Comparative morphology of the carpel in the Liliaceae: Baeometra, Burchardia and Walleria

The pistils in Baeometra, Burchardia and Walleria ate tricarpellate, and their ovules are mostly bitegmic. Baeometra has free styles and deep septal invaginations between the carpels. Its pistil is innervated by three dorsal bundles, three compound septal bundles (each of which may divide into two simple septal bundles above), six placental bundles, and six adjoining auxiliary placental bundles. The pistil of Burchardia resembles that of Baeometra , except that there are six simple septal bundles throughout and no auxiliary placental bundles. In Walleria the wings of adjoining carpels are completely fused (except for rare septal glands); there is a single compound style; additional vascular tissue is present in the central axis of the pistil up to the lowermost ovules; the carpels are fused with the floral cup above the base of the locules; and raphide idioblasts are present. Walleria has six "ventral" bundles, each of which appears to be the fusion product of a placental bundle with a simple septal bundle. Tribal affinities of these genera are discussed.     

Comparative morphology of the carpel in the Liliaceae: Wurmbaeae

The two genera of Buxbaum's tribe Wurmbaeae, Anguillaria and Wurmbea , have multiovulate carpels. There are deep septal indentations between the carpels of Anguillaria , but the wings of adjoining carpels are fused to solid septa in most species of Wurmbea. In Anguillaria the carpels have open sutures or prominent commissural markings; in Wurmbea the carpels generally lack these characteristics, and some species have a vascularized, columella-like axis in the centre of the pistil. In both genera there are a dorsal bundle, lateral bundles, and two placental bundles in each carpel. At the inner edge of the septum there are one or two septal bundles in Anguillaria and one or none in Wurmbea. The ovules are monotegmic, the integument and funiculus being partly fused in Anguillaria and mostly fused in Wurmbea. An obturator is present in Anguillaria but absent from most species of Wurmbea.     

Comparative morphology of the carpel in the Liliaceae: Veratreae

The genera of the Veratreae, a tribe of the Melanthioideae, have many features in common: there are usually many ovules, except for Amianthium (with 2 4), arranged in 2 -4 longitudinal placental rows per carpel; all are bitegmic, basipetal, and campylotropous. Of 37 species examined, only 2 have open sutures at the lowermost level of ovular insertion, but 13 species have holes in the centre of the pistil. These holes may represent possible stages in the evolutionary closure of previously open sutures. Most flowers were epigynous, only 11 being hypogynous-perigynous. The tribe as a whole is marked by the presence of 3 composite (heterologous) vascular bundles, composed of joined staminal and tepallary bundles alone and 3 composite bundles, as above, fused to a dorsal bundle. The bundles were united below the locular base in all genera except Schoenocaulon and Toxicoscordion. Two major kinds of central cylinder arrangement occurred at the level of the lowermost ovular insertion: either 6 inverted ventral bundles or 6 simple septal bundles, with normally arranged (or sometimes inverted) xylem and phloem centrifugally located and 6 simple placental bundles, with inverted xylem and phloem, at the centripetal end of the septum. Generally each septal bundle united with its nearest adjoining placental bundle about the mid-locular level.     

Comparative morphology of the carpel in the Liliaceae: Colchiceae (Androcymbium)

The pistil of Androcymbium closely resembles that of Colchicum : it is tricarpellate usually, syncarpous and multiovulate, and the carpels of most species have open sutures and bitegmic ovules. The only species with closed carpellary sutures, A. dregei has monotegmic ovules. There are always three dorsal bundles and three compound septal bundles, which latter may bifurcate into simple septal bundles. Six placental bundles (two per carpel) are differentiated, either separately from the compound septal bundles or as lateral branches of them. A statistical evaluation of 47 species (6 genera) of the hemisyncarpous Wurmbaeoideae shows a significant tendency for bitegmic ovules and two simple septal bundles per septum to be associated with open sutures and for monotegmic ovules and no septal bundles to be associated with closed sutures.     

Comparative morphology of the carpel in the Liliaceae: Neodregeae

The structure of the carpel has been studied in flowers of the Neodregeae ( Dipidax and Neodregea ). Except in D. triquetra , which is syncarpous, the carpels are united below and free above. A dorsal bundle, two or more lateral bundles, and two placental bundles supply each multiovulate carpel. The six placental bundles of the tricarpellate pistil are united by twos in the lower part of the pistil, forming three opposite compound placental bundles in most species of Dipidax and three alternate bundles in D. triquetra and Neodregea : In the latter, an additional septal bundle continues upward as a branch from the compound placental bundle. Sutural openings are usually short and restricted to the top of the locule. All the Neodregeae have monotegmic ovules.     

