Floral phenology and compatibility of sawgrass, Cladium jamaicense (Cyperaceae)

Sawgrass, Cladium jamaicense, is the dominant macrophyte in the Florida Everglades. We examined sawgrass flowering phenology and compatibility reactions in ex situ and in situ populations over 2 yr. Sawgrass flowers in May in southern Florida. Flower maturation was relatively synchronous within an inflorescence. Along the entire inflorescence, functionally male flowers emerged initially, followed by stigmas, then anthers of hermaphroditic flowers. Flowers of each sex expanded over 2 d with less than 1 d in between, totaling 6- 7 d for an inflorescence to complete flowering. Hand pollinations showed that sawgrass was self-compatible and not pollen-limited, because open pollinations produced fruit set similar to self- and cross-pollinations. Fruit set was low in autogamy and manipulation treatments. Manipulation treatments were used to study the effect of exposure to airborne pollen during hand pollinations. This treatment thus provides a useful technique for studies on the in situ compatibility of wind-pollinated graminoids. Sawgrass was able to self-fertilize, but the timing of flower maturation on an inflorescence promoted outcrossing. Actual outcrossing rates in sawgrass thus depend on clonal architecture and the timing of floral maturation on other inflorescences within a clone rather than on inflorescences of other genets in a population.     

Floral morphometrics and the evolution of sexual dimorphism in Lycium (Solanaceae)

Plants of Lycium californicum, L. exsertum, and L. fremontii produce flowers that are either male-sterile (female) or hermaphroditic, and populations are morphologically gynodioecious. As is commonly found in gynodioecious species, flowers on female plants are smaller than those on hermaphrodites for a number of floral traits. Floral size dimorphism has often been hypothesized to be the result of either a reduction in female flower size that allows reallocation to greater fruit and seed production, or an increase in hermaphroditic flower size due to the increased importance of pollinator attraction and pollen export for hermaphroditic flowers. We provide a test of these two alternatives by measuring 11 floral characters in eight species of Lycium and using a phylogeny to reconstruct the floral size shifts associated with the evolution of gender dimorphism. Our analyses suggest that female flowers are reduced in size relative to the ancestral condition, whereas flowers on hermaphrodites have changed only slightly in size. Female and hermaphroditic flowers have also diverged both from one another and from ancestral cosexual species in several shape characteristics. We expected sexual dimorphism to be similar among the three dimorphic taxa, as gender dimorphism evolved only a single time in the ancestor of the American dimorphic lineage. While the floral sexual dimorphism is broadly similar among the three dimorphic species, there are some species-specific differences. For example, L. exsertum has the greatest floral size dimorphism, whereas L. fremontii had the greatest size-independent dimorphism in pistil characters. To determine the degree to which phylogenetic uncertainty affected reconstruction of ancestral character states, we performed a sensitivity analysis by reconstructing ancestral character states on alternative topologies. We argue that investigations such as this one, that examine floral evolution from an explicitly phylogenetic perspective, provide new insights into the study of the evolution of floral sexual dimorphism.     

水烛的花部形态观察——兼论与长苞香蒲的分类学界定

通过腊叶标本研究、野外观察和文献考证,结合栽培试验,研究了香蒲科水烛（Typha angustifolia L.）花部结构特征,补充描述了该种丝状毛在子房柄上的着生方式、小穗不孕雌花数目等性状特征,观察了在成熟期不同阶段其孕性雌花柱头与小苞片的长度变异。结果显示,水烛孕性雌花小苞片呈宽披针形、匙形或条形,先端褐色,短于柱头,或与柱头近等长或稍长于柱头;子房柄上的丝状毛除少数散生外,多数基部合生呈鞘状或束状,在子房柄下部呈1～4轮排列;小穗不孕雌花常3（～4）枚。研究材料在7月中旬前后雌花小苞片明显短于柱头,随果穗成熟小苞片与柱头近等长。长苞香蒲（T.domingensis Pers.）子房柄上的丝状毛形态和着生方式与水烛中的情况基本一致,但小苞片白色透明,小穗不孕雌花常1（～2）枚。这表明水烛孕性雌花小苞片和柱头的长度比例与不同成熟阶段有关系,不宜作为与长苞香蒲的区别特征,而小穗不孕雌花数目和小苞片颜色等特征对两物种的划分有较重要的意义。     

