Floral structure and systematics in four orders of rosids, including a broad survey of floral mucilage cells

Phylogenetic studies have greatly impacted upon the circumscription of taxa within the rosid clade, resulting in novel relationships at all systematic levels. In many cases the floral structure of these taxa has never been compared, and in some families, even studies of their floral structure are lacking. Over the past five years we have compared floral structure in both new and novel orders of rosids. Four orders have been investigated including Celastrales, Oxalidales, Cucurbitales and Crossosomatales, and in this paper we attempt to summarize the salient results from these studies. The clades best supported by floral structure are: in Celastrales, the enlarged Celastraceae and the sister relationship between Celastraceae and Parnassiaceae; in Oxalidales, the sister relationship between Oxalidaceae and Connaraceae, and Tremandraceae embedded in Elaeocarpaceae; in Cucurbitales, the sister relationship between Corynocarpaceae plus Coriariaceae, and the grouping of the core Cucurbitales (Cucurbitaceae, Begoniaceae, Tetramelaceae, Datiscaceae); in Crossosomatales, the sister relationship between Ixerbaceae plus Strasburgeriaceae, and between this clade and Geissolomataceae. The core Crossosomatales (Crossosomataceae, Stachyuraceae, Staphyleaceae) and Celastrales as an order are not strongly supported by floral structure. In addition, a new floral feature of potential systematic interest is assessed. Specifically the presence of special cells in flowers with a thickened mucilaginous inner cell wall and a distinct, remaining cytoplasm is surveyed in 88 families and 321 genera (349 species) of basal angiosperms and eudicots. These cells were found to be most common in rosids, particulary fabids (Malpighiales, Oxalidales, Fabales, Rosales, Fagales, Cucurbitales), but were also found in some malvids (Malvales). They are notably absent or rare in asterids (present in campanulids: Aquifoliales, Stemonuraceae) and do not appear to occur in other eudicot clades or in basal angiosperms. Within the flower they are primarily found in the abaxial epidermis of sepal

First steps towards a floral structural characterization of the major rosid subclades

A survey of our own comparative studies on several larger clades of rosids and over 1400 original publications on rosid flowers shows that floral structural features support to various degrees the supraordinal relationships in rosids proposed by molecular phylogenetic studies. However, as many apparent relationships are not yet well resolved, the structural support also remains tentative. Some of the features that turned out to be of interest in the present study had not previously been considered in earlier supraordinal studies. The strongest floral structural support is for malvids (Brassicales, Malvales, Sapindales), which reflects the strong support of phylogenetic analyses. Somewhat less structurally supported are the COM (Celastrales, Oxalidales, Malpighiales) and the nitrogen-fixing (Cucurbitales, Fagales, Fabales, Rosales) clades of fabids, which are both also only weakly supported in phylogenetic analyses. The sister pairs, Cucurbitales plus Fagales, and Malvales plus Sapindales, are structurally only weakly supported, and for the entire fabids there is no clear support by the present floral structural data. However, an additional grouping, the COM clade plus malvids, shares some interesting features but does not appear as a clade in phylogenetic analyses. Thus it appears that the deepest split within eurosids-that between fabids and malvids - in molecular phylogenetic analyses (however weakly supported) is not matched by the present structural data. Features of ovules including thickness of integuments, thickness of nucellus, and degree of ovular curvature, appear to be especially interesting for higher level relationships and should be further explored. Although features of interest are not necessarily stable at the level of a large clade, they do show a considerable concentration in particular clades and are rare or lacking in others. This may be viewed as a special trend for this feature to evolve in this group or to be conserved as a synapomorphy (or a combination of both

