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 nesting biology of the bee, Osmia dimidiata Morawitz, 1870 (Hymenoptera, Megachilidae) in the Crimea

The structure and composition of Osmia dimidiata nests were studied based on the material of 133 nests obtained from trap-nests in five localities of the Crimea. The nests contained from 1 to 9 cells (on average, 2.2), separated with partitions made of masticated leaves. Most of the cells had a massive hemispherical bottom and a thin flat cover. The final plug had a structure similar to the cell bottom. The cells occurred singly in the nest cavity or were arranged in rows of 2 or 3 cells separated by galleries. The diameter of the occupied cavities was positively correlated with the fraction of the females in the progeny and with the mass of the cocoons, but negatively correlated with the cell length. The cells with females were 1.5 times as long as those with males. The cocoons with females were twice as heavy as those with males. The peculiar characters of the nest structure of O. dimidiata may mean that the typical nesting substrates of this species are abandoned cells of other bee and wasp species. The characteristic mechanism of estimating the mass of the pollen loaf stored in the cells and determining the brood sex ratio has evolved in the O. dimidiata due to its tenantry. This mechanism is based on two constants: the sex-linked cell length and the fixed distance between the pollen loaf and the cell cover. When choosing the nesting cavities, the female prefers broader and longer tubes for cells with females and narrower and shorter ones for cells with males. The overall sex ratio of the progeny was male-biased (1 ♀: 2.5 ♂) and depended on the food supply abundance. The adults of O. dimidiata fly in the Crimea from the last decade of May until the end of July. Hibernation occurs at the adult stage in the cocoons. Females collect pollen from flowers of Asteraceae or, rarely, Fabaceae. The nests may be destroyed by Melittobia acasta, Monodontomerus aereus, M. obscurus, Sapyga quinquepunctata, Stelis phaeoptera, and Trichodes apiarius; 21% of the progeny died from these enemies, and 22%, due to other factors. The previously proposed synonymy of Pseudosmia taurica Radoszkowski, 1874 (nec Osmia taurica Radoszkowski, 1887) and O. dimidiata is rejected.     

Few, small, and male: multiple effects of reduced nest space on the offspring of the solitary wasp, Euodynerus (Pareuodynerus) posticus (Hymenoptera: Vespidae)

The size of the preexisting wood cavities used as nests by aculeate Hymenoptera is expected to have consequences on fitness parameters such as offspring number and size. We evaluated the consequences of using small and large (three-times more voluminous) trap-nests by the solitary wasp, Euodynerus (Pareuodynerus) posticus (Herrich-Schaeffer). Following life-history and sex allocation theories, a number of non-mutually exclusive hypotheses were formulated: i.e. small nests either produce smaller or fewer offspring and/or more males, the cheaper sex. Wasps built about 28% more, but shorter brood cells in large nests, although their volume was still much higher in large nests. Adult males had smaller body size in small nests, but female size did not differ between large and small nests, possibly as an adaptive response against the future higher foraging costs of size-reduced females. Sex-ratio was often biased towards males in small nests. Mortality did not differ between large and small nests. We conclude that E.?(P.) posticus females would benefit from using larger nests, but that the sex-ratio would be probably overall unbalanced if females would not use also smaller, male-oriented tunnels.     

Phenotypic variability in nesting success among Osmia lignaria propinqua females in a glasshouse environment: (Hymenoptera: Megachilidae)

Abstract. 1. The phenotypic variability of several estimates of fitness among seventeen nesting female O.lignaria Cresson was examined in a glasshouse provided with abundant resources.
2. Females exhibited wide variation in: (a) number of nests and cells made and in their rate of construction, (b) the incidence of mortality of offspring, and (c) the percentage of male offspring produced (52–100%).
3. In confirmation of earlier field studies: (a) more offspring of both sexes were produced during the first half of the nesting season, (b) most female offspring were produced early in the nesting season, (c) most parental investment during the last half of the flight season was made in male offspring, (d) there was a decrease in offspring size with season, with females decreasing more than males, and (e) the ratio of female/male body weight was = 1.8. These results appear to be due to ageing rather than to a reduction in resource availability.
4. No relation between female size and fecundity was evident.
5. There was no evidence that small females tended to produce a greater proportion of the smaller sex (males) than did large females.
6. Smaller offspring of both sexes had a much greater probability of dying overwinter than did large offspring. However, where the sexes overlapped in size (large males, small females), almost all females died and almost all males survived. We hypothesize that as size of offspring produced declines with season, a greater proportion of males are produced because they have a much greater probability of surviving at small body sizes.     

