Phytotaxa 263 (3): 219–232 ISSN 1179-3155 (print edition)

http://www.mapress.com/j/pt/
Article PHYTOTAXA
Copyright © 2016 Magnolia Press ISSN 1179-3163 (online edition)

http://dx.doi.org/10.11646/phytotaxa.263.3.3

Phylogenetic placement and a new circumscription of Poteranthera (Microlicieae;

Melastomataceae)
MARIA JOSÉ REIS DA ROCHA¹*, PAULO JOSÉ FERNANDES GUIMARÃES2, FABIÁN A. MICHELANGELI3
& ROSANA ROMERO4
1
Universidade Federal de Minas Gerais, Departamento de Botânica, Av. Antônio Carlos 6627, Pampulha, 31270–901, Belo Horizonte,
Minas Gerais, Brazil
2
Instituto de Pesquisas Jardim Botânico de Rio de Janeiro, Rua Pacheco Leão 915, 22460–030, Rio de Janeiro, Rio de Janeiro, Brazil
3
The New York Botanical Garden, Bronx, NY 10458, USA
4
Universidade Federal de Uberlândia, Instituto de Biologia, Rua Ceará s.n., 38405–315, Uberlândia, Minas Gerais, Brazil
*Corresponding author: mariajosereisdarocha@gmail.com

Abstract

Poteranthera (Melastomataceae) is a genus of small, annual herbs with narrow leaves, that inhabit humid savannas in South
America and are visible only when in bloom. Due to their size and paucity of collections, the phylogenetic position of Po-
teranthera has been uncertain to this point. Poteranthera has been at times associated, confused, or placed in Acisanthera
and/or Siphanthera. Phylogenetic analyses based on DNA sequence data place Poteranthera unequivocally within the Mi-
crolicieae and not closely related to Acisanthera or Siphantera. These results are consistent with seed and anther morphol-
ogy. Putative morphological synapomorphies for this genus are the glandular hairs restricted to the leaf margins and the
apically constricted hypanthium. Based on morphology, Acisanthera leptalea seems better placed in Poteranthera and the
new combination Poteranthera leptalea is made. An expanded generic description is also provided.

Key words: Morphology; phylogenetic analysis; taxonomy

Resumo

Poteranthera (Melastomataceae) é um gênero de ervas anuais pequenas, com folhas estreitas, que habitam savanas úmidas
da América do Sul e cujos indivíduos são evidentes apenas quando em flor. Devido ao pequeno porte dos indivíduos e es-
cassez de boas coleções com detalhes morfológicos claros, o posicionamento supragenérico de Poteranthera permanecia
incerto. Poteranthera tem sido associada, confundida ou posicionada em Acisanthera e/ou Siphanthera. As análises filo-
genéticas baseadas em dados de sequência de DNA sugerem o posicionamento de Poteranthera em Microlicieae ao invés
de Acisanthera e/ou Siphanthera. Estes resultados são corroborados pela morfologia da semente e da antera. Prováveis
sinapomorfias para este gênero são os tricomas glandulares restritos às margens das folhas e a constrição apical do hipanto.
Baseado na morfologia, Acisanthera leptalea é melhor posicionada em Poteranthera e a nova combinação Poteranthera
leptalea é aqui proposta. Uma descrição genérica expandida é também fornecida.

Palavras chave: Morfologia, análise filogenética; taxonomia

Introduction

The German botanist August Gustav Bongard described in one article two of the rarest neotropical genera of
Melastomataceae: Lithobium Bongard (1838: 140) and Poteranthera Bongard (1838: 131). Samples of these plants
were sent from central Brazil to Russia by his German colleague Ludwig Riedel, who was part of the expedition of
Georg Heinrich von Langsdorff (Manizer 1967). Poteranthera was included by Naudin (1849) in the tribe Lasiandrales
Naudin (1849: 273) (=Melastomeae), and then transferred to the Microliciales Naudin (1849: 203) by Triana (1871),
an opinion maintained by Cogniaux (1885, 1891). Renner, in her classification of the Melastomataceae, placed
Poteranthera in the Melastomeae Renner (1993: 523) as earlier suggested by Wurdack (as Tibouchinae;1963).

Accepted by Renato Goldenberg: 21 May 2016; published: 3 Jun. 2016 219

 Cogniaux’s (1891) generic concept of Poteranthera was broader than the one used today, as it included several
species now in Acisanthera P. Browne (1756: 217) and Siphanthera Pohl ex de Candolle (1828: 121). Based on anther
and seed morphology, Cogniaux divided the genus in three sections. Wurdack later (1963) excluded from Poteranthera
the species of Poteranthera section Tulasnea Naudin (1844: 142; 1849: 276), which he transferred to Siphanthera
Pohl ex de Candolle (1828: 121), and the species of section Onoctonia Naudin (1849: 276), which he transferred or
considered as part of Acisanthera. Therefore, Wurdack (1963) considered Poteranthera as a monotypic genus closely
related to Acisanthera (Wurdack, 1973). Wurdack (1974) then added a second species at the genus based on collections
from Mato Grosso, Brasil.
The latest treatment of Poteranthera was a synopsis by Kriebel (2012), which also included a description of a
new species. Currently, Poteranthera consists of three herbaceous species that always grow in periodically inundated
to humid savannas with a disjunct distribution in Venezuela, Brazil and Bolivia (Fig.1).

