普通野生稻小种群的交配系统与遗传多样性

小种群的遗传动态是保育遗传学关注的核心问题之一,而种群遗传动态又与交配系统密切相关.普通野生稻(Oryza rufipogon Griff.)是具有重要经济价值的濒危物种,目前其种群规模都较小,研究其小种群交配系统与遗传变异性对普通野生稻的保护具有重要意义.运用7对SSR引物,对采自江西东乡普通野生稻小种群的36份种茎和其中20个家系共计601份子代进行了分析.结果显示:该种群的表观异交率为0.318,多位点法估计(MLTR)的多位点异交率为0.481;50%以上的子代共享亲本,非随机交配明显;东乡普通野生稻种群交配系统属于混合交配类型.比较亲本和子代种群的遗传变异性显示:子代种群比亲本种群遗传变异性更丰富;子代种群的杂合子不足与种群变小自交比例上升有关;而亲本种群杂合子过剩可能与杂合基因型的选择优势有关.这些结果说明创造条件扩大种群规模对普通野生稻的原生境保护显得尤为重要.     

马尾松二代无性系种子园子代父本分析及花粉散布

目前国内较早建立的马尾松(Pinus massoniana)二代种子园正陆续进入正常开花结实期。研究马尾松二代种子园花粉散布和自由授粉子代的父本组成, 可为生产上指导马尾松高世代种子园的规划设计和遗传管理提供理论依据。该文利用筛选的11对SSR引物, 对马尾松二代无性系种子园内8个无性系单株的320个自由授粉子代和48个候选父本进行了扩增, 并采用最大似然法对子代进行父本分析。结果表明: 11个位点共检测到61个等位基因, 每个位点的等位基因数在2-11之间, 平均为5.55个。试验亲本和子代群体的总平均观测杂合度(H_o)、期望杂合度(H_e)及多态信息含量(PIC)分别为0.428、0.433和0.387。在80%的可信度水平下可为232 (72.50%)个子代确定其父本。平均每个采种母树与19个父本产生子代。在自由授粉状态下, 马尾松二代种子园自交率为1.72%, 自交现象很弱, 其交配方式以异交为主。绝大多数亲本无性系的雄性繁殖适合度在1.00%-4.00%之间, 候选父本平均繁殖适合度为2.17%, 平均形成5个后代。马尾松有效花粉的散布距离和固定交配距离的父本繁殖适合度均符合正态分布, 两者呈极显著负相关, 其主要散布距离集中在0-100 m, 而检测到的最大散布距离为192 m。种子园花粉污染率较低, 仅为4.06%。总体看来, 树冠南面子代亲本交配距离较北面有增加的趋势, 但树冠南、北面子代父本组成数并未表现明显的规律。     

矮慈姑的传粉机制与交配系统

对矮慈姑（ＳａｇｉｔａｒｉａｐｙｇｍａｅａＭｉｑ）的传粉过程与花粉流作了观察,用同工酶遗传标记法对其一个自然居群的异交率作出定量估计。在自然及人工居群中均观察到虫媒传粉,其中有较大比例的近距离传粉。未检出风媒花粉流。自交可育,同时存在雌雄同株和花序内雌雄花异熟等异交机制。异交率估计结果（ｔ＝４９９％）表明其交配系统为异交／自交兼性系统。作者认为,其较大比例的自交应归因于同株之不同克隆分株间交配的频繁发生     

植物交配系统的进化、资源分配对策与遗传多样性

影响植物自交率进化的选择力量主要体现在两个方面：当外来花粉量不足时,自交可以提高植物的结实率,即雌性适合度（繁殖保障）;而如果进行自交的花粉比异交花粉更易获得使胚珠受精的机会,那么自交也可以提高植物的雄性适合度（自动选择优势）。但是,鉴别什么时候是繁殖保障、什么时候是自动选择优势导致了自交的进化却是极其困难的。花粉贴现降低了自交植物通过异交花粉途径获得的适合度,即减弱了自动选择优势,而近交衰退既减少了自动选择优势也减少了繁殖保障给自交者带来的利益。具有不同交配系统的植物种群将具有不同的资源分配对策。理论研究已经说明,自交率增加将减少植物对雄性功能的资源分配比例,但将使繁殖分配加大,而且在一定条件下交配系统的改变甚至可以导致植物生活史发生剧烈变化,即从多年生变为一年生。文献中支持自交减少植物雄性投入的证据有很多,但是对繁殖分配与自交率的关系目前还没有系统的研究。资源分配理论可以解释植物繁育系统的多样性,尤其是能够说明为什么大多数植物都是雌雄同体的。自交对植物种群遗传结构的影响是减少种群内的遗传变异,增加种群间的遗传分化。长期以来人们一直猜测,自交者可能会丢掉一些长期进化的潜能,目前这个假说得到了一些支持。     