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 seed

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

Abstract.: 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 contex

Floral structure of Kirkia (Kirkiaceae) and its position in Sapindales

BACKGROUND AND AIMS: The monogeneric Kirkiaceae (Sapindales) were formerly placed as Kirkioideae in Simaroubaceae. However, recent molecular phylogenetic studies indicate that they are not in Simaroubaceae and they appear to be sister to the clade of Anacardiaceae plus Burseraceae. Such affinity was never considered or discussed since the first description of Kirkia. The present study is the first detailed analysis of the floral structure of a representative of Kirkiaceae and the first comparison with other sapindalean families, especially Anacardiaceae and Burseraceae. METHODS: Floral structure of Kirkia wilmsii was studied using transversal and longitudinal microtome section series, scanning electron microscopy and light microscopy. KEY RESULTS: The flowers of Kirkia wilmsii are morphologically bisexual but functionally unisexual. They are polysymmetric, isomerous (tetramerous) and haplostemonous. The ovary is syncarpous and entirely synascidiate. The floral apex forms a hemispherical protrusion on top of the ovary. The styles are free but postgenitally united and apically form a stigmatic head with a compitum. Each carpel is uniovulate (biovulate in a few other species) and ovules are crassinucellar, bitegmic and slightly campylotropous. The micropyle is formed by both integuments and is unusually long. The unusual two radially disposed locules in each carpel in the former genus Pleiokirkia can be explained developmentally by the two offset and tightly contiguous lateral placentae. CONCLUSIONS: Paralleling the molecular results, a suite of floral features supports the position of Kirkiaceae close to the Anacardiaceae-Burseraceae clade, and not in Simaroubacea

A phylogenetic analysis of Apostasioideae (Orchidaceae) based on ITS, trn L-F and mat K sequences

Abstract.: The orchid subfamily Apostasioideae consists of two genera, Apostasia and Neuwiedia. To study the position of Apostasioideae within Orchidaceae and their intra- and intergeneric relationships, a molecular phylogenetic analysis has been conducted on the nuclear ITS region and the two plastid DNA regions trnL-F intron and matK. The two genera traditionally ascribed to Apostasioideae are each monophyletic. In Apostasia, A. nuda, with two stamens and no staminode, is sister to a clade comprising three species characterised by two stamens and one staminode. Within Neuwiedia, maximum parsimony analyses place N. zollingeri as sister to the clade formed by N. borneensis and N. veratrifolia. A family-wide phylogenetic analysis of matK sequences representing all proposed subfamilies of Orchidaceae produced five moderately to well-supported clades. One of these clades, Apostasioideae, is sister to the clade formed by Vanilloideae, Cypripedioideae, Orchidoideae and Epidendroideae. High transition-transversion ratio and the absence of stop codons in the individual sequences suggest that matK is at the transition from a possibly functional gene to a pseudogene in Apostasioideae, contrary to what is found in some other groups of Orchidacea

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 contex

A phylogenetic analysis of Apostasioideae (Orchidaceae) based on ITS, trn L-F and mat K sequences

The orchid subfamily Apostasioideae consists of two genera, Apostasia and Neuwiedia . To study the position of Apostasioideae within Orchidaceae and their intra- and intergeneric relationships, a molecular phylogenetic analysis has been conducted on the nuclear ITS region and the two plastid DNA regions trn L-F intron and mat K. The two genera traditionally ascribed to Apostasioideae are each monophyletic. In Apostasia , A. nuda , with two stamens and no staminode, is sister to a clade comprising three species characterised by two stamens and one staminode. Within Neuwiedia , maximum parsimony analyses place N. zollingeri as sister to the clade formed by N. borneensis and N. veratrifolia . A family-wide phylogenetic analysis of mat K sequences representing all proposed subfamilies of Orchidaceae produced five moderately to well-supported clades. One of these clades, Apostasioideae, is sister to the clade formed by Vanilloideae, Cypripedioideae, Orchidoideae and Epidendroideae. High transition-transversion ratio and the absence of stop codons in the individual sequences suggest that mat K is at the transition from a possibly functional gene to a pseudogene in Apostasioideae, contrary to what is found in some other groups of Orchidaceae.Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/41641/1/606_2004_Article_133.pd

Quantitative importance of staminodes for female reproductive success in Parnassia palustris under contrasting environmental conditions.

The five sterile stamens, or staminodes, in Parnassia palustris act both as false and as true nectaries. They attract pollinators with their conspicuous, but non-rewarding tips, and also produce nectar at the base. We removed staminodes experimentally and compared pollinator visitation rate and duration and seed set in flowers with and without staminodes in two different populations. We also examined the relative importance of the staminode size to other plant traits. Finally, we bagged, emasculated, and supplementary cross-pollinated flowers to determine the pollination strategy and whether reproduction was limited by pollen availability. Flowers in both populations were highly dependent on pollinator visitation for maximum seed set. In one population pollinators primarily cross-pollinated flowers, whereas in the other the pollinators facilitated self-pollination. The staminodes caused increased pollinator visitation rate and duration to flowers in both populations. The staminodes increased female reproductive success, but only when pollen availability constrained female reproduction. Simple linear regression indicated a strong selection on staminode size, multiple regression suggested that selection on staminode size was mainly caused by correlation with other traits that affected female fitness. [ABSTRACT FROM AUTHOR