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 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.     

Generative ontogeny in Tiquilia (Ehretiaceae: Boraginales) and phylogenetic implications

Tiquilia is very different from the other members of the Ehretiaceae (Boraginales) in many aspects of morphology and ecology. Because detailed knowledge about flower and fruit traits is necessary to reliably infer character evolution of and within Tiquilia, we investigated flower to fruit ontogeny in eight species of Tiquilia using light and electron microscopy. Tiquilia accumulated a number of autapomorphies such as the prostrate growth form, the lack of lateral and ventral bundles in the gynoecium, and the formation of nutlet‐like mericarpids as dispersal units instead of more or less succulent drupes. The internal architecture of the superior bicarpellate ovary resulted from the development of several secondary septa including apical, basal and false septa, as it has been reported also from other Boraginales. However, no character found in Tiquilia can be regarded as synapomorphic with any other taxon of the Ehretiaceae. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 520–534.     

Comparative floral structure and systematics in Ochnaceae s.l. (Ochnaceae,Quiinaceae and Medusagynaceae; Malpighiales)

Ochnaceae s.l. (Ochnaceae, Quiinaceae and Medusagynaceae), one of the well‐supported subclades of the large order Malpighiales retrieved so far in molecular phylogenetic studies, were comparatively studied with regard to floral structure using microtome section series and scanning electron microscopy (SEM). Floral morphology, anatomy and histology also strongly reflect this close relationship. Potential synapomorphies of the subclade include: flowers nectarless, sepals of different sizes within a flower, petals not retarded in development and forming the protective organs of advanced floral buds, petal aestivation contort, petals with three vascular traces, petals reflexed over the sepals and directed toward the pedicel, polystemony, anthers almost or completely basifixed, gynoecium often with more than five carpels, short gynophore present, styles separate for at least their uppermost part and radiating outwards, suction‐cup‐shaped stigmas, vasculature forming a dorsal band of bundles in the upper stylar region, gynoecium epidermis with large, radially elongate cells, ovules either weakly crassinucellar or incompletely tenuinucellar with an endothelium, abundance of tanniferous tissues and sclerenchyma in floral organs. The most strongly supported subclade of two of the three families in molecular analyses, Quiinaceae and Medusagynaceae, is also particularly well supported by floral structural features, including the presence of functionally and morphologically unisexual flowers, a massive thecal septum that persists after anther dehiscence, styles radiating outward from the ovary, two lateral ovules per carpel, positioned one above the other, conspicuous longitudinal ribs on the ovary wall at anthesis, and a ‘false endothelium’ on the nucellus at anthesis. Additionally, the group fits well in Malpighiales and further emphasizes the relationship of Malpighiales with Celastrales and Oxalidales, and thus the unity of the COM clade. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 299–392.     

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.     

Floral organogenesis of Chloranthus sessilifolius, with special emphasis on the morphological nature of the androecium of Chloranthus (Chloranthaceae)

 Floral organogenesis of Chloranthus sessilifolius K. F. Wu is described. The inflorescence primordium is dome-like in the beginning and then elongates, and bract primordia initiate almost decussately. Each floral primordium, arising from the axil of a bract, soon becomes a scale-like structure, with three primordia of androecial lobes originating from its abaxial part, and the gynoecial primordium in adaxial position. As the androecial lobes become more distinct, four thecae are already in differentiation, and the gynoecial primordium appears as a shallow disc. The androecial lobes do not extend their length until the thecae approach maturity and the stigma is differentiated. The androecial lobes are united at all the stages of development, and the entire androecium falls off as a unit at the end of anthesis. Based on these results, combined with published evidence from neobotany, palaeobotany and phylogenetic studies, the morphological nature of the androecium of Chloranthus is further discussed. Our studies support the viewpoint that the androecial structure of Chloranthus may have arisen by splitting of a single stamen with 2 marginal thecae. Received May 2, 2001 Accepted December 18, 2001     

Pseudodiplostemony, and its implications for the evolution of the androecium in the Caryophyllaceae

