Phenology of two territorial solitary bees, Anthidium manicatum and A. florentinum (Hymenoptera: Megachilidae)

Five years' data on phenology of an Anthidium manicatum population in southern Germany and comparative observations on A. manicatum and A. florentinum from southern France are analysed. Males and females had the same flight season, adult sex ratio was strongly female biased and males were larger than females in both species. This is the opposite pattern to most other solitary bees, where females generally are larger than males, sex ratio is male-biased, males emerge before females and males disappear long before females. We argue that two features of Anthidium female behaviour, namely prolonged sexual receptivity and use of resources easily defendable by males, explain male adaptations in behaviour, phenology and body size and, hence, population sex ratio.     

雌雄异熟植物露蕊乌头开花时间对雌雄功能期及表型性别的影响

开花时间决定了植物雌雄功能的交配机会, 最终影响繁殖成功。交配环境假说认为雌雄异熟植物开花时间的差异能引起植物表型性别的变异, 改变种群内的交配环境, 影响植物对雌雄功能的最佳性分配。为了研究开花时间对雌雄异熟植物的雌雄性别时期及表型性别的影响, 本文以毛茛科雄性先熟植物露蕊乌头(Aconitum gymnandrum)为实验材料, 记录了雄性和雌性功能期, 分析了植株开花时间、花的雌雄功能期和表型性别的关系。结果表明: 在植物同一花序内, 较晚开放的花有更长的雄性期和更短的雌性期, 性分配在时间上偏雄。雌雄功能期在时间上的相对分配随植物开花时间的变化表现出相似的趋势： 较晚开的花或较晚开花的个体, 花的雄性功能期相对于雌性功能期更长, 在时间上更偏向雄性功能。而且, 开花时间的差异影响种群内花的性比和植物个体的表型性别动态。随着开花时间由早到晚的变化, 种群内早期以雄花为主,末期以雌花为主, 种群内性别环境由偏雄向偏雌变化, 因此植株个体的平均表型性别则从偏雌转向偏雄。本文结果支持交配环境假说, 雄性先熟的露蕊乌头开花早期, 种群内花的性别比偏雄, 种群表型性别环境偏雄, 因而植物个体平均表型性别偏雌, 性别分配(即时间分配)偏向雌性功能, 而晚开花个体的平均性别偏雄, 更偏向雄性功能的分配。     

Lack of Sex-Biased Maternal Investment in spite of a Skewed Birth Sex Ratio in White-Headed Langurs (Trachypithecus leucocephalus)

Numerous hypotheses have been developed to explain sex allocation. In male-dispersing, female cooperatively breeding species, the local resource competition model predicts male-biased birth sex ratio, the local resource enhancement model predicts female-biased birth sex ratio, and the population adjustment model predicts that biased birth sex ratio should not be favored if the two sexes are equally costly to rear. The male quality model predicts that, in polygynous species, females in better physical condition will either produce more sons than daughters or invest more heavily in sons than in daughters. White-headed langurs are a female philopatry and female cooperatively breeding species. During a 11-yr study, a total of 133 births were recorded, among which birth sex ratio (M:F = 73:49) was significantly male-biased. This is consistent with the prediction of the local resource competition model. On the other hand, if mothers balanced their investment between the two sexes, according to Fisher's population adjustment model, males should be the less-costly-to-rear sex. However, we found no sex difference for infant mortality (12.3% in males and 12.2% in females), and sons induced slightly longer interbirth interval (son: 26.4 ± 1.1 mo, daughter: 24.1 ± 0.6 mo) and lactational period (son: 20.9 ± 1.0 mo, daughters: 19.6 ± 0.5 mo) for their mothers. Thus, the population adjustment model was not supported by this study. The local resource enhancement model was not supported because birth sex ratio did not bias to females who provided more reproductive assistance. On the individual level, probit regression showed no relation between birth sex ratio and group size. Because the group size was considered to be negatively related to female physical condition, our study did not support the male-quality model. We suggested several possibilities to explain these results.     

