Local Mating,Dispersal and Sex Ratio in a Gregarious Parasitoid Wasp

Parasitoid sex ratios can be greatly influenced by mating and dispersal behaviour. Many sex ratio models assume that mating is strictly local (only mated females disperse from the natal patch) and that a single male is sufficient to inseminate all females in a brood. Bethylids (aculeate parasitoids) have been used to test predictions of these models, but less attention has been paid to testing their underlying assumptions. We investigated the timing of eclosion, mating and dispersal in mixed-sex and single-sex broods of the bethylid wasp Goniozus nephantidis. In mixed-sex broods, almost all females mate before dispersal and a single male is sufficient to inseminate virtually all females, even when brood sizes are large. Males disperse from both mixed-sex and all-male broods, but males in all-male broods disperse more slowly. Virgin females disperse from all-female broods, which are common. Virgin females can produce a brood, mate with their own sons and subsequently produce mixed-sex broods, but their success rate is very low. Virgin females could potentially circumvent sex allocation constraints by superparasitizing mixed-sex broods, but when presented with hosts bearing mixed-sex broods they destroy all members of the initial brood before ovipositing. Because of the high prevalence of single-sex broods and dispersal of both sexes, the mating structure of G. nephantidis is unlikely to conform to the assumption of strict local mating.     

Unisexual broods and sex ratios in a polyembryonic encyrtid parasitoid (Copidosoma sp.: Hymenoptera)

Summary We examined brood sex ratios of an undetermined species of Copidosoma. Most broods (65%) were unisexual, with a greater proportion of female broods. Some mixed broods contained extremely small proportions of wasps of the opposite sex. Our results imply that the female-biased sex rations in this species cannot be explained by local mate competition theory.     

Co‐foundress confinement elicits kinship effects in a naturally sub‐social parasitoid

Kinship among interacting individuals is often associated with sociality and also with sex ratio effects. Parasitoids in the bethylid genus Goniozus are sub‐social, with single foundress females exhibiting post‐ovipositional maternal care via short‐term aggressive host and brood defence against conspecific females. Due to local mate competition (LMC) and broods normally being produced by a single foundress, sex ratios are female‐biased. Contests between adult females are, however, not normally fatal, and aggression is reduced when competing females are kin, raising the possibility of multi‐foundress reproduction on some hosts. Here, we screen for further life‐history effects of kinship by varying the numbers and relatedness of foundresses confined together with a host resource and also by varying the size of host. We confined groups of 1–8 Goniozus nephantidis females together with a host for 5+ days. Multi‐foundress groups were either all siblings or all nonsiblings. Our chief expectations included that competition for resources would be more intense among larger foundress groups but diminished by both larger host size and closer foundress relatedness, affecting both foundress mortality and reproductive output. From classical LMC theory, we expected that offspring group sex ratios would be less female‐biased when there were more foundresses, and from extended LMC theory, we expected that sex ratios would be more female‐biased when foundresses were close kin. We found that confinement led to the death of some females (11% overall) but only when host resources were most limiting. Mortality of foundresses was less common when foundresses were siblings. Developmental mortality among offspring was considerably higher in multi‐foundress clutches but was unaffected by foundress relatedness. Groups of sibling foundresses collectively produced similar numbers of offspring to nonsibling groups. There was little advantage for individual females to reproduce in multi‐foundress groups: single foundresses suppressed even the largest hosts presented and had the highest per capita production of adult offspring. Despite single foundress reproduction being the norm, G. nephantidis females in multi‐foundress groups appear to attune sex allocation according to both foundress number and foundress relatedness: broods produced by sibling foundresses had sex ratios similar to broods produced by single foundresses (ca. 11% males), whereas the sex ratios of broods produced by nonsibling females were approximately 20% higher and broadly increased with foundress number. We conclude that relatedness and host size may combine to reduce selection against communal reproduction on hosts and that, unlike other studied parasitoids, G. nephantidis sex ratios conform to predictions of both classical and extended LMC theories.     

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.     

