Comparative floral structure and systematics in Rhizophoraceae,Erythroxylaceae and the potentially related Ctenolophonaceae,Linaceae, Irvingiaceae and Caryocaraceae (Malpighiales)

Within the rosid order Malpighiales, Rhizophoraceae and Erythroxylaceae (1) are strongly supported as sisters in molecular phylogenetic studies and possibly form a clade with either Ctenolophonaceae (2) or with Linaceae, Irvingiaceae and Caryocaraceae (less well supported) (3). In order to assess the validity of these relationships from a floral structural point of view, these families are comparatively studied for the first time in terms of their floral morphology, anatomy and histology. Overall floral structure reflects the molecular results quite well and Rhizophoraceae and Erythroxylaceae are well supported as closely related. Ctenolophonaceae share some unusual floral features (potential synapomorphies) with Rhizophoraceae and Erythroxylaceae. In contrast, Linaceae, Irvingiaceae and Caryocaraceae are not clearly supported as a clade, or as closely related to Rhizophoraceae and Erythroxylaceae, as their shared features are probably mainly symplesiomorphies at the level of Malpighiales or a (still undefined) larger subclade of Malpighales, rather than synapomorphies. Rhizophoraceae and Erythroxylaceae share (among other features) conduplicate petals enwrapping stamens in bud, antepetalous stamens longer than antesepalous ones, a nectariferous androecial tube with attachment of the two stamen whorls at different positions: one whorl on the rim, the other below the rim of the tube, the ovary shortly and abruptly dorsally bulged and the presence of a layer of idioblasts (laticifers?) in the sepals and ovaries. Ctenolophonaceae share with Rhizophoraceae and/or Erythroxylaceae (among other features) sepals with less than three vascular traces, a short androgynophore, an ovary septum thin and severed or completely disintegrating during development, leading to a developmentally secondarily unilocular ovary, a zigzag‐shaped micropyle and seeds with an aril. Special features occurring in families of all three groupings studied here are, for example, synsepaly, petals not retarded and thus forming protective organs in floral bud, petals postgenitally fused or hooked together in bud, androecial tube and petals fusing above floral base, androecial corona, apocarpous unifacial styles, nucellus thin and long, early disintegrating (before embryo sac is mature), and nectaries on the androecial tube. Some of these features may be synapomorphies for the entire group, if it forms a supported clade in future molecular studies, or for subgroups thereof. Others may be plesiomorphies, as they also occur in other Malpighiales or also in Celastrales or Oxalidales (COM clade). The occurrence of these features within the COM clade is also discussed. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 331–416.     

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.     

Comparative floral structure and systematics of the clade of Lophopyxidaceae and Putranjivaceae (Malpighiales)

In molecular phylogenetic studies, Lophopyxidaceae and Putranjivaceae are well supported as sisters in the large rosid order Malpighiales. As the floral structure of both families is poorly known and the two families have never been compared, the present comparative study was carried out, as part of a larger project on the comparative floral structure of Malpighiales, using microtome section series and scanning electron microscopy (SEM) studies. Similar to other angiosperm clades, it appears that the structure of the ovules is a strong marker for suprafamilial relationships in Malpighiales. Both families have two collateral pendant antitropous ovules per carpel associated with obturators (as in some Euphorbiaceae s.l., to which Putranjivaceae belonged in earlier classifications). However, in contrast with Euphorbiaceae s.l., the ovules are not crassinucellar, but either incompletely tenuinucellar or only weakly crassinucellar with a long and conspicuously slender nucellus and an endothelium, and do not have a nucellar beak, but a normal micropyle, features they share with families other than Euphorbiaceae s.l. among Malpighiales. Other shared features of the two families include the following. The outer sepals tend to be smaller than the inner ones and the sepals do not protect the gynoecium in older buds. Sepals of some taxa have a single vascular trace. A short zone of synsepaly tends to be present. Stamens tend to be antesepalous in haplostemonous flowers. A short gynophore is present. The synascidiate zone extends up to above the placenta, but is restricted to the ovary in taxa with more than one carpel. The micropyle is formed by the inner integument. The ventral carpel slits extend down into the synascidiate zone as postgenitally fused furrows. The carpels have a broad dorsal band of vascular bundles in the style. The overall floral structure of the two families corroborates their sister position well and does not support the earlier association of Putranjivaceae with Euphorbiaceae s.l. or of Lophopyxidaceae with Geraniales–Sapindales–Celastrales, which rely on shared superficial floral similarities. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 404–448.     

