The effect of partial brood loss on male desertion in a cichlid fish: an experimental test

There is little experimental evidence testing whether currentbrood size and past brood mortality influence mate desertion.In the cichlid Aequidens coeruleopunctatus both parents initiallydefend offspring. In a field study, all experimental broods,irrespective of initial brood size (222.9 ± 60.4, mean± SD), were manipulated to a size of 100 fry. Neitherthe duration nor investment of females in parental care differed between control and brood reduced pairs, even though care seemedcostly. On average, females lost 5.1 ± 4.8% of initialweight while guarding a brood until independence. In contrast,males with experimentally reduced broods guarded fry for significantlyfewer days before deserting their mate than did males fromcontrol pairs with natural-sized broods (20.5 ± 7.5 vs. 14.2 ± 6.2 days). In at least 20% of cases (n = 9/45),the deserting male immediately mated with another female. Maleswith experimentally reduced broods also spent less time guardingfry before deserting and attacked fewer brood predators thandid males with control broods. For broods manipulated to have100 fry, there was a significant negative relationship betweenthe days until male desertion and the proportion of the initialbrood removed. This indicates that male assessment of the futuresuccess of the current brood (hence its reproductive value)is based on past mortality and/or that there is variation amongmales in the expected size of future broods. Both current broodsize and brood size relative to initial brood size are thereforepredictors of male, but not female, parental behavior and matedesertion. Female care may be unaffected by brood reductiondue to limited breeding opportunities and partial compensationfor reduced male care.     

Induced crossing-over in the presterile broods of drosophila melanogaster males

The results from two experiments on induced crossing-over in the germ cells of adultDrosophila melanogaster males, heterozygous for 12–13 third-chromosome mutant loci, support the premise that the F₁ offspring recovered in the 6–8 day broods are derived mainly from cells that had been primary spermatocytes at the time of irradiation. This conclusion is also rather well supported by the results from similar experiments reported in the literature. The frequency of exchange in the third-chromosome following irradiation of the adult male is in the order of 0.01% for the 1–3 and 4–5 day broods, 0.15% for the 6–7 day broods and 0.31% for the 8th day brood in the pooled data from 18 experiments. The spontaneous rate for the third-chromosome is in the order of 0.001%. The rather great variance among the experiments in time of recovery of the first crossovers and the frequency of exchange during the time of sampling the spermatocytes and definitive spermatogonia—ranging from none to a 20-fold increase (over the spontaneous rate) in the 6–8 day broods, and from none to a 10–50-fold increase from the 6–8 to the 9–11 day broods — can be attributed to the brood procedures or mating pressure, the effectiveness of ascertainment of crossing-over, particularly in the early broods, and the dosages used by the different investigators. The lack of a dosefrequency relationship in the broods prior to the period of peak sterility is due, at least in part, to lethal-loss of the cells with induced exchange. The evidence from the two experiments also supports the premise that there may be different types of exchange-induction with a causal relationship between the time of induction and the site in the chromosome. The higher relative frequency of multiple crossovers and noncentric exchanges among the recombinants recovered in the 1–7 day broods, as compared to their frequency in the spermatogonial broods, suggests that there may be a difference in the mode of induction or transmission of recombinant chromosomes in different stages of spermatogenesis. The relatively high incidence of F₁-sterility of the recombinants and homozygous lethality of the recombinant chromosomes points to the possibility that either the meiotically-induced exchanges are intrachromosomal aberrations frequently accompanied by duplications or deficiencies due to breakage at non-identical loci, or homologous chromosomes sensitive to the induction of exchange are more likely to be hypersensitive to the induction of genetic damage with detrimental fertility and viability effects.     