Mating system and sex allocation in the gregarious parasitoid Cotesia glomerata

The gregarious parasitoid Cotesia glomerata (L.) is often presumed to possess the characteristic attributes of a species that manifests local mate competition (LMC), as it commonly produces female-biased broods. However, our field surveys of sex ratio and laboratory observations of adult behaviour showed that this species is subject to partial local mate competition caused by natal dispersal. On average, 30% of males left their natal patch before mating, with the proportion of dispersing males increasing with an increase in the patch's sex ratio (i.e. proportion of males). Over 50% of females left their natal patch before mating, and only 27.5% of females mated with males emerging from the same natal patch. Although females showed no preference between males that were and were not their siblings, broods from females that mated with siblings had a significantly higher mean brood sex ratio (0.56) than broods from females that mated with nonsiblings (0.39). Furthermore, brood sex ratios increased as inbreeding was intensified over four generations. A field population of this wasp had a mean brood sex ratio of 0.35 over 3 years, which conformed well to the evolutionarily stable strategy sex ratio (r=0.34) predicted by Taylor's partial sibmating model for haplodiploid species. These results suggest that the sex allocation strategy of C. glomerata is based on both partial local mate competition in males and inbreeding avoidance in females. In turn, this mating system plays a role in the evolution of natal dispersal behaviour in this species.Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Brood sex ratio in the Kentish plover

How and why do the mating opportunities of males and femalesdiffer in natural population of animals? Previously we showedthat females have higher mating opportunities than males inthe Kentish plover Charadrius alexandrinus. Both parents incubatethe eggs, and males provide more brood care than females; thusit is not obvious why the females find new mates sooner thanthe males. In this study we investigated whether the sex-biasedmating opportunities stem from biased offspring sex ratios.We determined the sex of newly hatched, precocial chicks usingCHD gene markers. Among fully sexed broods, 0.461 ± 0.024(SE) of chicks (454 chicks in 158 broods) were male, and thissex ratio was not significantly different from unity. The proportionof males at hatching decreased significantly over the breedingseason, which occurred consistently in all 3 years of the study.Large chicks were more likely to be males than females. Neitherparental age nor body size of male and female parents was relatedto brood sex ratio. We also sexed a number of chicks that werecaught after they left their nest (range of estimated ages 0–17days) and found that the proportion of males increased withbrood age. This relationship remained highly significant whencontrolling statistically for hatching date. As brood size decreaseddue to mortality after the chicks left their nest, these resultssuggest that the mortality of daughters was higher than thatof the sons shortly after hatching. Taken together, our resultsshow that the female-biased mating opportunities in the Kentishplover are not due to biased brood sex ratio at hatching but,at least in part, are due to female-biased chick mortality soonafter hatching.     

Sex Ratios in a Socially Parasitic Bee and Implications for Host-Parasite Interactions

Obligate social parasites of Hymenoptera, known as inquilines, have received enormous attention due to the elaborate adaptations they exhibit for exploiting their hosts, and because they have frequently been used to infer sympatric speciation. Their population biology can be difficult to infer as they are both rare and difficult to extract from host nests. Sex allocation has been studied for very few inquilines of social Hymenoptera. Here we report sex ratio patterns in the allodapine bee Inquilina schwarzi, which is an obligate social parasite of another allodapine, Exoneura robusta. We show that the sex ratio of this inquiline varies with its brood number, it is female-biased in the smallest broods, but becomes more even in larger broods, where the population-wide sex ratio is close to parity. We argue that this pattern of bias is consistent with local resource competition, where inquiline females compete to inherit their natal colony. We also argue that extremely female-biased sex ratios of the host species, combined with overall sex ratio parity in the parasite, may help ameliorate disparity in effective population sizes between these two species which are locked in an evolutionary arms race.     

