Effect of sex ratio,habitat factors and neighborhood competition on stem growth in the dioecious tree Fraxinus mandshurica

Growth rates of male and female trees are often different in a dioecious species. In this study, we analyzed sex ratios and the effect of gender, neighborhood competition and habitat factors on the stem growth of dioecious Fraxinus mandshurica trees in a secondary conifer and broad-leaved mixed forest in the Changbai Mountains of northeastern China. The sex ratio in the 5.2-ha study area does not deviate significantly from the expected 1:1 ratio, except for trees in the large diameter classes. For dbh >40 cm, the sex ratio is male-biased. This result suggests that males have a faster rate of stem growth than females, which is usually explained by the higher cost of reproduction in the fruit-bearing females. An analysis of the dbh distributions of two successive measurements showed that the rate of stem growth of the (27) females drops off with increasing dbh and remains below that of the (35) males. A causal model was used to analyze factors affecting the rate of stem growth, showing that these rates are affected significantly and positively by soil moisture and tree size in both genders and that within-gender competition is mainly for nitrogen. Our study suggests that neighborhood competition does not affect stem growth significantly, which is a rather surprising result.     

Determinants of biased sex ratios and inter-sex costs of reproduction in dioecious tropical forest trees

Estimates of the sex ratio and cost of reproduction in plant populations have implications for resource use by animals, reserve design, and mechanisms of species coexistence, but may be biased unless all potentially reproductive individuals are censused over several flowering seasons. To investigate mechanisms maintaining dioecy in tropical forest trees, we recorded the flowering activity, sexual expression, and reproductive effort of all 2209 potentially reproductive individuals within 16 species of Myristicaceae over 4 years on a large forest plot in Amazonian Ecuador. Female trees invested >10 times more biomass than males in total reproduction. Flowering sex ratios were male-biased in four species in ≥1 year, and cumulative 4-year sex ratios were male-biased in two species and for the whole family, but different mechanisms were responsible for this in different species. Annual growth rates were equivalent for both sexes, implying that females can compensate for their greater reproductive investment. There was no strict spatial segregation of the sexes, but females were more often associated with specific habitats than males. We conclude that male-biased sex ratios are not manifested uniformly even after exhaustive sampling and that the mechanisms balancing the higher cost of female reproduction are extremely variable.     

Male-Biased Capture in Light Traps in <Emphasis Type="Italic">Spodoptera exigua</Emphasis> (Lepidoptera: Noctuidae): Results from the Studies of Reproductive Activities

Nocturnal insects show phototactic behavior. Interestingly, males are caught by light traps more often than females in many species. In the present study, the male-biased capture in light traps in the field was evaluated to test the hypothesis that male-biased capture was associated with the reproductive activities of Spodoptera exigua. During a 16-wk. period in 2011, a total of 5075 moths caught by light traps in the field revealed a significant male-biased capture. However, the sex ratio of the natural population was not significantly different from 1:1 from July to October 2011. The ratio of moths caught per hour by light traps showed that females were captured in the first half of the night, whereas the peak capture by light traps for males occurred in the second half of the night. Among the captured adults, the percentage of unmated females was significantly lower than that of the mated females and males. Experiments in the laboratory indicated that the time of activities for both sexes was consistent with the active patterns of oviposition and copulation. The percentage of flight-to-light of the unmated females was significantly lower than that of the mated females and males. Thus, the lower capture of the unmated females, which is associated with reduced positive phototactic responses of S. exigua, probably is one of the reasons resulting in the male-biased sex ratio in light traps.     

