Reevaluation of the perianth and androecium in Caryophyllales: implications for flower evolution

The Caryophyllales have the highest diversity in androecial patterns among flowering plants with stamen numbers ranging from 1 up to 4,000. Thanks to the recent progress in reconstructing the phylogeny of core Caryophyllales, questions of floral evolution, such as the origin and diversification of the androecium, can be readdressed. Caryophyllales are unique among core eudicots in sharing an androecial ring meristem or platform with centrifugal development of stamens and petals. Stamens are basically arranged in two whorls and evolution within the clade depends on the shift of either the antesepalous or the alternisepalous whorls to an upper position on the ring meristem and the reduction of the other. Four main developmental phenomena are responsible for the high diversity in androecial patterns: (1) the sterilisation of the outermost stamens through a division of common primordia; (2) the secondary addition of stamens by a centrifugal initiation of supernumerary stamens superimposed on a lower stamen number; (3) the pairwise displacement of alternisepalous stamens to the middle of the outer sepals and their potential fusion, or as part of a pluristaminate androecium; (4) the inversed sequence, reduction and loss of antesepalous stamens. Shifts in stamen numbers depend on pressures of the calyx and carpels and changes in the number of the latter. These patterns are expressed differently in the three main evolutionary lines of core Caryophyllales and are systematically relevant: (1) A basal grade of Caryophyllales, culminating with Caryophyllaceae, Amaranthaceae, Stegnosperma and Limeum, has the antesepalous stamens initiated in upper position on the ring meristem, and alternisepalous stamens are preferentially reduced. Among the antesepalous whorl there is a progressive loss of stamens following a sequence inversed to sepal initiation. Petaloid staminodes are formed by the radial division of outer stamens. (2) The raphide-clade and Molluginaceae are characterized by alternisepalous stamens in upper position on the ring meristem, with a trend to secondary stamen multiplication, and loss of antesepalous stamens. (3) The Portulacineae share the pattern of the raphide clade, but some taxa show shifts to an upper position on the ring meristem of either antesepalous or alternisepalous stamens, linked with secondary multiplications and reduction of either whorl. Different floral characters are plotted on a recent cladogram of Caryophyllales. The data show a consistent correlation between shifting carpel and stamen numbers independent of perianth evolution. Comparative data suggest that the basic androecium of Caryophyllales consists of two whorls of five stamens, linked with an absence of petals, and the evolution of the androecium is a combination of reductions and secondary multiplications of stamens with a highly predictive systematic value.     

MORPHOLOGICAL STUDIES IN ZYGOPHYLLACEAE. I. THE FLORAL DEVELOPMENT AND VASCULAR ANATOMY OF NITRARIA RETUSA

The floral development and vascular anatomy of Nitraria retusa were investigated in order to understand its characteristic androecium of 15 stamens and to clarify the systematic position of the genus relative to Zygophyllaceae. Sepals arise in a helical sequence and are relatively small at maturity. Petals are initiated almost simultaneously or in a rapid helical sequence. Five stamen primordia arise opposite the sepals. Next, two other antesepalous primordia are incepted centrifugally to the first primordia on the remaining receptacular surface. The outer stamens tend to be squeezed between the petals and upper stamens and appear to make up an antepetalous whorl of stamens. Three carpels arise from a low ringwall and grow into a hairy trilocular pistil. In each locule a single pendulous ovule is present. Disclike nectarial tissue is initiated in pits between the stamens and petals. Long trichomes develop on its surface. It is concluded that the androecium is linked with a haplostemonous condition because the stamens of each triplet develop on strictly localized sectors. The distinction between stamens arising on complex primordia and the inception of three independent units is explained by the “principle of variable proportions.” The vasculature also tends to confirm that the outer stamen pairs belong to antesepalous triplets.     

OVERLAPPING ORGAN INITIATION AND COMMON PRIMORDIA IN FLOWERS OF PISUM SATIVUM (LEGUMINOSAE: PAPILIONOIDEAE)

The floral ontogeny of Pisum sativum shows a vertical order of succession of sepals, petals plus carpel, antesepalous stamens, and antepetalous stamens. Within each whorl, unidirectional order is followed among the organs, beginning on the abaxial side of the flower, as in most papilionoids. Unusual features include the four common primordia which precede initiation of discrete petal and antesepalous stamen primordia, and the marked overlap of organ initiations between whorls which are usually separately initiated. The stamens arise in free condition, then become diadelphous by intercalary growth at the base of nine stamens, and finally become pseudomonadelphous by surface fusion between the vexillary stamen filament and the adjacent edges of the filament tube. The early initiation of the carpel is not unique among papilionoids, but is somewhat unusual.     

