The structure of phenotypic variation in gender and floral traits within and among populations of Spergularia marina (Caryophyllaceae)

We sampled four wild populations of the highly autogamous Spergularia marina (Caryophyllaceae) in California to detect and to measure the magnitude of within- and among-population sources of phenotypic variation in gender and floral traits. From flowers and fruits collected from field and greenhouse-raised plants, we measured ovule number, seed number, mean seed mass, pollen production (greenhouse families only), mean pollen grain volume (greenhouse families only), anther number, anther/ovule ratio, pollen/ovule ratio (estimated using different flowers for pollen than for ovules; greenhouse families only), petal number, and petal size. Using greenhouse-raised genotypes, variation among maternal families nested within populations was evaluated for each trait to determine whether populations differ in the degree of maternally transmitted phenotypic variation. For each population, we used 15 greenhouse-raised maternal families to estimate the broad-sense heritability and genetic coefficient of variation of each floral trait. The magnitude and statistical significance of broad-sense heritability estimates were trait- and population-specific. Each population was characterized by a different combination of floral traits that expressed significant maternally transmitted (presumably genetic) variation under greenhouse conditions. Flowers representing two populations expressed low levels of maternally transmitted variation (three or fewer of nine measured traits exhibited significant maternal family effects on phenotype), while flowers representing the other two populations exhibited significant maternal family effects on phenotype for five or more traits. Our ability to detect statistically significant differences among the four populations depended upon the conditions under which plants were grown (field vs. greenhouse) and on the floral trait observed. Field-collected flowers exhibited significant differences among population means for all traits except anther number. Flowers sampled from greenhouse-raised maternal families differed among populations for all traits except ovule number, seed number, and petal size. We detected negligible evidence that genetic correlations constrain selection on floral traits in Spergularia marina.     

毛茛状金莲花不同花期的花特征和访花昆虫的变化及表型选择

为了研究植物生长季内开花时间对花特征表型选择的影响,我们以青藏高原东缘高寒草地的毛茛状金莲花Trollius ranunculoides)为实验材料,在生长季内不同开花时间(花前期、花末期)测定花特征,观察访花昆虫的类群和访花频率,生长季结束后收集种子.根据昆虫访花的喜好和季节内类群与访花频率的变化,分析了不同开花时间毛茛状金莲花的花特征与昆虫的选择;并用种子产量表示雌性适合度,估计了毛茛状金莲花的花特征在不同开花时间所受的表型选择.结果表明:不同花期植物的花特征有显著差异,相应的访花昆虫的类群和频率也存在差异,不同类群昆虫访花喜好也不一样.蜂喜好花瓣和花萼较宽、花茎短和花茎数少的个体,这正符合花前期的特征,因而蜂的访花频率在花前期较高;蝇对花特征没有明显的偏好.而通过雌性适合度估计毛茛状金莲花花特征所受的表型选择则是:花前期,花茎较长和花茎数多的植株适合度大;花末期,花茎数多的植株适合度大.我们的研究表明:在植物生长季,花期的分化伴随着传粉昆虫活动的变化.不同花期,访花昆虫的变化可能对植物花特征的分化起了至关重要的作用.但是访花昆虫对花特征的选择与通过雌性适合度估计植物受到的选择不尽相同,这可能是由于其他因素造成的.     

Evolutionary analysis of a key floral trait in aquilegia canadensis (ranunculaceae): genetic variation in herkogamy and its effect on the mating system

