FLORAL DISPLAY IN PHYLA INCISA: CONSEQUENCES FOR MALE AND FEMALE REPRODUCTIVE SUCCESS

The inflorescences of Phyla incisa consist of flowers in two phases: younger, nectar-containing flowers that have yellow corolla throats and older, nectar-lacking flowers that have dark purple corolla throats. Observations of pollinator visitation patterns to both natural and manipulated inflorescences were made to determine the role of each flower phase in pollinator attraction. The effect of older-phase flowers on male and female reproductive success was determined by comparing stigmatic pollen loads and estimates of pollen removal from inflorescences having different numbers of these flowers. The pollinators of Phyla selected larger inflorescences more often than expected based upon the size distribution of inflorescences available to them. Both younger- and older-phase flowers contributed to the attraction of pollinators, but the latter were less effective in this function. The presence of older-phase flowers significantly increased the visitation rate to inflorescences and the amount of pollen removed but had little effect on pollen deposition on stigmas. The lack of correspondence between pollen deposition and pollinator-visitation rate was not due to stigma saturation, since stigma loads varied greatly. The data indicate that the deposition of pollen on stigmas in this species is a relatively stochastic process, whereas pollen removal from inflorescences occurs at a much more regular rate. Old-phase flower retention appears to contribute to reproductive success through increased pollen donation when pollinator activity is high and may also increase the probability of seed set when pollinators are rare.     

POPULATION STRUCTURE ALONG A STEEP ENVIRONMENTAL GRADIENT: CONSEQUENCES OF FLOWERING TIME AND HABITAT VARIATION IN THE SNOW BUTTERCUP,RANUNCULUS ADONEUS

Few studies have determined how gene flow and selection interact to generate population genetic structure in heterogeneous environments. One way to identify the potential role played by natural selection is to compare patterns of spatial genetic structure between different life cycle stages and among microenvironments. We examined patterns of spatial structure in a population of the snow buttercup (Ranunculus adoneus), using both adult plants and newly emerged seedlings. The study population spans a steep environmental gradient caused by gradual melting of snow within a permanent snowbed. Early-melting sites are characterized by denser vegetation, more fertile soils, and a longer growing season than late-melting sites tens of meters away. The flowering time of R. adoneus is controlled entirely by time of snowmelt, so the contiguous population is phenologically substructured into a series of successively flowering cohorts, reducing the opportunity for direct pollen transfer between early- and late-melting sites. For four highly polymorphic enzyme loci in this tetraploid species, there was subtle, but statistically significant, genetic differentiation between early, middle, and late-melting cohorts; adults usually showed greater differentiation among snowmelt zones than did seedlings. At two loci in adults and one locus in seedlings, homozygotes were more common than predicted at Hardy-Weinberg equilibrium, even when assuming maximum levels of double reduction during meiosis. This pattern suggests the occurrence of self-fertilization and/or population substructure. To determine how spatial isolation and phenological separation each contribute to genetic substructure, we used bivariate regression models to predict the numbers of allele differences between randomly paired individuals as a function of meters separation in space and days separation in flowering time. For newly emerged seedlings, we found that spatial separation was positively associated with genetic difference, but that the additional contribution of phenological separation to genetic difference was not significant. This implies that seeds and/or pollen move effectively across the snowmelt gradient, despite differences in flowering time. As was true for seedlings, spatial separation between paired adults contributed to greater genetic difference, but for a given spatial separation, the genetic difference between adult plants was reduced by phenological separation. This result implies that postemergence selection is favoring at least some seeds that migrate across the snowmelt gradient. Directional gene flow across the snowmelt gradient probably results from a genetic source-sink interaction, that is, the colonization of ecologically marginal late-melting sites by high quality seeds produced by the larger subpopulation in early-melting sites. Effective gene flow from high to low quality microenvironments is likely to impede adaptation to late-melting locations.     