Pair bond duration influences paternal provisioning and the primary sex ratio of brown thornbill broods

Parents should vary their level of investment in sons and daughters in response to the fitness costs and benefits accrued through male and female offspring. I investigated brood sex ratio biases and parental provisioning behaviour in the brown thornbill, Acanthiza pusilla, a sexually dimorphic Australian passserine. Parents delivered more food to male-biased than female-biased broods. However, factors determining parental provisioning rates differed between the sexes. Female provisioning rates were related to brood sex ratio in both natural and experimental broods with manipulated sex ratios. In contrast, male provisioning rates were not affected by brood sex ratio in either natural or experimental broods. However, males in established pairs provisioned at a higher rate than males in new pairs. Data on the sex ratio of 109 broods suggest that female brown thornbills adjust their primary sex ratio in response to pair bond duration. Females in new pairs produced broods with significantly fewer sons than females in established pairs. This pattern would be beneficial to females if the costs of rearing sons were higher for females in new than established pairs. This may be the case since females in new pairs provisioned experimental all-male broods at elevated rates. The condition of nestlings also tended to decline more in these all-male broods than in other experimental broods. This will have additional fitness consequences because nestling mass influences recruitment in thornbills. Female thornbills may therefore obtain significant fitness benefits from adjusting their brood sex ratio in response to the status of their pair bond. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Hatching order and size-dependent mortality in relation to brood sex ratio composition in chinstrap penguins

The differential environmental sensitivity of the sexes hasstrong implications in the evolutionary history of species asit can alter sexual size dimorphism, population sex ratios,and the faculty of parents to manipulate offspring sex in relationto environmental conditions. We studied sexual differences inhatching patterns and evaluated sex- and size-related mortalityin relation to hatching order and brood sex ratios in the chinstrappenguin Pygoscelis antarctica, a moderately size-dimorphic species,with a modal clutch size of 2 eggs. We found that male, second-hatched,and large eggs showed shorter hatching periods than female,first-hatched, and small eggs. We also found a male-biased mortalityof nestlings in the colony. However, male mortality patternsdiffered depending on the brood sex ratio composition. Mortalityof male chicks in all-male broods was higher than in mixed broodsand higher than female mortality in all-female broods. Contrary,females from mixed brood showed higher mortality than theirmale nest mates and higher too than females in all-female broods.Second-hatched chicks also suffered from higher mortality thanfirst-hatched chicks. Our results indicate that both the superiorcompetitive capacity and the higher energy demand of the largersex constitute 2 causal factors explaining patterns of sex-biasedmortality. Both factors occur in the same species and in differentsituations of sibling competition shaped by brood sex ratiocomposition. This study constitutes a good example of how patternsof sex-related mortality can vary depending on nest environmentalcircumstances. Furthermore, our study suggests that hatchingperiod can be a mechanism underlying sexual differences in theembryonic period of birds.     

Oviposition behavior and progeny allocation of the polyembryonic waspCopidosoma floridanum (Hymenoptera: Encyrtidae)

Oviposition behavior was used to determine the primary clutch size and sex ratio of the polyembryonic wasp Copidosoma floridanumAshmead (Hymenoptera: Encyrtidae) parasitizing Pseudoplusia includens(Walker) (Lepidoptera: Noctuidae). The laying of a female egg was associated with a pause in abdominal contractions during oviposition, while the laying of a male egg was associated with uninterrupted abdominal contractions. Although unmated females produced only male broods, they also displayed male and female egg oviposition movements. Wasps always laid a primary clutch of one or two eggs. For mated females if only one egg was laid, the emerging secondary clutch was all male or female, but if two eggs were laid a mixed brood of males and females was almost always produced. The secondary clutch of single sex broods was usually between 1000 and 1200 individuals, but the secondary clutch of mixed broods averaged 1143 females and 49 males. Thus, the primary sex ratio for mixed broods was 0.5 (frequency males), but the secondary sex ratio was 0.042. Manipulation of the sequence of male and female egg oviposition or of the primary clutch did not produce major alterations in the secondary clutch size or sex ratio.     

Brood guarding in a bethylid wasp

Abstract.