FIGURE 1. Poteranthera pusilla. A. Habit. B. Branches showing solitary flower. C. Details of the flower and reddish leaves. D. Flower. E.
Fruit showing the hypanthium constriction (arrow) and sessile leaves, margin beset with stout glandular trichomes. (Photos: A. R.Romero;
B–C. M.J.Rocha; D–E. W. Fernandes).

220 • Phytotaxa 263 (3) © 2016 Magnolia Press ROCHA ET AL.

 In summary, Poteranthera has been suggested to be related to species that are currently in Melastomeae, the
Marcetia alliance clade (see Michelangeli et al. 2013), and Microlicieae. However, while phylogenetic analyses
have been published for all three groups (respectively Michelangeli et al. 2013; Rocha et al. in press; Fritsch et al.
2004), Poteranthera has not been included in any of these studies, and thus, its phylogenetic placement within the
Melastomataceae remains uncertain.
In order to assess the phylogenetic position of Poteranthera whithin the Melastomataceae, samples of Poteranthera
pusilla Bongard (1838: 138) were collected in a humid savanna in Patrocínio municipality, Minas Gerais state, Brazil,
and using DNA sequence data analyzed within a broad sampling of capsular neotropical Melastomataceae.

Materials and Methods

In order to clarify the phylogenetic position of Poteranthera, we sampled 66 taxa from 27 genera belonging to the
Marcetia alliance, Melastomeae s.str., Microlicieae and Rhexieae de Candolle (1828: 114), all potential relatives of
Poteranthera. Acisanthera and Siphanthera were extensively sampled since these genera have been traditionally
associated with Poteranthera. A species of Cambessedesia (DC.) was included as an outgroup since this genus has
been shown to be outside of the clade formed by the Melastomeae, Rhexieae and the Marcetia alliance (Goldenberg
et al. 2012). Genomic DNA was extracted from silica-dried leaves collected in the field using the DNeasy plant mini
kit (Qiagen, Valencia, CA), with the modifications suggested by Alexander et al. (2007) and Martin et al. (2008).
Phylogenetic analyses were based on five molecular markers: two nuclear (nrITS, nrETS) and three plastid intergenic
spacers (accD-psaI, atpF-atpH, trnS-trnG). The nrITS region consisted of a small portion of the 3’ end of the 18S
ribosomal gene, the ribosomal internal transcribed spacers (ITS1 and ITS2), the intervening 5.8S ribosomal gene and
a section of about 241 bp on the 5’ end of the 26S ribosomal gene. The external transcribed spacer (nrETS) consisted
of a small portion of the 5’ end of the 18S ribosomal gene and ca 610 bp in the nrETS region. DNA amplification was
performed by polymerase chain reaction (PCR) in a final volume of 15 μL with the following reaction components:
0.5–0.7 μL of genomic DNA (ca. 30 ng μL-1), 7.5 μL 2x EconoTaq Plus Green (Lucigen Technologies Corporation,
Middleton, WI), 2 μL of each primer (3 μM), 0.75 μL of spermidine (4 mM), 2.5 μL of purified water. The nrITS
was amplified using primers designed by Stanford et al. (2000), Michelangeli et al. (2004), White et al. (1990) and
for nrETS designed by Kriebel et al. (2015). The plastid intergenic spacers (accD-psaI, atpF-atpH, trnS-trnG) were
amplified using primers designed by Shaw et al. (2005), Reginato et al. (2010) and Hamilton (1999), respectively.
Amplifications of nuclear and plastid markers were carried out with the same primers and conditions detailed in Rocha
et al. (in press). Cycle sequencing reactions were carried out with the same amplification primers using the sequencing
service at the University of Washington, USA (High Throughput Genomics Center-htSEQ). All data sets were analyzed
using maximum parsimony (MP) and Bayesian inference (BI). Parsimony analyses were performed in PAUP version 4
(Swofford 2002) using Fitch parsimony as the optimality criterion (Fitch 1971). Heuristic searches consisted of 10,000
replicates of random taxa addition, using the TBR algorithm (tree bisection-reconnection) and saving up to 10 MP trees
for each replication. Bayesian analyses were performed using the program MrBayes 3.1.2 run through the Cypress
Science Gateway (Miller et al. 2010). Analyses were performed with mixed models and independent parameters. The
analysis consisted of two independent runs, each with four Markov chains for 10 million generations, sampling one
tree every 1,000 generations. To achieve convergence the temperature parameter for heating the chains was lowered
to 0.05. Convergence between the chains was assessed by the average standard deviation of split frequencies (<0.01),
as well as the stationarity of the chains with the generated graphical outputs. Convergence was achieved after 245000
generations and the initial 25% of the resulting trees were discarded as burn-in. The remaining trees were used to
calculate the posterior probability (PP) in a majority-rule consensus. Phylogenetic trees were visualized and edited in
Figtree 1.4.2 and Coreldraw X6. GenBank accession numbers are listed in Appendix 1.
Seeds were imaged under scanning electron microscopy (SEM); these were selected from mature fruits from
herbarium specimens, mounted on double-sided tape on aluminium stubs, sputter coated with gold palladium for 4
minutes in a Hummer 6.2 sputter coater (Anatech, Springfield, VA) and visualized on JEOL-JSM 5410LV SEM and
imaged with JEOL ORION 5410, version 1.72.01.

