Floral ontogeny inMazus pumilus (Scrophulariaceae)

InMazus pumilus, all the floral appendages are initiated in acropetal sequence in the second cell layer (except stamens) of the floral primordium by periclinal divisions. The actinomorphic calyx tube is formed due to zonal growth. The zygomorphy in corolla is evident from the inception of petal primordia which arise sequentially as independent units in order of one anterior, a pair of anterio-lateral followed by a pair of posterio-lateral. Later these primordia exhibit differential growth because of which zygomorphy becomes more pronounced. The upper corolla tube is formed by interprimordial growth and lower corolla tube by zonal growth. Stamens are initiated in the third layer of the floral apex. Unlike sepals and petals, in the development of stamens (4) underlying cells of corpus also contribute. Posterior stamen is absent. The stamens become epipetalous because of interprimordial and zonal growth in the common region below the bases of petals as well as stamens. The two carpel primordia arise as crescent shaped structures which become continuous due to interprimordial growth. The ovary is formed by a ring of zonal meristem. The style develops later between stigma and ovary because of intercalary growth. The residual apex grows vertically along with the ovary and forms the septum of the ovary. All the floral appendages exhibit similar pattern of histogenesis and early growth suggesting thereby the appendicular nature of these appendages.     

Floral development of Iris decora Wall. (Iridaceae)

The outer tepal and stamen primordia arise as secondary primordia on the outer tepal-stamenprimordia, which are formed on the floral apex. The inner tepal primordia are formed directly on the floral apex. All the floral appendages are initiated in the second tunica layer and are homologous with regard to their origin and early development. A short perianth tube is formed as a result of intercalary growth in the common bases of the tepal primordia. The intercalary growth in the fused bases of the floral appendages elevates the peripheral zone. The floral apex thus appears as a shallow cup. Further intercalary growth results in the formation of an inferior ovary. The ovules are initiated as outgrowths on placental ridges from the lateral ovary wall, the trilocular appearance being the result of secondary cohesion of the parietal placentae.     

HISTOGENESIS OF THE ANDROECIUM AND GYNOECIUM IN DOWNINGIA BACIGALUPII

A histogenetic investigation of the synandrous androecium and syncarpous gynoecium in the flower of Downingia bacigalupii Weiler (Campanulaceae; Lobelioideae) was undertaken for the purpose of comparing the modes of initiation, early growth and fusion in these floral whorls with that reported previously for the perianth in this species. Stamens are initiated as separate organs from the second tunica layer and underlying corpus regions of the concave floral meristem. Subsequent growth of stamens involves apical and intercalary growth in length and rudimentary marginal growth in breadth. Tissues of the four microsporangia originate from hypodermal sporangial initial cells and the filament is formed by intercalary growth at the base of the anther. Lateral fusion of stamens is ontogenetic and involves cuticular fusion of adjacent epidermal layers. The two emergent carpel primordia arise as crescentic organs by periclinal divisions in the second tunica layer and corpus zones. Carpel primordia also undergo apical and intercalary growth in length as well as extensive marginal growth in breadth. Radial growth in carpels is mediated by an adaxial meristem which shows its greatest concentration of activity at the carpel margins. Carpel fusion appears to be partially ontogenetic accompanied by zonal growth. Closure of the stylar canal is by the formation of a transmitting tissue derived from the protodermal layers of the adaxial carpel surfaces. A discoid nectary is initiated around the base of the style and formation of the inferior ovary is by intercalary growth of the base of the concave floral bud. The two parietal placentae originate as longitudinal outgrowths from the walls of the floral cup. Ovule initiation is simultaneous at first and then intercalary during subsequent elongation of the ovary. The ovules are anatropous, unitegmic and tenuinucellate. Stamen and carpel procambium shows a slight delay in differentiation when compared to that reported for the perianth and bract, but in all other respects carpels resemble other floral organs in their patterns of histogenesis and early growth. Stamens diverge from the other floral organs in their early pattern of growth, but a consideration of all features of their histogenesis suggests an appendicular rather than an axial interpretation of these organs.     

