Floral development in Tribe Detarieae (Leguminosae: Caesalpinioideae): Amherstia, Brownea, and Tamarindus

Floral development was compared among three taxa in caesalpinioid tribe Detarieae sensu lato: Amherstia nobilis and Tamarindus indica have racemose, helically arranged inflorescences, while Brownea latifolia has cauliflorous capitate flower clusters that arise as racemes. All have acropetal flower order; initiation and development are sequential in all except Brownea, which is synchronous. All have paired persistent showy bracteoles. Floral symmetry is dorsiventral (zygomorphic) in all except Brownea, with radial symmetry at anthesis. Sepals initiate helically on a circular floral apex, starting with a median abaxial sepal, in all. Petals are initiated helically in Brownea, and unidirectionally in Amherstia and Tamarindus. Stamens are initiated unidirectionally in each stamen whorl in all except Amherstia, in which the outer whorl is bidirectional. The carpel initiates concurrently with the petals in Brownea, and with the outer stamens in the other taxa. The two upper (adaxial) sepal primordia become fused during development in all, so that the calyx appears tetramerous. Some reduced petals occur in Amherstia and Tamarindus, and some reduced stamens occur in all. All produce a hypanthium by zonal growth, and all except Tamarindus have the gynoecium attached adaxially to the hypanthial rim.     

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 in ficinia and isolepis (cyperaceae), with focus on the nature and origin of the gynophore

BACKGROUND AND AIMS: The generic delimitations of Ficinia and Isolepis, sister genera in the Cypereae, are blurred. Typical Ficinia flowers have a lobed gynophore, which envelops the base of the nutlet, whereas in Isolepis the character is considered to be absent. Some former species of Isolepis, lacking the gynophore, were recently included in Ficinia. The floral ontogeny of representative taxa in Ficinia and Isolepis were investigated with the aim of evaluating the origin and nature of the gynophore in the Cypereae. METHODS: The spikelet and floral ontogeny in inflorescences collected in the field was investigated using scanning electron microscopy (SEM) and light microscopy (LM). KEY RESULTS: SEM images of Isolepis setacea and I. antarctica, Ficinia brevifolia, F. minutiflora, F. zeyheri and F. gracilis, and LM sections of F. radiata, show that the gynoecium in Ficinia is elevated above the flower receptacle by the development of a hypogynous stalk. From its apex, a (often three-)lobed cup is formed, which envelopes the basal part of the later nutlet. In developing flowers of I. antarctica, a rudimentary hypogynous stalk appears. In I. setacea, rudiments of a hypogynous stalk can be observed at maturity. In F. radiata and F. zeyheri, intralocular hairs are present in the micropylar zone. At the surface of developing gynoecia in flowers of F. gracilis, star-shaped cuticular structures appear which disappear again at maturity. CONCLUSIONS: The overall floral ontogeny of all species studied occurs following a typical scirpoid pattern, though no perianth primordia are formed. The gynophore in Ficinia originates as a hypogynous stalk, from which the typical gynophore lobes develop. The gynophore is not homologous with the perianth.     

Chaotic Floral Phyllotaxis and Reduced Perianth in Achlys (Berberidaceae)

Among the 16 genera of the Berberidaceae Achlys is the only one with a reduced perianth, an irregular floral phyllotaxis, and variable stamen number. Early floral stages show an unstable (chaotic) arrangement of the organ primordia. Only the single carpel of the gynoecium has a more fixed position in that the placenta is formed in the adaxial half of the flower. The irregularities in the androecium may be caused by the lack of influence of a perianth on floral symmetry. On the other hand, the regular orientation of the carpel is perhaps due to the early polarity of the flower, whereby the abaxial half of the flower is larger (with further developed stamen primordia) at the time when carpel polarity is established.     