Poteranthera Phytotaxa 263 (3) © 2016 Magnolia Press • 221

FIGURE 2. Bayesian analysis of the combined nuclear (nrETS, nrITS) and plastid (accD-psaI, trnS-trnG, atpF-atpH) markers. Numbers
above the nodes are posterior probabilities.

222 • Phytotaxa 263 (3) © 2016 Magnolia Press ROCHA ET AL.

Results

Bayesian Inferecence (BI) and Maximum Parsimony analyses (MP; not shown) for individual loci and combined
datasets did not shown significant conflicts between well-supported clades (PP≥ 95%, BP ≥ 85%). Combined analyses
under both BI and MP unequivocally place Poteranthera within tribe Microlicieae with high support (PP=1.00;
BP=100%). Within Microlicieae (PP=1.00; BS=100%), Rhynchanthera de Candolle (1828: 106) was recovered as
the first divergent branch (PP=1.00; BS= 100%) followed by Lavoisiera bicolor Naudin (1849: 222) and L. imbricata
de Candolle (1828: 103) (Fig. 2). Poteranthera is resolved as sister to a clade with Chaetostoma de Candolle (1828:
112), Lavoisiera de Candolle (1828: 102), Microlicia Don (1823: 301), Stenodon Naudin (1849: 215) and Trembleya
de Candolle (1828: 125). None of the species of Acisanthera or Siphanthera sampled were resolved with Poteranthera
or elsewhere within the Microlicieae, but rather were all recovered in the Marcetia alliance clade, as previously shown
(Michelangeli et al. 2013, Rocha et al. in press).

Discussion

A careful evaluation of seed and anther characters of Poteranthera shows that its placement within Microlicieae is
consistent with molecular data. Naudin (1849), and later Triana (1866, 1871), characterized the Microlicieae based
on seed shape and testa sculpturing and these criteria have been followed in later studies, albeit with some minor
modifications (Renner 1993, Almeda & Martins 2001, Fritsch et al. 2004). Fritsch et al. (2004) describe the seeds of
Microlicieae as reniform, ellipsoid or elongate with a foveolate or lacunate-reticulate testa. Poteranthera clearly show
these same characteristics, as already mentioned by Bongard (1838) and Wurdack (1963) (Fig. 3). Anthers in all genera
currently in the Microlicieae (see Fritsch et al. 2004) have well developed pedoconnectives, with rostrate thecae with
a ventrally oriented pore. Anthers of Poteranthera share this morphology (Fig. 4), even if the pedoconectives are
proportionally shorter. It should be noted that the anthers in P. pusilla are rostrate before they open, but once the pore
dehisces is as broad as the thecae.
Much of the confusion in correctly placing Poteranthera has been due to its similarities to some species of
Acisanthera. Within Acisanthera the seeds were described as Microlicioid by Whiffin & Tomb (1972) due to their
elongate shape and foveolate surface. However, there is a great amount of variation in seed shape within Acisanthera
that spans from clearly cochleate , as in A. limnobios Schrank & Mart. ex de Candolle (1828: 117) Triana (1871: 33) to
straight, as in A. crassipes (Naudin 1849: 277) Wurdack (1963: 539) (Whiffin & Tomb 1972), with several intermediate
states with only slightly curved to obviously cochleate seeds in other species (Fig. 2). Due to this variability, Wurdack
(1963) agreed with Triana (1871) in that the reniform foveolate seeds of Poteranthera suggested a close relationship
with Acisanthera and that it should be placed in the Tibouchineae (= Melastomeae). Kriebel (2008) also pointed
out the simimilarities in the seed shape and testa morphology between Acisanthera and Poteranthera. However, our
phylogenetic results show that Acisanthera is not related to Poteranthera and thus, those cases in Acisanthera with
straight seeds should be interpreted as convergence or parallelism.
The anthers in some species of Acisanthera also have long pedoconnectives with or without rostrate apices
(Kriebel & Almeda 2013). However, in all these the pedoconnective is never as long as the thecae, and the thecae are
also proportionally narrower and more elongated than in the Microlicieae. The pedoconectives in P. pusilla and P.
annectans are also not as long as the thecae, but the thecae have ovate and usually expanded towards the bottom as in
most Microlicieae, and not subulate as in Acisanthera.
The seeds of Siphanthera are rather different from those of Poteranthera (or Acisanthera) with a lacrimiform
shape and areolate testa (Whiffin & Tomb 1972, Almeda & Robinson 2011), two characters that are considered
synapomorphies for this genus (Almeda & Robinson 2011). The anthers of Siphanthera are indeed rostrate, but the
pedoconnective is very short, the apical rostrum can be proportionally much longer, and the connective appendages
rather than ventral are usually lateral.
Kriebel (2012) mentioned that the most significant characters and possible synapomorphies of Poteranthera are
the conspicuous glandular hairs restricted to the leaf margins, and the constricted hypanthium. Among the species of
Acisanthera, A. leptalea Almeda (1999: 159) clearly stands out as having these same diagnostic characters. Moreover,
as already pointed out by Kriebel (2008, 2012), the seed and anthers of A. leptalea are like those of Poteranthera, In the
description of Acisanthera leptalea Almeda (1999) assigned it to Acisanthera sect. Dicrananthera Triana (1871: 33).
However, this placement did not seem to be unequivocal, as the author commented that A. leptalea differs markedly

