FLORAL ANATOMY IN THE SAXIFRAGACEAE SENSU LATO. II. SAXIFRAGOIDEAE AND ITEOIDEAE

The floral anatomy and morphology of 26 species from the Saxifragoideae and three from the Iteoideae are described and compared. The flowers of the Saxifragoideae are predominantly actinomorphic, partially epigynous and/or perigynous, and pentamerous, with two carpels which bear numerous ovules. There is usually some degree of independence between carpels, and the normally separate styles possess both a canal and transmitting tissue. Generally, staminodia are absent and nectariferous tissue, which is not vascularized, is present. The subfamily is characterized by large multicellular trichomes with globular, often glandular, heads. Placentation may be parietal, axile, or transitional between the two; parietal appears to be a derived condition in the subfamily. The vascular cylinder in the pedicel generally consists of several to many discrete bundles from which diverge ten compound traces at the base of the receptacle, leaving an inner cylinder of vascular strands that coalesce at a higher level into either as many ventral bundles as carpels or twice that number. In the former case, each ventral bundle consists of one-half of the vascular supply to each adjacent carpel and separates into individual ventral strands in the distal half of the ovary. The ventral bundles provide vascular traces to the ovules and, along with the dorsals, extend up the style to the stigma. Each trace diverging in a sepal plane typically supplies one or more carpel-wall bundles, a median sepal bundle, and a stamen bundle. Each petal-plane trace usually provides one or more carpel-wall bundles, a lateral trace to each adjacent sepal, a petal bundle and, in flowers with ten stamens, a stamen bundle. Dorsal carpel bundles are usually recognizable and may originate from traces in either perianth plane. While the position of Ribes remains problematical, its floral structure does not easily exclude it from the Saxifragoideae. Floral structure in the Iteoideae is remarkably similar to that in the Saxifragoideae, the main differences being a lesser degree of independence between carpels, generally narrower placentae with somewhat fewer ovules, and the presence of only unicellular, acutely pointed epidermal hairs as opposed to the relatively complex, multicellular trichomes prevalent in the Saxifragoideae.     

THE PALM GYNOECIUM

The morphology, anatomy, and histology of the gynoecia at or close to anthesis are described for 20 genera of palms selected to represent different taxonomic alliances and to include major gynoecial types within the family. Palms may have 1–10 carpels, but most have three. Fifteen genera, including 14 coryphoid palms and the monotypic Nypa fruticans, are apocarpous and the remainder, approximately 190, are syncarpous. Fusion of carpels in some gynoecia begins in the base, in others in the styles. Pseudomonomerous pistils occur in several different alliances: the ovarian parts of two carpels are reduced but three usually equal and functional styles and stigmas are present. The carpel is often follicular in shape with the ventral suture open or, more frequently, partially or completely closed. The carpel may be stipitate or sessile and usually has a conduplicate laminar part. Most carpels are spirally and laterally inserted on the receptacle, but the carpel in some unicarpellate genera (e.g., Thrinax) appears terminal. Stipes, ovarian parts, styles, and stigmas vary in structure and development. Septal nectaries which differ in size, in the presence or absence of specialized canals, and in position, characterize all genera of some groups but only some genera of others. Diverse vascular configurations in the bases of gynoecia vary according to the extent of the floral axis, the development of carpellary stipes, and the connation of the carpels and their adnation to the tip of the floral axis. Four types of carpellary vascular systems are present in the genera described: (1) most palm carpels have three major traces consisting of a dorsal bundle and two ventral bundles, and they may also have up to four pairs of lateral bundles or occasionally more; (2) in certain cocosoid palms no ventral bundles can be distinguished, but a dorsal bundle, many parallel lateral bundles, and a row of immature ventral strands vascularize each carpel; (3) carpels of Phytelephas have a dorsal bundle, two pairs of major lateral bundles and about four pairs of shorter lateral bundles, with no identifiable ventral bundles; (4) carpels of Nypa have many dichotomously branched bundles but none that are recognizable as dorsal, ventral, or lateral strands. Additional peripheral bundles or systems may be present in each of the above types. Ovules are supplied by 1–15 bundles. These are derived either from the carpellary stele; from ventral bundles only; from ventral, lateral, and dorsal bundles; or from a combination of these origins. Certain areas of the gynoecia or certain parts of dorsal carpellary walls in some genera are much less mature at anthesis than surrounding tissues. Implications for floral biology and relationships within the palms and of palms to other groups are discussed.     