植物交酸系统的进化、资源分配对策与遗传多样性

影响植物自交率进化的选择力量主要体现在两个方面：当外来花粉量不足时,自交可以提高植物的结实率,即雌性适合度（繁殖保障）;而如果进行自交的花粉比异交花粉更易获得使胚珠受精的机会,那么自交也可以提高植物的雄性适合度（自动选择优势）。但是,鉴别什么时候是繁殖保障、什么时候是自动选择优势导致了自交的进化却是极其困难的。花粉贴现降低了自交植物通过异交花粉途径获得的适合度,即减弱了自动选择优势,而近交衰退既减少了自动选择优势也减少了繁残给自交者带来的利益。具有不同交配系统的植物种群将具有不同的资源分配对策。理论研究已经说明,自交率增加将减少植物对雄性功能的资源分配比例,但将使繁殖分配加大,而且在一定条件下交配系统在改变甚至可以导致植物生活史发生剧烈变化,即从多年生变为一年生。文献中支持自交减少植物雄性投入的证据有很多,但是对繁殖分配与自交率的关系目前还没有系统的研究,资源分配理论可以解释植物繁育系统的多样性,尤其是能够3说明为什么大多数植物都是雌雄同体的,自交对植物种群遗传结构的影响是减少种群内的遗传变异,增加种群间的遗传分化,长期以来人们一直猜测,自交者可能会丢掉一些长期进化的潜能,目前这个假说得到了一些支持。     

鹅掌楸不同交配组合子代遗传多样性分析

揭示不同交配类型与子代遗传多样性的关系, 对于林木杂交育种及种子园管理具有理论与实践意义。文章选取来自鹅掌楸、北美鹅掌楸及杂交鹅掌楸的16个交配亲本, 共组配14个杂交组合, 分属5种交配类型, 分别为种间杂交、种内交配、多父本混合授粉、回交、以及自交。每个交配组合随机抽取30个子代, 利用SSR分子标记检测各子代群体遗传多样性以及16个交配亲本间的遗传距离。结果表明, 总体上, 鹅掌楸交配子代群体具有较高的遗传多样性。5种交配类型子代群体中, 遗传多样性水平由高至低的趋势为: 多父本混合授粉子代、种间交配子代、杂种F1与亲本的回交子代、种内交配子代, 自交子代。子代遗传多样性与亲本间遗传距离呈显著正相关, 表明亲本间遗传距离大, 则子代遗传多样性高。相同亲本正反交子代群体的遗传多样性差别不明显。     

二型花柱植物滇丁香二态和单态种群间花部形态变异模式（英文）

花部形态特征在植物交配系统的演化与维持过程中起关键作用,交配方式的转变将可能伴随着相应花部形态的调整。为探寻交配系统与花形态变异之间的联系,本研究对二型花柱植物滇丁香(Luculia pinceana)的二态和单态种群的花部形态特征进行比较分析。结果表明花部形态在两种不同花型构造的种群间分化程度显著大于花型构造相同的种群间变异。相对于二态种群而言,长花柱的单态种群中雌雄异位距离缩小,且个体间雌雄生殖器官在空间上的非法重叠程度增高,花部形态的调整能够同时促进自交和同型异交。在异型花柱物种内,异交为主的二型花柱种群向单态种群转变过程中自交比率预期上升,本研究表明花部形态的变异模式与交配系统转变相一致。     