The nest structure in four species of solitary wasps of the subfamily Eumeninae (Hymenoptera, Vespidae)

Nests of Syneuodynerus egregius, Euodynerus posticus, Ancistrocerus antilope, and A. nigricornis have a similar structure and consist of a linear row of cells separated by transverse partitions made of soil mastic. The number of cells in the nests is 1–11 (5.6 ± 0.8) in S. egregius, 1–7 (2.6 ± 0.2) in E. posticus, 1–10 (4.0 ± 0.7) in A. antilope, and 1–28 (7.9 ± 1.3) in A. nigricornis. Most nests consist of several cells in S. egregius and A. nigricornis and of one or two cells in E. posticus and A. antilope. The female to male ratio is 1.2: 1.0 in E. egregius, 1.5: 1.0 in E. posticus, 1.9: 1.0 in A. antilope, and 1.0: 1.0 in A. nigricornis. The nests of S. egregius usually contain brood of both sexes; those of A. antilope, that of one sex. The sex ratio is correlated with the diameter of the occupied nest cavities only in A. antilope. The volume of the cells with females exceeds that of the cells with males in all the species studied: by 23, 34, 54, and 98% in S. egregius, E. posticus, A. antilope, and A. nigricornis, respectively. The relative difference between the body masses of male and female prepupae is correlated with the relative difference between the volumes of their cells. The nest cells of S. egregius are separated by double partitions consisting of homologs of cell bottoms and lids. In the nests of E. posticus the partitions are single; each cell has a bottom and a lid, with “false cells” being located between them. In the nests of A. antilope and A. nigricornis, the partitions are single but all of them are homologs of the cell bottoms. The nest cells of S. egregius belong to the equilinear type; their length is relatively constant in the cavities of various diameters. The nest cells of A. antilope are of the equivolumetric type: their length is smaller in broad cavities and greater in narrow ones, the cell volume being relatively constant in the cavities of different diameters. The nest cells of E. posticus and A. nigricornis are of the intermediate type. Statistical models of nests reflecting their mean parameters and composition were built. The evolutionarily primitive and progressive features in the nest structure are discussed.     

Factors affecting female-biased sex ratio in a trap-nesting wasp, Trypoxylon malaisei

We examined the female-biased sex ratio of a trap-nesting wasp Trypoxylon malaisei considering the following factors: (1) local mate competition (LMC), (2) resource quality, (3) partial bivoltinism, and (4) presence of constrained females. The sex ratio (expressed as male ratio) at emergence was strongly female biased, i.e., 0.30 and 0.19, in terms of the number and investment, respectively. To evaluate the primary sex ratio, we analyzed the data from nests where all the offspring successfully emerged, excluding nests composed of single-sex offspring. The primary sex ratio was also female biased, at 0.33 and 0.21, in terms of the number and investment, respectively. LMC was highly responsible for the female-biased sex ratio because both the nonrandom oviposition sequence [females at inner cells and male(s) at outer cells] and earlier emergence of males allowed sib-matings to occur. In contrast, the other three factors little affected the female-biased sex ratio: the sex ratio was fairly constant when resource quality (nest size) varied, partial bivoltinism was extremely rare or absent, and constrained females were absent or did not reproduce at all. Received: June 19, 1998 / Accepted: January 18, 1999     

The nest structure in four wasp species of the genus Euodynerus Dalla Torre (Hymenoptera, Vespidae: Eumeninae)

