Vegetative anatomy of Ophiopogoneae (Convallariaceae)

CUTLER, D. F., 1992. Vegetative anatomy of Ophiopogoneae (Convallariaceae). The vegetative anatomy (particularly leaf) of species of Ophiopogon, Liriope and Peliosanthes is described from observations with light and scanning electron microscopy. A syndrome of leaf characters is present, including epidermal features; hypodermal fibre-like cells; raphides and unusual short, square-ended prismatic crystals arranged in plates; phloem with abundant sclerenchyma and frequent individual strands each composed of a sieve tube element and its associated companion cell; and vascular bundles with unusual orientation, which shows the very close inter-relationship between Ophiopogon and Liriope. Peliosanthes shares the phloem type, hypodermal fibre-like cells and raphides, but is less similar in epidermal characters and vascular bundle orientation. The significance of the unusual phloem type is considered in relation to similar types in other members of the Liliiflorae.     

Vegetative anatomy of Pachycortnus (Anacardiaceae)

Pachycormus discolor , an arborescent desert perennial endemic to Baja California, has small, pinnately compound, hypostomatic, bifacial leaves produced on short shoots and photosynthetic stem phelloderm covered by exfoliating translucent phellem. Tightly packed laminal palisade cells are filled with tannins and lack chloroplasts. Spongy mesophyll is the major photosynthetic tissue. Leaves possess unicellular trichomes with secondary walls and uniseriate trichomes with glandular heads. Schizogenous resin ducts occur in primary phloem of stems, leaves and roots as well as all living tissues of the bark. Developmental studies reveal that initiation and differentiation of foliar primordia resembles that of other dicotyledons except that tannin cells and secretory ducts arise precociously. Primary vasculature is an open sympodial system with three principal traces diverging toward each foliar primordium. The wood is highly specialized and comprises mostly unlignified cells packed with starch grains. Thick bark is mainly produced as annual layers of secondary phloem marked by a ring of secretory ducts each surrounded by tannin cells. The possible adaptive significance of these unusual anatomical features is discussed.     

Fossil forms of Amentiferae

Review of the procedures used in determining fossil plant organs indicates that the many Cretaceous records of extant genera of “Amentiferae” based on leaves should be rejected as theoretically unreliable. Palynological data, in combination with some valid megafossil data, indicate that most recognizable members of “Amentiferae” are no older than the later part of the Late Cretaceous. Juglandales appear to be derivatives of the ancient Normapolles complex and unrelated to other “Amentiferae.” A preliminary account of some of the comparative foliar morphology of extant “Amentiferae” indicates that some—particularly Betulaceae and Fagaceae—are closely related to Hamamelidales but that other families—notably Rhoipteleaceae, Juglandaceae, Didymelaceae, and Leitneriaceae—are unrelated to this order.     

Vegetative anatomy and systematics of Triphorinae (Orchidaceae)

Triphorinae represents a group of three anatomically simple genera, the structural features of which are unspecialized. The anomocytic stomatal pattern occurs in all genera; it predominates in Triphora. A foliar hypodermis, sclerenchyma, fibre bundles and stegmata are absent. The mesophyll is homogeneous. The exodermal and endodermal cells in the roots are entirely thin‐walled and tilosomes are absent. However, there are anatomical modifications that appear to be unique: root hairs in Monophyllorchis are borne on velamenal buttresses and, in Psilochilus, they arise endogenously. In the root vascular system of Psilochilus, the metaxylem occurs as a circumferential band. The surfaces of stems in Triphora are tuberculate. Mycorrhizae appear to characterize the root cortices of all genera. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 203–210.     

Chemical constituents and systematics of Amentiferae

The systematic value of various kinds of chemical constituents reported from Amentiferae is discussed in terms of chemical structural complexity and restricted distribution. Chemical evidence suggests the cohesiveness of Urticales, excluding Eucommiaceae. It also suggests that Garryaceae, Platanaceae, and Myricaceae are distinct from other families of Amentiferae. The absence of reports of elaborate, restricted naturalproduct structural types in Betulaceae, Fagaceae, Casuarinaceae, Juglandaceae, and Salicaceae means that there is no positive chemical evidence suggesting relatedness of these core families of Amentiferae.     

