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.     

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.     

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.     

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.     

FLORAL ONTOGENY OF ILLICIUM FLORIDANUM,WITH EMPHASIS ON STAMEN AND CARPEL DEVELOPMENT

The ontogeny of the flower and fruit of Illicium floridanum Ellis, the Star Anise, was investigated. Each of 5 or 6 bracts in each mixed terminal bud subtends either a vegetative or floral bud. The solitary flowers occur in terminal or axillary positions. Each flower has 3–6 subtending bracteoles arranged in a clockwise helix. The flowers in our material have 24–28 tepals, 30–39 stamens, and usually 13 (rarely 19) uniovulate carpels. Tepals and stamens are initiated in a low-pitched helix; carpels later appear whorled, but arise successively at different levels on the apical flanks. The floral apex is high-convex in outline with a tunica-corpus configuration; it increases in height and width throughout initiation of the floral appendages. Tepals, stamens, and carpels are initiated by one to several periclinal divisions in the subsurface layers low on the apical flanks, augmented by cell divisions in the outer layers of the corpus. The carpel develops as a conduplicate structure with appressed, connivent margins. Procambium development of floral appendages is acropetal and continuous. Bracteoles, tepals, stamens and carpels are each supplied by 1 trace; the carpellary trace splits into a dorsal and an ascending ventral sympodium. The latter bifurcates to form 2 ventral bundles. The ovular bundle diverges from the ventral sympodium. Ovule initiation occurs in a median axillary position to the carpel, an unusual type of ovule initiation. The fruit vasculature is greatly amplified as the receptacle and follicles enlarge. After carpel initiation an apical residuum persists which is not vascularized; a plate meristem develops over its surface to produce a papillate structure.     

Morphology of the gynoecium and systematic position of the Ochnaceae

The gynoecium is syncarpous in all Ochnaceae. In the Ochnoideae carpels are peltate with a conventional cross-zone bearing one ovule, or, in Lophira , a very broad cross-zone with an horizontal ovular row. In Ochna and Brackenridgea , the style is gynobasic, each carpel develops transmitting tissue on its morphologically dorsal surface, and this tissue lines a canal or originates a solid inner strand in each carpel at style level. The style is tubular, with an inner cuticle, and compound, each component with its own transmitting tissue. In Ouratea the style is solid with a single compound transmitting strand. In Lophira and Elvasia the transmitting tissue seems to be developed by the morphologically ventral carpellary surfaces. Ovules are unitegmic with a bivalent integument.
In the Sauvagesioideae carpels are peltate, but with ovules above the cross-zones, on margins of the symplicate zone. In Euthemis , there is one ovule on each side of, and close to, each cross-zone. The single stylar canal is bounded by the morphologically dorsal carpellary surfaces. In Sauvagesia ovules occur on both sides of the cross-zones but most of them are above on carpel margins, as are all ovules of Cespedesia. The stylar canal of Sauvagesia is bounded by the ventral carpel surfaces, three strips of the outer surface passing inside at the sutures and developing into transmitting tissue. The stylar canal of Cespedesia is bounded by the dorsal carpel surfaces. The gynoecium of Wallacea has two epeltate carpels with a laminar placentation, the carpel margins being displaced on to the topographically ventral carpel surfaces with a row of ovules along each margin. Ovules are bitegmic.
The Ochnoideae, which shows relationships with the Rutaceae, Meliaceae, Simaroubaceae and Hippocastanaceae, is more advanced than the Sauvagesioideae, which clearly belongs in the Violales. The Ochnaceae is to be placed in the Violales.     

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 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.     

Vascular floral anatomy of the east asianHeloniopsis orientalis (Thunb.) C. Tanaka (Liliaceae-Helonieae)

