A Variation of Secondary Sex Ratio in Blue Fox Alopex lagopus L.

The effects of several genotypic and paratypic factors on the secondary (at birth) sex ratio was analyzed in blue fox bred in captivity. In particular, variation of sex ratio was for the first time studied as dependent on sire's age (without considering dam's age), the ages of both sire and dam, and the lines of both parents. The initial data were obtained from the Pushkino breeding facility, Moscow oblast in 1985 through 1989. In total, 15 396 puppies were analyzed. The frequency of males ( ) in this population was 0.551 ± 0.004 (confidence interval 0.543 P 0.559). Parents' ages and litter size had no effect on the proportion of males in the progeny. In one of the two blue fox subpopulations under study, dam's line proved to be associated with a significant departure of sex ratio to a higher proportion of males, suggesting the effect of genotypic factors on the variation of secondary sex ratio in blue fox.     

Sex ratio in silver foxes: effects of domestication and the star gene

The course of changes in secondary sex ratio (proportion of males at birth) in silver foxes bred at the fur farm of this Institute was analyzed. Data collected over several years of breeding of a domesticated (experimental) population selected for amenability to domestication and of a commercial (control) were compared. A tendency to increase in secondary sex ratio was demonstrated in both populations. However, the proportion of males at birth was higher in domestic foxes. This proportion, calculated from the combined data for 1978–1993, was 0.538±0.005 and 0.511±0.007 in the selected and commercial populations, respectively. The minimal departure of the observed sex ratio from 0.5 was demonstrated for litters with five pups, which is close to the average litter size in fox populations. The proportion of males increases with both increasing and decreasing litter size. An analysis of secondary sex ratio with respect to maternal age revealed a minimal departure of sex ratio from the expected in offspring from foxes of optimal reproductive age (2–4 years). An effect of the autosomal semidominant coat color mutation star on male excess at birth was also found: secondary sex ratio was higher (0.583±0.015) in offspring of mothers heterozygous for the star mutation than from standard types of the domesticated population. The increase in secondary sex ratio in the analyzed fox populations is viewed as a correlated response to selection for domestication. The hormonal mechanisms mediating the effects of both this selection and the star mutation on sex ratio at birth are discussed.     

Effect of the hereditary characteristics of male blue foxes Alopex lagopus L. on the sex ratio of their offspring

Family analysis of a commercial population of the blue fox (the Pushkinskoe Breeding Fur Farm, Moscow oblast) with respect to secondary sex ratio has been performed. The offspring of each individual male or female involved in crossing between 1984 and 1988 was analyzed. The study of all families formed by every male and every female has made it possible to determine a group of "outstanding" fathers (23 out of 287 males), whose offspring was predominantly male (62.1% of the offspring were males, versus 53.9% in the total population). The results of subsequent detailed study on the pedigrees of male blue foxes in whose offspring the sex ratio significantly deviates from 1:1 indicate that this character is transmitted from fathers to sons without the deterioration of other commercially valuable characters. It is presumed that the significant deviation of sex ratio from 1:1 in the offspring of some male blue foxes is determined by genetic factors.     

Analysis of secondary sex ratio in the fox (<Emphasis Type="Italic">Vulpes vulpes</Emphasis> L.)

Secondary sex ratio and its variability in relation to some paratypic and genetic factors were studied in the silver fox by analysis of data obtained at the Pushkinskii fur farm (Moscow oblast) in 1980–1989. A total of 17285 whelps were examined. The mean proportion of males over the ten years of observation was 0.536 ± 0.004. No effect of parent age or litter size on this proportion was found. Individual analysis of the progeny of a single parent revealed 44 males and 49 females showing significant deviations from the expected sex ratio (1 ♂: 1 ♀). These results can be used for reconstruction of pedigrees whose progeny yields regular deviations from the expected sex ratio.     

Analysis of secondary sex ratio in the fox (Vulpes vulpes L.)

Secondary sex ratio and its variability in relation to some paratypic and genetic factors were studied in the silver fox by analysis of data obtained at the Pushkinskii fur farm (Moscow oblast) in 1980-1989. A total of 17285 whelps were examined. The mean proportion of males over the ten years of observation was 0.536+/-0.004. No effect of parent age or litter size on this proportion was found. Individual analysis of the progeny of a single parent revealed 44 males and 49 females showing significant deviations from the expected sex ratio (l male : 1 female). These results can be used for reconstruction of pedigrees whose progeny yields regular deviations from the expected sex ratio.     

