Floral ontogeny in tribe Dalbergieae (Leguminosae: Papilionoideae):Dalbergia brasiliensis, Machaerium villosum s. l.Platymiscium floribundum, andPterocarpus rotundifolius

Floral organogenesis and development of the tropical legume treesDalbergia brasiliensis, Machaerium villosum, Platymiscium floribundum, andPterocarpus rotundifolius were studied using scanning electron microscopy. The aims were to compare ontogenies and to elucidate if floral ontogenetic data will provide new character states diagnostic of the tribe Dalbergieae, which is considered a basal papilionoid tribe and primarily defined on fruit characters. Organ inception is principally acropetal in all taxa studied. Carpel inception is, however, consistently precocious. InD. brasiliensis sepals are initiated in an order not previously reported in Papilionoideae. It may be considered modified helical. InP. rotundifolius the inner whorl of stamens initiate in an unusual way, this is lateral two stamens first, then the two abaxial ones, and last the adaxial one, opposed to the unidirectional order usually seen in Papilionoideae. Generally the differences in flower development among the studied genera appear at initiation and late stage in ontogenesis, rather than at mid-stage.     

Leaflet secretory structures of five taxa of the genus Zornia J.F. Gmel. (Leguminosae, Papilionoideae, Dalbergieae) and their systematic significance

A survey of the secretory structures of leaflets has been carried out for five taxa of the genus Zornia J.F. Gmel. to assess possible taxonomic value of the glands, taking into account that these taxa, Zornia curvata Mohlenbr., Z. gemella Willd. ex Vogel, Z. glabra Desv., Z. latifolia Sm. and Z. reticulata Sm., have overlapping diagnostic characters. The leaflet secretory structures of the five studied taxa of Zornia were mucilage epidermis, mucilage cavities, and idioblasts secreting phenolic compounds. Mucilage epidermis is found in all five aforementioned taxa. Mucilage cavities are observed on both epidermal surfaces of the leaflets in four taxa, the exception being Z. glabra, whose cavities occur only on the abaxial surface. Idioblasts secreting phenolic compounds were detected only in the mesophyll of Z. latifolia and Z. reticulata. The joint occurrence of mucilage epidermis with mucilage cavities seems to be of unifying value for the five taxa analyzed. The position of mucilage cavities and the occurrence of secretory idioblasts in the mesophyll of the leaflets are of diagnostic value, providing a key to enable identification of the taxa studied. This study reveals that the secretory structures provide important information in support of systematic studies of the Leguminosae. In relation to the five taxa of Zornia studied, current results suggest recognition of four species: Z. curvata, Z. glabra, Z. latifolia, and Z. reticulata.     

Zornia subsessilis (Leguminosae: Papilionoideae: Dalbergieae), a new species from Serra do Cabral, Minas Gerais, Brazil

Zornia subsessilis, a new species from Serra do Cabral, Minas Gerais State, Brazil, is described and illustrated on the basis of field and herbarium studies. It belongs to Zornia section Zornia of subgenus Zornia and it appears to be restricted to “campo rupestre” (rocky fields) within the “cerrado” biome. This new species is characterized by its sessile to subsessile leaves, fruits with articles 4 to 5 mm long and with bristles up to 3.5 mm long.     

Leaf anatomy of the genus Psoralea sensu stricto (Psoraleeae, Papilionoideae, Leguminosae)

Leaf anatomy is described from 17 species of Psoralea sensu stricto which includes four species of the genus Hallia Thunberg. Species of the two genera share many characters including the presence of rod-shaped crystals, similar stipule structure and a possible continuum in secretory cavity anatomy ranging from a small and non-trabeculate to a large and trabci ulate form. Hallia species are distinguished b the presence of large tannin cells in the bundle sheaths and a narrow lcngllv.width ratio of palisade cells. In the light of other floral evidence this is considered insufficient to separate the two genera, a view confirmed by cluster analysis. Thus, Salter's proposal of 1939 for their amalgamation is supported. Leaf anatomy of herbaceous species is compared with that of seedlings of Psoralea sp. to assess the possibility that the herbaceous species may have arisen through neoteny from species that are trees or shrubs. Comparative venation studies between stipules and scale leaves suggest that the scale leaf form arose from pinnate-leaved ancestors by leaf reduction.     

