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 ROSACEAE. IV. POMOIDEAE: CHAMAEMELES,COTONEASTER, DICHOTOMANTHES,PYRACANTHA

The carpels of Chamaemeles, Cotoneaster, Dichotomanthes, and Pyracantha tend to be separate from one another, their sutures tend to be closed, and they become more or less bony at maturity. However, aside from having collaterally placed ovules, they do not appear to be structurally similar. There seem to be 2 different evolutionary trends in the ovular bundle–wing bundle relationship: in Pyracantha, progressive fusion between the ovular bundle and the wing bundle has led to the formation of a “ventral” bundle; in Cotoneaster, and possibly Chamaemeles, the wing bundle has become reduced and rather attenuated. A primitive pomoid state may be represented by the carpel of Dichotomanthes, which is completely free of the floral cup and in which wing and ovular bundles are separate. Differences in sutural closure appear only in Cotoneaster, and in species of that genus the wing bundles and ovular bundles tend to be fused if the suture is closed, and separate if it is open.     

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.     

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.     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE IX. SPIRAEOIDEAE: QUILLAJEAE,SORBARIEAE

A comparative study of carpellary structure in the spiraeoid subtribes Qui lajeae and Sorbarieae has shown that morphological inter-relationships are similar to those of other Rosaceae. When the suture is closed the carpels tend to be coherent and the ovular and wing bundles tend to be fused. These relationships are statistically significant in the Spiraeo deae as a whole. The construction of the gynoecium in Lindleya and in most species of Vauquelinia resembles that of a pomoid. Other features of a pomoid-spiraeoid affinity have been discussed.     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE. V. POMOIDEAE: AMELANCHIER,ARONIA, MALACOMELES,MALUS, PERAPHYLLUM,PYRUS, SORBUS

The carpels of 2 groups of pomoid genera, Amelanchier, Malacomeles, Peraphyllum and Aronia, Malus, Pyrus, and Sorbus, were analyzed morphologically. Open sutures are associated with a lesser extent of tegumentary fusion and ovular bundle–wing bundle fusion than are closed sutures. However, in the genera as a whole (and particularly in Aronia and Sorbus), the extent of sutural closure is inversely related with the amount of intercarpellary adhesion and with the fusion of carpels to the floral cup. In the Amelanchier group and in Malus and Pyrus, ovular- and wing-bundle fusion is directly related with intercarpellary adhesion. Malus and Pyrus have closer structural resemblances with one another than they have with Aronia and Sorbus.     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE. I. PRUNOIDEAE: PRUNUS

A study of the carpel in 27 species of Prunus has shown certain notable structural relationships associated with the extent of closure of the carpellary margins. These relationships involve the degree of fusion of the 2 integuments, the number of vegetative bundles in the base of the carpel, the extent of fusion of the ovular bundles with one another and with the wing bundles, the relative size of the ovular bundles, and the relative development of the central vascular plexus. The comparative evidence strongly supports a primitively separate state of ovular and vegetative bundles. The significance of this finding is discussed.     

THE AFFINITIES OF PRINSEPIA (ROSACEAE)

Sterling, C. (U. California, Davis.) The affinities of Prinsepia (Rosaceae). Amer. Jour. Bot. 50(7): 693–699. Illus. 1963.—Anatomical study of the carpels of 4 species of Prinsepia has shown that at flowering the 2 ovules are erect and pleurotropic. The funiculus is on the dorsal and lower side of the ovule; the micropyle faces a large obturator on the ventral side. The carpellary margins are separated by a fissure below the funicular insertion, but above this level they are fused. The style is laterally inserted on the ventral face of the carpel; it is vascularized only by the wing bundles and the recurving dorsal bundle. At the base of the ovary, 2 ovular bundles depart from the vascular cylinder and run separately, each to its respective ovule. In carpel morphology, ovular position, ovule structure, and vascular anatomy, Prinsepia is not a prunoid type. Although its features on the whole resemble those of chrysobalanoid plants, there are notable differences. Consequently, Prinsepia is assigned to a new subfamilial group in the Rosaceae, the Prinsepioideae. Some phylogenetic considerations are discussed briefly.     

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 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 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.     

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.     

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: 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.     

