Genetic microdifferentiation in the Semai Senoi of Malaysia.

Blood samples, demographic and cultural data were collected from seven settlements of Semai Senoi, a swidden farming ethnic group of Malaysia. Three genetic loci (ABO blood group, hereditary ovalcytosis, and hemoglobin) were analyzed in a total sample of 546 individuals. These data indicate a considerable degree of genetic microdifferentiation in this area of the Semai distribution. Parent-offspring birthplace data (analyzed by means of a migration matrix) and settlement histories show that settlements are not strongly isolated. Genetic differences in the study area demonstrate a reasonable correspondence with migration and the history of the settlements. Genetic convergence also occurs through the addition of migrant groups to established populations leading to new patterns of marriage between donor and recipient groups. The genetic structure of the total Semai population through time thus comprises a mosaic of shifiting allele frequencies in a series of semi-isolated local populations.     

Fission-fusion and lineal effect: aspects of the population structure of the Semai Senoi of Malaysia.

Analysis of histories and genealogies from seven relatively unacculturated, swidden-farming Semai settlements shows that the composition of local groups fluctuates through time. This instability is similar to a pattern which Neel and his colleagues have suggested is typical of primitive society, the fission-fusion model. In addition, the individuals comprising Semai fission groups are kinsmen which implies that the number of independent genomes represented is markedly less than the number of individual migrants (the lineal effect). Fission groups may form new villages or fuse with an established settlement. In either case, the genetic effects of such migration are more pronounced than would be expected on the basis of founder effect or random migration. Despite several conspicuous differences in social organization between the Semai and the South American Indians (e.g., bilateral vs. unilineal descent) whose population structure provided the empirical basis for the fission-fusion, lineal effect model, the basic similarities are striking. The Semai case thus lends support to the proposition that this pattern may be of some generality in technologically primitive populations.     

Patterns of sex ratio, virginity and developmental mortality in gregarious parasitoids

Theory considering sex ratio optima under ‘strict local mate competition with offspring groups produced by a single foundress’ makes a suite of predictions, one of which is a mean female bias. Treating individual offspring as discrete units, theory further predicts sex ratios to have low variance (precise sex ratio) and to equal the reciprocal of clutch size (one male per clutch). The maternal decision may be complicated by imperfect control of sex allocation, limited insemination capacity of sons and offspring developmental mortality: each can lead to virgin daughters (with zero fitness) and consequently select for less biased sex ratios. When clutches are small and/or developmental mortality is common, appreciable proportions of virgins are expected, even when control of sex allocation is perfect and the mating capacity of males is unlimited. This suite of predictions has been only partially tested. We provide further tests by examining sex ratios and developmental mortalities within and across species of locally mating parasitoids. We find a wide range of mean developmental mortalities (6–67%), but mortality distributions are consistendy overdispersed (have greater than binomial variance) and sexually differential mortality appears to be absent. Sex ratios are female biased and have low variance, but are not perfectly precise and variance is increased by mortality within species and (equivocally) across species. Sex ratios less biased than the reciprocal of clutch size are observed; probably due to a maternal response to developmental mortality in one species, and to limited insemination capacity in others. Cross species comparisons indicate that mean proportions of mortality and virginity are positively correlated. Virginity is more prevalent than predicted among species with higher mortalities but not among lower mortality species. Predicted relationships between virginity and clutch size are supported in species with lower mortalities but only partially supported when mortality rates are higher.     

Skewed birth sex ratio and premature mortality in elephants

Sex allocation theories predict equal offspring number of both sexes unless differential investment is required or some competition exists. Left undisturbed, elephants reproduce well and in approximately even numbers in the wild. We report an excess of males are born and substantial juvenile mortality occurs, perinatally, in captivity. Studbook data on captive births (CB, n = 487) and premature deaths (PD, <5 years of age; n = 164) in Asian and African elephants in Europe and North America were compared with data on Myanmar timber (Asian) elephants (CB, n = 3070; PD, n = 738). Growth in CB was found in three of the captive populations. A significant excess of male births occurred in European Asian elephants (ratio: 0.61, P = 0.044) and in births following artificial insemination (0.83, P = 0.003), and a numerical inclination in North American African elephants (0.6). While juvenile mortality in European African and Myanmar populations was 21–23%, it was almost double (40–45%) in all other captive populations. In zoo populations, 68–91% of PD were within 1 month of birth with stillbirth and infanticide being major causes. In Myanmar, 62% of juvenile deaths were at >6 months with maternal insufficient milk production, natural hazards and accidents being the main causes. European Asian and Myanmar elephants PD was biased towards males (0.71, P = 0.024 and 0.56, P < 0.001, respectively). The skewed birth sex ratio and high juvenile mortality hinder efforts to help captive populations become self-sustaining. Efforts should be invested to identify the mechanism behind these trends and seek solutions for them.     