Sex ratio from germination through maturity and its reproductive consequences in the liverwort Sphaerocarpos texanus

Summary The dioecious winter ephemeral liverwort, Sphaerocarpos texanus disperses spore tetrads consisting of two males and two females. I examined the subsequent sex ratio of S. texanus at different stages in its life cycle to detect possible mechanisms affecting deviations from a 1:1 sex ratio and the effect of sex ratio on reproductive success. As S. texanus occurs in pure male, pure female and mixed sex clumps, I examined the proportions and sizes of these, the reproductive success of pure female and mixed sex clumps in the field and the sex ratio of germinating plants in a growth chamber. In both the field and the growth chamber the most abundant clump type was pure female followed by mixed sex and pure male clumps. These abundance patterns suggest that males have a lower survival rate than females before emergence and this lower survival rate continues through the gametophytic stage. This disadvantage may be due to the higher susceptibility of males to environmental conditions, to their competitive inferiority to females, and/or to differential resource allocation to the sexes within the spore tetrad. The female biased sex ratio at germination is consistent with predictions from sex ratio theory. Further my field data indicate that males may gain a survival benefit from growing in a mixed sex clump and both males and females benefit reproductively when they occur in mixed sex clumps.     

Do secondary sexual dimorphism and female intolerance to drought influence the sex ratio and extinction risk of <Emphasis Type="Italic">Taxus baccata</Emphasis>?

Sex ratio and sexual dimorphism were studied in the dioecious tree Taxus baccata. We examined five populations of T. baccata in Poland and Ukraine to identify the differences between male and female individuals. The sex of all individuals, height and diameter, needle length and area, specific leaf area (SLA), the number of stomata rows, stomatal density, and content of carbon and nitrogen were measured to identify the differences between male and female individuals. The relationship between sex ratio and climatic conditions, age and population size were analysed using data collected from the field and the literature. Female trees were shorter than males, but needles of females were longer and had larger area. Although there were no differences among sexes in SLA, nitrogen and carbon concentration, we found a positive correlation between nitrogen concentration and SLA among females. The sex ratio changed with tree height within populations, and taller height classes were biased in favour of males. Regardless of population age, the percentage of females within populations was positively correlated with precipitation. Probably high reproductive effort caused female trees to lose in competition with males, and this loss may also be enhanced by lower drought tolerance in females and could contribute to risk of extinction for T. baccata. The continental geographic range of T. baccata may be restricted by limited occurrence of females, which demand higher water resources than males.     

不同林龄阔叶红松林林下簇毛槭的性比格局及雌雄个体的空间分布

为探讨不同发育阶段林分下的雌雄异株植物与性别相关的性比格局和空间分布, 以5.2 ha的中龄林和1.0 ha的老龄林固定监测样地内簇毛槭(Acer barbinerve)雌、雄植株的定位观测数据为基础, 对比分析了长白山不同林龄的阔叶红松林中的已花簇毛槭的性比格局、空间分布及其与环境因子间的关系。研究结果表明: 中龄林和老龄林中雌树的胸径均显著大于雄树, 总体上性比极显著偏离1:1。随着树木的生长, 性比由偏雄性转变为不再偏离1:1, 这可能是因为雄树始花胸径较小所致。O-ring单变量点格局分析显示中龄林样地中的雌树和雄树符合异质性泊松分布, 老龄林样地中的雌树和雄树均完全随机分布。O-ring双变量点格局分析显示, 在随机标签假设下, 中龄林中的雌树和雄树在1-4 m尺度上空间负相关, 在4-100 m尺度上空间独立, 老龄林中的雌树和雄树在所有尺度上空间独立。簇毛槭在中龄林和老龄林中不同的空间分布格局说明中龄林中簇毛槭分布的斑块性相对明显, 随着林分的发育, 郁闭度较高的老龄林样地中环境异质性降低, 环境因子对簇毛槭分布的影响减弱。典范冗余分析(redundancy analysis, RDA)表明在中龄林中, 林分密度只能解释3.73%的雌树分布的变异, 与雄树分布的相关性不显著, 叶面积指数和非生物因子对雌树和雄树的影响均较弱; 老龄林中簇毛槭的分布与生物因子和非生物因子的相关性均不显著。     

Sexual differences in reproductive phenology and their consequences for the demography of Baccharis dracunculifolia (Asteraceae), a dioecious tropical shrub