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 in Oxalidales (Oxalidaceae, Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae, Elaeocarpaceae, Tremandraceae)

Floral morphology, anatomy and histology were studied in representatives of all families of current Oxalidales, which were recently constituted as a result of molecular systematic studies by other authors, and are composed of families of different positions in traditional classifications (Oxalidaceae, Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae, Elaeocarpaceae, Tremandraceae). Two of the three pairs of sister (or nested) families that come out in molecular analyses are highly supported by floral structure: Oxalidaceae/Connaraceae and Elaeocarpaceae/Tremandraceae, whereas Cephalotaceae/Cunoniaceae are not especially similar at the level of Oxalidales. Oxalidaceae and Connaraceae share petals that are postgenitally united into a basal tube (although they are imbricate in both) but free at the insertion zone, stamens that are congenitally united at the base, uniseriate glandular hairs on the stamen filaments, and ovules that are hemianatropous to almost orthotropous. The sharing of a special type of sieve-tube plastids and of trimorphic heterostyly, studied by other authors, should also be mentioned. With Brunelliaceae, the two families share an androgynophore and nectaries at the base of the stamens in alternisepalous sectors. Elaeocarpaceae and Tremandraceae share buzz-pollinated flowers and a syndrome of features functionally connected with it. In addition, petals are larger than sepals in advanced bud, they are valvate, involute and enwrap part of the adjacent stamens, they have three vascular traces. Lignified hairs are common on the anthers and are found in the ovary locules and on the ovules (not lignified) of representatives of both families. Ovules have a chalazal appendage, and the inner integument is much thicker than the outer.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 321–381.     

Comparative floral structure and systematics in Cucurbitales (Corynocarpaceae, Coriariaceae, Tetramelaceae, Datiscaceae, Begoniaceae, Cucurbitaceae, Anisophylleaceae)

Floral structure, including morphology, anatomy and histology, was comparatively studied in representatives of all seven families of Cucurbitales as currently circumscribed by other authors based on molecular analyses and including Corynocarpaceae, Coriariaceae, Tetramelaceae, Datiscaceae, Begoniaceae, Cucurbitaceae and Anisophylleaceae. Three superfamilial clades are supported by floral structure: Tetramelaceae/Datiscaceae, Tetramelaceae/Datiscaceae/Begoniaceae and Corynocarpaceae/Coriariaceae. Anisophylleaceae appear most isolated in Cucurbitales, and show more similarities with Oxalidales, especially Cunoniaceae, although some features of interest are shared with other Cucurbitales and not Oxalidales. Tetramelaceae and Datiscaceae share dioecy, completely isomerous (but not regularly pentamerous) flowers (not in male Datiscaceae), only small sepals, lacking petals (not in male Octomeles). Tetramelaceae, Datiscaceae and Begoniaceae share the presence of numerous small ovules and seeds with a large‐celled surface, 2‐cell‐layered integuments, and a collar around the funicle by an extension of the outer integument. Corynocarpaceae and Coriariaceae share thick petals, unifacial stigmas, superior ovaries with a single, median, pendant syntropous ovule per carpel, and annular outer integuments with vasculature at the base. The four classical core families of Cucurbitales: Tetramelaceae, Datiscaceae, Begoniaceae and Cucurbitaceae (relationship unresolved, not retrieved as a clade as yet in molecular studies) share in various combinations androdioecy, basifixed and extrorse or latrorse anthers, trimerous gynoecia, bifurcate free carpel parts, an extended roof over the ovary formed by the ventral parts of the carpels, and parietal placentae. Trends of interest at the order level are unisexual flowers, thick, pointed petals (if present) that do not conform to the model in other rosids or basal core eudicots, a 2‐cell‐layered inner integument, which is delayed in development, and lacking or scant tanniferous tissues in flowers. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145 , 129–185.     