The mating system of flowering plant populations evolves through selection on genetically based phenotypic variation in floral traits. The physical separation of anthers and stigmas within flowers (herkogamy) is expected to be an important target of selection to limit self-fertilization. We investigated the pattern of phenotypic and genetic variation in herkogamy and its effect of self-fertilization in a broad sample of natural populations of Aquilegia canadensis, a species that is highly selfing despite strong inbreeding depression. Within natural populations, plants exhibit substantial phenotypic variation in herkogamy caused primarily by variation in pistil length rather than stamen length. Compared to other floral traits, herkogamy is much more variable and a greater proportion of variation is distributed among rather than within individuals. We tested for a genetic component of this marked phenotypic variation by growing naturally pollinated seed families from five populations in a common greenhouse environment. For three populations, we detected a significant variation in herkogamy among families, and a positive regression between parental herkogamy measured in the field and progeny herkogamy in the greenhouse, suggesting that there is often genetic variation in herkogamy within natural populations. We estimated levels of self-fertilization for groups of flowers that differed in herkogamy and show that, as expected, herkogamy was associated with reduced selfing in 13 of 19 populations. In six of these populations, we performed floral emasculations to show that this decrease in selfing is due to decreased autogamy (within-flower selfing), the mode of selfing that herkogamy should most directly influence. Taken together, these results suggest that increased herkogamy should be selected to reduce the production of low-quality selfed seed. The combination of high selfing and substantial genetic variation for herkogamy in A. canadensis is enigmatic, and reconciling this observation will require a more integrated analysis of how herkogamy influences not only self-fertilization, but also patterns of outcross pollen import and export.     

Plasticity of inflorescence traits in Lobelia siphilitica (Lobeliaceae) in response to soil water availability

Many workers have demonstrated a genetic basis for variation in inflorescence traits, but this variation can also have an environmental component. Because flowering can incur significant water costs, I estimated plasticity of inflorescence traits of three populations of Lobelia siphilitica in response to drought. I manipulated soil water availability in the greenhouse and measured seven inflorescence traits. Under drought conditions, plants from one population flowered later and produced fewer flowers with shorter corollas and narrower landing pads. In contrast, the height of the flowering stalk decreased in response to drought in all three populations. Consequently, pollinator-mediated natural selection on these plastic traits may depend on soil water availability. Plastic responses differed between genotypes only for the height of the flowering stalk and the length of the corolla tube and only in one or two populations. This suggests that genotype × environment interactions would not limit the evolution of inflorescence traits in L. siphilitica. The strength and sign of phenotypic correlations among inflorescence traits did not respond plastically to drought, suggesting that indirect selection on inflorescence traits of L. siphilitica will not vary strongly with water availability. My results suggest that plasticity of inflorescence traits may influence their evolution, but the effects are population- and trait-specific.     

Floral Display in Narcissus: Variation in Flower Size and Number at the Species, Population, and Individual Levels

Floral display (the size, number, and arrangement of open flowers) influences pollinator visitation to animal-pollinated plants and should be an important determinant of reproductive success. We examined variation in the size and number of open flowers in wild daffodils (Narcissus). Our analysis of published data on 45 taxa showed that flower number varied negatively with flower diameter among Narcissus species, which supports the widespread assumption that there is a trade-off between these traits. In contrast, field measurements indicated a positive relation between flower number and diameter within two populations of Narcissus dubius, and no relation was evident after we controlled for variation in bulb size. The discrepancy between inter- and intraspecific patterns may have occurred because variable resource levels obscure trade-offs when variation in flower size is low (e.g., within species). Size-related increases in floral tube length were half as great as corresponding increases in flower diameter, a result that is consistent with stronger stabilizing selection on tube length. Staggered flowering within N. dubius inflorescences limited the mean number of open flowers to <66% of total flower number, and slow expansion by later opening flowers resulted in significant differences in flower size throughout flowering. Although pollinators preferred large flowers, experimental reductions in flower diameter did not affect seed production. Our results illustrate how the relative importance of the factors influencing floral display can vary among levels of biological organization. Interspecific variation in flower size and number appeared to be constrained by allocation trade-offs, but intraspecific variation in both traits was more greatly influenced by plant resource status. Within plants, the size and number of open flowers reflected the relative age of individual flowers and floral longevity.     

