Support for an expanded family concept of Malvaceae within a recircumscribed order Mai vales: a combined analysis of plastid atpB and rbcL DNA sequences

Sequence analyses of the plastid genes atpB and rbcL support an expanded order Malvales. Within this alliance, core Malvales are clearly supported and comprise most genera that have previously been included in Sterculiaceae, Tiliaceae, Bombacaceae, and Malvaceae. Additional well supported malvalean alliances include the bixalean clade (Bixaceae, Diego-dendraceae, and Cochlospermaceae), the cistalean clade (Cistaceae, Dipterocarpaceae, and Sarcolaenaceae) and Thymelaeaceae (including Gonystyloideae and Aquilarioideae). Our results indicate sister-group relationships between (1) Neuradaceae and the cistalean clade; (2) Sphaerosepalaceae and Thymelaeaceae; (3) these two clades (1 and 2); and (4) all these and an alliance comprising the bixalean clade and core Malvales, but this pattern is weakly supported by the bootstrap. The affinities of Muntingiaceae and Petenaea are especially ambiguous, although almost certainly they are Malvales s.l. The traditional delimitation of families within core Malvales is untenable. Instead, we propose to merge Sterculiaceae, Tiliaceae and Bombacaceae with Malvaceae and subdivide this enlarged family Malvaceae into nine subfamilies based on molecular, morphological, and biogeographical data: (1) Byttnerioideae, including tribes Byttnerieae, Lasiopetaleae and Theobromeae (all of which have cucullate petals) and Hermannieae; (2) Grewioideae, including most genera of former Tiliaceae; (3) Tilioideae, monogeneric in our analysis; (4) Helicteroideae, comprising most of the taxa previously included in Helictereae, plus Mansonia, Triplochiton (indicating that apocarpy evolved at least twice within Malvaceae) and possibly Durioneae; (5) Sterculioideae, defined by apetalous, apocarpous, usually unisexual flowers with androgynophores; (6) Brownlowioideae, circumscribed as in previous classifications; (7) Dombeyoideae, expanded to include Burretiodendron, Eriolaena, Pterospermum, and Schoutmia; (8) Bombacoideae, corresponding to former Bombacaceae (without Durioneae) but including Fremontodendreae     

Phylogeny of the core Malvales: evidence from ndhF sequence data

The monophyly of the group comprising the core malvalean families, Bombacaceae, Malvaceae, Sterculiaceae, and Tiliaceae, was recently confirmed by molecular studies, but the internal structure of this clade is poorly understood. In this study, we examined sequences of the chloroplast ndhF gene (aligned length 2226 bp) from 70 exemplars representing 35 of the 39 putative tribes of core Malvales. The monophyly of one traditional family, the Malvaceae, was supported in the trees resulting from these data, but the other three families, as traditionally circumscribed, are nonmonophyletic. In addition, the following relationships were well supported: (1) a clade, /Malvatheca, consisting of traditional Malvaceae and Bombacaceae (except some members of tribe Durioneae), plus Fremontodendron and Chiranthodendron, which are usually treated as Sterculiaceae; (2) a clade, /Malvadendrina, supported by a unique 21-bp (base pair) deletion and consisting of /Malvatheca, plus five additional subclades, including representatives of Sterculiaceae and Tiliaceae, and Durionieae; (3) a clade, /Byttneriina, with genera traditionally assigned to several tribes of Tiliaceae, plus exemplars of tribes Byttnerieae, Hermannieae, and Lasiopetaleae of Sterculiaceae. The most striking departures from traditional classifications are the following: Durio and relatives appear to be more closely related to Helicteres and Reevesia (Sterculiaceae) than to Bombacaceae; several genera traditionally considered as Bombacaceae (Camptostemon, Matisia, Phragmotheca, and Quararibea) or Sterculiaceae (Chiranthodendron and Fremontodendron) appear as sister lineages to the traditional Malvaceae; the traditional tribe Helictereae (Sterculiaceae) is polyphyletic; and Sterculiaceae and Tiliaceae, as traditionally circumscribed, represent polyphyletic groups that cannot sensibly be maintained with their traditional limits for purposes of classification. We discuss morphological characters and conclude that there has been extensive homoplasy in characters previously used to delineate major taxonomic groups in core Malvales. The topologies here also suggest that /Malvatheca do not have as a synapormophy monothecate anthers, as has been previously supposed but, instead, may be united by dithecate, transversely septate (polysporangiate) anthers, as found in basal members of both /Bombacoideae and /Malvoideae. Thus, “monothecate” anthers may have been derived at least twice, independently, within the /Bombacoideae (core Bombacaceae) and /Malvoideae (traditional Malvaceae).     

