Vidal de Freitas Mansano

Floral ontogeny of Lecointea, Zollernia, Exostyles, and Harleyodendron (Leguminosae: Papilionoideae: Swartzieae S.L.)

Floral initiation and development were examined using scanning electron microscopy in Exostyles venusta, Harleyodendron unifoliolatum, Lecointea hatschbachii, and Zollernia ilicifolia. Common features include (1) unidirectional sepal initiation, (2) simultaneous petal initiation, (3) unidirectional initiation of each stamen whorl (except in the antesepalous whorl in Lecointea and Exostyles), (4) overlap in time of initiation of the two stamen whorls, and (5) initiation of the carpel concurrently with petals. Significant developmental features include (1) the first sepal median abaxial in all except Lecointea where it is non-median abaxial; (2) intraspecific variation in petal aestivation in Exostyles, Harleyodendron, and Lecointea; (3) initiation of antepetalous stamens before the antesepalous ones in Zollernia, Exostyles, and Lecointea; and (4) ovule initiation before the carpel margins are fused in Exostyles. The stamen sequence has not been found in any other legumes. The following late developmental events distinguish the four genera from each other: copious hairs hold the anthers together as a domelike structure at anthesis in Harleyodendron; zygomorphy in Zollernia results from differing petal reflexion; late hypanthium in Exostyles, Lecointea, and Holocalyx (no hypanthium in Harleyodendron or Zollernia); and reflexed sepal lobes in Exostyles, Harleyodendron, and Zollernia but not in Holocalyx and Lecointea. The genera studied here are ontogenetically more similar to taxa of Sophoreae than to other Swartzieae that have been investigated. None of the taxa studied here has a ring meristem, the structure that characterizes the remaining swartzioid taxa studied elsewhere.

The taxonomy of some Swartzieae (Leguminosae, subfam. Papilionoideae) from southeastern Brazil

Three new species of Swartzieae are described and illustrated:Swartzia alternifoliolata, S. capixabensis, andZollernia cowanii. Swartzia apetala var.blanchetii and var.subcordata are considered to be synonyms ofS. apetala var.apetala, andS. grazielana a synonym ofS. macrostachya var.macrostachya. Keys to southeastern Brazil members ofSwartzia andZollernia are provided.

Composition of the Lecointea clade (Leguminosae, Papilionoideae, Swartzieae), a re-evaluation based on combined evidence from morphology and molecular data

We investigated morphological and DNA sequence data for studying the composition and phylogenetic relationships of the Lecointea clade sensu Herendeen and for testing its monophyly. Twenty nine representing 14 genera wereanalyzed, including all members of the Lecointea clade and some species of the genera Acosmium, Luetzelburgia, Sweetia, Vatairea and Vataireopsis, Ateleia herbert-smithii, Bocoa mollis, and Myrocarpus frondosus were included as outgroup. Matrices were analyzed using maximum parsimony. Analyses of morphological data, chloroplast DNA trnL sequence data, and combined datasets resulted in similar tree topologies. The Lecointea clade sensu Herendeen, with the additional inclusion of Uribea, is monophyletic in all analyses. Exostyles and Harleyodendron belong to the Lecointea clade and not to the Vatairea clade as recently proposed by Pennington & al.

Swartzia Schreb. (Leguminosae: Papilionoideae: Swartzieae): a taxonomic study of the Swartzia acutifolia complex including a new name and a new species from southeastern Brazil

A taxonomic survey of the Swartzia acutifolia complex and a description of a new species, Swartzia linharensis, are presented. Swartzia acutifolia var. leiogyna is treated as S. submarginata var. leiogyna, S. acutifolia var. parvipetala as S. parvipetala, S. acutifolia var. submarginata as S. submarginata var. submarginata and S. acutifolia var. ynesiana as S. oblata. A key to the species from southeastern Brazil is provided.

Comparative development of rare cases of a polycarpellate gynoecium in an otherwise monocarpellate family, Leguminosae

PREMISE OF THE STUDY Apocarpy (i.e., free carpels) is considered to be the basal condition for ovary development in angiosperms. Yet it only occurs in 10% of angiosperm species, of which another 10% are monocarpellate. Most legume flowers are monocarpellate. Species with polycarpellate gynoecia occur in about 15 genera with most representatives in Mimosoideae. In the present study, we analyze legumes with polycarpellate flowers with the aim of improving our understanding of gynoecium evolution. METHODS Flowers of nine legume species from five genera were analyzed using a scanning electron microscope (SEM). KEY RESULTS In Leguminosae, carpels usually form as individual primordia or protuberances. Inga congesta differs slightly from this pattern in that the central apex bulges outward before the formation of individual carpel primordia. While legumes usually develop entirely plicate carpels, flowers of Acacia celastrifolia and Inga bella show an intermediate type of carpel morphology with a distal plicate zone and a small proximal ascidiate zone. Carpels in Inga congesta and Archidendron glabrum are sometimes slightly fused at the ovary base. The orientation of carpel clefts seems to reflect the floral symmetry. They are directed to the floral center in mimosoids and caesalpinioids, whereas in Swartzia dipetala carpel clefts are oriented to the adaxial side. CONCLUSIONS Polycarpelly arose at least seven times independently in Leguminosae. The polycarpellate condition appears to be correlated with polyandry, and in most instances, it is accompanied by a profound change in floral organization from a closed to an open system.

