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TAXON 00 (00) • 1–66 Guimarães & al.

• Systematics of Tibouchina and allies

SYSTEMATICS AND PHYLOGENY

Systematics of Tibouchina and allies (Melastomataceae:


Melastomateae): A new taxonomic classification
Paulo José Fernandes Guimarães,1 Fabián A. Michelangeli,2 Karla Sosa3 &
Jesús Ricardo de Santiago Gómez4
1 Instituto de Pesquisas Jardim Botânico de Rio de Janeiro, Rua Pacheco Leão 915, 22460-030, R.J., Rio de Janeiro, Brazil
2 The New York Botanical Garden, Bronx, New York, U.S.A.
3 Department of Biology, Duke University, Durham, North Carolina, U.S.A.
4 Departamento de Biología Comparada, Universidad Nacional Autónoma de México, Ciudad de México, México
Address for correspondence: Paulo José Fernandes Guimarães, paulojose.guimaraes@gmail.com
DOI https://doi.org/10.1002/tax.12151

Abstract Tibouchina (Melastomataceae: Melastomateae) is an exclusively Neotropical genus that currently includes close to 310 spe-
cies. These were first described or assigned to 16 other genera, a reflection of the complex taxonomic and phylogenetic history of the
genus. Previous phylogenetic analyses have shown that the genus is polyphyletic. Here we present an expanded analysis that includes
representatives of 28 genera of New World Melastomateae, as well as representatives of Old World Melastomateae and members of
the Marcetieae and Microlicieae. Over 300 species were sequenced for nrITS and the plastid spacers accD-psaI and psbK-psbL. We
recovered a large clade of Neotropical Melastomateae that contains Tibouchina along with at least 13 other Neotropical genera. More-
over, Tibouchina is not recovered as monophyletic because three mostly Central American and five mostly Andean genera are nested
within it, albeit each these genera are recovered as monophyletic. Many of the previously recognized sections of Tibouchina are recov-
ered as monophyletic or paraphyletic providing a basis for a new classification. Tibouchina is here re-circumscribed into four genera
that are each monophyletic and easily diagnosable. This results in a narrowly circumscribed Tibouchina and the re-establishment of
Pleroma and Chaetogastra, and the description of a new genus, Andesanthus. Pleroma now also includes Itatiaia, Microlepis and Svi-
tramia. Brachyotum is maintained until its relationship to Chaetogastra can be further examined. This new taxonomic arrangement is
more informative and convenient than merging eight other genera into Tibouchina. An identification key, descriptions of genera, and
lists of taxa with their current generic assignment and necessary nomenclatural novelties are also presented here.
Keywords Andesanthus; Brachyotum; Chaetogastra; Neotropical; Svitramia

Supporting information may be found online in the Supporting Information section at the end of the article.

■ INTRODUCTION The Neotropical Tibouchina, with almost 310 species, is


the argest genus of Melastomataceae with capsular fruits.
Recent phylogenetic analyses have greatly increased our While Tibouchina species range in habit from herbs to trees
understanding about the evolutionary relationships of Neotropi- over 20 meters, most are shrubs. Tibouchina has been tradi-
cal capsular-fruited Melastomataceae Juss. (Goldenberg & al., tionally characterized by its pentamerous flowers (rarely
2012, 2015; Michelangeli & al., 2013, 2016; Almeda & al., tetramerous) with a corresponding number of locules in the
2016; Rocha & al., 2016). As a result of these studies, the ovary, terete hypanthia, stamen connectives often prolonged
boundaries of the tribe Melastomateae are now well established, with two ventral appendages, setose or pilose ovary apex
excluding Marcetieae, Rhexieae and Microlicieae (Michelangeli and cochleate and tuberculate seeds (Cogniaux, 1885, 1891).
& al., 2013; Rocha & al., 2016, 2018). These analyses have The closely related Brachyotum with 56 species is the
shown that the Melastomateae includes both New World and second-largest Neotropical genus of the Melastomateae. It dif-
Old World elements with the African and Asian species forming fers from Tibouchina by having flowers with petals that do not
a monophyletic group nested within the Neotropical Melasto- spread at anthesis, thus forming a tube, and that are pollinated
mateae (Michelangeli & al., 2013). These same analyses showed by hummingbirds. Tibouchina has a widespread distribution,
that although most Neotropical genera of Melastomateae are ranging from western Mexico and the West Indies to northern
monophyletic (or could be rendered monophyletic with small Argentina and Paraguay, and it is highly diverse in Brazil
taxonomic changes), the two largest Neotropical genera in the where two-thirds of the species occur. Brachyotum is largely
tribe, Tibouchina Aubl. and Brachyotum (DC.) Triana, are poly- restricted to páramo and subpáramo zones from Colombia to
phyletic and paraphyletic, respectively. northern Argentina.