Survival for specific life intervals of smallmouth bass, Micropterus dolomieu, during parental care

We tested whether daily mortality rates (DMR) of smallmouth bass offspring were influenced by life interval, offspring density and growth, parental male attributes, and selected mortality factors during parental care in a regulated Virginia stream. Mortality averaged 9.5% per day (range 5.2–13.9%) and 94.1% total (range 80.9–99.5%) from egg deposition to the juvenile period (29–36 d) for individual broods. Offspring losses were primarily attributed to fungus (Saprolegnia parasitica) infection of eggs and to American eel, Anguilla rostrata, predation. DMR were significantly higher for the interval from swim-up of larvae to metamorphosis relative to earlier and later intervals. There was no significant autocorrelation of DMR among life intervals for individual broods, indicating that relative mortality rates were inconsistent among broods through time. DMR were also uncorrelated with the number of offspring per brood, offspring growth rates, and parental male attributes, except during egg and embryo intervals. Daily egg mortality was negatively related to male size and positively related to the number of eggs per nest, suggesting that density-dependent egg mortality may have been partially offset in nests of larger males. Larger males received more eggs, tended to maintain larger broods throughout parental care, and contributed a high proportion of the total number of juveniles reared. This revised version was published online in August 2006 with corrections to the Cover Date.     

Brood parasitism increases provisioning rate, and reduces offspring recruitment and adult return rates, in a cowbird host

Interspecific brood parasitism in birds presents a special problem for the host because the parasitic offspring exploit their foster parents, causing them to invest more energy in their current reproductive effort. Nestling brown-headed cowbirds (Molothrus ater) are a burden to relatively small hosts and may reduce fledgling quality and adult survival. We documented food-provisioning rates of one small host, the prothonotary warbler (Protonotaria citrea), at broods that were similar in age (containing nestlings 8–9 days old), but that varied in composition (number of warbler and cowbird nestlings) and mass, and measured the effect of brood parasitism on offspring recruitment and adult returns in the host. The rate of food provisioning increased with brood mass, and males and females contributed equally to feeding nestlings. Controlling for brood mass, the provisioning rate was higher for nests with cowbirds than those without. Recruitment of warbler fledglings from unparasitized nests was 1.6 and 3.7 times higher than that of fledglings from nests containing one or two cowbirds, respectively. Returns of double-brooded adult male and female warblers decreased with an increase in the number of cowbirds raised, but the decrease was more pronounced in males. Reduced returns of warbler adults and recruitment of warbler fledglings with increased cowbird parasitism was likely a result of reduced survival. Cowbird parasitism increased the warblers’ investment in current reproductive effort, while exerting additional costs to current reproduction and residual reproductive value. Our study provides the strongest evidence to date for negative effects of cowbird parasitism on recruitment of host fledglings and survival of host adults.     

Male death resulting from hybridization between subspecies of the gypsy moth,Lymantria dispar

We explored the origin of all-female broods resulting from male death in a Hokkaido population of Lymantria dispar through genetic crosses based on the earlier experiments done by Goldschmidt and by testing for the presence of endosymbionts that are known to cause male killing in some insect species. The mitochondrial DNA haplotypes of the all-female broods in Hokkaido were different from those of normal Hokkaido females and were the same as those widely distributed in Asia, including Tokyo (TK). Goldschmidt obtained all-female broods through backcrossing, that is, F1 females obtained by a cross between TK females (L. dispar japonica) and Hokkaido males (L. dispar praeterea) mated with Hokkaido males. He also obtained all-male broods by mating Hokkaido females with TK males. Goldschmidt inferred that female- and male-determining factors were weakest in the Hokkaido subspecies and stronger in the Honshu (TK) subspecies. According to his theory, the females of all-female broods mated with Honshu males should produce normal sex-ratio broods, whereas weaker Hokkaido sexes would be expected to disappear in F1 or F2 generations after crossing with the Honshu subspecies. We confirmed both of Goldschmidt''s results: in the case of all-female broods mated with Honshu males, normal sex-ratio broods were produced, but we obtained only all-female broods in the Goldschmidt backcross and obtained an all-male brood in the F1 generation of a Hokkaido female crossed with a TK male. We found no endosymbionts in all-female broods by 4,′6-diamidino-2-phenylindole (DAPI) staining. Therefore, the all-female broods observed in L. dispar are caused by some incompatibilities between Honshu and Hokkaido subspecies.     