雌全同株植物三脉紫菀花期偏雄的个体大小依赖的性别分配

经典的虫媒传粉植物个体大小依赖的性别分配模型通常预期:分配给雌性功能的资源比例将随着个体大小的增大而增加;但一些研究表明,花期个体大小依赖的性别分配模式表现出随个体大小增大而偏雄的趋势.我们以植株高度衡量个体大小,从花和花序两个水平上研究了雌花、两性花同株植物三脉紫菀(Aster ageratoides)花期个体大小依赖的性别分配策略.随着植株高度的增大,植株产生的头状花序数量增加,表明三脉紫菀投入到繁殖的资源不是固定不变的,而是随个体大小增大而增加的.在花和花序水平上,繁殖资源在雌雄性别功能之间的分配均表现为随个体大小的增大而更偏雄的模式,即花粉/胚珠比增加,产生花粉的两性花占两性花和雌花总花数的比例升高.这些结果与花期个体越大、性别分配越偏雄的预期一致.花期更偏雄的性别分配可能有助于植物在花期通过输出花粉提高雄性适合度,从而实现个体适合度的最大化.     

Size-dependent gender modification in Lilium apertum (Liliaceae): does this species exhibit gender diphasy?

Background and Aims

Variation in the relative female and male reproductive success of flowering plants is widespread, despite the fundamental hermaphroditic condition of the majority of species. In many hermaphroditic populations, environmental conditions and their influence on development and size can influence the gender expression of individuals through the formation of hermaphroditic and unisexual flowers. This study investigates the hypothesis that the bulbous, animal-pollinated, perennial Lilium apertum (Liliaceae) exhibits a form of size-dependent gender modification known as gender diphasy, in which the sexual expression of individuals depends on their size, with plants often changing sex between seasons.

Methods

Variation in floral traits was examined in relation to their size using marked individuals in natural populations, and also under glasshouse conditions. Measurements were taken of the height, flower number, floral sex expression, flower size, flower biomass and pollen production of individuals over consecutive years between 2009 and 2012 in seven populations in south-west China.

Key Results

Flowers of L. apertum are either perfect (hermaphroditic) or staminate (male) and, in any given season, plants exhibit one of three sex phenotypes: only hermaphrodite flowers, a mixture of hermaphroditic and male flowers, or only male flowers. Transitions between each of these sex phenotypes were observed over consecutive years and were commonly size-dependent, particularly transitions from small plants bearing only male flowers to those that were taller with hermaphroditic flowers. Hermaphroditic flowers were significantly larger, heavier and produced more pollen than male flowers.

Conclusions

The results for L. apertum are consistent with the ‘size advantage hypothesis’ developed for animal species with sex change. The theory predicts that when individuals are small they should exhibit the sex for which the costs of reproduction are less, and this usually involves the male phase. L. apertum provides an example of gender diphasy, a rare sexual system in flowering plants.     

Flower or spikelet? Understanding the morphology and development of reproductive structures in Exocarya (Cyperaceae, Mapanioideae, Chrysitricheae)

Fundamental questions of floral morphology remain unresolved in the grasslike monocots in order Poales, including what constitutes a flower and what constitutes a spikelet. The mapaniid sedges have particularly complex spikeletlike structures, variously interpreted as clusters of flowers or spikelets. Recent phylogenetic studies of Cyperaceae have identified the mapaniid clade as sister to the rest of the family, but the homology of mapaniid reproductive units (RUs) and spikeletlike units (SLUs) to other sedge flowers and spikelets is unclear. We examined reproductive development in the mapaniid Exocarya sclerioides. Inflorescence branches terminated in a SLU with bracts and 1-4 RUs. RUs had four small leaflike structures (LLSs): two lateral LLSs, each associated with a stamen, an abaxial LLS associated with a stamen, and an adaxial LLS. The gynoecium terminated the RU. All RUs were axillary to bracts, and unexpanded bracts and RUs were produced beyond expanded RUs, so SLUs were racemose. RUs developed from a single primordium that initiated two lateral LLSs, then two lateral stamens, then the gynoecium. Initiation of the abaxial LLS and stamen and the adaxial LLS followed. We hypothesize that the RU is a sympodial branch that terminates in a hermaphroditic flower with two stamens and a gynoecium; the two lateral LLSs are halves of a deeply divided prophyll.     

Spikelet structure and development in Cyperoideae (Cyperaceae): a monopodial general model based on ontogenetic evidence

Background and Aims

In Cyperoideae, one of the two subfamilies in Cyperaceae, unresolved homology questions about spikelets remained. This was particularly the case in taxa with distichously organized spikelets and in Cariceae, a tribe with complex compound inflorescences comprising male (co)florescences and deciduous female single-flowered lateral spikelets. Using ontogenetic techniques, a wide range of taxa were investigated, including some controversial ones, in order to find morphological arguments to understand the nature of the spikelet in Cyperoideae. This paper presents a review of both new ontogenetic data and current knowledge, discussing a cyperoid, general, monopodial spikelet model.