STUDIES ON THE FLORAL ANATOMY OF THE CUNONIACEAE

Twenty-two genera representing sixty-two species of Cunoniaceae and Davidsonia were examined with respect to floral anatomy. Sepals are vascularized by three traces with the lateral traces of adjacent sepals united. Pancheria is unique for the family with species in which the sepals are vascularized by a single, undivided bundle. Petals, when present, and stamens, are uniformly one-trace structures. A general tendency exists within the family for the principal floral bundles to unite in various ways, with fusions evident between calyx, corolla, and androecial vascular supplies. Carpel number ranges from two to five and the gynoecium is generally surrounded by a prominent disc. Gynoecia of Ceratopetalum and Pullea are “half-inferior.” The number of ovules per carpel locule ranges from one to numerous. Ventral carpel sutures range from open to completely sealed at the level of placentation. Carpels of the apocarpous genus Spiraeanthemum (incl. Acsmithia) are vascularized by a dorsal bundle and either three or four bundles constituting the ovular and wing vasculation in the ventral position, a condition unlike other members of the family. Ovules are supplied by the median ventral bundle. More advanced bicarpellate gynoecia within the family are predominately vascularized by a dorsal and two ventral bundles although a variable number of additional lateral wall traces may be present. A major trend exists toward fusion of the ventral bundles of adjacent carpels in the ovary of both bicarpellate and multicarpellate plants. At the base of the styles the fused ventral strands separate and extend along with the dorsal carpellary bundles into styles of adjacent carpels. In Pullea the ventral bundles terminate within the ovules. The united ventral carpellary bundles in Aphanopetalum, Gillbeea, and Aistopetalum lie in the plane of the septa separating adjacent carpels. Ovules are vascularized by traces originating from the vascular cylinder at the base of the gynoecium or by traces branching from the ventral bundles. Ovular traces in each carpel are united, or remain as discrete bundles, prior to entering the placenta. Tannin and druses are common throughout all floral parts. Although floral anatomy generally supports the position of Cunoniaceae near Saxifragaceae and Davidsoniaceae, the evolutionary relationship of the Cunoniaceae to the Dilleniaceae is uncertain.     

Comparative anatomy and systematics of woody Saxifragaceae. Aphanopetalum Endl.

The floral and vegetative anatomy of the small Australian genus Aphanopetalum were studied. Wood is described for the first time and is characterized by predominantly solitary pores, scalariform vessel element perforation plates with low bar numbers, imperforate tracheary elements with distinctly bordered pits, sparse axial parenchyma, and a combination of homocellular and heterocellular rayS. Starch occurs in both axial and ray parenchyma of the wood. Stems possess unilacunar, one-trace nodes and the uncommon feature of an endodermis with well-defined Casparian stripS. Leaves have anomocytic stomata, a bifacial mesophyll and semicraspedodromous venation or a combination of semicraspedodromous and brochidodromous venation. The tetramerous flowers are apetalous or have minute petals. The compound, half-inferior gynoecium consists of essentially totally united carpels. The pattern of floral vascularization resembles different Saxifragaceae sensu lalo in that the compound sepal-plane and petal-plane traces give rise to staman bundles as well as sepal, petal, and carpel wall venation in their respective planes. The ventral ovarian bundles are fused into a single ventral complex that subdivides at the top of the ovary to form ventral bundles and to supply the one ovule in each locule. Vegetative and floral features provide compelling evidence to suggest that Aphanopetalum has its nearest relatives among the Saxifragaceae sensu lato rather than Cunoniaceae. The genus is probably best treated as forming its own subfamily (or family) among the saxifragaean alliance.     