基于生物学物种定义探讨物种形成理论与验证的研究进展

物种形成是进化生物学研究的一个永恒主题, 由于生物群体进化是连续和动态的, 物种界限变得难于界定。本文首先讨论了3种地理物种形成模式(同域、邻域及异域), 并分析了近期报道的研究证据。其次, 评述了合子后生殖隔离机制的分子遗传基础和应用群体基因组数据分析的证据, 包括BDMI模型(Bateson-Dobzhansky-Muller incompatibility)、QTLs (quantitative trait loci)、霍尔丹法则及大X染色体效应。最后, 探讨了交配系统作为合子前隔离机制之一与物种形成的关系, 认为近交或自交通过扩大种群遗传结构分化, 增强不同交配系统的种群间不对称基因渐渗, 或种群间无基因渐渗等途径, 促进新物种形成。已知植物交配系统的演化更倾向于从异交(或自交不亲和)向自交(或近交亲和)方式, 花性状和基因组的分化推动形成所谓的自交综合征, 研究交配系统驱动或强化物种形成模式对认识植物物种形成机制有重要意义。     

鹅掌楸配子选择与雄性繁殖适合度

在多父本等量花粉混合授粉的交配实验设计基础上, 利用SSR分子标记对其子代进行父本分析, 研究鹅掌楸的配子选择与雄性繁殖适合度。参试亲本为鹅掌楸2个种的5个单株（鹅掌楸（Liriodendron chinense）2株, 即FY和LS; 北美鹅掌楸（L. tulipifera）3株, 即LYS、MSL和NK）。结果表明： 鹅掌楸的配子选择个体间差异较大。作为母本, NK和LYS倾向于选择异种雄配子, 而MSL和LS则倾向于选择同种雄配子; 在同种雄配子的选择中, NK、LYS和LS倾向于自交, 而MSL则倾向于异交。以北美鹅掌楸为母本时, 北美鹅掌楸与鹅掌楸的雄性繁殖贡献率分别为45.5%和54.5%, 北美鹅掌楸的雄性繁殖适合度为鹅掌楸的0.556倍。以鹅掌楸为母本时, 二者的繁殖贡献率分别为15.6%和84.4%, 北美鹅掌楸的雄性繁殖适合度为鹅掌楸的0.123倍。总体上看, 鹅掌楸的雄性繁殖适合度高于北美鹅掌楸, 鹅掌楸与北美鹅掌楸均表现为自交亲和。     

增温条件下花粉来源对红雉凤仙花生殖成功的影响

花粉来源影响着植物的生殖成功和种群的遗传结构, 母体效应则有利于子代适应母代经历的环境。然而, 气候变暖背景下, 花粉来源联合母体效应对植物生殖成功的影响仍不明了。该研究以兼性异交的中国特有植物红雉凤仙花(Impatiens oxyanthera)为研究对象, 采用双因素随机区组设计, 设置母代增温(+0 ℃和+2 ℃)嵌套子代增温(+0 ℃、+2 ℃和+4 ℃)共6种实验处理, 探讨不同母代增温条件下子代增温对红雉凤仙花花粉来源限制的影响。结果显示: 红雉凤仙花异交的结实率和种子数均高于自交的相应值, 且随着子代增温幅度提升, 这2个值均呈现不同程度降低, 特别是自交结实率和种子数随子代增温明显降低, 而母代增温则减缓了自交引起的生殖成功率降低的效应。上述结果说明, 红雉凤仙花异交生殖成功率大于自交。子代增温显著降低了兼性异交植物自交的适合度, 母代增温可以减缓这种不利影响, 从而表现出对增温的跨代适应性。     

Parentage versus two-generation analyses for estimating pollen-mediated gene flow in plant populations

Assessment of contemporary pollen-mediated gene flow in plants is important for various aspects of plant population biology, genetic conservation and breeding. Here, through simulations we compare the two alternative approaches for measuring pollen-mediated gene flow: (i) the NEIGHBORHOOD model--a representative of parentage analyses, and (ii) the recently developed TWOGENER analysis of pollen pool structure. We investigate their properties in estimating the effective number of pollen parents (N(ep)) and the mean pollen dispersal distance (delta). We demonstrate that both methods provide very congruent estimates of N(ep) and delta, when the methods' assumptions considering the shape of pollen dispersal curve and the mating system follow those used in data simulations, although the NEIGHBORHOOD model exhibits generally lower variances of the estimates. The violations of the assumptions, especially increased selfing or long-distance pollen dispersal, affect the two methods to a different degree; however, they are still capable to provide comparable estimates of N(ep). The NEIGHBORHOOD model inherently allows to estimate both self-fertilization and outcrossing due to the long-distance pollen dispersal; however, the TWOGENER method is particularly sensitive to inflated selfing levels, which in turn may confound and suppress the effects of distant pollen movement. As a solution we demonstrate that in case of TWOGENER it is possible to extract the fraction of intraclass correlation that results from outcrossing only, which seems to be very relevant for measuring pollen-mediated gene flow. The two approaches differ in estimation precision and experimental efforts but they seem to be complementary depending on the main research focus and type of a population studied.     