Vegetative anatomy of subtribe Habenariinae (Orchidaceae)

Leaves of Habenariinae are characterized by anomocytic stomatal apparatuses, homogeneous mesophyll, collateral vascular bundles in a single series, and thin-walled bundle sheath cells. There is no foliar sclerenchyma nor a hypodermis. Cauline cortex consists of thin-walled living cells among which are large and numerous intercellular spaces. The ground tissue is bordered externally by a layer of thick-walled living cells, except in Habenaria repens. Central ground tissue cells are living, and usually thin-walled surrounding intercellular spaces of various dimensions. These are conspicuously large in H. repens. Collateral vascular bundles are scattered across the ground tissue. Sclerenchyma is absent. Absorbing roots are generally velamentous, exodermal dead cells are diin-walled, and passage cells usually have a thickened outer wall. A regular vascular cylinder is present, and vascular tissue is embedded in parenchyma. Root tubers are velamentous, exodermal cells are usually thin-walled, and passage cells frequently have thickened outer walls. Vascular tissue of root tubers is organized into two classes: (1) those with a single vascular cylinder surrounded by a cortex and (2) those with a series of meristeles dispersed throughout the ground tissue. In group (1) cortex is homogeneous either with or without mucilage cells except in Stenoglattis where the cortex is heterogeneous, consisting of water-storage and assimilatory cells, and lacks mucilage cells. In group (2) the ground tissue consists of larger mucilage-containing cells and smaller assimilatory cells.     

Vegetative anatomy of subtribe Orchidinae (Orchidaceae)

Leaves in Orchidinae are essentially glabrous; anticlinal walls of foliar epidermal cells arc basically straight-sided to curvilinear, and cells arc fundamentally polygonal on both surfaces; adaxial cells are larger than abaxial cells. Stomata arc anomocytic and usually only abaxial and superficial; substomatal chambers are small to moderate; outer and inner guard cell ledges are mostly small. There is no hypodermis nor are there fibre bundles. Mesophyll is homogeneous, chlorcnchyma cells arc thin-walled, and intercellular spaces numerous. Crystalliferous idioblasts abound. Vascular bundles are collateral, organized in a single series. and lack associated sclerenchyma. Bundle sheath cells are thin-walled and chlorophyllous. Stems are glabrous; stomata arc frequent in one species, lacking in others. Cortical cells are oval to circular, thick-walled, and interspersed with triangular intercellular spaces. Ground-tissue cells are circular, and triangular intercellular spaces are present. Vascular bundles arc collateral and scattered throughout the ground-tissue or are absent from the central ground-tissue. Epidermis in absorbing roots is one-layered and non-velamcntous. Exodcrmal cells are thin-walled and dead cell walls bear tenuous scalariform bars; some species lack an exodermis. Outer cortical cells are polygonal and lack intercellular spaces; middle layer cortical cells are rounded with triangular intercellular spaces; inner layer cells are polygonal and lack intercellular spaces. Endodermis and pericycle are thin-walled and one-layered. Vascular cylinder is mostly 7–9-arch with xylcm and phloem components alternating regularly; vascular tissue is embedded in parenchyma; pith cells are parenchymatous, polygonal, thin-walled and lack intercellular spaces. Root tubers generally bear a velamen of variable thickness; bulbous-based unicellular hairs frequently form a dense mat; exodermal cells are thin-walled; dead cells have scalariform bars, passage cells are sparse. Ground-tissue consists of rounded water-storage and assimilatory cells interspersed with triangular or quadrangular intercellular spaces; peripheral cells arc polygonal lacking intercellular spaces. Vascular tissue consists of monarch to pentarch meristeles distributed thoughout the ground-tissue each surrounded by a uniscriale endodermis of thin-walled cells. Thin roots ofPlalanthera exhibit a typical central cylinder surrounded by a homogeneous cortex uninterrupted by meristeles; thicker roots show a central vascular cylinder and cortex in which meristeles are also present; in globoid root tubers there is no central cylinder, and the ground-tissue is replete with scattered meristeles. Because the central vascular cylinder in Platanthera gives rise to branches (meristeles), these represent components of a single vascular system and are not separate stelar entities as implied by the use of the term ‘polystele’.     