Patterns of phenological variation and reproductive investment were studied in the dioecious shrub Baccharis dracunculifolia DC (Asteraceae), and possible consequences on survivorship were evaluated. The sex ratio was determined in a natural field population (n = 921) of B. dracunculifolia in Belo Horizonte, Brazil. Fifty-two males and 56 females were sampled at random from this population. During the reproductive season of 1999, inflorescence production, shoot growth and mortality, and xylem water potential were recorded for each individual. The population sex ratio was male-biased (1.27 : 1, P < 0.05), and was associated with a higher mortality of female shoots (38.4 vs. 23.1 %, P < 0.05), and individuals (17.8 vs. 11.5 %, P < 0.1), despite lower water stress in female plants. Flowering phenology also differed between the sexes, with males producing more inflorescences, and earlier, than females. Owing to fruit maturation, the number of inflorescences supported by females was higher than that supported by males later in the reproductive season. This occurred during the dry season, and drought stress may have been responsible for the greater female mortality. Thus, the male-biased sex ratio in this population of B. dracunculifolia is probably due to different reproductive functions of males and females. Intersexual differences in reproductive phenology had consequences for plant demography.     

Increased density and male-male interactions reduce male longevity in the medfly, Ceratitis capitata

We investigated the effect of adult density on male and female longevity and behavioural interactions in a mass-reared strain of the medfly. Male survival decreased significantly, and male-male behavioural interactions increased significantly, with increasing male density (males were kept at 1, 2, 6 and 10 flies per 285 cm²in pots of 5.5 cm in diameter and 12 cm high). No such effects were observed in females over the same density range. This suggests that male survival decreased in response to the deleterious effects of male-male behavioural interactions. At the highest density, courtship wing fanning directed towards other males decreased significantly over time. In addition, as either density or male-biased sex ratio increased, males were increasingly likely to be engaged in courtship wing fanning that was not directed towards females. These results suggest that high adult density may significantly reduce male reproductive success, by decreasing survival and mating success. The results highlight the potentially high survival costs of male-male interactions and indicate differences in the nature of reproductive costs in the two sexes.     

Reproductive barriers between populations of the cereal rust mite <Emphasis Type="Italic">Abacarus hystrix</Emphasis> confirm their host specialization

Recent studies have shown that certain host populations of the cereal rust mite Abacarus hystrix are highly specialized in their host use and it is likely that reproductive isolation exists between them. Here I verified this expectation by testing for reproductive barriers between ryegrass and quackgrass populations of A. hystrix. I performed reciprocal crosses between individuals from both populations and observed results of crosses. Leaves of the three grass species, ryegrass, quackgrass and wheat, were used as mating arenas. I used two criteria to determine reproductive barriers between strains: the proportion of female progeny and viability of progeny. If studied populations of this haplodiploid species are reproductively isolated a male-biased sex ratio and/or hybrid progeny of reduced viability would be expected. I found that in the presence of quackgrass and ryegrass pre-zygotic barriers between studied populations exists. On wheat I observed asymmetry in reproductive barriers. Between females from quackgrass and males from ryegrass a pre-zygotic barrier existed (only males obtained). However, the opposite reciprocal cross (females from ryegrass and males from quackgrass) produced progeny of both sexes. A male-biased sex ratio and low adult emergence suggests that post-zygotic mechanisms acted here. Low viability of progeny obtained from crosses in which females from ryegrass were engaged suggests that the origin of the female nymph acted as a predictor of hybrid inviability. The pattern of sterility observed in the cereal rust mite indicates that in the presence of three hosts (ryegrass, quackgrass and wheat) pre- and post-zygotic reproductive barriers between quackgrass and ryegrass populations of this mite exist. In addition to host fidelity (which acts as pre-zygotic barrier) there are post-zygotic barriers to gene flow.     

