André Olmos Simões

PHYLOGENY AND SYSTEMATICS OF THE RAUVOLFIOIDEAE (APOCYNACEAE) BASED ON MOLECULAR AND MORPHOLOGICAL EVIDENCE1

Abstract To elucidate deeper relationships within Rauvolfioideae (Apocynaceae), a phylogenetic analysis was conducted using sequences from five DNA regions of the chloroplast genome (matK, rbcL, rpl16 intron, rps16 intron, and 3′ trnK intron), as well as morphology. Bayesian and parsimony analyses were performed on sequences from 50 taxa of Rauvolfioideae and 16 taxa from Apocynoideae. Neither subfamily is monophyletic, Rauvolfioideae because it is a grade and Apocynoideae because the subfamilies Periplocoideae, Secamonoideae, and Asclepiadoideae nest within it. In addition, three of the nine currently recognized tribes of Rauvolfioideae (Alstonieae, Melodineae, and Vinceae) are polyphyletic. We discuss morphological characters and identify pervasive homoplasy, particularly among fruit and seed characters previously used to delimit tribes in Rauvolfioideae, as the major source of incongruence between traditional classifications and our phylogenetic results. Based on our phylogeny, simple style-heads, syncarpous ovaries, indehiscent fruits, and winged seeds have evolved in parallel numerous times. A revised classification is offered for the subfamily, its tribes, and inclusive genera.

IS MANDEVILLA (APOCYNACEAE, MESECHITEAE) MONOPHYLETIC? EVIDENCE FROM FIVE PLASTID DNA LOCI AND MORPHOLOGY1

Abstract In order to test the monophyly of Mandevilla Lindl., the largest genus in tribe Mesechiteae (Apocynaceae, Apocynoideae), and its affinities to other genera in the tribe, maximum parsimony analysis was conducted on a data set comprising DNA sequences from five plastid loci (rpl16, rps16, and trnK introns; trnSGCU-trnGUUC intergenic spacer; and matK gene), as well as morphological data for 65 taxa of Mesechiteae (48, Mandevilla) and nine taxa from other tribes of the subfamily. Mandevilla, as circumscribed by Pichon, was found to be monophyletic, whereas Woodsons circumscription proved to be polyphyletic. Thus defined, Mandevilla forms a strongly supported clade that can be divided into six clades of species groups. Most of the infrageneric taxa of Mandevilla proposed by Woodson and Pichon are polyphyletic. Many of the diagnostic characters previously used to define taxonomic groups are shown to have arisen multiple times, rendering them unsuitable for classificatory purposes. The similar growth form and tubular flowers of Macrosiphonia Müll. Arg. and Telosiphonia (Woodson) Henr., two geographically disjunct segregates, represent the most extreme case of parallel evolution within Mandevilla, with their striking similarities most likely correlated to colonization of open, dry habitats and pollination by hawkmoths.

Tribal and intergeneric relationships of Mesechiteae (Apocynoideae, Apocynaceae): evidence from three noncoding plastid DNA regions and morphology

The Neotropical tribe Mesechiteae (Apocynaceae) is currently considered to include nine genera: Allomarkgrafia, Galactophora, Macrosiphonia, Mandevilla, Mesechites, Quiotania, Secondatia, Telosiphonia, and Tintinnabularia. Tribal and intergeneric relationships, however, are in dispute. To test the monophyly of the tribe and evaluate intratribal relationships, a maximum parsimony analysis was conducted based on DNA sequences from the plastid rpl16 intron, rps16 intron, and trnS-G intergenic spacer region as well as morphological data for 23 taxa of Mesechiteae and 11 taxa from other tribes of Apocynoideae. Mesechiteae, as currently circumscribed, was found to be polyphyletic. Only removal of Secondatia and Galactophora and inclusion of Forsteronia rendered the tribe monophyletic. Thus defined, Mesechiteae forms a strongly supported clade including seven genera in three subclades: the Mesechites subclade (comprising Tintinnabularia, Allomarkgrafia, and Mesechites), the Forsteronia subclade (containing only Forsteronia) and the Mandevilla subclade (comprising Macrosiphonia, Mandevilla, and Telosiphonia). Allomarkgrafia is nested in Mesechites. Macrosiphonia and Telosiphonia form two distinct monophyletic clades. Both, however, are nested in Mandevilla. Results suggest upholding the following genera in Mesechiteae: Allomarkgrafia, Forsteronia, Mandevilla, Mesechites, and Tintinnabularia. The status of Quiotania could not be evaluated.