Comparative floral structure and systematics in Celastrales (Celastraceae, Parnassiaceae, Lepidobotryaceae)

Floral morphology, anatomy and histology in the newly circumscribed order Celastrales, comprising Celastraceae, Parnassiaceae and Lepidobotryaceae are studied comparatively. Several genera of Celastraceae and Lepidobotrys (Lepidobotryaceae) were studied for the first time in this respect. Celastraceae are well supported as a group by floral structure (including genera that were in separate families in earlier classifications); they have dorsally bulged‐up locules (and thus apical septa) and contain oxalate druses in their floral tissues. The group of Celastraceae and Parnassiaceae is also well supported. They share completely syncarpous gynoecia with commissural stigmatic lobes (and strong concomitant development of the commissural vascular bundles but weak median carpel bundles), only weakly crassinucellar or incompletely tenuinucellar ovules with an endothelium, partly fringed sepals and petals, protandry in bisexual flowers combined with herkogamy by the movement of stamens and anther abscission, and stamens fused with the ovary. In contrast, Lepidobotryaceae are more distant from the other two families, sharing only a handful of features with Celastraceae (not Parnassiaceae), such as pseudohermaphroditic flowers, united stamen bases forming a collar around the gynoecium and seeds with a conspicuous aril. However, all three families together are also somewhat supported as a group and share petals that are not retarded in late floral bud development, 3‐carpellate gynoecia, ventral slits of carpels closed by long interlocking epidermal cells and pollen tube transmitting tissue encompassing several cell layers, both integuments usually more than two cell layers thick, and only weak or lacking floral indumentum. In some molecular analyses Celastrales form an unsupported clade with Malpighiales and Oxalidales. This association is supported by floral structure, especially between Celastrales and Malpighiales. Among Celastrales, Lepidobotryaceae especially share special features with Malpighiales, including a diplostemonous androecium with ten fertile stamens, epitropous ovules with an obturator and strong vascularization around the chalaza. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149 , 129–194.     

Floral anatomy and development of <Emphasis Type="Italic">Saxofridericia aculeata</Emphasis> (Rapateaceae) and its taxonomic and phylogenetic significance

Floral anatomy and development of Saxofridericia aculeata Körn was studied in a comparative approach to contribute to the understanding of the family. Flowers at different developmental stages were analysed with light and scanning electron microscopy, and the nature of the exudate secreted by the floral trichomes was investigated by histochemical tests. The anatomical characteristics observed in S. aculeata flowers were compared with those from other Rapateaceae species by a cluster analysis (UPGMA). The dendrogram generated reflects the groupings that emerged in phylogenetic molecular analyses, highlighting the usefulness of floral anatomy for taxonomy and for the understanding of infrafamilial relationships. The exudate secreted by the trichomes has a polysaccharidic composition. Such trichomes (colleters) occur in the sepals, petals, filaments and around the gynoecium; they are initiated at mid-stage of floral development and are an apomorphy of the family. The flowers are pentacyclic, presenting three initially free sepals, petals, stamens and carpels that mature in a centripetal order. The connate portion of the corolla, which is also adnate to the stamens, has a late development by zonal growth. Gynoecium formation is a combination of postgenital and congenital fusion processes. Data on floral organogenesis of Rapateaceae are first reported here and support the early diverging position of the family in Poales, close to Bromeliaceae.     