Comparative morphology of the carpel in the Liliaceae: Helonieae

Most Helonieae have only slight septal indentations between the three carpels: in Xerophyllum deep septal clefts extend centripetally and completely enclosed, narrow septal pockets occur in Metanarthecium . Other unique generic features are found: tepallary-staminal nectarial glands in Heloniopsis , zygomorphy in Chionographis , and dioecism in Chamaelirium . The carpels are biovulate in Chionographis; there are two to several ovules per carpel in Xerophyllum; 8–12 ovules occur in the carpel of Chamaelirium; and numerous bitegmic ovules are borne in many longitudinal rows on enlarged placentae in Helonias, Heloniopsis, Metanarthecium , and Ypsilandra . Except for Metanarthecium , this last-named group of genera displays a near ring composed of 'accessory' placental bundles and a compound septal bundle (with normally oriented xylem and phloem) in cross-section at the inner edge of each septum. Ventral bundles occur in the other four genera.     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE VI. POMOIDEAE: ERIOBOTRYA,HETEROMELES, PHOTINIA,POURTHIAEA, RAPHIOLEPIS,STRANVAESIA

The pomoid genera, Eriobotrya, Photinia, Pourthiaea, Raphiolepis, Stranvaesia, and Heteromeles, have compound inflorescences and biovulate carpels which become papery at maturity. The carpels of all of these except Heteromeles are fused with one another. There are open sutures in the carpels of Heteromeles, Photinia, Pourthiaea, and Raphiolepis, and in these four genera the extent of fusion of the ovular bundle with the wing bundle is related directly to the state of tegumentary fusion and to the extent of fusion of the carpel with the floral cup. In those species of Eriobotrya and Stranvaesia with closed sutures the integuments tend to be fused, as do the ovular and wing bundles, and the carpels are adnate with the floral cup for a considerable distance; in species with open sutures the integuments tend to be free, the ovular and wing bundles tend to be separate, and the extent of fusion of carpel with floral cup tends to be shorter. In genera with connate carpels the wing bundles of adjoining carpels may also be fused. The greatest extent of fusion occurs in Eriobotrya and Raphiolepis, in which there may also be attenuation and disappearance of the wing bundles above the region of ovular insertion and even reduction and disappearance of the carpellary margin.     

Comparative morphology of the carpel in the Liliaceae: Glorioseae

The pistils of the Glorioseae (Gloriosa, Littonia, Sandersonia) are generally tricarpellate and alike. Virtually all have closed sutures at flowering; they have many ovules, some of which are barely bitegmic, with inner integuments often nearly fused with nucellar remnants; and there is usually but one compound septal bundle in the inner edge of a septum. In two species of Littonia , the compound septal bundle divided to form two simple septal bundles; but in many other plants it remained undivided, and in some it died out, still undivided, below the locular apex. Most of the placental and septal bundles are vascularized in large part by three alternate (compound septal) bundles at the base of the locules and sometimes by branches from the lateral bundles. Three large (compound) placental bundles are formed just below the lowermost ovular insertion, and each then divides in two to furnish ovular branches along their ascent. Occasional auxiliary placental bundles lie between the septal bundle and the placental bundles in the septum (Gloriosa, Sandersonia).     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE VII. POMOIDEAE: CHAENOMELES,CYDONIA, DOCYNIA

The multi-ovulate pomoids, Chaenomeles, Cydonia, and Docynia, all have closed sutures and extensive fusion between carpel and floral cup and between ovular and wing bundles. Although the ovules in Docynia are generally apotropic and few in number (4–7), the ovules in the other two genera are pleurotropic and numerous (15–48). A statistical treatment of the whole tribe of Pomoideae shows that in carpels with open sutures ovular and wing bundles definitely tend to be separate while in those with closed sutures these bundles tend to be fused. To a lesser degree carpels with open sutures also tend to have bitegmic ovules, separate carpels, and a lesser extent of fusion between carpel and floral cup, while carpels with closed sutures tend to have monotegmic ovules, united carpels, and a greater extent of fusion between carpel and floral cup.     