Effect of the hereditary characteristics of male blue foxes <Emphasis Type="Italic">Alopex lagopus</Emphasis> L. on the sex ratio of their offspring

Family analysis of a commercial population of the blue fox (the Pushkinskoe Breeding Fur Farm, Moscow oblast) with respect to secondary sex ratio has been performed. The offspring of each individual male or female involved in crossing between 1984 and 1988 was analyzed. The study of all families formed by every male and every female has made it possible to determine a group of outstanding fathers (23 out of 287 males), whose offspring was predominantly male (62.1% of the offspring were males, versus 53.9% in the total population). The results of subsequent detailed study on the pedigrees of male blue foxes in whose offspring the sex ratio significantly deviates from 1 : 1 indicate that this character is transmitted from fathers to sons without the deterioration of other commercially valuable characters. It is presumed that the significant deviation of sex ratio from 1 : 1 in the offspring of some male blue foxes is determined by genetic factors.Translated from Genetika, Vol. 41, No. 3, 2005, pp. 422–426.Original Russian Text Copyright © 2005 by Beketov, Kashtanov.     

Milk intake in blue fox (Alopex lagopus) and silver fox (Vulpes vulpes) cubs in the early suckling period

Milk intake of fox cubs (2-16 days of age; body weight, 96-649 g) in ten blue fox litters and ten silver fox litters were measured by the water isotope dilution (WID) technique following a single intraperitoneal injection of tritiated water (3HHO). Litter size varied from four to 14 in blue foxes and from three to eight in silver foxes. Silver fox cubs had higher birth weights than blue foxes. Inter-species body weights and growth rates were apparently dependent on litter size and the dam's constitution. In both species growth rate increased with age and body weight (7-35 g per day). In the cubs, the biological half-life of body water turnover (BWT) rose from 1.5 days at 2-3 days of age to 2.5 days at 13-16 days of age, although a considerable scatter was seen. The mean daily milk intake of the cubs varied with body weight, from 31 to 193 g per day, whereas daily milk intake per unit of body mass remained stable at 30-35 g per 100 g body weight. The ratio of milk intake to body weight gain varied considerably among cubs, averaging 4.5 g/g during the 3-week experimental period. In suckling fox cubs, the calculated daily intake of metabolically energy (ME) corresponded fairly with the estimated energy requirements for growth and maintenance of the young. Finally, the applicability and the accuracy of the WID technique was evaluated in ten 3-week-old fox cubs, by tube-feeding with a milk replacer for 48 h, which documented that the daily rates of milk intake and water turnover can be accurately measured in suckling fox cubs by the WID technique following a single injection of 3HHO.     

Offspring sexual dimorphism and sex-allocation in relation to parental age and paternal ornamentation in the barn swallow

We analysed the morphology of nestling barn swallows (Hirundo rustica) in relation to their sex, and laying and hatching order. In addition, we studied sex-allocation in relation to parentage, parental age and expression of a secondary sexual character of fathers. Molecular sexing was conducted using the sex chromosome-linked avian CHD1 gene. Sex of the offspring was not associated with laying or hatching order. None of nine morphological, serological and immunological variables varied in relation to offspring sex. Sexual dimorphism did not vary in relation to parental age and expression of a paternal secondary sexual character. The proportion of sons declined with brood size. Individual males and females had a similar proportion of sons during consecutive breeding years. The proportion of sons of individual females declined with age, but increased with the expression of a secondary sexual character of their current mate. The generalized lack of variation in sexual dimorphism among nestlings may suggest that barn swallows do not differentially invest in sons vs. daughters. Alternatively, male offspring may require different parental effort compared to their female siblings in order to attain the same morphological state. The lack of variation in offspring sexual dimorphism with paternal ornamentation suggests no adjustment of overall parental effort in relation to reproductive value of the two sexes. However, male-biased sex ratio among offspring of highly ornamented males may represent an adaptive sex-allocation strategy because the expression of male ornaments is heritable and highly ornamented males are at a sexual selection advantage.     