Circumscription of the Malvales and relationships to other Rosidae: evidence from rbcL sequence data

The order Malvales remains poorly circumscribed, despite its seemingly indisputable core constituents: Bombacaceae, Malvaceae, Sterculiaceae, and Tiliaceae. We conducted a two-step parsimony analysis on 125 rbcL sequences to clarify the composition of Malvales, to determine the relationships of some controversial families, and to identify the placement of the Malvales within Rosidae. We sampled taxa that have been previously suggested to be within, or close to, Malvales (83 sequences), plus additional rosids (26 sequences) and nonrosid eudicots (16 sequences) to provide a broader framework for the analysis. The resulting trees strongly support the monophyly of the core malvalean families, listed above. In addition, these data serve to identify a broader group of taxa that are closely associated with the core families. This expanded malvalean clade is composed of four major subclades: (1) the core families (Bombacaceae, Malvaceae, Sterculiaceae, Tiliaceae); (2) Bixaceae, Cochlospermaceae, and Sphaerosepalaceae (Rhopalocarpaceae); (3) Thymelaeaceae sensu lato (s.l.); and (4) Cistaceae, Dipterocarpaceae s.l., Sarcolaenaceae (Chlaenaceae), and Muntingia. In addition, Neurada (Neuradaceae or Rosaceae) falls in the expanded malvalean clade but not clearly within any of the four major subclades. This expanded malvalean clade is sister to either the expanded capparalean clade of Rodman et al. or the sapindalean clade of Gadek et al. Members of Elaeocarpaceae, hypothesized by most authors as a sister group to the four core malvalean families, are shown to not fall close to these taxa. Also excluded as members of, or sister groups to, the expanded malvalean clade were the families Aextoxicaceae, Barbeyaceae, Cannabinaceae, Cecropiaceae, Dichapetalaceae, Elaeagnaceae, Euphorbiaceae s.l., Huaceae, Lecythidaceae, Moraceae s.l., Pandaceae, Plagiopteraceae, Rhamnaceae, Scytopetalaceae, Ulmaceae, and Urticaceae.     

梧桐科一些属的分类位置探讨

梧桐科（Ｓｔｅｒｃｕｌｉａｃｅａｅ）是锦葵目中的一个多型科,科的特征比较比样化。自从Ｅ．Ｐ．Ｖｅｎｔｅｎａｔ（１８３０）建立该科以来,对于该科范围和包含的属种数目,各国学者至今尚存在各种不同的看法。作者认为,火桐属（Ｅｒｙｔｈｒｏｐｓｉｓ）应当从梧桐属（Ｆｉｒｍｉａｎａ）中分出成为单独的属;午时花属（Ｐｅｎｔａｐｅｔｅｓ）不应归入锦葵科（Ｍａｌｖａｃｅａｅ）,应当置于梧桐科;田麻属（Ｃｏｒｃｈｏｒ     

Phylogenetic Position of <Emphasis Type="Italic">Corchoropsis</Emphasis> Siebold &; Zucc. (Malvaceae s.l.) Inferred from Plastid DNA Sequences