福建山樱花天然居群表型变异研究

以闽、赣、粤三省天然分布的福建山樱花(Cerasus campanulata Maxim.)为研究对象,调查了11个居群226个单株的17个表型性状指标,运用变异系数和单因子方差分析研究了居群间和居群内的表型变异;应用相关分析揭示了表型性状间及其与地理、气候因子间的相关性以及表型变异的地理格局;并且运用聚类分析进行了居群分类.结果表明,福建山樱花天然居群内的变异远大于居群间的变异,居群分化较小;花色、花量与经度、纬度间显著或极显著相关,花瓣宽与纬度、海拔显著或极显著相关,花色、花量与年降雨量极显著相关,花量、花瓣宽、开花习性与年日照时数显著相关,花瓣宽与年均温、无霜期显著或极显著相关,存在较明显的地理变异趋势.而居群间马氏距离聚类分析结果与参试居群分布地域性呈现高度一致,二者之间相关显著.     

濒危植物夏蜡梅花的形态变异

研究了不同地理位置的3个夏蜡梅自然种群间以及种群内花部形态变异,并对3个种群进行聚类分析。结果表明：夏蜡梅花部表型性状在种群间和种群内都存在一定程度的变异;各性状总的平均变异系数在6.293（内瓣长）~46.90（花柄长）之间,种群间总的平均变异系数在13.430（DMS）~15.720 8（DLS）,因此认为夏蜡梅花的表型性状无论是在物种水平还是在种群水平都具有较丰富的表型多样性,其中又以大雷山种群最高;单因素方差分析显示：部分指标分化显著或极显著,13个指标在种群内的变异占了总变异的82.9%,这说明夏蜡梅种群内变异是表型变异的主要来源,由此可以推测遗传上的多样性是形成夏蜡梅花部形态变异的主导因素;种群间平均变异组分占了17.1%,表明花部形态在种群间存在一定程度的表型分化;聚类分析显示距离较近的大明山种群和龙须山种群先聚在一起,表型差异与地理距离之间存在显著正相关,经分析认为微生境的异质性和种群隔离以及遗传漂变均可能导致夏蜡梅种群间的表型分化;相关性分析显示部分性状之间存在显著的相关性。     

Within‐plant variation in reproductive investment: consequences for selection on flowering time

Variation among the leaves, flowers or fruit produced by a plant is often regarded as a nuisance to the experimenter and an impediment to selection. Here, we suggest that within‐plant variation can drive selection on other plant‐level traits. We examine within‐plant variation in floral sex allocation and in fruit set and predict that such variation generates variation in male success among plants, thereby driving selection on flowering time. We tested this prediction in a simulation model estimating selection on flowering time through male fitness when floral sex allocation and/or fruit set vary directionally among flowers on plants. We parameterized the model through a quantitative literature survey of within‐plant change in sex allocation. As predicted, within‐plant variation in floral sex allocation and in fruit set probability can generate selection on flowering time through male fitness. Declining fruit set from first to last flowers on plants, as occurs in many species, selected for early flowering onset through male fitness. This result was robust to self‐incompatibility and to varying returns on male versus female investment. Selection caused by declining fruit set was strong enough to reverse the selection for late flowering that can be caused by intrafloral protandry. Our model provides testable predictions regarding selection on flowering time through male fitness. The model also establishes the intriguing possibility that within‐plant variation may influence selection on other traits, regardless of whether that variation is under selection itself.     

