Synorganisation without organ fusion in the flowers of Geranium robertianum (Geraniaceae) and its not so trivial obdiplostemony

Background and Aims Synorganisation of floral organs, an important means in angiosperm flower evolution, is mostly realized by congenital or post-genital organ fusion. Intimate synorganisation of many floral organs without fusion, as present in Geranium robertianum, is poorly known and needs to be studied. Obdiplostemony, the seemingly reversed position of two stamen whorls, widely distributed in core eudicots, has been the subject of much attention, but there is confusion in the literature. Obdiplostemony occurs in Geranium and whether and how it is involved in this synorganisation is explored here. Methods Floral development and architecture were studied with light microscopy based on microtome section series and with scanning electron microscopy. Key Results Intimate synorganisation of floral organs is effected by the formation of five separate nectar canals for the proboscis of pollinators. Each nectar canal is formed by six adjacent organs from four organ whorls. In addition, the sepals are hooked together by the formation of longitudinal ribs and grooves, and provide a firm scaffold for the canals. Obdiplostemony provides a guide rail within each canal formed by the flanks of the antepetalous stamen filaments. Conclusions Intimate synorganisation in flowers can be realized without any fusion, and obdiplostemony may play a role in this synorganisatio

Carpels in Brasenia (Cabombaceae) are Completely Ascidiate Despite a Long Stigmatic Crest

• Background and Aims The morphological structure of anthetic carpels of Brasenia (Cabombaceae), a member of the phylogenetically basal ANITA grade, has not been studied before. The carpel has a long stigmatic crest on the ventral side and could give the impression of a conduplicate structure. This is in contrast to the carpel structure in other genera of the ANITA grade. Therefore, a study of carpel development and carpel structure at anthesis was carried out. • Methods Carpels of Brasenia schreberi were studied at different developmental stages up to anthesis by means of microtome section series and SEM to analyse and reconstruct the outer and inner carpel morphology. • Key Results Carpels of Brasenia are extremely ascidiate up to anthesis. The elongate stigma originates around the mouth of the young carpel, which is slightly curved toward the centre of the flower. Subsequently, the stigmatic zone below the mouth expands by massive intercalary elongation. • Conclusions In their ascidiate shape, carpels of Brasenia are similar to carpels of Cabomba, the other genus of Cabombaceae, which, in contrast, has a short stigma restricted to the tip of the carpel. Thus, the morphological structure is independent of the extent (and one-sidedness) of the stigma. The outer shape of carpels at anthesis does not allow the inference of the inner morphological surface. If an angiosperm carpel has a one-sided stigma it can be extremely conduplicate or extremely ascidiate. Therefore, caution has to be used in the interpretion of the structure of fossil carpel

Angiosperm ovules: diversity, development, evolution

Background Ovules as developmental precursors of seeds are organs of central importance in angiosperm flowers and can be traced back in evolution to the earliest seed plants. Angiosperm ovules are diverse in their position in the ovary, nucellus thickness, number and thickness of integuments, degree and direction of curvature, and histological differentiations. There is a large body of literature on this diversity, and various views on its evolution have been proposed over the course of time. Most recently evo-devo studies have been concentrated on molecular developmental genetics in ovules of model plants. Scope The present review provides a synthetic treatment of several aspects of the sporophytic part of ovule diversity, development and evolution, based on extensive research on the vast original literature and on experience from my own comparative studies in a broad range of angiosperm clades. Conclusions In angiosperms the presence of an outer integument appears to be instrumental for ovule curvature, as indicated from studies on ovule diversity through the major clades of angiosperms, molecular developmental genetics in model species, abnormal ovules in a broad range of angiosperms, and comparison with gymnosperms with curved ovules. Lobation of integuments is not an atavism indicating evolution from telomes, but simply a morphogenetic constraint from the necessity of closure of the micropyle. Ovule shape is partly dependent on locule architecture, which is especially indicated by the occurrence of orthotropous ovules. Some ovule features are even more conservative than earlier assumed and thus of special interest in angiosperm macrosystematic

Floral development and floral phyllotaxis in Anaxagorea (Annonaceae)