Floral development of Justicia gendarussa (Acanthaceae)

The sequence of primordial initiation is acropetal and the primordia develop in the same order in which they appear. The floral apex has a two-layered tunica in all stages of development. There is no significant difference in the initiation of any of the floral appendages and thus all floral organs are homologous with respect to their histogenetic origin. The short calyx tube is formed by ontogenetic fusion of the bases of sepals; but the corolla tube arises partly by ontogenetic union of originally free parts and partly by zonal or intercalary growth. Each primordium receives a single procambium strand shortly after its initiation, except those of the posterior pair of stamens, which do not receive any vascular supply. This provides another example which goes against the doctrine of "conservatism of vascular bundles". The placentation in Justicia is parietal ontogenetically as well as anatomically.     

INFLORESCENCE AND FLORAL DEVELOPMENT IN HOUTTUYNIA CORDATA (SAURURACEAE)

The inflorescence of Houttuynia cordata produces 45–70 sessile bracteate flowers in acropetal succession. The inflorescence apical meristem has a mantle-core configuration and produces “common” or uncommitted primordia, each of which bifurcates to form a floral apex above, a bract primordium below. This pattern of organogenesis is similar to that in another saururaceous plant, Saururus cernuus. Exceptions to this unusual development, however, occur in H. cordata at the beginning of inflorescence activity when four to eight petaloid bract primordia are initiated before the initiation of floral apices in their axils. “Common” primordia also are lacking toward the cessation of inflorescence apical activity in H. cordata when primordia become bracts which may precede the initiation of an axillary floral apex. Many of these last-formed bracts are sterile. The inflorescence terminates with maturation of the meristem as an apical residuum. No terminal flowers or terminal gynoecia were found, although subterminal gynoecia or flowers in subterminal position may overtop the actual apex and obscure it. Individual flowers have a tricarpellate syncarpous gynoecium and three stamens adnate to the carpels; petals and sepals are lacking. The order of succession of organs is: two lateral stamens, median stamen, two lateral carpels, median carpel. The three carpel primordia almost immediately are elevated as part of a gynoecial ring by zonal growth of the receptacle below the attachment of the carpels. The same growth elevates the stamen bases so that they appear adnate to the carpels. The trimerous condition in Houttuynia is the result of paired or solitary initiations rather than trimerous whorls. Symmetry is bilateral and zygomorphic rather than radial. No evidence of spiral arrangement in the flower was found.     

Inflorescence and flower development in the Hedychieae (Zingiberaceae): Hedychium coccineum Smith

The inflorescence of Hedychium coccineum Smith is thyrse, and the primary bracts are initiated in a spiral phyllotactic pattern on the sides of the inflorescence dome. Cincinnus primordia are initiated on the flank of the inflorescence apex, in the axils of primary bracts. This primordium subsequently develops a bract and a floral primordium. Then, the floral primordium enlarges, flattens apically, and becomes rounded. Sepals are initiated sequentially from the rounded corner of the primordium ring sepal initiation, and the floral primordium continues to enlarge and produces a ring primordium. Later, this ring primordium separates three common primordia surrounding a central cavity. The adaxial common primordium is the first separation. This primordium produces the posterior petal and the fertile stamen. The remaining two common primordia separate and produce respectively a petal and a petaloid, the inner androecial member. As the flower enlarges, the cavity of the floral cup becomes a rounded–triangular apex; these apices are the sites of outer androecial primordium initiation. The abaxial outer androecial member slightly forms before the two adaxial members develop. But this primordium ceases growth soon after initiation, while the two posterior primordia continue growth to produce the lateral petaloid staminodes. During this stage, gynoecial initiates in the floral cup and continues to grow until extending beyond the labellum.     

FLORAL DEVELOPMENT IN MYRISTICA (MYRISTICACEAE)

Myristica fragrans and M. malabarica are dioecious. Both staminate and pistillate plants produce axillary flowering structures. Each pistillate flower is solitary, borne terminally on a short, second-order shoot that bears a pair of ephemeral bracts. Each staminate inflorescence similarly produces a terminal flower and, usually, a third-order, racemose axis in the axil of each pair of bracts. Each flower on these indeterminate axes is in the axil of a bract. On the abaxial side immediately below the perianth, each flower has a bracteole, which is produced by the floral apex. Three tepal primordia are initiated on the margins of the floral apex in an acyclic pattern. Subsequent intercalary growth produces a perianth tube. Alternate with the tepals, three anther primordia arise on the margins of a broadened floral apex in an acyclic or helical pattern. Usually two more anther primordia arise adjacent to each of the first three primordia, producing a total of nine primordia. At this stage the floral apex begins to lose its meristematic appearance, but the residuum persists. Intercalary growth below the floral apex produces a columnar receptacle. The anther primordia remain adnate to the receptacle and grow longitudinally as the receptacle elongates. Each primordium develops into an anther with two pairs of septate, elongate microsporangia. In pistillate flowers, a carpel primordium encircles the floral apex eventually producing an ascidiate carpel with a cleft on the oblique apex and upper adaxial wall. The floral ontogeny supports the morphological interpretation of myristicaceous flowers as trimerous with either four-sporangiate anthers or monocarpellate pistils.     