Environmental- and parental condition-related variation in sex ratio of kestrel broods

Variation of brood sex ratio was studied in a Finnish population of Eurasian Kestrels Falco tinnunculus breeding in an unpredictably variable environment. From those young that survived until 2–4 weeks of age, blood was collected and their sex determined from polymorphic DNA profiles produced by hybridisation with a human minisatellite probe. The sex ratio was male-biased during a year of food (vole) scarcity. Furthermore, in broods without mortality, contrasting seasonal trends in sex ratios emerged. In this subsample, the proportion of males increased with later laying date during years of low and moderate food supply, whereas the opposite was true in a year of relatively high food supply. These trends may indicate circumstances that favour the raising of different sex. The proportion of males in the brood was negatively correlated with body condition of both male and female parents, also reflecting an adaptive condition-dependent sex-ratio adjustment, or alternatively the inability of the parents to meet the requirements of the more energetically expensive female offspring. We discuss the limitations that unpredictable conditions during brood raising can impose on adaptive sex-ratio manipulation, particularly in species with sexual size dimorphism and consequent differences in the cost of raising the two sexes.     

Brood desertion in Kentish plover: sex differences in remating opportunities

To understand the evolution of parental care, one needs to estimatethe payoffs from providing care for the offspring and the payoffsfrom terminating care and deserting them. These payoffs arerarely known. In this study we experimentally estimated therewards from brood desertion in a species that has a variablepattern of parental care. In particular, either the female or themale parent may desert the brood in Kentish plover Charadrius alexandrinus,so some broods are attended by one parent of either sex, whereasin other broods both parents stay with the brood until the chicks fledge.We created single males and single females by experimentallyremoving the other parent and the clutch. The expected rematingtime of males was significantly higher (median: 25.4 days) thanthat of the females (5.3 days, p <.0001). The expected rematingtime tended to increase over the breeding season in both sexes,although the increase was significant only in females. The newnest of remated males was closer to their previous territory (mean± SE, 46 ± 8 m) than that of the remated females(289 ± 57 m, p <.001). Hatching success of new nestswas not different between remated males and females. Our resultsdemonstrate that the remating opportunities are different formale and female Kentish plovers and these opportunities varyover the season. We propose that the remating opportunitieswere influenced by the male-biased adult sex ratio and the seasonaldecrease in the number of breeders. However, we stress thatmeasuring remating times is a more direct measure of matingopportunities than calculating the operational sex ratio.     

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.     

High prevalence of multiple paternity in the invasive crayfish species,Procambarus clarkii

Reproductive strategy is a central feature of the ecology of invasive species as it determines the potential for population increase and range expansion. The red swamp crayfish, Procambarus clarkii, has invaded many countries and caused serious problems in freshwater ecosystems. However, little is known about the effects of environmental conditions on crayfish paternity and offspring traits in the wild. We studied these reproductive characteristics of P. clarkii in wild populations from two different habitats (ponds and ditches) in three locations with different environmental conditions in China. Genotyping of 1,436 offspring and 30 mothers of 30 broods was conducted by using four microsatellites. An analysis of genotyping results revealed that gravid females were the exclusive mother of the progeny they tended. Twenty-nine of 30 mothers had mated with multiple (2-4) males, each of which contributed differently to the number of offspring in a brood. The average number of fathers per brood and the number of offspring per brood were similar (P > 0.05) among six sampling sites, indicating that in P. clarkii multiple paternity and offspring number per brood are independent of environmental conditions studied. Indirect benefits from increasing the genetic diversity of broods, male and sperm competition, and cryptic female choice are a possible explanation for the high level multiple paternity and different contribution of fathers to offspring in this species.     

Reproduction characteristics,survival rate and sex-ratio of four brine shrimp Artemia salina (Linnaeus, 1758) populations from Tunisia cultured under laboratory conditions