Methods

Scanning electron microscopy and light microscopy were used to examine spikelets of 106 species from 33 cyperoid genera.

Results

Ontogenetic data presented allow a consistent cyperoid spikelet model to be defined. Scanning and light microscopic images in controversial taxa such as Schoenus nigricans, Cariceae and Cypereae are interpreted accordingly.

Conclusions

Spikelets in all species studied consist of an indeterminate rachilla, and one to many spirally to distichously arranged glumes, each subtending a flower or empty. Lateral spikelets are subtended by a bract and have a spikelet prophyll. In distichously organized spikelets, combined concaulescence of the flowers and epicaulescence (a newly defined metatopic displacement) of the glumes has caused interpretational controversy in the past. In Cariceae, the male (co)florescences are terminal spikelets. Female single-flowered spikelets are positioned proximally on the rachis. To explain both this and the secondary spikelets in some Cypereae, the existence of an ontogenetic switch determining the development of a primordium into flower, or lateral axis is postulated.     

Evidence of gynodioecy and sex ratio variation in Wurmbea biglandulosa (Colchicaceae)

 A valuable approach to understanding the evolution of gender dimorphism involves studies of single species that exhibit intraspecific variation in sexual systems. Here we survey sex ratios in 35 populations of Wurmbea biglandulosa, previously described as hermaphroditic. We found pronounced intraspecific variation in sexual systems; populations in the northeastern part of the species' range were hermaphroditic, whereas other populations were gynodioecious and contained 2–44% females. Populations with lower annual rainfall were more likely to be gynodioecious, supporting the view that gender dimorphism evolves more frequently in harsher environments. In gynodioecious populations, however, female frequency was not related to either annual rainfall or habitat, indicating that other factors are important in determining sex ratio variation. Females had smaller flowers and shorter stems than did hermaphrodites, potentially providing a basis for resource compensation. A female fecundity advantage may contribute to the maintenance of females in populations because females produced more ovuliferous flowers and had more ovules per flower than did hermaphrodites. Received March 2, 2001 Accepted February 25, 2002     

Polygamy in Atriplex halimus L. (Chenopodiaceae)

It appears that up to now the inflorescence and flower morphologies of Atriplex halimus have been described incompletely. This species has been classified as monoecious or dioecious. Numerous observations and ontogenic studies have pointed to types of flowers morphologically and functionally hermaphroditic, never described until now. One specimen of this species presents both unisexual, male and female flowers and bisexual flowers, so Atriplex halimus is polygam and more precisely trimonoecious. Observation of inflorescences reveals a structure based on the spike and the dichasium. The sex distribution along the inflorescence axis was studied and the existence of a physiological gradient controlling its expression is discussed.     

新疆郁金香一居群个体性别7年的动态变化

在雌雄同株植物中, 花性别被认为是两性资源分配对环境条件的响应, 了解个体性别随时间的变化对探讨植物性系统的变异有重要意义。早春短命植物新疆郁金香(Tulipa sinkiangensis)多为两性花居群, 前期调查显示一些居群由雄花和两性花构成。为了探讨新疆郁金香雄花的发生与变化, 我们连续7年对一个居群的花性别及其变化动态进行了跟踪。结果显示: (1)该居群主要是由一朵花植株和两朵花植株构成, 分别占居群植株总数的74.5%和23.0%。两种性别的花随机分布在不同类型的单株上, 形成了以两性花植株为主, 含有雄株、雄花与两性花同株(andromonoecy)的居群结构。(2)雄花相对较小, 子房退化, 无胚珠发育, 在植株或居群中花期较晚出现。与相同大小的两性花相比, 雄花在单花花粉量、花粉败育率、花粉粒大小及形态、雄性功能等方面没有差异。(3) 7年间, 雄花在居群中的比例经历了2011-2014年间的显著下降(23.4-3.1%)和2015-2017年间相对稳定的零星分布(1.5-1.0%)两个阶段。居群中一朵花植株数量和两朵花植株数量在年份间呈波动性变化。(4)雄花的出现可能是植株受自身资源状况及环境条件等因素的影响在花性别选择上作出的可塑性反应。     

Sympodial structure of spikelets in the tribe Schoeneae (Cyperaceae)