Correlations between ultraviolet coloration, overwinter survival and offspring sex ratio in the blue tit

We studied the correlations between offspring sex ratio, UV coloration and overwinter survival in a population of blue tits, breeding in Gotland, Sweden, over three consecutive breeding seasons. In 2 of 3 years, we found that females paired to males with relatively brighter UV-coloration produced a greater proportion of sons in their broods, and that this effect was significant with all 3 years combined, despite a significant year by male UV interaction. In addition, we found other correlates of sex ratio (breeding time, female age and clutch size) in some, but not all years, and some of these showed significantly different relationships with sex ratio between years. In both years for which data were available, there were indications that males with relatively brighter UV coloration, and that paired with females that produced male-biased clutches, were more likely to survive to the next year. In addition, we also found that in both males and females, individuals produced similar sex ratios in consecutive years. Because correlations with the sex ratio may be expected to be weak, variation in results between years within the same population may be explained by low statistical power or genuine biological differences. Our results suggest that conclusions about sex ratio variation in birds should be based on multiple years. The correlations that we found in some years of this study are consistent with models of adaptive sex ratio adjustment in response to mate quality. However, careful experimental work is required to provide tests of the assumptions of these models, and should be a priority for future work.     

Temperature and sex allocation in a spider mite

Although temperature is the most important environmental factor regulating arthropod development and reproduction, its influence on sex allocation in haplodiploid arthropods remains largely unexplored. We investigated under laboratory conditions how maternal age and temperature mediate offspring sex ratio of the spider mite Tetranychus mcdanieli (Acari: Tetranychidae). Over nine temperature regimes, female-biased sex ratios were consistently observed, varying from 57 to 87% among progeny produced over lifetime. Spider mite sex ratio was affected by maternal age: more male progeny were produced at both the beginning and the end of the female lifespan, yielding a dome shaped curvilinear relationship. This pattern of variation with age probably results from constraints on using sperm at young ages and sperm depletion or viability at older ages. We found a significant curvilinear relationship between temperature and sex ratio. The proportion of female offspring was lowest at intermediate temperatures and highest at extreme temperatures. We suggest that increased female-biased sex ratio at extreme temperatures is an evolutionary response of spider mites to deteriorating habitats as, in the Tetranychidae, females have better capacities than males to disperse and survive under harsh conditions.     

Female bias in the adult sex ratio of African annual fishes: interspecific differences,seasonal trends and environmental predictors

Many populations have consistently biased adult sex ratios with important demographic and evolutionary consequences. However, geographical variation, the mechanisms, temporal dynamics and predictors of biased sex ratios are notoriously difficult to explain. We studied 334 wild populations of four species of African annual fish (Nothobranchius furzeri, N. kadleci, N. orthonotus, N. rachovii) across their ranges to compare their adult sex ratio, its seasonal dynamics, interpopulation variation and environmental predictors. Nothobranchius populations comprise a single age cohort and inhabit discrete isolated pools, with wide-ranging environmental conditions (habitat size, water turbidity, structural complexity, predators), making them ideal to study adult sex ratio variation. In captivity adult sex ratios were equal. In natural populations, adult sex ratios were biased 1:2 toward females in three study species while N. kadleci had sex ratios at unity. There was a decline in the proportion of males with age in one species, but not in the other species, implying most severe male mortality early after maturation, declining later perhaps with a decrease in male abundance. In general, the populations at vegetated sites had relatively more males than populations at sites with turbid water and little vegetation. Selective avian predation on brightly coloured male fish likely contributed to female dominance and vegetation cover may have protected males from birds. In addition, an aquatic predator, a large belastomid hemipteran, decreased the proportion of males in populations, possibly due to greater male activity rather than conspicuous colouration. Alternative explanations for a sex ratio bias, stemming from male–male contests for matings, are discussed. We conclude that the effect of environmental conditions on adult sex ratio varies dramatically even in closely related and ecologically similar sympatric species. Therefore, difficulties in explaining the ecological predictors of sex ratio biases are likely due to high stochasticity rather than limited sample size.     