About the sex ratio in connection with early embryonic mortality in man

The problem of the functioning specificity of sex chromosomes during the early stages of embryogenesis in man and the associated problem of the sex ratio in spontaneous and induced abortions, as well as in newborns, remains open. We have conducted a cytogenetic examination of 342 spontaneous abortions divided into three clinical groups on the basis of the severity of the developmental disturbances of the embryo: spontaneous abortionssensu stricto with a developed embryo without any significant intrauterine delay of development (n=100), nondeveloping pregnancies (n=176), and anembryonic fetuses (n=66). The frequency of chromosomal mutations in these groups was 22.0, 48.3, and 48.5%, respectively. Statistical analysis has demonstrated significant differences between the studied groups in the frequencies of the normal and abnormal karyotypes: the major contributions to these differences were associated with autosomal trisomy, triploidy, and the 46.XY karyotype. The presence of 46.XY may reflect the specific genetic mechanisms of the prenatal mortality of embryos with the normal karyotype, associated with sex and/or with the imprinting of X-chromosomes. The sex ratio in spontaneous abortions with the normal karyotype was as follows: 0.77 for spontaneous abortions with well-developed embryos without any significant intrauterine delay of development; 0.60 for nondeveloping pregnancies; and 0.31 for anembryonic fetuses. An analysis of DNA from the embryos and their parents has demonstrated a low probability of contamination of cell cultures with mother cells as a possible source of the prevalence of embryos with the 46.XX karyotype among spontaneous abortions. Nondeveloping pregnancies and anembryonic fetuses showed statistically significant differences in the sex ratio from the control group consisting of medical abortions (1,11). Differences in the sex ratio were due to an increasingly lower proportion of embryos with karyotype 46.XY (relative to the expected one) among the fetuses with an increased severity of developmental disturbances. The statistical “chances ratio” index also provided evidence that embryos with the 46.XY karyotype had a higher propensity to produce a well-formed fetus as compared with the female embryos. We propose that the expression of genes of the maternal X-chromosome in XY embryos supports a more stable development during early embryogenesis as compared with XX embryos. In the latter case, normal development is coupled with the operation of an additional mechanism for compensation of the dose of X-linked genes. Operation of this mechanism increases the probability of disturbances in female embryos. A higher viability of XY embryos during the early stages of ontogenesis in man appears to explain their underrepresentation in samples of spontaneously aborted embryos and appears to be the major factor responsible for the deviation of the sex ratio from the theoretically expected value.     

Opposing effects of mortality factors on progeny operational sex ratio may thwart adaptive manipulation of primary sex ratio

Despite extensive research on mechanisms generating biases in sex ratios, the capacity of natural enemies to shift or further skew operational sex ratios following sex allocation and parental care remains largely unstudied in natural populations. Male cocoons of the sawfly Neodiprion abietis (Hymenoptera: Diprionidae) are consistently smaller than those of females, with very little overlap, and thus, we were able to use cocoon size to sex cocoons. We studied three consecutive cohorts of N. abietis in six forest stands to detect cocoon volume‐associated biases in the attack of predators, pathogens, and parasitoids and examine how the combined effect of natural enemies shapes the realized operational sex ratio. Neodiprion abietis mortality during the cocoon stage was sex‐biased, being 1.6 times greater for males than females. Greater net mortality in males occurred because male‐biased mortality caused by a pteromalid parasitic wasp and a baculovirus was greater and more skewed than female‐biased mortality caused by ichneumonid parasitic wasps. Variation in the susceptibility of each sex to each family of parasitoids was associated with differences in size and life histories of male and female hosts. A simulation based on the data indicated that shifts in the nature of differential mortality have different effects on the sex ratio and fitness of survivors. Because previous work has indicated that reduced host plant foliage quality induces female‐biased mortality in this species, bottom‐up and top‐down factors acting on populations can affect operational sex ratios in similar or opposite ways. Shifts in ecological conditions therefore have the potential to alter progeny fitness and produce extreme sex ratio skews, even in the absence of unbalanced sex allocation. This would limit the capacity of females to anticipate the operational sex ratio and reliably predict the reproductive success of each gender at sex allocation.     