New Combinations in Mandevilla Lindley (Apocynaceae)

ABSTRACT Historically, taxonomic delimitation of Mandevilla Lindley, a large Neotropical genus of Apocynaceae, has been controversial. Two competing classifications were proposed by Woodson (1933) and Pichon (1948), the former now being widely accepted by taxonomists in the family. Recent phylogenetic studies, however, support the monophyly of Mandevilla sensu Pichon rather than sensu Woodson, the latter being paraphyletic with respect to Quiotania Zarucchi, Macrosiphonia Müller Argoviensis, and Telosiphonia (Woodson) Henrickson. Based on the principle of monophyly, we here re-establish Pichons classification and include Quiotania, Macrosiphonia, and Telosiphonia as synonyms of Mandevilla. Three new combinations are made here: Mandevilla hesperia (I. M. Johnston) A. O. Simões, L. S. Kinoshita-Gouvêa & M. E. Endress, M. nacalpulensis (Felger & Henrickson) A. O. Simões, L. S. Kinoshita-Gouvêa & M. E. Endress, and M. undulata (C. Ezcurra) A. O. Simões, L. S. Kinoshita-Gouvêa & M. E. Endress; Q. colombiana Zarucchi is newly synonymized with M. ligustriflora Woodson; and a lectotype is designated for M. martii (Müller Argoviensis) Pichon.

Eight New Species of Mandevilla Lindley (Apocynaceae, Apocynoideae) from Brazil

ABSTRACT Eight new species of Mandevilla Lindley are described and illustrated. The new species, six of them rare and endemic, occur in eastern Brazil. Three of the new species (M. fistulosa, M. grazielae, and M. harleyi) are shrubs growing on the upper slopes of granitic inselbergs in Minas Gerais and Espírito Santo in southeastern Brazil. Two species, M. semirii and M. rubra, are woody lianas growing on rock outcrops of quartzite in campo rupestre vegetation of Minas Gerais. Mandevilla hatschbachii, a distinctive shrub known only from three collections, occurs in the campo rupestre formations in eastern Brazil in Bahia. Mandevilla guanabarica, a suffrutescent liana generally misidentified as M. funiformis (Vellozo) K. Schumman, is found at the borders of the Atlantic rainforest in Espírito Santo and Rio de Janeiro. Mandevilla dardanoi is a small shrub endemic to Maciço da Borborema, in northeastern Brazil.

Systematics and character evolution of Vinceae (Apocynaceae)

Vinceae comprises 9 genera and about 153 species. The genera differ greatly in a number of morphological traits, especially those relating to fruit and seeds, which resulted in their placement in different tribes and/or subtribes by earlier taxonomists. Here, 378 new chloroplast DNA sequences from 98 species from 5 tribes of the rauvolfioid grade were analyzed phylogenetically to test the monophyly of Vinceae, its 6 subtribes, and constituent genera, as well as the sectional treatments of Rauvolfia, and to find morphological synapomorphies for major clades. Vinceae, as currently circumscribed, was found to be monophyletic as were four of the six subtribes. Kopsia and Ochrosia were confirmed as successive sister groups to the rest. All nine genera currently recognized in the tribe were resolved as monophyletic, Laxoplumeria and Tonduzia for the first time. The broad circumscription of Petchia and Ochrosia proposed by previous authors was supported. The pantropical genus Rauvolfia is divided into two major lineages: one comprised entirely of paleotropical species, and the other the neotropical species, a recurring geographic pattern in the family. Surprisingly, the Hawaiian species of Vinceae have probably arisen by two different trans-oceanic routes, either by long-distance dispersal from Australasian (Ochrosia spp.) or Caribbean (Rauvolfia sandwicensis) ancestors. Within Rauvolfia, most of the sections, series and subseries recognized in previous classifications are paraphyletic. Only 2 of the 14 sections of Rauvolfia proposed by Pichon and 1 of the 11 sections proposed by Rao that are not monospecific were found to be monophyletic. Mapping of 30 morphological characters onto the molecular tree identified an unlobed upper wreath on the style head as an unambigous synapomorphy characterizing the tribe as well as synapomorphies for each of its genera. The two earliest-branching genera, Kopsia and Ochrosia, have dextrorse corolla lobe aestivation, in contrast to the sinistrorse corollas of the other Vinceae and the great majority of genera throughout the rauvolfioid grade. Drupes have arisen in parallel at least three times within Vinceae. The adaptations of the endocarp for water dispersal are possibly ancestral in the tribe, and the observed morphological differences between the drupaceous fruits of Kopsia/Ochrosia versus Petchia and Rauvolfia may be due to their abiotic versus biotic dispersal mode, respectively.