Recent phylogenetic research suggests that Malvaceae s.l. comprises formerly Tiliaceae, Byttneriaceae, Bombacaceae, and Sterculiaceae. Corchoropsis is traditionally included in Tiliaceae or Sterculiaceae and is distributed in China, Korea, and Japan. One to three species have been recognized for this genus. Phylogenetic relationships among the Malvacean taxa have been intensively studied with molecular data, and the evolution of their morphological characteristics has been re-interpreted accordingly. However, no Corchoropsis species have been included for their phylogenetic position. Here, three chloroplast coding regions—rbcL, atpB, and ndhF, from Corchoropsis psilocarpa and Corchoropsis crenata—were amplified and sequenced, then compared with other Malvacean taxa. This analysis of the three plastid gene sequences now places Corchoropsis species in Dombeyoideae, as previously proposed by Takeda (Bull Misc Inform Kew 365, 1912), Tang (Cathaya 4:131–150, 1992), and Bayer and Kubitzki (2003). Within Dombeyoideae, Corchoropsis forms a strongly supported sister relationship with the Dombeya–Ruizia clade.     

利用叶绿体基因组数据解析锦葵科梧桐亚科的系统位置和属间关系

分子系统学研究将传统梧桐科与锦葵科、木棉科和椴树科合并为广义锦葵科,并进一步分为9个亚科.然而,9个亚科之间的关系尚未完全明确,且梧桐亚科内的属间关系也未得到解决.为了明确梧桐亚科在锦葵科中的系统发育位置,厘清梧桐亚科内部属间系统发育关系,该研究对锦葵科8个亚科进行取样,共选取55个样本,基于叶绿体基因组数据,采用最大...     

The bicolor unit — homology and transformation of an inflorescence structure unique to coreMalvales

A broad comparative analysis reveals that the inflorescences of coreMalvales, familiesSterculiaceae, Tiliaceae, Bombacaceae andMalvaceae, include characteristic repeating units. The basic repeating unit is called bicolor unit (afterTheobroma bicolor, where it was first observed). It is determinate and bears three bracts, one of which is invariably sterile, whereas the others subtend lateral cymes or single flowers. Through the demonstration of intermediate steps in closely related taxa the triad of bracts within a bicolor unit and the trimerous malvalean epicalyx are shown to be homologous. Various possibilities for an origin of the bicolor unit are discussed. Bicolor units are variously arranged to form complete inflorescences. In many taxa they are terminal on modules that comprise two (or fewer) prophylls. These modules may be arranged in elongated anthocladia or condensed sympodia, which in turn may constitute components of higher order inflorescence structures. The presence of the bicolor unit or its derivatives linksSterculiaceae, Tiliaceae, Bombacaceae andMalvaceae. It is absent from all other families included in a broader defined orderMalvales and represents one of the rare morphological synapomorphies of coreMalvales. Furthermore, inflorescence morphology provides characters of systematic significance for various taxa within coreMalvales.     

Structure and occurrence of unusual fatty acids in minor seed oils

This review deals with the literature survey of the structure and occurrence of unusual and novel fatty acids in minor seed oils belonging to 37-botanical plant families. The plant families include—Anacardiaceae, Apocyanaceae, Araliaceae, Asclepediaceae, Bignoniaceae, Bombacaceae, Celastraceae, Chailletaceae, Compositae, Connaraceae, Coriaraceae, Cruciferae, Eleocarpaceae, Euphorbiaceae, Flacourtiaceae, Labiateae, Lauraceae, Leguminosae, Lythraceae, Malphighiaceae, Malvaceae, Olacaceae, Papaveraceae, Polygonaceae Protaceae, Rosaceae, Rutaceae, Santalaceae, Sapindaceae, Sapotaceae, Simarubiaceae, Sterculiaceae, Tiliaceae, Thymalaceae, Ulmaceae, Umbelliferae and Valerianaceae.     

