Sex ratio and maternal investment in the multivoltine large carpenter bee Xylocopa sulcatipes (Apoideh: Anthophoridae)

Abstract. 1. Females of the multivoltine carpenter bee Xylocopa sulcutipes (Maa) (Hymenoptera: Anthophoridae) usually excavate a straight tunnel in dead twigs and mass provision a linear array of up to ten brood cells with pollen and nectar. An egg is deposited upon each food mass within one cell.
2. Female offspring generally receive a higher provisioning mass (0.180 ± 0.048 g) than males, a significant difference ( P > 0.001). There are, however, male larvae that receive as much food or more as their sisters or female larvae reared in another nest.
3. There is a close positive association between the size of a mother and the weight of provisions for individual daughters, but not for sons.
4. Female offspring are positioned in the innermost brood cells (Gositions 1, 2 and 3). The sex ratio of the outer cells is either significantly male biased (positions 4–6) or skewed towards males (positions 8 and 9). Positions 7 and 10 are in equilibrium.
5. Solitary females produce a significantly female biased sex ratio ( P < 0.01). Sex ratio in social nests is skewed toward females, but not significantly so ( P < 0.2). There is no significant difference between the sex ratio of solitary and social nests ( P = 0.361). The population sex ratio (pooled sex ratio of all broods produced) is significantly female biased ( P = 0.003).
6. Females kept in the laboratory produced female biased sex ratios whilst unmated females produced all-male broods indicating that insemination and ovarian development are not causally related.
7. The expected sex ratio (ESR) under equal investment, calculated as 1/CR (CR = mean male provision weight/mean female provision weight), is 137.5:117.5 (males:females), and differs significantly from that observed, 104:151 (males:females) ( P < 0.001). The 'Local Resource Enhlancement' hypothesis best explains the female biased sex ratio found in X.sulcatipes and its maintenance in the population.     

The Influence of Maternal Quality on Brood Sex Allocation in the Small Carpenter Bee,Ceratina calcarata

In the twig‐nesting carpenter bee, Ceratina calcarata, body size is an important component of maternal quality, smaller mothers producing significantly fewer and smaller offspring than larger mothers. As mothers precisely control the sex and size of each offspring, smaller mothers might compensate by preferentially allocating their investment towards sons. We investigated whether variation in maternal quality leads to variation in sex allocation patterns. At the population level, the numerical sex ratio was 57% male‐biased (1.31 M/F), but the investment between the sexes was balanced (1.02 M/F), because females are 38% larger than males (1.28 F/M). Maternal body size explained both sex allocation pattern and size variation among offspring: larger mothers invested more in individual progeny and produced more female offspring than smaller mothers. Maternal investment in offspring of both sexes decreased throughout the season, probably as a result of increasing maternal wear and age. The exception to this pattern was the curious production of dwarf females in the first two brood cell positions. We suggest that the sex ratio distribution reflects the maternal body size distribution and a constraint on small mothers to produce small broods. This leads to male‐biased allocation by small females, to which large mothers respond by biasing their allocation towards daughters.     

Testing the predictions of sex allocation hypotheses in dimorphic,cooperatively breeding riflemen

Evolutionary theory predicts that parents should invest equally in the two sexes. If one sex is more costly, a production bias is predicted in favour of the other. Two well‐studied causes of differential costs are size dimorphism, in which the larger sex should be more costly, and sex‐biased helping in cooperative breeders, in which the more helpful sex should be less costly because future helping “repays” some of its parents’ investment. We studied a bird species in which both processes should favor production of males. Female riflemen Acanthisitta chloris are larger than males, and we documented greater provisioning effort in more female‐biased broods indicating they are likely costlier to raise. Riflemen are also cooperative breeders, and males provide more help than females. Contrary to expectations, we observed no male bias in brood sex ratios, which did not differ significantly from parity. We tested whether the lack of a population‐wide pattern was a result of facultative sex allocation by individual females, but this hypothesis was not supported either. Our results show an absence of adaptive patterns despite a clear directional hypothesis derived from theory. This appears to be associated with a suboptimal female‐biased investment ratio. We conclude that predictions of adaptive sex allocation may falter because of mechanistic constraint, unrecognized costs and benefits, or weak selection.     

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.     