Divergent selection on flowering phenology but not on floral morphology between two closely related orchids

Closely related species often differ in traits that influence reproductive success, suggesting that divergent selection on such traits contribute to the maintenance of species boundaries. Gymnadenia conopsea ss. and Gymnadenia densiflora are two closely related, perennial orchid species that differ in (a) floral traits important for pollination, including flowering phenology, floral display, and spur length, and (b) dominant pollinators. If plant–pollinator interactions contribute to the maintenance of trait differences between these two taxa, we expect current divergent selection on flowering phenology and floral morphology between the two species. We quantified phenotypic selection via female fitness in one year on flowering start, three floral display traits (plant height, number of flowers, and corolla size) and spur length, in six populations of G. conopsea s.s. and in four populations of G. densiflora. There was indication of divergent selection on flowering start in the expected direction, with selection for earlier flowering in two populations of the early‐flowering G. conopsea s.s. and for later flowering in one population of the late‐flowering G. densiflora. No divergent selection on floral morphology was detected, and there was no significant stabilizing selection on any trait in the two species. The results suggest ongoing adaptive differentiation of flowering phenology, strengthening this premating reproductive barrier between the two species. Synthesis: This study is among the first to test whether divergent selection on floral traits contribute to the maintenance of species differences between closely related plants. Phenological isolation confers a substantial potential for reproductive isolation, and divergent selection on flowering time can thus greatly influence reproductive isolation and adaptive differentiation.     

POLLINATOR-MEDIATED SELECTION ON FLORAL DISPLAY AND FLOWERING TIME IN THE PERENNIAL HERB ARABIDOPSIS LYRATA

The evolution of floral display and flowering time in animal-pollinated plants is commonly attributed to pollinator-mediated selection. Yet, the causes of selection on flowering phenology and traits contributing to floral display have rarely been tested experimentally in natural populations. We quantified phenotypic selection on morphological and phenological characters in the perennial, outcrossing herb Arabidopsis lyrata in two years using female reproductive success as a proxy of fitness. To determine whether selection on floral display and flowering phenology can be attributed to interactions with pollinators, selection was quantified both for open-pollinated controls and for plants receiving supplemental hand-pollination. We documented directional selection for many flowers, large petals, late start of flowering, and early end of flowering. Seed output was pollen-limited in both years and supplemental hand-pollination reduced the magnitude of selection on number of flowers, and reversed the direction of selection on end of flowering. The results demonstrate that interactions with pollinators may affect the strength of selection on floral display and the direction of selection on phenology of flowering in natural plant populations. They thus support the contention that pollinators can drive the evolution of both floral display and flowering time.     

Selection for floral integration and trait variation in zygomorphic flowers of Aconitum japonicum ssp. subcuneatum (Ranunculaceae)

Pollinator-mediated selection might lead to among-trait differences in the degree and pattern of floral integration and intra-flower variation. To examine the patterns of intra-flower variation in floral traits, including nectar volume, we performed a field study using the zygomorphic flowers of Aconitum japonicum ssp. subcuneatum. We investigated (1) correlations between the sizes of the left and right sepals and petals, (2) variation in floral traits among plants, within plants and within flowers, (3) effects of sexual phases on floral integration variation in floral and nectar traits, and (4) the effect of size and intra-flower variation in traits of the left and right sepals and petals on pollen removal by pollinators. Lateral sepal area, but not lower sepal area, was highly correlated between the left and right sepals. Floral traits were more integrated during the male phase than during the female phase. Nectar standing crop in male-phase flowers correlated with helmet height and lateral and lower sepal area, but in female-phase flowers it only correlated with spur length. While intra-flower variance in lateral sepal area accounted for approximately 10% of the overall variance in these traits, the variance in lower sepal area accounted for 70% of the overall variance. Lateral sepal area had a negative effect on the number of pollen grains remaining after pollinator visits. Low variance in lateral sepals within flowers and measurements of pollen removal suggest that lateral sepals play a more important role in pollen export than the other traits. Left and right sepals may be the targets of selection for symmetry in zygomorphic flowers.