Background and Aims Anaxagorea is the phylogenetically basalmost genus in the large tropical Annonaceae (custard apple family) of Magnoliales, but its floral structure is unknown in many respects. The aim of this study is to analyse evolutionarily interesting floral features in comparison with other genera of the Annonaceae and the sister family Eupomatiaceae. Methods Live flowers of Anaxagorea crassipetala were examined in the field with vital staining, liquid-fixed material was studied with scanning electron microscopy, and microtome section series were studied with light microscopy. In addition, herbarium material of two other Anaxagorea species was cursorily studied with the dissecting microscope. Key Results Floral phyllotaxis in Anaxagorea is regularly whorled (with complex whorls) as in all other Annonaceae with a low or medium number of floral organs studied so far (in those with numerous stamens and carpels, phyllotaxis becoming irregular in the androecium and gynoecium). The carpels are completely plicate as in almost all other Annonaceae. In these features Anaxagorea differs sharply from the sister family Eupomatiaceae, which has spiral floral phyllotaxis and ascidiate carpels. Flat stamens and the presence of inner staminodes differ from most other Annonaceae and may be plesiomorphic in Anaxagorea. However, the inner staminodes appear to be non-secretory in most Anaxagorea species, which differs from inner staminodes in other families of Magnoliales (Eupomatiaceae, Degeneriacae, Himantandraceae), which are secretory. Conclusions Floral phyllotaxis in Anaxagorea shows that there is no signature of a basal spiral pattern in Annonaceae and that complex whorls are an apomorphy not just for a part of the family but for the family in its entirety, and irregular phyllotaxis is derived. This and the presence of completely plicate carpels in Anaxagorea makes the family homogeneous and distinguishes it from the closest relatives in Magnoliale

Comparative floral structure and systematics in Apodanthaceae (Rafflesiales)

Abstract.: Comparative studies on floral morphology, anatomy, and histology were performed to identify shared features of the genera of Apodanthaceae (Rafflesiales): Apodanthes, Pilostyles, and Berlinianche. Berlinianche was studied for the first time in detail and its affinity to Apodanthaceae was confirmed. It has a previously undescribed hair cushion on the inner perianth organs and inaperturate pollen. Shared features of members of Apodanthaceae are: unisexual flowers; three (or four) alternating di-/tetra- or tri-/hexamerous whorls of scales of which the inner one or two correspond to a perianth; a synandrium with pollen sacs typically arranged in two rings; opening by a dehiscence line between the two rings of pollen sacs; large vesicular hairs above the synandrium; a gynoecium with four united carpels; inferior and unilocular ovaries with four parietal placentae, ovules tenuinucellate, anatropous with two well developed integuments, oriented in various directions; a nectary disk. Apodanthaceae share some special structural features with Malvale

Advances in the floral structural characterization of the major subclades of Malpighiales, one of the largest orders of flowering plants

Background and Aims Malpighiales are one of the largest angiosperm orders and have undergone radical systematic restructuring based on molecular phylogenetic studies. The clade has been recalcitrant to molecular phylogenetic reconstruction, but has become much more resolved at the suprafamilial level. It now contains so many newly identified clades that there is an urgent need for comparative studies to understand their structure, biology and evolution. This is especially true because the order contains a disproportionally large diversity of rain forest species and includes numerous agriculturally important plants. This study is a first broad systematic step in this endeavour. It focuses on a comparative structural overview of the flowers across all recently identified suprafamilial clades of Malpighiales, and points towards areas that desperately need attention. Methods The phylogenetic comparative analysis of floral structure for the order is based on our previously published studies on four suprafamilial clades of Malpighiales, including also four related rosid orders (Celastrales, Crossosomatales, Cucurbitales, Oxalidales). In addition, the results are compiled from a survey of over 3000 publications on macrosystematics, floral structure and embryology across all orders of the core eudicots. Key Results Most new suprafamilial clades within Malpighiales are well supported by floral structural features. Inner morphological structures of the gynoecium (i.e. stigmatic lobes, inner shape of the locules, placentation, presence of obturators) and ovules (i.e. structure of the nucellus, thickness of the integuments, presence of vascular bundles in the integuments, presence of an endothelium in the inner integument) appear to be especially suitable for characterizing suprafamilial clades within Malpighiales. Conclusions Although the current phylogenetic reconstruction of Malpighiales is much improved compared with earlier versions, it is incomplete, and further focused phylogenetic and morphological studies are needed. Once all major subclades of Malpighiales are elucidated, more in-depth studies on promising structural features can be conducted. In addition, once the phylogenetic tree of Malpighiales, including closely related orders, is more fully resolved, character optimization studies will be possible to reconstruct evolution of structural and biological features within the orde

Floral Morphogenesis in Euptelea (Eupteleaceae, Ranunculales)