Sex ratio and sexual dimorphism in the dioecious Borderea pyrenaica (Dioscoreaceae)

Sex ratio and sexual dimorphism of Borderea pyrenaica, a long-lived dioecious geophyte endemic to the Pyrenees (north-east Iberian Peninsula), were examined in three alpine populations. In this species, age can be estimated and the sex of nonreproductive adult plants identified. Male plants attain sexual maturity earlier, flower more frequently and grow faster than female plants, whereas females allocate a higher biomass to reproduction than males. These results support the hypothesis that female plants incur a higher cost of sexual reproduction and that this higher cost is measurable as reduced vegetative growth and lower flowering frequency. Variation of sex ratio among young, intermediate and old adults within populations suggests, however, that this higher female reproductive investment does not result in sexual differences in mortality. The overall male-biased sex ratio in B. pyrenaica is mainly a consequence of the tendency of males to reproduce at an earlier age and more frequently than females.     

Male-biased sex allocation in late-blooming flowers driven by resource limitation in the clonal perennial Aconitum kusnezoffii (Ranunculaceae)

Flowering phenology and clonal growth are known to affect resource and pollen availability, and therefore select for adaptive or constrained sex allocation strategies to some degree. However, the consequences of temporal sex allocation patterns for reproductive fitness across the flower, inflorescence, and genet levels have rarely been examined. Moreover, experimental tests of the underlying regulatory mechanisms are scarce. We examined the association of flowering phenology and inflorescence position with temporal sex allocation and reproductive success in the protandrous perennial clonal herb, Aconitum kusnezoffii, over four consecutive growing seasons by examining more than 39 000 flowers. We also conducted controlled experiments to test the effects of resource and pollen limitation on the female reproductive success of lateral inflorescences. We found that some male functions were positively correlated with flowering phenology, whereas female reproductive success was negatively correlated with flowering phenology and inflorescence position. Lateral inflorescences invested more in male function than terminal inflorescences and therefore yielded fewer and smaller seeds. Resource limitation may serve as the key mechanism underlying this differentiated pattern. Decreased female reproductive success was consistently observed at the flower and inflorescence levels as flowering occurred later in the growth season. Late-blooming lateral inflorescences specialized in the male function, and their female reproductive success was constrained by early-blooming terminal inflorescences. This might be the first attempt to systematically demonstrate sex allocation strategy differentiation in a protandrous plant species at the inflorescence level. In addition, our study provides empirical evidence of dichogamy selecting for specialized sex allocation strategies among inflorescences.     

Home range size and activity of the wood lemming, Myopus schisticolor

We present the results from a radiotelemetric study on space use and activity in a natural population of the wood lemming Myopus schisticolor. Male home ranges were larger than females, and the ratio female home range to male home range was smaller than expected compared to other small rodents. Males moved distances 4-12 times longer than females. We argue that this difference in mobility gives a higher probability of capturing males than females in snap trap studies. If there is a sex biased trappability, this might explain the increasing female biased sex ratio during the snow free season (1:1 in spring, 3:1 [female:male| in autumn), as a higher mortality of males during the reproductive season.     

Conflict over multiple-partner mating between males and females of the polygynandrous common lizards

The optimal number of mate partners for females rarely coincides with that for males, leading to a potential sexual conflict over multiple-partner mating. This suggests that the population sex ratio may affect multiple-partner mating and thus multiple paternity. We investigate the relationship between multiple paternity and the population sex ratio in the polygynandrous common lizard (Lacerta vivipara). In six populations the adult sex ratio was biased toward males, and in another six populations the adult sex ratio was biased toward females, the latter corresponding to the average adult sex ratio encountered in natural populations. In males the frequency and the degree of polygyny were lower in male-biased populations, as expected if competition among males determines polygyny. In females the frequency of polyandry was not different between treatments, and polyandrous females produced larger clutches, suggesting that polyandry might be adaptive. However, in male-biased populations females suffered from reduced reproductive success compared to female-biased populations, and the number of mate partners increased with female body size in polyandrous females. Polyandrous females of male-biased populations showed disproportionately more mating scars, indicating that polyandrous females of male-biased populations had more interactions with males and suggesting that the degree of multiple paternity is controlled by male sexual harassment. Our results thus imply that polyandry may be hierarchically controlled, with females controlling when to mate with multiple partners and male sexual harassment being a proximate determinant of the degree of multiple paternity. The results are also consistent with a sexual conflict in which male behaviors are harmful to females.     