The floral development of Popowia whitei (Annonaceae)

A study of the floral ontogeny of Popowia was carried out to investigate the phyllotactic arrangement of the floral organs and occurring trends in the androecium of Annonaceae. The flower buds arise on a common stalk in the axil of a bract. Three sepals emerge in quick succession and are rapidly overrun in size by two whorls of petals. The androecium is initiated centripetally in successive whorls. A first whorl of three pairs of outer staminodes emerges opposite the outer petals and is followed by nine staminodes. Next a whorl of nine fertile stamens arises in alternation with the second whorl of staminodes. The carpels arise in three alternating whorls of nine. The nature of the perianth parts is morphologically identical. The process of cyclisation of the androecium from a spiral is discussed for Annonaceae and Magnoliidae in general. The inception of the three outer stamen pairs is a widespread reductive step for multistaminate androecia in the process of oligomerization. It is proposed to define the cyclic inception of numerous stamens as whorled polyandry, being an intermediate step between true polyandry and a reduced stamen number in whorls. The absence of a cup-like shape in the carpel development is related to the flattened receptacle.     

Floral ontogeny of Sinojackia xylocarpa (Styracaceae), with special reference to the development of the androecium

Using scanning electron microscopy, we studied the floral ontogeny of Sinojackia xylocarpa. There are 6–7 (–8) sepals. Sepal initiation is staggered; adaxial sepals arise later than abaxial and lateral ones. There are (5–) 6–7 (–8) petals, initiated simultaneously. Petals alternate with the sepals, and occasionally there are two petals instead of one between two sepals. The (10–) 12–14 (–16) stamens are initiated centripetally in two sets (whorls). These floral organ numbers deviate from those of the otherwise mostly pentamerous family Styracaceae. The ovary consists of three (rarely four) locules. In each of the locules, two rows of ovules are differentiated basipetally. Placentation is axile with (5–) 6 (–7) ovules in each locule. Ovules are unitegmic and are ascending with the micropyle directed downwards. Intra‐ovarian trichomes are present as in other representatives of Styracaceae and seem to be an apomorphic character of the family as they are absent in the closely related Symplocaceae and Diapensiaceae. Various levels of organ union occur in anthetic S. xylocarpa. The calyx is synsepalous and the ovary syncarpous. Possibly, the basal connation of petals and stamens is postgenital (and not congenital), but this needs further study. The outward curvature of the young anthers of the inner stamen whorl superficially simulates an obdiplostemonous androecium. However, the sequence of stamen initiation shows a diplostemonous pattern.     

Trends in evolution of floral ontogeny in Cassia sensu stricto,Senna, and Chamaecrista (Leguminosae: Caesalpinioideae: Cassieae: Cassiinae); a study in convergence

Distinctions in floral ontogeny among three segregate genera (Cassia sensu stricto, Chamaecrista, and Senna) of Cassia L. support their separation. In all species studied, the order of floral organ initiation is: sepals, petals, antesepalous stamens plus carpel, and lastly antepetalous stamens. Sepal initiation is helical in all three genera, which however differ in whether the first sepal is initiated in median abaxial position (Senna), or abaxial and off-median (Cassia javanica), a rare character state among legumes. Order of petal initiation varies: helical in Senna vs. unidirectional in Cassia and Chamaecrista. Both stamen whorls are uniformly unidirectional. Intergeneric ontogenetic differences occur in phyllotaxy, inflorescence architecture, bracteole formation, overlap of initiation among organ whorls (calyx/corolla in Cassia; two stamen whorls in Chamaecrista), eccentric initiation on one side of a flower, anther attachment, anther pore structure, and precocious carpel initiation in Senna. The asymmetric corolla and androecium in Chamaecrista arise by precocious organ initiation on one side (left or right). The poricidal anther character can result from differing developmental pathways: lateral slits vs. sealing of lateral sutures; clasping hairs vs. sutural ridges; terminal pores (one or two) vs. none; and clamp layer formation internally that prevents lateral dehiscence. Genera differ in corolla aestivation patterns and in stigma type. Convergence is shown among the three genera, based on intergeneric dissimilarities in early floral ontogeny (floral position in the inflorescence, bracteole presence, position of the first sepal initiated, order of petal initiation, asymmetric initiation, overlap between whorls, anther morphology, and time of carpel initiation) resulting in similarities at anthesis (showy, mostly yellow salverform flowers, heteromorphic stamens, poricidal anther dehiscence, bee pollination, and chambered stigma).     