A Revision of the South American Genus Zollernia Wied-Neuw. & Nees (Leguminosae, Papilionoideae, Swartzieae)

Zollern, in southern Germany (Lewis & Foster 1997). The genus is characterised by simple leaves, corolla generally with 5 petals, anthers subulate or lanceolate, longer than the filaments, and the lack of a hypanthium. The fruits are indehiscent, and generally globose and fleshy. The genus originally contained 2 species: Z. falcata Wied-Neuw. & Nees and Z. splendens Wied-Neuw. & Nees. Vogel (1837) described Z. discolor and transferred Coquebertia ilicifolia Brongn. to Zollernia. Two other species, Z. houlletiana and Z. vogelii, were described by Tulasne (1844). Of these six species of Zollernia, Bentham (1870) recognised five as occurring in Brazil. In addition he described Z. latifolia and Z. securidacifolia as new species, placed Z. discolor as a synonym of Z. falcata, and Z. houlletiana and Z. vogelii as synonyms of Z. ilicifolia. He did not, however, provide a key to the species. Huber (1910) described Z. paraensis as a new species from Parai, Brazil. Cowan (1952), described Z. kanukuensis from Guyana, and mentioned the difficulty in delimiting the species of Zollernia. Yakovlev (1976) presented a taxonomic survey of Zollernia with a key to the species recognised by him. His key was based, however, on vegetative characters which vary intraspecifically and are not mutually exclusive. Yakovlev (1976) described Z. glaziovii Yakovlev and placed Z. falcata as a synonym of Z. glabra, a species first described by Sprengel (1821) as Krameria glabra. He also considered Z. mocitayba Allemao ex Emygdio as a synonym of Z. glabra and Z. securidacifolia Benth. as a synonym of Z. ilicifolia (Brongn.) Vogel. Yakovlev remarked that he had no information about Z. grandifolia Schery, a species described for the Flora of Venezuela by Schery (1952). Carvalho & Barneby (1993), in a study of the genus Zollernia in Bahia, Brazil described Z. magnifica A. M. de Carvalho & Barneby and Z. modesta A. M. de Carvalho & Barneby. Z. cowanii Mansano was described from the state of Minas Gerais, Brazil by Mansano (Mansano & Tozzi 1999a). Wied-Neuwied & Nees (1827) did not designate a type species for Zollernia. Cowan (1959) selected Z. falcata as the type species. This is currently considered to be a synonym of Z. glabra. Although there have been various studies of Zollernia (e.g., Yakovlev 1976; Carvalho & Barneby 1993; Mansano & Tozzi 1999a), the genus is in need of revision, several species need to be newly circumscribed, and a key to all recognised taxa is required. We address these points in the present study. Prior to this work herbarium specimens were often misidentified.

A Molecular Phylogeny and New Infrageneric Classification of Mucuna Adans. (Leguminosae-Papilionoideae) including Insights from Morphology and Hypotheses about Biogeography

Premise of research. The genus Mucuna has a pantropical distribution and comprises approximately 105 species, many of which show great economic value for forage, ornament, and medicine. To date, phylogenetic relationships within Mucuna have not been investigated using molecular data. The aim of this study was to build a phylogenetic framework for Mucuna to address questions about its monophyly, infrageneric relationships, divergence times, and biogeography. Methodology. We sequenced plastid (trnL-F) and nuclear ribosomal (internal transcribed spacer) regions and applied Bayesian and maximum likelihood analyses. An ancestral area reconstruction coupled with a divergence time analysis was used to investigate the historical biogeography of the genus. Pivotal results. Our results show that Mucuna is a monophyletic genus and that subgenus Stizolobium is a monophyletic group within it. We present here the analyses and results that support the need to recircumscribe subgenus Mucuna and to segregate a small group of species with large fruits into a newly proposed subgenus (to be described formally elsewhere after additional investigations). Conclusions. On the basis of ancestral area reconstruction and divergence time analyses, we conclude that the genus Mucuna originated and first diversified in the Paleotropics around 29.2 Ma and achieved a pantropical distribution through multiple long-distance dispersal events, which were facilitated by the occurrence of seeds adapted to oceanic dispersal.