     

Floral symmetry is associated with flower size and pollen production but not insect visitation rates in Geranium robertianum (Geraniaceae)

The degree to which fine‐scaled variation in floral symmetry is associated with variation in plant fitness remains unresolved, as does the question of whether floral symmetry is in itself a target of pollinator‐mediated selection. Geranium robertianum (Geraniaceae) is a broadly distributed species whose five‐petaled flowers vary widely with respect to their degree of rotational asymmetry. In this study, we used a naturally occurring population of plants to investigate whether floral rotational asymmetry and leaf bilateral symmetry were phenotypically correlated with a series of fitness‐related traits, and also used an experimental array with model flowers to investigate the preference of insect visitors for varying degrees of floral size and symmetry. We found that leaf asymmetry was not associated with any of the phenotypic traits measured, and that the degree of floral rotational asymmetry was strongly associated with decreased flower size and decreased pollen production. Our experimental arrays showed that insect visitors did not discriminate among model flowers on the basis of size or symmetry alone; however, insect visitors preferentially visited smaller, symmetric model flowers over larger, severely asymmetric model flowers. Taken together, our results suggest that floral and leaf symmetry in G. robertianum are not likely strong indicators of phenotypic quality, and that floral symmetry is unlikely to be a target of pollinator‐mediated selection. However, the relationship between floral asymmetry and pollen production may provide a role for fecundity selection on symmetry in this species. These data importantly add to the growing literature on the adaptive nature of floral symmetry in the wild.     

浙江楠种群表型变异

以探讨浙江楠(Phoebe chekiangensis)种群间和种群内的表型变异程度和变异规律为目的, 对其分布区9个种群的10个表型性状进行了研究。采用巢式方差分析、变异系数、多重比较和相关分析等多种分析方法, 对种群间和种群内的表型多样性及其与地理生态因子的相关性进行了讨论。研究结果表明: 浙江楠叶片、种子等表型性状在种群间和种群内均存在着丰富的变异, 10个性状在种群间、种群内的差异均达极显著水平; 种群内的变异(21.74%)大于种群间的变异(18.45%), 表型性状的平均分化系数为41.43%, 种群内变异是浙江楠表型变异的主要变异来源。各性状平均变异系数为12.78%, 变化幅度为6.50%-19.38%。种群间叶片的平均变异系数(16.99%)高于种群间种子的平均变异系数(8.58%), 表明种子的稳定性高于叶片。叶宽、叶面积与叶片其他性状间多呈显著或极显著相关, 种子千粒重与种子宽、种子体积显著正相关; 而种子和叶片的各性状间多无显著相关性。种子宽、种子体积和种子千粒重随着海拔的增加出现减小的趋势, 种子体积受纬度控制, 随着纬度的增大表现出增大的趋势。     

浙江省野生蜡梅花部形态变异及其与环境因子的相关性

以浙江省蜡梅自然分布区的5个居群为研究对象,采用巢式方差分析、主成分分析、相关分析、聚类分析等多种分析方法,探究蜡梅花部的形态变异及其与环境因子相关性以及居群间和居群内的表型多样性。结果表明:蜡梅花部性状中除雄蕊长度、雌蕊数、雌蕊长度、内被片数以外,其他12个性状在居群间均有极显著差异。胚珠数的平均变异系数最大,花冠内径的平均变异系数最小,各性状的平均变异系数为15.03%(10.81%—23.38%)。五尖山的表型多样性最丰富,碧东山居群最小,种群间平均表型分化系数为44.38%(1.57%—89.62%),种群内变异大于种群间变异,种群内变异是花部变异的主要来源。主成分分析显示花冠直径、花冠内径、花筒深度和中被片长对种群变异起主要贡献作用。花部多数性状间存在显著或极显著的相关关系,生态因子中花部性状与土壤酸度相关性最大,土壤中花部性状与大量元素相关性最大,微量元素次之,中量元素最小。通过UPGMA聚类分析可以将5个种群分成两组。浙江省野生蜡梅花部形态存在丰富的变异和多样性,花部部分性状与土壤中钙、铁、铜含量和海拔有显著或极显著相关关系。     

Covariation among floral traits in Spergularia marina (Caryophyllaceae): geographic and temporal variation in phenotypic and among-family correlations