Comparative floral structure and systematics in Crossosomatales (Crossosomataceae, Stachyuraceae, Staphyleaceae, Aphloiaceae, Geissolomataceae, Ixerbaceae, Strasburgeriaceae)

Floral structure of all putative families of Crossosomatales as suggested by molecular studies was comparatively studied. The seven comprise Crossosomataceae, Stachyuraceae, Staphyleaceae, Aphloiaceae, Geissolomataceae, Ixerbaceae, and Strasburgeriaceae. The entire clade (1) is highly supported by floral structure, also the clades (in sequence of diminishing structural support): Ixerbaceae/Strasburgeriaceae (2), Geissolomataceae/Ixerbaceae/Strasburgeriaceae (3), Aphloiaceae/Geissolomataceae/Ixerbaceae/Strasburgeriaceae (4), and Crossosomataceae/Stachyuraceae/Staphyleaceae (5). Among the prominent floral features of Crossosomatales (1) are solitary flowers, presence of a floral cup, imbricate sepals with outermost smaller than inner, pollen grains with horizontally extended endoapertures, shortly stalked gynoecium, postgenitally united carpel tips forming a compitum, stigmatic papillae two‐ or more‐cellular, ovary locules tapering upwards, long integuments forming zigzag micropyles, cell clusters with bundles of long yellow crystals, mucilage cells, seeds with smooth, sclerified testa and without a differentiated tegmen. Clade (2) is characterized by large flowers, petals forming a tight, pointed cone in bud, stamens with long, stout filaments and sagittate anthers, streamlined, conical gynoecium, antitropous ovules, rudimentary aril, lignified, unicellular, T‐shaped hairs and idioblasts with striate mucilaginous cell walls. Clade (3) is characterized by alternisepalous carpels, punctiform stigma formed by postgenitally united and twisted carpel tips, synascidiate ovary, only one or two pendant ovules per carpel, nectary recesses between androecium and gynoecium. Clade (4) is characterized by pronounced ‘pollen buds’. Clade (5) is characterized by polygamous or functionally unisexual flowers, x‐shaped anthers, free and follicular carpels (not in Stachyuraceae). Crossosomataceae and Aphloiaceae, although not retrieved as a clade in molecular studies, share several special floral features: polystemonous androecium; basifixed anthers without a connective protrusion; stigma with two more or less decurrent crests; camplyotropous ovules and reniform seeds; simple, disc‐shaped nectaries and absence of hairs. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 147 , 1–46.     

Comparative floral structure and systematics in the balsaminoid clade including Balsaminaceae,Marcgraviaceae and Tetrameristaceae (Ericales)

Based on molecular phylogenetic studies, Balsaminaceae, Tetrameristaceae (including Pellicieraceae) and Marcgraviaceae form the strongly supported first branching clade in the asterid order Ericales. Marcgraviaceae and Tetrameristaceae were proposed to be closely related in pre‐molecular studies, but the systematic position of Balsaminaceae has been controversial for some time and a relationship with the other two families was never suggested in pre‐molecular/pre‐cladistic times. However, interfamilial relationships in the clade are still unclear because of conflicting phylogenetic hypotheses from molecular analyses. In order to assess the validity of these molecular hypotheses from a morphological point of view, the floral morphology, anatomy and histology of Balsaminaceae, Tetrameristaceae and Marcgraviaceae are comparatively studied in detail. In addition, earlier literature is reviewed. The monophyly of the balsaminoid clade is strongly supported by floral structure, and a series of potential floral synapomorphies is identified for the clade. Prominent features shared by the three families include broad and dorsiventrally flattened filaments, thread‐like structures lining the stomia of dehisced anthers, secretory inner morphological surfaces of the gynoecium, ovules intermediate between uni‐ and bitegmic, incompletely tenuinucellar ovules, fruits with persistent style and stigma, seeds lacking endosperm and several anatomical/histological traits. The families are also distinctive because the bracts and/or sepals are petaloid and nectariferous. Further, the floral structure supports a sister group relationship between Balsaminaceae and Tetrameristaceae rather than any of the other possible interfamilial relationships. These two families share a caducous calyx, post‐genital fusion/coherence of filaments and ovary surface, latrorse anther dehiscence, commissural carpel lobes and ovules with a thickened funiculus and a constricted chalazal region. The occurrence of these features in Ericales is discussed. Future structural studies in other ericalean lineages and additional molecular studies are needed to further test these features with respect to their systematic value for the balsaminoid clade. Some may turn out to be true synapomorphies, whereas others may be recognized as plesiomorphies, as they may be more widely spread in Ericales than currently thought. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 325–386.     