Vegetative anatomy and systematics of subtribe Dendrobiinae (Orchidaceae)

Anatomy of leaf, stem, and root of more than 100 species in subtribe Dendrobiinae (Orchidaceae) was studied with the light microscope to provide a comparative anatomical treatment of these organs, to serve as an independent source of evidence that might be taxonomically important, and to recommend such reinterpretations of existing classifications as are suggested by a phylogenetic assessment of data. We based our classification on that of Rudolf Schlechter as the most complete and widely accepted today. We found that the anatomy of plants in subtribe Dendrobiinae reflects a high degree of morphological diversity, and many of the anatomical characters appear to be homoplasous. When these anatomical data are used to interpret the systematic relationships among the genera, they indicate that Dendrobium is not monophyletic and that Cadetia and Pseuderia are apparently nested within the structure of Dendrobium when section Grastidium is chosen as a functional outgroup. Lack of resolution in the strict consensus tree illustrates the difficulty of determining the phylogenetic relationships of many of Schlechter's sections using anatomical characters. Nevertheless, we recommend that his sectional classification, with appropriate modifications based on available data, be retained for the present, pending a more detailed understanding of the phylogeny of Dendrobiinae based on morphology, micromorphology, anatomy, and DNA studies.     

Seed morphology and anatomy in someAnnonaceae

Twelve species ofAnnonaceae, namelyAncana sp.,A. stenopetala, Annona cherimola, A. montana, A. muricata, A. squamosa, Bocagea sp.,Bocageopsis canescens, two species ofUnonopsis, Xylopia aromatica, andX. emarginata, were investigated with respect to the morphology and anatomy of the seed. They show the basic structural pattern characteristic of annonaceous seeds: perichalaza, fibrous mesotesta, and rumination developed by both integuments. However, several differential characters, some of them never previously described, were found: All taxa exhibit an additional endotestal mechanical layer which forms the micropylar plug except inAncana. InAncana the plug is built by the inner integument. Aril, sarcotesta and pits on the seed surface are present in some taxa and show different origin and structure. Special anatomical adaptations possibly functioning during germination are described. The results obtained from the investigated taxa are discussed and compared with published data on seed structure inAnnonaceae.     

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.     

Vegetative morphology and trait correlations in 54 species of Commelinaceae

The morphospace of 54 species of Commelinaceae from nine genera was examined with simultaneous attention to constraints, adaptive hypotheses and relatedness. Eleven morphological traits, including leaf length and width, angle between the leaves and internode distances, were measured for each species and analysed by principal components analysis and nested analysis of variance. The results revealed a significant signal of relatedness in vegetative morphology; genus explained 20–50% of the variance in a single trait. The relationships between some traits are consistent with adaptive explanations. The findings are consistent with the prediction that evolution for optimal phyllotaxis should be relaxed as self‐shading decreases, and that light availability governs leaf size and branching patterns. Constraints potentially explain some trait correlations, and support was found for the hypothesis that structural constraints govern leaf size and internode size correlations. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 257–268.     

Comparative anatomy and morphology of nectar-producing Melastomataceae

Background and Aims

Most neotropical Melastomataceae have bee-pollinated flowers with poricidal anthers. However, nectar rewards are known to be produced in about 80 species in eight genera from four different tribes. These nectar-producing species are pollinated by both vertebrates and invertebrates.

Methods

The floral morphology and anatomy of 14 species was studied in six genera of nectar-producing Melastomataceae (Blakea, Brachyotum, Charianthus, Huilaea, Meriania and Miconia). Anatomical methods included scanning electron microscopy, and serial sections of paraffin-embedded flowers.

Key Results

All vertebrate-pollinated melastome flowers have petals that do not open completely at anthesis, thus forming a pseudo-tubular corolla, while closely related species that are bee pollinated have rotate or reflexed corollas. In most species, nectar secretion is related to stomatal or epidermal nectaries and not filament slits as previously reported. Moreover, the nectar is probably supplied by large vascular bundles near the release area. Blakea and Huilaea have nectary stomata located upon the dorsal anther connective appendages. Brachyotum also has nectary stomata on the anther connectives, but these are distributed lengthwise along most of the connective. Meriania may release nectar through the anther connective, but has additional nectary stomata on the inner walls of the hypanthium. Miconia has nectary stomata on the ovary apex. Charianthus nectaries were not found, but there is circumstantial evidence that nectar release occurs through the epidermis at the apex of the ovary and the lower portions of the inner wall of the hypanthium.