Floral ontogeny and anatomy inKoelreuteria with special emphasis on monosymmetry and septal cavities

The floral ontogeny and anatomy ofKoelreuteria paniculata have been investigated to understand the developmental basis for the occurring monosymmetry and the origin of the septal cavities. Petals arise sequentially and one petal is missing between sepals 3 and 5, or rarely between sepals 2 and 5. The eight stamens arise sequentially before petal initiation is completed. The last formed petal and one stamen arise on a common primordium. Two stamen positions are empty (opposite the petal between the sepals 2 and 5, and the petal between sepal 1 and 3); consequently two antesepalous stamens have become displaced. The derivation of octandry from a diplostemonous ancestry, and reduction of the petal are discussed. The triangular gynoecium has a strong impact in obliquely reorganizing the symmetry of the flower, loss of organs, and shifts of stamens. The so-called septal slits occurring within the style are a deepreaching non-nectariferous extension of the stigma. Alternating locular furrows are present which could play a role as pollen transmitting tissue and in the loculicid dehiscence of the capsule.     

FLORAL DEVELOPMENT IN CAESALPINIA (LEGUMINOSAE)

Utilizing scanning electron microscopy, we studied the early floral ontogeny of three species of Caesalpinia (Leguminosae: Caesalpinioideae): C. cassioides, C. pulcherrima, and C. vesicaria. Interspecific differences among the three are minor at early and middle stages of floral development. Members of the calyx, corolla, first stamen whorl, and second stamen whorl appear in acropetal order, except that the carpel is present before appearance of the last three inner stamens. Sepals are formed in generally unidirectional succession, beginning with one on the abaxial side next to the subtending bracts, followed by the two lateral sepals and adaxial sepal, then lastly the other adaxial sepal. In one flower of C. vesicaria, sepals were helically initiated. In the calyx, the first-initiated sepal maintains a size advantage over the other four sepals and eventually becomes cucullate, enveloping the remaining parts of the flower. The cucullate abaxial sepal is found in the majority of species of the genus Caesalpinia. Petals, outer stamens, and inner stamens are formed unidirectionally in each whorl from the abaxial to the adaxial sides of the flower. Abaxial stamens are present before the last petals are visible as mounds on the adaxial side, so that the floral apex is engaged in initiation of different categories of floral organs at the same time.     

太原黄耆(豆科)花器官及胚珠发育研究

太原黄耆是新近发表的物种,分布于中国陕西和山西。该实验利用扫描电子显微镜对太原黄耆的花器官发生和发育过程进行观察研究。结果显示:(1)太原黄耆的各轮花器官都是从远轴端向近轴端单向连续发生,在不同轮之间存在花器官重叠发生的现象。(2)在花的发育过程中,出现2种共同原基,即初级共同原基和次级共同原基,由初级共同原基发育成对萼雄蕊原基和次级共同原基,再由次级共同原基发育成花瓣原基和对瓣雄蕊原基。(3)雄蕊管近轴端基部开口是在进化过程中产生的特殊结构,是一种对传粉者的适应机制,从而有利于传粉活动的进行。(4)胚珠为倒生胚珠,具有2层珠被,认为倒生胚珠是内外2层珠被共同作用的结果。     

NORMAL FLORAL ONTOGENY AND COOL TEMPERATURE-INDUCED ABERRANT FLORAL DEVELOPMENT IN GLYCINE MAX (FABACEAE)