Phylogenetic analysis of the Malvadendrina clade (Malvaceae s.l.) based on plastid DNA sequences

Phylogenetic relationships within Malvaceae s.l., a clade that includes the traditional families Bombacaceae, Malvaceae s.str., Sterculiaceae, and Tiliaceae, have become greatly clarified thanks to recent molecular systematic research. In this paper, we use DNA sequences of four plastid regions (atpB, matK, ndhF, and rbcL) to study relationships within Malvadendrina, one of the two major clades of Malvaceae s.l. The four data sets were generally in agreement, but five terminal taxa manifested highly unexpected affinities in the rbcL partition, and the non-coding sequences of the trnK intron were found to provide limited phylogenetic information for resolving relationships at the base of Malvadendrina. The remaining data strongly support the existence of six major clades within Malvadendrina: Brownlowioideae, Dombeyoideae, Helicteroideae, Malvatheca (comprising Bombacoideae and Malvoideae), Sterculioideae, and Tilioideae. These data also resolve the placement of two problematic taxa: Nesogordonia (in Dombeyoideae) and Mortoniodendron (in Tilioideae). The relationships among the six clades are not definitively resolved, but the best-supported topology has Dombeyoideae as sister to the remainder of Malvadendrina (posterior probability PP=80%) and Sterculioideae as sister to Malvatheca (PP=86%). This early branching position of Dombeyoideae is supported by similarities in floral characters between members of that clade and outgroup taxa in Byttnerioideae. Similarly, the sister-group relationship of Sterculioideae and Malvatheca receives support from androecial characteristics, like subsessile or sessile anthers and an absence of staminodes, shared by these two clades.     

Engineering aspects of biological ion channels—from biosensors to computational models for permeation

Molecular sequencing has helped resolve the phylogenetic relationships amongst the diverse groups of algal, fungal-like and protist organisms that constitute the Chromalveolate “superkingdom” clade. It is thought that the whole clade evolved from a photosynthetic ancestor and that there have been at least three independent plastid losses during their evolutionary history. The fungal-like oomycetes and hyphochytrids, together with the marine flagellates Pirsonia and Developayella, form part of the clade defined by Cavalier-Smith and Chao (2006) as the phylum “Pseudofungi”, which is a sister to the photosynthetic chromistan algae (phylum Ochrophyta). Within the oomycetes, a number of predominantly marine holocarpic genera appear to diverge before the main “saprolegnian” and “peronosporalean” lines, into which all oomycetes had been traditionally placed. It is now clear that oomycetes have their evolutionary roots in the sea. The earliest diverging oomycete genera so far documented, Eurychasma and Haptoglossa, are both obligate parasites that show a high degree of complexity and sophistication in their host parasite interactions and infection structures. Key morphological and cytological features of the oomycetes will be reviewed in the context of our revised understanding of their likely phylogeny. Recent genomic studies have revealed a number of intriguing similarities in host–pathogen interactions between the oomycetes with their distant apicocomplexan cousins. Therefore, the earlier view that oomycetes evolved from the largely saprotrophic “saprolegnian line” is not supported and current evidence shows these organisms evolved from simple holocarpic marine parasites. Both the hyphal-like pattern of growth and the acquisition of oogamous sexual reproduction probably developed largely after the migration of these organisms from the sea to land.     

Wood and bark anatomy of Muntingiaceae: A phylogenetic comparison within Malvales s. l.

Quantitative and qualitative data on wood and bark anatomy are given for Muntingia calabura L. and Dicraspidia donnell-smithii Standley. These data are compared with phylogenetic schemes, based on DNA analysis, in which Muntingiaceae belong to the “dipterocarp clade” within Malvales. The data are consistent with this hypothesis, although Muntingiaceae lack pit vestures in vessels, which are seen in the other malvalean families (Cistaceae, Dipterocarpaceae, Neuradaceae, Sarcolaenaceae, Thymeleaceae), and this may represent a loss of pit vestures. All families of the dipterocarp clade agree with both genera of Muntingiaceae in having tracheids as the imperforate tracheary element type (at least ancestrally), although fiber-tracheids also occur in some Dipterocarpaceae and Thymeleaceae. The large size of some malvalean families (with attendant greater diversity in character states) and a paucity of wood studies in those families make for difficulty in comparison of features such as axial parenchyma and ray types with those of Muntingiaceae; character states of these features are consistent with placement of Muntingiaceae in the dipterocarp clade of Malvales. Banded phloem fibers in bark of Muntingiaceae are much like those of other Malvales. Wood of Muntingiaceae is highly mesomorphic according to quantitative vessel features.     