Conclusions

Nectar release in Melastomataceae is apparently related to nectary stomata and not filament slits. The presence of nectary stomata on stamens and on ovary apices in different lineages suggests that the acquisition of nectaries is a derived condition. Nectary location also supports a derived condition, because location is strongly consistent within each genus, but differs between genera.Key words: Blakea, Brachyotum, Charianthus, Huilaea, Meriania, Melastomataceae, Miconia, nectaries, nectary stomata, pollination     

Flowers and inflorescences of the “Amentiferae”

The floral and inflorescence morphology of the major genera of the Myricaceae, Betulaceae (including Corylaceae), Fagaceae, Leitneriaceae, and Juglandaceae are reviewed. Major problems in interpretation of morphology are examined in the light of various comparative morphological studies as well as ontogenetic and vascular anatomical studies. Basically similar phenomena associated with miniaturization of the partial inflorescence have led to superficially similar morphological patterns. The partial inflorescences in the various families, in spite of their reduced size, can be adequately analyzed in most cases on the basis of the bract-branch relationship. The highly modified morphology of the floret is clarified by the application of the general tenets of the leaf-stem relationship in the frame of reference of the minute absolute size of the floret. Numerous problems remain to be attacked. The total and partial inflorescences and the florets of the Myricaceae, Betulaceae, Fagaceae, Leitneriaceae and Juglandaceae are reviewed in terms of external morphology, vascular anatomy and ontogeny as reported in the more objective literature on the subject. The total inflorescences in these families range from the complex, androgynous panicles of stiff spikes of such genera asCastanopsis to the condensed, bud-like pistillate spikes of some Myricaceae andCorylus on the one hand, to the simple staminate floret in the axil of the foliage leaf in some species ofNothofagus on the other. In many species of these families the inflorescence is the apparently simple spike with a flaccid axis, the ament, but so often is this not the case that the designation “Amentiferae” for this artificial assemblage must be considered a misnomer. Whether the total inflorescences are composed of racemose or cymose partial inflorescences is a question not completely answered in all the families. In the Betulaceae the partial inflorescence has long been taken to be a cymule. But, a re-interpretation of the vascular anatomy suggests the alternative that the most distal floret in a short raceme has overtopped the axis of the partial inflorescence thus producing a pseudo-cymule. This is similar to a recent interpretation of the staminate partial inflorescence ofMyrica esculenta, where the individual floret is composed of a single stamen. The partial inflorescence in the more reduced species ofMyrica is thus a pseudanthium. Recent ontogenetic studies in the Betulaceae dramatically corroborate the earlier interpretation, based on vascular anatomy, that the staminate partial inflorescence ofOstrya is three-flowered. On similar grounds it has recently been shown that the spiny “involucre” of pistillateComptonia is composed of tertiary bracts. The structure of the staminate partial inflorescences in the Fagaceae seems reasonably clear except in certain species ofNothofagus where it may well be a synanthium, although the alternative of chorisis exists. The interpretation of the pistillate partial inflorescence inLeitneria requires re-study; the unvascularized tepal-like structures subtending the ovary have been alternatively treated as bracts -an ontogenetic study is badly needed. The organization of the staminate partial inflorescence of the Juglandaceae remains equivocal, although recent ontogenetic work on one species ofJuglans shows that the primordia of the secondary bracts are readily distinguished from tepal primordia, although at anthesis they are very similar. At present the number of florets in the partial inflorescence of the Juglandaceae remains an open question in spite of a fragmentary study of the vascular system. The cupule ofLithocarpus andQuercus continues to present a major morphological problem. The valves of the husk in other genera of the Fagaceae seem, on the basis of the vascular anatomy and some ontogenetic information, to be axes of the ultimate order of branching. A thorough study of these complex structures is needed. Staminate florets which are set off by tepals are readily identified with the reservation that those of some species ofNothofagus and ofJuglans, for instance, may be more complex than they seem. The absence of tepals creates major difficulties which have been resolved in some instances by the study of the vascular anatomy and/or ontogeny. But many problems remain. The pistillate floret seems clearly delimited in the various families. There continues to be the usual conflict concerning the proper interpretation of the wall of the inferior ovary, whether on the basis of ontogeny it should be considered cauline or on the basis of the vascular anatomy it is to be considered appendicular. Oddly enough there are also diametrically opposed interpretations of placentation -is it axile or parietal in one and the same species. This perhaps results from a conceptual conflict. The basal ovule, as in the Myricaceae, or even the ovules perched on a partial septum, as in the Juglandaceae, are similarly much discussed. The ontogenists tend to agree that such ovules are cauline, while the anatomists find that the complex vascular system is not that of a stele. There is a multitude of discrepancies, as yet, in observations, and even when there is mutually accepted fact, there are often conflicting interpretations. Above all, there is a massive lack of knowledge of the vascular anatomy and ontogeny of these miniature and modified flowers and inflorescences.     