Experimental evolution exposes female and male responses to sexual selection and conflict in Tribolium castaneum

Between-individual variance in potential reproductive rate theoretically creates a load in reproducing populations by driving sexual selection of male traits for winning competitions, and female traits for resisting the costs of multiple mating. Here, using replicated experimental evolution under divergent operational sex ratios (OSR, 9:1 or 1:6 ♀:♂) we empirically identified the parallel reproductive fitness consequences for females and males in the promiscuous flour beetle Tribolium castaneum. Our results revealed clear evidence that sexual conflict resides within the T. castaneum mating system. After 20 generations of selection, females from female-biased OSRs became vulnerable to multiple mating, and showed a steep decrease in reproductive fitness with an increasing number of control males. In contrast, females from male-biased OSRs showed no change in reproductive fitness, irrespective of male numbers. The divergence in reproductive output was not explained by variation in female mortality. Parallel assays revealed that males also responded to experimental evolution: individuals from male-biased OSRs obtained 27% greater reproductive success across 7-day competition for females with a control male rival, compared to males from the female-biased lines. Subsequent assays suggest that these differences were not due to postcopulatory sperm competitiveness, but to precopulatory/copulatory competitive male mating behavior.     

Behavioral and physiological female responses to male sex ratio bias in a pond-breeding amphibian

Introduction

The phenomenon of sexual conflict has been well documented, and in populations with biased operational sex ratios the consequences for the rarer sex can be severe. Females are typically a limited resource and males often evolve aggressive mating behaviors, which can improve individual fitness for the male while negatively impacting female condition and fitness. In response, females can adjust their behavior to minimize exposure to aggressive mating tactics or minimize the costs of mating harassment. While male-male competition is common in amphibian mating systems, little is known about the consequences or responses of females. The red-spotted newt (Notophthalmus viridescens) is a common pond-breeding amphibian with a complex, well-studied mating system where males aggressively court females. Breeding populations across much of its range have male-biased sex ratios and we predicted that female newts would have behavioral mechanisms to mitigate mating pressure from males. We conducted four experiments examining the costs and behavioral responses of female N. viridescens exposed to a male-biased environment.

Results

In field enclosures, we found that female newts exposed to a male-biased environment during the five-month breeding season ended with lower body condition compared to those in a female-biased environment. Shorter-term exposure to a male-biased environment for five weeks caused a decrease in circulating total leukocyte and lymphocyte abundance in blood, which suggests females experienced physiological stress. In behavioral experiments, we found that females were more agitated in the presence of male chemical cues and females in a male-biased environment spent more time in refuge than those in a female-biased environment.

Conclusions

Our results indicate that male-biased conditions can incur costs to females of decreased condition and potentially increased risk of infection. However, we found that females can also alter their behavior and microhabitat use under a male-biased sex ratio. Consistent with surveys showing reduced detection probabilities for females, our research suggests that females avoid male encounters using edge and substrate habitat. Our work illustrates the integrated suite of impacts that sexual conflict can have on the structure and ecology of a population.     

EFFECT OF DENSITY ON SECONDARY SEX CHARACTERISTICS AND SEX RATIO IN SILENE ALBA (CARYOPHYLLACEAE)

