Variably male-biased sex ratio in a marine bird with females larger than males

When the costs of rearing males and females differ progeny sex ratios are expected to be biased toward the less expensive sex. Blue-footed booby (Sula nebouxii) females are larger and roughly 32% heavier than males, thus presumably more costly to rear. We recorded hatching and fledging sex ratios in 1989, and fledging sex ratios during the next 5 years. In 1989, the sample of 751 chicks showed male bias at hatching (56%) and at fledging (57% at ˜90 days). Fledging sex ratios during the five subsequent reproductive seasons were at unity (1 year) or male-biased, varying from 56% to 70%. Male bias was greater during years when mean sea surface temperature was warmer and food was presumably in short supply. During two warm-water years (only) fledging sex ratio varied with hatching date. Proportions of male fledglings increased with date from 0.48 to 0.73 in 1994, and from 0.33 to 0.79 in 1995. Similar results were obtained when the analysis was repeated using only broods with no nestling mortality, suggesting that the overall increase in the proportion of males over the season was the result of sex ratio adjustments at hatching. The male-biased sex ratio, and the increased male bias during poor breeding conditions supports the idea that daughters may be more costly than sons, and that their relative cost increases in poor conditions. Received: 3 February 1998 / Accepted: 12 September 1998     

Reversed sexual size dimorphism in microtines: Are females larger than males or are males smaller than females?

Summary We analysed sexual size dimorphism for 21 populations of microtine rodents. Female to male size ratio varied considerably among populations from females significantly larger than males (ratio=1.18) to males larger than females (ratio=0.78). In a multiple regression analysis female to male home range size ratio explained 94% of the total variation in body size dimorphism and was the only one of eight independent variables that was selected in a stepwise regression procedure. When females are the larger sex, males have home range sizes much larger than females. We suggest that the relationship between home range size ratio and body weight size dimorphism reflects different selection pressures on males and females in competition for resources and mates.     

Persistence of an extreme male-biased adult sex ratio in a natural population of polyandrous bird

In a number of insects, fishes and birds, the conventional sex roles are reversed: males are the main care provider, whereas females focus on matings. The reversal of typical sex roles is an evolutionary puzzle, because it challenges the foundations of sex roles, sexual selection and parental investment theory. Recent theoretical models predict that biased parental care may be a response to biased adult sex ratios (ASRs). However, estimating ASR is challenging in natural populations, because males and females often have different detectabilities. Here, we use demographic modelling with field data from 2101 individuals, including 579 molecularly sexed offspring, to provide evidence that ASR is strongly male biased in a polyandrous bird with male-biased care. The model predicts 6.1 times more adult males than females (ASR=0.860, proportion of males) in the Kentish plover Charadrius alexandrinus. The extreme male bias is consistent between years and concordant with experimental results showing strongly biased mating opportunity towards females. Based on these results, we conjecture that parental sex-role reversal may occur in populations that exhibit extreme male-biased ASR.     

Temperature-dependent sex ratio in a bird

To our knowledge, there is, so far, no evidence that incubation temperature can affect sex ratios in birds, although this is common in reptiles. Here, we show that incubation temperature does affect sex ratios in megapodes, which are exceptional among birds because they use environmental heat sources for incubation. In the Australian brush-turkey Alectura lathami, a mound-building megapode, more males hatch at low incubation temperatures and more females hatch at high temperatures, whereas the proportion is 1:1 at the average temperature found in natural mounds. Chicks from lower temperatures weigh less, which probably affects offspring survival, but are not smaller. Megapodes possess heteromorphic sex chromosomes like other birds, which eliminates temperature-dependent sex determination, as described for reptiles, as the mechanism behind the skewed sex ratios at high and low temperatures. Instead, our data suggest a sex-biased temperature-sensitive embryo mortality because mortality was greater at the lower and higher temperatures, and minimal at the middle temperature where the sex ratio was 1:1.     