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.     

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.     

Comparative morphology of the carpel in the Liliaceae: Iphigenieae

The pistil in the flowers of the Iphigenieae (Camptorrhiza, Iphigenia, Omithoglossum) is usually tricarpellate. The carpels are coherent generally, with closed sutures and seemingly bitegmic ovules. Camptorrhiza differs from the others in having a single compound style. The pistils of most species of these genera have a common vascular structure: three dorsal bundles which run into the style(s), a number of lateral bundles, six placental bundles, and up to three compound septal bundles. The latter nine bundles usually differentiate from a central vascular plexus above the base of the locules. There may be fewer than three septal bundles in some specieS. When present, the septal bundles usually die out in the ovuliferous region, but in some cases they persist to the apex of the locules.     

THE COMPARATIVE MORPHOLOGY OF THE CANELLACEAE. IV. FLORAL MORPHOLOGY AND CONCLUSIONS

The morphology and anatomy of 105 flowers representing 13 species and 6 genera of the Canellaceae are summarized. The flowers are borne in axillary or terminal racemes, cymes, or small groups, or solitary, in an axillary or terminal position. The flowers are characterized as follows: bisexual, hypogynous; sepals 3, thick and leathery; petals, 5–12, free or united into tube at base, rather thick, in 1 or 2 whorls and/or spirals; androecium of 6–12 stamens united by their filaments forming a tube, anthers with longitudinal extrorse dehiscence; gynoecium of 2–6 carpels fused by their ventral margins; 2–6 placentae. There are 2 vascular bundles (rarely 3) to each sepal, 3 to each petal (some of the inner petals have only 1), 1 to each stamen and 1 trace to each carpel. The petal and stamen bundles have a common origin. All the data accumulated in this series on the Canellaceae indicate that the correct systematic placement of the Canellaceae is in the woody Ranales, perhaps in a complex with the Myristicaceae.     

ANATOMY OF THE GYNOECIUM IN TWO SPECIES OF BAKERIDESIA (MALVACEAE)

Structure of the gynoecium is described in two species of Bakeridesia, subgenus Bakeridesia (Malvaceae, tribe Malveae). The dorsal wall of each carpel bears a winglike projection with a marginal pair of pubescent, bluntly dentate wings. The projection arises as a single, solid ridge of tissue after the ovules are initiated and after the ventral carpellary margins are fused with the receptacle. Two multiseriate layers of fiber-sclereids line each locule and continue into the winglike projection where they are separated by parenchyma. Gynoecial vascularization is described in detail. The richly vascularized carpels are supplied by five traces: a median dorsal trace, which bifurcates into two dorsal bundles; two lateral traces; and two ventral traces. Adjacent ventral traces, lateral traces, and septal bundles are fused—i.e., they are held in common by neighboring carpels. The presence of lateral carpellary traces may be a primitive character in the tribe Malveae.     

Comparative morphology of the carpel in the Liliaceae: Colchiceae (Colchicum)

The pistil of Colchicum is syncarpous, the carpels having open sutures or well-marked commissures and many bitegmic ovules of variable orientation. Although the vascularization of the carpel is also variable, there are usually three dorsal bundles and three alternate, septal bundles at the base of the pistil, with occasionally some placental bundles at that level. More often the placental bundles, differentiating basipetally, appear to establish connections with the septal bundles higher up, at the lowermost ovular insertion level. The septal bundles divide in two more frequently in pistils in which the carpellary suture is open than in those in which it is closed.     