Brood sex ratio variance,developmental mortality and virginity in a gregarious parasitoid wasp

Females of the parasitoid wasp Goniozus nephantidis paralyse host caterpillars and lay a clutch of up to 18 eggs onto the host integument. The known biology of G. nephantidis suggests that matings occur exclusively between siblings from the same brood. This leads to the prediction that brood sex ratios should be highly female-biased and have low variance. Sex ratios are indeed female-biased, with the mean proportion of males equal to 0.093. However, while sex ratio variance is significantly less than binomial, many broods contain no males at emergence. During development 28% of G. nephantidis offspring die. Male mortality offers a potential explanation for all-female (= virgin) broods. For the clutch sizes and mortality observed, theory predicts that <10% of females will emerge from all-female broods but the empirical value is much higher. The prediction that the prevalence of virginity decreases with increasing clutch size is, however, supported. We consider alternative explanations for the observed proportion of all-female broods, but this appears to be neither an artefact of the laboratory environment nor due to incorrect assumptions about G. nephantidis life history. Although its reproductive biology has been much investigated and its sex ratio matches some theoretical predictions, we conclude that a fuller understanding of G. nephantidis sex ratio requires a deeper knowledge of its field biology.     

Hatching sex ratio and sex specific chick mortality in common terns Sterna hirundo

Bias in sex ratios at hatching and sex specific post hatching mortality in size dimorphic species has been frequently detected, and is usually skewed towards the production and survival of the smaller sex. Since common terns Sterna hirundo show a limited sexual size dimorphism, with males being only about 1–6% larger than females in a few measurements, we would expect to find small or no differences in production and survival of sons and daughters. To test this prediction, we carried out a 2-year observational study on sex ratio variation in common terns at hatching and on sex specific post hatching mortality. Sons and daughters hatched from eggs of similar volume. Post hatching mortality was heavily influenced by hatching sequence. In addition, we detected a sex specific mortality bias towards sons. Overall, hatching sex ratio and sex specific mortality resulted in fledging sex ratios 8% biased towards females. Thus, other reasons than body size may be influencing the costs of rearing sons. Son mortality was not homogeneous between brood sizes, but greater for two-chick broods. Since adults rearing two-chick broods were younger, lighter and bred consistently later than those rearing three-chick broods, it is suggested that lower capacity of two-chick brood parents adversely affected offspring survival of sons. Though not significantly, two-chick broods tended to be female biased at hatching, perhaps to counteract the greater male-biased nestling mortality. Thus, population bias in secondary sex ratio is not limited to strongly size dimorphic species, but species with a slight sexual size dimorphism can also show sex ratio bias through a combination of differential production and mortality of sons and daughters.     

Parasitoid developmental mortality in the field: patterns, causes and consequences for sex ratio and virginity

1. Sex ratio theory predicts that developmental mortality can affect sex ratio optima under Local Mate Competition and also lead to 'virgin' broods containing only females with no sibling-mating opportunities on maturity. 2. Estimates of developmental mortality and its sex ratio effects have been laboratory based, and both models and laboratory studies have treated mortality as a phenomenon without identifying its biological causes. 3. We contribute a large set of field data on Metaphycus luteolus Timberlake (Hymenoptera: Encyrtidae), an endoparasitoid of soft scale insects (Hemiptera: Coccidae), which has sex allocation conditional on host quality and female-biased brood sex ratios. Developmental mortality within broods can be both assessed and attributed to distinct causes, including encapsulation by the host and larval-larval competition. 4. Thirty per cent of M. luteolus offspring die during development with 65% of this mortality because of encapsulation and 28% because of larval competition. The distributions of mortality overall and for each cause of mortality separately were overdispersed. 5. The probability of an individual being encapsulated increased with clutch size, while the probability of being killed by a brood mate declined with increasing clutch size and with increasing per capita availability of resources. 6. The sexual compositions of broods at emergence were influenced by both the degree and the type of mortality operating. At higher levels of mortality, single sex broods were more common and sex ratios were less precise. Overall, virginity was more prevalent than predicted and was more greatly affected by the occurrence of competition than by other sources of mortality, almost certainly because competition tended to eliminate males. 7. The reproductive and developmental biology of M. luteolus appears to be influenced by a complex interplay of maternal clutch size and sex allocation strategies, offspring-offspring developmental interactions, host defence mechanisms and postemergence mating behaviour. Despite the great sophistication of sex ratio theory, it has not yet evolved to the point where it is capable of considering all of these influences simultaneously.     