Unusual allometry for sexual size dimorphism in a cichlid where males are extremely larger than females

When males are the larger sex, a positive allometric relationship between male and female sizes is often found across populations of a single species (i.e. Rensch’s rule). This pattern is typically explained by a sexual selection pressure on males. Here, we report that the allometric relationship was negative across populations of a shell-brooding cichlid fish Lamprologus callipterus, although males are extremely larger than females. Male L. callipterus collect and defend empty snail shells in each of which a female breeds. We found that, across six populations, male and female sizes are positively correlated with not only sexual and fecundity selection indices, but also with shell sizes. Given their different reproductive behaviours, these correlations mean that males are required to be more powerful, and thus larger, to transport larger shells, while female bodies are reduced to the shell size to enable them to enter the shells. Among the three size selections (sexual selection, fecundity selection and shell size), shell size explained the allometry, suggesting that females are more strongly subject to size selection associated with shell size availability than males. However, the allometry was violated when considering an additional population where size-selection regimes of males differed from that of other populations. Therefore, sexual size allometry will be violated by body size divergence induced by multiple selection regimes.     

Predictable males and unpredictable females: sex difference in repeatability of parental care in a wild bird population

Repeatability of parental care, let alone heritability of care, has been rarely measured, although there has been much research linking sexual selection to male parental care and also examining biparental care in relation to game theory models. We investigated within- and between-year repeatabilities of incubation and nestling provisioning and how these two types of parental care were related in a sexually dimorphic species, the house sparrow, Passer domesticus. We found that between- and within-year repeatabilities of feeding rate were high in males and low to moderate in females, but that between- and within-year repeatabilities of incubation time were low to moderate in both sexes. Interestingly, the amount of time during which neither sex incubated significantly predicted the subsequent male feeding rate but not the female feeding rate. Our results suggest a need for a new theoretical framework that encompasses variation in the predictability and plasticity of parental investment by individuals.     

Sex-biased environmental sensitivity: natural and experimental evidence from a bird species with larger females

The larger sex is often more vulnerable, in terms of developmentand survival, to poor conditions during early life. Differentialvulnerability has implications for parental investment strategiessuch as sex ratio theory. When males are larger, it is not possibleto separate the effects of larger size per se and other aspectsof the male phenotype on vulnerability. Furthermore, offspringcompetition might favor the larger sex and thereby mask intrinsic,size-related effects. We studied sex-specific mortality in abird species with reversed size dimorphism, the great skua Stercorariusskua, under natural and experimentally created poor conditions.Small eggs from extended laying sequences were used to createpoor early conditions for the offspring, which were raised assingletons. Daughters had a lower survival in all treatmentgroups. Survival in natural broods was additionally affectedby hatch date and position. Hatch weight was not different forsons and daughters but was lower in experimental than in naturalnests. In natural nests, daughters fledged 10% heavier thansons, but in experimental nests, they did not reach a highermass. The average survival difference between sons and daughterswas not increased in experimental broods. However, hatch weighthad a strong sex-specific effect. Very light females never survived,and survival probability of daughters increased with increasinghatch weight. By contrast, survival of sons over the same rangeof hatch weights was not related to weight. These findings supportthe hypothesis that larger (final) size per se is related tosex-specific offspring vulnerability during early life.     

Brood sex ratio variation in a cooperatively breeding bird

In cooperatively breeding species, the fitness consequences of producing sons or daughters depend upon the fitness impacts of positive (repayment hypothesis) and negative (local competition hypothesis) social interactions among relatives. In this study, we examine brood sex allocation in relation to the predictions of both the repayment and the local competition hypotheses in the cooperatively breeding long-tailed tit Aegithalos caudatus. At the population level, we found that annual brood sex ratio was negatively related to the number of male survivors across years, as predicted by the local competition hypothesis. At an individual level, in contrast to predictions of the repayment hypothesis, there was no evidence for facultative control of brood sex ratio. However, immigrant females produced a greater proportion of sons than resident females, a result consistent with both hypotheses. We conclude that female long-tailed tits make adaptive decisions about brood sex allocation.     