Artemia has a remarkable genetic variability that can be expressed in various phenotypic characteristics, such as morphometry, growth rate, reproductive isolation or molecular composition. This study presents reproduction characteristics, survival rate and sex-ratio of four Artemia salina populations from Tunisia cultured under standard conditions. Results show that both low (17.8% and 30.6% in Megrine saltwork and Sabkhet Sijoumi, respectively) and high (83.9% for Sahline saltwork) values were recorded for the percentage of total oviparous offspring. The offspring per brood varied from 70.3?±?41.9 (Megrine saltwork) to 73.6?±?51.4 (Sabkhet El Adhibet). Broods per female varied between 3.1?±?0.9 (Sahline saltwork) and 4.5?±?1.2 (Megrine saltwork), and time between broods ranged from 5.7?±?1.1 to 6.5?±?2.2 days for Megrine and Sahline, respectively. Statistical analysis (one-way analysis of variance, Tukey's HSD test, p?<?0.05) revealed no significant differences between reproductive traits of the four studied populations except for mean of oviparous and oviviparous offspring per female (F?=?9.158, p?<?0.05), and brood per female (F?=?4.779, p?<?0.05). The survival rate of the four studied A. salina populations fluctuated between 31.4% and 64.5% for Megrine and Sahline saltwork, respectively. However, the sex-ratio showed that for Megrine saltwork and Sabkhet El Adhibet, males predominated with 1?:?0.90 and 1?:?0.97 (males?:?females), respectively. The comparison between our results and those reported of other Artemia populations showed that the offspring per brood of Tunisian Artemia are comparable to A. persimilis but different to A. salina from Abu Kammash (Libya) and A. urmiana (Urmia Lake, Iran), and that days between brood, are similar to A. salina from Abu Kammash, A. sinica and polyploid A. parthenogenetica, but different from A. persimilis and A. franciscana.     

Nest intrusions, infanticide, and parental care in the burying beetle, Nicrophorus orbicollis (Coleoptera: Silphidae)

Duration of paternal care in the burying beetle Nicrophorus orbicollis Say is highly variable. Both parents bury and defend mouse-sized vertebrate carcasses as food resources for their offspring, but males abandon their broods several days before females. Nests defended by single female parents were taken over by aggressive conspecifics in live of nine cases, whereas only six of 16 nests defended by both parents were taken over. In the event of a takeover, the intruding beetle replaced the resident beetle of the same sex, destroyed any eggs that were present, and paired with the remaining resident to produce a new clutch. Broods raised by usurpers following takeovers were less successful than broods raised by initial residents on unused carcasses. The majority of takeovers occurred 35 days after carcass burial. The occurrence of nest intrusions by conspecifics did not significantly influence duration of male parental care; when conspecific intruders were excluded from nests males remained with their broods (± S.E.) 11·2 ± 0·8 days ( n = 15), and when intruders were added to nests males remained with their broods 12·2 ± 0·6 days ( n = 8). Conflict for carcasses intensified in response to larger brood mass, but duration of male care was unaffected by brood mass. Overall. brood mass and the presence or absence of intruders explained only 5% of the variance associated with brood abandonment by males.     

Sexual conflict among polygynous pied flycatchers feeding young

In the polygynous pied flycatcher, Ficedula hypoleuca, reproductivesuccess of females is constrained by male food provisioningduring the nestling period. Hence, there will be conflictinginterests among the male and each of his mates as to how malefeeding effort should be shared among broods. This paper describesthree experiments designed to examine the parental behaviorof the members of a bigynous trio, i.e., the male and his twomates, in light of these conflicts. In all experiments, primaryand secondary broods were manipulated to hatch on the same dayto reduce the difference in brood-reproductive value due toage. Males divided their effort equally when the two broodswere the same size. However, males did not allocate their investmentin proportion to brood size when brood sizes differed, but investedmore heavily per young in the larger broods. This finding suggeststhat males tried to optimize the joint effort of their two mates.Males and females showed similar responses to experimental reductionin brood demands, which indicates no difference in their willingnessto invest in offspring. When one of the male’s mates wasremoved temporarily, the male increased his total feeding rateand provided proportionately more food to the "motherless" brood.Through flexible allocation of parental investment, males seemable to optimize their reproductive interests in the two broods.The only way a polygynously mated female might successfullyincrease the amount of male assistance at her nest is to makeher own brood more valuable for the male, relative to the otherbroods he might have. We discuss some ways this might be achieved.[Behav Ecol 1991;2:106–115]     

Reproductive success of polygynous male corn buntings (Miliaria calandra) as confirmed by DNA fingerprinting