Floral morphogenesis of Caltha and Trollius (Ranunculaceae) and its systematic significance

The floral morphogenesis of Caltha palustris L. and Trollius buddae Schipcz. was observed with a scanning electron microscope (SEM). The primordia of all floral organs initiate spirally and centripetally and develop centripetally. The spiral initiation sequence may be a basic pattern in Ranunculaceae. The primordia of bracts, sepals, and other floral organs are different in shape: the bract primordia are triangle, the sepal primordia crescent, and the petal (in Trollius), stamen, and carpel primordia hemispheric. This may indicate that the bracts, the sepals and other floral organs are different in origin. The petals are retarded in early developmental stages in Trollius buddae Schipcz, and have purses at the base. The retarded petals are very common in Ranunculaceae and the purse of the petal is similar to that of some Aquilegia species. The microspores in a longitudinal series of stamens develop centripetally in Caltha and Trollius; this may be a basic pattern in Ranunculaceae. The carpel primordia are plicate. In the developmental process of the carpels, the stigmatic tissue appears from the apex of the style and is decurrent along the ventral suture in Caltha, but there is no obvious stigmatic tissue in Trollius. Based on floral morphogenesis characteristics as well as the results from molecular systematics, comparative morphology and palynology studies, we consider that Caltha is not closely related to Trollius and that these two genera should not be treated in the same tribe.     

驴蹄草属和金莲花属(毛茛科)花器官的形态发生及系统学意义

利用扫描电镜观察了驴蹄草Caltha palustris L.和川陕金莲花Trollius buddae Schipcz.花器官的发生和发育过程。结果显示:驴蹄草和川陕金莲花的所有花器官均螺旋状向心式发生、向心式发育,花器官的螺旋状发生方式在毛茛科Ranunculaceae可能是一种基本式样;苞片、萼片与其他花器官原基的形状明显不同,显示苞片、萼片与其他花器官在系统发生上有所不同;川陕金莲花的花瓣在早期延迟发育且基部具囊,花瓣的延迟发育在毛茛科具花瓣的属中非常普遍,而花瓣基部的囊类似于耧斗菜属Aquilegia一些植物;两个属雄蕊群一纵列雄蕊中的小孢子均向心式发育,这种发育方式在毛茛科可能为基本类型。两个属植物的心皮原基均为对折式,在发育过程中,驴蹄草心皮顶端沿腹缝线形成下延的柱头组织,川陕金莲花不形成明显的柱头组织。根据花形态发生和发育特点,并结合其他研究成果,认为这两个属不应当属于同一个族。     

Inflorescence and Flower Development of Globba barthei ( Zingiberaceae)

Inflorescence of Globba barthei is a thyrse . Primary bracts are initiated in a spiral phyllotactic pattern on the inflorescence apex . Cincinnus primordia are initiated in the axils of primary bracts . These promordia develop secondarybracts and floral primordia . The floral primordium continues to enlarge and produce a ring primordium . Sepals are initiated sequentially from the rounded corner of the primordium . The ring primordium separates three common primordium surrounding a central cavity . The adaxial common primordium is the first to separate . This primordium divides transversely and producespetal and fertile stamen . The remaining two common primordium transversely separate and produce respectively a petal and a petaloid . As the flower developing , the cavity of the floral cup becomes triangular . The angles of this triangle are the sites of outer androecial primordium . The abaxial androecia forms slightly earlier than the two adaxial ones, and then this primordium ceases growth soon . The two posterior primordia continue growth to produce the lateral petaloid staminodes . During this stage , gynoecia initiate from the floral cup and continue to fuse and develop into style and stigma. In addition ,Initiation of the bulbil primordium is observed at base of inflorescence axis during the early floral development . The bulbil primordium initiates in the axil of primary bract . The evolutionary significance of six androecia is discussed .     