The androecium of the Caryophyllaceae is varied, ranging from a two-whorled condition to a single stamen. A number of species belonging to the three subfamilies, Caryophyl-loideae, Alsinoideae and Paronychioideae have been studied ontogenetically with the SEM to understand their peculiar androecial development in the broader context of the Caryophyllales alliance. Although patterns of initiation are highly variable among species, there are three ontogenetic modes of stamen initiation: all stamens simultaneous within a whorl, the antepetalous stamens simultaneous and the antesepalous sequentially with a reversed direction, or both whorls sequentially with or without a reversed direction. The most common floral (ontogenetic) sequence of the Caryophyllaceae runs as follows: five sepals (in a 2/5 sequence), the stamens in front of the three inner sepals successively, stamens opposite the two outermost sepals, five antepetalous stamens (simultaneously or in a reversed spiral superimposed on the spiral of the antesepalous stamens), five outer sterile (petaloid) organs arising before, simultaneously or after the antesepalous stamens, often by the division of common primordia. A comparison with the floral configurations of the Phytolaccaceae and Molluginaceae indicates that the outer petaline whorl of the Caryophyllaceae corresponds positionally to the alternisepalous stamens of somePhytolacca, such asP. dodecandra. The difference withP. dodecandra lies in the fact that an extra inner or outer whorl is formed in the Caryophyl-laceae, in alternation with the sepals. A comparable arrangement exists in the Molluginaceae, though the initiation of stamens is centrifugal. A comparison of floral ontogenies and the presence of reduction series in the Caryophyllaceae support the idea that the pentamerous arrangement is derived from a trimerous prototype. Petals correspond to sterillized stamens and are comparable to two stamen pairs opposite the outer sepals and a single stamen alternating with the third and fifth sepals. Petals are often in a state of reduction; they may be confused with staminodes and they often arise from common stamenpetal primordia. The antesepalous stamen whorl represents an amalgamation of two whorls: initiation is reversed with the stamens opposite the fourth and fifth formed sepals arising before the other, while the stamens opposite the first and second formed sepals are frequently reduced or lost. Reductive trends are correlated with the mode of initiation of the androecium, as well as changes in the number of carpels, and affect the antesepalous and antepetalous whorls in different proportions. It is concluded that the androecium of the Caryophyllaceae is pseudodiplos-temonous and is not comparable to diplostemonous forms in the Dilleniidae and Rosidae. The basic floral formula of Caryophyllaceae is as follows: sepals 5—petals 5 (sterile stamens)—antesepalous stamens 3+2—antepetalous stamens 5 gynoecium 5.     

The evolution of staminodes in angiosperms: patterns of stamen reduction, loss, and functional re-invention

Stamens that have lost their primary function of pollen production, or staminodes, occur uncommonly within angiosperms, but frequently fulfill important secondary floral functions. The phylogenetic distribution of staminodes suggests that they typically arise during evolutionary reduction of the androecium. Differences in the genetic control and patterns of stamen loss between actinomorphic and zygomorphic flowers shape staminode development. In clades with actinomorphic flowers, staminodes generally replace an entire stamen whorl and staminode loss seems irreversible. In contrast, in clades with zygomorphic flowers staminodes evolve from a subset of the stamens in a whorl and staminodes can reappear in a lineage after being lost (e.g., Cheloneae, Scrophulariaceae). If staminodes do not adopt new functions during androecium reduction they are lost quickly, so that nonfunctional staminodes appear only in recently derived taxa. Alternatively, when staminodes assume new floral roles, either directly or indirectly after a nonfunctional period, they can become integral floral components which perpetuate within clades (e.g., Orchidaceae). Indirect evolution of staminode function allows greater flexibility of function by allowing staminodes to take over roles not performed by stamens, such as involvement in mechanisms to prevent self-pollination and mechanisms of explosive pollination. Multifunctional staminodes characterize lineages with universal or widespread staminodes.     

