Gametic, morphometric, and physiological variables influencing clutch size in the Chilean oyster, Ostrea chilensis (Philippi, 1845)

A number of marine bivalve taxa, including species of the genus Ostrea, have adopted brooding of the young in the mantle cavity as a reproductive mechanism. In spite of the importance of brooding in the reproductive success of such species, little is known about the most important variables influencing the process, including those limiting clutch size. This study addresses the regulation of brood size in the hermaphroditic oyster Ostrea chilensis. During spawning, oysters released all their oocytes into the mantle cavity. No residual oocytes remained in the gonad, so a second spawning during the same brooding season was not possible. There was a weak correlation between the number of embryos incubated during the early phase of brooding and the dry tissue weight of the brooding oyster, and between the number incubated and the shell length of the brooding adult during the later phases of brooding. The number of embryos was also correlated with the area of the labial palps of the brooder during the later stages, suggesting that the loss of veligers observed at this time may be at least partially attributable to a limitation of space around the palps, which manipulate the larvae and with which the larvae are closely associated. The oxygen consumption rate of brooders incubating a normal clutch of embryos was not significantly different from that of oysters in which clutch size had been experimentally reduced by 50%. Experimental increase of the normal clutch size by 100% significantly increased the oxygen consumption of the brooder, suggesting that there is a physiological as well as a spatial limit to brood size. Thus the number of embryos brooded by an oyster is initially dependent on its production of oocytes and secondarily by the high metabolic costs of incubating large numbers of embryos. As development proceeds, space available for brooding apparently becomes a limiting factor as the larvae grow. The fate of the excess larvae is not known at present, but any larvae released prematurely cannot be competent to settle, since development is synchronous and there is a complete release of all pediveligers at the end of the brooding period.     

Ecological and phenological covariates of offspring sex ratio in barn swallows

Non-random sex allocation in relation to parental, ecological and phenological factors has been investigated in several correlational studies of birds, mostly based on few breeding seasons and relatively small sample sizes, which have led to different results. We investigated sex ratio of nestling barn swallows (Hirundo rustica) in relation to adult sex ratio, laying date, clutch size, colony size and meteorological conditions in a sample of 553 broods (>2200 nestlings) during 10 years. At the population level, nestling sex ratio varied among years and deviated from parity in two years. Sex ratio among adults did not predict offspring sex ratio in the current or the following year. At the within-family level, the proportion of sons increased with laying date in large clutches, did not vary among clutches of intermediate size, and tended to decline with laying date in small clutches. Large colonies harbored more sons. The proportion of males increased with temperature during laying whereas the effects of temperature during the pre- or post-laying periods and that of rainfall were non-significant. These patterns of variation of offspring sex ratio did not differ between years. Thus, we identified several potential causal sources of variation in barn swallow offspring sex ratio, including temporal, phenological and ecological factors. The observation of an association of offspring sex with temperature during laying is novel for birds and may be mediated by effects on maternal steroid hormones profile. The ecological and evolutionary implications of present findings are discussed in the light of adaptive sex allocation theory.     

Male reproductive skew, paternal relatedness, and female social relationships

Female social relationships among primates are thought to be shaped by socio-ecological factors and phylogenetic constraints. We suggest that patterns of paternal relatedness among females influence measures of social tolerance that have been used to classify species into different social relationship categories. As kin support and kin preference have only been measured for matrilineal kin and related individuals exchange less aggression and have a higher conciliatory tendency, the observed low nepotism levels and high tolerance levels may be an artifact of hidden paternal relatedness among the nonkin category. Using comparative data on macaques, we investigate this hypothesis using male reproductive skew as a proxy for paternal relatedness. Within the limitations of the study we show that populations classified as being less nepotistic, and more tolerant exhibit higher levels of reproductive skew. This first result and the reasoning behind may motivate future students of social relationships to take paternal relatedness into consideration. Potential implications of this finding if repeated with larger samples include that variation in aspects of macaque social relationships may be explained without considering phylogeny or the strength of between-group contest competition for food.     

The joint effects of kin, multilevel selection and indirect genetic effects on response to genetic selection

Kin and levels-of-selection models are common approaches for modelling social evolution. Indirect genetic effect (IGE) models represent a different approach, specifying social effects on trait values rather than fitness. We investigate the joint effect of relatedness, multilevel selection and IGEs on response to selection. We present a measure for the degree of multilevel selection, which is the natural partner of relatedness in expressions for response. Response depends on both relatedness and the degree of multilevel selection, rather than only one or the other factor. Moreover, response is symmetric in relatedness and the degree of multilevel selection, indicating that both factors have exactly the same effect. Without IGEs, the key parameter is the product of relatedness and the degree of multilevel selection. With IGEs, however, multilevel selection without relatedness can explain evolution of social traits. Thus, next to relatedness and multilevel selection, IGEs are a key element in the genetical theory of social evolution.     

TECHNICAL ADVANCES: A maximum-likelihood relatedness estimator allowing for negative relatedness values

Previously reported maximum-likelihood pairwise relatedness (r) estimator of Thompson and Milligan (M) was extended to allow for negative r estimates under the regression interpretation of r. This was achieved by establishing the equivalency of the likelihoods used in the kinship program and the likelihoods of Thompson. The new maximum-likelihood (ML) estimator was evaluated by Monte Carlo simulations. It was found that the new ML estimator became unbiased significantly faster compared to the original M estimator when the amount of genotype information was increased. The effects of allele frequency estimation errors on the new and existing relatedness estimators were also considered.     