Despite a sex ratio approximating to unity, female corn buntingswere not equally distributed among males. In 1989 and 1990,41.2% of 50 males were monogamously paired, 29.4% were polygynous,and 23.5% were unpaired. Polygynous males usually paired withtwo females, although in 1990 three males were trigamous. Polygynousmales fledged more offspring from their territories than didmonogamous males, mainly because they had more mates. The fledgingsuccess per nesting female was slightly higher in territoriesof polygynous males, but not significantly so. DNA fingerprintingwas used to confirm the true paternity of 44 offspring from15 broods and the true maternity of 50 offspring from 16 broods.A further 12 offspring from three broods for which neither putativeparent was available were also fingerprinted. Actual reproductivesuccess of parents was close to that inferred from observationsof number of young raised. There was only one brood, containingtwo chicks (4.5% of offspring, or in 6.7% of broods), wherethe chicks were not fathered by the male defending the territory.However, this nest was close to the territory boundary, andthe defending male may have been assigned incorrectly. Therewere no cases of intraspecific brood parasitism (n = 16 broods).The copulation rate was low, and extrapair copulation attemptswere rare, probably because of the poor chances of sneakingonto a neighbor's territory undetected and the costs of leavinga territory unguarded.     

Scale and seasonal sex-ratio trends in northern goshawk Accipiter gentilis broods

In many populations of size dimorphic birds, brood sex ratios change with advancing laying date. The slopes of these trends, however, vary in time and space, both between and within species. We studied brood sex ratios (proportion of males) of northern goshawks Accipiter gentilis in Finland in relation to laying date using ringing data from 1989 to 1998. At the nationwide scale, i.e. the whole of Finland, between-year variation in offspring sex ratio was moderate, and the sex ratio did not change with later laying date. At a regional scale, the sex ratio was seasonally constant in one region but decreased in another, although the laying-date/brood-size relationship was identical. Hence, the size and sex composition of goshawk broods are locally two uncoupled facets of reproduction. Both the national and regional patterns differ drastically from the pattern of a Dutch population, where sex ratio increased seasonally. We suggest that spatial variation in inter-annual seasonal sex-ratio trends might be indicative of the scale at which sex-ratio feedback functions. The sex ratio of breeders is a factor that could add to the understanding of the observed geographical differences in seasonal sex-ratio patterns.     

Cumulative effects matter: multi‐brood responses of Daphnia to hypoxia

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Survivorship,growth and reproduction of the non-native <Emphasis Type="Italic">Caprella mutica</Emphasis> Schurin, 1935 (Crustacea: Amphipoda)

Caprella mutica Schurin is an epifaunal amphipod crustacean which originates in north-east Asia and has spread throughout the world, yet very little is known about fundamental aspects of this species biology. This paper examined the survivorship of C. mutica reared under laboratory conditions at 13–14 °C, 14 h light: 10 h dark photoperiod and fed commercial salmon feed, the diatom Cylindrotheca fusiformis Reumann and Lewin, the macroalgae, Fucus vesiculosus L. and given no additional feed. In addition, growth, maturation and reproduction of C. mutica fed C. fusiformis were assessed. No significant difference in survivorship of C. mutica was observed for the diet types over the experimental period. C. mutica was able to survive for upto 20 days without additional food. Average survival time of males and females fed the diatom, C. fusiformis was 68.8 d (range = 62–73 d) and 82.0 d (range = 76–92 d). Juvenile C. mutica emerged from the brood pouch at a body length of 1.33 mm and moulted at 5.0–11.0 day intervals. Males exhibited faster growth rates than females after Instar VII. Females produced their first brood at Instar VII, 24–26 days post-hatching and with an average body length of 8.5 mm. Each female had an average of two broods sequentially and these were released at 20.2 day intervals. Brood size for a single female increased from 11.3 (±9.9) hatchlings at Instar VII to 25.5 (±11.5) at Instar IX and the maximum number of hatchlings produced by a single female was 82. The results suggest that C. mutica exhibits a number of life-history traits that would potentially enable it to withstand global transportation and to rapidly become established in an introduced region, if environmental conditions are suitable.     

Genetic parentage assessment in the crayfish Orconectes placidus,a high-fecundity invertebrate with extended maternal brood care

Microsatellite data have recently been introduced in the context of genetic maternity and paternity assignments in high-fecundity fish species with single-parent-tended broods. Here we extend such analyses to an aquatic invertebrate, the crayfish Orconectes placidus, in which gravid females carry large numbers of offspring. Genetic parentage analyses of more than 900 progeny from 15 wild crayfish broods revealed that gravid females were invariably the exclusive dams of the offspring they tended (i.e. there was no allomaternal care), and that most of the females had mated with multiple (usually two) males who contributed sometimes highly skewed numbers of offspring to a brood. Within any multiply sired brood, the unhatched eggs (or the hatched juveniles) from different fathers were randomly distributed across the mother's brood space. All of these genetic findings are discussed in the light of observations on the mating behaviours and reproductive biology of crayfish.     