Early floral development and androecium organization in Fouquieriaceae (Ericales)

Early floral development with focus on the androecium was studied with the help of scanning electron microscopy and serial microtome sectioning in Fouquieria columnaris and F. splendens. Perianth organs appear in a spiral pattern on the floral apex. The spiral may be a clockwise or anti-clockwise. The androecium is best interpreted as two-whorled with all the stamens arranged in a single series. In F. splendens, two or more of the five epipetalous stamen positions are doubled, i.e. they are occupied by stamen pairs. Unusual features in the floral development of Fouquieriaceae include (1) a strong spiral component even in whorled organ categories and (2) a pronouncedly asymmetric floral apex during an early phase of floral development. From a phylogenetic point of view, it seems plausible that the common ancestor of Fouquieriaceae and its sister family Polemoniaceae was characterized by two alternating, pentamerous stamen-whorls.     

Reevaluation of the perianth and androecium in Caryophyllales: implications for flower evolution

The Caryophyllales have the highest diversity in androecial patterns among flowering plants with stamen numbers ranging from 1 up to 4,000. Thanks to the recent progress in reconstructing the phylogeny of core Caryophyllales, questions of floral evolution, such as the origin and diversification of the androecium, can be readdressed. Caryophyllales are unique among core eudicots in sharing an androecial ring meristem or platform with centrifugal development of stamens and petals. Stamens are basically arranged in two whorls and evolution within the clade depends on the shift of either the antesepalous or the alternisepalous whorls to an upper position on the ring meristem and the reduction of the other. Four main developmental phenomena are responsible for the high diversity in androecial patterns: (1) the sterilisation of the outermost stamens through a division of common primordia; (2) the secondary addition of stamens by a centrifugal initiation of supernumerary stamens superimposed on a lower stamen number; (3) the pairwise displacement of alternisepalous stamens to the middle of the outer sepals and their potential fusion, or as part of a pluristaminate androecium; (4) the inversed sequence, reduction and loss of antesepalous stamens. Shifts in stamen numbers depend on pressures of the calyx and carpels and changes in the number of the latter. These patterns are expressed differently in the three main evolutionary lines of core Caryophyllales and are systematically relevant: (1) A basal grade of Caryophyllales, culminating with Caryophyllaceae, Amaranthaceae, Stegnosperma and Limeum, has the antesepalous stamens initiated in upper position on the ring meristem, and alternisepalous stamens are preferentially reduced. Among the antesepalous whorl there is a progressive loss of stamens following a sequence inversed to sepal initiation. Petaloid staminodes are formed by the radial division of outer stamens. (2) The raphide-clade and Molluginaceae are characterized by alternisepalous stamens in upper position on the ring meristem, with a trend to secondary stamen multiplication, and loss of antesepalous stamens. (3) The Portulacineae share the pattern of the raphide clade, but some taxa show shifts to an upper position on the ring meristem of either antesepalous or alternisepalous stamens, linked with secondary multiplications and reduction of either whorl. Different floral characters are plotted on a recent cladogram of Caryophyllales. The data show a consistent correlation between shifting carpel and stamen numbers independent of perianth evolution. Comparative data suggest that the basic androecium of Caryophyllales consists of two whorls of five stamens, linked with an absence of petals, and the evolution of the androecium is a combination of reductions and secondary multiplications of stamens with a highly predictive systematic value.     

Variation and evolution of herkogamy in Exochaenium (Gentianaceae): implications for the evolution of distyly

Backgrounds and Aims

The spatial separation of stigmas and anthers (herkogamy) in flowering plants functions to reduce self-pollination and avoid interference between pollen dispersal and receipt. Little is known about the evolutionary relationships among the three main forms of herkogamy – approach, reverse and reciprocal herkogamy (distyly) – or about transitions to and from a non-herkogamous condition. This problem was examined in Exochaenium (Gentianaceae), a genus of African herbs that exhibits considerable variation in floral morphology, including the three forms of herkogamy.

Methods

Using maximum parsimony and maximum likelihood methods, the evolutionary history of herkogamic and non-herkogamic conditions was reconstructed from a molecular phylogeny of 15 species of Exochaenium and four outgroup taxa, based on three chloroplast regions, the nuclear ribosomal internal transcribed spacer (ITS1 and 2) and the 5·8S gene. Ancestral character states were determined and the reconstructions were used to evaluate competing models for the origin of reciprocal herkogamy.

