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.     

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.     

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

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

Evaluation of the floral vasculature of the Janusia,Mascagnia and Tetrapterys species as a tool to explain the decrease of floral organs in Malpighiaceae

The family Malpighiaceae is considered monophyletic, but the intra-family classification is conflicting. Analyses of floral vasculature allow the identification of reductions, connations and adnations and can even reveal evolutionary steps prior to current floral morphology. The present work analysed the floral vasculature of Janusia mediterranea, Mascagnia cordifolia and Tetrapterys chamaecerasifolia using material processed by traditional methods for light microscopy. A general pattern was observed of three bundle traces supplying each sepal and one trace per petal and stamen; Mascagnia is an exception, as its eglandular sepal has only a median trace but shares lateral traces with adjacent sepals. No dorsal traces are emitted to the carpels; however, three intercarpellary complexes are emitted that divide into six ventral bundles, supplying the ovule. Mascagnia demonstrates connation between the anterior and adjacent sepal glands; reductions of the anterior sepal glands were registered in Tetrapterys and Janusia. This work reveals two distinct processes for gland loss in non-related groups of the family that resulted in similar present appearances. Our evaluation of the number of calyx glands and the processes of glandular loss in species with less than ten glands improves our understanding of the evolution of calyx glands in Malpighiaceae.     

Floral vascular anatomy ofConvallaria majalis L. andC. keiskei miq. (liliaceae-convallariinae)

In comparing the floral vascular anatomy ofConvallaria majalis andC. keiskei a similar pattern of vasculature was shown. Both have pedicels with six (3 large +3 small) bundles which via radial division and fusion form the tepal, stamen and ovary traces. The outer tepal and outer stamen traces, the dorsals and placentals (i.e. ventral supply) arise from the larger three pedicel bundles, while the inner tepal and inner stamen traces and the septal axials arise from the smaller three. The dorsals, septal axials, and all of the stamen and tepal bundles are fusion products, while the placentals are free, though arising from compound bundles. The overy vasculature lacks both lateral peripherals and terminal cross-connections between the inner bundles and the outer dorsals. The placentation is only axile basally, since the three septa are freed at the mid-ovary level, and the resulting common, upper carpellary cavity is continuous with the hollow style. Normally four ovules are observed in each carpel, with the lower tier associated with the lower solid central axis, and the upper tier associated with the freed septa. The orientation of the ovules is varied (heterotropic). An internal system of stigmatoidal tissue is continuous from the base of each locule to the stigma, and involves micropylar associated obturators. Raphides characterize mature ovaries of both species, though both lack septal glands and septal grooves.     

大钟花属和黄秦艽属花部解剖

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

MORPHOLOGICAL STUDIES OF THE NYMPHAEACEAE SENSU LATO. XVII. FLORAL ANATOMY OF ONDINEA PURPUREA SUBSPECIES PURPUREA (NYMPHAEACEAE)

Classification and phylogeny of the Nymphaeaceae are unresolved. This study provides floral anatomical data that will assist in elucidating generic interrelationships and systematic relationships to other taxa of angiosperms. The floral anatomy of Ondinea purpurea den Hartog subsp. purpurea has been examined utilizing light microscopy. The peduncle possesses stelar vascular bundle complexes and cortical vascular bundles. Cortical bundles terminate within the peduncle. Each bundle complex consists of 2 collateral bundles on the same radius, the inner bundle inverted; 2 protoxylary lacunae occur yet differ in structure and function. Progressing acropetally, the inner xylary lacunae become discrete mesarch strands surrounded centrifugally by a vascular cylinder formed by divisions and anastomosing of the bundle complexes. Together these become the massive receptacular vascular plexus. The plexus provides collateral traces to the floral organs. Each sepal receives 3 traces that separate from the plexus as 1–3 lateral traces. Petals are absent and no vestigial petal traces have been observed. Distally, the plexus forms several large strands of connate gynoecial and androecial traces termed the principal vascular bundles (PVBs). Ventral veins separate from the PVBs and the latter extend acropetally through the outer ovary wall. Branches of the ventrals and PVBs contribute to septal vascular reticula from which each ovule is supplied by one vascular bundle. Each stamen receives 1 trace from branches of the PVBs. The ventrals and PVBs terminate within the carpellary lobes. A comparative anatomical study is offered that supports the inclusion of Ondinea in the Nymphaeaceae sensu stricto.     