The influence of increased line-fishing mortality on the sex ratio and age of sex reversal of the venus tusk fish

The age of sex reversal of the venus tusk fish Choerodon venustus , caught by line fishing at various locations on the southern Great Barrier Reef, indicated that C. venustus is capable of modifying its life cycle in response to increased mortality. The evidence suggests Masthead Reef fish, which experience the highest mortality, underwent sex reversal at a smaller size and younger age than at the other sites. The largest female fish, sexually transitional fish and males were smaller at Masthead Reef than at the Swains Reefs or One Tree Reef at Masthead Reef. There was also considerable overlap in the size of males and females within the exploited populations indicating that sex reversal is not initiated at a particular length but may have a social cause. The sex ratio of fish was essentially the same for fish fully susceptible to line fishing in the Swains and Masthead samples. Circumstantial evidence suggested that the absence of large males in a population may initiate sex reversal, indicating the maintenance of a constant sex ratio may have a social basis.     

Demography of the human sex ratio in some Latin American countries, 1967-1986.

A sample based on hospital births recorded for the Latin American Collaborative Study on Congenital Malformations (ECLAMC) program was used in the present study to determine sex ratios for live births and for stillbirths. Sixty-four cities and 147 hospitals in 11 countries (Uruguay, Chile, Argentina, Brazil, Bolivia, Peru, Paraguay, Ecuador, Venezuela, Colombia, and Costa Rica) were included in the present analyses. The number of live births was 1,886,653 in the period 1967-1986, and the number of stillbirths was 24,818 in the period 1978-1986. The sex ratio for the total sample was 0.5112 for live births and 0.5477 for stillbirths. The sex ratio as a whole is decreasing with time in a parabolic fashion. Each country in our study behaved differently. Except for Peru and Uruguay, the countries experienced a significant decrease in the sex ratio after 1978 for live births; only Brazil did not show a temporal trend for the sex ratio for stillbirths.     

The secondary sex ratio in Israel: 1980-1995

This study deals with models and hypotheses that attempt to explain the underlying mechanisms determining the sex ratios at birth in human populations. Since the factors responsible are still questionable and research results are contradictory, we examine data available in Israel on the sex ratios at birth among two different sociodemographic groups, Jews and Moslems. Results suggest a difference between Jewish and Moslem patterns of secondary sex ratios with respect to parental age, education, and birth number. The difference may be described as a more regular and, by existing models, a more predictable pattern of secondary sex ratio among Moslems than among Jews. The possibility that Jewish religious laws play a role in this difference is discussed.     

Air pollution and daily mortality in London: 1987-92.

OBJECTIVE--To investigate whether outdoor air pollution levels in London influence daily mortality. DESIGN--Poisson regression analysis of daily counts of deaths, with adjustment for effects of secular trend, seasonal and other cyclical factors, day of the week, holidays, influenza epidemic, temperature, humidity, and autocorrelation, from April 1987 to March 1992. Pollution variables were particles (black smoke), sulphur dioxide, ozone, and nitrogen dioxide, lagged 0-3 days. SETTING--Greater London. OUTCOME MEASURES--Relative risk of death from all causes (excluding accidents), respiratory disease, and cardiovascular disease. RESULTS--Ozone levels (same day) were associated with a significant increase in all cause, cardiovascular, and respiratory mortality; the effects were greater in the warm seasons (April to September) and were independent of the effects of other pollutants. In the warm season an increase of the eight hour ozone concentration from the 10th to the 90th centile of the seasonal change (7-36 ppb) was associated with an increase of 3.5% (95% confidence interval 1.7 to 5.3), 3.6% (1.04 to 6.1), and 5.4% (0.4 to 10.7) in all cause, cardiovascular, and respiratory mortality respectively. Black smoke concentrations on the previous day were significantly associated with all cause mortality, and this effect was also greater in the warm season and was independent of the effects of other pollutants. For black smoke an increase from the 10th to 90th centile in the warm season (7-19 microg/m3) was associated with an increase of 2.5% (0.9 to 4.1) in all cause mortality. Significant but smaller and less consistent effects were also observed for nitrogen dioxide and sulphur dioxide. CONCLUSION--Daily variations in air pollution within the range currently occurring in London may have an adverse effect on daily mortality.     

ABO and Rh blood groups in relation to sex ratio, mean number and mortality of sibs.

Data are presented on the sex ratio, mean number and mortality of the sibs of 17,060 schoolchildren, and on the sex ratio and mean number of the sibs of 5,785 blood donors, in relation to the children's and donors' sex and ABO and Rh blood groups. The sex ratio is significantly higher for the sibs of AB + B than for those of A + O schoolboys, and for the sibs of Rh-negative than for those of Rh-positive male blood donors, but in both cases the mean number of sibs is exactly the same for the first-mentioned as for the second-mentioned category.