A comparative analysis of the main parameters of the nest structure and composition in Euodynerus quadrifasciatus, Eu. dantici, Eu. disconotatus, and Eu. velutinus in the Crimea is given. Nests of all the studied species were obtained from trap-nests made of reed stems; the nests of Eu. dantici and Eu. disconotatus were additionally obtained from the abandoned cells of the nests of Sceliphron destillatorium. The nest of Eu. quadrifasciatus consists of a consecutive row of cells, each cell having its own bottom and lid with empty space between them; the nests of three other species consist of an uninterrupted row of cells without spaces, so that the cell bottoms act as partitions. The data on the diameter and length of nest cavities occupied by the studied species, the number of cells in nests, the sex ratio and the length of the cells with prospective females and males are reported. The correlations between the length of the cells and the diameter of the occupied cavities and between the length of the rear empty nest space and vestibule and the length of the occupied nest cavity are analyzed. The structure of cocoons and final nest plugs are described. The difference in the evolution of nest building instincts in members of the subgenera Pareuodynerus and Euodynerus s. str. and the role of adaptation to nest tenantry in the evolution of nest building instincts of Eu. dantici and Eu. disconotatus are discussed.     

Nesting biology of four Tetrapedia species in trap-nests (Hymenoptera: Apidae: Tetrapediini)

The nests used in this study were obtained from trap-nests (tubes of cardboard and cut bamboo stems) placed on Santa Carlota Farm (Itaoca Section-IS, Santana Section-SS and Cerrado-Ce), Cajuru, SP, Brazil. The number of nests and corresponding species obtained were as follows: 516 nests of T. curvitarsis, 104 of T. rugulosa, 399 of T. diversipes and 98 of T. gamfaloi. The most abundant species from SS and Ce was T. curvitarsis, and from IS it was T. diversipes. In general, most nests were collected during the hot and wet season (September to April). The nests were constructed with sand and an oily substance, and a single female established them. The cells were constructed in a linear series, sometimes followed by a vestibular cell. The number of brood cells ranged from 1 to 10 in T. curvitarsis (n=200), and in T. garofaloi (n-51), from 1 to 8 (n-30) in T. rugulosa, and from 1 to 6 (n=37) in T. diversipes. The pollen mass (pollen + oily substance) contained a hollow, sometimes divided by a transverse ridge, on the exposed face of the pollen mass. The egg was vertically positioned in the lower part of the hollow. At times, the closing of a cell was initiated before provisioning was completed, with a construction of a collar at the cell limit. In some nests the final cellular partition also acted as a closure plug. Females began activities at 6:18 a.m. and ended between 3:31 and 6:26 p.m. Some females (T. curvitarsis, T. rugiulosa and T. ganrfaloi) did not spend the nights at their nests, returning to them only the following morning with additional material. In general, the development period (for males and females) was greater in nests collected near the end of the hot and wet season than it was for nests collected in other months. Sex ratios for each species were as follows: T. curvitarsis. 1:1: T. rugulosa, 1.6:1 female; T. diversipes, 1.9:1: T. garofaloi, 2.8:1. Males and females of T. diversipes exhibited statistically similar sizes and in the other three species the females were larger than the males. The mortality rates were statistically similar: 33.2% for T. curvitarsis, 25.8% for T. rugulosa, 26.8% for T. diversipes and 38.2% for T. garnfaloi. The parasitoids were: Coelioxoides exulans, Leucospis cayenensis, Anthrax sp., Coelioxys sp., Coelioxoides sp. and individuals of the family Meloidae.     

Simultaneous spawning by female stream goby Rhinogobius sp. and the association with brood cannibalism by nesting males

A laboratory experiment was conducted by varying the undersurface area of nesting substratum and the number of females in an experimental tank to elucidate the determinants of the mating pattern in the stream goby, Rhinogobius sp. cross‐band type. Males with larger nests tended to attract two or more females to their nest in a tank. Moreover, males spawned simultaneously with multiple females and entire brood cannibalism by males was rarely observed under a female‐biased sex ratio. When males spawned with a single female with low fecundity, however, entire brood cannibalism occurred at a high frequency, suggesting that a male guarding a nest with fewer eggs consumes the brood. Therefore, spawning behaviour of females that leads to a large egg mass would decrease the risk of entire brood cannibalism. In this species, simultaneous spawning by multiple females in a nest serves as a female counter‐measure against entire brood cannibalism. These results suggest that a conflict of interest between the sexes through brood cannibalism is a major determinant of simultaneous spawning.     