Endosymbionts moderate constrained sex allocation in a haplodiploid thrips species in a temperature-sensitive way

Maternally inherited bacterial endosymbionts that affect host fitness are common in nature. Some endosymbionts colonise host populations by reproductive manipulations (such as cytoplasmic incompatibility; CI) that increase the reproductive fitness of infected over uninfected females. Theory predicts that CI-inducing endosymbionts in haplodiploid hosts may also influence sex allocation, including in compatible crosses, however, empirical evidence for this is scarce. We examined the role of two common CI-inducing endosymbionts, Cardinium and Wolbachia, in the sex allocation of Pezothrips kellyanus, a haplodiploid thrips species with a split sex ratio. In this species, irrespective of infection status, some mated females are constrained to produce extremely male-biased broods, whereas other females produce extremely female-biased broods. We analysed brood sex ratio of females mated with males of the same infection status at two temperatures. We found that at 20 °C the frequency of constrained sex allocation in coinfected pairs was reduced by 27% when compared to uninfected pairs. However, at 25 °C the constrained sex allocation frequency increased and became similar between coinfected and uninfected pairs, resulting in more male-biased population sex ratios at the higher temperature. This temperature-dependent pattern occurred without changes in endosymbiont densities and compatibility. Our findings indicate that endosymbionts affect sex ratios of haplodiploid hosts beyond the commonly recognised reproductive manipulations by causing female-biased sex allocation in a temperature-dependent fashion. This may contribute to a higher transmission efficiency of CI-inducing endosymbionts and is consistent with previous models that predict that CI by itself is less efficient in driving endosymbiont invasions in haplodiploid hosts.Subject terms: Evolutionary genetics, Evolutionary ecology, Parasitology     

Phenology, sex ratio, and spatial distribution among dioecious species of Trichilia (Meliaceae)

The flowering, sex ratio, and spatial distribution of four dioecious species of Trichilia (Meliaceae) were studied in a semi-deciduous forest in southeastern Brazil. All reproductive trees (T. clausseni, T. pallida and T. catigua) with dbh > or = 5 cm within a 1-ha plot were collected, sexed, mapped and, for individuals of each species, the distances to the nearest neighbour of the same and opposite sex were measured. For the shrub species T. elegans (dbh < 5 cm), all reproductive individuals were sampled randomly in 10 samples of 10 x 10 m. The reproductive phenology was observed at weekly to monthly intervals from May 1988 to January 1990. The species are strictly dioecious, did not present any sex-mixed trees or sex switching during the study, and sex ratio did not differ significantly from 1 : 1. The size distributions and the relative size variation were not significantly different between sexes. There was no significant segregation or clumping between individuals of either sex and no fruit production without pollination. Onset of flowering and flowering peak were synchronous between male and female plants for all species studied. Flower synchrony was related to outcrossing and pollinator attraction rather than climatic factors.     

Population structure and reproduction in the neotropical dioecious tree Compsoneura sprucei

Summary In rain forest study plots, the sexes of Compsoneura sprucei (Myristicaceae) were radomly distributed and similar in vegetative dimensions. The sex ratio among adults was estimated as 1.25 male: female. The population showed two flowering episodes per year, of unequal intensity. Sexual dimorphisms in order of increasing difference included the frequency of flowering, the number of flowers per inflorescence and the number of inflorescences per tree. Most females matured only 0–10 seeds per tree per flowering episode. Tree size was a better indicator of fecundity in males than females. Reproduction in both sexes was dominated by a very few prolific trees.     

Sexual reproductive biology of the endangered Japanese red maple (<Emphasis Type="Italic">Acer pycnanthum</Emphasis>)