A field survey of plant and flower sex ratio and secondary sex characteristics was made in Silene alba. Female-biased plant sex ratios were found, as seems typical for the species. Sex ratio distribution correlated with a gradient of soil moisture (with the more moist area having a more female-biased ratio) and with changes in the density of Silene (intermediate and higher density areas having greater female bias). The floral sex ratio was significantly female-biased only at the site that was most female-biased in terms of plant sex ratio. Otherwise the population of flowers was significantly male-biased. Male and female plants harvested from the field differed in secondary sexual characteristics. Males had more flowers and invested proportionately more biomass in leaf, but less in root, stem and reproductive tissue than did females. Although both males and females were larger in terms of total dry weight at the moist site, males produced more flowers at the driest (high density) site. Here the female bias in plant sex ratio was intermediate, but the floral sex ratio was significantly male-biased. A glasshouse experiment was performed in which plants were grown at four densities. Density significantly influenced plant survivorship and the probability of flowering, and increased female bias in the pots, but it did not affect patterns of biomass allocation in flowering plants. Patterns of male and female biomass allocation did not differ in the experiment, except in terms of reproductive allocation (greater in females) and allocation to leaf, greater in males, but only at the lowest density. This work urges caution in interpreting differences between males and females in the field as secondary sex characteristics, since we find such properties to be overlapping under experimental conditions. It supports the idea that males and females of a species may sustain different reproductive output under differing conditions.     

Causes of a male-biased operational sex ratio in the fiddler crab<Emphasis Type="Italic"> Uca crenulata</Emphasis>

This research investigates the causes of a male-biased operational sex ratio in a population of the California fiddler crab, Uca crenulata. Mensurative studies revealed there were almost twice as many adult males as females, mating occurred across half of the days within the breeding season, and females had much longer individual reproductive cycles than males. Therefore, many more males than females were available for mating on each breeding day. Perhaps as a consequence, males spent a large proportion of their time fighting with neighbors and rapidly waving their large claws when females passed by. Electronic Publication     

Sex change towards female in dying Acer rufinerve trees

BACKGROUND AND AIMS: Sex changes within the genus Acer (Aceraceae) may occur because of associations of sex expression and plant health. In this study, a natural population of Acer rufinerve was monitored to clarify the sex change patterns, the relationship between sex expression and plant health, and the causal environmental conditions that precede sex changes. METHODS: Sex expression, growth rate and mortality of A. rufinerve trees in a natural population were monitored from 1992 to 1997. KEY RESULTS: Three types of sex expression were observed among A. rufinerve: male, female and bisexual. Among the three types of sex expression, sex changes occurred in all directions. In the growing season of 1994, precipitation was reduced. Stem growth rate decreased and mortality was high in 1994. In the spring of 1995, a drastic sex change from male to female or to bisexual occurred. As a result, the sex ratio became female-biased in 1995, although it had been male-biased from 1992 to 1994. In 1996 and 1997, the proportion of males in the population increased, partly as a result of female mortality and partly as a result of female-to-male sex changes. Sex expression of A. rufinerve was associated with their growth rate and mortality. The growth rate decreased for trees whose sex changed from male to female or to bisexual, and increased for trees whose sex changed from female to male or to bisexual. Dead trees reproduced as females before they died, except for those that died as males in 1994. CONCLUSIONS: One explanation for the sex change towards increasing femaleness for this A. rufinerve population in 1995 was the deterioration of plant health in the previous growing season, because of reduced precipitation. Sex changes of unhealthy and dying A. rufinerve towards femaleness may facilitate re-occupancy by offspring in gaps created by the death of A. rufinerve trees.     

Determinants of natural mating success in the cannibalistic orb-web spider Argiope bruennichi

Monogynous mating systems (low male mating rates) occur in various taxa and have evolved several times independently in spiders. Monogyny is associated with remarkable male mating strategies and predicted to evolve under a male-biased sex ratio. While male reproductive strategies are well documented and male mating rates are easy to quantify, especially in sexually cannibalistic species, female reproductive strategies, the optimal female mating rate, and the factors that affect the evolution of female mating rates are still unclear. In this study, we examined natural female mating rates and tested the assumption of a male-biased sex ratio and female polyandry in a natural population of Argiope bruennichi in which we controlled female mating status prior to observations. We predicted variation in female mating frequencies as a result of spatial and temporal heterogeneity in the distribution of mature females and males. Females had a low average mating rate of 1.3 and the majority copulated only once. Polyandry did not entirely result from a male-biased sex-ratio but closely matched the rate of male bigamy. Male activity and the probability of polyandry correlated with factors affecting pheromone presence such as virgin females' density. We conclude that a strong sex ratio bias and high female mating rates are not necessary components of monogynous mating systems as long as males protect their paternity effectively and certain frequencies of bigyny stabilise the mating system.     