Poteranthera Phytotaxa 263 (3) © 2016 Magnolia Press • 223

from Acisanthera hedyotidea (Presl 1832: 70) Triana (1871: 33) and A. tetramera (Naudin 1849: 280) Triana (1871:
33) based on vegetative and floral characters. Most notably, although many species of Acisanthera have a developed
pedoconnective, in none of them the pedoconnective is longer than the thecae as in A. leptalea, P. annectans, or any
other Microliceae. It should be noted that A. leptalea has 4-merous flowers and 2-locular ovaries, characters not
known in Poteranthera up until now, presumably leading to its placement in Acisanthera. However, in Microlicieae
the number of ovary locules is variable within its genera, and this character is often not completely stable within some
species of Chaetostoma (Koschnitzke 1997), Lavoisiera (Romero 2000), Microlicia (Romero 2000), Rhynchanthera
(Renner 1990) and Trembleya (Martins 1997). Given anther, leaf and hypanthium morphology, we conclude that A.
leptalea is better placed within Poteranthera and the appropriate transfer is made below. With the combination of A.
leptalea in Poteranthera the description of the genus needs to be amended to reflect these changes.

FIGURE 3. Scanning Electron Microscopy of seeds (lateral view). A. Siphanthera paludosa. B. Acisanthera quadrata. C. Acisanthera
crassipes. D. Rhynchanthera grandiflora. E. Poteranthera pusilla. F. Microlicia chrysantha. (A. Rocha 621; B. Michelangeli 826; C.
Rocha 833; D. Elcoso 623; E. Rocha 940; F. Ganev 2181).

224 • Phytotaxa 263 (3) © 2016 Magnolia Press ROCHA ET AL.

FIGURE 4. Stamens of Poteranthera and putative relatives. A–B. Acisanthera alsinaefolia. A. Antepetalous (larger) stamen. B.
Antesepalous (smaller) stamen. C. Poteranthera annectans, antesepalous stamen. D. Microlicia fasciculata, antepetalous (larger) stamen.
E. Rhynchanthera dichotoma, lower antesepalous (single largest) stamen. (A–B. Souza 2355 (RB); B. Sick 478 (RB); C. Almeda 5387
(RB); D. Baitello 2415(RB)).

Poteranthera Phytotaxa 263 (3) © 2016 Magnolia Press • 225

 Based on the molecular phylogeny produced here and corroborated by morphological data, we conclude that
Poteranthera should be placed in the Microlicieae. With this new circumscription, Microlicieae has now seven genera:
Chaetostoma, Lavoisiera, Microlicia, Poteranthera, Rhynchanthera, Stenodon and Trembleya.

Taxonomy

Poteranthera Bong. (1838: 131)

Small herbs; leaves sessile, 1-nerved, mostly linear, and glabrous with the margin entire or serrulate beset with stout
glandular trichomes; flowers 4–5-merous with a campanulate to globose hypanthium somewhat constricted at the
apex, haplostemonous or diplostemonous; antesepalous stamens with a prolonged connective, shorter or longer than
the thecae, and modified ventrally into a bilobed yellow appendage at the junction with the filament, the thecae ovate
and with a rostrate apex, with a cucullate or broad pore; antepetalous stamens absent, staminodial, or if fertile either
just smaller than the larger anthers or with the connective less developed than in the larger stamens; ovary 2–3-locular,
glabrous; fruits capsular; seeds reniform to subcochleate, the testa foveolate .

Poteranthera leptalea (Almeda) M.J.Rocha, P.J.Guim. & R.Romero, comb. nov.

Basionym:—Acisanthera leptalea Almeda (1999: 159–162). Type:—BOLIVIA. Santa Cruz: Velasco, Noel Kempff Mercado National
Park, Serranía de Huanchaca, north and northeast of Noel Kempff Mercado trail, 13°54’22”S, 60°048’52”W, 360 m, 8 June 1994
(fl., fr.), B. Mostacedo et al. 2127 (holotype: MO 038864-image!; isotypes: CAS 0000053-image!, USZ).

Acknowledgments

The first author thanks Wesley Fernandes for help her on the field expedition and by the images, the scholarships
provided by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior—CAPES, Conselho Nacional de
Desenvolvimento Científico e Tecnológico—CNPq and Fundação de Amparo a Pesquisa do Estado de Minas Gerais—
FAPEMIG. This research was partially supported by NSF grants to Fabian A. Michelangeli (DEB-1140409 and DEB-
1343612), FAPEMIG (PPM-00226-12) and CAPES (PNADB/UFMG-922/2010). We thank Matthew Sewell, Marcelo
Reginato and Ricardo Kriebel for their assistance in the laboratory of the Cullman Program for Molecular Systematics
at NYBG. We are grateful to anonymous reviewers and Frank Almeda to provide valuable suggestions.