Japanese red maple, Acer pycnanthum, is an endangered wetland species having dioecious sex expression. Prior studies conclude that land development has caused severe loss of habitat, and seedling regeneration is rare. However, information is lacking on sexual reproduction, which is the first critical stage of regeneration. My research objective was to describe flowering and fruiting characteristics of Japanese red maple, focusing on tree sizes at the onset of flowering, factors affecting flowering regularity and fruit abundance, periodicity of fruit production, and sex ratio. In a study of a young population, Japanese red maple initiated flowering at a small size under 100% light intensity. Initiation of flowering occurred at relatively lower height in males than females, and males flowered more regularly than females. A survey of 1,106 clones from 46 natural populations revealed that male clones were significantly more abundant than female clones. In a 4-year study of mature populations, nearly all clones flowered regularly, and fruit abundance of dominant females was typically high. Fruit abundance at the population level also remained high, although fruit abundance of individual female clones varied year by year. Therefore, sexual reproductive ability is highly vigorous. Sex ratio exhibited a major deviation from a 1:1 ratio in populations with a small number of clones. In conservation planning, we should prioritize avoidance of an unbalanced sex ratio by increasing population sizes. In addition, populations having many clones >40 cm dbh with large, well-lighted crowns have a high potential of regular and abundant seed production.     

Sex ratio variation and spatial distribution of Siparuna grandiflora, a tropical dioecious shrub

Populations of dioecious plant species often exhibit biased sex ratios. Such biases may arise as a result of sex-based differences in life history traits, or as a result of spatial segregation of the sexes. Of these, sex-based differentiation in life history traits is likely to be the most common cause of bias. In dioecious species, selection can act upon the sexes in a somewhat independent way, leading to differentiation and evolution toward sex-specific ecological optima. I examined sex ratio variation and spatial distribution of the tropical dioecious shrub Siparuna grandiflora to determine whether populations exhibited a biased sex ratio, and if so, whether the bias could be explained in terms of non-random spatial distribution or sex-based differentiation in life history traits. Sex ratio bias was tested using contingency tables, a logistic regression approach was utilized to examine variation in life history traits, and spatial distributions were analyzed using Ripley's K, a second-order neighborhood analysis. I found that although populations of S. grandiflora have a male-biased sex ratio within and among years, there was no evidence of spatial segregation of the sexes. Rather, the sex ratio bias was shown to result primarily from sex-based differentiation in life history traits; males reproduce at a smaller size and more frequently than females. The sexes also differ in the relationship between plant size and reproductive frequency. Light availability was shown to affect reproductive activity in both sexes, though among infrequently flowering plants, females require higher light levels than males to flower. The results of this study demonstrate that ecologically significant sex-based differentiation has evolved in S. grandiflora. Received: 30 July 1997 / Accepted: 16 December 1997     

Costs of reproduction in Nyssa sylvatica: sexual dimorphism in reproductive frequency and nutrient flux

Summary We examined the influence of differential reproductive frequency between the sexes on tertiary (phenotypic) sex ratios in the the dioecious tree Nyssa sylvatica (Nyssaceae). Reproduction was evaluated in relation to sex, size and canopy exposure using flowering data collected from 1229 marked trees over a four year period. For subsets of each population we used data on flower number, fruit crop size, fruit/flower ratios, and individual flower and fruit mass to compare biomass invested in reproductive structures of males and females. We also examined seasonal changes in stem nitrogen and soluble carbohydrate content in relation to flower and fruit production for trees of each sex. Our results indicate that: 1) Male-biased tertiary sex ratios could be explained by more frequent reproduction by male trees; 2) Estimated secondary sex ratios based on sums of all known males and females were not significantly different from 1:1; 3) Flowering frequency of males and females was significantly related to plant size (DBH) and exposure of the canopy to light; 4) Estimtes of reproductive biomass allocation ranged from 1.36 to 10.8 times greater for females relative to males; 5) Flower production was related to stem nutrient status for both sexes, but nutrient depletion and its effect on subsequent flowering was much more pronounced for female trees. We conclude that less frequent flowering by female trees may result from depletion of stored reserves, and that differential flowering frequency in N. sylvatica may ultimately reduce apparent sexual differences in the costs of reproduction.     