Spikelet morphology of 250 specimens of 47 species of Schoeneae was examined using light microscopy and scanning electron microscopy. We confirmed that spikelet structure in Schoeneae is cymose with a sympodial "rachilla." Monopodial spikelets, as described by most current literature, were not found in Schoeneae. Prophylls are not always present in sympodial spikelets and cannot be used to determine whether a spikelet is sympodial or monopodial. Spikelets of Schoeneae develop acropetally, and the uppermost glume may or may not produce a flower. The last feature may be variable within species and within individual plants, so presence or absence of this flower is not an indication of sympodial or monopodial spikelet structure in Schoeneae. Relative position of flower, glume, and axis is a reliable criterion to judge whether spikelets are sympodial or monopodial. In some species of Schoenus and in Ptilothrix, formation of the arch-shaped base of the fertile glume relates to the shape of the inclined nodes on which the glume grows. This study highlights the need to reinvestigate spikelet structure in other tribes of Cyperaceae.     

Architectural effects mimic floral sexual dimorphism in Solanum (Solanaceae)

Factors underlying apparent floral sexual dimorphism were examined in six species of andromonoecious Solanum section Lasiocarpa (Solanaceae). Both multivariate and univariate analyses show that hermaphroditic flowers are significantly larger than staminate flowers for all features measured. Thus, flowers could be characterized as sexually size dimorphic. However, when size variation due to flower position (architecture) is controlled experimentally, differences between the floral genders for the nongynoecial characters disappear; there is no difference in corolla or androecium size. Staminate flowers appear to be generally smaller than hermaphroditic flowers, not because of any difference related to primary sexual function, but because they tend to occur in the distal regions of each inflorescence. In contrast, significant differences between hermaphroditic and staminate flowers for primary female traits (ovary, style, and stigma) remain after controlling for position: the two floral types are truly dimorphic for these characters. We show that consideration of architectural effects can direct and refine hypotheses concerning the evolution of andromonoecy. More generally, if architectural effects on flower size are common among taxa with unisexual flowers, then these effects may contribute to the common perception of size dimorphism in taxa with unisexual flowers.     

The ontogeny of flowering and sex expression in divergent populations of Lomatium grayi

Summary Populations of Lomatium grayi (Umbelliferae), an andromonoecious, perennial herb, differe in growth rates, flowering frequency, and survivorship. Effects of these different life histories on the ontogeny of sex expression were analyzed for plants from two populations grown from seedlings in a common garden and monitored for six years. Plants from Smoot Hill, Washington grew faster, had a higher probability of flowering at each age and size after the first year of growth, and a higher probability of flowering repeatedly among years than did plants from Clarkston, Washington. The proportion of plants producing some hermaphroditic flowers increased with plant size in both populations. Smoot Hill plants, however, were more likely to begin flowering as small, staminate plants than Clarkston plants. Clarkston plants did not begin flowering until they were older and larger, and most of these plants produced some hermaphroditic flowers when they began reproduction. The positive association between production of hermaphroditic flowers and both plant size and age was consistent with the hypothesis that hermaphroditic flowers are more costly to produce than staminate flowers.Although the populations did not differ in the total number of flowers per plant produced at any age or size, Smoot Hill plants consistently produced a lower percentage of hermaphroditic flowers than Clarkston plants at larger sizes and later ages. Consequently, selection for faster growth rates and higher flowering frequency at small sizes and early ages may have favored the more staminate-biased sex ratios in the Smoot Hill population.     

The roles of female and hermaphroditic flowers in the gynodioecious–gynomonoecious Silene littorea: insights into the phenology of sex expression

Some gynodioecious species have intermediate individuals that bear both female and hermaphroditic flowers. This phenomenon is known as a gynodioecious–gynomonoecious sexual system. Gender expression in such species has received little attention in the past, and the phenologies of male and female functions have also yet to be explored. In this study, we examined variations in gender patterns, their effects on female reproductive success and sex expression in depth throughout the flowering period in two populations. The studied populations of Silene littorea contained mostly gynomonoecious plants and the number of pure females was very low. The gynomonoecious plants showed high variability in the total proportion of female flowers. In addition, the proportion of female flowers in each plant varied widely across the flowering season. Although there was a trend towards maleness, our measures of functional gender suggested that most plants transmit their genes via both pollen and ovules. Fruit set and seed set were not significantly different among populations; in contrast, flower production significantly varied between the two populations – and among plants – with consequent variation in total seed production. Conversely, gender and sex expression were similar in both populations. Plants with higher phenotypic femaleness did not have higher fruit set, seed set or total female fecundity. The mating environment fluctuated little across the flowering period, but fluctuations were higher in the population with low flower production. We therefore conclude that the high proportion of gynomonoecious individuals in our studied populations of S. littorea may be advantageous for the species, providing the benefits of both hermaphroditic and female flowers.     