Observations on the vascular floral anatomy, carpel morphology and floral biology ofHeloniopsis orientalis are presented. The lower flowering pedicel has six large bundles which lack an enclosing sclerenchymatous sheath. At mid-pedicel, branch bundles originate via radial divisions from each of these bundles. Subsequently, there is a vascular ring of 12 bundles below the receptacle. The six smaller bundles which are derived from alternate pedicel bundles eventually establish all of the ventral gynoecium supply. The six larger bundles supply the tepals, stamens and dorsal gynoecial vasculature. The simple dorsals do not branch or fuse in their vertical ascent. The ventral and placental supplies are far more complex. Fusion occurs between paired sets of the six smaller pedicel bundles along the septal radii and results in a submarginal laminal ventral network. An independent ventral plexus is formed in each septum and from each plexus two septal axials, of which the innermost has a reversed xylem-phloem disposition, and four placental bundles are derived. Two placental bundles are associated with each septal axial. Basally the septa are fused centrally, but are freed at mid-gymoecial height. The broadly tri-lobed, tri-carpellate gynoecium is depressed terminally where the erect, hollow style with its capitate stigma is attached. Dorsal grooves are present: the fruit is loculicidally dehiscent. There are no septal glands due to complete lateral fusion of the septal wings. Basally each of the six equal tepals has a saccate nectary. The similarity in vascular anatomy and carpel morphology of the AsianHeloniopsis and eastern North American endemic,Helonias bullata, justifies their position in the same tribe. Research and publication supported in part by the M. Graham Netting Research Fund through a grant from the Cordelia Scaife May Charitable Trust, the U. S.—Japan Cooperative Science Program Grant GF-41367, the Japan Society for the Promotion of Science, and Grant-in-Aid No. 934053 from the Ministry of Education, Japan.     

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 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.     

CARPELLARY VASCULATURE AND THE OVULAR VASCULAR SUPPLY IN DRIMYS

The ovules in Drimys winteri var. chilensis and D. lanceolata are consistently vascularized entirely by the ventral bundles, without contribution from the dorsal bundle(s) as generally assumed. The ovules are initiated in two rows, without any in “median” position. Post-initiatory differential growth of the carpel wall brings the lowermost ovules into apparently median position at maturity. The anomalous vascular supply to the lowermost ovules is thought to be related to concurrence of delayed initiation and development of these ovules with delayed differentiation of the vascular supply.     

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 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.     

Le Basi Anatomiche Dei Rapporti Nutritivi Tra Pianta Madre,Seme Ed Embrione

Abstract

The anatomical basis of the nutritive relationships between mother plant, seed, and embryo. — The morphology and anatomy of the fruits and seeds of the Angiosperms show a great variety of structures and adaptments, even within the same family, and one must be cautious in drawing generalized conclusions.

If we first examine the ovary we see that the single carpel receives three vascular traces from which the three main bundles originate, a dorsal and two ventral ones, all more or less reduced. Except in the case of laminar placentation the ovule traces are connected to the ventral vascular system, but often the entire vascular system of the ovary is anastomosed and therefore reticular. However especially when the placentae are at the centre or at the basis of the ovary, it is possible to detect a tendency towards a separation between the vascular system of the ovarian wall and that of the placentae.

The ovular bundle runs through the funicle reaching the chalaza, where it can either end or continue towards the micropyle with a single bundle or with a few branches or even forming a complete reticular envelope surrounding the ovule. The ovular vascular bundles are normally found in the outer integument.

The ovule is made of an inner part (nucellus), and an outer one (integuments). The integuments play a very important role in the processes of seed maturation, dormancy, and germination. They are isolated from the interior of the seed by a cuticle which is a common production of the inner integumentary epidermis, and of the nucellus. The cuticle is not present in the chalaza and can be dissolved in the micropylar region: through these two apertures nutrients can penetrate into the seed or haustoria can grow out of it. During the course of maturation these openings become closed by various means, often through the formation of a new cuticle or of a suberised chalazal plate.

The nutrients which pass through the chalaza penetrate into the nucellus where in some cases one can find some structures which facilitate the communications between the chalaza and embryo sac. The endosperm feeds at the expense of the nucellus but often it can establish a direct contact with the chalaza or the integuments or even the placentae. This occurs often thanks to haustoria.

The embryo is normally surrounded at first by a more or less liquid endosperm: in a second stage the endosperm becomes cellular and the embryo grows at its expense through the digestive activity of the cotyledonar epidermis.

From an anatomo-physiological point of view the following points seem of particular interest:

(I) The endosperm and the embryo show a remarkable autonomy in respect of the mother plant: from an anatomical point of view this is shown by the isolation of the endosperm and embryo by means of a cuticular covering or substitutive structures and by the interposition of nutritive tissues between the vascular system of the mother plant and the endosperm.

(II) Given the importance of the inner cuticle its presence and its constitution should be ascertained in the various species having also in mind the properties of selective permeability shown by the testa.