Article history: Received: 8 Mar 2018 | returned for (first) revision: 4 May 2018 | (last) revision received: 12 May 2019 | accepted: 10 Jun 2019
Associate Editor: Eric H. Roalson | © 2019 International Association for Plant Taxonomy

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Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

The absence of constant or well-defined floral morpho- distinctifs” (In their organization, there is no response to the
logical features (Fig. 1) in Tibouchina and many other genera idea of genus, or any other division that consists of a grouping
of Melastomateae has resulted in different generic circum- of species that could be isolated by a special character and dis-
scriptions and taxonomic classifications. Triana (1872: 3) tinctive features). The Colombian botanist José Jerónimo Tri-
wrote: “Dans leur organisation, rien ne répond à l’idée de ana Silva (1826–1890) had extensive experience in the field
genre, ni de toute autre division composée d’une réunion d’es- and carried out an in-depth study of major collections in
pèces, ou s’isolant par un caractère spécial et par des signes European herbaria. At the Botanical Congress in Amsterdam

Fig. 1. Photographs of Tibouchina and allies: A, Andesanthus lepidotus; B, Bucquetia glutinosa; C, Castratella piloselloides; D, Centradenia inae-
quilateralis; E, Chaetogastra longifolia; F, Brachyotum radula; G, Chaetolepis microphylla; H, Desmoscelis villosa; I, Heterocentron elegans;
J, Monochaetum myrtoideum; K, Pilocosta nana; L, Pleroma bandeirae (= Svitramia pulchra); M, P. candolleanum; N, Schwackaea cupheoides;
O, Tibouchina aspera; P, T. johnwurdackiana. — A–D, F, G, I–K & N by Fabián A. Michelangeli; E by Ricardo Kriebel; H, L, M, O & P by Paulo
J.F. Guimarães.

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TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