澜沧荛花(瑞香科)的花部形态发生 及其系统学意义

通过扫描电镜对澜沧荛花Wikstroemia delavayi花部的形态发生过程进行了观察和分析,旨在为该属的系统学研究提供花部发育形态学资料。澜沧尧花花部的发生和早期发育呈远轴面向近轴面的顺序,但这一式样由于近轴面的器官在早期发育之后生长加速发生了转变。因此,花开放时所表现的所谓辐射对称,显然是由同一轮器官的异率生长所导致的次生现象。花盘发生于花萼筒基部的远轴面上,与花萼、雄蕊的发生间隔时间较长。花盘原基在下轮雄蕊着生处凹陷或间断,与之相对应,花盘裂片与下轮雄蕊呈互生。由此,花盘显然不是花托的一部分,也不是象花萼、雄蕊和心皮一样的独立结构,将其解释为雄蕊群的一部分更合理。花盘的发生和早期发育及其着生位置同其他花部器官的发生和发育式样具有明显的相关性,这种相关性对进一步阐明瑞香属Daphne和荛花属Wikstroemia的系统发育关系具有—定意义。根据对雌蕊群的发生和发育过程观察,该种的子房是由一个近轴面的可育心皮和一个远轴面的不育心皮融合而成的单室子房,为假单心皮雌蕊。尽管荛花属和瑞香属均属于单室产房,但澜沧荛花的子房维管束中的腹束排列于中轴位置,而目前资料显示瑞香属植物的腹束接近于侧膜位置,这方面仍需进一步研究。     

Floral morphogenesis of Wikstroemia delavayi (Thymelaeaceae) and its phylogenetic implication

Floral morphogenesis of Wikstroemia delavayi Lecomte was investigated by scanning electron microscope (SEM) and compared with its allied groups. Initiation and early development of floral parts in W. delavayi followed unidirectional sequences from the abaxial side to the adaxial side. Because the floral parts grew faster at the adaxial side than at the abaxial one in following development, the zygomorphic pattern in the early development changed and finally became an almost actinomorphic form at anthesis. The disc was initiated from the abaxial base of the floral tube and itslobes were alternate with lower whorl stamens. According to this initiatial and developmental pattern, it is reasonable to interpret the disc as a part of the androecium rather than a modification of the receptacle. The located position and development of the disc was correlative with the development of other floral organs, which might provide insight to delimit Wikstroemia and Daphne based on different floral developmental pattern that might exist between the two genera. The developmental process of W. delavayi indicated that the syncarpous and uniloculate gynoecium was in fact bicarpellate, which consisted of a fertile carpel and a sterile one. It was pseudomonomerous. Even though the ovary in both Wikstroemia and Daphne was uniloculate, the location of the ventral bundles in the ovary was obviously different from each other according to data to date. In this respect, further investigation is undertaken between the two genera.     

澜沧荛花(瑞香科)的花部形态发生yh及其系统学意义(英文)

通过扫描电镜对澜沧荛花Wikstroemiadelavayi花部的形态发生过程进行了观察和分析 ,旨在为该属的系统学研究提供花部发育形态学资料。澜沧荛花花部的发生和早期发育呈远轴面向近轴面的顺序 ,但这一式样由于近轴面的器官在早期发育之后生长加速发生了转变。因此 ,花开放时所表现的所谓辐射对称 ,显然是由同一轮器官的异率生长所导致的次生现象。花盘发生于花萼筒基部的远轴面上 ,与花萼、雄蕊的发生间隔时间较长。花盘原基在下轮雄蕊着生处凹陷或间断 ,与之相对应 ,花盘裂片与下轮雄蕊呈互生。由此 ,花盘显然不是花托的一部分 ,也不是象花萼、雄蕊和心皮一样的独立结构 ,将其解释为雄蕊群的一部分更合理。花盘的发生和早期发育及其着生位置同其他花部器官的发生和发育式样具有明显的相关性 ,这种相关性对进一步阐明瑞香属Daphne和荛花属Wikstroemia的系统发育关系具有一定意义。根据对雌蕊群的发生和发育过程观察 ,该种的子房是由一个近轴面的可育心皮和一个远轴面的不育心皮融合而成的单室子房 ,为假单心皮雌蕊。尽管荛花属和瑞香属均属于单室子房 ,但澜沧荛花的子房维管束中的腹束排列于中轴位置 ,而目前资料显示瑞香属植物的腹束接近于侧膜位置 ,这方面仍需进一步研究     