Manipulation of offspring sex ratio by second-mated female house wrens

In 1973, Trivers and Willard proposed that offspring sex ratio should be associated with the quality of parental care likely to be provided to the offspring. We tested this hypothesis by comparing fledgling sex ratios in nests of first- and second-mated female house wrens (Troglodytes aedon). In our Wyoming population, second-mated females typically receive little or no male parental assistance and fledge fewer and lower-quality young compared with first-mated females. Assuming that being of lower quality has stronger negative effects on the future reproductive success of males than that of females in this polygynous population, we predicted that fledgling sex ratios in the nests of second-mated females would be female-biased compared with the fledgling sex ratios of first-mated females. Additionally, we asked whether any sex bias at fledging could have resulted from male-biased nestling mortality caused by sex-biased parental provisioning. As predicted, mean fledgling sex ratios in nests of second-mated females were more female-biased than fledgling sex ratios in nests of first-mated females. However, we found no evidence of either sex-biased nestling mortality or sex-biased parental provisioning. These findings suggest that females are responding to their status as second-mated females and to the associated low-quality parental care that their young are likely to receive by producing female-biased clutches rather than manipulating the offspring sex ratio through sex-biased nestling mortality.     

Size structure, sex ratio and onset of sexual maturity of two species of Hypostomus

Length-frequency analyses showed that male Hypostomus affinis are bigger than females, but H. luetkeni showed no such differences. The sex ratio was estimated at 1:1 for both species. The temporal variation of sex ratio indicated a decrease in the male population when GSI was at its maximum value, suggesting that males are less prone to capture during the spawning season. According to the decrease of males during the reproductive period, paternal behaviour is proposed for both species. H. affinis reached sexual maturity at a smaller size than H. luetkeni. Considering the length weight relationship and the mortality rate of the reproductive females in both species, the physiological consequences of delay or precocity of the first maturation are discussed.     

饲养林麝的种群特征和幼麝的存活研究

对饲养在上海崇明岛东平林场的林麝种群特征以及1998～2000 年间该种群所出生的74 只幼麝存活情况进行分析研究。结果表明: 该林麝种群中能繁殖的个体占51.1 % , 种群数量逐渐增长。种群存活率曲线呈C 型,说明林麝的死亡主要发生在幼麝时期。从不同年龄段性比变化来看, 初生幼麝的性比接近1∶1 , 由于雌幼麝比雄幼麝具有更高的存活率( P < 0.05) , 进入亚成体以后雄性比例逐渐下降, 显示幼麝时期存在偏雄性死亡。早出生幼麝的存活率明显高于晚出生幼麝( P < 0.05) 。引起幼麝死亡的主要原因是疾病(45. 9 %) 。     

花尾榛鸡的非对称性比和成体存活率

1999至2002年,在法国东南部的阿尔卑斯山,通过无线电追踪方法研究了花尾榛鸡的性比问题。首先,我们通过直接观测无线电标记雄性的配对比例,在消除可能的捕捉误差后,发现繁殖种群性比为1．26（即未配对的雄性占21％）。然后,我们估计了标记个体的存活率和繁殖成功率。在假设窝扩散时幼体的性比平衡的前提下,通过两性和两个年龄组（幼体,〈10月龄的个体;成体〉10月龄的个体）的确定性统计模型,得到的理论性比为1．22（即未配对的雄性占18％）,这一结果与实际观测的性比十分接近。我们认为,雌性死亡率高导致的两性存活率差异可以解释雄性性比偏高现象。我们进而讨论了花尾榛鸡整个分布区内居间强度的非对称性比的适应意义,以及在衰退的花尾榛鸡种群中雌性数量极端不足的后果[动物学报52（4）：655—662,2006]。     

No differential consequences of reproduction according to sex in Juniperus communis var. depressa (Cupressaceae)