PROGENY SEX RATIOS IN DIOECIOUS SILENE LATIFOLIA (CARYOPHYLLACEAE)

Most sex ratios reported for Silene latifolia are female biased. As a result of experiments performed by Correns in the early 1900s, pollen tube competition has generally been accepted as the primary cause of these skewed ratios. We did four sets of hand pollinations in which we varied the size of pollen loads and placement of pollen along the filamentous stigma. The effect of pollen load size on progeny sex ratios was not statistically significant. Of 32 maternal families, 17 contained more females than males (one ratio deviated statistically from 1:1), and 13 contained more males than females. Paternal families exhibited a greater range of sex ratios, including three with a significant female bias and one with a significant male bias. Within experiments, neither the maternal parent nor where pollen was placed had a statistically significant effect on progeny sex ratios; the paternal effect was significant in one experiment. We suggest that sex ratios in Silene latifolia are not necessarily affected by the level of pollen competition. Other factors, including variation among males and sex-linked mortality, may help explain the skewed sex ratios that characterize populations of this species. Further, Correns' observations of excess females may have resulted from his use of interspecific hybrids.     

Morphometics and Gonadal Development of the Hagfish Eptatretus cirrhatus in New Zealand

Hagfishes have been the target of commercial fisheries in many areas of the world, with the catch processed for leather and for human consumption. A fishery has been operating in New Zealand waters for the last six years, harvesting the bearded hagfish, Eptatretus cirrhatus. The fishery has thus far been unregulated. Based on samples collected dockside over a two-year period, this report expands the morphometric database for this species, provides information on the size and weight of the harvested animals, determines the sizes at the onset of gonadal development and the minimum sizes at sexual maturation for males and females, and indicates that E. cirrhatus, like most other hagfish species, has no specific breeding season. Although females appear in the population at smaller sizes, the sex ratio for mature animals is 1:1 and the sizes of the largest males and females are comparable. The changes observed in sex ratio as a function of TL suggest differences in the timing and rates of gonadal development in females versus males rather than protogyny. Based on the size of the eggs, the number of eggs per female, the proportion of the population that contains large eggs, and the number of postovulatory females, it is clear that E. cirrhatus, like other hagfish species, are potentially vulnerable to overexploitation.     

Age dependent sex allocation in the polygynous spotless starling

Theoretical and empirical evidence suggests that avian females are able to manipulate the offspring sex ratio at birth. Although mating with an attractive male may induce females to skew the sex ratio toward males, the balance between the benefits of producing attractive sons and the costs of competing with other females for mates could vary with female age, a possibility that had not been previously explored. In this paper we increment experimentally the attractiveness of males of the polygynous spotless starling (Sturnus unicolor) by adding green plants to their nests, a trait involved in courtship, and look for female age-differential effects on offspring primary sex ratio. Young and middle aged females produced more sons in experimental than in control nests, as expected, but old females showed the opposite tendency. To explain this novel result, we speculate that older females are limited to produce the most costly sex because the physiological drawbacks imposed by ageing reduce their ability to compete with younger ones for the non-shareable resources offered by males. We discuss that this evolutionary scenario may be widespread in avian polygynous systems.     