Floral onset in soybean (Glycine max cv. Ransom) is characterized by precocious initiation of axillary meristems in the axils of the most recently initiated leaf primordium. During floral transition, leaf morphology changes from trifoliolate leaf with stipules, to a three-lobed bract, to an unlobed bract. Soybean flowers initiated at 26/22 C day/night temperatures are normal, papilionaceous, and pentamerous. Sepal, petal, and stamen whorls are initiated unidirectionally from the abaxial to adaxial side of the floral apex. The median sepal is located abaxially and the median petal adaxially on the meristem. The organogeny of ‘Ransom’ flowers was found to be: sepals, petals, outer stamens plus carpel, inner stamens; or, sepals, petals, carpel, outer stamens, inner stamens. The outer stamen whorl and the carpel show possible overlap in time of initiation. Equalization of organ size occurs only within the stamen whorls. The sepals retain distinction in size, and the petals exhibit an inverse size to age relationship. The keel petals postgenitally fuse along part of their abaxial margins; their bases, however, remain free. Soybean flowers initiated at cool day/night temperatures of 18/14 C exhibited abnormalities and intermediate organs in all whorls. The gynoecium consisted of one to ten carpels (usually three or four), and carpel connation varied. Fusion of keel petals was often lacking, and stamen filaments fused erratically. Multiple carpellate flowers developed into multiple pods that were separate or variously connate. Intermediate type organs had characteristics only of organs in adjacent whorls. These aberrant flowers demonstrate that the floral meristem of soybean is not fixed or limited in its developmental capabilities and that it has the potential to produce alternate morphological patterns.     

Floral ontogeny of Ruteae (Rutaceae) and its systematic implications

Floral development was investigated in Ruta graveolens and Psilopeganum sinense, representing two genera in the tribe Ruteae. Special attention was paid to the sequence of initiation of organ whorls in the androecium and gynoecium. The antepetalous stamens arise at the same level as the antesepalous stamens in both species. The carpels are antepetalous in both taxa, indicating the androecium in both genera is obdiplostemonous. Compared with floral ontogeny of the ancestral genus Phellodendron (Toddalioideae), the obdiplostemonous androecium is a derived condition. The floral apex in P. sinense is quadrangular before initiation of the two carpels. Additionally, there are four dorsal and four ventral traces in the ovary. Integrated morphological and anatomical evidence indicates that the bicarpellate gynoecium in Psilopeganum most likely evolved from a tetracarpellate ancestor. Considering the similarities in morphological, geographical and chromosomal features, the ancestor may be Ruta‐like. Further molecular phylogenetic and genetic studies are needed to verify this assumption.     

Notes on the Evolution of Androecial Organisation in the Magnoliophytina (Angiosperms)

This paper aims to summarize briefly and to update our ideas about androecial architecture formulated in earlier publications. Ontogenetic evidence of stamen development, viz. the initiation, arrangement and relationship of stamens to other floral morphomes, can be translated into a semophyletic scheme reflecting the phylogeny of the androecium. The ancestral androecium is discussed in the light of recent theoriesabout angiosperm phylogeny. Two divergent androecial processes are proposed for the angiosperms starting from a spiral androecium with a moderate number of stamens. However, transitions exist between spiral polyandry, numerous stamens in whorls, and chaotic polyandry. From an androecium with several alternating whorls of paired and single stamens, outer stamen pairs are retained following the successive loss of inner stamen whorls. Single stamens instead of pairs occur at the very end of this line and represent a more advanced condition. This line is mostly present in tri- and dimerous flowers. From the same starting point diplostemony (with two alternating whorls of single stamens) originated, again giving rise to various states usually present in pentamerous or tetramerous flowers.     

Early development of androecia in polystemonous Hydrangeaceae

Polystemonous androecia are diverse in both number and position of stamens. This investigation of polystemonous Hydrangeaceae uses developmental data to characterize (1) the range of developmental variations that account for the diverse androecial patterns and (2) how the expressions of polystemony among Hydrangeaceae compare to those found generally among other angiosperms and especially in their sister family, the Loasaceae, some of which have particularly complex androecia. All polystemonous Hydrangeaceae share the common element of stamen clusters in antesepalous positions. In each of these taxa, the first stamens are initiated opposite the medians of the sepals. Subsequently, stamens form laterally on the flanks of the initial antesepalous stamens, giving rise to the clusters designated as antesepalous triplets. The simplest elaborations based on those common initial developmental steps include (1) adding additional lateral flanking stamens and (2) adding a single stamen in each antepetalous position between adjacent antesepalous groups. More complex elaborations are characteristic of (1) Carpenteria and Philadelphus, which form common primordia at the beginning of androecial development and, subsequently, have stamen primordia form on them, and (2) Deinanthe, which has an elongate hypanthial region on which numerous whorls of stamens are initiated. Carpenteria is unique among Hydrangeaceae in having groups of stamens that are initiated centrifugally in antepetalous positions, and this is similar to complex elements found among some Loasaceae. Generally, the polystemony of Hydrangeaceae that is based in the formation of antesepalous triplets is very similar to that found to evolve in parallel among various clades of rosids and asterids.     

