Monophyletic subgroups of the tribe Millettieae (Leguminosae) as revealed by phytochrome nucleotide sequence data

Phylogenetic analysis of phytochrome (PHY) genes reveals the identity and relationships of four PHY loci among papilionoid Leguminosae. A phylogenetic analysis of loci combined according to species suggests that most of the tribe Millettieae belongs to one of two monophyletic clades: the Derris–Lonchocarpus or the Tephrosia clade. Together these two form a monophyletic group that is sister to a lineage represented by Millettia grandis of Millettia sect. Compresso-gemmatae. Collectively, this large monophyletic group is referred to as the Millettieae-core group, which based on our sampling, includes species of Millettieae that do not accumulate the nonprotein amino acid canavanine and that mostly have pseudoracemose or pseudopaniculate inflorescences. This new phylogenetic framework assists in targeting additional taxa for future sampling. For example, the “American Derris” (Deguelia), which accumulate canavanine, might not be members of the Millettieae core group. Afgekia is also predicted not to be a member because it accumulates canavanine and has an inflorescence of terminal racemes. PHY gene analysis specifically reveals that certain genera traditionally classified in Millettieae are actually distantly related to the Millettieae core group, such as Austrosteensia, Callerya, Craibia, Cyclolobium, Fordia, Platycyamus, Poecilanthe, and Wisteria.     

Phylogenetic systematics of the tribe Millettieae (Leguminosae) based on chloroplast trnK/matK sequences and its implications for evolutionary patterns in Papilionoideae

Phylogenetic relationships in the tribe Millettieae and allies in the subfamily Papilionoideae (Leguminosae) were reconstructed from chloroplast trnK/matK sequences. Sixty-two accessions representing 57 traditionally recognized genera of Papilionoideae were sampled, including 27 samples from Millettieae. Phylogenies were constructed using maximum parsimony and are well resolved and supported by high bootstrap values. A well-supported "core Millettieae" clade is recognized, comprising the four large genera Millettia, Lonchocarpus, Derris, and Tephrosia. Several other small genera of Millettieae are not in the core Millettieae clade. Platycyamus is grouped with Phaseoleae (in part). Ostryocarpus, Austrosteenisia, and Dalbergiella are neither in the core Millettieae or Phaseoleae clade. These taxa, along with core Millettieae and Phaseoleae, form a monophyletic sister group to Indigofereae. Cyclolobium and Poecilanthe are close to Brongniartieae. Callerya and Wisteria belong to a large clade that includes all the legumes that lack the inverted repeat in their chloroplast genome, which confirms previous rbcL and phytochrome gene family phylogenies. The evolutionary history of four characters was examined in Millettieae and allies: the presence of canavanine, inflorescence types, the dehiscence of pods, and the presence of winged pods. trnK/matK sequence analysis suggests that the presence of a pseudoraceme or pseudopanicle and the accumulation of nonprotein amino acids are phylogenetically informative for Millettieae and allies with only a few exceptions.     

Leontidine-type quinolizidine alkaloids in Orphanodendron (Leguminosae)

Herbarium leaf fragments of the two known species of Orphanodendron (Leguminosae), O. bernalii and O. grandiflorum, were found to contain the quinolizidine alkaloids camoensine (1), camoensidine (2) and guianodendrine (3), supporting the recent phylogenetic placement of the genus in the genistoid clade of subfamily Papilionoideae rather than its traditional placement in subfamily Caesalpinioideae.     

Four new Brazilian species of Lonchocarpus (Leguminosae, Papilionoideae, Millettieae)

Lonchocarpus castaneifolius, L. grazielae, L. longiunguiculatus, and L. montanus from Brazil are described and illustrated based on field and herbarium studies. The first species belongs to Lonchocarpus sect. Punctati and is restricted to northeastern Brazil (Bahia state). The other three are included in sect. Laxiflori. Lonchocarpus grazielae is restricted to the southern coastal region (Santa Catarina state), L. montanus is found in northeastern Brazil (Bahia and Piauí states), the west-central region (Goiás and Tocantins states) and the southeast (Minas Gerais state), and L. longiunguiculatus occurs in northeastern (Bahia state) and southeastern (Minas Gerais state) Brazil.     

Millettia sapindifolia is a synonym of Derris yunnanensis (Leguminosae: Millettieae)

Millettia and Derris are two taxonomically complicated genera of the tribe Millettieae (Leguminosae), and when only flowering material is available they are not easily distinguished from each other. A critical examination of literature and specimens as well as a field survey show that Millettia sapindifolia T. C. Chen, a Chinese species described on the basis of two flowering collections, is conspecific with Derris yunnanensis Chun & F. C. How. The two are closely similar in various characters, such as the inflorescence type, monadelphous stamens, branches with longitudinal ridges, and size, shape, number and hairiness of leaflets. Therefore, the former is here reduced to a synonym of the latter.     