Vegetative morphology and anatomy of Maundia (Maundiaceae: Alismatales) and patterns of peripheral bundle orientation in angiosperm leaves with three‐dimensional venation

Collateral bundles with external position of the phloem characterize the stem vasculature of most seed plants. An earlier study highlighted the occurrence of inverted peripheral bundles in the leafless inflorescence peduncle of the rare Australian aquatic Maundia triglochinoides. This unusual feature and other morphological and molecular data supported the recognition of the monogeneric Maundiaceae, but the anatomy of the leaves, rhizomes and roots of Maundia remained unknown and is studied here. Maundia has an iterative sympodial growth with all shoots bearing five tubular cataphylls splitting longitudinally and simulating open sheaths at maturity and two (or three) linear foliage leaves without a conspicuous basal sheath. This morphology distinguishes Maundiaceae from all other Alismatales. The rhizome has an atactostele with collateral bundles of normal orientation; peripheral bundles are absent. Cataphylls have a series of normally oriented bundles. Foliage leaves are thick, bifacial, semi‐elliptical in cross‐section, with a thin subepidermal layer of chlorenchyma on both sides, accompanied by peripheral bundles with xylem facing outwards (thus abaxial peripheral bundles are inverted) and central large bundles of normal orientation. Strong anatomical similarity between leaves and peduncles is related to their shared function as assimilatory organs. As in angiosperm succulents, the three‐dimensional leaf venation in thick aquatic and helophyte leaves of Alismatales serves to reduce transport distances between veins and photosynthetic cells. In both cases, the patterns of orientation of peripheral bundles (with inverted adaxial or abaxial bundles) are unstable in large clades. These slender bundles cannot be used for the identification of unifacial leaves. Some anatomical characters express homoplastic similarities between Maundiaceae and Aponogetonaceae.     

Studies in the floral morphology and anatomy of nymphaeales

Floral morphology ofBrasenia schreberi Gmel. andCabomba caroliniana A. Gray was observed chiefly from an anatomical point of view. The receptacle ofB. schreberi is rather flat and a vascular plexus is observable in the mature flower. The vasculature in this plexus is so complex taht it is not easy to trace its structure in detail. by observation on small buds, it can be seen that the receptacular vasculature consists of a girdling bundle in the basal area and usually nine receptacular strands from which traces to the petals and stamens branch off. The vasculature in the receptacle is reconstructed and diagramatically shown as though split longitudinally and spread out in one plane. Floral vasculature inCabomba caroliniana is simpler, and is probably related to the smaller number of stamens and carpels. It also has a girdling bundle at the bottom of receptacle and this vasculature is suggested to be derived by simplification from aBrasenia-type vasculature. Evidence from floral anatomy suggests that these two genera are closely related. InNymphaea, a vascular plexus in the receptacle is also observed (Moseley, 1961; Ito 1983). The plexus ofBrasenia andNymphaea are not the same in their construction. Nevertheless, their fundamental floral vasculature is comparable and it is preferable to place them in the same family or same order.     