REPRODUCTIVE SUCCESS OF FUCUS VESICULOSUS (PHAEOPHYCEAE) IN THE BALTIC SEA

Marine organisms colonizing brackish habitats such as the Baltic Sea must cope with the negative effects of low salinities on reproductive success because these may reduce gamete viability and/or increase polyspermy. Reproductive characteristics of the marine seaweed Fucus vesiculosus L. were studied in several brackish habitats, particularly in the northern Baltic Sea, to understand its ability to reproduce where few other marine species survive. Polyspermy and fertilization success were variable at the boundary of the continuous distribution of F. vesiculosus in the Baltic Sea, and polyspermy was high (10%–30%) when fertilization was successful. A strong female bias (80%–86%, ca. 5.5:1) was found at the northernmost limit of Baltic F. vesiculosus. Electrophysiological studies showed that many eggs have a high input resistance (519 ± 150 MΩ[mean ± SE, n = 14] at Drivan, 1995), which may be helpful in preventing polyspermy in this brackish habitat. The polyspermy block remains sodium-dependent in the northern Baltic. Sperm bound quickly to northern Baltic eggs in natural water, but fertilization was delayed compared to marine F. vesiculosus. A subset of northern Baltic eggs studied during an optimal reproductive period (7–11 July 1995) had a membrane potential (E_m) of ca. −100 mV and an effective fertilization potential (FP) of ca. 2 min with a plateau of −25 mV, but repolarized too rapidly for the FP to be protective. Pronuclear migration and cell wall secretion occurred more slowly in Baltic than in marine zygotes. The reproductive success of theseboundary populations may be dependent upon windows of opportunity when there are favorable combinations of the levels of salinity, water motion, population density, and sex ratio. These factors and the short duration of the reproductive season in the northern Baltic Sea may result in reproductive failure in some years.     

PHENOLOGY AND REPRODUCTIVE ALLOCATION IN ANDROPOGON SCOPARIUS (GRAMINEAE) POPULATIONS IN COMMUNITIES OF DIFFERENT SUCCESSIONAL STAGES

Successional populations of Andropogon scoparius in Franklin Township, Somerset County, New Jersey, were compared as to their phenology and reproductive allocation. Field data from six populations in old-fields ranging in age from 2 to 40 years indicated significant differences in both phenology and reproductive effort (ratio of reproductive to total shoot biomass). Mean date of first anthesis was consistently later with increasing age of the old-field, and the later the first anthesis date for a population, the lower was the reproductive effort. Population data on mean maximum height and seed weight showed no relation to successional age. Greenhouse experiments on three of the populations revealed that most of the field differences were attributable to local habitat effects on phenotypic expression. The possibility of some genetic differentiation paralleling successional age was raised by the consistent (though sometimes statistically not significant) differences between the early (3-yr) and late (40-yr) successional populations in date of first anthesis and in reproductive effort under two light intensities and uniform greenhouse conditions.     

VARIATION IN INDIVIDUAL FLOWERING TIME AND REPRODUCTIVE SUCCESS OF AGALINIS STRICTIFOLIA (SCROPHULARIACEAE)

Field studies on two populations of Agalinis strictifolia were conducted over a 3-year period to investigate the relationship between flowering time of individuals and plant size, flowering duration, flower and fruit production, fruit predation, and growth rate. Seasonal patterns of pollinator visitation were compared with those of individual flowering time, flower density, percent fruit production, and mean seeds/fruit. In general, early and middle flowering individuals (as determined by either first flowering date or peak flowering) were larger, flowered longer, and produced more flowers and fruits than late flowering individuals. Early and middle flowering individuals (based on first flowering date) also grew faster than late flowering individuals. Although early and middle flowering individuals produced more fruits, fruit predators did not damage a disproportionate number of fruits compared to late flowering individuals. Patterns of bee visitation showed no association with seasonal patterns of flower density, percent fruit production, mean seed/fruit, or individual flowering time. In populations of A. strictifolia, it would seem that biotic or environmental determinants of growth rate (hence size and reproductive success) may be more important in generating variation in individual flowering time than patterns of pollinator visitation or fruit predation.     

REPRODUCTIVE SUCCESS AND INFLORESCENCE SIZE OF CALOPOGON TUBEROSUS (ORCHIDACEAE)

Reproductive success of Calopogon tuberosus, which produces no nectar, was investigated in relation to inflorescence size and dispersion pattern. Mean inflorescence size was 2.56 (range 1–10). A bagging experiment showed that insects are required for pollen transfer and that fruits are produced from self-, geitonogamous, and cross-pollinations; fruit set was not 100%. Fruit set of nonmanipulated plants was limited by the number of pollinator visits. Reproductive success increased with increasing inflorescence size, although not above theoretical predictions. However, the probability of producing no fruit or contributing no pollinia decreased with increasing inflorescence size since sequential flowering increased the probability of a pollinator visit to the inflorescence over the blooming period. Large inflorescences did not provide a greater pollinator attraction than small ones, because inflorescences only presented a few open flowers at a time. In addition, flowers on plants growing in clumps of 2–8 plants had a higher probability of setting fruit, apparently because of increased pollinator attraction. Although there are obvious selective advantages for large inflorescences, the sequential flowering habit, and low resource availability may reduce the advantages of large inflorescence size at our study site.     