Literature Cited

Alexander, P.J., Rajanikanth, G., Bacon, C.D. & Bailey, C.D. (2007) Recovery of plant DNA using a reciprocating saw and silica–based
columns. Molecular Ecology Notes 7: 5–9.
http://dx.doi.org/10.1111/j.1471–8286.2006.01549
Almeda, F. & Martins, A.B. (2001) New combinations and new names in some Brazilian Microlicieae (Melastomataceae), with notes on
the delimitation of Lavoisiera, Microlicia, and Trembleya. Novon 11: 1–7.
http://dx.doi.org/10.2307/3393198
Almeda, F. (1999) A New Acisanthera (Melastomataceae: Melastomeae) from Noel Kempff Mercado National Park, Bolivia. Brittonia
51: 159–162.
http://dx.doi.org/10.2307/2666623
Almeda, F. & Robinson, O.R. (2011) Systematics and Phylogeny of Siphanthera (Melastomataceae). Systematic Botany Monographs 93:
1–101.
http://dx.doi.org/10.1600/036364415X688862
Bongard, H.G. (1838) Genera duo Melastomacearum ordine nova. Mémoires de l’académie Impériale des Sciences de Saint–Pétersbourg,
Sér. 6, Sciences Naturelles 4 (2): 137−142.
Browne, P. (1756) The civil and natural history of Jamaica in three parts. London.
Candolle, A.P. de (1828) Melastomaceae. In: Prodromus systematis naturalis regni vegetabilis 3. Treuttel & Würtz, Paris, pp. 99–202.

226 • Phytotaxa 263 (3) © 2016 Magnolia Press ROCHA ET AL.

Cogniaux, A.C. (1885) Poteranthera. In: Martius, C.F.P., Eichler, A.W. & Urban, I. (Eds.) Flora Brasiliensis 14 (3). Fleischer, Leipzig,
pp. 201–204.
Cogniaux, A. (1891) Mélastomacées. In: de Candolle, A. & de Candolle, C. (Eds.) Monographieae Phanerogamarum Vol. 7. Masson,
Paris, pp. 1–1256.
Don, D. (1823) An illustration of the natural family of plants called Melastomataceae. Memoirs of the Wernerian Natural History Society
4: 276–329.
Fitch, W.M. (1971) Toward defining the course of evolution: minimum change for a specific tree topology. Systematic Zoology 20:
406–416.
http://dx.doi.org/10.2307/2412116
Fritsch, P.W., Almeda, F., Renner, S.S., Martins, A.B. & Cuz, B.C (2004) Phylogeny and circumscription of the near–endemic Brazilian
tribe Microlicieae (Melastomataceae). American Journal of Botany 91: 1105–1114.
http://dx.doi.org/10.3732/ajb.91.7.1105
Goldenberg, R., de Fraga, C.N., Fontana, A.P., Nicolas, A.N. & Michelangeli, F.A. (2012) Taxonomy and Phylogeny of Merianthera
(Melastomataceae). Taxon 61: 1040–1056.
Hamilton, M.B. (1999) Four primer pairs for the amplification of chloroplast intergenic regions with intraspecific variation. Molecular
Ecology 8: 513–525.
http://dx.doi.org/10.1046/j.1365–294X.1999.00510.x
Kriebel, R. (2008) Systematics and biogeography of the neotropical genus Acisanthera (Melastomataceae). Master of Science in Biology:
Ecology and Systematic Biology. San Francisco, California, pp. 1–125.
Kriebel, R. (2012) A synopsis of the genus Poteranthera (Melastomeae: Melastomataceae) with the description of a new, apparently
pollinator deceiving species. Brittonia 64: 6–14.
http://dx.doi.org/ 10.1007/s12228–011–9192–2
Kriebel, R. & Almeda, F. (2013) Clinal variation and the decoupling of vegetative and reproductive characters in Acisanthera section
Acisanthera (Melastomataceae). Harvard Papers in Botany 18: 157–172.
http://dx.doi.org/10.3100/025.018.0211
Kriebel, R., Michelangeli, F.A. & Kelly, L.M. (2015) Discovery of unusual anatomical and continuous characters in the evolutionary
history of Conostegia (Miconieae: Melastomataceae). Molecular Phylogenetics and Evolution 82: 289–313.
http://dx.doi.org/10.1016/j.ympev.2014.09.021
Manizer, G.G. (1967) A Expedição do acadêmico G. I. Langsdorff ao Brasil: 1821-1828. São Paulo: Cia. Ed. Nacional, 244 pp.
Martin, C.V., Little, D.P., Goldenberg, R. & Michelangeli, F. (2008) A phylogenetic evaluation of Leandra (Miconieae, Melastomataceae):
a polyphyletic genus where the seeds tell the story, not the petals. Cladistics 24: 315–327.
http://dx.doi.org/10.1111/j.1096-0031.2007.00185.x
Michelangeli, F.A., Penneys, D.S., Giza, J., Soltis, D., Hils, M.H. & Skean Jr., J.D. (2004) A preliminary phylogeny of the tribe Miconieae
(Melastomataceae) based on nrITS sequence data and its implications on inflorescence position. Taxon 53: 279–290.
http://dx.doi.org/10.2307/4135608
Michelangeli, F.A., Guimarães, P.J.F., Penneys, D.S., Almeda, F. & Kriebel, R. (2013) Phylogenetic relationships and distribution of New
World Melastomeae (Melastomataceae). Botanical Journal of the Linnean Society 171: 38–60.
http://dx.doi.org/10.1111/j.1095–8339.2012.01295.x
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In:
Proceedings of the Gateway Computing Environments Workshop (GCE). New Orleans, LA, pp. 1–8.
http://dx.doi.org/10.1109/GCE.2010.5676129
Naudin, C.V. (1844) Additions á la flore du Brésil méridional. Annales des Sciences Naturalles, Botanique, sér. 3 2: 140–156.
Naudin, C.V. (1849) Melastomacearum quae in Museuo Parisiensi continentur monographicae et secundum affinitates distributionis
tentamen. Annales des Sciences Naturalles, Botanique, sér. 3 12: 196–284.
Presl, C.B. (1832) Descriptiones et icones plantarum novarum aut minus cognitarum Symbolae botanicae 1: 75.
Reginato, M., Michelangeli, F.A. & Goldenberg, R. (2010) Phylogeny of Pleiochiton (Melastomataceae, Miconieae): total evidence.
Botanical Journal of the Linnean Society 162: 423–434.
http://dx.doi.org/10.1111/j.1095–8339.2009.01022.x
Renner, S.S. (1993) Phylogeny and classification of the Melastomataceae and Memecylaceae. Nordic Journal of Botany 13: 519–540.
http://dx.doi.org/10.1111 / j.1756–1051.1993.tb00096.x
Rocha, M.J.R., Batista, J.A.N., Guimarães, P.J.F. & Michelangeli, F.A. (In press) Phylogenetic relationships in the Marcetia alliance
(Melastomeae, Melastomataceae) and implications for generic circumscription. Botanical Journal of the Linnean Society.
Shaw, J., Lickey, E.B., Beck, J.T., Farmer, S.B., Liu, W., Miller, J., Siripun, K.C., Winder, C.T., Schilling, E.E. & Small, R.L. (2005) The
tortoise and the hare II: relative utility of 21 noncoding chloroplast DNA sequences for phylogenetic analysis. American Journal of