STORM: software for testing hypotheses of relatedness and mating patterns

Storm is a software package that allows users to test a variety of hypotheses regarding patterns of relatedness and patterns of mate choice and/or mate compatibility within a population. These functions are based on four main calculations that can be conducted either independently or in the hypothesis-testing framework: internal relatedness; homozygosity by loci; pairwise relatedness; and a new metric called allele inheritance, which calculates the proportion of loci at which an offspring inherits a paternal allele different from that inherited from its mother. STORM allows users to test four hypotheses based on these calculations and Monte Carlo simulations: (i) are individuals within observed associations or groupings more/less related than expected; (ii) do observed offspring have more/less genetic variability (based on internal relatedness or homozygosity by loci) than expected from the gene pool; (iii) are observed mating pairs more/less related than expected if mating is random with respect to relatedness; and (iv) do observed offspring inherit paternal alleles different from those inherited from the mother more/less often than expected based on Mendelian inheritance.     

Genetic relatedness in groups is sex-specific and declines with age of helpers in a cooperatively breeding cichlid

Kin selection can explain the evolution of cooperative breeding and the distribution of relatives within a population may influence the benefits of cooperative behaviour. We provide genetic data on relatedness in the cooperatively breeding cichlid Neolamprologus pulcher. Helper to breeder relatedness decreased steeply with increasing helper age, particularly to the breeding males. Helper to helper relatedness was age‐assortative and also declined with age. These patterns of relatedness could be attributed to territory take‐overs by outsiders when breeders had disappeared (more in breeding males), between‐group dispersal of helpers and reproductive parasitism. In six of 31 groups females inherited the breeding position of their mother or sister. These matrilines were more likely to occur in large groups. We conclude that the relative fitness benefits of helping gained through kin selection vs. those gained through direct selection depend on helper age and sex.     

Different effects of age on reproductive performance in relation to breeding stage in Bull-headed Shrikes

The difference in the reproductive performance of males and females of the Bull-headed Shrike (Lanius bucephalus) according to age class, i.e. yearling and adult, was studied, and the age-related difference was examined according to parental feeding behavior. The clutch initiation date was not affected by the age class. Females that paired with an adult male laid more eggs per clutch than those paired with a yearling male. The age class of males affected the mass of nestlings at 6 days old, and the age class of females affected the mass of nestlings at 12 days old. The effects of the age of either parent independently were observed at different breeding stages. A change in the degree of nestling feeding peformed by the male and female parents occurred at some point between when the broods were 6 days and 12 days old. It is likely that this caused an effect of age at different stages of the breeding cycle. The effects of the age of the male parent are consistent with accounts of age-related reproduction in raptors where males provide resources to offspring. Individual improvements in foraging skills and/or courtship feeding rate are suggested to be possible explanations. Electronic Publication     

Variation in reproductive output of marine turtles

Most marine turtle species are non-annual breeders and show variation in both the number of eggs laid per clutch and the number of clutches laid in a season. Large levels of inter-annual variation in the number of nesting females have been well documented in green turtle nesting populations and may be linked to environmental conditions. Other species of marine turtle exhibit less variation in nesting numbers. This inter-specific difference is thought to be linked to trophic status. To examine whether individual reproductive output is more variable in the herbivorous green turtle (Chelonia mydas Linneaeus 1758) than the carnivorous loggerhead (Caretta caretta Linneaeus 1758), we examined the nesting of both species in Cyprus over nine seasons. Green turtles showed slower annual growth rates (0.11 cm year⁻¹ curved carapace length (CCL) and 0.27 cm year⁻¹ curved carapace width (CCW)) than loggerhead turtles (0.36 cm year⁻¹ CCL, 0.51 cm year⁻¹ CCW). CCL was highly correlated to mean clutch size in both green (R²=0.51) and loggerhead turtles (R²=0.61) and maximal clutch size of green turtles (R²=0.58). Larger females did not lay a greater number of clutches or have a shorter remigration interval than smaller females of either species. On average, the size of green turtle clutches increased and that of loggerhead turtles decreased as the season progressed. Individual green turtles, however, produced more eggs per clutch through the season to a maximum in the third or fourth clutch. In loggerhead turtles, clutches 1-4 were very similar in size but the fifth clutch was 38% smaller than the first. No individuals of either species were recorded laying more than five clutches. Green turtles may not be able to achieve their maximum reproductive output with respect to clutch size throughout the season, whereas only loggerhead turtles laying five clutches (n=5) appear to become resource depleted. Green turtles nesting in years when large numbers of nests were recorded laid a greater number of clutches than females nesting in years with lower levels of nesting.     

黑斑侧褶蛙的两性异形和雌性繁殖特征

测定了黑斑侧褶蛙成体的体长、体重、头长、头宽、眼径、鼓膜径、前肢长、后肢长等形态指标以及雌体的怀卵数量。黑斑侧褶蛙雌体的体长和体重显著大于雄体。其它局部形态特征指标与体长呈正相关,协方差分析表明,雄体的鼓膜径大于雌体,其余形态指标不存在明显的两性差异。黑斑侧褶蛙雌体怀卵数量与体长和体重皆成正相关关系,表明黑斑侧褶蛙通过增加个体大小增加繁殖输出。