Primary sex ratio of 125 males to 100 females? Analysis of an artifact

A longitudinal study sponsored by the German Research Society has focused on pregnancy, attendant complications, and resulting fetal and infant development in a total of 13,834 cases from 42 clinics. Data from this project on abortions, miscarriages and stillbirths are evaluated for their respective sex ratios and by several diagnostic procedures, in an attempt to recalculate the primary sex ratio. It is shown that karyotype analysis in the representative sample leads to an estimated primary sex ratio of I to 1, rather than suggesting an excess of male conceptions as formerly believed.     

Reproductive success in reindeer males in a herd with varying sex ratio

In polygynous species, male reproductive success is often correlated with dominance status of individual males and sex ratio in the population. Reindeer, Rangifer tarandus, is a polygynous species, and here we compared the variation in male reproductive success and dominance status during two successive years in a herd with a male:female sex ratio of 1:7 and 1:3. Copulations were recorded, together with data on male dominance hierarchy and size of mating groups. Male reproductive success was estimated by paternity analysis of calves using microsatellite DNA markers. The distribution of reproductive success among the males was highly skewed for both years with the most dominant male also being the most successful. The largest mating group was established in the herd with the least skew in sex ratio. In this herd some of the adult males present were less reproductively successful than some of the more subordinate younger males. Estimates of the mating group size of males, correcting for dominance status when more than one male is present in the groups, gave good prediction of individual males' reproductive success.     

Differential allocation in a lekking bird: females lay larger eggs and are more likely to have male chicks when they mate with less related males

The differential allocation hypothesis predicts increased investment in offspring when females mate with high-quality males. Few studies have tested whether investment varies with mate relatedness, despite evidence that non-additive gene action influences mate and offspring genetic quality. We tested whether female lekking lance-tailed manakins (Chiroxiphia lanceolata) adjust offspring sex and egg volume in response to mate attractiveness (annual reproductive success, ARS), heterozygosity and relatedness. Across 968 offspring, the probability of being male decreased with increasing parental relatedness but not father ARS or heterozygosity. This correlation tended to diminish with increasing lay-date. Across 162 offspring, egg volume correlated negatively with parental relatedness and varied with lay-date, but was unrelated to father ARS or heterozygosity. Offspring sex and egg size were unrelated to maternal age. Comparisons of maternal half-siblings in broods with no mortality produced similar results, indicating differential allocation rather than covariation between female quality and relatedness or sex-specific inbreeding depression in survival. As males suffer greater inbreeding depression, overproducing females after mating with related males may reduce fitness costs of inbreeding in a system with no inbreeding avoidance, while biasing the sex of outbred offspring towards males may maximize fitness via increased mating success of outbred sons.     

Sex-different response in growth traits to resource heterogeneity explains male-biased sex ratio

In dioecious plants, differences in growth traits between sexes in a response to micro-environmental heterogeneity may affect sex ratio bias and spatial distributions. Here, we examined sex ratios, stem growth traits and spatial distribution patterns in the dioecious clonal shrub Aucuba japonica var. borealis, in stands with varying light intensities. We found that male stems were significantly more decumbent (lower height/length ratio) but female stems were upright (higher height/length ratio). Moreover, we found sex-different response in stem density (no. of stems per unit area) along a light intensity gradient; in males the stem density increased with increases in canopy openness, but not in females. The higher sensitivity of males in increasing stem density to light intensity correlated with male-biased sex ratio; fine-scale sex ratio was strongly male-biased as canopy openness increased. There were also differences between sexes in spatial distributions of stems. Spatial segregation of sexes and male patches occupying larger areas than female patches might result from vigorous growth of males under well-lit environments. In summary, females and males showed different growth responses to environmental variation, and this seemed to be one of possible causes for the sex-differential spatial distributions and locally biased sex ratios.     

Why do females care more than males?

Females tend to provide more parental care than males. Previous efforts to account for this have been confused because it is difficult to express the costs of care for males and females in the same currency. Here I propose a null model that does so, using the Fisherian constraint that total male and female reproduction must be equal. The model shows that, contrary to a number of recent analyses, lower probability of parentage for males does tend to make males less likely than females to provide care. It also shows how sexual selection stemming from premating asymmetries in investment promotes similar post-mating asymmetries.     