THE FLORAL ANATOMY OF KOEBERLINIA ZUCC: SYSTEMATIC IMPLICATIONS

The in toto pattern of the floral vasculature in Koeberlinia Zucc, is distinctive. The median vascular trace to each sepal is concrescent with the antesepalous stamen trace forming a trace complex. Each petal trace is concrescent with the nearest antestaminal trace, and this common trace is in turn concrescent basally with the common basal supply to the adjacent sepal margins. The ventral carpellary bundles and the ovular traces of the two carpels are arranged for part of the ventral carpellary system into an essentially continuous hollow stele-like cylinder and many of the ovular vascular supplies originate from this strand. All vascular concrescences are congenital. Comparisons of the morphological and floral vasculature characters of Koeberlinia with those of its various putative allies revealed that there are no substantial reasons for linking Koeberlinia with Canotia, Celastraceae, Rutaceae, Simaroubaceae, or Zygophyllaceae. The in toto floral vascular structure of Koeberlinia is closely similar to that of the Caryophyllaceae and dissimilar to that of the Capparaceae. Several qualitative characters of the secondary xylem of Koeberlinia differ from those of the Capparaceae, yet certain important ones are similar. Many of the morphological characters of Koeberlinia are similar to those of the Capparaceae as well as the Caryophyllaceae, yet certain critically important ones strongly indicate a relation of Koeberlinia to the Capparaceae: occurrence of myrosin cells, capparaceous pollen, capparaceous ovular characters. To include Koeberlinia within either of these families is unwise, but the writers are inclined to retain Koeberlinia in a monogeneric family within the larger Capparales.     

THE MORPHOLOGY OF THE MYRISTICACEAE. I. FLOWERS OF MYRISTICA FRAGRANS AND M. MALABARICA

Floral histology and vascular anatomy of Myristica fragrans Van Houtt. and M. malabarica Lam. have been investigated from serial sections and specimens cleared in chloro-lacto-phenol. The flowers are unisexual. The androecium is considered to consist of a whorl of laterally concrescent anthers. The bisporangiate anthers are attached by a ridge of tissue to the terminal part of the androphore. In many cases the number of vascular bundles in the androphore is half the number of anthers. The gynoecium consists of a monocarpellate pistil with basal placentation and a single anatropous ovule. Of the many vascular bundles that enter the base of the carpel, two, because of their position and because they provide vascular traces to the ovule, are designated as ventral bundles. Additional ovular traces are provided by the carpel wall vascular system. These additional traces originate at the top of the locule and descend to the ovule. The similarity between the androecia of these two species and the androecium of the ćnellaceae is noted.     

Comparative morphology of the carpel in the Liliaceae: Uvularieae

Three genera of the Uvularieae (Kreysigia, Schelhammera, Uvularia) have tricarpellate, syncarpous pistils. Ventral bundles (presumably the united simple septal and placental bundles of a carpellary wing) may be present in Kreysigia and Schelhammera. In Kreysigia the two presumptive ventral bundles from adjoining carpels are fused basipetally in each septum. The septal bundles of the other two genera are either simple (Schelhammera) or in part compound (united) below and simple (separate) above (Uvularia) , hence fused acropetally. In Uvularia , the dorsal bundle of the carpel and the median bundle of the tepal are uniquely tripartite and probably homologous. No raphides were found in the carpels of these genera.     

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.     

Typification of some Maloideae (Rosaceae) commonly cultivated in Norden

In connection with a contribution of various genera of Maloideae (Rosaceae) to "Flora Nordica" lectotypes are designated for Crataegus sanguinea, C. punctata, Pyrus japonica (‵ Chaenomeles japonica ), Crataegus pyrifolia (‵ Photinia pyrifolia ), and Pyrus floribunda (‵ Photinia floribunda ) . A lectotype and an epitype is selected for Mespilus flabellata (‵ Crataegus flabellata ) . — Flora Nordica note no. 23.     

POSTGENITAL CARPEL FUSION IN CATHARANTHUS ROSEUS (APOCYNACEAE). I. LIGHT AND SCANNING ELECTRON MICROSCOPIC STUDY OF GYNOECIAL ONTOGENY

The detailed ontogeny of postgenital fusions within the gynoecium of Catharanthus roseus was investigated. The basal margins of the young carpel primordia infold and fuse together to seal shut the loculi. Independently, the opposing distal tips of the two carpels also unite, with the fusion region subsequently developing into the stigma, style, and a small distal region of the compound ovary. The basal ovary regions of the two opposing carpels remain unfused, thus leaving the tip fusion spatially restricted. In the region of contact, cells with distinctively epidermal features progressively lose their epidermal character after their participation in the fusion. In the fused stigma these former epidermal cells redifferentiate into transmitting and secretory tissues; in the fused style these cells undergo a tremendous expansion in length while forming stylar transmitting tissue; but in the compound ovary region corresponding cells experience little expansion or redifferentiation. It is concluded that the loss of epidermal features or the occurrence of periclinal cell divisions in the epidermis is a definitive indication that cells have fused postgenitally. However, studies with the transmission electron microscope are necessary to detect the first indications of a postgenital fusion. The compound ovary region within the gynoecium of C. roseus is a tissue appropriate for a high resolution ultrastructural study of the cytological events accompanying postgential tissue fusion because the fusion occurs quickly and little subsequent cell expansion takes place within this region.