Poteranthera Phytotaxa 263 (3) © 2016 Magnolia Press • 227

 Botany 92: 142–166.
http://dx.doi.org/10.3732/ajb.92.1.142
Stanford, A.M., Harden, R. & Parks, C.R. (2000) Phylogeny and biogeography of Juglans (Juglandaceae) based on matK and ITS sequence
data. American Journal of Botany 87: 872–882.
http://dx.doi.org/10.2307/2656895
Swofford, D.L. (2002) PAUP* Phylogenetic analysis using parsimony (*and other methods). Version 4. Sinauer Associates, Inc.,
Sunderland, pp.1–142.
Triana, J.J. (1866) Dispositio Melastomacearum. Bulletin du Congres International de Botanique et d’Horticulture reuni a Amsterdam
1865, pp. 457–461.
Triana, J.J. (1871) Les Melastomacées. Transactions of the Linnean Society of London 28: 1–188.
http://dx.doi.org/10.1111/j.1096-3642.1871.tb00222.x
Whiffin, T. & Tomb, A.S. (1972) The systematic significance of seed morphology in the neotropical capsular–fruited Melastomataceae.
American Journal of Botany 59: 411–422.
http://dx.doi.org/10.2307/2441552
White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics.
In: Innis, M.A., Gelfand, D.H., Sninsky, J.J. & White, T.J. (Eds.) PCR Protocols: A Guide to Methods and Applications. Academic
Press, San Diego, pp. 315–322.
http://dx.doi.org/10.1016/b978-0-12-372180-8.50042-1
Wurdack, J.J. (1963) An evaluation of the genus Poteranthera. Fieldiana Botany 29: 535–541.
http://dx.doi.org/10.5962/bhl.title.2417

228 • Phytotaxa 263 (3) © 2016 Magnolia Press ROCHA ET AL.

 APPENDIX 1. List of species, voucher information, provenance and GenBank accession numbers for taxa included in this study. A dash (–) indicates that the DNA region was not sequenced.

