Comparative morphology of female flowers and systematics in Geonomeae (Arecaceae)

Female floral structure is compared in Geonomeae (Arecaceae). A perianth is formed by two alternate whorls of three basally congenitally united and imbricate sepals and three basally congenitally united and apically valvate petals. A sterile androecium is formed by a variable number of staminodes, which are united into a tube. The gynoecium shows three more or less equally developed carpels or is pseudomonomerous (Geonoma). The single anatropous ovule per carpel is median, either basal or at mid-height of the ovary. A septal nectary is present at the base and mid-height of the ovaries and exits at different levels of the ovary. Carpels in pseudomonomerous gynoecia seem to be basistylous, but the styles are more lateral or apical in gynoecia with all three carpels equally developed. Stigmas expose unicellular or multicellular (Welfia) papillae at anthesis. Pollen tube transmitting tracts and a compitum are present in the ventral slits of the postgenitally united styles. Floral structure in Geonomeae is compared with other Arecaceae, especially Arecoideae, in a morphological and systematic context.     

Expression of ethylene biosynthetic and receptor genes in rose floral tissues during ethylene-enhanced flower opening

Ethylene production, as well as the expression of ethylene biosynthetic (Rh-ACS1-4 and Rh-ACO1) and receptor (Rh-ETR1-5) genes, was determined in five different floral tissues (sepals, petals, stamens, gynoecia, and receptacles) of cut rose (Rosa hybrida cv. Samantha upon treatment with ethylene or the ethylene inhibitor 1-methylcyclopropene (1-MCP). Ethylene-enhanced ethylene production occurred only in gynoecia, petals, and receptacles, with gynoecia showing the greatest enhancement in the early stage of ethylene treatment. However, 1-MCP did not suppress ethylene production in these three tissues. In sepals, ethylene production was highly decreased by ethylene treatment, and increased dramatically by 1-MCP. Ethylene production in stamens remained unchanged after ethylene or 1-MCP treatment. Induction of certain ethylene biosynthetic genes by ethylene in different floral tissues was positively correlated with the ethylene production, and this induction was also not suppressed by 1-MCP. The expression of Rh-ACS2 and Rh-ACS3 was quickly induced by ethylene in gynoecia, but neither Rh-ACS1 nor Rh-ACS4 was induced by ethylene in any of the five tissues. In addition, Rh-ACO1 was induced by ethylene in all floral tissues except sepals. The induced expression of ethylene receptor genes by ethylene was much faster in gynoecia than in petals, and the expression of Rh-ETR3 was strongly suppressed by 1-MCP in all floral tissues. These results indicate that ethylene biosynthesis in gynoecia is regulated developmentally, rather than autocatalytically. The response of rose flowers to ethylene occurs initially in gynoecia, and ethylene may regulate flower opening mainly through the Rh-ETR3 gene in gynoecia.     

The cooling of convolvulaceous flowers in a tropical environment

The temperatures of flowers of Ipomoea pes‐caprae ssp. brasilensis, Ipomoea aquatica and Merremia borneensis in bright sunshine, were studied to determine the role of corollas and sepals in cooling the gynoecium. The corollas and sepals were prevented from transpiring by greasing, to investigate the extent of evaporative cooling. In the exposed natural habitats of these flowers the maximum temperatures of air and soil were high (32 and 42 °C, respectively) and corolla, sepal and gynoecium temperatures were often intermediate. Despite being almost astomatous, significant evaporative cooling was observed in the corolla. Between 20 and 80% of the energy absorbed by the corollas was dissipated as evaporation. The sepals were stomatous and their evaporative cooling was very important in reducing the temperature of the gynoecium. The temperatures of the non‐transpiring gynoecia and corollas were significantly higher than the temperatures of the normally transpiring corollas and gynoecia. Furthermore, the gynoecia temperatures were significantly higher with non‐transpiring corollas than with normally transpiring corollas, suggesting that the corollas alone play a role in maintaining the gynoecium within optimal temperatures levels. It was shown in an incubation experiment that temperatures exceeding 32 °C may damage the carpels, and temperatures exceeding 42 °C may damage sepals. Pollen grains were killed after 200 min of exposure to temperatures in the range 32 to 47 °C. It is concluded that the cooling mechanisms (evaporation and self‐shading) are critical for the reproductive success of these flowers in their natural environment.     