Stemodia cipoensis (Plantaginaceae): A New Species from Serra Do Cipó, Minas Gerais, Brazil

Abstract The Serra do Cipó, part of the central portion of the Espinhaço Range in Minas Gerais, Southeast Brazil, is known to have been extensively investigated over the past three decades, especially because of the “Flora da Serra do Cipó” project. Nevertheless, some areas have been poorly collected as the efforts were largely concentrated near the roads. Various new species, in several families, have been recently discovered in the campos rupestres of the Serra do Cipó. Here we report the discovery of a new species of Stemodia (Plantaginaceae) from the northwestern slopes. We describe and illustrate the new species based on morphological and micro-morphological evidence, provide information on its taxonomy, habitat, geographical distribution, and conservation status, along with a key to distinguish it from the related species. Stemodia cipoensis is similar to S. harleyi, S. lobata, and S. stellata, being characterized by the presence of long, stiff, curved, eglandular trichomes. It has been assessed as CE (critically endangered) under criteria B1ab(iii) and B2ab(iii), following IUCN guidelines.

Phylogeny and biogeography of the genus Zornia (Leguminosae: Papilionoideae: Dalbergieae)

The genus Zornia has a pantropical distribution and comprises 80 species in two subgenera (Myriadena, Zornia) and three sections (Zornia, Isophylla, Anisophylla). The genus is characterized mainly by flowers arranged in a spiciform inflorescence, with paired peltate bracteoles protecting each flower, and stipules that resemble the bracteoles. Phylogenetically Zornia is a member of the informal Adesmia clade of the Dalbergioid group, being most closely related to the predominantly South American genera Poiretia and Amicia. Phylogenetic analyses of DNA sequence data from the internal transcribed spacer region of nuclear ribosomal DNA and the plastid trnL-trnF intron and spacer, including multiple accessions of species of all subgenera and sections, support the monophyly of Zornia and indicate that Poiretia is sister to Zornia. All traditional subgenera and sections of Zornia are resolved as paraor polyphyletic in our study. Nevertheless, two major clades can be recognized. Time-calibrated phylogenetic analyses suggest that most Zornia species diverged 8–10 Ma and reveal that only one migratory event occurred within the genus ca. 5 Ma from America to the Old World probably via long-distance dispersal.

Gynostegium Morphology of Mesechiteae Miers (Apocynaceae, Apocynoideae) as It Pertains to the Classification of the Tribe

In order to investigate morphological patterns in Mesechiteae and test the new circumscription of the tribe, the gynostegium structure of nine species belonging to Forsteronia, Mandevilla, and Mesechites, with two species from Secondatia as outgroup, were comparatively studied. Our results support the most recent circumscription of Mesechiteae, including Forsteronia, Mandevilla, and Mesechites and excluding Secondatia. The gynostegia of Forsteronia, Mandevilla, and Mesechites have the same basic organization: a proliferation of parenchyma cells in the style head forms five projecting ribs that are adnate to an expanded portion of the connective below the thecae, here named the “staminal retinacle.” This adnation is very tenuous in Forsteronia (few cells of the staminal retinacle are effectively connected to the style head) but much stronger in Mandevilla and Mesechites (several cells of the staminal retinacle are connected to the style head). The observed differences in the size of the style head ribs and the degree of adnation to the stamens also support the three major clades recognized within the tribe. The gynostegium of Secondatia has a different organization, of the type reported for representatives of the tribe Apocyneae: the style head has no projecting ribs, and its epidermal cells are interdigitated with, but not adnate to, unicellular trichomes of the staminal retinacle.

Microsatellite Markers for Studies with the Carnivorous Plant Philcoxia minensis (Plantaginaceae)

Premise of the study: Microsatellite markers were developed for the critically endangered carnivorous species Philcoxia minensis (Plantaginaceae) for further population genetic studies aiming at its conservation. Methods and Results: We identified 29 clones containing 40 microsatellites from a genomic enriched library. A total of 27 primer pairs were developed and evaluated in 30 individuals of a natural P. minensis population. Seventeen markers successfully presented amplification products within the expected size range, of which 12 were polymorphic. The expected and observed heterozygosities ranged from 0.03 to 0.65 and from 0.00 to 0.77, respectively. Positive transferability with the related species P. bahiensis was observed for the same 17 markers. Conclusions: The 12 polymorphic microsatellite markers are suitable for studies in genetic diversity and structure, mating system, and gene flow in P. minensis and also may be useful for similar issues regarding the related species P. bahiensis.