The morphology of styles and stigmas inVicia (Leguminosae), and its systematic implications

The morphology and development of styles and stigmas ofVicia, one of the five genera of tribe Vicieae, were studied by scanning electron microscopy to contribute to a revision of infrageneric classification. Among the 22 species examined, variations were found as to: (1) whether a boundary between stigma and style is V-shaped when viewed from the adaxial side or not; (2) whether stylar hairs sprout on all sides of the style or only on the abaxial side; (3) whether a hairy area of the style connects with the stigma area first at lateral sides or at an adaxial side; (4) whether styles are tapered or bottle-neck-shaped; (5) whether styles are straight or bend abaxially. Based on these variations, five morphological types of styles and stigmas were provided with a diagnostic key. Comparisons with related tribes Cicereae, Coronilleae, Galegeae, Hedysareae, Loteae and Trifolieae as possible outgroups suggested that the V-shaped boundary between stigma and style is a synapomorphy of some species withinVicia, and that a dorsally compressed and abaxially tufted style, which was distributed in separate subgenera and sections sensu Kupicha but is now known to occur together with the V-shaped boundary, has evolved only one time in the genus, rather than more than once as supposed earlier.     

马先蒿属植物花冠分化与繁殖适应的研究进展

结合已有的研究报道和作者近年来的工作,对马先蒿属（Pedicularis）植物的花冠多样化成因与繁殖适应特性进行了总结和探讨。通过对该属4种进化花冠型的花器官发生和分化的研究发现,花部各器官在发生和发育初期基本一致,后期上唇形态的分化是导致成熟花形态结构产生较大差异的重要阶段。孢粉学研究认为,花冠类型与花粉萌发孔类型之间具有显著相关性;萌发沟的演化可能与繁殖适应有一定的关系。分子系统学研究表明,多样化的花冠类型在不同的谱系内经过若干次的独立进化而表现出了高度的平行演化（parallelism）。传粉生物学研究证实,该属植物花冠多样化与其主要传粉者熊蜂属（Bombus）昆虫的传粉行为存在较为密切的关系。具有相同（似）花冠类型的马先蒿可能被同种或不同种的熊蜂以相同的方式访问,但在花粉落置位置上存在显著差异,这可能有助于同域分布重叠的物种间在生殖上的机械隔离,而花冠的分化在一定程度上促进了新的物种形成。     

Towards unlocking the deep nodes of Leguminosae: Floral development and morphology of the enigmatic Duparquetia orchidacea (Leguminosae, Caesalpinioideae)

Duparquetia orchidacea (Caesalpinioideae-Cassieae-Duparquetiinae) is a monotypic liana from tropical West Africa. Its highly unusual, zygomorphic flowers, the unique pollen morphology, and the lack of vestured pits in the wood correspond with previous phylogenetic studies that resolved the position of the species to an isolated position among the early-branching Leguminosae. Here we present a detailed analysis of floral morphology and development to clarify open questions of its floral organization. We provide new data that can be useful in clarifying phylogenetic relationships among early branching Leguminosae and improve our understanding of floral evolution in this large and important plant family. For comparison, we also present developmental data for other Fabales. Our analysis reveals some unusual and in parts unique developmental patterns, such as strict acropetal organ formation, loss and suppression of floral organs, and early petal enlargement. We interpret alternating left-right symmetries in floral development as clues to a spiral organ formation in ancestral taxa. Early asymmetry of the young carpel helps to interpret enantiostyly of other Leguminosae as an example of imprinted shape. Finally, we show that cochlear-descending petal aestivation in Duparquetia and in Papilionoideae is based on different ontogenetic patterns and therefore is most probably nonhomologous.     