Updates to the taxonomy of Swartzia (Leguminosae) in extra-Amazonian Brazil, with descriptions of five new species and a regional key to the genus

In advance of an updated monographic revision of Swartzia (Leguminosae), we discuss the taxonomy of the genus in extra-Amazonian Brazil and present descriptions of five new species. All of the new species are narrowly distributed in eastern Brazil in threatened Atlantic wet forest and coastal scrub habitats. Four of them—S. alagoensis, S. arenophila, S. revoluta, and S. submontana—belong to the diverse and taxonomically challenging section Acutifoliae, which has undergone extensive evolutionary radiation in the region. The fifth, S. thomasii, is a member of the otherwise Amazonian section Glabriplantae and is only subtly distinct from the Amazonian species S. reticulata. The new species and other recent additions to the genus are incorporated in a key to the species of Swartzia of extra-Amazonian Brazil.ResumoComo parte do estudo de atualização da monografia de Swartzia (Leguminosae), são apresentados dados sobre os táxons do Brasil extra-amazônico com a descrição de cinco espécies novas. Todas as novas espécies propostas neste trabalho apresentam distribuição restrita em habitats ameaçados na floresta atlântica brasileira e vegetação arbustiva costeira. Quatro novos táxons—S. alagoensis, S. arenophila, S. revoluta e S. submontana—pertencem à seção Acutifoliae, um grupo diverso e taxonomicamente complexo que teve alta radiação evolutiva na região. A quinta espécie, S. thomasii, pertence à seção Glabriplantae, um grupo mais comum na amazônia, e é apenas sutilmente distinta de S. reticulata, uma espécie amazônica. Devido ao fato da inclusão destes novos táxons e outras alterações recentes na taxonomia de Swartzia no Brasil extra-amazônico, é apresentada uma chave para distinguir as espécies de Swartzia desta área.

A família Leguminosae na Serra de Baturité, Ceará, uma área de Floresta Atlântica no semiárido brasileiro

The Baturite mountain range is one of the largest remnants of Atlantic forest located in the the Brazilian semiarid region. The area presents a high diversity, being considered of extreme biological importance for the conservation. The species of family Leguminosae were collected from February/2007 to February/2009 and the species list was complemented with herbarium analysis. The Leguminosae was represented by 56 genera and 132 taxa. The subfamily Papilionoideae contributed with higher richness of genus and species (31 genera and 58 taxa) followed by Mimosoideae (17 genera and 41 taxa) and Caesalpinioideae (eight genera and 33 taxa). Among the species studied in the area, 55% of them are trees and shrubs. The most diverse genera are Mimosa L. (15), Senna Mill. (12), Desmodium Desv. (seven) e Chamaecrista Moench, Bauhinia L., Inga Mill. and Aeschynomene L. (six each)

Evidence for Division of Labor and Division of Function Related to the Pollen Release in Papilionoideae (Leguminosae) with a Heteromorphic Androecium

Premise of research. Heterostemony occurs in all three traditional subfamilies of Leguminosae, but it is most commonly found in the Papilionoideae, which are characterized by papilionaceous corollas and complex pollen-releasing mechanisms. Our objective was to evaluate evolutionary and reproductive advantages of heterostemony in papilionoid legumes. Methodology. We investigated the functionality of pollen grains and floral ontogeny of two heterostemonous papilionoid species, Lupinus “The Governor” and Cytisus scoparius, using SEM, LM, and TEM techniques. Pivotal results. Lupinus showed differences in pollen viability between the two stamen morphs. The antesepalous stamens (producing fertile pollen grains) are initiated in reversed unidirectional order and develop before the antepetalous stamens (producing sterile pollen grains). During the development, the keel petals fuse at their ventral and dorsal regions, forming a tube, which surrounds the reproductive column, leaving an apical opening through which the antepetalous stamens outgrow the antesepalous stamens and push their pollen grains out of the keel. In C. scoparius, the keel petals have lateral projections that fit into depressions in the wing petals, facilitating explosive pollen release. Stamen formation and elongation occur in a unidirectional order. The two distinct sets of stamens have mixed initiation, forming abaxial stamens for pollination and adaxial stamens for feeding. Conclusions. The androecium development corresponds with differences in the pollen structure and functionality, and there is strong evidence for a division of functions in both studied species. Cytisus scoparius shows a classic case of division of labor, with some stamens producing pollen for food and others for pollination. In Lupinus, the heterostemony is related to the pollen-releasing mechanism, with the smaller stamens producing sterile pollen and in much lower proportion.