Andromonoecy in Solanum lycocarpum A. St. -Hil. (Solanaceae): Floral attributes,visitors and variation in sexual expression over time

Sexual expression in andromonoecious species—those in which a single individual can bear both staminate and hermaphroditic flowers—may vary among reproductive events in the same plant, among individuals and across populations. This variation influences, in turn, the individual contribution of hermaphroditic plants via male and female fitness functions (i.e., Lloydʼs phenotypic gender). However, temporal variation in sexual expression in andromonoecious species and its relationship with seasonal changes in climatic conditions remain poorly understood. Here we analyze floral attributes, visitors and variation in sexual expression in three populations of Solanum lycocarpum A. St. -Hil. Seasonality in the production of floral types, the mating system and floral visitors were also investigated. Hermaphroditic flowers produced more pollen grains, but the pollen of staminate flowers had higher viability. Only hermaphroditic flowers produced fruits, and ovules in staminate flowers were sterile. Solanum lycocarpum is mainly pollinated by large bees with the ability to vibrate flowers. Phenotypic gender varied throughout the year, and the seasonal production of staminate flowers is associated with the local climate. We suggest that the higher and seasonally variable relative abundance of staminate flowers compared to the low and uniform production of hermaphroditic flowers may be explained by (a) the very high energetic costs incurred in producing large fruits, which in turn make hermaphroditic flower production very costly, and (b) the potentially lower energy expenditure of the smaller staminate flowers with reduced pistils and non-viable ovules that allow them to rapidly respond to climate variability.     

Pollen and Resource Limitation in Veratrum nigrum L. （Liliaceae）, an Andromonoecious Herb

Pollen limitation and resource limitation were invoked to account for the pattern that flowering plants produce more flowers and ovules than fruits and seeds. This study aimed to determine their relative importance in Veratrum nigrum, a self-compatible, perennial, andromonoecious herb. In order to determine whether female production was limited by pollen grains on stigmas or by available resources, we performed supplemental hand pollination in three populations, male-flower-bud removal in three other populations, and emasculation of hermaphroditic flowers in still another population, resulting in a total of seven populations experimentally manipulated. Across the three populations, supplemental hand pollination did not significantly increase fruit set, seed number per fruit, and total seed production per individual, nor did emasculation of hermaphroditic flowers. Taken together, our results suggest that pollen grains deposited on stigmas were abundant enough to fertilize all the ovules. Male-flower-bud removal significantly increased the mean size of hermaphroditic flowers in all three populations. Female reproductive success was increased in one population, but not in the other two populations possibly due to heavy flower/seed predation. We concluded that the female reproductive success of V. nigrum was not limited by pollen grains but by available resources, which is consistent with Bateman＇s principle. Furthermore, the female reproduction increase of male-flower-bud removal individuals might suggest a trade-off between male and female sexual functions.     

Differences in sex expression and mating systems in three pollination morphs of Cimicifuga simplex

Cimicifuga simplex (Ranunculaceae) has three genetically distinct pollination morphs. Here, we report that each of the three pollination morphs of C. simplex differs from the others with regard to sex expression and mating system: morph I consists mostly of ramets with hermaphroditic flowers and ramets with only female flowers, morph II consists of ramets with hermaphroditic flowers and ramets with hermaphroditic and male flowers, and morph III consists mostly of ramets with hermaphroditic flowers. A microsatellite analysis of seed DNA showed that morph III has a high self-fertilization rate. Flowering season and flower visitor assemblages, which also differ among the three morphs, may influence the evolution and maintenance of the differences in sex expression and mating systems in the morphs.     