Sex ratio of some long-lived dioecious plants in a sand dune area

In dioecious plants the fraction of males among flowering plants in the field (the secondary sex ratio) is the result of the fraction of males in the seeds (the primary sex ratio) and the subsequent survival and age at first reproduction of the two genders. It has been assumed that survival and age at first reproduction are the main determinants of biased secondary sex ratio but, especially for long-lived perennials, few data are available. We address this issue for natural populations of four long-lived perennials in a dune area. In Asparagus officinale and Bryonia dioica, the secondary sex ratio was unbiased. In Salix repens the secondary sex ratio was female-biased (0.337). Hippophae rhamnoides populations were male-biased; the average sex ratio of flowering plants was 0.658, while the fraction of males varied between 0.39 near the sea to 0.84 at the inland side of the dunes. The primary sex ratio was estimated by germinating seeds and growing plants under favourable conditions with minimal mortality. In S. repens the primary sex ratio in seeds was variable among mother plants and was, on average, female-biased (0.289). This is close to the secondary sex ratio, suggesting that the female bias already originates in the seed stage. In Hippophae rhamnoides the primary sex ratio was slightly male-biased (0.564). We argue that in this species, apart from the primary sex ratio, higher mortality and a later age at first reproduction for females contribute to the strong male bias among flowering plants in the field.     

Factors affecting recruitment age and recruitment probability in the Western Gull Larus occidentalis

We studied recruitment age (i.e. the age that a gull bred for the first time) and the probability that an adult reproduced during its life in three cohorts of Western Gulls Larus occidentalis breeding on southeast Farallon Island. We examined the effects of sex, hatching date, hatching order, brood size and annual variation in food supply during the recruitment period. Modal recruitment ages of males and females were 4 and 5 years, respectively, a difference that was significant. None of the factors examined, including hatching date, hatching order and brood size, had a significant effect on recruitment age. The probability of breeding in males who experienced low food supply in their fourth and fifth years was lower than that of males who experienced favourable food supply in their fourth and fifth years. Recruitment probability (the proportion of the sample group that recruited among those available to recruit during a given year) of 4- and 5-year-old males was lower in the food-poor 1983 El Nino than in years of favourable food supply. The only variable affecting recruitment probability of females was age: recruitment probability was greater among 5-year-olds than 4-year-olds. Recruitment age in males was more sensitive to annual variation in food supply, probably because females have more difficulty recruiting due to an excess of adult females in the population and because of the different roles in reproduction in which males are primarily responsible for provisioning chicks and maintaining the territory.     

Causes and consequences of adaptive seasonal sex ratio variation in house sparrows

1. Here we examine how sex ratio variation in house sparrow broods interacts with other demographic traits and parental characteristics to improve the understanding of adaptive significance and demographic effects on variation in sex ratio. 2. The sex ratio in complete broods did not deviate significantly from parity (54.9% males). 3. There was sex-specific seasonal variation in the probability of recruitment. Male nestlings that hatched late in the breeding season had larger probability of surviving than early hatched males. 4. An adaptive adjustment of sex ratio should favour production of an excess of males late in the breeding season. Accordingly, the proportion of male offspring increased throughout the breeding season. 5. A significant nonlinear relationship was present between sex ratio and age of the female. However, there was no relationship between parental phenotype and standardized hatch day that could explain the observed seasonal change in sex ratio. 6. The sex-specific number of offspring recruited by a pair to subsequent generations was closely related to the brood sex ratio. 7. These results indicate an adaptive adjustment of sex ratio to seasonal variation in environmental conditions that affects the offspring fitness of the two sexes differently. Our results also suggest that such a sex ratio variation can strongly influence the demography and structural composition of small passerine populations.     

Primary sex ratio adjustment to experimentally reduced male UV attractiveness in blue tits

The study of primary sex ratio adjustment in birds is notoriousfor inconsistency of results among studies. To develop our understandingof avian sex ratio variation, experiments that test a prioripredictions and the replication of previous studies are essential.We tested if female blue tits Parus caeruleus adjust the sexratio of their offspring to the sexual attractiveness of theirmates, as was suggested by a previous benchmark study on thesame species. In 2 years, we reduced the ultraviolet (UV) reflectanceof the crown feathers of males in the period before egg layingto decrease their attractiveness. In contrast to the simpleprediction from sex allocation theory, we found that the overallproportion of male offspring did not differ between broods ofUV-reduced and control-treated males. However, in 1 year, theUV treatment influenced offspring sex ratio depending on thenatural crown UV reflectance of males before the treatment.The last result confirms the pattern found in the previous bluetit study, which suggests that these complex patterns of primarysex ratio variation are repeatable in this bird species, warrantingfurther research into the adaptive value of blue tit sex ratioadjustment to male UV coloration.     