Egg sex ratio and paternal traits: using within-individual comparisons

Empirical studies of sex ratios in birds have been limited dueto difficulties in determining offspring sex. Since molecularsexing techniques removed this constraint, the last 5 yearshas seen a great increase in studies of clutch sex ratio manipulationby female birds. Typically these studies investigate variationin clutch sex ratios across individuals in relation to environmentalcharacteristics or parental traits, and often they find no relationships. In this study we also found that clutch sex ratiosdid not vary in relation to a number of biological and environmentalfactors for 238 great tit Parus major nests. However, interestingsex ratio biases were revealed when variation in clutch sexratios was analyzed within individual females breeding in successiveyears. There was a significant positive relationship betweenthe change in sex ratio of a female's clutch from one yearto the next and the relative body condition of her partner.Females mating with males of higher body condition in yearx + 1 produced relatively male-biased sex ratios, and the oppositewas true for females mated with lower condition males. Within-individualanalysis also allowed investigations of sex ratio in relationto partner change. There was no change in sex ratios of femalespairing with the same male; however, females pairing with anew male produced clutches significantly more female biased. Comparisons of clutch sex ratios within individuals may be apowerful method for detecting sex ratio variation, and perhapsfemale birds may indeed manipulate egg sex but require personalcontextual experience for such decisions.     

Male mating success and female mate choice in the river sculpin,Cottus nozawae (Cottidae)

Synopsis Mating success of males and its correlates were investigated in a natural population of the polygynous fluvial sculpinCottus nozawae. Furthermore, the female mate preference of this species was examined experimentally under alternative conditions for mating in a stream. The mating success of individual males (the number of females with which a male mated) ranged between 0 and 8 with a mean of 2.41 in 1983 and 2.52 in 1989, in a population of which the sex ratio was about 1 : 2 in both years, skewed toward females. Mainly due to the excess of nests without egg masses and the few nests with one egg mass, the distribution of male mating success did not fit a Poisson distribution, indicating its non-randomness. Male mating success was not correlated either with the size of the nest rocks or with the male size, suggesting that these two variables are not determinants of mating success. The mate choice experiments demonstrated that females of this species more frequently chose smaller males as mates whose nests already contained eggs than large males without eggs. Additionally, an analysis of stomach contents of guarding males suggested that the parental males ate their own eggs during egg guarding (filial-cannibalism). Based on these results and on a comparison of reproductive characteristics with congeneric species, it is suggested that one of the most important determinants for female mate choice inCottus species may be whether or not parental males are filial egg cannibals.     

Polygyny in the Willow Warbler Phylloscopus trochilus in Swedish Lapland

The breeding sex ratio of Willow Warblers Phylloscopus trochilus in a subalpine birch forest was strongly female biased. Very few territorial males remained unmated and most became polygynous. Bigyny was the rule, but some males probably had three females. Male assistance at secondary nests varied and was probably dependent on temporal distance between the females' nests. There was no significant reduction in reproductive output for these females. Variation in the sex ratio of breeding birds can be explained, at least partly, by variation in the timing of male settlement.     

Is egg carrying attractive? Mate choice in the golden egg bug (Coreidae,Heteroptera)

In several species of fish, females select males that are already guarding eggs in their nests. It is a matter of debate as to whether a female selects a good nest site for her offspring (natural selection) or a male for his attractiveness (sexual selection). The golden egg bug, Phyllomorpha laciniata Vill, resembles fish in the sense that mating males carry more eggs than single males, but in the bugs, female mate choice is decoupled from egg site choice. The sexual selection hypothesis predicts that if females select males using male egg load as a cue for male quality, they should not mate with a male when eggs are removed, regardless of his mating attempts. When individual females were enclosed with an egg-loaded male and an unloaded male, they mated equally often with both males, although the loaded males courted more. In addition, when only successful males were used, females mated equally often with the loaded male and the unloaded male irrespective of sex ratio. Male choice rather than female choice affected mating frequency when sex ratio was equal. Therefore, females do not select the male by the eggs he carries, but successful males may receive many eggs due to egg dumping by alien females while they mate or as a consequence of mate guarding.     

Intranidal mating,emergence, and sex ratio in a communal beeAndrena jacobi Perkins 1921 (Hymenoptera: Andrenidae)

The extent of preemergence intranidal mating, schedules of emergence, and sex ratio at emergence were documented forAndrena jacobi, a communal, univoltine bee, by collecting and dissecting adults as they emerged from their fossorial nests in 1994. Over 70% of females had mated intranidally with nestmate males, thus potentially incestuously, before emerging. Preemergence intranidal mating did not vary during a day or between nests within a day, though it was less frequent at the start and end of the period of emergence. It was independent of a female's size.A. jacobi was protandrous, though some males emerged after all females. The sex ratio at emergence was remarkably female biased, possibly a consequence of local mate competition. Intranidal mating may represent a characteristic trait of communal bees where a high density of receptive females are predictably aggregated within a nest.     