Reduced sexual size dimorphism in a pipefish population where males do not prefer larger females

Within a species' distribution, populations are often exposed to diverse environments and may thus experience different sources of both natural and sexual selection. These differences are likely to impact the balance between costs and benefits to individuals seeking reproduction, thus entailing evolutionary repercussions. Here, we look into an unusual population (Baltic Sea) of the broadnosed pipefish, Syngnathus typhle, where males do not seem to select females based on size and hypothesize that this pattern may derive from a reduction in direct benefits to the male. We further hypothesize that if larger females do not persistently secure a higher reproductive success, either through pre‐ or postcopulatory sexual selection, a decrease in sexual size dimorphism in the Baltic population should be apparent, especially when contrasted with a well‐studied population, inhabiting similar latitudes (Swedish west coast), where males prefer larger females. We found that, in the Baltic population, variation in female quality is low. We were unable to find differences in abortion rates or protein concentration in oocytes produced by females of contrasting sizes. Direct benefits from mating with large partners seem, thus, reduced in the Baltic population. We also found no evidence of any postcopulatory mechanism that could favor larger mothers as embryo development was unrelated to female size. While female size can still be selected through intrasexual competition or fecundity selection, the pressure for large female body size seems to be lower in the Baltic. Accordingly, we found a noticeable decrease in sexual size dimorphism in the Baltic population. We conclude that, although far from negating the significance of other selective processes, sexual selection seems to have a decisive role in supporting pipefish sexual size asymmetries.     

Offspring sex ratio allocation in the parasitic jaeger: selection for pale females and melanic males?

The maintenance of plumage color polymorphism in the parasiticjaeger (Stercorarius parasiticus) is still not well understood.Earlier studies indicated that selection may favor pale femalesand melanic males. If so, females would maximize their fitness,producing pale female and melanic male offspring. We thereforepredicted that females might bias their offspring sex ratiotoward daughters in pale pairs and toward sons in melanic pairs.Females might also choose to mate assortatively in relationto plumage color, thereby maximizing the probability of producingeither pale or melanic offspring. Because females are largerthan males, differential rearing costs may affect the offspringsex ratio independent of parental plumage color. We examinedoffspring sex ratio allocation, breeding variables indicativeof parental quality, and mating pattern in relation to plumagecolor in a colony of parasitic jaegers in northern Norway. Jaegerstended to mate assortatively in relation to plumage color. Thereproductive performance declined with season, and matched pairsappeared to be of lower quality than mixed pairs. The proportionof male offspring increased with hatching date in matched paleand mixed pairs, whereas the situation was reversed in matchedmelanic pairs. Matched pale pairs produced an overall surplusof favorable pale but costly daughters despite their lower quality,while melanic pairs produced a surplus of favorable melanicsons. However, differential offspring rearing costs and parentalrearing capacity may have additionally affected the realizedoffspring sex ratio. Mixed pairs producing an overall surplusof pale and melanic daughters allocated their resources accordingto differential rearing costs and parental quality only. Wesuggest that both strategies of sex ratio allocation togetherwith differences in reproductive success in matched versus mixedpairs may have a balancing effect on the mating pattern betweenplumage morphs and may contribute to the maintenance of thecolor polymorphism in this species.     

Adaptive secondary sex ratio adjustments via sex-specific infanticide in a bird

Infanticide is easiest to understand when it involves killing the offspring of others [1], but a parent may also kill its own offspring if the sacrifice of currently dependent young leads to higher survival of brood mates [2] or an improvement in the parent's likely future reproduction [3]. However, sex-specific infanticide by parents of their own offspring, although occurring in some human societies [4], is rare across species. Its rarity may be because killing one sex combines wasted parental effort with consequent biases in population sex ratios that are detrimental for the fitness of the overproduced sex [5-7]. We show that killing male offspring can be advantageous to Eclectus parrot (Eclectus roratus) mothers even though frequency-dependent selection then elevates the reproductive value of sons above that of daughters. In poorer-quality nest hollows, broods with a single female nestling had higher reproductive value than broods in which the female had a younger brother. Our data demonstrate frequent targeted removal of male nestlings within 3 days of hatching in these specific brood types and nesting conditions. The ability of Eclectus parrots to perceive the sex of their offspring relatively early may favor decisions to kill one sex before further investment in parental care.     