Key results

Reciprocal herkogamy originated once in Exochaenium from an ancestor with approach herkogamy. Reverse herkogamy and the non-herkogamic condition homostyly were derived from heterostyly. Distylous species possessed pendent, slightly zygomorphic flowers, and the single transition to reverse herkogamy was associated with the hawkmoth pollination syndrome. Reductions in flower size characterized three of four independent transitions from reciprocal herkogamy to homostyly.

Conclusions

The results support Lloyd and Webb''s model in which distyly originated from an ancestor with approach herkogamy. They also demonstrate the lability of sex organ deployment and implicate pollinators, or their absence, as playing an important role in driving transitions among herkogamic and non-herkogamic conditions.     

Floral biology and histochemical analysis of Vanilla edwallii Hoehne (Orchidaceae: Vanilloideae): an orchid pollinated by Epicharis (Apidae: Centridini)

The genus Vanilla is the most diverse in Vanilloideae, with ca 90 species distributed among tropical regions. Despite their economic importance, studies on pollination of Vanilla are very scarce and data on pollinators of species endemic to Brazil are lacking. Based on fieldwork and laboratory investigations, the floral biology of V. edwallii was studied. The pollinators and pollination process were recorded at the Serra do Japi reserve, state of São Paulo, southeastern Brazil, and the presence of floral reward was also investigated. Vanilla edwallii blooms in summer. The lateral inflorescences produce up to four pale green flowers. The white labellum is united to the base of the column forming a mentum. In the studied population V. edwallii is pollinated by Epicharis (Hoplepicharis) affinis, where the males exhibit a territorial behavior, defending flowers from other possible flower visitors. The pollen is deposited on the scutellum of bees when they abandon the flower. The mentum region is dry, suggesting no nectar production. The only secretory structures are osmophores dispersed on the inner surface of the lip responsible for production of a sweet fragrance, which together with color and morphology of flowers is related to bee attraction. The labellum is rich in mucilaginous cells, while the mucilaginous substance is retained inside the cells. The histochemical analysis also detected the presence of phenolic compounds and starch concentrated mainly at the adaxial surface of the lip and around the vascular bundles.     

The adaptive value of young leaves being tightly folded or rolled on monocotyledons in tropical lowland rain forest: an hypothesis in two parts

In tropical lowland rain forest, we find that species with the leaves tightly folded or rolled until they reach at least 50% of final length occur in 10 of the 15 monocot families with >100 species, and in 12 of the 24 monocot families with <100 species, but in only seven of the 212 dicot families (eudicots and magnoliids). Earlier researchers have described how examples of tightly folded and rolled leaves develop, but most have not considered the potentially adaptive value of this pattern of growth. We hypothesize that it is a protection against herbivorous invertebrates. For tropical and temperate dicots, the young leaves have been found to suffer much smaller losses to herbivores while folded and rolled than after they are unfolded or unrolled. Being folded or rolled until a late stage involves an ‘opportunity cost’ in the loss of photosynthesis. Among dicots, defences involving such a cost (notably late development of photosynthetic systems in pendent soft young leaves) are typical of shade-tolerant species, which have longer-lived leaves than light-demanders. In contrast, among monocots late folding and rolling are found in both shade-tolerators and light-demanders. We hypothesize that late folding and rolling bring a net advantage to monocots in general, whether shade-tolerant or light-demanding, despite the opportunity cost, because they mostly have fewer leaves per plant of a given size, and therefore an individual leaf is relatively more valuable to the plant. As a coda, we suggest that the ‘sleep movements’ of some tropical plants, and the circinate vernation of ferns and some cycads, provide protection against invertebrate herbivores through the apposition of two or more layers of leaf.     