• 1 Atypical of the parasitoid Hymenoptera, female Goniozus nephantidis Muesebeck remain with offspring until they pupate. This behaviour will only have evolved if the consequent reduction in fecundity is outweighed by fitness returns.
• 2 G.nephantidis competes for hosts with conspecific and non-conspecific parasitoids. The effectiveness of G.nephantidis at deterring superparasitism and multiparasitism is tested. Brood survivorships were compared when G.nephantidis and Bracon hebetor Say intrude on hosts with the mother present and absent and with offspring at different developmental stages.
• 3 Laying by the intruder reduced brood survivorship. Guarding reduced oviposition on unparasitized hosts by intruding females, and prevented superparasitism of hosts with egg broods. Hosts with larval broods were rarely superparasitized. Multiparasitism was common except on hosts with guarded larval broods, but even here survivorship was reduced.
• 4 Competitive asymmetries determined the outcome of contests for possession of host resources.
• 5 The costs of remaining seem at least partially offset by the prevention of oviposition by competing parasitoids.

     

Sex allocation in the sexually monomorphic fairy martin

Offspring sex ratios were examined at the population and family level in the sexually monomorphic, socially monogamous fairy martin Petrochelidon ariel at five colony sites over a 4-year period (1993–1996). The sex of 465 nestlings from 169 broods was determined using sex-specific PCR at the CHD locus. In accordance with predicted sex allocation patterns, population sex ratios at hatching and fledging did not differ from parity in any year and the variance in brood sex ratios did not deviate from the binomial distribution. Further, brood sex ratio did not vary with hatching date during the season, brood number, brood size or colony size. The sex ratio of broods with extra-pair young did not differ from those without, while the sex ratio of broods fathered by males that gained extra-pair fertilizations did not differ from broods fathered by other males. Extra-pair chicks were as likely to be male as female. Neither the total number of feeding visits to the brood nor the relative feeding contribution by the sexes varied significantly with brood sex ratio. Brood sex ratios were also unrelated to paternal size, condition and breeding experience or maternal condition and breeding experience. However, contrary to our prediction, brood sex ratio was negatively correlated with maternal size. Generally, these results were consistent with our expectations that brood sex ratios would not vary with environmental factors or parental characteristics, and would not influence the level of parental provisioning. However, the finding that females with longer tarsi produced an excess of daughters is difficult to reconcile with our current understanding of fairy martin life history and breeding ecology.     

Offspring sex ratio variation in relation to brood size and mortality in a promiscuous species: the Aquatic Warbler Acrocephalus paludicola

We analyse nestling sex ratio variation in the Aquatic Warbler Acrocephalus paludicola to test for predictions from sex allocation theory that the brood sex ratio is close to parity. We also tested Fiala's (1980) prediction that there is no difference in sex ratio between broods affected and not affected by mortality, and whether a shift in primary sex ratios or simple differential mortality by sex underlies that difference. Furthermore, we explore additional analytical possibilities for inferring proximate mechanisms through simulation modelling. In the Aquatic Warbler, which is promiscuous, the overall sex ratio determined by molecular sexing of nestlings at 8–11 days of age did not deviate significantly from parity (proportion of females 0.509), nor did we find any predictive effect of brood size, maternal body mass, fat condition, wing and bill length, laying date, mean daily temperature, and multiple-male mating. However, extensive simulation suggested that the whole pattern of sex ratio variation is unlikely to arise purely by chance: (1) there is a diverging sex ratio between complete and partial broods, (2) large broods tended to be female-biased and small broods male-biased, and (3) low ambient temperature prior to the laying period seemed to increase the proportion of female offspring in complete broods. We conclude that most variation in nestling sex ratio is non-adaptive in nature, and results from variation in female nestlings mortality dependent on brood size and sex ratio.     