毛舞花姜花器官的发生与发育

通过扫描电镜观察了毛舞花姜(Globba barthei Gagne p.)的花序及花器官的发生与发育。3枚萼片原基首先于花顶连续发生,随后花顶的中心凹陷形成环状原基,环状原基进一步分化形成三枚花瓣—雄蕊共同原基,并在花顶的中心形成花杯。共同原基分化形成花瓣和三枚内轮雄蕊,紧接着外轮雄蕊在花杯的顶点发生。远轴的两枚内轮雄蕊延伸生长并相互融合形成了唇瓣,近轴的一枚形成了可育雄蕊;近轴的两枚外轮雄蕊发育形成了成熟花结构中的侧生退化雄蕊,而远轴的一枚缺失。近轴的两枚外轮雄蕊原基起始的同时,3枚心皮原基也在中心花杯的内侧发生而后与外轮雄蕊相间排列。对毛舞花姜花序的发生和发育的观察发现,在花序轴的头几片初级苞片中产生的是珠芽原基而非蝎尾状小花序原基,其形态特征类似于早期的蝎尾状小花序原基,由此推测珠芽很可能是蝎尾状小花序的变异。     

Floral development in Nymphaea tetragona (Nymphaeaceae)

We describe in detail the floral ontogeny of Nymphaea tetragona from a wild population to provide evidence regarding the phylogenetic position of Nymphaea and to reveal evolutionary trends of flowers in Nymphaeaceae by comparison with that of the other genera. Four sepals are initiated unidirectionally. The basal petals are initiated unidirectionally and alternate with the sepals. The dome‐shaped floral apex continues to expand and produces more petal and stamen primordia. The remaining petals and all stamens are initiated in spirals or whorls. Later, the periphery of the floral apex grows more quickly than the centre and results in a depression in the centre of the apex after all stamens have been initiated. Carpels are simultaneously initiated in a cycle at the periphery of the depression. They are ascidiate. After all organs have been initiated, the centre of the depression on the floral apex grows and develops into a globular structure. The connected inferior ovary, stigma caps and the globular floral apex together form an extragynoecial compitum. Within Nymphaeaceae, the floral ontogeny of Nymphaea is most similar to that of Euryale and Victoria. It differs more from Ondinea and Barclaya, and differs most from Nuphar. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 211–221.     

FLORAL DEVELOPMENT IN MANGROVE RHIZOPHORACEAE

The flowers of mangrove Rhizophoraceae (tribe Rhizophoreae) are adapted to three different pollination mechanisms. Floral development of representative species of all four genera suggests that the ancestral flower of the tribe was unspecialized, with successively initiated whorls of separate sepals, petals, antisepalous stamens, and antipetalous stamens; at its inception, the gynoecium had a united, half-inferior ovary and separate stigmatic lobes. This developmental pattern is found in Rhizophora mangle (wind-pollinated) and Ceriops decandra (insect-pollinated). In Kandelia, all floral organs distal to the sepals are initiated simultaneously, and there has apparently been an evolutionary amplification in the number of stamens to about six times the number of petals. Explosive pollen release evolved independently in C. tagal and in Bruguiera. In the former, all stamens belong to one whorl and arise simultaneously upon a very weakly differentiated androecial ring primordium. In Bruguiera, the androecial ring is pronounced, and two whorls of stamens arise upon it; the primordia of the antisepalous whorl arise first but are closer to the center of the apex than the antipetalous stamen primordia. The antisepalous stamens bend toward and are enclosed by the petals early in development. In all genera, the inferior ovary develops by zonal growth of receptacular tissue; additional intercalary growth above the placenta occurs in Bruguiera. In general, floral specialization is accompanied by an increase in the width of the floral apex compared to the size of the primordia, increasing fusion of the stylar primordia, and decreasing prominence of the superior portion of the ovary. Apparent specializations of petal appendages for water storage, including the presence of sub-terminal hydathodes (previously unreported in any angiosperm), were found in two species in which flowers remain open during the day but were absent from two species normally pollinated at night or at dawn. Distinctive tribal characteristics that may aid in phylogenetic analysis include the mode of development of the inferior ovary; the aristate, bifid, usually fringed petals that individually enclose one or more stamens; the intrastaminal floral disc; and the initially subepidermal laticiferous cell layer in the sepals and ovary.     