Taxonomic studies ofAsarum sensu lato

The floral vascular anatomy of 12 species representing each ofAsarum s. str.,Asiasarum, Geotaenium, Heterotropa andHexastylis are compared to clarify intergeneric relationships. The five genera have basically similar structures in floral morphology and vasculature, and consistently have a six-carpelled compound ovary and the associated similar placental vasculature. They show, however, a significant difference in the position and the constituent of the “ventral” carpellary bundles in the placental axis betweenAsiasarum-Heterotropa-Hexastylis andAsarum-Geotaenium. InAsiasarum, Heterotropa andHexastylis the ventral bundles of each carpel are basically free and antilocular as expected in the least specialized compound ovary of angiosperms; in contrast, inAsarum andGeotaenium the ventral carpellary bundles are antiseptal and heterogenous (i.e., formed by the lateral fusion of ventral bundles of adjacent carpels). Shared probable apomorphic floral vasculature, as well as shared single style-column, suggests the closest mutual relationships betweenAsarum andGeotaenium. In terms of floral morphology and anatomy,Asiasarum, Heterotropa andHexastylis retain plesiomorphies. Possible chromosomal evolution in the related genera is also discussed.     

大钟花属和黄秦艽属花部解剖

对大钟花属和黄秦艽属进行了花部解剖学研究,并以此讨论了它们的系统演化关系。大钟花属和黄秦艽属的雌花部分花萼维管束与花冠维管束来源于同一维管束迹,而雄蕊维管束来源于雄蕊迹,每心皮具1条背维管束2条腹维管束,因此,花被维管束为融合型;黄秦艽属的雄花每个花萼、花瓣和雄蕊的维管束均来源于单个维管束迹,每心皮具1条背维管束2条腹维管束,属于基本型。从花部解剖结构看出,大钟花属与假龙胆演化支关系较近;黄秦艽属较獐牙菜属进化。     

FLORAL ANATOMY IN THE SAXIFRAGACEAE SENSU LATO. I. INTRODUCTION,PARNASSIOIDEAE AND BREXIOIDEAE

The floral morphology and anatomy of one representative of the Parnassioideae and two of the Brexioideae are described, and some of the recent literature dealing with the Saxifragaceae sensu lato is reviewed. Comparison of the floral structure in Parnassia to that typical of the Saxifragoideae, the subfamily constituting the Saxifragaceae sensu stricto and which, therefore, may be considered to show the basic saxifragaceous characteristics, reveals little similarity. Parnassia differs in pattern of both sepal and androecial vascularization, vascularization and degree of connation of the carpels, height in the gynoecium to which ventral bundles remain compound, possession of nectariferous staminodia, and the absence of epidermal appendages. Brexia and Ixerba (both of the Brexioideae) are strikingly dissimilar in floral structure and probably should be dissociated. While the position of Ixerba is problematical, it shares more floral characters with the Escallonioideae than with either Brexia or the Saxifragoideae and is better associated with that taxon. In both Parnassia and Brexia the vascular pattern suggests derivation of the androecium from a fascicled condition: the vascular supply of each filament consists of a cylinder of closely associated collateral bundles, and each staminodial set receives a single vascular complex which subsequently divides into as many vascular strands as there are staminodia in the set.     

Anatomy of the floral nectaries of some neotropical Salacioideae (Celastraceae)