The evolutionary response of mating system to heterosis

Isolation allows populations to diverge and to fix different alleles. Deleterious alleles that reach locally high frequencies contribute to genetic load, especially in inbred or selfing populations, in which selection is relaxed. In the event of secondary contact, the recessive portion of the genetic load is masked in the hybrid offspring, producing heterosis. This advantage, only attainable through outcrossing, should favour evolution of greater outcrossing even if inbreeding depression has been purged from the contributing populations. Why, then, are selfing‐to‐outcrossing transitions not more common? To evaluate the evolutionary response of mating system to heterosis, we model two monomorphic populations of entirely selfing individuals, introduce a modifier allele that increases the rate of outcrossing and investigate whether the heterosis among populations is sufficient for the modifier to invade and fix. We find that the outcrossing mutation invades for many parameter choices, but it rarely fixes unless populations harbour extremely large unique fixed genetic loads. Reversions to outcrossing become more likely as the load becomes more polygenic, or when the modifier appears on a rare background, such as by dispersal of an outcrossing genotype into a selfing population. More often, the outcrossing mutation instead rises to moderate frequency, which allows recombination in hybrids to produce superior haplotypes that can spread without the mutation's further assistance. The transience of heterosis can therefore explain why secondary contact does not commonly yield selfing‐to‐outcrossing transitions.     

Inbreeding depression in Vriesea gigantea,a perennial bromeliad from southern Brazil

Inbreeding depression is a reduction of fitness in the progeny of closely related individuals and its effects are assigned to selfing or biparental inbreeding. Vriesea gigantea is a self‐compatible bromeliad species distributed in the Brazilian Atlantic rainforest and habitat destruction and fragmentation and collection have decreased the natural populations. We aim to describe the occurrence of inbreeding depression (δ) in three natural populations of V. gigantea and to correlate this phenomenon with previous studies of fertility, genetic diversity, population genetic structure, gene flow, mating system and seed dispersal in this species. Fifty‐four adult plants were sampled and 108 flowers were used for pollination treatments (selfing, outcrossing and control). For adult plants, we analysed plant and inflorescence height, flower numbers and seed set. In the progenies, evaluated parameters included seed germination and seedling survival rate. The results indicated low to moderate levels of inbreeding depression in V. gigantea (δ = 0.02 to 0.39), in agreement with molecular data from a previous study. Vriesea gigantea populations tolerate some degree of inbreeding, which is consistent with previous results on fertility, mating system, genetic diversity and gene flow. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169 , 312–319.     

Bayesian approach reveals confounding effects of population size and seasonality on outcrossing rates in a fragmented subalpine conifer

Mating systems have long been recognized as key factors determining genetic structure within and between populations. Outcrossing promotes genetic diversity and gene flow between populations, while inbreeding, on the other hand, decreases recombination rates, facilitating fixation of co-adapted genes. In small populations, selfing moderates pollen limitation because of low mate availability, but at the cost of increased inbreeding depression. These conflicts are of more than theoretical interest; they are critical for the management of endangered species. In order to help designing conservation strategies for the management of the gene pool of fragmented populations of Pinus cembra, a protected species in Poland, we have characterized pollen flow and mating structure using nuclear microsatellite markers. We demonstrated that P. cembra in the studied stands of the Tatra Mts. is characterized by an average outcrossing rate (t) of 0.72. Unlike with the existing approaches, using the newly developed Bayesian method, we found that population size and seasonal variation had confounding effects on outcrossing rates. In concordance with predictions, large populations showed significantly higher outcrossing rates (t?=?0.89) than smaller ones (t?=?0.51). Temporal variation revealed in the outcrossing rate might be linked with masting behavior of the species. On the other hand, we showed that outcrossing rates were not associated with a trunk diameter of a mother tree. Our study also demonstrated that biparental inbreeding is a significant component of mating system. However, we further show that pollen dispersal follows a fat-tailed distribution (with the average dispersal distance of 1,267 m) so that at least some long-distance pollen dispersal must be occurring. Overall, we conclude that the high inbreeding (both selfing and mating between relatives) found in P. cembra buffers for pollen limitation. We argue that small, isolated stands can be at risk of gene pool erosion, despite the potential for long-distance pollen and seed dispersal.     