Sex allocation and clutch size in parasitoid wasps that produce single-sex broods

The parasitoid wasp genus Achrysocharoides (Eulophidae) is unusual in that many of its species lay male and female eggs in single-sex clutches. The average clutch size of female broods is always greater than that of male broods, and in some species male clutch size is always one. We constructed models that predicted that severely egg-limited wasps should produce equal numbers of male and female eggs while severely host-limited wasps should produce equal numbers of male and female broods (and hence an overall female-biased sex ratio). Theory is developed to predict clutch size and sex ratio across the complete spectrum of host and egg limitation. A comparison of 19 surveys of clutch composition in seven species of Achrysocharoides showed a general pattern of equal numbers of male and female broods with a female-biased sex ratio (suggesting host limitation) although with considerable heterogeneity amongst collections and with a number of cases of unexpectedly low frequencies of male broods. Using a previous estimate of the relationship between fitness and size in the field, we predicted the maximally productive (Lack) clutch size for female broods of Achrysocharoides zwoelferi to be three. Of clutches observed in nature, 95% were equal to or smaller in size than the predicted Lack clutch size. When we manipulated local host density in the field, and as predicted by our models, clutch size and the proportion of female broods of A. zwoelferi decreased as hosts became more common, but the absolute frequency of male clutches was lower than expected. Copyright 1998 The Association for the Study of Animal Behaviour.     

Sex ratio dynamics of Gonatocerus ashmeadi,a parasitoid of the glassy‐winged sharpshooter in California

We examined whether Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae), a quasi‐gregarious egg parasitoid of Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), produces precise sex ratios under a field setting. Under laboratory conditions, previous studies have shown that G. ashmeadi exhibits strongly female‐biased sex ratios with low variance in the number of males produced per host. Field‐collected G. ashmeadi tend to produce much less female‐biased sex ratios with high variance in male numbers. We found significant positive effects of proportion parasitism and host density on sex ratio. Proportion parasitism also had a positive effect on sex ratio variance. The findings of this study are discussed in the context of theoretical predictions.     

Family planning in a stemborer parasitoid: sex ratio, brood size and size-fitness relationships in Parallorhogas pyralophagus (Hymenoptera: Braconidae), and implications for biological control

Various aspects were studied of the brood size and sex allocation strategies, and of size-fitness relationships in Parallorhogas pyralophagus (Marsh), a gregarious ectoparasitoid of Eoreuma loftini Dyar. Brood size was significantly correlated with host size; larger hosts were allocated larger broods. Brood sex ratios were fixed precisely at 1 male per 4 females, and eggs were likely to be deposited in that order; differential mortality did not contribute to this precise sex ratio. The sex allocation strategy of P. pyralophagus is likely to conform to strict, i.e. single foundress, local mate competition. Adoption of this strategy is probably influenced by a limited insemination capacity of males; a smaller proportion of females (0.09 vs. 0.21) remained virgin in broods with precise or higher sex ratios (> or = 0.20 males) relative to broods with lower than precise sex ratios (< 0.20 males). Moreover, all females were inseminated in most broods (60%) with precise or higher sex ratios, whereas this did not occur in broods with lower than precise sex ratios. The hypothesized occurrence of strict local mate competition in P. pyralophagus was supported also by observations that: (i) offspring brood sex ratios were independent of maternal brood sex ratios and number of parental females concurrently allocating offspring to a group of hosts, and; (ii) the rate of superparasitism under no-choice conditions was low (approximately 20%), suggesting that rates of outbreeding in the field are low. Other results suggested that fitness in P. pyralophagus was correlated with adult size; longevity and reproductive capacity both increased with adult size in males and females. However, adult size may be more important for females than for males because the differences in reproductive capacity between the largest and smallest individuals was up to 7.3 times greater in females versus < 2 times in males.     