(III) Two nutritive mechanisms exist: translocation of nutrients via the vascular system and the nutritive tissues, and digestion of surrounding cells. In the digestive phenomena it is important to explain the mechanisms by which only the right cells are digested and not the others.

(IV) The embryo very frequently is immersed at first in a more or less liquid endosperm and is later surrounded by a compact tissue; the nutritive mechanisms are probably different in the two cases.

(V) Two endospermic zones are often distinguishable: one having an haustorial or at least a digestive or elaborative function, and being typically non cellular; another zone, typically cellular, forms a tissue which is sooner or later absorbed by the embryo. The cellularization of this zone seems to coincide with the establishment of polarity and with the beginning of maximum growth of the embryo.

(VI) The relationships between the inner seed and the integuments is complex and there is a correlation between the histoanatomical and biochemical changes of these two parts during seed development. The modifications undergone by the integuments are important steps also towards the preparation of the seed to the processes of dispersal, dormancy, and germination.     

VASCULAR SUPPLY AND STRUCTURE OF THE OVULE AND ARIL IN PEONY AND OF THE ARIL IN NUTMEG

Camp , Wendell H., and Mary M. Hubbard . (U. Connecticut, Storrs.) Vascular supply and structure of the ovule and aril in peony and of the aril in nutmeg. Amer. Jour. Bot. 50(2): 174–178. Illus. 1963.—Examination of the placental region in the carpel of Paeonia indicates a complexity and super-abundance of vascular supply beyond that usually found in angiosperms and certainly more than is necessary for adequate nutrition and water supply of the ovules. From this it is concluded that the ovules once were borne on a larger and more complex structure than the present carpel. Vascular strands leading to the aril and the hypostase are interpreted as being relictual. The large multifid aril of Myristica has a well-developed vascular system composed of several sizes of branched bundles.     

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 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.     

Floral ontogeny of Zippelia begoniaefolia and its familial affinity: Saururaceae or Piperaceae?

Zippelia begoniaefolia Bl., a monotypic species having characteristics of both Piperaceae and Saururaceae, has racemes of about 20 small flowers lacking a perianth, each with six free stamens and a four-carpellate syncarpous gynoecium. The inflorescence apical meristem initiates bracts acropetally and helically, each of which subtends a later initiated single floral apex; there are no “common” primordia. The six stamens are initiated as two lateral pairs and two solitary successive primordia, the latter two opposite in median sagittal positions. Four carpel primordia are initiated as a lateral pair and two successively initiated in the median sagittal plane. This order of organ inception is unique among Piperaceae and Saururaceae. Intercalary growth below carpellary attachment raises them up on a common cylindrical base that becomes the syncarpous ovary, covered with unique glochidiate hairs and containing a single basal ovule. The free portions of the carpels become the reflexed papillate stigmas. The floral vascular system has a single bundle at base that branches to supply the bract and flower traces. The floral vasculature is similar but not identical to that of Saururus (Saururaceae) and some Piper species (Piperaceae). Plesiomorphic character states of Zippelia that are shared with Saururus include hypogyny, free stamens, cleft stigma, and a similar floral groundplan. Synapomorphies, derived shared character states that unite Zippelia with Piperaceae, include syncarpy, solitary ovule, basal placentation, fused ventral carpellary bundles, and a double vascular cylinder in the stem. Cladistic analysis aligns Zippelia with Piperaceae because they share apomorphies, and because Zippelia shares only plesiomorphies with Saururus.     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE. III. POMOIDEAE: CRATAEGUS,HESPEROMELES, MESPILUS,OSTEOMELES

The carpels in Crataegus, Hesperomeles, Mespilus, and Osteomeles appear to constitute a morphologically related group: all have bony pits, ovules that tend to be acollateral (usually superposed), and clearly separate ovular and wing bundles, i.e., no “ventral” bundles, at the level of ovular insertion. In species whose carpels have no sutural opening, the integuments are more extensively fused with one another, the degree of intercarpellary fusion tends to be greater, and the carpels are fused with the floral cup to relatively higher levels than in those species whose carpels have a sutural opening. In the few cases in which wing and ovular bundles are adnate at the locular base (Crataegus monogyna, Mespilus, Osteomeles anthyllidifolia, O. Schwerinae), the extent of inter- and extracarpellary fusion and sutural closure is among the most advanced.