In dioecious plant species, males and females are thought to have dissimilar allocation patterns. Females are believed to invest more in reproduction and less in growth and maintenance than males. This differential investment between sexes could result in distinct growth patterns and contrasting survival rates, thereby affecting the sex ratio of a population and the age and size distribution of males and females, possibly leading to habitat segregation according to sex. These effects might become more apparent under particularly limiting conditions, such as in nutrient-deficient soils or in climatically stressed environments. To verify these predictions, growth patterns, microsite characteristics, and age and size distribution of male and female individuals were compared, and population sex ratio was determined in three populations of the dioecious shrub Juniperus communis var. depressa (Cupressaceae, Pinophyta) along a short latitudinal gradient on the eastern coast of Hudson Bay (Northern Québec, Canada). We found that the northernmost population had a male-biased sex ratio, but that the southernmost one had a higher proportion of females. Our results failed to reveal any significant differences in radial growth patterns, mean sensitivity, annual elongation of the main axis, and size and age frequency distribution between males and females in any population. Furthermore, there was no evidence of microhabitat segregation according to sex as indicated by the lack of differences in the physicochemical characteristics of the substrate under males and females. Clearly, the expected ecological consequences of a presumed greater investment of females in reproduction were not apparent even under the very stressful conditions prevailing on subarctic dunes. Many factors could reduce differences in the cost of reproduction between males and females, such as the number and quality of reproductive structures produced annually by individuals of each sex, the possible photosynthetic activity of the immature female cones, and the complexity of the source/sink relationship within individuals. Alternatively, there may be no differences between sexes in their reproductive investment.     

The effect of drought stress on the sex ratio variation ofSilene otites

Under dry environmental conditions the sex ratio of many dioecious plants is male-biased, which is usually explained by the higher susceptibility of females to drought stress. We investigated if spatio-temporal variation in the sex ratio ofSilene otites could be explained by the higher sensitivity of female plants to drought stress as compared to males. Long-term field observations, however, did not support this hypothesis. The sex ratio in 34 patches at the study site in Central Germany changed from slightly female biased in 1994 to strongly male-biased in 1997 and 1998. The interannual change in the proportion of plants that were female was positively correlated with the number of days with soil-water deficit in the late summer, suggesting higher mortality in males than in females under drought stress. In two closely studied patches, mortality in males was also higher than in females, although this difference could not be related to drought stress. These field observations were supported by an experiment with potted plants in two climate chambers, in which male mortality was higher during a three-week period without water supply. We conclude that the often reported male bias in patches ofS. otites is not caused by sexual differences in the sensitivity to drought stress. Field data in this study, however, suggest that maleS. otites plants flower earlier than females, which causes a shift in sex ratio to more male bias among flowering plants.     

Male‐skewed adult sex ratio,survival, mating opportunity and annual productivity in the Snowy Plover Charadrius alexandrinus

Sex differences in adult mortality may be responsible for male‐skewed adult sex ratios and male‐skewed parental care in some birds. Because a surplus of breeding males has been reported in serially polyandrous populations of Snowy Plover Charadrius alexandrinus, we examined sex ratio, early‐season nesting opportunities, adult survival and annual reproductive success of a Snowy Plover population at Monterey Bay, California. We tested the hypotheses that male adult survival was greater than female survival and that a sex difference in adult survival led to a skewed adult sex ratio, different mating opportunities and different annual productivity between the sexes. Virtually all females left chicks from their first broods to the care of the male and re‐nested with a new mate. As a result, females had time to parent three successful nesting attempts during the lengthy breeding season, whereas males had time for only two successful attempts. Among years, the median population of nesting Plovers was 96 males and 84 females (median difference = 9), resulting in one extra male per eight pairs. The number of potential breeders without mates during the early nesting period each year was higher in males than in females. Adult male survival (0.734 ± 0.028 se) was higher than female survival (0.693 ± 0.030 se) in top‐ranked models. Annually, females parented more successful clutches and fledged more chicks than their first mates of the season. Our results suggest that in C. alexandrinus a sex difference in adult survival results in a male‐skewed sex ratio, which creates more nesting opportunities and greater annual productivity for females than for males.