FLORAL ANATOMY OF KRAMERIA LANCEOLATA

The anatomy of each of the series of floral organs of Krameria lanceolata was examined. The sepals are characterized by three main veins each, an undifferentiated mesophyll, and stomata on the upper epidermis. The fleshy petals are distinguished by their numerous veins as well as by palisade-like epidermal cells on the outer surface. The three partially united petals have each a single vein and long, narrow epidermal cells similar to those on other floral organs. The stamens are united at their bases and bear tetra-sporangiate, conical anthers. The gynoecium includes a sterile and a fertile carpel. In the receptacle the veins to the sepals and petals are separated by a wide gap; those to the petals and stamens, by a narrow gap. Anatomical characteristics of the flower dissociate Krameriaceae from the legumes with which they have frequently been thought to be allied.     

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.     

Convergent elaboration of apocarpous gynoecia in higher advanced dicotyledons (Sapindales, Malvales, Gentianales)

The apocarpous gynoecia of three separate groups of higher advanced dicotyledons show postgenital fusion of their apical parts. In this fused region the pollen tube transmitting tissue of the carpels is united into a compitum, which provides advantages of a syncarpous to the apocarpous gynoecium. It is supposed that in at least some of these groups the general evolutionary trend of the angiosperms from apocarpy towards syncarpy is reversed.     

Comparative floral structure and systematics in Celastrales (Celastraceae, Parnassiaceae, Lepidobotryaceae)

Floral morphology, anatomy and histology in the newly circumscribed order Celastrales, comprising Celastraceae, Parnassiaceae and Lepidobotryaceae are studied comparatively. Several genera of Celastraceae and Lepidobotrys (Lepidobotryaceae) were studied for the first time in this respect. Celastraceae are well supported as a group by floral structure (including genera that were in separate families in earlier classifications); they have dorsally bulged‐up locules (and thus apical septa) and contain oxalate druses in their floral tissues. The group of Celastraceae and Parnassiaceae is also well supported. They share completely syncarpous gynoecia with commissural stigmatic lobes (and strong concomitant development of the commissural vascular bundles but weak median carpel bundles), only weakly crassinucellar or incompletely tenuinucellar ovules with an endothelium, partly fringed sepals and petals, protandry in bisexual flowers combined with herkogamy by the movement of stamens and anther abscission, and stamens fused with the ovary. In contrast, Lepidobotryaceae are more distant from the other two families, sharing only a handful of features with Celastraceae (not Parnassiaceae), such as pseudohermaphroditic flowers, united stamen bases forming a collar around the gynoecium and seeds with a conspicuous aril. However, all three families together are also somewhat supported as a group and share petals that are not retarded in late floral bud development, 3‐carpellate gynoecia, ventral slits of carpels closed by long interlocking epidermal cells and pollen tube transmitting tissue encompassing several cell layers, both integuments usually more than two cell layers thick, and only weak or lacking floral indumentum. In some molecular analyses Celastrales form an unsupported clade with Malpighiales and Oxalidales. This association is supported by floral structure, especially between Celastrales and Malpighiales. Among Celastrales, Lepidobotryaceae especially share special features with Malpighiales, including a diplostemonous androecium with ten fertile stamens, epitropous ovules with an obturator and strong vascularization around the chalaza. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149 , 129–194.     

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

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

Involvement of CUP-SHAPED COTYLEDON genes in gynoecium and ovule development in Arabidopsis thaliana

When mutations in CUP-SHAPED COTYLEDON1 (CUC1) and CUC2 are combined, severe defects involving fusion of sepals and of stamens occur in Arabidopsis flowers. In addition, septa of gynoecia do not fuse along the length of the ovaries and many ovules have their growth arrested. CUC2 is expressed at the tips of septal primordia during gynoecium development and at the boundary between nucellus and chalaza during ovule development. These expression patterns are partially consistent with the phenotype of the mutant gynoecium. CUC2 mRNA is also shown to be expressed at the boundaries between meristems and organ primordia during both the vegetative and reproductive phases. This expression pattern indicates that CUC2 is generally involved in organ separation in shoot and floral meristems.     