Using genetic markers to directly estimate gene flow and reproductive success parameters in plants on the basis of naturally regenerated seedlings

Estimating seed and pollen gene flow in plants on the basis of samples of naturally regenerated seedlings can provide much needed information about "realized gene flow," but seems to be one of the greatest challenges in plant population biology. Traditional parentage methods, because of their inability to discriminate between male and female parentage of seedlings, unless supported by uniparentally inherited markers, are not capable of precisely describing seed and pollen aspects of gene flow realized in seedlings. Here, we describe a maximum-likelihood method for modeling female and male parentage in a local plant population on the basis of genotypic data from naturally established seedlings and when the location and genotypes of all potential parents within the population are known. The method models female and male reproductive success of individuals as a function of factors likely to influence reproductive success (e.g., distance of seed dispersal, distance between mates, and relative fecundity--i.e., female and male selection gradients). The method is designed to account for levels of seed and pollen gene flow into the local population from unsampled adults; therefore, it is well suited to isolated, but also wide-spread natural populations, where extensive seed and pollen dispersal complicates traditional parentage analyses. Computer simulations were performed to evaluate the utility and robustness of the model and estimation procedure and to assess how the exclusion power of genetic markers (isozymes or microsatellites) affects the accuracy of the parameter estimation. In addition, the method was applied to genotypic data collected in Scots pine (isozymes) and oak (microsatellites) populations to obtain preliminary estimates of long-distance seed and pollen gene flow and the patterns of local seed and pollen dispersal in these species.     

Postglacial ecotype formation under outcrossing and self‐fertilization in Arabidopsis lyrata

The formation of ecotypes has been invoked as an important driver of postglacial biodiversity, because many species colonized heterogeneous habitats and experienced divergent selection. Ecotype formation has been predominantly studied in outcrossing taxa, while far less attention has been paid to the implications of mating system shifts. Here, we addressed whether substrate‐related ecotypes exist in selfing and outcrossing populations of Arabidopsis lyrata subsp. lyrata and whether the genomic footprint differs between mating systems. The North American subspecies colonized both rocky and sandy habitats during postglacial range expansion and shifted the mating system from predominantly outcrossing to predominantly selfing in a number of regions. We performed an association study on pooled whole‐genome sequence data of 20 selfing or outcrossing populations, which suggested genes involved in adaptation to substrate. Motivated by enriched gene ontology terms, we compared root growth between plants from the two substrates in a common environment and found that plants originating from sand grew roots faster and produced more side roots, independent of mating system. Furthermore, single nucleotide polymorphisms associated with substrate‐related ecotypes were more clustered among selfing populations. Our study provides evidence for substrate‐related ecotypes in A. lyrata and divergence in the genomic footprint between mating systems. The latter is the likely result of selfing populations having experienced divergent selection on larger genomic regions due to higher genome‐wide linkage disequilibrium.     

Allee effect and self-fertilization in hermaphrodites: reproductive assurance in a structured metapopulation

Reproductive assurance through selfing during colonization events or when population densities are low has often been put forward as a mechanism selecting for the evolution of self-fertilization. Such arguments emphasize on the role of both local demography and metapopulation processes. We developed a model for the evolution of self-fertilization in a structured metapopulation in which local densities are not steady because of population growth. Reproduction by selfing is density-independent (reproductive assurance) but selfed seeds endure inbreeding depression, whereas reproduction by outcrossing is density-dependent (Allee effect). First, we derived an analytical criterion for metapopulation viability as a function of the selfing rate and metapopulation parameters. We show that outcrossers can develop a viable metapopulation when they produce a high amount of dispersal seeds that counterbalances their incapacity to found new populations from low densities. Second, the model shows there is a positive feedback between demography and outcrossing rates, leading to either complete outcrossing or selfing. Specifically, we illustrate that inbreeding depression can paradoxically favor the evolution of selfing because of its negative effect on density. Also, complete outcrossing can be selected despite pollen limitation, although it does not provide a full seed set. This model underlines the influence of the mating system both on demography and gene dynamics in a metapopulation context.     