Floral nectaries have contributed to the systematics of different taxonomic groups. Since those of the neotropical genera included in subfamily Salacioideae—Cheiloclinium Miers, Peritassa Miers, Salacia L. and Tontelea Aubl.—have different forms and positions, we explored their anatomy to delimit more precisely the genera of subfamily Salacioideae. Buds and open flowers of six species were treated following the usual techniques in plant anatomy. The obtained data were helpful in characterizing the floral nectary anatomy of the studied species. Furthermore, some features such as form, position and surface of nectaries; form of their epidermal cells; presence and distribution of stomata; occurrence of idioblasts containing druses in the nectariferous parenchyma; and absence of nectary vascularization can contribute to the taxonomy and phylogeny of the Salacioideae studied. In most of the studied species the nectar is probably released by both the stomata and the nectary epidermal surface. In Cheiloclinium cognatum, the structure acknowledged as nectary is actually a vestigial tissue and the functions of attracting and rewarding pollinators has phylogenetically migrated to the stigmatic region. The druses and phenolic substances observed in the nectariferous parenchyma probably help defend flowers against herbivore attacks. The minute size of the nectaries of Salacioideae may explain the absence of vascularization. The floral nectaries of Salacia elliptica are epithelial while those of the other species are mesenchymal.     

FLORAL ONTOGENY IN CYCLAMEN PERSICUM ‘F-1 ROSEMUNDE ROSE’ (PRIMULACEAE)

Floral ontogeny was examined in Cyclamen persicum ‘F-1 Rosemunde Rose’ using a combination of light and scanning electron microscopy. The leaf plastochron index (LPI), earlier calculated for leaf elongation, was used to determine the length of each stage of floral development. LPI will provide a useful tool for selecting flowers of a given stage from large plant populations or from plants where flowers are small or inaccessible during early ontogenetic stages. Most features of floral development are similar to those previously described for other primulaceous genera. The petal-stamen relationship, however, is unusual; stamens arise through periclinal cell divisions in the adaxial surface layers of common petal-stamen primordia. Anatomical evidence suggests that the placenta is formed both by appendicular initials which give rise to the ovules and ventral carpellary bundles, and receptacular cells which form some, if not all, of the central axis.     

马蹄香属营养器官解剖及其分类位置的探讨

对马蹄香属Ｓａｒｕｍａ Ｏｌｉｖ．营养器官的形态解剖进行了研究,首次报道该属植物叶片宏观结构;叶表皮和叶表面的微观结构;叶柄和茎的初生结构特征与次生结构特征,并与近缘属植物细辛属Ａｓａｒｕｍ Ｌ．的解剖学资料作了比较研究,发现两属的叶形均为心形;叶脉都是掌状脉序;叶沿无齿都具毛;且都是单毛;气孔器都是“毛茛科”型;叶柄维管束都呈“Ｖ”字形排列;从近轴端到远轴端都是呈３－４－４束的变化;茎的初生结构     

Floral anatomy of Bromeliaceae, with particular reference to the evolution of epigyny and septal nectaries in commelinid monocots

Floral anatomy is described in ten genera of Bromeliaceae, including three members of subfamily Bromelioideae, three Tillandsioideae, and four genera of the polyphyletic subfamily Pitcairnioideae (including Brocchinia, the putatively basal genus of Bromeliaceae). Bromeliaceae are probably unique in the order Poales in possessing septal nectaries and epigynous or semi-epigynous flowers. Evidence presented here from floral ontogeny, vasculature, and the relative positions of nectary and ovules indicates that there could have been one or more reversals to apparent hypogyny in Bromeliaceae, although this hypothesis requires a better-resolved phylogeny. Such evolutionary reversals probably evolved in response to specialist pollinators, and in conjunction with other aspects of floral morphology of Bromeliaceae, such as the petal appendages of some species. The ovary is initiated in an inferior position even in semi-epigynous or hypogynous species. The ovary of all so-called hypogynous Bromeliaceae is actually semi-inferior, because the septal nectary is infralocular; in these species the nectaries have a labyrinthine surface and many vascular bundles. Brocchinia differs from most other fully epigynous species in that each carpel is secretory at the apex and reproductive, rather than secretory, at the base.     

Floral anatomy and morphology in thePolygalaceae

Floral morphology and anatomy of 15 genera in thePolygalaceae have been studied. The pentamerous origin of the polygalaceous flower is confirmed and shown to apply to all genera in the family. The keel is interpreted as a single petal, and the androecium as of bimeric origin. Vascular structure in the receptacles ofCarpolobia andMonnina subg.Monnina is described in detail, and a compilation of results, focusing on the vascular supply for the androecium and gynoecium, is given for all genera. Based on similarities and differences in vascularization it is concluded that present taxonomy, in particular the tribal system, needs to be reviewed.     