Floral ontogeny in<Emphasis Type="Italic"> Lespedeza thunbergii</Emphasis> (Leguminosae: Papilionoideae: Desmodieae): variations from the unidirectional mode of organ formation

Floral ontogeny of Lespedeza thunbergii was studied with the use of scanning electron microscopy (SEM). The ontogeny varies in all whorls from the undirectional mode, which has been long held to be the rule in Leguminosae. In the sepal whorl, the lateral and the adaxial sepals are formed simultaneously, which is interpreted as a tendency towards whorled organ formation. Whorled organ formation is shown in the petal whorl. The antesepalous stamen whorl varies least from the unidirectional mode. Here, the adaxial stamens are formed successively. This is seen as a remnant of an original helical organ formation in Papilionoideae. Within the antepetalous stamen whorl, the two abaxial stamens and the adaxial stamen are formed first, followed by the two lateral stamens. This is a rarely found phenomenon, which is hard to interpret at the present state of knowledge. Concerning the mature flower, it is shown that nectar stomata are found in a distinct area on the adaxial side of the flower. The presented new characteristics should be an initial step toward further work on taxa of the tribe Desmodieae. These studies will broaden the data set and enable a detailed phylogenetic analysis.     

Morphological studies in Zygophyllaceae. II. The floral development and vascular anatomy of Peganum harmala

Floral development and vascular anatomy are investigated in Peganum harmala, emphasizing its unusual androccium with 15 stamens. Sepals arise successively; petals emerge simultaneously with five antesepalous stamens. The five stamen pairs arise in the space between the petals and the antesepalous stamens. The gynoecium arises from three carpel primordia with evidence of two reduced carpels. Placentae are axile and each bears two double rows of ovules. A weakly developed nectary surrounds the base of the ovary. The antepetalous stamen traces diverge from a common supply to petals and sepal laterals, independent of the antesepalous stamen traces. The androecium of Peganum is described as a derived obdiploste-monous form, differing from the complex haplostemonous androecium of Nitraria. “Congenital dédoublement” cannot adequately explain the origin of the paired antepetalous stamens; two stamens can arise either by the splitting of a common primordium or independently, and both ways of inception are best understood as extremes of a gradation. The systematic position of Peganum is discussed in relation to other Zygophyllaceae using a cladistic analysis with Ptelea (Rutaceae) and Quassia (Simaroubaceae) as outgroups. The basal division in the Zygophyllaceae is between Peganum and the rest of the family.     

大戟科麻疯树属三种植物花器官发生

利用扫描电子显微镜观察了大戟科Euphorbiaceae麻疯树属Jatropha麻疯树J. curcas L.、佛肚树J. podagrica Hook.和棉叶麻疯树J. gossypifolia L.花器官发生。结果表明: 麻疯树、佛肚树和棉叶麻疯树花萼原基均为2/5型螺旋发生。在同一个种不同的花蕾中, 花萼的发生有两种顺序: 逆时针方向和顺时针方向。远轴面非正中位的1枚先发生。5枚花瓣原基几乎同时发生。雄花中雄蕊两轮, 外轮对瓣, 内轮对萼。研究的3种麻疯树属植物雄蕊发生方式有两种类型: 麻疯树亚属麻疯树的5枚外轮雄蕊先同时发生, 5枚内轮雄蕊后同时发生, 佛肚树亚属佛肚树和棉叶麻疯树雄蕊8-9枚, 排成两轮, 内外轮雄蕊同时发生。雌花的3枚心皮原基为同时发生。麻疯树属单性花, 雌花的子房膨大而雄蕊退化, 雄花的雄蕊正常发育, 子房缺失。根据雄蕊发生方式, 支持将麻疯树属分为麻疯树亚属subgen. Jatropha和佛肚树亚属subgen. Curcas。     

Quantitative genetics of stamen number in the selfing Scleranthus annuus (Caryophyllaceae)

Heritability of stamen fertility—different scores were given to sterile stamens developed to different degrees as well as to fertile stamens with one or two pollen sacs—was studied in Scleranthus annuus (Caryophyllaceae), a selling annual that shows extensive phenotypic variation in stamen fertility. Variation within and among 172 maternal families, derived from plants representing 20 natural populations from southern Sweden, was used to estimate heritabilities of stamen fertility for stamens/staminoids at each of the ten stamen positions in the flower. The hierarchical design of the study allowed partitioning of variation at four levels of organization using nested analysis of variance. Heritabilities ranged from 0.631 to 0.714 for stamen positions in the outer whorl of stamens and from 0.235 to 0.555 for positions in the inner whorl. When stamen fertility was pooled across all stamen positions of a flower, the heritability was 0.807. The nested ANOVA indicated that stamen positions in the outer whorl have comparatively higher proportions of among-family and among-population variation than those in the inner whorl. Furthermore, highly significant genetic correlations exist among stamen positions within the inner whorl and among positions within the outer whorl, but not so between positions from each of the two whorls.     