Female-biased operational sex ratio of sexual host fish in a gynogenetic complex of Carassius auratus

To study the coexistence of sexual and gynogenetic forms, we examined the population structure of a gynogenetic complex of the Japanese crucian carp, Carassius auratus Temminck et Schlegel, during the April–June reproductive season by collecting 1225 mature fish that migrated from Lake Suwa to a tributary river for spawning. There were more sexual fish (about 80%) than gynogenetic fish in this complex, and the operational sex ratio in the sexual form was female biased (males were about 20%). Mean standard length and body weight of sexual females were larger than those of sexual males. Sex ratio was male biased in smaller fish (standard length, <8.5 cm) but female biased in larger fish (standard length, ≥8.5 cm). We determined age by scale ring marks; the average age of sexual females was higher than that of males, but there was no significant difference in the average age between sexual and gynogenetic females. Sex ratio in the sexual form was more female biased for old than for young fish, and the mean size of sexual females was larger than that of males of the same age. The clear female-biased sex ratio and age difference between sexual females and males can be explained either by (1) higher mortality of males or by (2) female-biased sex allocation. The latter process reduces the disadvantage of sex and contributes to the coexistence of sexual and gynogenetic forms. Received: November 24, 2000 / Accepted: March 6, 2001     

Partial local mate competition and the sex ratio: A study on non-pollinating fig wasps

In many species, mating takes place in temporary patches where only a small number of females produce offspring. In this situation Local Mate Competition (LMC) theory predicts that the optimal sex ratio (defined as proportion males) should become increasingly female biased as the number of females contributing offspring to a patch decreases. However, in a large number of these species, some mating is also likely to occur away from the natal patch (termed partial LMC). In this case the degree of LMC is reduced, and theory predicts a relatively less female biased sex ratio. We tested these two predictions with field data from 17 species of New World non-pollinating fig wasps representing three genera. We present a model which suggests that the average number of females ovipositing in a fruit (i.e. patch) is positively correlated with the proportion of fruit of a given tree species in which that species of wasp occurs. Across species, the overall sex ratio was positively correlated with the proportion of fruit in which that species occurs. Furthermore, the males of some species are wingless, and in these species all mating must take place before females disperse from their natal fruit. In contrast, the males of other species are winged, and in these species mating may also take place away from the natal fruit. Species with winged males had less female biased sex ratios than species with wingless males that occurred in a similar proportion of fruit. Finally, the correlation between sex ratio and the proportion of fruit in which a species occurs was also observed within species when comparing between the fruit crops of different trees. This suggests that individual females facultatively adjust the sex ratio of their offspring in response to variable LMC.     

Operational sex ratio and resource defence as predictors of the mating system in European bitterling

Operational sex ratio (OSR), the ratio of sexually active males to fertilizable females in a population, plays a central role in the theory of mating systems by predicting that the intensity of male–male competition and the degree of sexual selection increases as the OSR becomes increasingly male biased. At high values of OSR, however, resource defence theory predicts the breakdown of territoriality and a shift towards scramble competition with a decrease in sexual selection. The direction that correlations between OSR and resource competition and variance in mating success will take depends on the biology of the species of interest. We investigated the effects of male population density and male‐biased operational sex ratio on male mating tactics shown by a freshwater fish, the European bitterling, Rhodeus sericeus . This species spawns inside living unioneid mussels. Large males defended territories, were aggressive towards conspecifics under equal sex ratios and monopolized pair spawnings with females. The mating tactic, however, changed at high male density where large males ceased to be territorial and instead competed with groups of smaller males to release sperm when females spawned. This change in male behaviour from pair to group spawning has two ramifications for sexual selection. The intensity of sexual selection and variance in male mating success decrease, and the form of sexual competition changes from resource‐ to sperm competition. Thus, the use of alternative mating tactics renders the OSR unable to predict the direction of resource competition and variance in male mating success at high densities.     

Seasonal variation in the sex allocation of a neotropical solitary bee

We carried out a field study on the life history and sex allocationof the ground-nesting solitary bee Diadasina distincta (Hymenoptera: Anthophoridae).This species is multivoltine, undergoing five generations a yearbetween February and September. The numerical sex ratio of thisspecies was female biased overall (approximately 38% males)and showed a strong and consistent seasonal pattern. The numericalsex ratio was extremely female biased (approximately 20% males)from February until May, and then slightly male biased (approximately60% males) from June until September. Females were 3.26 timesthe size of males, and so the overall investment ratio was female biasedthroughout the year. The overall female bias and seasonal variationin sex allocation is unlikely to be explained by models thatinvoke overlapping generations or competition between brothersfor mates (local mate competition). We suggest that a possibleexplanation for the female bias in the early part of the seasonis local resource enhancement (LRE): nesting near larger numbersof sisters reduces parasitism. LRE is likely to decrease in importancein the later part of the season, when the biased numerical and investmentratios may be explained by models in which male and female offspringgain different fitness returns from resources invested.     