Poteranthera
Herbarium acronyms according to Thiers (2016).
Species Voucher (herbarium) Provenance nrITS nrETS accD–psaI atpH–atpF trnS–trnG
Aciotis circaeifolia (Bonpl.) Triana Caddah, M.K., 621 (NY) Brazil JQ730038 KF462812 JQ730249 KU501108 KU500926
Aciotis paludosa (Mart. ex DC.) Triana Guimarães, P., 317 (RB) Brazil JQ730040 KF462814 JQ730251 – KU500925
Acisanthera alsinaefolia (DC.) Triana Goldenberg, R., 826 (NY) Brazil JQ730043 KF462817 JQ730254 KU501112 KU500930
Acisanthera bivalvis (Aubl.) Cogn. Rocha, M.J.R., 871 (BHCB) Brazil KU501064 KU501003 KU501172 KU501117 KU500936
Acisanthera boissieriana Cogn. Maguire, B., 35919 (NY) Venezuela – KU501002 – – –
Acisanthera crassipes (Naudin) Wurdack Rocha, M.J.R., 833 (BHCB) Brazil KU501065 KU501004 – KU501118 KU500937
Acisanthera genliseoides (Hoehne) Wurdack Rocha, M.J.R., 942 (BHCB) Brazil KU501066 KU501005 KU501173 KU501119 KU500938
Acisanthera hedyotidea (Presl) Triana Popovick, A., s.n. (BHCB) Brazil – KU501001 – – –
Acisanthera limnobios (Schrank & Mart. ex DC.) Moreira, S.N., 312 (BHCB) Brazil – KU501006 KU501174 – –
Triana
Acisanthera paraguayensis (Hook.f.) Cogn Krapovickas, A., 45640 (NY) Paraguai KU501058 KU500994 – KU501113 KU500931
Acisanthera quadrata Pers. Rocha, M.J.R., 729 (BHCB) Brazil KU501060 KU500996 KU501167 – –
Acisanthera tetraptera (Cogn.) Gleason Rocha, M.J.R., 757 (BHCB) Brazil KU501067 KU501007 KU501175 KU501120 KU500939
Acisanthera variabilis (DC.) Triana Rocha, M.J.R., 611 (BHCB) Brazil – KU500998 KU501170 KU501115 KU500934
Brachyotum microdon (Naudin) Triana Nee, M., 55327 (NY) Bolivia JQ730059 KF462828 JQ730270 – KU500984
Bucquetia glutinosa DC. Alvear, M., 1398 (CAS) Colombia JQ730061 – JQ730272 – –
Cambessedesia hilariana (A.St.Hil. ex Bonpl.) DC. Guimarães, P., 405 (RB) Brazil JQ730063 KF462835 JQ730274 - KU500987
Centradenia grandiflora (Schltdl.) Endl. ex Walp. Penneys, D.S., 1544 (FLAS) Costa Rica JQ730065 KF462837 – – –
...Continued on next page

Phytotaxa 263 (3) © 2016 Magnolia Press • 229

 APPENDIX 1. (Continued)
Species Voucher (herbarium) Provenance nrITS nrETS accD–psaI atpH–atpF trnS–trnG
Chaetolepis cufodontisii Standl. Michelangeli, F.A., 1160 (NY) Costa Rica JQ730067 – JQ730278 – –
Chaetostoma armatum (Spreng.) Cogn. Guimarães, P., 396 (RB) Brazil JQ730069 KF462840 – – –
Comolia microphylla Benth. Redden, K.M., 1454 (NY) Guyana JQ730070 KF462841 JQ730281 KU501125 KU500945
Comolia ovalifolia Triana Rocha, M.J.R., 719 (BHCB) Brazil KU501077 KU501020 – – –

230 • Phytotaxa 263 (3) © 2016 Magnolia Press

Comolia villosa (Aubl.) Triana Rocha, M.J.R., 739 (BHCB) Brazil KU501073 KU501019 – KU501126 KU500946
Desmocelis villosa (Aubl.) Naudin Zenteno, F., 8902 (NY) Bolivia JQ730073 KF462844 - - -
Dichaetanthera africana (Hook.f.) Jacq.–Fél. Smith, S., 1885 (US) Gabon JQ730074 KF462845 JQ730285 – KU845163
Fritzschia erecta Cham. Guimarães, P., 406 (RB) Brazil JQ730083 KF462849 JQ730296 – –
Fritzschia anisostemon Cham. Mota, N., 2694 (BHCB) Brazil KU501070 KU501012 KU501179 – KU500942
Heterotis decumbens (P.Beauv.) Triana Smith, S., 1705 (US) Brazil JQ730088 KF462853 JQ730302 - KU500985
Lavoisiera bicolor Naudin Guimarães, P., 345 (RB) Brazil JQ730091 KF462855 JQ730304 KU501157 KU500981
Lavoisiera imbricata (Thunb.) DC. Goldenberg, R., 820 (NY) Brazil JQ730091 – JQ730304 – –
Lavoisiera mucorifera DC. Guimarães, P., 345 (RB) Brazil JQ730092 – JQ730305 – –
Lavoisiera phyllocalycina Cogn. Guimarães, P., 345 (RB) Brazil JQ730092 KF462856 JQ730305 – –
Lavoisiera pulchella Cham. Goldenberg, R., 814 (NY) Brazil JQ730093 KF462857 JQ730306 – –
...Continued on next page

ROCHA ET AL.
 APPENDIX 1. (Continued)
Species Voucher (herbarium) Provenance nrITS nrETS accD–psaI atpH–atpF trnS–trnG