Effects of population structure and density on calf sex ratio in red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>)—implications for management

There is ongoing interest to assess what factors affect offspring sex ratio, especially in ungulates. Wildlife managers might be interested in influencing this sex ratio for two reasons: either in order to limit population growth more effectively by reducing the proportion of females born or to increase revenues by a higher proportion of trophy bearing males in the population. While previous studies mostly focused on how maternal traits affect offspring sex ratio, we included here also male traits in our analysis. We achieved this by investigating data from 30 areas covering entire Lower Austria, collected over the past 12 years from both hunted red deer and those killed in road accidents. We focused our analyses on parameters that can be easily assessed by managers on the population and individual level, i.e. the numbers of animals culled in different age/sex classes and their body mass. We found that the proportion of females among calves increased with population density. Furthermore, we found that calf sex ratio (i.e. the proportion males among calves aged between 2 and 7 months) increased with increasing proportions of adult females and males older than 10 years, independent of the density effect. We conclude that wildlife managers interested in the effective reduction of red deer abundance and/or increasing the proportion of males among offspring should select a culling regime leading to a low population density dominated by adult, prime-aged females and males. This can be achieved by over-proportional removal of young females and warranting that a high number of strong males reach an age of at least 10 years.     

Age ratio and survival of Common Rosefinch (<Emphasis Type="Italic">Carpodacus erythrinus</Emphasis>, Passeriformes,Fringillidae) in a local population based on ringing data

The ringing data on Common Rosefinch obtained during thirteen field seasons (2002–2014) at the National Park “Russky Sever” (Vologda region) are analyzed. Birds were trapped in two sites 59 km apart. The age of the birds (yearlings or first breeders and adults) was determined according to the plumage color in every male captured with mist nets. The apparent survival rate of adult individuals was determined using two methods: according to the ratio of males’ age and the CJS model (Lebreton et al., 1992; Bursky, 2011). A total of 713 captures of 657 individuals were analyzed. The ratios of ages in the males from the sites studied did not differ; yearlings amounted to 56% of all males. The rate of survival in the males based on the age ratios was estimated at 44 ± 2%. The apparent survival rate obtained using the CJS model was significantly higher (61 ± 6%). The survival rate of adult rosefinches in two sites did not differ and did not depend on sex. Literature materials on the age ratio and survival of rosefinches in local populations are analyzed. There is a suggestion that the mismatch of the survival rates obtained by two methods and the high proportion of yearling males in the banding captures can be explained by the fact that the majority of individuals of that age category did not participate in nesting.     

The MHC and non-random mating in a captive population of Chinook salmon

Detailed analysis of variation in reproductive success can provide an understanding of the selective pressures that drive the evolution of adaptations. Here, we use experimental spawning channels to assess phenotypic and genotypic correlates of reproductive success in Chinook salmon (Oncorhynchus tshawytscha). Groups of 36 fish in three different sex ratios (1:2, 1:1 and 2:1) were allowed to spawn and the offspring were collected after emergence from the gravel. Microsatellite genetic markers were used to assign parentage of each offspring, and the parents were also typed at the major histocompatibility class IIB locus (MHC). We found that large males, and males with brighter coloration and a more green/blue hue on their lateral integument sired more offspring, albeit only body size and brightness had independent effects. There was no similar relationship between these variables and female reproductive success. Furthermore, there was no effect of sex ratio on the strength or significance of any of the correlations. Females mated non-randomly at the MHC, appearing to select mates that produced offspring with greater genetic diversity as measured by amino-acid divergence. Females mated randomly with respect to male genetic relatedness and males mated randomly with respect to both MHC and genetic relatedness. These results indicate that sexual selection favours increased body size and perhaps integument coloration in males as well as increases genetic diversity at the MHC by female mate choice.     

A two-generation study of human sex-ratio variation.

We report here the first vertical population study of human sex-ratio variation. Sex-ratio data for 2 generations from Akita, Japan, have been analyzed. Parental age, birth order, sequences of the sexes at birth, and generations have no statistically significant effect on sex ratio. There is a slight excess of males at birth, as is typical for human sex-ratio studies. There is evidence of sex-ratio-dependent family planning. An analysis of vertical transmission of sex-ratio modifying factors that excludes effects of birth order in both the parental and offspring generations has detected a marginally significant paternal effect. Genetic variability of the sex ratio, if present at all, is of a very minor magnitude.