Comparative floral morphology and anatomy of Anacardiaceae and Burseraceae (Sapindales), with a special focus on gynoecium structure and evolution

Anacardiaceae and Burseraceae are traditionally distinguished by the number of ovules (1 vs. 2) per locule and the direction of ovule curvature (syntropous vs. antitropous). Recent molecular phylogenetic studies have shown that these families are sister groups in Sapindales after having been separated in different orders for a long time. We present a comparative morphological study of the flower structure in both families. The major clades, usually supported in molecular phylogenetic analyses, are well supported by floral structure. In Anacardiaceae, there is a tendency to gynoecium reduction to a single fertile carpel (particularly in Anacardioideae). The single ovule has a long and unusually differentiated funicle, which connects with the stylar pollen tube transmitting tract in all representatives studied. In Anacardiaceae–Spondiadoideae, there is a tendency to form an extensive synascidiate zone, with a massive remnant of the floral apex in the centre; these features are also present in Beiselia (Burseraceae) and Kirkiaceae (sister to Anacardiaceae plus Burseraceae) and may represent a synapomorphy or apomorphic tendency for the three families. In core Burseraceae, gynoecium structure is much less diverse than in Anacardiaceae and has probably retained more plesiomorphies. Differences in proportions of parts of the ovules in Anacardiaceae and Burseraceae are linked with the different direction of ovule curvature. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 499–571.     

Inflorescence and floral morphology of Haptanthus hazlettii (Buxaceae,Buxales)

The enigmatic Central American tree Haptanthus hazlettii has recently been placed in Buxaceae (Buxales) by molecular evidence. However, Haptanthus appears morphologically to be fundamentally different from other Buxales in having pluriovular carpels with parietal placentation and reduced male reproductive units of an obscure morphological nature. The latter have been interpreted to be pairs of unistaminate flowers, or single flowers, either bearing two stamens or a pair of phyllomes with adnate introrse anthers. We (re‐)investigated the structure of the inflorescences and flowers of Haptanthus in order to clarify their homologies with reproductive structures of Buxales. We found that, despite some distinctive traits of flower morphology, Haptanthus shares many floral characters, including the opposite and pairwise arrangement of floral organs and the fusion between perianth members and stamens, with some Buxales and other early‐branching eudicots. The plicate and pluriovular gynoecium of Haptanthus may be the result of a drastic elongation of the symplicate zone, accompanied by an increase in ovule number, and is thus a derived trait in Buxales. The anther‐bearing structures are phyllomes with adnate anthers rather than stamens or unistaminate flowers. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 190–200.     