Floral development and vasculature in Eriocaulon (Eriocaulaceae) provide insights into the evolution of Poales

Background and AimsFloral developmental studies are crucial for understanding the evolution of floral structures and sexual systems in angiosperms. Within the monocot order Poales, both subfamilies of Eriocaulaceae have unisexual flowers bearing unusual nectaries. Few previous studies have investigated floral development in subfamily Eriocauloideae, which includes the large, diverse and widespread genus Eriocaulon. To understand floral variation and the evolution of the androecium, gynoecium and floral nectaries of Eriocaulaceae, we analysed floral development and vasculature in Eriocaulon and compared it with that of subfamily Paepalanthoideae and the related family Xyridaceae in a phylogenetic context.MethodsThirteen species of Eriocaulon were studied. Developmental analysis was carried out using scanning electron microscopy, and vasculature analysis was carried out using light microscopy. Fresh material was also analysed using scanning electron microscopy with a cryo function. Character evolution was reconstructed over well-resolved phylogenies.Key ResultsPerianth reductions can occur due to delayed development that can also result in loss of the vascular bundles of the median sepals. Nectariferous petal glands cease development and remain vestigial in some species. In staminate flowers, the inner stamens can emerge before the outer ones, and carpels are transformed into nectariferous carpellodes. In pistillate flowers, stamens are reduced to staminodes and the gynoecium has dorsal stigmas.ConclusionsFloral morphology is highly diverse in Eriocaulon, as a result of fusion, reduction or loss of perianth parts. The nectariferous carpellodes of staminate flowers originated first in the ancestor of Eriocaulaceae; petal glands and nectariferous branches of pistillate flowers originated independently in Eriocaulaceae through transfer of function. We present a hypothesis of floral evolution for the family, illustrating a shift from bisexuality to unisexuality and the evolution of nectaries in a complex monocot family, which can contribute to future studies on reproductive biology and floral evolution in other groups.     

Floral development of Hydrocera and Impatiens reveals evolutionary trends in the most early diverged lineages of the Balsaminaceae

Background and Aims

Balsaminaceae consist of two genera, the monospecific Hydrocera and its species-rich sister Impatiens. Although both genera are seemingly rather similar in overall appearance, they differ in ecology, distribution range, habitat preference and morphology. Because morphological support for the current molecular phylogenetic hypothesis of Impatiens is low, a developmental study is necessary in order to obtain better insights into the evolutionary history of the family. Therefore, the floral development of H. triflora and I. omeiana was investigated, representing the most early-diverged lineage of Impatiens, and the observations were compared with the literature.

Methods

Flowers at all developmental stages were examined using scanning electron microscopy and light microscopy.

Key results

In Hydrocera, two whorls of five free perianth primordia develop into a less zygomorphic perianth compared with its sister genus. The androecial cap originates from five individual stamen primordia. Post-genital fusion of the upper parts of the filaments result in a filament ring below the anthers. The anthers fuse forming connivent anther-like units. The gynoecium of Hydrocera is pentamerous; it is largely synascidiate in early development. Only then is a symplicate zone formed resulting in style and stigmas. In I. omeiana, the perianth is formed as in Hydrocera. Five individual stamen primordia develop into five stamens, of which the upper part of the filaments converge with each other. The gynoecium of I. omeiana is tetramerous; it appears annular in early development.

Conclusions

Comparison of the present results with developmental data from the literature confirms the perianth morphocline hypothesis in which a congenital fusion of the parts of the perianth results in a shift from pentasepalous to trisepalous flowers. In addition, the development of the androecial cap and the gynoecium follows several distinct ontogenetic sequences within the family.     