in 1865, Triana introduced a classification system for the Me- Tibouchina was established by Aublet (1775) in his Flora
lastomataceae that was followed by most other specialists until of French Guiana with the description of a single species, T.
its re-evaluation based on phylogenetic concepts by Ren- aspera. Scopoli (1777) subsequently established Savastania
ner (1993). to accommodate T. aspera. However, Tibouchina has priority
This lack of understanding the importance of any given because it was validly published first, and Savastania is a
diagnostic character is evidenced by how the generic bound- superfluous name.
aries of Tibouchina have changed since its original description Jussieu (1789), in the description and circumscription of
(Aublet, 1775). This has resulted in a large number the “Ordo Melastomae”, included Tibouchina among the spe-
of nomenclatural synonyms in some genera. For example, cies with a superior ovary. Desrousseaux (1797) argued that
Baillon (1877) suggested that 20 other genera should be the bracteoles at the base of the hypanthium are common
included in Tibouchina; Cogniaux (1885, 1891) later reversed among the Melastoma L. species, as well as the hypanthium
these changes but transferred to Tibouchina species that were covered by scale-like trichomes, and thus, he did not consider
initially described in 16 other genera. These two examples Tibouchina sufficiently distinct from Melastoma to merit
demonstrate the lack of agreement with respect to the circum- generic rank.
scription of Tibouchina. The generic limits of Tibouchina Ventenat later (1807) advocated that the diversity of Me-
were more recently questioned by Renner (1989: 69), who lastomataceae was best represented by only four genera. Fol-
stated that “It is possible that the more than 250 species of lowing this concept, he transferred Melastoma and
Tibouchina form a paraphyletic group; the problem of circum- Tibouchina to Meriana based on the presence of dry capsular
scribing Tibouchina is confounded by several medium-sized fruits (note that in this study, there was an orthographic change
genera, for instance Brachyotum, Desmoscelis, Pterolepis of Meriania Sw., with the suppression of the letter i from
and Pterogastra, which appear to be segregates of the larger the name).
group.” David Don (1823) did not follow Ventenat’s (1807) cri-
Although it has been clear for some time that the taxon- teria and considered Tibouchina aspera as part of Melastoma
omy of Neotropical Melastomateae should be revised, this due to the closer resemblance of this Neotropical species to
was not possible in the absence of molecular phylogenetic species of Melastoma from the Republic of Sierra Leone
analyses that would allow an evaluation of relationships of (West Africa) he had seen in the herbarium of Aylmer Bourke
all constituent genera in the tribe and the potential taxonomic Lambert. Simultaneously, Don (1823) established Pleroma,
value of different morphological characters. The analysis of which he considered to be close to Melastoma, but distin-
Neotropical Melastomateae (Michelangeli & al., 2013) guished it by the deciduous calyx lobes that fall at about the
showed that although Tibouchina is not monophyletic, several same time as the petals, and the subisomorphic anthers with
of the previously defined sections or groups of sections (see an arched base lacking bifurcated appendages. Under this con-
Cogniaux, 1885, 1891) could be rendered monophyletic, thus cept of Pleroma, he grouped species previously described
shedding some light on which characters could be used in an under Melastoma and Rhexia L., and added another unpub-
updated classification. lished one, totaling five species. Don (1823) also described
Here we provide an updated phylogenetic analysis for Diplostegium D.Don, characterized by the presence of a dou-
Tibouchina and allies together with an updated classification. ble calyptra, and validated Arthrostemma. The genus Arthros-
temma was initially suggested by Pavón in two unpublished
illustrations of Flora Peruviana, et Chilensis: number
■ TAXONOMIC AND NOMENCLATURAL 326, Arthrostemma ciliatum, and 327, A. caulialatum (= Acio-
HISTORY tis D.Don. in Triana, 1872). Don (1823) published A. ciliatum
but rejected the other species. However, he described a second
The taxonomic and nomenclatural history of Tibouchina section for Arthrostemma, expanding his concept of this
and Brachyotum is one of constant changes regarding the per- genus. This second section was characterized by anthers with
ceived importance of various diagnostic characters. Thus, an single appendages, and it included species that are currently in
almost continuous flow of changes in the generic limits the Neotropical Monochaetum (DC.) Naudin and Oxyspora
ensued for most of the 19th century. The species currently DC., an Asian genus of Sonerileae.
placed in Tibouchina were at different times assigned to Ple- Candolle, in his Prodromus (1828a), greatly advanced the
roma D.Don, Lasiandra DC. and Chaetogastra DC. Mean- taxonomy of Melastomataceae, erecting new genera or recir-
while, those species belonging to Brachyotum were at cumscribing previously described ones. He restored Tibou-
different points assigned to either Arthrostemma Pav. ex china and considered it a monotypic genus. Candolle
D.Don or Chaetogastra. The circumscription of Tibouchina (1828b) established the genus Lasiandra to which he trans-
and its infrageneric classification are associated with the his- ferred some species in Pleroma. At the same time, he also
torical discussion of the different concepts used for the limits described in this genus several species suggested by Schrank
of this taxon, discussed in more detail by Souza (1984) and and Martius. Lasiandra, unlike Pleroma, according to Can-
Guimarães (1992). A historical review of Brachyotum was dolle (1828b), are never glabrous but rather hispid, the repro-
presented by Wurdack (1953) in his monograph of the genus. ductive organs are covered with trichomes, and the dry