Evolution of social parasitism in ants: size of sexuals,sex ratio and mechanisms of caste determination

Social parasitism, one of the most intriguing phenomena in ants, has evolved to various levels, the most extreme form being parasites that have lost the worker caste and rely completely on the host''s worker force to raise their brood. A remarkable feature of workerless social parasites is the small size of sexuals. It has been suggested that reduced size evolved as a means to take advantage of the host''s caste-determination system, so that parasite larvae develop into sexuals with less food than is required to produce host workers. An important consequence of size reduction is that it might restrict the host workers'' ability to discriminate between the brood of the social parasite and their own brood and might protect parasite sexuals from elimination. We found that sexuals of the workerless inquiline ant Plagiolepis xene were significantly smaller than the sexuals of their host Plagiolepis pygmaea, but remarkably similar to the host workers. The size variance of parasite sexuals was much lower than that of their host; this result possibly suggests that there is very stabilizing selection acting on size of the parasite sexuals. Comparison of the primary (egg) and secondary (adult) sex ratios of the parasite and host showed that miniaturization of P. xene sexuals has been accompanied by their ability to develop into sexuals even when the host P. pygmaea actively prevents production of its own sexuals. These results suggest that the inquiline''s size and caste threshold have been reduced such that all individuals in a parasite brood will develop into sexuals. We also found that the adult sex ratio of P. xene was heavily female-biased. This bias probably stems from local mate competition that arises from sexuals mating within the nest. There was no significant difference between the proportion of haploid eggs and adult males produced; this observation indicates that a female-biased sex ratio is achieved by queens producing a higher proportion of diploid eggs rather than by a higher mortality of haploid males.     

Resource quality and spatial variation in sex ratios of a free-living solitary sawfly,Dineura virididorsata (Hym., Tenthredinidae)

The sex ratio in final-instar larvae of a birch-feeding, free-living solitary sawfly, Dineura virididorsata, was investigated in Finnish Lapland. The prepupal proportion of females, pooled over ten sites, was 56%, and at four individual sites the sex ratio was significantly female-biased. Larval survival from egg to prepupae did not differ between the sexes. This suggests a femalebiased primary sex ratio in the field. The sex ratio varied among the sites but not among host trees within sites. Contrary to previous results with hymenopterans, we did not find that differences in the sex ratio depended on forage quality: site-specific or tree-specific sex ratios did not correlate with the average prepupal weight. A literature search indicated that female-biased sex ratios are also common in other free-living sawflies. We are unable to explain sex ratios of Dineura virididorsata or other free-living sawflies with existing general models.     

Stochastic effects in LMC models.

LMC (local mate competition) was first introduced by W. D. Hamilton to explain extraordinary female-biased sex ratios observed in a variety of insects and mites. In the original model, the population is subdivided into an infinite number of colonies founded by a fixed number of inseminated females producing the same very large number of offspring. The male offspring compete within the colonies to inseminate the female offspring and then these disperse at random to found new colonies. An unbeatable sex ratio strategy is found to be female-biased. In this paper, the effects of having colonies of random size and foundresses producing a random finite number of offspring are considered. The exact evolutionarily stable strategy (ESS) sex ratio is deduced and comparisons with previous approximate or numerical results are made. As the mean or the variance of brood size increases, the ESS sex ratio becomes more female-biased. An increase in the variance of colony size increases the ESS proportion of males when the mean brood size and colony size are both small, but decreases this proportion when the mean brood size or the mean colony size is large.     

Forest fragmentation is associated with primary brood sex ratio in the treecreeper (Certhia familiaris)

We studied the primary brood sex ratio of an old-growth forest passerine, the Eurasian treecreeper (Certhia familiaris), along a gradient of forest fragmentation. We found evidence that male nestlings were more costly to produce, since they suffered twofold higher nestling mortality and were larger in body size than females. Furthermore, the proportion of males in the brood was positively associated with the provisioning rate and the amount of food delivered to the nestlings. During the first broods, a high edge density and a high proportion of pine forests around the nests were related to a decreased production of males. The densities of spiders, the main food of the treecreeper, were 38% higher on spruce trunks than on pine trunks. This suggests that pine-dominated territories with female-biased broods may have contained less food during the first broods. The observation was further supported by the fact that the feeding frequencies were lower in territories with high proportions of pines. In the second broods, territories with a high forest patch density produced female-biased broods, whereas high-quality territories with a large amount of deciduous trees and mixed forests produced male-biased broods. Our results suggest that habitat quality as measured by habitat characteristics is associated with sex allocation in free-living birds.