On the Structural Regularity in Nucleobases and Amino Acids and Relationship to the Origin and Evolution of the Genetic Code

To explore how chemical structures of both nucleobases and amino acids may have played a role in shaping the genetic code, numbers of sp² hybrid nitrogen atoms in nucleobases were taken as a determinative measure for empirical stereo-electronic property to analyze the genetic code. Results revealed that amino acid hydropathy correlates strongly with the sp² nitrogen atom numbers in nucleobases rather than with the overall electronic property such as redox potentials of the bases, reflecting that stereo-electronic property of bases may play a role. In the rearranged code, five simple but stereo-structurally distinctive amino acids (Gly, Pro, Val, Thr and Ala) and their codon quartets form a crossed intersection “core”. Secondly, a re-categorization of the amino acids according to their β-carbon stereochemistry, verified by charge density (at β-carbon) calculation, results in five groups of stereo-structurally distinctive amino acids, the group leaders of which are Gly, Pro, Val, Thr and Ala, remarkably overlapping the above “core”. These two lines of independent observations provide empirical arguments for a contention that a seemingly “frozen” “core” could have formed at a certain evolutionary stage. The possible existence of this codon “core” is in conformity with a previous evolutionary model whereby stereochemical interactions may have shaped the code. Moreover, the genetic code listed in UCGA succession together with this codon “core” has recently facilitated an identification of the unprecedented icosikaioctagon symmetry and bi-pyramidal nature of the genetic code.     

New Mexican species of Byttneria (Sterculiaceae), Bakeridesia (Malvaceae), and Triumfetta (Tiliaceae)

Recent collections have brought to light the following new species from the states of Tabasco and Oaxaca, Mexico:Byttneria fluvialis (Sterculiaceae) from Tabasco;Bakeridesia amoena (Malvaceae), andTriumfetta calzadae (Tiliaceae) from Oaxaca.     

Pollen Morphology of Craigia with Reference to Its Systematic Position

Due to the unique floral and pollen morphology of the genus Craigia its systematic position, whether in the family Sterculiaceae or in the family Tiliaceae, has often been debated. In order to add more evidence for solving this problem, a comparative study on pollen morphology of the genus and 15 species of 3 tiliaceous genera, viz.: Tilia Linn., Hainania Merr. and Excentrodendron Chang. et Miau. was carried out under the light and scanning electron microscope. The result shows that the polarity, the structure and position of the aperture of Craigia pollen differ from those in Tiliaceae but are similar to those in Sterculiaceae. From palynology, it is more appropriate to place the genus under discussion in Sterculiaceae than in Tiliaceae.     

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.     

The host range and vector species of viruses from Cola chlamydantha K. Schum., Adansonia digitata L. and Theobroma cacao L

Seventeen species in the Tiliaceae, Sterculiaceae and Bombacaceae were tested for susceptibility to three virus isolates from naturally infected Cola chlamydantha trees and three from Adansonia digitata trees in Ghana. Seven species of Pseudococcidae were tested as vectors of the six isolates. These studies indicate that the Cola isolates should be classified with cocoa swollen shoot virus and those from Adansonia with cocoa mottle leaf virus.     

Homomorphic Self‐Incompatibility,with Stylar Rejection Site,in Luehea grandiflora Mart. and Zucc. (Tiliaceae‐Malvaceae s.l.)

Abstract: The breeding system of Luehea grandiflora (Tiliaceae‐Malvaceae s.l.) was investigated using hand pollinations and fluorescence microscopy studies of pollen tube growth. Although selfed flowers persisted for some 10 days, our study indicates that L. grandiflora is self‐incompatible, with self pollen tube inhibition in the upper style, as occurs in many taxa with homomorphic, gametophytic self‐incompatibility (GSI). L. grandiflora is only the second species reported within the Malvales with homomorphic stylar inhibition. This result is discussed within the context of a report for self‐compatibility in this species, and we also consider the phylogenetic implications for the occurrence of GSI in the family Malvaceae s.l.