Strong covariation among traits suggests the presence of constraints on their independent evolution due to pleiotropy, to linkage, or to selective forces that maintain particular trait combinations. We examined floral trait covariation among individuals, among maternal families within and across populations, and over time, in greenhouse-raised plants of the autogamous Spergularia marina. We had three aims. First, since the phenotype of traits expressed by modular organs often changes as individuals age, estimates of the degree of genetic covariation between such traits may also change over time. To seek evidence for this, we measured weekly (for five weeks) an array of floral traits among plants representing ～ 10 maternal families from each of four populations. The statistical significance of the phenotypic and among-family correlations among traits changed over time. Second, we compared populations with respect to trait covariation to determine whether populations or traits appear to be evolving independently of one another. Differences observed among populations suggest that they have diverged genetically. Third, we sought correlations that might reflect constraints on the independent evolution of floral traits. Investment in another and ovule production per flower vary independently among maternal families; there was no evidence for a “trade-off” between male and female investment. We propose that in autogamous taxa one should not find a negative correlation between pollen and ovule production per flower, as such taxa cannot evolve sexual specialization and should be under strong selection to maintain an efficient pollen:ovule ratio, preventing the evolution of male-biased or female-biased genotypes. We found that other pairs of floral traits, however, expressed highly signficant correlation coefficients, suggesting the presence of some evolutionary constraints, at least within some populations, although their strength depended on exactly when flowers were sampled.     

Phenotypic selection on flowering phenology and pollination efficiency traits between Primula populations with different pollinator assemblages

Floral traits have largely been attributed to phenotypic selection in plant–pollinator interactions. However, the strength of this link has rarely been ascertained with real pollinators. We conducted pollinator observations and estimated selection through female fitness on flowering phenology and floral traits between two Primula secundiflora populations. We quantified pollinator‐mediated selection by subtracting estimates of selection gradients of plants receiving supplemental hand pollination from those of plants receiving open pollination. There was net directional selection for an earlier flowering start date at populations where the dominant pollinators were syrphid flies, and flowering phenology was also subjected to stabilized quadratic selection. However, a later flowering start date was significantly selected at populations where the dominant pollinators were legitimate (normal pollination through the corolla tube entrance) and illegitimate bumblebees (abnormal pollination through nectar robbing hole which located at the corolla tube), and flowering phenology was subjected to disruptive quadratic selection. Wider corolla tube entrance diameter was selected at both populations. Furthermore, the strength of net directional selection on flowering start date and corolla tube entrance diameter was stronger at the population where the dominant pollinators were syrphid flies. Pollinator‐mediated selection explained most of the between‐population variations in the net directional selection on flowering phenology and corolla tube entrance diameter. Our results suggested the important influence of pollinator‐mediated selection on floral evolution. Variations in pollinator assemblages not only resulted in variation in the direction of selection but also the strength of selection on floral traits.     

Similarity in G matrix structure among natural populations of Arabidopsis lyrata

Understanding the stability of the G matrix in natural populations is fundamental for predicting evolutionary trajectories; yet, the extent of its spatial variation and how this impacts responses to selection remain open questions. With a nested paternal half‐sib crossing design and plants grown in a field experiment, we examined differences in the genetic architecture of flowering time, floral display, and plant size among four Scandinavian populations of Arabidopsis lyrata. Using a multivariate Bayesian framework, we compared the size, shape, and orientation of G matrices and assessed their potential to facilitate or constrain trait evolution. Flowering time, floral display and rosette size varied among populations and significant additive genetic variation within populations indicated potential to evolve in response to selection. Yet, some characters, including flowering start and number of flowers, may not evolve independently because of genetic correlations. Using a multivariate framework, we found few differences in the genetic architecture of traits among populations. G matrices varied mostly in size rather than shape or orientation. Differences in multivariate responses to selection predicted from differences in G were small, suggesting overall matrix similarity and shared constraints to trait evolution among populations.     