ORGAN INITIATION AND DEVELOPMENT OF INFLORESCENCES AND FLOWERS OF ACACIA BAILEYANA

Inflorescence and floral ontogeny are described in the mimosoid Acacia baileyana F. Muell., using scanning electron microscopy and light microscopy. The panicle includes first-order and second-order inflorescences. The first-order inflorescence meristem produces first-order bracts in acropetal order; these bracts each subtend a second-order inflorescence meristem, commonly called a head. Each second-order inflorescence meristem initiates an acropetally sequential series of second-order bracts. After all bracts are formed, their subtended floral meristems are initiated synchronously. The sepals and petals of the radially symmetrical flowers are arranged in alternating pentamerous whorls. There are 30–40 stamens and a unicarpellate gynoecium. In most flowers, the sepals are initiated helically, with the first-formed sepal varying in position. Petal primordia are initiated simultaneously, alternate to the sepals. Three to five individual stamen primordia are initiated in each of five altemipetalous sectorial clusters. Additional stamen primordia are initiated between adjacent clusters, followed by other stamens initiated basipetally as well as centripetally. The apical configuration shifts from a tunica-corpus cellular arrangement before organogenesis to a mantle-core arrangement at sepal initiation. All floral organs are initiated by periclinal divisions of the subsurface mantle cells. The receptacle expands radially by numerous anticlinal divisions in the mantle at the summit, concurrently with proliferation of stamen primordia. The carpel primordium develops in terminal position by conversion of the floral apex.     

A SCANNING ELECTRON MICROSCOPIC STUDY ON THE INITIATION AND DEVELOPMENT OF FLORAL ORGANS OF BRASSICA NAPUS (CV. WESTAR)

The initiation and development of the floral organs of Brassica napus L. (cv. Westar) were examined using the scanning electron microscope. After transition of the vegetative apex into an inflorescence apex, flower primordia were initiated in a helical phyllotactic pattern. The sequence of initiation of the floral organs in a flower bud was that of sepals, stamens, petals and gynoecium. Of the four sepal primordia, the abaxial was initiated first, followed by the two lateral and finally the adaxial primordium. The four long stamens were initiated simultaneously in positions alternating with the sepals. The two short stamens were initiated basipetal to and outside the long stamens, and opposite the lateral sepals. The petals arose on either side of the two short stamens and the gynoecium was produced from the remainder of the apex. During development, the sepal primordia curved sharply at the tips and tightly enclosed the other organs. Stamen primordia developed tetralobed anthers at an early stage while filament elongation occurred just prior to anthesis. A unique pattern of bulbous cells was present on the abaxial surface of the anther. Growth of petal primordia lagged relative to the other floral organs but expansion was rapid prior to anthesis. The gynoecium primordium was characterized by an invagination early in development. At maturity, there was differentiation of a papillate stigma, an elongated style and a long ovary marked externally by sutures and divided internally by a septum. Distinct patterns of cuticular thickenings were observed on the abaxial and adaxial surfaces of the petals and stamens and on the surface of the style. The patterns were less obvious on the sepals and ovary. Stomata were present on both surfaces of the mature sepals, on the style and restricted areas on the abaxial surface of the anthers and nectaries but were absent from the petals, the adaxial surface of the stamens and the ovary. No hairs were present on any of the floral organs.     

弯齿盾果草(紫草科)花序及花的形态发生

以弯齿盾果草不同发育时期的花芽为材料,在体视显微镜解剖观察的基础上使用扫描电镜对弯齿盾果草花序、花及果实的发育过程进行了观察。结果显示:(1)弯齿盾果草的花序是由最初的一个球形花序原基经过多次分裂形成的,且花序发生式样符合蝎尾状聚伞花序结构,而非通常所描述的镰状或螺状聚伞花序;花序发生过程中无单一主轴,花序轴是由侧枝连接而成,每一朵花原基有其对应的1枚苞片,下一花原基是从相邻的上一枚苞腋里发生,相邻两花原基交错互生。(2)花器官的发生是按照花萼原基、花冠原基、雄蕊原基和雌蕊原基的顺序发育,但雄蕊原基的花药部分发育速度要比花冠原基快,所以花器官的发育是按照花萼、雄蕊、花冠和雌蕊的顺序发育。(3)子房四深裂结构是由4个原基分别发育,而后相互靠拢而成。(4)小坚果表面的附属结构发生于子房发育后期,其背面的内外层突起分别是由生长较快的外部组织的边缘通过上部内缩和下部向外环状生长形成。     