Sex-different response in growth traits to resource heterogeneity explains male-biased sex ratio

In dioecious plants, differences in growth traits between sexes in a response to micro-environmental heterogeneity may affect sex ratio bias and spatial distributions. Here, we examined sex ratios, stem growth traits and spatial distribution patterns in the dioecious clonal shrub Aucuba japonica var. borealis, in stands with varying light intensities. We found that male stems were significantly more decumbent (lower height/length ratio) but female stems were upright (higher height/length ratio). Moreover, we found sex-different response in stem density (no. of stems per unit area) along a light intensity gradient; in males the stem density increased with increases in canopy openness, but not in females. The higher sensitivity of males in increasing stem density to light intensity correlated with male-biased sex ratio; fine-scale sex ratio was strongly male-biased as canopy openness increased. There were also differences between sexes in spatial distributions of stems. Spatial segregation of sexes and male patches occupying larger areas than female patches might result from vigorous growth of males under well-lit environments. In summary, females and males showed different growth responses to environmental variation, and this seemed to be one of possible causes for the sex-differential spatial distributions and locally biased sex ratios.     

Size-related flowering and fecundity in the tropical canopy tree species, Shorea acuminata (Dipterocarpaceae) during two consecutive general flowerings

We monitored the reproductive status of all trees with diameters at breast height (dbh) >30 cm in a 40-ha plot at Pasoh, west Malaysia, and investigated the individual fecundity of 15 Shorea acuminata Dyer (Dipterocarpaceae) trees using seed-trapping methods during two consecutive general flowering periods in 2001 (GF2001) and 2002 (GF2002). The proportion of flowering trees was higher, and not dependent on size, in GF2002 (84.2%), than in GF2001 (54.5%), when flowering mainly occurred in trees with a dbh ≤70 cm. Fecundity parameters of individual trees per event varied widely (221,000–35,200,000 flowers, 0–139,000 mature seeds, and 1.04–177 kg total dry matter mass of fruit (TDM) per tree). Monotonic increases with increasing tree size were observed for flower production and TDM amongst trees up to 90 cm in dbh, but not for mature seed production or for any of these parameters amongst larger trees. The pattern of reproductive investment during the two consecutive reproductive events clearly differed between medium-sized and large trees; the former concentrated their reproductive investment in one of the reproductive events whereas the latter allocated their investment more evenly to both reproductive events. Our results suggest size-related differences in the resource allocation pattern for reproduction. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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     

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]。     

SEX-RATIO AND REPRODUCTIVE VARIATION IN THE MISTLETOE PHORADENDRON JUNIPERINUM (VISCACEAE)

Despite the fact that many parasitic and hemiparasitic plant species such as mistletoes are dioecious and occur in both the new and the old world, few data exist on variation in the sex ratio and allocation to reproduction in these taxa. We investigated 1) the sex-ratio of the xylem-tapping mistletoe Phoradendron juniperinum in relation to its age and position within the canopy of its host tree Juniperus osteosperma, and 2) reproductive effort in relation to the gender and age of mistletoe plants. Our surveys showed that P. juniperinum has a male-biased sex ratio. Despite this predominance of male individuals, females lived longer and had a greater reproductive effort than did males. A statistical analysis of the age distribution data indicated that the peak in the frequency of reproductively mature individuals was later in females than in males. These gender-specific distributions may have resulted 1) from sequential hermaphroditism (age-specific sex switching), or 2) because the average age of peak reproduction is later in female individuals. Because sex is genetically determined in a closely related genus of mistletoe and because we have no data to indicate sex switching in this species, we feel that our data support the interpretation that female individuals, on average, show a peak in reproductive vigor at an older age relative to males. While delayed reproduction in females may be favored because reproductive effort and success appear to be age-dependent in females of this species, both sexes can become reproductively mature relatively early in life. Further, because 1) allocation to reproduction as a function of age increases more rapidly for females of this species relative to males, and 2) because there may be a higher resource cost associated with reproduction in females, we hypothesized that female individuals would be more abundant in the best quality locations within the host tree so as to maximize the opportunity to meet those costs. In spite of the association between gender and some host characteristics, there was no indication that female plants were located in sites most favorable to either their carbon or water balance. We discuss reasons why this may be the case.