Floral morphology and ontogeny in Orchidaceae subtribe Disinae

Floral morphology and ontogeny in Orchidaceae subtribe Disinae. The flower structure and development of 24 species of the orchid subtribe Disinae are described and illustrated by drawings and scanning electron micrographs with special attention being paid to the gynostemium. The morphogenesis of this subtribe is fundamentally similar to that of the closely related tribe Orchideae. This includes the initiation of the auricles on the anther base in a dorsolateral position, and hence their interpretation as being mere appendages of the filament. The keel connecting the petals and the gynostemium plus its protrusion is considered homologous to the inner lateral staminodeS. Presumed vestiges of the adaxial staminodes were detected in one specieS. A peculiarity of the Disinae is that the entire apex of the median carpel develops into the rostellum, whereas its stigmatic portion emerges from the median carpel below the rostellum in later stages. The main diagnostic feature of the group is the reflexed position of the mature anther. However, it is shown here that this anther movement occurs in the later stages and that the initial anther is erect.     

Floral ontogeny in Scirpus, Eriophorum and Dulichium (Cyperaceae), with special reference to the perianth

BACKGROUND AND AIMS: Based on molecular phylogenetic analysis, it has been suggested recently that the Cyperaceae comprises only two subfamilies: the Mapanioideae and the Cyperoideae. In most flowers of the Cyperoideae, the whorl of inner stamens is reduced, resulting in tetracyclic flowers. In the more primitive (scirpoid) genera within the Cyperoideae, the perianth consists of two polysymmetric whorls, whereas the perianth parts in the more derived genera have been subject to modifications and/or reduction. Comparative studies of the many silky hairs of Eriophorum and of the eight bristles of Dulichium have given rise to much discussion about their homology. METHODS: The spikelet and floral ontogeny in freshly collected inflorescences was investigated using scanning electron microscopy. KEY RESULTS: Complete floral ontogenies are presented for Scirpus sylvaticus L., Eriophorum latifolium Hoppe and Dulichium arundinaceum (L.) Britton, with special reference to the perianth. The results in S. sylvaticus confirm the trimerous monocot-like organization of the flower. It is used as a model for floral development in Cyperoideae. In the early developmental stages, the androecium of E. latifolium is surrounded by a massive perigonial primordium, from which the many hair-like bristles originate. Consequently, the stamens develop among the hair primordia, more or less simultaneously. The hairs are arranged in whorls, which develop centripetally. The development of the perianth in D. arundinaceum starts with the formation of three initial perianth primordia opposite the stamens. Subsequently, two more abaxial bristle primordia, alternating with the stamens, originate simultaneously with the appearance of three adaxial bristle primordia in the zone where an adaxial inner perianth primordium is expected. CONCLUSIONS: The floral development in E. latifolium and D. arundinaceum can be considered as variations upon the scirpoid floral ontogenetic theme.     

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.     

Floral structure, development and diversity in Thunbergia (Acanthaceae)

Floral structure and development of 18 species of Thunbergia (Thunbergioideae si , Acanthaceae) were studied comparatively. The flowers of Thunbergia are highly diverse and show a wide range of pollination syndromes. In general they are large and showy. Their pollination apparatus is highly elaborate, floral organs are often synorganized, and floral architecture is complex. In contrast to the high diversity of the anthetic flowers, their bauplan is uniform and their early development shows no major differences, i.e. in all species studied, the calyx arises as a ring primordium, the corolla is 'late sympetalous', and petals and stamens are initiated more or less simultaneously. Some differences are found in further calyx development, where several developmental patterns are present. More significant differences arise only later during development and mainly concern the structures of the calyx, the anthers, the stigma, and corolla aestivation. In the anthetic flowers there are many special characters that are present in all or the majority of the species studied, e.g. the calyx is reduced, the corolla tube is subdivided into two compartments and the anthers lack an endothecium. The present results on development and morphology of the flowers of Thunbergia are compatible with an earlier subdivision of the genus into eight subgenera.     

A revised classification of the tribe Phaseoleae (Leguminosae: Papilionoideae), and its relation to canavanine distribution

The subtribal classification of the tribe Phaseoleae has drifted into disorder since Bentham's (1865) scheme in the Genera Plantarum. A revised classification is here proposed, which for the first time accounts for all of the currently recognized genera. This classification has seven subtribes: Cajaninae, Diocleinae, Kennediinae, Phaseolinae, Glycininae, Ophrestiinae, and Erythrininae, of which the Ophrestiinae is new. A survey for the free amino acid, canavanine, shows that it is most often present in the Diocleinae and Kennediinae and frequently absent in the rest of the tribe.     