Background and Aims Based on molecular phylogenetic studies, the unigeneric family Eupteleaceae has a prominent phylogenetic position at or near the base of Ranunculales, which, in turn, appear at the base of eudicots. The aim of the present paper is to reveal developmental features of the flowers and to put the genus in a morphological context with other basal eudicots. Methods Flowers in all developmental stages of Euptelea pleiosperma were collected in the wild at intervals of 7-10 d in the critical stages and studied with a scanning electron microscope. Key Results Remnants of a perianth are lacking throughout flower development. Floral symmetry changes from monosymmetric to asymmetric to disymmetric during development. Asymmetry is expressed in that the sequence of stamen initiation is from the centre to both lateral sides on the adaxial side of the flower but starting from one lateral side and proceeding to the other on the abaxial side. Despite the pronounced floral disymmetry, a dimerous pattern of floral organs was not found. The carpel primordia arise between the already large stamens and alternate with them. Stamens and carpels each form a somewhat irregular whorl. The carpels are ascidiate from the beginning. The stigma differentiates as two crests along the ventral slit of the ovary. The few lateral ovules alternate with each other. Conclusions Although the flowers have some unusual autapomorphies (wind pollination, lack of a perianth, pronounced disymmetry of the floral base, long connective protrusion, long temporal gap between androecium and gynoecium initiation, small space for carpel initiation), they show some plesiomorphies at the level of basal eudicots (free carpels, basifixed anthers, whorled phyllotaxis), and thus fit well in Ranunculale

Fusion within and between whorls of floral organs in Galipeinae (Rutaceae): structural features and evolutionary implications

Background and Aims Most genera of the neotropical Galipeinae (tribe Galipeeae, Rutoideae) exhibit several forms and degrees of fusion between the floral organs, including the union of petals into an apparently sympetalous corolla, the joining of the stamens among themselves and to the corolla, and the partial to complete connation of carpels. Though these and others floral traits are currently used in the circumscription of species in Galipeinae, few studies have shown in detail in which way (postgenital or congenital) and to what extent these fusions occur. To elucidate these anatomical conditions, a structural study of the flowers of the Galipeinae species was carried out. Methods Flowers of six species from three genera of Galipeinae were studied in their morphology, anatomy and development with stereomicroscopy, light microscopy and scanning electron microscopy (SEM). Key Results The floral tube is formed by synorganization of stamens with petals in all species, and exhibits three main patterns: (1) Conchocarpus heterophyllus and C. minutiflorus have a floral tube formed by marginal coherence/adherence of petals and filaments due to interwining trichomes (postgenital connection); (2) Erythrochiton brasiliensis has a tube formed by congenital fusion of petals and filaments; and (3) Galipea jasminiflora and Conchocarpus macrophyllus have a tube formed distally with the first pattern, and proximally with the second pattern. Although floral tubes seem to be homologous within Galipeinae, this is not true at the level of the family: the floral tube of Correa (from an only distantly related clade of the family) is formed by postgenital union of the petals representing a convergent structure. The gynoecium of the studied species of Galipeinae shows a great variability in the extent of fusion of carpel flanks. Even though different structures for the mature gynoecium were found in each genus, all genera show postgenitally fused carpel apices, which is related to the formation of a compitum, as described earlier for other members of Rutaceae. Conclusions The degree and diversity of fusions of floral organs in Galipeinae is unique within the order Sapindales. A study of the amount of diversification of Galipeinae in South America and comparison with other clades of Rutaceae would be of interes

Heterodichogamy in Kingdonia (Circaeasteraceae, Ranunculales)

Background and Aims Preliminary field observations in 2001 and 2002 suggested that Kingdonia uniflora (Circaeasteraceae, Ranunculales) exhibits heterodichogamy, an unusual kind of reproductive heteromorphy, hitherto unreported in Ranunculales and known from only one other genus in basal eudicots. Methods During several subsequent years flowers were observed in the field. Flowers were fixed in FAA and studied with microtome sections series and with the scanning electron microscope. Key Results The flowers proved to be heterodichogamous, with protandrous and protogynous morphs, which have a 1 : 1 ratio. Both morphs equally set fruit. Each year a single flower is formed at the tip of a rhizome or more rarely two flowers. The flowers are already open when they appear at the soil surface, before they are receptive and before pollen is dispersed. In both floral morphs the styles elongate early and the stigmas are positioned above the anthers before anthesis begins. In protogynous flowers the stigmas become receptive in this position; later the styles become reflexed and then the anthers dehisce. In contrast, in protandrous flowers the stamen filaments elongate during early anthesis such that the dehiscing anthers come to lie above the (still unreceptive) stigmas; after dehiscence of all anthers in a flower the styles begin to elongate and become receptive. Conclusions This is the first record of heterodichogamy in a representative of Ranunculales, in an herbaceous eudicot, and in a plant with uniflorous ramets. The occurrence of heterodichogamy in Kingdonia in which clonal reproduction appears to be dominant might be an adaptation to avoid mating between the ramets from a common mother individual (genet