SEXUAL PARTNERS FOR THE STRESSED: FACULTATIVE OUTCROSSING IN THE SELF-FERTILIZING NEMATODE CAENORHABDITIS ELEGANS

Sexual reproduction shuffles genetic variation, potentially enhancing the evolutionary response to environmental change. Many asexual organisms respond to stress by generating facultative sexual reproduction, presumably as a means of escaping the trap of low genetic diversity. Self-fertilizing organisms are subject to similar genetic limitations: the consistent loss of genetic diversity within lineages restricts the production of variation through recombination. Selfing organisms may therefore benefit from a similar shift in mating strategy during periods of stress. We determined the effects of environmental stress via starvation and passage through the stress-resistant dauer stage on mating system dynamics of Caenorhabditis elegans , which reproduces predominantly through self-fertilization but is capable of outcrossing in the presence of males. Starvation elevated male frequencies in a strain-specific manner through differential male survival during dauer exposure and increased outcrossing rates after dauer exposure. In the most responsive strain, the mating system changed from predominantly selfing to almost exclusively outcrossing. Like facultative sex in asexual organisms, facultative outcrossing in C. elegans may periodically facilitate adaptation under stress. Such a shift in reproductive strategy should have a major impact on evolutionary change within these populations and may be a previously unrecognized feature of other highly selfing organisms.     

Fine-scale estimation of outcrossing in western redcedar with microsatellite assay of bulked DNA

Western redcedar (Thuja plicata, Cupressaceae) is a self-fertile conifer with a mixed mating system and significant variation for outcrossing among populations. In this paper, we conducted a fine-scale study of mating system variation to identify correlates of outcrossing in natural populations. We examined variation for outcrossing within and among individual trees, and describe a new method to estimate outcrossing using bulked DNA samples. Bulking (assaying DNA tissues from several individuals simultaneously) increases the experimental power without increasing the experimental effort. We sampled 80 trees from four natural populations in southwestern British Columbia. From each tree, we sampled from up to six crown positions (three heights and inner vs outer branches). From each position, two samples of three seedlings each were bulked before DNA extractions. Using four microsatellite loci, we obtained outcrossing rates for each tree and for each of the six crown positions. We found individual tree selfing rates to increase with tree height in all four populations, but selfing rates did not differ among crown positions. The higher selfing rate of larger trees is probably due to their greater proportional contribution to local pollen clouds. Individual tree outcrossing rates ranged from 22 to 100% and the population outcrossing rates from 66 to 78%. Missed alleles due to bulking and the estimation method used both cause a downward bias in outcrossing rates, so that these estimates are probably lower than the actual outcrossing rates. Nevertheless, the trends we observed are not affected by systematic biases of estimation.     

Consequences of multiple mating‐system shifts for population and range‐wide genetic structure in a coastal dune plant

Evolutionary transitions from outcrossing to selfing can strongly affect the genetic diversity and structure of species at multiple spatial scales. We investigated the genetic consequences of mating‐system shifts in the North American, Pacific coast dune endemic plant Camissoniopsis cheiranthifolia (Onagraceae) by assaying variation at 13 nuclear (n) and six chloroplast (cp) microsatellite (SSR) loci for 38 populations across the species range. As predicted from the expected reduction in effective population size (N_e) caused by selfing, small‐flowered, predominantly selfing (SF) populations had much lower nSSR diversity (but not cpSSR) than large‐flowered, predominantly outcrossing (LF) populations. The reduction in nSSR diversity was greater than expected from the effects of selfing on N_e alone, but could not be accounted for by indirect effects of selfing on population density. Although selfing should reduce gene flow, SF populations were not more genetically differentiated than LF populations. We detected five clusters of nSSR genotypes and three groups of cpSSR haplotypes across the species range consisting of parapatric groups of populations that usually (but not always) differed in mating system, suggesting that selfing may often initiate ecogeographic isolation. However, lineage‐wide genetic variation was not lower for selfing clusters, failing to support the hypothesis that selection for reproductive assurance spurred the evolution of selfing in this species. Within three populations where LF and SF plants coexist, we detected genetic differentiation among diverged floral phenotypes suggesting that reproductive isolation (probably postzygotic) may help maintain the striking mating‐system differentiation observed across the range of this species.