The evolution of beetle pollination in a South African orchid

The pollination biology of the orchid Ceratandra grandiflora was investigated in order to determine whether the partial loss of a specialized floral reward (i.e., oil) was the result of an incomplete shift from one specialized pollinator to another. In the three-species clade of section Ceratandra, there has been a progressive loss of the oil-secreting callus. lt is always present in C atrata, sometimes present in C. grandiflora, and never present in C. globosa. Thirty-nine to 67% of individuals in populations of C. grandiflora bear the callus gland, but gland presence has no signifikant effect on the proportion of flowers pollinated. Pollinator observations show that the shift in pollinators is complete and that the oil-secreting callus is a vestige of the ancestral oil-bee pollination system that no longer plays a role in pollination. C grandiflora is pollinated almost exclusively by a single species of hopliine beetle (Scarabaeidae). Experiments with artificial flower traps indicate that color alone can explain the attraction of beetles to C. grandiflora, despite the absence of a floral reward. The proportion of C. grandiflora flowers pollinated (50.2 and 61.1%; N = 524 and 324 flowers, respectively) is unusually high for a plant that relies on generalized food deception and is probably due to the use of inflorescences as mating sites (i.e., rendezvous pollination).     

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.     

Models of floral pattern in detached flowers of Silene coeli-rosa (L) Godr. (Caryophyllaceae)

Organ number per whorl was analysed in aberrant flowers of the long-day (LD) plant , Silene coeli-rosa , to test a hypothesis that organ number in a whorl takes its cue from an adjacent outer whorl and that perturbed organ number per whorl is not random but defaults to that of closely related taxa or genera of the Caryophyllaceae. When plants were grown under short-days (SD), transferred to LD and the shoot meristem excised and cultured in vitro under SD, the normal pattern of flower development was often disrupted. For example, we observed flowers which comprised floral whorls with an aberrant number of floral organs. In part, this was an effect of tissue culture; however, the over-and-above effect was the establishment of an alternative pattern of development. Our data indicate that two distinct and recurrent patterns occurred in the aberrant flowers we observed in five separate experiments. First, pairs of floral whorls were linked so that aberration in one whorl resulted in the next whorl being more aberrant than normal. Second, the number of organs in aberrant whorls was not random, but defaulted to an organ number which mimicked the flowers of closely related species of Silene or related genera in the Caryophyllaceae.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 140 , 229−235.     

The floral scales in Hellmuthia (Cyperaceae, Cyperoideae) and Paramapania (Cyperaceae, Mapanioideae): an ontogenetic study

BACKGROUND AND AIMS: In 1976 the monotypic genus Hellmuthia was placed in the Hypolytreae s.l., but was subsequently ascribed to the Mapanioideae, tribe Chrysitricheae, mainly because of the presence in Hellmuthia of two lateral, mapanioid-like floral scales with ciliated keels, the anatomy of the nutlet, the embryo and the inflorescence. Recently, based on cladistic analyses and supported by pollen ontogenetic evidence, Hellmuthia was transferred to a Cyperaceae, tribe Cypereae, clade mainly consisting of Ficinia and Isolepis. In this study, the floral ontogeny in Hellmuthia was investigated and compared with the floral ontogeny in Paramapania, with special attention for the floral scales. METHODS: Freshly collected inflorescences of Hellmuthia membranacea and Paramapania parvibractea were investigated using scanning electron and light microscopy. KEY RESULTS: In the conical 'spikelet' in Hellmuthia, proximal bracts occur, each axillating an axis with empty glume-like structures, or a reduced spikelet. Hence, it is a reduced partial inflorescence. In Hellmuthia, the stamen primordia originate before the primordia of the perianth-gynoecium appear. Moreover, a third adaxially positioned 'floral scale' was observed for the first time. The position and relative time of appearance of the floral scales in Hellmuthia are typical for perianth parts in Cyperoideae. The basal position of Hellmuthia within a clade of species with usually perianthless flowers, allows the presence of rudiments of a perianth in Hellmuthia to be interpreted as a primitive state. Development of the lateral 'scales' in Paramapania follows a different pattern. Therefore, it was decided that the lateral 'scales' in Paramapania are different from the lateral perianth parts in Hellmuthia. The pollen grains in Hellmuthia are cyperoid, with one polar and five lateral apertures, of which the membrane is covered with sexinous bodies. The pollen surface is granulate and perforate with microspines. CONCLUSIONS: The floral ontogeny in Hellmuthia occurs according to the general cyperoid pattern. The lateral scales in Hellmuthia are perianth parts, and they are not homologous to the lateral 'scales' in Paramapania.