Colony structure, provisioning and sex allocation in the sweat bee Halictus ligatus (Hymenoptera: Halictidae)

Population and colony-level sex allocation and nest productivity in the eusocial sweat bee Halictus ligatus Say were studied by excavating nests during one season. The emphasis was on measuring the provision masses, which differ in size and shape depending on the sex of the egg to be laid on them (male-producing provision masses are smaller and more or less round, whereas gyne-producing provision masses are larger and 'loaf-shaped). The primary aim of this study was to test theoretical predictions about female-bias of the sex ratio in the summer brood, both on the population level and on the colony level.
The overall sex ratio of the summer brood was moderately biased towards gynes. A significant positive correlation between the overall size of provision masses (as an estimate for the degree of female bias of the nest sex ratio) and the number of eusocial workers was found. This relationship further improved in partial analyses in which the provision mass weights were adjusted for sampling date, removing the effect of protandry. Foundress size, however, had no effect on the second brood provision masses and neither was there an effect of worker number on the size of gynes and males separately. In the first brood only the size of the foundress had a consistently positive effect on the size of the provision masses and on the size of the emerging daughter workers.
The observed increase of female bias in the nest sex ratio with increasing numbers of eusocial worker bees conforms to optimization predictions following from kin-selection theory.     

Local resource competition and sex ratio in the ant Cataglyphis cursor

The local resource competition (LRC) hypothesis predicts thatwherever philopatric offspring compete for resources with theirmothers, offspring sex ratios should be biased in favor of thedispersing sex. In ants, LRC is typically found in polygynous(multiple queen) species where foundation of new nests occursby budding, which results in a strong population structure anda male-biased population-wide sex ratio. However, under polygyny,the effect of LRC on sex allocation is often blurred by theeffect of lowered relatedness asymmetries among colony members.Moreover, environmental factors, such as the availability ofresources, have also been shown to deeply influence sex ratioin ants. We investigated sex allocation in the monogynous (singlequeen) ant Cataglyphis cursor, a species where colonies reproduceby budding and both male and female sexuals are produced throughparthenogenesis, so that between-colony variations in relatednessasymmetries should be reduced. Our results show that sex allocationin C. cursor is highly male biased both at the colony and populationlevels. Genetic analyses indicate a significant isolation-by-distancein the study population, consistent with limited dispersal offemales. As expected from asexual reproduction, only weak variationsin relatedness asymmetry of workers toward sexual offspringoccur across colonies, and they are not associated with colonysex ratio. Inconsistent with the predictions of the resourceavailability hypothesis, the male bias significantly increaseswith colony size, and investment in males, but not in females,is positively correlated with total investment in sexuals. Overall,our results are consistent with the predictions of the LRC hypothesisto account for sex ratio variation in this species.     

Sex ratio determination by queens and workers in the ant Pheidole desertorum

Because workers in colonies of eusocial Hymenoptera are more closely related to sisters than to brothers, theory predicts workers should bias investment in reproductive broods to favour reproductive females over males. However, conflict between queens and workers is predicted. Queens are equally related to daughters and sons, and should act to prevent workers from biasing investment. Previous study of the ant Pheidole desertorum showed that workers are nearly three times more closely related to reproductive females than males; however, the investment sex ratio is very near equal, consistent with substantial queen control of workers. Near-equal investment is produced by an equal frequency of colonies whose reproductive broods consist of only females (female specialists) and colonies whose reproductive broods consist of only males or whose sex ratios are extremely male biased (male specialists). Because natural selection should act on P. desertorum workers to bias investment in favour of reproductive females, why do workers in male-specialist colonies rear only (or mostly) males? We tested the hypothesis that queens prevent workers from rearing reproductive females by experimentally providing workers with immature reproductive broods of both sexes. Workers reared available reproductive females, while failing to rear available males. Worker preference for rearing reproductive females is consistent with queens preventing their occurrence in colonies of male specialists. These results provide evidence that queens and workers will act in opposition to determine the sex ratio, a fundamental prediction of queen-worker conflict theory. Copyright 2000 The Association for the Study of Animal Behaviour.     