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capsule is never adnate to the hypanthium. Candolle (1828a) species from 21 to 108 and the number of sections from 3 to
also described Chaetogastra and subdivided it into three sec- 17. He transferred Lasiandra oleifolia to Ancistrodesmus
tions: Chaetogastra sect. Bractearia DC., C. sect. Dio- Naudin (= Microlepis).
tanthera DC. and C. sect. Nonocentra DC. Chaetogastra Naudin (1849, 1850) was responsible for a significant
differed from Lasiandra by the presence of glabrous stamens realignment of the taxonomy of the Melastomataceae, includ-
and from Arthrostemma by the presence of pentamerous ing novel generic limits for Arthrostemma and Chaetogastra.
flowers. Furthermore, Chaetogastra differed from In his system, he carved out of Chaetogastra four new genera:
Osbeckia L. by the absence of appendages between the lobes Desmoscelis, Pterogastra Naudin, Hephestionia Naudin, and
of the calyx, and from Melastoma by its free capsular fruit. Oreocosmus Naudin (the latter two currently part of Tibou-
Under this concept, Chaetogastra comprises species that china). Chaetogastra was restricted to those species with per-
today belong to Brachyotum, Meriania, Purpurella Naudin, sistent calyx lobes, shortly prolonged connectives, and
Tibouchina sect. Diotanthera and T. sect. Lepidotae. glabrous filaments (Naudin, 1850). Naudin established two
Candolle (1828a) also recognized Pleroma, and he sections for Chaetogastra based on the morphology of stamens.
divided the species into two groups. The first consisted of Chaetogastra sect. Lasiandrellae included species that are now
Pleroma laxum (= T. laxa, T. sect. Diotanthera), P. lendifo- in Tibouchina (T. bicolor, T. longifolia, T. panicularis), while
lium (= Macairea lendifolium = M. radula) and P. trichopo- in C. sect. Dicenrae there were species of Brachyotum and
dum (= T. trichopoda, T. sect. Pleroma). The second group, Tibouchina (B. campanulare, B. cernuum, B. confertum,
which accounted for 60% of the genus (Candolle, 1828a: B. ledifolium, B. lutescens, B. microdon, B quinquenervia, B. ros-
151–152), was made up of species whose generic placement marinifolium, B. rostratum, B. sanguinolentum, B. strigosum,
he was unsure of (currently four of these belong to Tibouchina T. lepidota). Naudin placed C. nigritella (= Bucquetia nigritella
and one to Desmoscelis Naudin). (Naudin) Triana) in this group, but he expressed doubts due to
Candolle (1828a) recognized Arthrostemma and inter- the differences in habit and leaves and the shorter anther connec-
preted it in a very broad sense, distinguished mostly by the tive. Naudin’s (1850) concept of C. sect. Dicenrae is the closest
presence of tetramerous flowers. In this expanded context, to the modern concept of Brachyotum.
Arthrostemma included species that had been assigned to Bra- After the changes proposed by Naudin (1850), Arthro-
chyotum, Castratella Naudin, Comolia DC., Monochaetum, stemma was essentially restricted to the species that are today
Pterolepis (DC.) Miq. and Tibouchina. in Pterolepis, plus Tibouchina herbacea (DC.) Cogn. The
Shortly thereafter, Martius (1829) recognized Lasiandra type, Arthrostemma ciliatum Pav. ex D.Don was transferred
by the deciduous calyx lobes and often villous filaments. In to Heteronoma DC., which created a nomenclatural problem.
this same publication, he transferred Osbeckia oleifolia At the same time, Naudin elevated Arthrostemma sect. Mono-
DC. to Lasiandra. chaetum DC. to generic status. In this work, Naudin described
Chamisso (1835) expanded the generic limits of Lasiandra, Tetrameris to accommodate six species currently placed in
and he included in this genus most species that up until then had Castratella, Comolia, Macairea DC. and Tibouchina.
been placed in Pleroma. Under this concept, he also included Triana (1872) recognized that Candolle’s distinction
Tibouchina, which has scale-like trichomes and glabrous fila- between Lasiandra and Pleroma was based on a highly vari-
ments, and Diplostegium, with calyptra-shaped bracteoles. He able character with little taxonomic importance: the presence
kept in this genus Lasiandra oleifolia (DC.) Mart. (= Microlepis of trichomes on the filament. Triana (1872) mentioned that
oleifolia (DC.) Triana). Chamisso (1835) transferred some Neo- there was already a consensus that considered Lasiandra a
tropical species of Osbeckia DC. to Chaetogastra, which are synonym of Pleroma. Furthermore, he also recognized that
currently in Desmoscelis, Pterolepis and Tibouchina. He estab- Tibouchina was part of this group of species, but because the
lished Svitramia Cham. to accommodate S. pulchra Cham., latter was little known at the time and contained only a few
which he considered distinct from Chaetogastra due to its calyx, species, the use of this name would create more confusion.
stamens and habit. Wurdack (1973a) described a second species Triana (1872: 12) argued that “thanks to its euphony and also
of Svitramia, and Romero & Martins (2003) published four new because it indicates a very general character, which is common
species; thus, Svitramia currently comprises six species. to most species” (trans. from French) he adopted the generic
Miquel (1840) established the genus Microlepis to accom- name Pleroma.
modate Osbeckia sect. Microlepis DC. This section consisted Triana (1872) had a much narrower concept of Arthro-
of four species that currently belong to Cambessedesia stemma while considering Heteronoma as a synonym. He con-
DC. (O. pityrophylla DC.), Miconia Ruiz & Pav. (O. jucunda sidered that Chaetogastra was not a natural group and
DC.), Microlepis (O. oleifolia DC.) and Tibouchina transferred its species to Brachyotum, Bucquetia Naudin, Des-
(O. urceolaris DC.). However, Miquel (1840) did not specifi- moscelis, Macairea, Meriania, Monochaetum, Pleroma and
cally transfer any species. Miquel (1849) eventually formally Pterolepis.
transferred Osbeckia oleifolia var. quaternifolia DC. to Micro- The name Chaetogastra resurfaced again when Goyena
lepis quaternifolia (DC.) Miq. (1909: 413) in the Flora of Nicaragua used it to accommodate
Naudin (1849) adopted the circumscription of Lasiandra C. dichotoma Ram.Goyena. This species is now considered a
proposed by Chamisso (1835), increasing the number of synonym of Tibouchina longifolia (Vahl) Baill.