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.     

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 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.     

MORPHOLOGICAL STUDIES OF THE NYMPHAEACEAE (SENSU LATO). XIII. CONTRIBUTIONS TO THE VEGETATIVE AND FLORAL STRUCTURE OF CABOMBA

Six species of Cabomba have been examined although the anatomy of the vegetative axes is based on the study of only C. caroliniana and C. palaeformis. A plant consists of an erect short shoot with decussate leaves which bears axillary flowering shoots and rhizomes. A rhizome bears decussate leaves and may also form axillary flowering shoots or turn upward and become a new short shoot. The phyllotaxies of the flowering shoots are proximately decussate or ternate (C. piauhyensis). The flowering shoots with decussate phyllotaxy change to 1/3 phyllotaxy distally; they bear axillary flowers proximally, and extra-axillary flowers distally. Flowering shoots with ternate phyllotaxy do not change distally but each produces first axillary and then extra-axillary flowers. Decussate vegetative axes and flowering shoots have four vascular bundles; ternate vegetative axes and flowering shoots have six vascular bundles, distantly paired into two or three vascular bundle-pairs, respectively. An elliptical vascular plexus occurs at each node. Each leaf receives one bundle-pair from one trace and each flower three bundle-pairs. A two-level receptacular vascular plexus occurs in flowers; the proximal, larger portion provides traces to perianth and stamens and the distal, smaller portion becomes carpellary traces. Each of the three sepals typically receives five branch traces from a basal principal trace, and each of the three petals receives, typically, three branch traces from a basal principal trace. Sepals and petals generally occur in a single, basally connate whorl. Each stamen receives one trace. Each stamen of three-stamen flowers is opposite a petal; each stamen of six-stamen flowers is aligned with an interval between a petal and adjacent sepal. Each staminal trace, which is just above the principal petal trace, in a three-petal flower, is frequently adnate to the latter trace. Each carpel receives one principal trace from the distal, small extension of the receptacular plexus, and each principal trace becomes three conventional veins of a carpel. Ovules may be borne directly over one of the veins or in any position between veins and are supplied by branches of the nearest vein or nearest two veins. All traces, ovular supply veins and the proximal portions of all veins are amphicribral. The several anatomical and morphological differences in vegetative axes and flowers between Cabomba and Brasenia suggest a greater taxonomic distance between the two genera than commonly supposed. It is suggested that extra-axillary flowers in 1/3 helical and ternate flowering shoots of Cabomba might be advantageous in preventing anthesis of flowers beneath peltate leaves. The aberrant position might be the initial evolutionary step toward what, in other nymphaeaceous genera, has shifted each flower to an adjacent helix. It is proposed that the zigzag stem accompanying the trigonal and sympodial flowering shoots may offer greater stability and floatability in water than the monopodial form. Several suggestions are offered for the variability of ovular positions: 1) the variability is a vestige of former laminar placentation in conduplicate carpels; 2) it is a vestige of a primitive condition antedating the current close association of ovules with ventral carpellary veins; 3) it is an early stage of evolution which might have terminated in laminar placentation and cantharophily, but which was replaced by a trend toward myophily.     

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.     

Morphological studies on the genusClematis Linn

The basic pattern of the vascular supply to stamens and carpels in the flowers ofClematis is discussed on the basis of serial sections. The bundles of the receptacular stele show fairly regular fusions and divisions in relation to the origin of the vascular supply, giving off a single trace for each stamen or carpel. In many cases the trace arises by the trifurcation of the “fused bundle” and the subsequent departure of the median strand. This is the pattern basic to the structure of the receptacular stele of the genus. Although the basic pattern involves a variety of modifications, each of the diverging traces fundamentally leaves a single independent gap in the stele, contrary to the conclusion of previous authors. Similarities and differences between a group of stamens and carpels and that of sepals and foliage leaves are also discussed based on the results of the present and previous studies on the vascular anatomy of the floral receptacle and the inflorescence axis.     