Does constrained oviposition influence offspring sex ratio in the solitary parasitoid wasp Venturia canescens?

Abstract.  1. In haplodiploid organisms, virgin or sperm-depleted females can reproduce but are constrained to produce only male progeny. According to Godfray's constrained model, when p , the proportion of females constrained to produce only male progeny, is not null in a panmictic population, unconstrained females should bias their sex allocation towards females to compensate for the excess of males. These unconstrained females should be able to adjust the sex ratio in response to local variation of p .
2. In this paper an experimental approach is used to test the hypotheses of this model in the solitary endoparasitoid Venturia canescens under both field and laboratory conditions. Specifically, it is tested whether unconstrained females use their encounters with conspecifics (either male or female) to estimate p and then adjust their sex ratio accordingly.
3. As assumed by Godfray's model, constrained females actively search for host patches in the field and under laboratory conditions produce the same number of offspring during their lifetime as unconstrained females. As predicted by the model, unconstrained females produce a sex ratio biased towards females both in the laboratory and in the field.
4. The results show that this bias is not a response to encounters with conspecifics previous to oviposition. The hypothesis that the bias is due to differential mortality between sexes during ontogeny is also rejected. The proportions of constrained ovipositions estimated in two natural populations explain only a small fraction of the sex ratio bias observed in V. canescens.     

The effects of temperature and body size on the mating pattern of a gregariously nesting bee, Colletes cunicularius (Hymenoptera: Colletidae)

Abstract. 1. In a 3-year study of the solitary bee Colletes cunicularius L. in Sweden, average body size and population density fluctuated greatly between years.
2. In this protandrous population, females mated just once and the sex ratio was slightly male biased. Males were smaller than females.
3. Size assortative mating (homogamy), associated with an increase in population density during the central days of female emergence and mating, was observed in two out of three years. Homogamy was also observed in pairs with remating males.
4. Most of the mating males had emerged the day they mated, but 42% were older. We found no support for a general large-male mating advantage.
5. Weight of emerging females and mating males were negatively correlated with ground temperature, indicating thermoregulatory influence on the process of sexual selection in this species.     

Facultative Sex Allocation and Sex‐Specific Offspring Survival in Barrow's Goldeneyes

Sex allocation theory predicts that females should bias their reproductive investment towards the sex generating the greatest fitness returns. The fitness of male offspring is often more dependent upon maternal investment, and therefore, high‐quality mothers should invest in sons. However, the local resource competition hypothesis postulates that when offspring quality is determined by maternal quality or when nest site and maternal quality are related, high‐quality females should invest in the philopatric sex. Waterfowl – showing male‐biased size dimorphism but female‐biased philopatry – are ideal for differentiating between these alternatives. We utilized molecular sexing methods and high‐resolution maternity tests to study the occurrence and fitness consequences of facultative sex allocation in Barrow's goldeneyes (Bucephala islandica). We determined how female structural size, body condition, nest‐site safety and timing of reproduction affected sex allocation and offspring survival. We found that the overall sex ratio was unbiased, but in line with the local resource competition hypothesis, larger females produced female‐biased broods and their broods survived better than those of smaller females. This bias occurred despite male offspring being larger and tending to have lower post‐hatching survival. The species shows strong female breeding territoriality, so the benefit of inheriting maternal quality by philopatric daughters may exceed the potential mating benefit for sons of high‐quality females.     

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.     

Birth sex ratio in captive mammals: Patterns,biases, and the implications for management and conservation

Sex allocation theory predicts that a female should produce the offspring of the sex that most increases her own fitness. For polygynous species, this means that females in superior condition should bias offspring production toward the sex with greater variation in lifetime reproductive success, which is typically males. Captive mammal populations are generally kept in good nutritional condition with low levels of stress, and thus populations of polygynous species might be expected to have birth sex ratios biased toward males. Sex allocation theory also predicts that when competition reduces reproductive success of the mother, she should bias offspring toward whichever sex disperses. These predicted biases would have a large impact on captive breeding programs because unbalanced sex ratios may compromise use of limited space in zoos. We examined 66 species of mammals from three taxonomic orders (primates, ungulates, and carnivores) maintained in North American zoos for evidence of birth sex ratio bias. Contrary to our expectations, we found no evidence of bias toward male births in polygynous populations. We did find evidence that birth sex ratios of primates are male biased and that, within primates, offspring sex was biased toward the naturally dispersing sex. We also found that most species experienced long contiguous periods of at least 7 years with either male‐ or female‐biased sex ratios, owing in part to patterns of dispersal (for primates) and/or to stochastic causes. Population managers must be ready to compensate for significant biases in birth sex ratio based on dispersal and stochasticity. Zoo Biol 19:11–25, 2000. © 2000 Wiley‐Liss, Inc.     