A widespread Neotropical Bradyrhizobium lineage associated with Machaerium and Desmodium (Papilionoideae)

Multilocus enzyme electrophoresis and ribosomal RNA gene sequencing were used to compare bacterial strains from the liana Machaerium milliflorum and the herb Desmodium axillare at two lowland forests 440 km apart in Panama and Costa Rica. Out of 27 multilocus genotypes (electrophoretic types or ETs) detected in the isozyme analysis, only two ETs were found in both locations. Nevertheless, these shared ETs were closely related to a cluster including 11 other common ETs that represented 72% and 80%, respectively, of all bacterial isolates sampled in Panama and Costa Rica. Isolates in this group from both locations and from both legume hosts all had identical sequences for a highly polymorphic 5 portion of 23S rRNA, while nine other ETs outside this cluster all had unique sequences. This indicates that both sites are occupied by one primary lineage of nodule symbionts that is both locally predominant and geographically widespread. However, this group coexists with numerous other rare bacterial lineages whose ecology, distribution, and relationships require further study.     

Cladistic analysis of chloroplast DNA restriction site characters in Andira (Leguminosae: Dalbergieae)

Chloroplast DNA (cpDNA) restriction site variation was examined in 29 accessions, representing 29 populations of 17 species of Andira (Leguminosae, Papilionoideae, tribe Dalbergieae). This sample spans the morphological and geographical diversity of this poorly known, woody genus of approximately 30 species, which occurs throughout the neotropics, with one species in Africa. Cladistic analysis of 38 restriction site characters generated 182 equally most-parsimonious trees. The strict consensus tree preserved four well-supported clades, which are incongruent with previous infrageneric classifications of Andira. None of these clades has been recognized by previous workers, and three are not marked by any major morphological innovations. There were few restriction site differences between closely related species, but two (possibly three) species showed intraspecific cpDNA polymorphism, which probably reflects processes of hybridization with subsequent introgression, or lineage sorting.     

Micropropagation of adult Swainsona formosa (Leguminosae: Papilionoideae: Galegeae)

Summary Explants of axillary buds excised from mature adult stems of Swainsona formosa (G. Don) J. Thompson (syn. Clianthus formosus) were cultured on Murashige and Skoog medium supplemented with a range of auxins, cytokinins, and sucrose concentrations. Auxins did not increase shoot or bud numbers above controls, and 2,4-dichlorophenoxyacetic acid was the only auxin to significantly increase callus production. Benzyladenine or thidiazuron incorporated into the medium at 0.1 μM stimulated shoot and bud production, and shoot growth occurred following removal of cytokinins from the medium after 4 wk. Shoot number increased linearly with sucrose concentration up to 40 g l⁻¹, but shoot height and the number of cytokinin-induced buds were optimal at sucrose levels of 20–30 g l⁻¹. Roots were initiated in vitro following treatment of cuttings with 0.1% indole-3-butyric acid and 0.1% α-naphthaleneactic acid. Plantlets were successfully established in soil but were plagiotropic and exhibited distichous phyllotaxy.     

Pollen morphology in relation to pollinators in Papilionoideae (Leguminosae)

The surface sculpturing of the pollen of some species of the taxonomically widely separated genera Harpalyce (tribe Brongniartieae), Camoensia (tribe Sophoreae), Millettia (tribe Tephrosieae), and of the monotypic Dahlstedtia (tribe Tephrosieae) which have large red or white flowers adapted for pollination by birds or bats, is coarsely rugulate or verrucate. Related taxa with small insect pollinated flowers have pollen with simple reticulate or perforate surface sculpturing. The exine stratification of Alexa and Castanospermum (tribe Sophoreae), genera with large red bird-flowers, is complex with a layer of tectal columellae and differs from that of other genera in the tribe Sophoreae which have a normal pollen wall structure. These modifications of pollen structure and sculpture appear to be the result of convergent evolution and a secondary adaptation to pollination. The taxonomic and functional significance of the observations are briefly discussed.     

The genus Cremosperma (Gesneriaceae) in Ecuador

A key to the 10 species of Cremosperma occurring in Ecuador is presented, including four new species that are described: C. ecuadoranum, C. humidum, C. muscicola and C. reldioides . A new variety is C. hirsutissimum var. glabrum. Cremosperma demissum and C. album , originally described from Ecuador and Colombia, respectively, are reduced to C. hirsutissimum var. demissum and C. hirsutissimum var. album. Cremosperma pusillum var. ecuadorense is transferred to C. hirsutissimum var. hirsutissimum . The distribution of Cremosperma is discussed.     