Quinolizidine alkaloid status of Styphnolobium and Cladrastis (Leguminosae)

Reports of quinolizidine alkaloids in Styphnolobium Schott and Cladrastis Raf. (Leguminosae) conflict with their position in recent molecular phylogenies because they are not members of a major clade of quinolizidine alkaloid-accumulating taxa. The alkaloid status of these two genera was therefore re-investigated using gas chromatography–mass spectrometry. Quinolizidine alkaloids could not be detected in extracts of leaves, flowers or seeds of S. japonicum (L.) Schott, nor in leaves of S. affine (Torrey & A. Gray) Walp., C. delavayi (Franch.) Prain, C. kentukea (Dum.-Cours.) Rudd or C. platycarpa Mak. In contrast, Calia secundiflora (Ortega) Yakovlev, also currently placed outside the major clade of quinolizidine alkaloid-producing genera in molecular phylogenies, was confirmed to accumulate a range of quinolizidine alkaloids.     

Observations on the vascular anatomy of the tribe Vicieae (Leguminosae)

The tribe Vicieae (Adanson) DC. ex Ser. is characterized by an unusual type of stele in which the lateral leaf-traces are present as cortical bundles in the internode below the insertion of their leaf. In most members of the tribe a completely fresh pair of cortical bundles is substituted at each node, but in ten species of Vicia L. there is only partial replacement. The group thus delimited also possesses some distinctive non-anatomical features.
The behaviour of the three leaf-traces at the node also varies throughout the tribe. This variation is shown to be correlated with differences in the shape of the stipules and their size in relation to the rest of the leaf.     

Stirtonia, a new genus of the tribe Podalyrieae (Leguminosae) from South Africa

The genus Stirtonia is described to accommodate three anomalous species, hitherto placed in Podalyria . The new genus differs from Podalyria mainly in the decussate (geminate) inflorescence structure (racemose in Podalyria ), the yellow flower colour (purple, pink or white in Podalyria ), the non-fleshy rim-aril of the seeds (seed aril invariably fleshy and collar-like in Podalyria ) and in the diagnostically different combination of quinolizidine alkaloids. The major alkaloidal metabolites are carboxylic acid esters of lupanine and virgiline, but these compounds are totally absent in Podalyria . The morphology of inflorescences, leaves, flowers and seeds are described and illustrated. Chromosome numbers (2n = 18) are reported for the first time. A formal taxonomic treatment of the three species, S. tayloriana, S. chrysantha and S. insignis , is presented.     

Seed morphology: an addition to the taxonomy of Tephrosia (Leguminosae, Papilionoideae, Millettieae) from South America

Seed morphology was studied using scanning electron microscopy in 13 species, one subspecies and one forma of Tephrosia Pers. (Leguminosae, Papilionoideae, Millettieae) occurring in South America. Macromorphological and micromorphological characters were examined, including seed form, colour and size, testa pattern, reticulum anticlinal wall, boundaries of anticlinal wall and hilum form. Crested and simple-reticulate testa patterns were predominant. A foveolate pattern, multifoveolate pattern and subgrooved pattern are all recorded for the first time in this genus. The macromorphological characters displayed continuous variation in shape and size and are thus are not significant for species separation. The data obtained in this study together with data from the literature provide additional characters to help classify the genus. A key to the taxa under study is presented.     

Stirtonanthus, a new name for Stirtonia (Leguminosae, tribe Podalyrieae)

The new generic name Stirtonanthus is proposed to replace the illegitimate Stirtonia Van Wyk & Schutte (1994), which proves to be a later homonym of Stirtonia A. L . Smith (1926) and Stirtonia R. Brown (1900).     

Distribution of quinolizidine alkaloid types in nine Ormosia species (Leguminosae-Papilionoideae)

Quinolizidine alkaloids were surveyed in 22 plant samples, representing nineOrmosia species and up to five different plant parts per species, using combined gas chromatography and mass spectroscopy. The detected alkaloids were classified into 40 structural types. There was a remarkable degree of dissimilarity of alkaloid-type profiles between any two plant samples, including those obtained from the same species and even from a single tree. The similarity of alkaloid-type profiles among the studied samples varied between 0% and 79% (Jaccard similarity coefficient). Of chemotaxonomic interest was the finding of acosmine inO. isthmensis, which previously had been reported only from the related genusAcosmium. Furthermore, the alkaloid-type profile ofO. panamensis seeds was distinct from that of all other samples, supporting the hypothesis that this species is only distantly related to the other Latin AmericanOrmosia species.     

Inflorescence and floral development of Dahlstedtia species (Leguminosae: Papilionoideae: Millettieae)

Inflorescence and floral development of two tropical legume trees, Dahlstedtia pinnata and Dahlstedtia pentaphylla, occurring in the Atlantic Forest of south-eastern and southern Brazil, were investigated and compared with other papilionoids. Few studies have been made of floral development in tribe Millettieae, and this paper is intended to fill that gap in our knowledge. Dahlstedtia species have an unusual inflorescence type among legumes, the pseudoraceme, which comprises axillary units of three or more flowers, each with a subtending bract. Each flower exhibits a pair of opposite bracteoles. The order of flower initiation is acropetal; inception of the floral organs is as follows: sepals (5), petals (5), carpel (1) plus outer stamens (5) and finally inner stamens (5). Organ initiation in sepal, petal and inner stamen whorls is unidirectional; the carpel cleft is adaxial. The vexillum originates from a tubular-shaped primordium in mid-development and is larger than other petals at maturity, covering the keels. The filament tube develops later after initiation of inner-stamen primordia. Floral development in Dahlstedtia is almost always similar to other papilionoids, especially species of Phaseoleae and Sophoreae. But one important difference is the precocious ovule initiation (open carpel with ovules) in Dahlstedtia, the third citation of this phenomenon for papilionoids. No suppression, organ loss or anomalies occur in the order of primordia initiation or structure. Infra-generic differences in the first stages of ontogeny are rare; however, different species of Dahlstedtia are distinguished by the differing distribution pattern of secretory cavities in the flower.     