Female body condition and brood sex ratio in Yellow-legged Gulls Larus cachinnans

In the Yellow-legged Gull Larus cachinnans , males are the larger sex, and show more reproductive variance than females. We predicted that the proportion of male chicks in a brood should increase with female body condition. We investigated brood sex ratio by using DNA markers taken from samples of hatchlings or dead embryos, and female body condition using plasma cholesterol concentration as a reliable indicator. The brood sex ratio of females in good condition was male biased and the sex ratio of females in poor condition was female biased. This relationship was also significant in those nests where all the eggs laid were sexed. Thus, manipulation of embryo mortality cannot explain the biases reported in this study, suggesting that the sex ratio of the eggs was biased prior to laying. These results confirm that sex-ratio manipulation in gulls operates under natural conditions, and supports earlier experimental findings.     

Secondary sex ratio and maternal dominance rank among wild yellow baboons (Papio cynocephalus) of Mikumi National Park,Tanzania

The Trivers and Willard model predicts that offspring of dominant mothers will be biased toward males and offspring of subordinate mothers towards females, whereas a local-resource-competition hypothesis predicts the reverse. Available data bearing upon these alternative predictions are inconsistent. It has been suggested that the local-resource-competition hypothesis will predominate when resources are scarce, and the Trivers-Willard hypothesis when resources are abundant. The relationship between maternal dominance and secondary sex ratio for 214 offspring of 61 females was examined using four troops of wild yellow baboons living in Mikumi National Park, where the population was increasing in a resource-rich habitat. For all troops combined, no significant relationship was found between offspring sex ratio and maternal rank. The four troops separately showed inconsistent trends, and in no case did the relationship reach conventional levels of statistical significance. In contrast, the local resource-competition hypothesis was supported by a study of yellow baboons done with one troop and 80 offspring in Amboseli National Park, where a massive population decline had occurred. The contrasting Amboseli and Mikumi results may be due to differences in resource-competition at the two study sites, or to stochastic variation. © 1992 Wiley-Liss, Inc.     

The sex ratio in ectoparasitic insects

Abstract. 1. The sex ratio in 379 collections, each consisting of more than 100 individuals, of 250 different species of ectoparasite belonging to seven orders of insects is recorded. The data are gathered from a broad survey of published literature.
2. In general, ectoparasitic insects emerge as adults in equal numbers of each sex, although there are certain exceptions.
3. The sex ratio in natural populations may alter with numerous factors, but in general it is unbalanced, females usually predominating. Out of 359 collections from the host's body or home, the sex ratio of 30% showed no significant departure from unity, 63% contained significantly more females, and 7% significantly more males.
4. These imbalances could be due to sampling methods but this is rare. Generally they are due to one sex, usually the male, being shorter lived than the other. This may be because males are often more active and smaller than females, and thus more likely to be separated from the host's body or home, succumb to host predation, or be killed by adverse climatic or nutritional conditions.     

Female size and fitness in the leaf-cutter bee Megachile apicalis

1. The effects of female size on fitness of the leaf-cutter bee Megachile apicalis Spinola (Hymenoptera: Megachilidae) were examined using artificial nesting sites in the field.
2. Although female size had no significant effect on female longevity or sex ratio of progeny, it did have a significant effect on fecundity; larger females attained greater realized fecundity than did smaller females.
3. This significant effect of female size on fecundity occurred because larger females produced cells faster than did smaller females.
4. Female size also had a significant effect on egg size; larger females laid larger eggs than did smaller females.
5. Female size had a significant effect on the size of investment in each progeny. The size of investment estimated by brood cell weight was greater for larger females than for smaller females. This pattern was largely absent, however, when the size of investment was estimated by adult progeny weight.
6. Female size had a significant effect on nest usurpation behaviour; larger females had a higher capacity to usurp nests than did smaller females.