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.     

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.     

Floral vasculature in Alpinia hainanensis in relation to the nature of the labellum in gingers

Observations presented here on floral vasculature in Alpinia hainanensis indicate that the labellum incorporates elements of five androecial members rather than two or three, as suggested by previous authors for Zngiberaceae flowers. The pedicel contains an outer ring and a central region of vascular bundles. Three carpellary dorsal bundles (CDs) and three alternatively arranged parietal bundles (PBs) separate from the central region successively. The remaining bundles of the central region run upwards and become the placental bundles to supply ovules. The placental bundles terminate between the top of the locular region and the base of the prolongation. The three PBs divides into about five strands respectively. Of which the outer strand enters into the petal being its midrib and the remaining strands move into the stamen adaxially being the vasculature of the functional stamen and the labellum abaxially being the lateral strands of the labellum. The three CDs divide into about five traces, of which the outer strand becomes the midrib of each sepal and the inner strand runs into the style. The remaining traces re‐unite, re‐divide again in the course up and the two adaxial sets of carpellary dorsals finally enter into the labellum being the marginal traces of it while the abaxial single strand enters into the labellum being its midrib. The two antero‐lateral glands receive small traces without lignified tube elements from the vascular plexus, which fonn in prolongation from both PBs and CDs and a few small strands in the ovary wall. There are no subulate appendages differentiated in the flower of Alpinia hainanensis. Hereby, the median of the sepals, both the marginal portions and the median of labellum, and the style have the same origin in vasculature from the CDs and so do the stamen, the lateral portions of labellum and the median of the petals from PBs. The labellum is supposed to represent three members of the outer androecial whorl by its two marginal portions and the median and two members of the inner whorl by its two lateral parts except the median.     

COMPARATIVE ANATOMY AND SYSTEMATICS OF WOODY SAXIFRAGACEAE: TETRACARPAEA

Vegetative and reproductive anatomy and morphology are described for the first time for Tetracarpaea tasmannica Hook., a small shrub endemic to Tasmania. Tetracarpaea, a monotypic genus, has many characteristics of other woody Saxifragaceae, such as wood with solitary pores, scalariform perforation plates, sparse axial xylem parenchyma, tracheids, spiral thickenings in tracheary elements, and perforated ray cells. The tracheary elements of Tetracarpaea are much smaller than those characteristic of the Escallonioideae, a feature probably related to its montane forest habitat. Other features of Tetracarpaea inconsistent with most Escallonioideae include dark-staining deposits in the ray cells; a unilacunar, one-trace nodal pattern; lack of unicellular foliar trichomes; simple craspedodromous venation; areole development that is lacking or incomplete; straight and tapered veinlets; abaxial fibers associated with the foliar vascular bundles; and lack of bundle sheaths. The genus is further characterized by complete, hypogynous, tetramerous flowers. The essentially apocarpous gynoecium has multiovulate carpels, each supplied by three veins that reach the stigma. Ovules are anatropous, bitegmic, and crassinucellate. Lateral sepal bundles are derived either from the sepal midrib or from the petal-plane bundles; stamens are supplied by independent traces or by bundles originating from compound traces in both sepal- and petal-planes. The follicular fruits possess a sclerenchymatous endocarp and contain winged seeds that have a membranous testa, a ridged surface, and a cellular endosperm. Reproductive morphological and anatomical features are more consistent with features of the Saxifragoideae than with the Escallonioideae or the Cunoniaceae, although the essentially apocarpous gynoecium with multiovulate carpels is not found in these groups. Vegetative and reproductive characteristics indicate that Tetracarpaea is more closely related to Saxifragaceae than to Cunoniaceae. It is possible this isolated genus should have separate familial status.