Flower development in Abrus precatorius (Leguminosae: Papilionoideae: Abreae) and a review of androecial characters in Papilionoideae

The jequirity bean (Abrus precatorius) is well known because of its shiny black and red coloured seeds and because of the poison (abrin) it contains. The genus Abrus is placed in a monogeneric tribe Abreae which is placed in a relatively isolated systematic position at the base of Millettieae. To contribute to a better understanding of this taxon, a detailed ontogenetic and morphologic analysis of its flowers is presented. Floral primordia are subtended by an abaxial bract and preceded by two lateral bracteoles which are formed in short succession. Sepal formation is unidirectional starting abaxially. All petals are formed simultaneously. The carpel is formed concomitantly with the outer (antesepalous) stamen whorl, which arises unidirectionally, starting in an abaxial position. In the inner, antepetalous stamen whorl two abaxial stamens are formed first, followed by two lateral stamen primordia. The adaxial, antepetalous position remains organ free (i.e. this stamen is lost). Later in development the nine stamen filaments fuse to form an adaxially open sheath. The filament bases of the two adaxial outer-whorl stamens grow inwards, possibly to provide stability and to compensate for the lost stamen. In the mature flower a basal outgrowth can be found in the position of the lost stamen. However this is more likely to be an outgrowth of the filament sheath rather than a remnant of the lost stamen. These ontogenetic patterns match in parts those found in other Millettieae (unidirectional formation of sepals and stamens, simultaneous petal formation). In contrast, the complete loss of a stamen is rather unusual and supports the isolated position of Abreae and probably justifies (among other characters) its tribal status. A review of androecial characters shows that androecial merosity is on the one hand extremely variable among Leguminosae, varying from a single stamen per flower to more than 500. On the other hand it is noteworthy that the number of stamens becomes stabilised in more derived Papilionoideae such as the large non-protein-amino-acid-accumulating clade (NPAAA clade). This indicates that the androecium has played an important role in the success of a major part of Leguminosae.     

Developmental study on the inflorescence and flower of Caldesia grandis Samuel (Alismataceae)

The inflorescence and floral development of Caldesia grandis Samuel is reported for the first time in this paper. The basic units of the large cymo‐thyrsus inflorescence are short panicles that are arranged in a pseudowhorl. Each panicle gives rise spirally to three bract primordia also arranged in a pseudowhorl. The branch primordia arise at the axils of the bracts. Each panicle produces spirally three bract primordia with triradiate symmetry (or in a pseudowhorl) and three floral primordia in the axils of the bract primordia. The apex of the panicle becomes a terminal floral primordium after the initiations of lateral bract primordia and floral primordia. Three sepal primordia are initiated approximately in a single whorl from the floral primordium. Three petal primordia are initiated alternate to the sepal primordia, but their subsequent development is much delayed. The first six stamen primordia are initiated as three pairs in a single whorl and each pair appears to be antipetalous as in other genera of the Alismataceae. The stamen primordia of the second whorl are initiated trimerously and opposite to the petals. Usually, 9–12 stamens are initiated in a flower. There is successive transition between the initiation of stamen and carpel primordia. The six first‐initiated carpel primordia rise simultaneously in a whorl and alternate with the trimerous stamens, but the succeeding ones are initiated in irregular spirals, and there are 15–21 carpels developed in a flower. Petals begin to enlarge and expand when anthers of stamens have differentiated microsporangia. Such features do not occur in C. parnassifolia. In the latter, six stamen primordia are initiated in two whorls of three, carpel primordia are initiated in 1–3 whorls, and there is no delay in the development of petals. C. grandis is thus considered more primitive and C. parnassifolia more derived. C. grandis shares more similarities in features of floral development with Alsma, Echinodorus, Luronium and Sagittaria. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140 , 39–47.