Impatiens angulata (Balsaminaceae), a new species from Guangxi, China

Impatiens angulata S. X. Yu, Y. L. Chen et H. N. Qin sp. nova (Balsaminaceae), a new species from Guangxi, China, is described and illustrated. This species is close to I. hainanensis in morphological characters . Both have succulent stems, 4 lateral sepal connected upper lobes of lateral united petals, but are distinguished by the base of stems with 6–9 ridges, leaves oblong or oblanceolate, the outer lateral sepals with 9 veins, inner lateral sepals ovate and dorsal sepal with deep bilobate spur.     

Ingenious floral structure drives explosive pollination in Hydrilla verticillata (Hydrocharitaceae)

• In explosive pollination, many structures and mechanisms have evolved to achieve high‐speed stamen movement. The male flower of the submerged plant Hydrilla verticillata is reported to be able to release pollen explosively some time after leaving the mother plant time, but the mechanism of stamen movement and the related functional structure in this species are unclear.
• In this study, we observed the male flower structure and pollen dispersal process of H. verticillata. We analysed the stamen movements during the pollen dispersal process and conducted several controlled experiments to study the process of storage and release of elastic potential energy in explosive pollination.
• When the male flower of H. verticillata is bound to the united bracts, the sepals accumulate elastic potential energy through the expansion of basal extensor cells. After the male flower is liberated from the mother plant, the stamens unfold rapidly with the sepals under adhesion and transfer the elastic potential energy to the filament in seconds. Once stamens unfold to a critical angle, at which the elasticity of the filament just exceeds the adhesion between sepals and anthers, the stamens automatically rebound and release pollen in milliseconds.
• These results reveal that Catapult‐like stamens, spoon‐shaped sepals and enclosed united bracts in the spathe together constitute the functional structure in rapid stamen movement of H. verticillata. They ensure that the pollen can be released on the water surface, and thus adapt successfully to the pollen‐epihydrophilous pollination.

     

DIFFERENTIAL SENSITIVITY OF “DOUBLE” AND “SINGLE” FLOWERS OF NIGELLA DAMASCENA (RANUNCULACEAE) TO EMASCULATION AND TO GA3

In Nigella damascena “double” flower is inherited as a single-gene recessive to “single” flower. Early emasculation of “double” flowers greatly inhibits gynoecium development and, to a lesser degree, development of sepals and bracts. No comparable inhibition of gynoecia was induced in “single” flowers though some sepal and bract inhibition was detected. A differential nutritional requirement for organ initiation on cultured flower apices was also detected. Apices of both varieties initiated stamens and carpels if kinetin was added to the basal medium. However, in the absence of kinetin, GA₃ was required for organ initiation in “doubles” but completely inhibited stamen initiation in “singles.” Both these differential effects are thought to reflect rather different patterns of gibberellin metabolism in the two genetic strains.     

Comparative floral structure and systematics in Chrysobalanaceae s.l. (Chrysobalanaceae, Dichapetalaceae, Euphroniaceae, Trigoniaceae; Malpighiales)

Chrysobalanaceae s.l. , one of the few suprafamilial subclades of Malpighiales that is supported by molecular phylogenetic analyses, and containing Chrysobalanaceae, Dichapetalaceae, Euphroniaceae, and Trigoniaceae, was comparatively studied with regard to floral structure. The subclade is well supported by floral structure. Potential synapomorphies for Chrysobalanaceae s.l. are the following shared features: floral cup; flowers obliquely monosymmetric; sepals congenitally united at base; sepals of unequal size (outer two shorter); fertile stamens concentrated on the anterior side of the flower and sometimes united into a strap; staminodes absent in the posteriormost antepetalous position; anthers extremely introrse, with thecae almost in one plane; endothecium continuous over the dorsal side of the connective; dorsal anther pit; gynoecium completely syncarpous up to the stigma; carpel flanks slightly bulged out transversely and thus carpels demarcated from each other by a longitudinal furrow; flowers with dense unicellular, non-lignified hairs, especially on the gynoecium; light-coloured, dense indumentum on young shoots and inflorescences. Potential synapomorphies for Chrysobalanaceae + Euphroniaceae include: spur in floral cup; clawed petals; lignified hairs on petals; nectary without lobes or scales and mostly annular. Potential synapomorphies for Dichapetalaceae + Trigoniaceae include: special mucilage cells in sepals in mesophyll (in addition to epidermis); anthers almost basifixed; gynoecium synascidiate up to lower style; nectary with lobes or scales and semi-annular.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 249–309.     