Female flowers and systematic position of Picrodendraceae (Euphorbiaceae s.l., Malpighiales)

This is the first comparative study of floral structure of the recently established new family Picrodendraceae (part of Euphorbiaceae s.l.) in Malpighiales. Nine species of eight (out of ca. 28) genera were studied. Female flowers are mainly completely trimerous, and in such flowers the perianth consists of one or two whorls of sepals. A floral disc (which probably functions as a nectary) is mostly present. The free parts of the carpels are simple (unbranched) in all ten species studied. Each carpel contains two crassinucellar, anatropous or hemitropous, epitropous (antitropous) ovules, which are covered by a large obturator. The inner integument is thicker than the outer (equally thick in two species studied), and commonly both integuments form the micropyle. In mature ovules the vascular bundle commonly branches in the chalaza, with the branches extending to the base of the inner integument but not entering it. A nucellar cap and, less often, a nucellar beak is formed. Floral structure supports the close relationship of Picrodendraceae with Phyllanthaceae and Euphorbiaceae s.str. within Malpighiales, as suggested (but not yet strongly supported) by some recent published molecular analyses. These three families share a unique combination of characters, including (1) unisexual, apetalous trimerous flowers, (2) crassinucellar ovules with a nucellar beak, (3) a large obturator, and (4) explosive fruits with carunculate seeds.     

Flower development schedule and AGAMOUS‐like gene expression patterns in two morphs of Nigella damascena (Ranunculaceae) differing in floral architecture

Love‐in‐a‐mist (Nigella damascena) is an annual species of Ranunculaceae native to the Mediterranean Basin, characterized by delicate flowers lying on long lacy bracts. Two floral morphs of N. damascena, designated [P] and [T], differ in the identity and number of perianth organs and in the position of the perianth–androecium boundary on the meristem. They both occur in the wild. Here we describe a precise comparative schedule of floral development in the two morphs. We divided the sequence of developmental events affecting the floral meristem into six stages and related them to the height of the elongating stem and to the time elapsed after the beginning of stem elongation. In addition, we characterized the expression pattern of C‐class genes in floral organs of both morphs in an attempt to better characterize the differences between the two floral groundplans. In the [T] morph an expansion of the expression domain of AGAMOUS (AG) paralogues outside the fertile organs was observed, correlating with the change in identity of the inner perianth organs. Expression of AG‐like genes in the sepal‐like organs suggests these are not identical to true sepals at the molecular level. The morpho‐temporal framework we have defined will allow us to compare various gene expression profiles at targeted developmental stages in both morphs, providing further insight into the molecular control of the floral dimorphism in N. damascena and into the processes underlying the transition from a differentiated (bipartite) to an undifferentiated (unipartite) perianth. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 608–619.     

Structure and evolution of the androecium in the Malvatheca clade (Malvaceae s.l.) and implications for Malvaceae and Malvales

Androecial development and structure as well as floral vasculature of six selected species of Bombacoideae and of several smaller lineages of the Malvatheca clade (Malvaceae s.l.) were studied. All studied taxa share a similar pattern of androecial development: initially, five antepetalous/antetepalous and five alternipetalous/alternitepalous primary androecial primordia develop on a ring wall. Two elongate secondary androecial primordia form on each antepetalous/antetepalous sector. At anthesis the androecium consists of an androecial tube crowned by five androecial lobes. Each of these lobes is the developmental product of an alternipetalous/alternitepalous primary androecial primordium and its two neighbouring antepetalous/antetepalous secondary androecial primordia. The elongate, sessile androecial units are positioned along the lateral margins of the androecial lobes and in the distal part of the androecial tube. Seen in the light of the most recent studies of floral development and phylogeny of the Malvaceae and the Malvales as a whole, our data indicate that i) elongate, sessile androecial units are ancestral in the Malvatheca clade, that ii) an obdiplostemonous floral ground plan is a synapomorphy for the Malvaceae, and that iii) diplostemony is most likely ancestral in the Malvales.     

Comparative floral structure and systematics in Apodanthaceae (Rafflesiales)

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 Malvales.     

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.     