Seed-coat anatomy of the non-pleurogrammic seeds in the tribe Ingeae (Leguminosae,Mimosoideae)

An anatomically complex structure of the seed-coat is a general characteristic of all members of the Leguminosae. Nonetheless, certain genera exhibit a particular type of seed, the “overgrown” seen that has been defined by a developmental criterion and by more simple anatomical features of the seed-coat. Only the last criterion seems suitable for unambiguously distinguishing this type of seed, which is here referred to as “overgrown-like.” This type appears to be apomorphic in the Ingeae (and probably in all the Leguminosae) and likely results from a heterochronic loss of tissue differentiation. Variations in this character may be useful at the generic level, and detailed anatomical observations reveal the occurrence of three distinct patterns. The high degree of correlation with other characters suggests that overgrown-like seeds have evolved separately at least three times in the Ingeae and thatPithecellobium s.s. may be polyphyletic. The overgrown-like seeds are likely to be an adaptive response to wet tropical climates.     

Comparative biosynthesis of quinolizidine alkaloids from dl-lysine in five species of Leguminosae

Five species of the leguminosae produced radioactive lupine alkaloids after feeding with dl-lysine-[2-¹⁴C]. Saturated alkaloids and compounds with a pyridone ring were radioactive. The specific radioactivity of the isolated compounds provides evidence that conversion of lysine into the saturated alkaloids, and by further oxidation to compounds both with a pyridone ring and without a d ring.     

Genome relationship among nine species of Millettieae (Leguminosae: Papilionoideae) based on random amplified polymorphic DNA (RAPD)

Random amplified polymorphic DNA (RAPD) marker was used to establish intergeneric classification and phylogeny of the tribe Millettieae sensu Geesink (1984) (Leguminosae: Papilionoideae) and to assess genetic relationship between 9 constituent species belonging to 5 traditionally recognized genera under the tribe. DNA from pooled leaf samples was isolated and RAPD analysis performed using 25 decamer primers. The genetic similarities were derived from the dendrogram constructed by the pooled RAPD data using a similarity index, which supported clear grouping of species under their respective genera, inter- and intra-generic classification and phylogeny and also merger of Pongamia with Millettia. Elevation of Tephrosia purpurea var. pumila to the rank of a species (T. pumila) based on morphological characteristics is also supported through this study of molecular markers.     

The delimitation of the tribe Vicieae (Leguminosae) and the relationships of Cicer L.

A survey of morphological, anatomical, karyological and chemical characters has been carried out, centred on the Vicieae but extending to the neighbouring tribes Trifolieae and Ononideae. The results show that Cicer , traditionally a member of the Vicieae, has more in common with genera of the Trifolieae and Ononideae than with the rest of the Vicieae. It is proposed that Cicer should be removed from the Vicieae and recognized as the monogeneric tribe Cicereae Alef. The tribe Vicieae sensu stricto, a well-defined natural group, is delimited and described. Phylogenetic relationships of the Cicereae are discussed.     

A revised classification of the tribe Phaseoleae (Leguminosae: Papilionoideae), and its relation to canavanine distribution

The subtribal classification of the tribe Phaseoleae has drifted into disorder since Bentham's (1865) scheme in the Genera Plantarum. A revised classification is here proposed, which for the first time accounts for all of the currently recognized genera. This classification has seven subtribes: Cajaninae, Diocleinae, Kennediinae, Phaseolinae, Glycininae, Ophrestiinae, and Erythrininae, of which the Ophrestiinae is new. A survey for the free amino acid, canavanine, shows that it is most often present in the Diocleinae and Kennediinae and frequently absent in the rest of the tribe.     

The morphology of the wet, non-papillate (WN) stigma form in the tribe Caesalpinieae (Gaesalpinioideae: Leguminosae)

Stigmas in 24 species of 11 genera, classified in three groups, Caesalpinia, Peltophorum and Dimorphandra, of the tribe Caesalpinieae have been examined by light and scanning electron microscopy. All are of the WN (wet, non-papillate) form. The receptive stigmatic surface comprises a crater, generally at the apex of the style, which in fresh flowers is full of clear fluid. The crater rim may be fringed with non-receptive hairs, and its shape and depth are variable between species. Extreme forms are found in Caesalpinia sappan where crater hairs are short and the crater funnel-shaped and very deep, and C. vesicaria and Delonix regia where crater hairs are long and the crater appears very shallow.