FLOWER DEVELOPMENT IN DIOECIOUS SPINACIA OLERACEA (CHENOPODIACEAE)

Spinacia oleracea (Chenopodiaceae) is a potential model system for studies of mechanisms of sex expression and environmental influences on gender in dioecious species. Development of the male and female flowers and inflorescences of spinach were studied to determine when the two sex types can be distinguished. We found that female inflorescence apices are significantly larger than those of the male. Flower primordia are similar in size prior to perianth initiation, but the male primordia develop at a faster rate. Another distinguishing feature at this early stage is the larger bract subtending the female primordium. The two flower types become readily distinguishable when the perianth initiates. Male flowers produce four sepals and four stamens in a spiral pattern in close succession. Female flowers produce two alternate perianth parts that enlarge somewhat before the gynoecium becomes visible. There are no traces of gynoecia in male flowers or of stamens in female flowers. We propose that plant sex type is determined before inflorescence development, prior to or at evocation.     

INFLORESCENCE AND FLORAL DEVELOPMENT IN HOUTTUYNIA CORDATA (SAURURACEAE)

The inflorescence of Houttuynia cordata produces 45–70 sessile bracteate flowers in acropetal succession. The inflorescence apical meristem has a mantle-core configuration and produces “common” or uncommitted primordia, each of which bifurcates to form a floral apex above, a bract primordium below. This pattern of organogenesis is similar to that in another saururaceous plant, Saururus cernuus. Exceptions to this unusual development, however, occur in H. cordata at the beginning of inflorescence activity when four to eight petaloid bract primordia are initiated before the initiation of floral apices in their axils. “Common” primordia also are lacking toward the cessation of inflorescence apical activity in H. cordata when primordia become bracts which may precede the initiation of an axillary floral apex. Many of these last-formed bracts are sterile. The inflorescence terminates with maturation of the meristem as an apical residuum. No terminal flowers or terminal gynoecia were found, although subterminal gynoecia or flowers in subterminal position may overtop the actual apex and obscure it. Individual flowers have a tricarpellate syncarpous gynoecium and three stamens adnate to the carpels; petals and sepals are lacking. The order of succession of organs is: two lateral stamens, median stamen, two lateral carpels, median carpel. The three carpel primordia almost immediately are elevated as part of a gynoecial ring by zonal growth of the receptacle below the attachment of the carpels. The same growth elevates the stamen bases so that they appear adnate to the carpels. The trimerous condition in Houttuynia is the result of paired or solitary initiations rather than trimerous whorls. Symmetry is bilateral and zygomorphic rather than radial. No evidence of spiral arrangement in the flower was found.     

Multicarpellate gynoecia in angiosperms: occurrence,development, organization and architectural constraints

Most angiosperms have gynoecia with two to five carpels. However, more than five carpels (here termed ‘multicarpellate condition’) are present in some representatives of all larger subclades of angiosperms. In such multicarpellate gynoecia, the carpels are in either one or more than one whorl (or series). I focus especially on gynoecia in which the carpels are in a single whorl (or series). In such multicarpellate syncarpous gynoecia, the closure in the centre of the gynoecium is imprecise as a result of slightly irregular development of the carpel flanks. Irregular bumps appear to stuff the remaining holes. In multicarpellate gynoecia, the centre of the remaining floral apex is not involved in carpel morphogenesis, so that this unspent part of the floral apex remains morphologically undifferentiated. It usually becomes enclosed within the gynoecium, but, in some cases, remains exposed and may or may not form simple excrescences. The area within the remaining floral apex is histologically characterized by a parenchyma of simple longitudinal cell rows. In highly multicarpellate gynoecia with the carpels in a whorl, the whorl tends to be deformed into an H‐shaped or star‐shaped structure by differential growth of the floral sectors, so that carpels become aligned in parallel rows, in which they face each other with the ventral sides. In this way, a fractionated compitum may still be functional. Multicarpellate gynoecia (with the carpels in one whorl or series) occur in at least one species in 37 of the 63 angiosperm orders. In contrast, non‐multicarpellate gynoecia are present in at least one species of all 63 orders. The basal condition in angiosperms is more likely non‐multicarpellate. Multicarpellate gynoecia are restricted to flowers that are not highly synorganized. In groups with synorganized androecium and gynoecium and in groups with elaborate monosymmetric flowers, multicarpellate gynoecia are lacking. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 1–43.     