Gynoecium and perianth inZanthoxylum s.l. (Rutaceae)

The question whether the uniseriate perianth ofZanthoxylum L. s. str. is homologous with the calyx or the corolla of taxa included inFagara, or of an independent origin, has been controversial for a long time, but the arguments mostly have remained theoretical. The present investigation of floral structures indicates that there are two different types of uniseriate perianth inZanthoxylum s. str. Therefore, this taxon does not represent a natural group and should be united withFagara asZanthoxylum s.l. The infrageneric taxonomy of this genus is still very ambiguous. It is shown that differences in indumentum, number of sepals and petals (5-4-3) resp. perianth segments (4–9), stamens (3–6), and free carpels (1–5) are of systematic relevance. Particularly important but so far neglected is carpel shape, where an acrostylous and an anacrostylous-basistylous type can be recognized. Stigmata of 2 or more carpels mostly fuse to form a compitum. 4–5-merous flowers with calyx and corolla, and acrostylous carpels are considered as plesiomorphic character states in the genus. On the basis of ± corresponding morphological and phytochemical progressions a working hypothesis about the relationships withinZanthoxylum s.l. is presented in graphical form (Fig. 9).Adapted from a lecture held at the 10th Symposion on Morphology, Anatomy, and Systematics in Göttingen, February 1991.     

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.     

Fruit structure and systematics of Monimiaceae s.s. (Laurales)

Fruit structure (anatomy) was studied in 27 species of 15 genera of Monimiaceae s.s. Almost all have apocarpous gynoecia, with the carpels more or less surrounded by a floral cup. The fruitlets are presented on the opened floral cup, which, depending on its pre‐ and post‐floral development, differentially contributes to the attractive part of the mature fruit. Morphologically similar fruits may differ conspicuously in anatomical structure. Based on anatomical characters two different fruit forms were found: drupe(let)s (with compact sclerenchymatic endocarp forming a stone: putamen) and berry(let)s (with parenchymatic endocarp, and mesocarp parenchyma containing isolated sclereid nests). Four types of drupelets differing by the endocarp structure were tentatively distinguished: (1) the Monimia‐type has a many‐cell‐layered putamen of large isodiametric sclereids, interrupted on the ventral side by few radial rows of small sclereids; (2) the Hortonia‐type has a few‐cell‐layered putamen of isodiametric, especially thick‐walled sclereids – it may be composed of two lateral halves, i.e. with the sclerenchyma partially interrupted on the ventral and dorsal sides (but without rows of small sclereids); (3) the Mollinedia‐type has a few‐cell‐layered putamen, with more or less radially elongate sclereids with wavy cell walls; and (4) the Hedycarya‐type has a one‐cell‐layered putamen of pronouncedly radially elongate sclereids with wavy cell walls. Drupelets of some taxa with a single‐cell‐layered endocarp with only weakly thickened cell walls may represent a transition from drupelets to berrylets. The fruit structure supports three major clades recognized earlier by morphological studies and by molecular phylogenetic analyses: (1) Monimioideae (Monimia‐type drupelets), (2) Hortonieae of Mollinedioideae (Hortonia‐type drupelets), and (3) the remainder of Mollinedioideae (Hedycarya‐ and Mollinedia‐types) and berrylets. Fruit structure also supports the close relationship of Monimiaceae and Lauraceae. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 265–285.     

Floral morphology,micromorphology and palinology of selected Sedum s.l. species (Crassulaceae)

Sedum L. (Crassulaceae) is a large and taxonomically difficult genus whose delimitation and classification are under debate. Due to the controversial results of previous cytological, morphological, and molecular studies, further investigations are needed in order to gain a shared taxonomy of the current recognized species clades. In the present paper, morphological and micromorphological features of 23 selected Mediterranean species of Sedum s.l. – collected from exsiccata or fresh specimens throughout Italy – were investigated, in order to provide additional data toward their classification above species level. In particular, the study focused on flower structure and morphology, floral epidermal surfaces and pollen morphology. The distribution pattern of the examined microcharacters across the species revealed a wide range of variation and different combinations of the single characters. NMDS analysis allowed individuating discrete groups that showed a general consistency with the current systematic delimitation of species groups. Our study also evidenced for the first time the wide morphological variability of nectaries and of the glandular indumentum, not previously investigated in detail. In addition, we proposed the combined use of floral diagrams and floral formulae as valuable tools in studying the variability of flower structure at genus level.