch height between males. Thus, patch height would not be a good indicator of male quality. Furthermore, patch size was also not related to male ability to feed nestlings, indicating that females would not obtain direct benefits by choosing males with large patches. However, patch height could be a Fisher trait, but this requires heritability and there was no significant father-son resemblance in patch height. Comparisons of the males visited by each female during the mate sampling period indicated that chosen males did not have larger forehead patches than rejected males. Experimental manipulation of patch height did not affect male mating success. These results indicate that females do not use patch size as a mate choice cue. Finally, patch height did not predict the outcome of male contests for nestboxes, suggesting that the forehead patch is not an intrasexually selected cue of status. Norwegian pied flycatchers have smaller forehead patches than both Spanish pied flycatchers and Swedish collared flycatchers. We suggest that this pattern may be explained by the lack of sexual selection on the forehead patch in the Norwegian population as compared to the other populations, where the patch is apparently sexually selected. We discuss possible reasons for these population divergences, such as female choice on an alternative secondary sexual character (general plumage color) and speciation among Ficedula flycatchers.

GENETIC, ENVIRONMENTAL, AND CONDITION-DEPENDENT EFFECTS ON FEMALE AND MALE ORNAMENTATION IN THE BARN OWL TYTO ALBA.

Secondary sexual characters are thought to indicate individual quality. Expression of sex-limited traits in an extravagant state may require both the underlying genes and the available nutrient resources. The assessment of the relative contribution of genes, environment, and body condition is relevant for understanding to that extent the extravagant trait may signal genotypic or phenotypic quality of the individual. In birds, usually only the males are ornamented. In the barn owl, Tyto alba, both females and males display sex-limited plumage traits. Males are commonly lighter colored and females spottier. In an experiment with combined cross-fostering and brood size manipulation we determined the relative contribution of genes, environment, and body condition to the variation in plumage coloration and plumage spottiness. The partial cross-fostering experiment tested the relative importance of shared genes and a shared environment for the resemblance of related birds. Siblings raised in different nests converged toward similar trait values, offspring resembled the true but not the foster parents, and plumage traits of unrelated nestlings sharing the same nest were not correlated. Results were not inflated by maternal effects detectable in the mother's phenotype, because middaughter to mother resemblance was not higher than midson to father resemblance. This suggests that plumage coloration and spottiness are largely genetically inherited traits, and that the rearing environment does not have a strong impact on the expression of these traits. To further investigate whether the two sex-limited traits are condition dependent, brood sizes were manipulated. Enlargement or reduction of broods by two nestlings resulted in lower and higher body mass of nestlings, respectively. However, nestlings raised in enlarged or reduced broods did not show either a significantly darker or lighter or a more or less spotted plumage. We did not detect any genotype-by-environment interaction. In conclusion, simultaneous cross-fostering and brood size manipulation demonstrate that additive genetic variance for plumage coloration and spottiness is maintained and that both the rearing environment and body condition do not account for a large proportion of the phenotypic variance in female and male ornamentations.

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

Ornamental tail feathers of male barn swallows (Hirundo rustica) confer an advantage in sexual selection because long-tailed males are preferred by females. However, the size of tail ornaments exceeds the natural selection optimum and males are predicted to pay an energetic cost for flying, directly related to tail length. An increase in hematocrit is an adaptive response to enhance oxygen uptake, for example during periods of intense locomotory activity. In this study, we analyzed the effect of experimental manipulation of tail length on the hematocrit of male barn swallows from an Italian and a Spanish population. We predicted that the natural decrease in hematocrit during the breeding season would be reduced by experimental elongation and enhanced by experimental shortening of tail ornaments. The results showed that the decrease in hematocrit was significantly different among tail treatments, and tail-elongated males had the smallest hematocrit reduction. In Italy, the hematocrit of tail-elongated males did not change after tail manipulation, while that of two control groups and tail-shortened males decreased. A comparatively high hematocrit in males with experimentally enlarged tail ornaments may be a response to increased energetic requirements and, hence, to oxygen demands for flying imposed by their tail morphology.

AN EXPERIMENTAL STUDY OF PATERNITY AND TAIL ORNAMENTATION IN THE BARN SWALLOW (HIRUNDO RUSTICA).

Previous studies of the socially monogamous barn swallow (Hirundo rustica) have shown that males that most frequently engage in extrapair copulations and whose partners are least involved in copulations with extrapair males are those with long tail ornaments. In this study, through the use of three highly polymorphic microsatellite markers, we analyze the relationships between length of tail ornaments of male barn swallows and proportion of nestlings fathered in own broods, number of offspring fathered in broods of other pairs, and total number of offspring fathered, using both a correlational and an experimental approach. Consistent with our predictions, we show that males with either naturally long or experimentally elongated tails have higher paternity (proportion of biological offspring in own broods), and they produce more biological offspring during the whole breeding season than males with naturally short or experimentally shortened tails. Males with naturally long tails also had more offspring in extrapair broods than short-tailed males, but the effect of tail manipulation on the number of offspring fathered in extrapair broods, although being in the predicted direction, was not statistically significant. Cuckolded males that did not fertilize extrapair females had smaller postmanipulation tail length than cuckolders. We conclude that there is a causal, positive relationship between male tail length and paternity. Since female barn swallows have extensive control over copulation partners and heritability of tail length is high, this study shows that female choice is a component of selection for larger male ornaments. Benefits from extrapair fertilizations to females may arise because they acquire "good" genes for sexual attractiveness or high viability for their offspring.

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.