Resources and pollinators contribute to population sex-ratio bias and pollen limitation in Fragaria virginiana (Rosaceae)

Populations of gynodioecious species vary in the ratio of female versus hermaphroditic individuals they contain, and many exhibit higher frequencies of females under poor resource conditions. One important factor limiting female frequencies within populations is predicted to be pollen limitation of seed production, caused by either low abundance of pollen donors or insufficient pollen transfer. However, empirical studies measuring variation in pollen limitation with population sex ratios or resource gradients in gynodioecious plants are inconsistent. Part of this inconsistency may be that pollen limitation and its causes are context-dependent. Another possibility is that sex-specific daily flower production and/or sex-biased visitation are more relevant to the likelihood of pollen limitation than sex ratio based on counting individual plants. In this study, we examined context-dependent pollen limitation in gynodioecious/subdioecious Fragaria virginiana . We specifically examined the potential for resource availability to influence sex-specific daily flower production, sex-biased pollinator visitation, and their relationships with pollen limitation in experimental populations that contained either high or low frequencies of female plants. High resource availability reduced apparent female frequency by increasing daily flower production by hermaphrodites relative to females. This is important because pollinators increasingly discriminated against female flowers as floral sex ratios became more female-biased. Contrary to expectation, females in high-female populations were not consistently more pollen limited than those in low-female populations. The level of pollen limitation of females was better explained by sex–biased pollinator foraging and visitation frequency than by the plant sex ratio or floral sex ratio. Thus, negative frequency dependence of female pollen limitation was evident only considering sex ratio bias mediated by pollinator visitation.     

SEX ALLOCATION IN THE MONOECIOUS HERB BEGONIA SEMIOVATA

Sex-allocation models predict that the evolution of self-fertilization should result in a reduced allocation to male function and pollinator attraction in plants. The evolution of sex allocation may be constrained by both functional and genetic factors, however. We studied sex allocation and genetic variation for floral sex ratio and other reproductive traits in a Costa Rica population of the monoecious, highly selfing annual Begonia semiovata. Data on biomass of floral structures, flower sex ratios, and fruit set in the source population were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and genetic correlations for floral sex ratio and for floral traits related to male and female function were estimated from the greenhouse-grown progeny of field-collected maternal families. The proportion of reproductive biomass invested in male function was low (0.34 at flowering, and 0.07 for total reproductive allocation). Significant among-family variation was detected in the size (mass) of individual male and female flowers, in the proportion of male flowers produced, and in the proportion of total flower mass invested in male flowers. Significant among-family variation was also found in flower number per inflorescence, petal length of male and female flowers, and petal number of female flowers. Except for female petal length, we found no difference in the mean value of these characters between selfed and outcrossed progeny, indicating that, with the possible exception of female petal length, the among-family variation detected was not the result of variation among families in the level of inbreeding. Significant positive phenotypic and broad-sense genetic correlations were detected between the mass of individual male and female flowers, between male and female petal length, and between number of male and number of female flowers per inflorescence. The ratio of stamen-to-pistil mass (0.33) was low compared to published data for autogamous species with hermaphroditic flowers, suggesting that highly efficient selfing mechanisms may evolve in monoecious species. Our results indicate that the study population harbors substantial genetic variation for reproductive characters. The positive genetic correlation between investment in male and female flowers may reflect selection for maximum pollination efficiency, because in this self-pollinating species, each female flower requires a neighboring male flower to provide pollen.     

Evidence for distinct roles of the SEPALLATA gene LEAFY HULL STERILE1 in Eleusine indica and Megathyrsus maximus (Poaceae)

LEAFY HULL STERILE1 (LHS1) is an MIKC-type MADS-box gene in the SEPALLATA class. Expression patterns of LHS1 homologs vary among species of grasses, and may be involved in determining palea and lemma morphology, specifying the terminal floret of the spikelet, and sex determination. Here we present LHS1 expression data from Eleusine indica (subfamily Chloridoideae) and Megathyrsus maximus (subfamily Panicoideae) to provide further insights into the hypothesized roles of the gene. E. indica has spikelets with three to eight florets that mature acropetally; E. indica LHS1 (EiLHS1) is expressed in the palea and lemma of all florets. In contrast, M. maximus has spikelets with two florets that mature basipetally; M. maximus LHS1 (MmLHS1) is expressed in the palea and lemma of the distal floret only. These data are consistent with the hypothesis that LHS1 plays a role in determining palea and lemma morphology and specifies the terminal floret of basipetally maturing grass spikelets. However, LHS1 expression does not correlate with floret sex expression; MmLHS1 is restricted to the bisexual distal floret, whereas EiLHS1 is expressed in both sterile and bisexual floret meristems. Phylogenetic analyses reconstruct a complex pattern of LHS1 expression evolution in grasses. LHS1 expression within the gynoecium has apparently been lost twice, once before diversification of a major clade within tribe Paniceae, and once in subfamily Chloridoideae. These data suggest that LHS1 has multiple roles during spikelet development and may have played a role in the diversification of spikelet morphology.