The morphology and anatomy of the stigma of Petunia hybrida

Summary A mature stigma of Petunia hybrida ready for pollination shows 4-6 large shining drops of the exudate along with numerous smaller ones. A developing style and stigma have a columnar tissue that flares at the top. In the stigma there can be distinguished a secretory and a storage zone. In the former, schizogenous cavities are formed which are filled with the exudate. The mode of formation and secretion of the drop has been studied with light and electron microscope. The nature of reserves has been studied histochemically.The exudation takes place in two stages. In the first stage, the epidermal and papillae cells release out the oily exudate upon rupture of the cuticle. The second phase of exudation begins with anthesis. The exudate from the schizogenous cavities is released between the epidermal cells. There are distinct loci on the stigma surface where more exudate is given out than at other places.

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Zusammenfassung Bei der Durchsicht der einschlägigen Literatur ergab sich, daß die Kenntnisse über die Entwicklungsgeschichte und den Sekretionsmechanismus des Narben-Schleimes äußerst lückenhaft sind, obgleich dieser im Verlauf des Bestäubungs-Vorganges bei vielen Pflanzen eine bedeutende Rolle spielt. Die vorliegende Untersuchung versuchte daher eine detaillierte Einsicht in die Morphologie und Anatomie der Narbe von Petunia hybrida zu geben, da diese Art als Objekt für zahlreiche physiologische Untersuchungen auf dem Gebiet der Befruchtungs-inkompatibilität dient.Die Blüten von Petunia sind zwittrig, zygomorph und fünfzählig. Sie besitzen fünf epipetale Antheren mit drei verschiedenen Filament-Längen; diese öffnen sich zu verschiedenen Zeitpunkten. Die reife Narbe zeigt zum Zeitpunkt der Bestäubung 4-6 große sowie zahlreiche kleinere, transparente Flüssigkeits-Tropfen.Der sich entwickelnde Griffel mit der Narbe besteht aus einem säulenartigen Gewebe, das sich an der Spitze verbreitert. Bei der Narbe kann eine Sekretions-und eine Reservematerial-Zone unterschieden werden. In der Sekretionszone werden schizogen Hohlräume gebildet, die sich mit dem Exsudat füllen. Die Bildung und Sekretion der Narbenflüssigkeit wurde licht-und elektronenmikroskopisch untersucht. Mittels histochemischer Methoden wurden die Substanzen der Reservematerial-Zone näher charakterisiert.Die Freisetzung der Narben-Flüssigkeit findet in zwei Schritten statt: Während der ersten Phase wird das ölige Exsudat von den epidermalen Zellen und den Papillen der Narbe nach Zerreißen der Narben-Cuticula freigesetzt. In der zweiten Phase jedoch tritt das Exsudat aus den schizogenen Hohlräumen durch die Zwischenräume in der Narben-Epidermis aus. Auf der Narben-Oberfläche gibt es Bezirke, welche offensichtlich in reichlicherem Maße Flüssigkeit produzieren als andere Bezirke.

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Dedicated to the memory of the late Prof. P. Maheshwari FRS, our teacher (R. N. K.) and friend (H. F. L.) who read through the Ms in April at Paris, a month before his death.     

Floral morphology and anatomy of Pittosporum tobira (Pittosporaceae)

Comparative studies are made on floral morphology and anatomy of female and male flowers of Pittosporum tobira. The two types of flower differ little from each other in structure at the early stage of floral development, but appear dimorphic towards anthesis. The male flower becomes cryptically bisexual, although its pistil is slender compared to that of the female flower. The stigmas of the male flower are receptive and can induce pollen germination. The structure of the style in the male flower is identical to that in the female flower. Ovules are produced on the protruded parietal placenta in the male flower, but their development is arrested at the stage of the 4–nucleate embryo sac. The female flower is clearly unisexual, with obviously aborted and sagittate anthers. Its pistil is rather plump and can produce darkish red seeds immersed in sticky pulp. The male and female flowers are similar in vascular anatomy. A conspicuous difference between the two types of flower lies in the stamens. Variation of sexual organs in the genus Pittosporum is reviewed. We assume that the flowers of Pittosporum are derived from the hermaphrodite-flowered ancestor and the female flower has become unisexual through partial reduction of sexual organs at a faster rate than the male flower.