Identification and characterization of microsatellite loci in Intsia palembanica (Leguminosae), a valuable tropical timber species

We present 25 polymorphic microsatellite loci for Intsia palembanica, a highly valued timber species in the Indo-Malayan region, Australia and western Pacific islands. Microsatellite loci were tested for polymorphism across a total of 76 individuals from three natural populations from Irian Jaya, Indonesia. The average number of alleles for these microsatellites was 12.1 per locus, ranging from four to 19. The observed heterozygosity within the natural populations ranged from 0.01 to 0.96. The markers will enable us to evaluate the spatial-temporal population genetic structure and gene flow dynamics among populations. They can also be used for tracking and tracing wood from legally logged concessions.     

Analysis of late stage flower development in Primula vulgaris reveals novel differences in cell morphology and temporal aspects of floral heteromorphy

Heterostyly in Primula is characterized by the development of long-styled pin and short-styled thrum flowers, with anthers midway down the corolla tube in pin flowers, and at its mouth in thrum flowers. Other differences include pollen size and stigmatic papillae length. Several linked genes at the S locus control these differences.In this study we have analyzed pin and thrum flowers through the temporal development of heteromorphy.These studies indicate that the S locus linked genes that orchestrate heteromorphic flower development act in coordination, but with different temporal and spatial dynamics. Style length is differentiated by longer style cells in pin than thrum. However, our studies on cell shape and size within the corolla tube show that a different mechanism mediates the dissimilar elevation of anthers between pin and thrum types. These studies have also revealed that upper corolla tube cells in thrum flowers are wider than those in pin flowers. This results in a larger corolla tube mouth in thrum flowers and represents a new and previously undocumented heteromorphic variation between pin and thrum flowers.     

Floral ontogeny of the Afro-Madagascan genus Mitrasacmopsis with comments on the development of superior ovaries in Rubiaceae

BACKGROUND AND AIMS: Members of Rubiaceae are generally characterized by an inferior ovary. However, Mitrasacmopsis is cited in the literature as having a semi-inferior to superior ovary. It has previously been hypothesized that the gynoecial development of Rubiaceae with semi-inferior to superior ovaries takes place in the same way as in Gaertnera, one of the most commonly cited rubiaceous genera with a superior ovary. To test this hypothesis, a floral ontogenetic study of Mitrasacmopsis was carried out with special attention paid to the gynoecial development. METHODS: Floral ontogeny and anatomy of Mitrasacmopsis were examined using scanning electron and light microscopy. KEY RESULTS: At an early developmental stage, a concavity becomes visible in the centre of the floral apex simultaneously with the perianth initiation. A ring primordium surrounding this concavity expands vertically forming the corolla tube (early sympetaly). Stamen primordia develop inside the corolla. From the bicarpellate gynoecium only two carpel tips are visible because the ovary is formed by a gynoecial hypanthium. In the basal part of each carpel, a placenta primordium is initiated. Two septa divide the ovary into two locules. In each locule, the placenta becomes mushroom shaped and distinctly stalked. Eventually, the inferior ovary of Mitrasacmopsis develops into a beaked capsule. Only very late in the fruiting stage, the continuously expanding ovary is raised above the insertion point of the persistent calyx, changing the ovary position of Mitrasacmopsis from basically inferior to secondarily semi-inferior. CONCLUSIONS: Mitrasacmopsis follows an epigynous pattern of floral development. However, the presence of a prominent beak in the fruiting stage gives the whole ovary a semi-inferior appearance. This kind of secondarily semi-inferior ovary is shown to be different from the secondarily superior ovary observed in Gaertnera.