Poteranthera
Macairea radula (Bonpl.) DC. Lima, J., 715 (UPCB) Brazil JQ730095 KF462859 JQ730307 KU501133 KU500954
Macairea thyrsiflora DC. Wurdack, J.J., 4153 (NY) Guyana JQ730096 KF462860 KU501184 KU501132 KU500953
Marcetia taxifolia (A. St. Hil.) DC. Michelangeli, F.A., 680 (BH) Venezuela JQ730102 KU501041 JQ730311 KU501148 KU500970
Marcetia mucugensis Wurdack Rocha, M. J. R., 285 (BHCB) Brazil KU501088 KU501033 KU501192 KU501142 KU500963
Microlicia fulva (Spreng.) Cham. Michelangeli, F.A. 1576 Brazil KU501105 KU501051 KF407963 KU501156 KU500980
(UPCB)
Microlicia isophylla DC. Michelangeli, F.A., 1564 Brazil KU845166 – KF407965 KU845169 KU845162
(UPCB)
Microlicia cordata (Spreng.) Cham. Michelangeli, F.A., 1578 Brazil KU845165 – KF407962 KU845168 KU845161
(UPCB)
Poteranthera pussila Bong. Rocha, M.J.R., 940 (BHCB) Brazil KU845164 KU845167 – – –
Pterogastra divaricata (Bonpl.) Naudin Michelangeli, F. A., 540 (BH) Venezuela JQ730126 KF462875 JQ730337 KU501159 -
Pterolepis glomerata (Rottb.) Miq. Martin, C. V., 419 (NY) French JQ730129 KF462876 JQ730340 KU501158 KU500982
Guyana
Rhynchanthera bracteata Triana Zenteno, F., 8801 (NY) Bolivia JQ730137 KF462880 JQ730347 – –
Rhynchanthera grandiflora (Aubl.) DC. Michelangeli, F.A., 541 (BH) Venezuela JQ730138 KF462881 JQ730348 – –
Rhynchanthera serrulata (Rich.) DC. Michelangeli, F.A., 828 (BH) Venezuela AY460435 JQ730349 – –
Rhexia aristosa Britton Naczi, R. F. C., 12065 (NY) USA JQ730134 KF462878 – – KU500986
Siphanthera arenaria (DC.) Cogn. Mota, N., 2656 (BHCB) Brazil KU501099 KU501045 KU501203 – KU500974
Siphanthera cordata Pohl ex DC. Rocha, M J.R., 964 (BHCB) Brazil KU501102 KU501048 KU501208 – KU500977
KU501212
Siphanthera cordifolia (Benth.) Gleason Rocha, M.J.R., 882 (BHCB) Brazil KU501096 KU501042 KU501201 KU501151 KU500973
...Continued on next page

Phytotaxa 263 (3) © 2016 Magnolia Press • 231

 APPENDIX 1. (Continued)
Species Voucher (herbarium) Provenance nrITS nrETS accD–psaI atpH–atpF trnS–trnG
Siphanthera dawsonii Wurdack Versiane, A.F., 641 (HUFU) Brazil KU501103 KU501049 KU501207 – –
KU501211
Siphanthera fasciculata (Gleason) Almeda & O.R.Rob. Wurdack, J.J., 1959 (NY) Venezuela KU501097 KU501043 – – –
Siphanthera foliosa (Naudin) Wurdack Rocha, M.J.R., 728 (BHCB) Brazil KU501098 KU501044 KU501202 KU501152 –
Siphanthera gracillima (Naudin) Wurdack Rocha, M.J.R., 941 (BHCB) Brazil KU501101 KU501047 KU501206 KU501154 KU500976
KU501210

232 • Phytotaxa 263 (3) © 2016 Magnolia Press

Siphanthera hostmannii Cogn. Wurdack, K.J., 4142 (NY) Guyana JQ730142 KF462883 JQ730353 KU501155 KU500978
Siphanthera paludosa Cogn. Rocha, M.J.R., 621 (BHCB) Brazil KU501100 KU501046 KU501204 KU501153 KU500975
Siphanthera subtilis Pohl ex DC. Moreira, S.N., 878 (BHCB) Brazil KU501104 KU501050 KU501209 – KU500979
KU501213
Stenodon suberosus Naudin Almeda, F. A. 7861 (CAS) Brazil AY553725 – – – –
Svitramia wurdackiana R.Romero & A.B.Martins Guimarães, P., 395 (RB) Brazil JQ730148 KF462889 JQ730359 – –
Tibouchina cardinalis (Bonpl.) Cogn. Guimarães, P., 407 (RB) Brazil JQ730165 KF462908 JQ730376 – –
Tibouchina granulosa (Desr.) Cogn. Guimarães, P., 378 (RB) Brazil JQ730191 KF462934 JQ730399 – –
Tibouchina heteromalla (D.Don) Cogn. Almeda, F., 9868 (CAS) Brazil JQ730193 KF462936 JQ730401 – KU500983
Tibouchina lepidota (Bonpl.) Baill. Morales, M.E., s.n. (UPTC) Colombia JQ730201 KF462943 JQ730409 – –
Tibouchina longifolia (Vahl) Baill. Michelangeli, F.A., 816 (BH) Venezuela JQ730204 – JQ730412 – –
Trembleya laniflora (D.Don.) Cogn. Almeda et al. 8549 (CAS) Brazil AY553744 – – – –
Trembleya parviflora Cogn. Goldenberg, R., 824 (NY) Brazil JQ730242 KF462987 JQ730451 – –
Trembleya pentagona Naudin Almeda et al. 8555 (CAS) Brazil AY553745 – – – –

ROCHA ET AL.