Ontogeny and morphology of the fertile flowers of Hydrangea and allied genera of tribe Hydrangeeae (Hydrangeaceae)

The monophyletic Hydrangeeae (Hydrangeaceae) consists of the clade Cardiandra + Deinanthe and its sister, the Hydrangea clade, which includes the paraphyletic Hydrangea as well as Broussaisia, Decumaria, Dichroa, Pileostegia, Platycrater and Schizophragma. The plesiomorphic imbricate corolla aestivation and polystemony of Cardiandra and Deinanthe distinguish these two genera from most members of the Hydrangea clade. Deinanthe has postgenitally fused styles and cushion-like, dorsally positioned stigmas. The Hydrangea clade is notable because most species of Hydrangea share a floral morphology characterized by small size; tetramerous-pentamerous perianths; inconspicuous sepals; reflexed petals; diplostemony; stamens that are longer than the styles; completely inferior ovaries; separate styles; terminal, papillate stigmas; and dimerous-tetramerous gynoecia. This suite of states is termed the 'Hydrangea floral syndrome' (HFS). Various members of the Hydrangea clade lack the HFS, including (1) Platycrater; (2) Hydrangea anomala; (3) H. paniculata + H. heteromalla; (4) the Schizophragma clade (Schizophragma, Pileostegia and Decumaria); and (5) the macrophylla clade (H. macrophylla, H. scandens, H. hirta, Dichroa and Broussaisia). The meristic uniqueness of Decumaria reflects mutations observed in Arabidopsis (clavata) and Lycopersicon (fasciated) that cause organ number increases because of changes in meristem capital. The modification of early perianth development to form a prominent corolline torus at a point when sepals are diminutive is present in H. anomala and Hydrangea section Cornidia and may be synapomorphic for them. Various transformations in the perianth, androecium, and gynoecium lie behind the floral diversity of Hydrangeeae. Some morphological transformations have been homoplastic, including shifts to polystemony, calyptrate corollas, and synstyly.     

Homeotic, Meristic and Cytological Floral Mutants of Thryptomene calycina (family Myrtaceae)

Twenty plants with various phenotypic abnormalities to the flowerswere selected from very large populations of Thryptomene calycinain the Grampian and Black Ranges. Most of these had impairedreproductive function. Normal flowers were epigynous with fivesepals, five petals, five anthers, a single style and two anatropousovules. The mutants were two partially male sterile, tetraploidplants with large flowers, one of which occasionally producedadditional flowers from the leaf axils with peduncles as wellas pedicels; one plant which produced a proportion of hexapetaloidflowers with six stamens; three gross mutants with fleshy, bracteoidpointed petals and sepals, no stamens, vestigial styles andstigmas, exposed ovules and no inferior ovary; one plant withfleshly, bracteoid pointed sepals, vestigial style and stigmabut with exposed ovular structures replaced by four to fivesterile ovules generally inside an abnormal ovary; two plantswith reduced ovary diameter and sterile ovules, shortened style,five reduced sepals and petals and five to eight anthers; threeanthocyanin-free plants; three plants with pink sepals; twoplants with half-sized flowers which produced a proportion offasciated stems; one plant which occasionally produced flowerswithout pedicels which virtually resulted in organs which wereleaf-flower composites; two plants which produced sepals andpetals which contained chlorophyll and prematurely senesced,and had partial substitution of petals by anthers.Copyright1993, 1999 Academic Press Thryptomene calycina, Myrtaceae, Victorian lace flower, floral mutations, mutants, homeotic, meristic, tetraploid, fasciation, male sterility, cut flowers     

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.