Flower development in Abrus precatorius (Leguminosae: Papilionoideae: Abreae) and a review of androecial characters in Papilionoideae

The jequirity bean (Abrus precatorius) is well known because of its shiny black and red coloured seeds and because of the poison (abrin) it contains. The genus Abrus is placed in a monogeneric tribe Abreae which is placed in a relatively isolated systematic position at the base of Millettieae. To contribute to a better understanding of this taxon, a detailed ontogenetic and morphologic analysis of its flowers is presented. Floral primordia are subtended by an abaxial bract and preceded by two lateral bracteoles which are formed in short succession. Sepal formation is unidirectional starting abaxially. All petals are formed simultaneously. The carpel is formed concomitantly with the outer (antesepalous) stamen whorl, which arises unidirectionally, starting in an abaxial position. In the inner, antepetalous stamen whorl two abaxial stamens are formed first, followed by two lateral stamen primordia. The adaxial, antepetalous position remains organ free (i.e. this stamen is lost). Later in development the nine stamen filaments fuse to form an adaxially open sheath. The filament bases of the two adaxial outer-whorl stamens grow inwards, possibly to provide stability and to compensate for the lost stamen. In the mature flower a basal outgrowth can be found in the position of the lost stamen. However this is more likely to be an outgrowth of the filament sheath rather than a remnant of the lost stamen. These ontogenetic patterns match in parts those found in other Millettieae (unidirectional formation of sepals and stamens, simultaneous petal formation). In contrast, the complete loss of a stamen is rather unusual and supports the isolated position of Abreae and probably justifies (among other characters) its tribal status. A review of androecial characters shows that androecial merosity is on the one hand extremely variable among Leguminosae, varying from a single stamen per flower to more than 500. On the other hand it is noteworthy that the number of stamens becomes stabilised in more derived Papilionoideae such as the large non-protein-amino-acid-accumulating clade (NPAAA clade). This indicates that the androecium has played an important role in the success of a major part of Leguminosae.     

Development and cytology of pollen in Dahlstedtia Malme (Leguminosae: Papilionoideae)

Microsporogenesis, chromosome number, meiotic behaviour and meiotic index were investigated in Dahlstedtia pinnata and D . pentaphylla , two legume species occurring largely in Brazil, in order to ascertain whether the pollen could limit fertilization events. Archesporial cells originate primary sporogenous and anther wall precursor cells, the tapetum is uniseriate, uninucleate and glandular. Tetrads are tetrahedric or decussate, and cytokinesis is of the simultaneous type. Mature pollen grains are tricolpate and bicellular. No abnormalities in microsporogenesis were found. In both species the chromosome number is n = 11, a number not reported previously. The base number for Dahlstedtia is also 11, because cytological observations include both species of Dahlstedtia . D. pentaphylla has a higher meiotic index and lower individual variation values, and it is considered meiotically stable. Its pollen grains do not limit fertilization. D. pinnata has a lower meiotic index, and the pollen is one of the factors which limit fertilization. Furthermore, D. pinnata has numerous adventitious shoots, which suggest that vegetative propagation is important in its reproductive process. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 461–471.     

The genus Archidendron (Leguminosae - Mimosoideae)

An account is given of the morphology and the taxonomy of the Asian, Australian and Pacific genus Archidendron (Leguminosae - Mimosoideae). A new infrageneric classification based on morphological data is presented, the genus being subdivided in 8 series. The phylogeny of the genus is discussed, the base of discussion being all available morphological, palynological and wood-anatomical characters. The presence/absence of stipules, the length of the staminal tube compared with that of the corolla-tube, the sessile/stipitate ovary(-ies), the morphology of the pods and the wood-anatomy have been particularly useful in determining the evolutionary trends within the genus. Analyses of the geographical range of selected character states are presented. The data suggest a Central - W. Malesian origin of the genus. The series endemic to the E. Malesian - Australian area have probably evolved more recently. The pluricarpellate condition of the flowers in several species endemic to the E. Malesian and Australian area is considered to be a derived character state. The following new taxa are proposed: Ser. Calycinae Nielsen, ser. Ptenopae Nielsen, ser. Bellae Nielsen, Archidendron falcatum Nielsen, A. cockburnii Nielsen, A. sabahense Nielsen, A. fagifolium (Bl. ex Miq.) Nielsen var. borneense Nielsen, A. kunstleri (Prain) Nielsen subsp. ashtonii Nielsen, A. ellipticum (Bl.) Nielsen subsp. cordifoliolatum Nielsen. New combinations are proposed in the Malesian species formerly referred to Abarema, Zygia and Morolobium by Kostermans. Keys to and an enumeration of the species are presented.     