Variation in pollinator abundance and selection on fragrance phenotypes in an epiphytic orchid

Extraordinary floral variation is common among some orchids that employ deception to attract pollinators. This variation may be maintained by frequency-dependent selection where rare phenotypes are preferred. Over a 2-yr period, 1993-1994, we monitored the reproductive success of Tolumnia variegata, an obligately outcrossing epiphytic orchid, at three localities in Puerto Rico that differed in pollinator service. Plants varied in floral morphology and fragrance characteristics. Artificial arrays of varying frequencies of scentless and fragrant phenotypes were established to test for frequency-dependent selection. Where pollinators were rare (Cambalache, range of census average = 0-0.2 bees/h), 0.9-1.2% of the flowers were effectively visited (pollinarium removals and pollinations). At Tortuguero where 0.4-1.1 bees/h were observed, 4-9.2% of the flowers were visited. At Pi;atnones where bees were the most abundant (1.4-5.2 bees/h), 20.9-25.0% of the flowers were visited. A significant portion of the variance in all measures of reproductive success (male, female, and combined) was explained by differences among populations, which we attribute mostly to variation in pollinator abundance. Neither the fragrance phenotype nor its frequency had a significant effect on success as revealed by a split-plot ANOVA. There was a significant interaction between population and phenotypic frequencies in all our measures of reproductive success, but only for the 1994 flowering season. Thus, variation in floral fragrance phenotypes is not likely maintained by frequency-dependent selection. High levels of variation remain unexplained.     

Floral trait differentiation in Anacamptis coriophora: Phenotypic selection on scents,but not on colour

Current divergent selection may promote floral trait differentiation among conspecific populations in flowering plants. However, whether this applies to complex traits such as colour or scents has been little studied, even though these traits often vary within species. In this study, we compared floral colour and odour as well as selective pressures imposed upon these traits among seven populations belonging to three subspecies of the widespread, generalist orchid Anacamptis coriophora. Colour was characterized using calibrated photographs, and scents were sampled using dynamic headspace extraction and analysed using gas chromatography–mass spectrometry. We then quantified phenotypic selection exerted on these traits by regressing fruit set values on floral trait values. We showed that the three studied subspecies were characterized by different floral colour and odour, with one of the two predominant floral volatiles emitted by each subspecies being taxon‐specific. Plant size was positively correlated with fruit set in most populations, whereas we found no apparent link between floral colour and female reproductive success. We detected positive selection on several taxon‐specific compounds in A. coriophora subsp. fragrans, whereas no selection was found on floral volatiles of A. coriophora subsp. coriophora and A. coriophora subsp. martrinii. This study is one of the first to document variation in phenotypic selection exerted on floral scents among conspecific populations. Our results suggest that selection could contribute to ongoing chemical divergence among A. coriophora subspecies.     

Sources of variation in floral nectar production rate in Epilobium canum (Onagraceae): implications for natural selection

Sources of variation in floral nectar production were investigated in a natural population of Epilobium canum (Onagraceae), a hummingbird-pollinated herbaceous shrub. Field measurements showed significant phenotypic variation among plants in floral nectar production rates. Average variance among flowers within plants was approximately one-third to one-half as great as variance among plants, with coefficients of variation among flowers ranging from 6.5% to 116.7%. A greenhouse experiment using clonally propagated ramets from field plants showed significant genetic variation for nectar production rates; broad sense heritability was estimated to have a maximum value of 0.64. In the greenhouse, plants grown under low water or low light conditions produced approximately 25% less nectar on average than those grown under control conditions. However, significant genotype-environment interactions indicated that genets differed in their responses to the changes in conditions. Rank correlations for genet mean nectar production rates across environmental conditions were low, and in two out of three comparisons were not different from zero. It is concluded that although the opportunity for natural selection on nectar production rates exits in this population, the response to selection will likely be slow, and the opportunity for selection of a narrow-optimum nectar production phenotype may be limited. Received: 9 January 1996 / Accepted: 18 December 1996