LIFETIME REPRODUCTIVE SUCCESS AND THE OPPORTUNITY FOR SELECTION IN A NONTERRITORIAL DAMSELFLY (ODONATA: COENAGRIONIDAE)

Major components of male and female lifetime reproductive success (LRS) were quantified for a damselfly that exhibits “scramble competition” for mates. The opportunity for selection on male reproduction was potentially 2.9 times that for females. Differential fertility/clutch and survivorship each accounted for about half of the total variation in female reproductive success. Variation in fertilization efficiency accounted for 7% of the total opportunity for selection on males. Although differences in survivorship and mating efficiency each contributed to about a third of the total opportunity for selection on male reproduction, both components appeared to be influenced by random factors. Survivorship was age-independent, and the mating distributions among males with equal mating opportunities were indistinguishable from those expected if matings were random with respect to male phenotype. Because the proportion of the standarized variance (I) in LRS that was attributed to sexual selection depended on the way the selective episodes were defined, the sample of individuals included in the partitioning analysis, and the degree of sexual selection on mated males that could be detected, my results caution against drawing conclusions about the dynamics of sexual selection on populations based on a superficial comparison of I values.     

VARIATION IN MALE REPRODUCTIVE INVESTMENT AND MALE REPRODUCTIVE SUCCESS IN WHITE SPRUCE

An experimental population composed of ramets of white spruce clones was studied to determine the influence of different levels of clonal male reproductive investment on clonal male reproductive success, where the male reproductive success of different clones in the population was defined as the proportion of the seed crop sired by each clone. We present a multilocus estimation procedure which provides unbiased estimates of the proportion of seed sired by each parental genotypic class from pollen gamete frequency data, whenever gametic segregation frequencies are known. Significant linear and quadratic components were found for the relationship between the number of male cones produced and the estimated proportion of the seed crop sired. A clone that produced an exceptionally large number of male cones, however, did not have the greatest male reproductive success. In general, the results support an assumption made in theoretical models of selection on mating system modifiers regarding the relationship between male investment and male reproductive success. They also support the hypothesis that male reproductive success in wind-pollinated species continues to increase substantially with increasing male reproductive investment.     

AUTOGAMY AND COMPETITION FOR POLLINATORS IN HEPATICA AMERICANA (RANUNCULACEAE)

Hepatica americana is one of the earliest flowering vernal herbs in the deciduous forests of piedmont North Carolina. Unlike most other members of the spring wildflower community, its flowers are nectarless and autogamous. The number of insect visits a flower receives varies markedly through the blooming season and is lowest when H. americana is in full bloom. Potential for outcrossing as measured by the seed set of emasculated flowers follows the same pattern. The mid-season decline in insect-pollination is most likely the result of competition with Erythronium umbilicatum, which flowers concurrently with H. americana and is much more attractive to the solitary bees that are the principal early season pollen vectors. Because of this competitive disadvantage, seed set of H. americana in piedmont North Carolina would be pollination-limited if the flowers were not autogamous. Autogamy may also be favored by flowering early in the spring before pollinators are available. Opportunities for outcrossing in H. americana are primarily the result of protogyny and blooming earlier than competitors.     

THE MORPHOLOGY OF THE EXINE IN NIGELLA (RANUNCULACEAE)

The pollen morphology of eight species of Nigella (Ranunculaceae) was examined by scanning and transmission electron microscopy. The exomorphology of all species was identical: 3-colpate, spinulose, and punctate, but thin sections revealed two structural patterns. The ektexine structure of Nigella integrifolia, consisting of thickened foot layer, columellae, and thin tectum, is typical for the family as well as the order Ranunculales in general. In contrast, the remaining seven species, N. arvensis, N. damascena, N. elata, N. hispanica, N. sativa, N. segetalis, and N. stellaris, have an ektexine with an additional unit, a horizontal layer with shorter columellae, placed between the foot layer and tectum. Of all genera examined in the Ranunculaceae, only Nigella had this unusual stratification. This difference in the exine structure would add support to the treatment of N. integrifolia as a monotypic genus, Komaroffia integrifolia (Regel) Lemos Pereira.     

CHROMOSOMES OF OREITHALES (RANUNCULACEAE)

A chromosome count of 2n =48 is reported for Oreithales integrifolia from Peru. This represents the first chromosome report for this genus. This chromosome report is discussed in relation to other genera in the Ranunculaceae.     