Breeding systems and pollination inVigna minima (Leguminosae,Papilionoideae)

Four types of floral breeding systems—(i) chasmogamy, (ii) aerial pseudocleistogamy, (iii) subterranean pseudocleistogamy and (iv) obligate subterranean true cleistogamy—are observed in the populations ofVigna minima inhabiting the Western Ghats (India). Five categories of phenotypes are recognized based on the number and kinds of floral breeding systems found in a given individual. The frequencies of different categories of phenotypes not only show intra- and interpopulation variation, but also fluctuate from generation to generation suggesting differences in the genetic structure of populations. This polymorphism in the breeding system of a single species is unique and may be adaptive. Obligate subterranean true cleistogamy and amphicarpy appear to be adaptations to jungle fires and soil erosion.—The flowers are of the flag-blossom type and insect visitors act as tripping agents. The tripping mechanism together with the polymorphic floral breeding system result in a balanced mixture of selfing and outcrossing. Such a recombination system may enhance the fitness ofV. minima which is essentially a colonizing species.     

The genus Archidendron (Leguminosae – Mimosoideae)

An account is given of the morphology and the taxonomy of the Asian, Australian and Pacific genus Archidendron (Leguminosae – Mimosoideae). A new infrageneric classification based on morphological data is presented, the genus being subdivided in 8 series. The phylogeny of the genus is discussed, the base of discussion being all available morphological, palynological and wood–anatomical characters. The presence/absence of stipules, the length of the staminal tube compared with that of the corolla–tube, the sessile/stipitate ovary(–ies), the morphology of the pods and the wood–anatomy have been particularly useful in determining the evolutionary trends within the genus. Analyses of the geographical range of selected character states are presented. The data suggest a Central – W. Malesian origin of the genus. The series endemic to the E. Malesian – Australian area have probably evolved more recently. The pluricarpellate condition of the flowers in several species endemic to the E. Malesian and Australian area is considered to be a derived character state. The following new taxa are proposed: Ser. Calycinae Nielsen, ser. Ptenopae Nielsen, ser. Bellae Nielsen, Archidendron falcatum Nielsen, A. cockburnii Nielsen, A. sabahense Nielsen, A. fagifolium (Bl. ex Miq.) Nielsen var. borneense Nielsen, A. kunsrteri (Prain) Nielsen subsp. ashtonii Nielsen, A. ellipticum (Bl.) Nielsen subsp. cordifoliolatum Nielsen. New combinations are proposed in the Malesian species formerly referred to Abarema, Zygia and Morolobium by Kostermans. Keys to and an enumeration of the species are presented.     

Geographical diversification of the genus Cicer (Leguminosae: Papilionoideae) inferred from molecular phylogenetic analyses of chloroplast and nuclear DNA sequences

Cicer L. (Leguminosae: Papilionoideae) consists of 42 species of herbaceous or semi-shrubby annuals and perennials distributed throughout the temperate zones of the Northern Hemisphere. The origin and geographical relationships of the genus are poorly understood. We studied the geographical diversification and phylogenetic relationships of Cicer using DNA sequence data sampled from two plastid regions, trnK / matK and trnS - trnG , and two nuclear regions, the internal transcribed spacer (ITS) and external transcribed spacer (ETS) regions of nuclear ribosomal DNA, from 30 species. The results from the phylogenetic analyses of combined nuclear and chloroplast sequence data revealed four well-supported geographical groups: a Middle Eastern group, a West-Central Asian group, an Aegean–Mediterranean group, and an African group. Age estimates for Cicer based on methods that do not assume a molecular clock (for example, penalized likelihood) demonstrate that the genus has a Mediterranean origin with considerable diversification in the Miocene/Pliocene epochs. Geological events, such as mountain orogenesis and environmental changes, are major factors for the dispersal of Cicer species. The early divergence of African species and their geographically distinct region in the genus suggest a broader distribution pattern of the genus in the past than at present.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 154 , 175–186.