COMPARATIVE STUDY OF THE FLORAL VASCULATURE IN SAURURACEAE

The floral vascular systems are compared among all six taxa of Saururaceae, including the two species of Gymnotheca which have not been studied previously. All are zygomorphic (dorsiventrally symmetrical), not radial as sometimes reported, in conformity with dorsiventral symmetry during organogenesis. Apocarpy in the two species of Saururus (with four carpels and six free stamens) is accompanied by a vascular system of four sympodia, each of which supplies a dorsal carpellary bundle, two ventral carpellary bundles, and one or two stamen traces. The level at which the ventral bundles diverge is the major difference in vasculature between the two species. The other four taxa are all syncarpous, and share some degree of stamen adnation and/or connation. The vascular systems also show varying degrees of fusion. The two species of Gymnotheca (with four carpels and six stamens) are very similar to each other; in both, the ventral traces of adjacent carpels fuse to form a placental bundle, which supplies the ovules and then splits into a pair of ventral strands. The flowers of Houttuynia cordata (with only three carpels and three adnate stamens) are sessile. Each flower is vascularized by three sympodia; the median adaxial sympodium is longer than the other two sympodia before it diverges to supply the adaxial organs. Three placental bundles also are formed in Houttuynia, but the three bundles differ in their origin. The median abaxial placental bundle diverges at the same level as the three sympodial bundles of the flower, while the other two lateral placental bundles diverge at a higher level from the median adaxial sympodium. Anemopsis californica, with an inferior ovary of three carpels, sunken in the inflorescence axis, and six stamens adnate to the carpels, has a vascular system very similar to that of Houttuynia cordata. The modular theory of floral evolution is criticized, on the bases of the known behavior of apical meristems and properties of vascular systems. The hypothesis is supported that saururaceous plants may represent a line of angiosperms which diverged very early.     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE. II. PRUNOIDEAE: MADDENIA,PYGEUM; OSMARONIA

A survey of species of the prunoid genera, Maddenia and Pygeum, and of the genus Osmaronia has been made. The ovules of all are pendent, campylotropous, and epitropic. In the prunoids, the ovular supply is intimately connected with a central vascular plexus in the base of the carpel; that plexus is absent from Osmaronia. The prunoid carpels are marked by an extensive degree of fusion among the ovular and wing bundles, by fusion of the sutural margins, by fusion of the 2 integuments of the ovule to a single massive one, and by the presence of 3 or 5 well-developed bundles in the base. The carpel of Osmaronia also has a strongly fused bipartite ovular supply, separate bundles of which, however, become very much attenuated before reaching the funiculus; it has independent ovular and wing bundles, completely separate carpellary margins, 2 clearly separate integuments in the ovule, and 6 distinctive bundles in the carpel base. At the funiculus, the wing bundle of Osmaronia is connected with the adjoining weak ovular bundle by a well-developed vascular branch. Various particularities in the morphology of Osmaronia lend support to its segregation into a unique tribe, the Osmaronieae of Rydberg.     

兰花蕉花的形态解剖学

兰花蕉（Orchidantha chinensis)的子房室顶部闭合后向上延长成延长部,实心,但有花柱沟和隔膜蜜腺管通过,隔膜蜜腺管,可分为中央蜜腺管和三条侧蜜腺管;中央蜜腺管位于三个心皮连接处,自子房室区下部产生,向上于延长部的部顶端终止;三条侧管分别位于两个心皮连接处,于子房室区近中部产生,开口于花柱基部。兰花蕉子房室区与延长部均具6枚雄蕊的维管束系统,即3枚心皮背束的伴束与3枚隔膜束,近轴面1枚事膜向上进入唇瓣的维管束系统,位于唇瓣的中央,致使兰花蕉仅具5枚功能雄蕊,唇瓣具双重结构,本文还讨论了兰花蕉科的系统发育位置。     

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.     

Comparative floral structure and systematics of Pelagodoxa and Sommieria (Arecaceae)

Floral structure is compared in Pelagodoxa and Sommieria (Arecaceae, Arecoideae). Male flowers have three free, imbricate sepals, three basally congenitally united and apically valvate petals, and six stamens. Anthers are dorsifixed and dehiscence introrse. The sterile gynoecium is tricarpellate. Female flowers have three free, imbricate sepals and three free, imbricate petals, which are slightly fused with the sepals at the base. Four to six staminodes are congenitally united at the base and fused with the ovary for a short distance. The gynoecium is syncarpous. Carpels are almost equal in early development; later the gynoecium becomes pseudomonomerous. The three stigmatic branches are equally developed, apical and sessile. The carpels are (syn-)ascidiate up to the level of the placenta and (sym-)plicate above. Each carpel has one ovule, in the sterile carpels it is aborted at anthesis. The fertile ovule is erect up to anthesis and pendant afterwards because of the bulging out of the ovary. Pollen tube transmitting tracts (PTTT) encompass the secretory epidermis of the ventral slits of each carpel. Floral structure in Pelagodoxa and Sommieria supports the sister group relationship between the two genera suggested in recent molecular phylogenies and reflects their close relationships to a major clade of pseudomonomerous arecoid palms from the Indo-Pacific region.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 27–39.