FLORAL MORPHOLOGY OF HORSFIELDIA (MYRISTICACEAE)

The pistillate flowers of Horsfieldia are morphologically similar to those of Myristica and Knema, and are composed of a single whorl of thick, fleshy tepals, and an unsealed, monocarpellate pistil bearing a single ovule. The carpel is vascularized by two ventral bundles, a pair of dorsal bundles, and several supernumerary bundles. The ovule vascularization is derived from the supernumerary bundles. Paired dorsal vascular bundles are an uncommon feature of uncertain significance. Carpels of Myristica and Knema lack any clearly defined dorsal vasculature, and the ovule vascular supply is derived from both the ventral and supernumerary bundles. The organization of the staminate flowers of Horsfieldia agrees with the myristicaceous pattern observed in Myristica and Knema. Each androecium consists of a single whorl of anthers fused or partially fused to a massive connective column. Each anther consists of a pair of bisporangiate lobes and a single vascular bundle. The androecial forms observed are interpreted as forming a series of intermediates between the monadelphous type of androecia of two South American genera, Compsoneura and Dialyanthera, and one African genus, Brochneura, and the solid, columnar androecia which are predominate in the family. Accumulating evidence supports a proposed South American or west Gondwanaland origin of the Myristicaceae.     

Anatomy and histochemistry of the nectaries of Rodriguezia venusta (Lindl.) Rchb. f. (Orchidaceae)

Orchidaceae is one of the largest angiosperm families. Although extensively studied, reports of anatomy of secretory structures of orchids are relatively scarce. Rodriguezia venusta is an epiphytic orchid occurring in Brazil and Peru that has floral and extrafloral nectaries. This study describes the structure and the histochemistry of these secretory structures. Floral and extrafloral nectary samples were obtained from R. venusta plants that were collected in a gallery forest in the State of Bahia, Brazil, and grown in a greenhouse. Theses samples were fixed and processed according to routine procedures in plant anatomy and histochemistry or for scanning electron microscopy. The extrafloral nectaries occur on the edge and sub-edge of young leaves and at the basal portion of bracts that subtend the floral buds. They are structurally very similar, being formed by a nectary parenchyma and a simple epidermis with stomata (“non-structured nectaries”). The floral nectary is inserted at the floral receptacle fused with the labellum base, between this structure and the two inferior connate sepals. This nectary consists of an epidermis with numerous specific nectar secreting trichomes, a subnectary and a nectary parenchyma abundantly supplied by vascular terminations. Its structure is complex and distinct from other floral nectaries described for Orchidaceae.     

THE DELIMITATION OF BIGNONIACEAE AND SCROPHULARIACEAE BASED ON FLORAL ANATOMY,AND THE PLACEMENT OF PROBLEM GENERA

The delimitation of Bignoniaceae and Scrophulariaceae has long been a taxonomic problem. Several genera, including Paulownia, Schlegelia, Gibsoniothamnus, and Synapsis, have been variously placed in one or the other family. Differences between these two families have been noted with regard to the presence of endosperm, embryo and seed morphology, and placentation; however, the lack of comprehensive data on the distribution of such characters within these two families left the delimitation problem unsolved. A comprehensive study of floral anatomy confirmed a basic difference in the placentation of these two families, as well as a basic difference in gynoecial vascularization. Paulownia has a floral anatomy, embryo morphology, and seed morphology consistent with placement in Scrophulariaceae. While reminiscent of Bignoniaceae, Paulownia is not an intermediate genus linking the two families. Schlegelia and Gibsoniothamnus have a floral anatomy consistent with placement in Scrophulariaceae. Schlegelia also has a scrophulariaceous seed morphology. Considered anomalous in the Bignoniaceae, the Schlegelieae similarly are distinct in the Scrophulariaceae.