Insemination Capacity and Dispersal in Relation to Sex Allocation Decisions in Goniozus legneri (Hymenoptera: Bethylidae): Why Are There More Males in Larger Broods?

Models considering sex ratio optima under single foundress strict local mate competition predict that female bias will be reduced by stochasticity in sex allocation, developmental mortality of males and limited insemination capacity of males. In all three cases the number of males per brood is expected to increase with brood size. Sex ratio optima may also be less female biased when several mothers contribute offspring to local mating groups or if non‐local mating occurs between members of different broods; again more males are expected in larger broods. In the parasitoid wasp Goniozus legneri (Hymenoptera: Bethylidae), sex allocation has only a small stochastic component, developmental mortality is low and non‐siblings are unlikely to develop in the same brood. However, the number of males per brood increases with the size of the brood (produced by a single mother). We investigated the further possibilities of limited insemination capacity and non‐local mating using a naturalistic experimental protocol. We found that limited insemination capacity is an unlikely general explanation for the increase in number of males with brood size. All males and females dispersed from both mixed and single sex broods. Although most females in mixed sex broods mated prior to dispersal, these data suggest that non‐local mating is possible, for instance via male immigration to broods containing virgin females. This may influence sex ratio optima and account for the trend in male number.     

Is the small clutch size of a Corsican blue tit population optimal?

In an attempt to test predictions of the optimisation hypothesis of life history traits in birds, we estimated fitness consequences of brood size manipulations. Experiments were carried out over a period of 4 years in a Mediterranean population of blue tits Parus caeruleus which is confronted with a particular set of environmental constraints. Effects of brood size manipulation were investigated in relation to year-to-year variation in environmental conditions, especially caterpillar abundance. There was a strong variation in the effects of brood size manipulation depending on year. Most effects were on offspring quality (fledging mass, tarsus length). The absolute number of recruits did not significantly differ among categories (reduced, control, enlarged broods) but varied considerably among years. Females recruited from enlarged broods were of lower quality, started to breed later and laid fewer eggs than those recruited from control and reduced broods. Neither parental survival nor reproductive performances of adults in year n + 1 was affected by brood size manipulation in year n. Thus there was no evidence for a cost of reproduction in this population. Since the number of recruits did not depend on brood size manipulation (recruitment rates were higher in reduced broods), but recruits from reduced broods were of better quality compared with other groups, we conclude that adults lay a clutch that is larger than that which is predicted by the optimisation hypothesis. Producing more young could incur some penalties because offspring from large broods are of lower quality and less likely to recruit in the population. Two possible reasons why decision rules in this population seem to be suboptimal are discussed. Received: 10 March 1998 / Accepted: 1 July 1998     

Sex-ratio meiotic drive in Drosophila simulans is related to equational nondisjunction of the Y chromosome

The sex-ratio trait, an example of naturally occurring X-linked meiotic drive, has been reported in a dozen Drosophila species. Males carrying a sex-ratio X chromosome produce an excess of female offspring caused by a deficiency of Y-bearing sperm. In Drosophila simulans, such males produce approximately 70-90% female offspring, and 15-30% of the male offspring are sterile. Here, we investigate the cytological basis of the drive in this species. We show that the sex-ratio trait is associated with nondisjunction of Y chromatids in meiosis II. Fluorescence in situ hybridization (FISH) using sex-chromosome-specific probes provides direct evidence that the drive is caused by the failure of the resulting spermatids to develop into functional sperm. XYY progeny were not observed, indicating that few or no YY spermatids escape failure. The recovery of XO males among the progeny of sex-ratio males shows that some nullo-XY spermatids become functional sperm and likely explains the male sterility. A review of the cytological data in other species shows that aberrant behavior of the Y chromosome may be a common basis of sex-ratio meiotic drive in Drosophila and the signal that triggers differential spermiogenesis failure.