The Growth of the Shoot Apex and the Apical Dome of Barley During Ear Initiation

The growth of the floral main shoot apex of spring barley wasstudied during the period of ear initiation (that is, from initiationof the collar primordium until maximum primordium number wasattained). While floral primordia were being initiated the relativelength growth rate of the shoot apex was low. After maximumprimordium number there was about a twofold increase in relativelength growth rate. Estimates of the volume, fresh and dry weightof the floral apex indicated that the relative weight growthrate was also low at first and increased after maximum primordiumnumber. The rates of growth and the size at initiation of thefloral primordia was affected by their position on the floralshoot apex. The relative volume growth rate increased acropetallyfrom the first initiated (collar) primordium. The collar wasthe smallest and each subsequently-initiated primordium increasedin length. The diameter of the newly-initiated primordium alsoincreased until more than half the primordia had been initiatedand then it declined. The apical dome increased in both lengthand diameter and both were at a maximum at the time of the double-ridgestage and then both measurements declined. Length and diameterwere at a minimum at maximum primordium number. Subsequentlythere was an increase in the length of the dome, after whichboth the dome and some of the last formed, distal primordiadied. The period of spikelet initiation therefore is a stage duringwhich the relative growth rate of the floral shoot apex is low,there are changes in the size of the dome and the primordiashow a progression of increasing relative growth rates acropetallyalong the shoot apex. These changes produce the embryo ear inwhich the most advanced spikelets are in the lower mid-partof the ear. Changes in size of the apical dome prior to maximumprimordium number may be related to the subsequent death ofspikelet primordia and therefore also to grain number in themature ear.     

RE-EVALUATION OF CERTAIN KEY RELATIONSHIPS IN THE ALISMATIDAE: FLORAL ORGANOGENESIS OF SCHEUCHZERIA PALUSTRIS (SCHEUCHZERIACEAE)

Transition to flowering in the North-temperate bog plant Scheuchzeria palustris occurs in early May and results in the formation of a simple raceme with six flowers. Five of the flowers are subtended by large foliar bracts, while the sixth and last-formed flower on the inflorescence remains ebracteate. The individual flowers develop along a clearly trimerous pattern. The three outer tepals develop first, arising almost simultaneously at the periphery of the triangular floral apex. They are followed closely by the development of the three anti-tepalous outer stamens. The three inner tepals are next in the developmental sequence, alternating with the outer whorl of tepal-stamen pairs but arising at a slightly higher level on the floral meristem. Three inner stamens are initiated opposite the inner tepal primordia. Finally, three gynoecial primordia are initiated on the remaining central portion of the floral apex and alternating with the inner whorl of tepal-stamen pairs. Each carpel develops at first as a horseshoe-shaped structure. Two ovules form in each carpel, initiating on the adaxial margin of the carpel wall. Histogenesis of all floral appendages involves initially periclinal divisions in the second tunica layer followed by corresponding anticlinal divisions in the first tunica layer and concurrent activity in the underlying corpus. Separate procambial strands differentiate acropetally from the inflorescence axis to each tepal-stamen pair and then bifurcate. The vascular connection to the gynoecium develops directly from the strands in the tepal-stamen pairs. The results of this developmental study of the flower of S. palustris have a significant bearing on the positioning of this and related taxa within the Alismatidae and on the speculation of the phylogeny of the monocotyledon flower.     

Floral development in Astragalus caspicus Bieb. (Leguminosae: Papilionoideae: Galegeae)

Floral organogenesis and development of the bushy perennial legume Astragalus caspicus were studied using epi-illumination light microscopy techniques. Based on our observations, flowers are in axillary two-flowered racemes, initiate all 21 floral organs and show precocious appearance of zygomorphy. The order of floral organ initiation is unidirectional in whorls starting from the abaxial position of the flower with a high degree of overlap. Another important ontogenetic feature is the existence of two successive common primordial stages categorized as primary and secondary. The primary common primordia produce antesepalous stamens and secondary common primordia. In contrast, the five secondary common primordia subdivide into a petal and an antepetalous stamen primordia. Our findings on floral ontogeny of A. caspicus provide new evidence for the complex and variable floral initiation and development in legumes. The floral apex with strong overlapping initiation of different organs illustrates a paradox in which different capabilities must be presumed to exist simultaneously. Moreover, two extraordinary types of common primordia represent possibly an advanced evolutionary trend where time intervals between the initiations of different floral organs in Papilionoideae are shortened.