Surface electromyography reveals males have a slower patellar reflex than females

In humans the cross sectional area of spinal motor neurons at L3 is larger in males than in females. Since these contribute to the control of the quadriceps femoris muscle group and are involved in the patellar reflex (PR), gender differences in the PR are expected. We have investigated this possibility using a group of 28 young subjects (14 male and 14 female) aged 20–22 years. The PR was quantified by the muscle compound action potential (MCAP) from the surface electromyogram (sEMG) of the vastus lateralis muscle. We found that the PR latency in females (17 ± 0.19 ms), was significantly (p < 0.001) faster than in males (21 ± 0.37 ms). This 4 ms difference in latency could not be ascribed to differences in stature or thigh length. In conclusion, for the age range tested females posses a significantly faster patellar reflex than males. We suggest that the slower PR latency of male subjects may arise in part from their larger α-motorneurons: such that longer integration times are required for the summation of postsynaptic excitation to be sufficient to excite α-motorneurons.     

Maternal investment in a bee species with facultative nest guarding and males heavier than females

1. Maternal investment can be influenced by several factors, especially maternal quality and possibilities for future reproduction. Mass provisioning Hymenoptera are an excellent group for measuring maternal investment because mothers distribute food sources to each brood cell for each offspring separately. Generally in aculeate Hymenoptera, larger females produce larger offspring and invest more in female offspring than in male offspring. 2. This study investigated patterns of maternal investment in Ceratina chalcites, which has an uncommon type of sexual size dimorphism in Hymenoptera: on average, males are heavier than females. It was found that larger females produce a significantly higher proportion of male offspring, as males are the costlier sex in this species. 3. Facultative nest guarding by females was observed. Females can guard offspring until adulthood, as is typical for bees of genus Ceratina (34.43% of nests); however, in the majority of cases (65.56% of nests), females plug and abandon the nest. Significant differences were found in the amount of investment between guarded and unguarded nests. Guarded nests had a greater number of provisioned brood cells and a higher proportion of male offspring. It is suggested that mothers have two facultative strategies – either she makes a large investment in the offspring of one nest or she abandons the first nest and carries out a second nesting elsewhere.     

Woody elements benefit bird diversity to a larger extent than semi-natural grasslands in cereal-dominated landscapes

Increasing landscape complexity can mitigate negative effects of agricultural intensification on biodiversity by offering resources complementary to those provided in arable fields. In particular, grazed semi-natural grasslands and woody elements support farmland birds, but little is known about their relative effects on bird diversity and community composition. In addition, the relative importance of local habitat versus landscape composition remains unclear. We investigated how the presence of semi-natural grasslands, the number of woody elements and the composition of the wider agricultural landscape affect bird species richness, true diversity (exponential Shannon diversity) and species composition. Bird communities were surveyed four times on 16 paired transects of 250 m each with 8 transects placed between a crop field and a semi-natural grassland and 8 transects between two crop fields with no semi-natural grasslands in the vicinity. The number of woody elements around transects was selected as an important predictor in all models, having a positive effect on species richness and true diversity, while the local presence of semi-natural grasslands was not selected in the best models. However, species richness and true diversity increased with increasing cover of ley and semi-natural grasslands, whereas species composition was modified by the coverage of winter wheat at the landscape scale. Furthermore, bird species richness, true diversity and species composition differed between sampling dates. As bird diversity benefited from woody elements, rather than from the local presence of semi-natural grasslands as such, it is important to maintain woody structures in farmland. However, the positive effect of grassland at the landscape scale highlights the importance of habitat variability at multiple scales. Because species richness and true diversity were affected by different landscape components compared to species composition, a mosaic of land-use types is needed to achieve multiple conservation goals across agricultural landscapes.