Factors affecting sex ratio in rearing ofCoeloides sordidator (Hym.: Braconidae)

Five factors known to affect the sex ratio (% of males) in parasitic Hymenoptera were investigated forCoeloides sordidator, a parasitoid ofPissodes weevils. The host age, the age of ovipositing females, and the host of origin had a significant impact on the sex ratio of offspring. In contrast, the number of ovipositing females had an insignificant effect on sex ratio whereas the effect of host density could not be clearly defined. The sex ratio decreased with host age, probably because, like many other hymenopteran parasitoids, females tend to lay male eggs on small hosts and female eggs on larger hosts in order to maximize the size and fitness of their female offspring. The sex ratio also varied with the age of the mother, younger females laying more male eggs and older females more female eggs. The host of origin also had an influence on sex ratio. The strain fromPissodes castaneus was significantly more male-biased than the strain fromP. validirostris, which corroborates previous observations made on field populations     

Nesting patterns of European hedgehogs (Erinaceus europaeus) under northern conditions

Little information exists on the nesting of hedgehogs (Erinaceus europaeus) in the harsh northern environment, although this aspect has specific importance for the species’ survival in this region. We studied the nest types and nesting site selection of radio tagged hedgehogs in urban areas in Eastern Finland. Altogether, four different nest types were discovered and one of them (the pre-hibernation nest) is described here for the first time. Hedgehogs preferred residential areas for nesting during the mating and post-mating seasons while in the pre-hibernation season and hibernation period nesting was more common in forest areas. Nest constructions were closely related to materials found in the nest vicinity. During the active period, males used on average 24 nests and changed them 30 times, whereas females used 17 nests and changed them 26 times. The number of female nests and frequency of nest change increased from post-mating to pre-hibernation seasons, while in males both decreased. In general, female and male nests were situated within their established long-term home range, but in the pre-hibernation season over 30 % of male nests were situated outside of it. In post-mating and pre-hibernation seasons, females avoided other individuals of the same sex; this was demonstrated by the lack of nests in other female core areas. Hedgehogs hibernate for around 223 days, which is the longest reported hibernation period for this species. The results of the present study emphasise the importance of forest and diverse residential areas including forest patches for the hedgehog’s nesting habitat.     

Growth and maturity of the spider crab Halicarcinus planatus (Brachyura: Hymenosomatidae) females in the southwestern Atlantic Ocean. Can these parameters be influenced by the population sex ratio?

The Halicarcinus planatus populations of the southwestern Atlantic Ocean show a highly variable sex ratio and a large size overlapping between females in the last immature instar (ADO) and mature (MAT) females. We hypothesized that these facts are related and that female impregnation has a central role in this relation. Non-impregnated ADO females delay maturity and would continue to grow, leading to size overlapping. This scenario is most probable in populations with a scarcity of males and could affect the growth, maturity and population structure. The sex ratio and female size distribution of several populations of the southwestern Atlantic Ocean were analysed. Growth and maturity were studied in two populations with different sex ratios (Camarones, a population with males, and Rada Tilly, a population where no males were found). Size overlapping was not related to the population sex ratio. Size overlapping was due to a variable moult increment and, to a lesser extent, to delayed maturity. Females mated before terminal moult in both populations. However, in Camarones, physiological maturity was not related to impregnation, while in Rada Tilly, vitellogenic oocytes were observed only in impregnated females. Also, differences in the number of spermatophores in female spermathecae were related to the sex ratio. Our results showed that the morphological maturity pattern of females was not influenced by the population sex ratio. However, physiological maturity was related to female impregnation and sex ratio. Also, the variation in the number of spermatophores suggests differences in the mating system related to the sex ratio.