长白山花尾榛鸡繁殖成功率的研究

１９８６～１９９５年,对长白山花尾榛鸡的繁殖率进行了研究。无线电遥测的结果表明,花尾榛鸡的孵化率为６５５％（ｎ＝５）,繁殖成功率为２０９％（ｎ＝４）,幼鸟成活率３７５％（ｎ＝３）。花尾榛鸡家族群的幼鸟数随日龄增加而减少,幼鸟日龄为０～２５日时死亡率较高,２５日龄后的死亡率显著减少。１０年花尾榛鸡的繁殖成功率指数的变化与６月的降水量及＞１ｍｍ的降水天数呈显著的负相关（Ｐ＜００５,Ｐ＜００２５）,说明育雏初期降水对幼鸟成活率的影响较大。     

BREEDING SYSTEM EVOLUTION IN LEAVENWORTHIA: BREEDING SYSTEM VARIATION AND REPRODUCTIVE SUCCESS IN NATURAL POPULATIONS OF LEAVENWORTHIA CRASSA (CRUCIFERAE)

Populations of Leavenworthia crassa (Cruciferae) studied for 3 years exhibited among- and within-population genetic variation for a suite of floral and reproductive traits (flower width, petal length, anther position, ability to set seeds in the absence of pollinators, time to first flowering) associated with breeding system. We used electrophoretic markers to show that a population with small, monomorphically colored flowers with introrse anthers had a significantly lower outcrossing rate (t = 0.03) than did a population with larger, polymorphically colored flowers with extrorse anthers (t = 0.33). In the more-outcrossing population the correlation between higher maternal plant outcrossing rate and the suite of six traits approached significance (P < 0.067), with greater petal size, greater flower width, and reduced ability to set seeds in the absence of pollinators contributing significantly. Plants in selfing populations had a generally higher reproductive success, with a higher number of flowers per plant, a smaller proportion of unfertilized ovules, a smaller proportion of fertilized ovules aborted, a higher rate of fruit set, and overall a larger number of seeds matured than did plants from the more outcrossing populations. Pollen limitation did not appear to account for lower reproductive success in outcrossing populations. Resource limitation did not differ substantially between populations. However, within-ovary patterns of fertilization, abortion, and seed weight were significantly less random in outcrossing populations than in selfing populations, suggesting that differential gamete and embryo success may be responsible for lower reproductive success in outcrossing populations.     

FACTORS INFLUENCING THE REPRODUCTIVE SUCCESS OF IPOMOEA PES-CAPRAE (CONVOLVULACEAE) AROUND THE GULF OF MEXICO

The reproductive biology of Ipomoea pes-caprae, a pantropical beach morning glory, was studied at five sites around the Gulf of Mexico. The primary pollinators were Xylocopa species (carpenter bees) which dispersed pollen up to 90 m. Exclusion experiments demonstrated that ants feeding on extrafloral nectaries increased seed set but did not protect seeds from predation by the bruchid beetle Megacerus. The water-dispersed seed and long-range dispersal of pollen may function to counter the sporophytic incompatability mechanism of the plant; populations exhibit a large neighborhood size. Key reproductive factors in the life cycle of I. pes-caprae are the long-range pollen flow and mass germination of water-dispersed seeds.     

THE EFFECT OF FLOWER PRODUCTION ON MALE REPRODUCTIVE SUCCESS IN WILD RADISH POPULATIONS

Flower production is the major determinant of pollen yield and an important component in pollinator attraction. Consequently differences among plants in flower production are expected to have a substantial impact on their relative success at fathering seed. We examined this prediction using one natural and three structured populations of wild radish. We found that a plant's relative success at fathering seed on another plant in the population (male fertility) increased with flower production. Nonetheless, the increase in fertility exhibited a diminishing marginal gain, with the relationship varying among populations. The relationship between the estimates of total number of seeds sired and flower production varied substantially among the populations examined, ranging from a weakly linear to strongly negative quadratic. Not surprisingly, the spatial structure of the population with respect to seed yield had a powerful effect on the total number of seeds sired because male fertility decreased exponentially with intermate distance. This exponential relationship occurred in all populations examined. Other covariates important to male fertility were flower color, time, the specific identity of the male parent, and male by female interaction. The identity of the male parent consistently accounted for a large portion of the variation in male fertility, indicating that other unmeasured features of the plant influenced its success.     

濒危植物南川升麻生殖特性的研究

研究了南川升麻的生殖生物学特性,并探讨了其致濒的原因。南川升麻只分布在中国四川省金佛山地区,仅１个居群,个体总数６０００￣８０００株,其中开花植株约占１９％左右;花序圆锥状,花期８月中至１０月初,１１月中下旬果实成熟。南川升麻雌雄异熟,表现为大约花谢７ｄ后柱头才发育成熟;自然条件下可正常受精,但频率低,结籽率平均仅为５．８％￣７．０％,可授期内人工异株授粉,结籽率可提高到２８．０％;胚乳核型,种子