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The current concept of Tibouchina can be traced back to In total, ca. 470 taxa have been assigned to Tibouchina.
Baillon (1877). He considered the morphology of stamens, Only two groups have been revised in the last 30 years: Todzia
especially the presence of a pedoconnective, as a diagnostic & Almeda (1991) published a revision of T. sect Lepidotae
character to restructure the genus Tibouchina. However, at with nine species, and Guimarães (1997) revised T. sect. Ple-
the time he did not make any of the necessary taxonomic roma with 41 recognized species.
changes. Moreover, the presence of a pedoconnective is a syn- The consequences of the phylogenetic analyses of Miche-
apomorphy shared with Microlicieae (Clausing & Renner, langeli & al. (2013) for the taxonomy and nomenclature of
2001; Fritsch & al., 2004). Melastomateae can already be seen, even though no compre-
Cogniaux (1885), in his treatment for Flora brasiliensis, hensive taxonomic realignment has been proposed until now.
mostly followed Baillon’s concept of an expanded Tibou- Pleroma was recently reestablished to accommodate P. cleis-
china, and he transferred into it many of the species described toflora (Ule) P.J.F.Guim. & al. (Oliveira da Silva & al.,
or placed at that time in Chaetogastra, Diplostegium, Lasian- 2014). This species of Itatiaia was transferred to Pleroma to
dra, Pleroma and Purpurella, among others. Simultaneously, reflect the findings of recent phylogenetic analyses
Cogniaux (1885) adopted into Tibouchina most of the sec- (Michelangeli & al., 2013). Similarly, Romero & Versiane
tional classification that Triana (1872) had employed for Ple- (2014) transferred Microlepis oleifolia to Pleroma. Addition-
roma. He divided Tibouchina into ten sections, restoring ally, 13 new species have been described in Pleroma, and
sections proposed by Candolle (1828a), Naudin (1849, 3 species of Tibouchina have been transferred (Fraga & Gui-
1850) and Triana (1872). Later, in his global monograph for marães, 2014; Meyer & Goldenberg, 2014; Oliveira & al.,
the family Melastomataceae, Cogniaux (1891) added an elev- 2014; Guimarães & Freitas, 2015; Guimarães & Oliveira da
enth section, T. sect. Octomeris Cogn. Silva, 2015; Meyer & al., 2016; Goldenberg & Kollmann,
When transferring Purpurella to Tibouchina, Cogniaux 2016; Freitas & Van Den Berg, 2016; Meyer & Matos,
(1885) also expanded its limits by including species with 2017). Meyer & Goldenberg (2016), to avoid future changes,
anthers that have a truncated apex, not attenuated, and dehis- described four new species from southern Brazil inside
cent by a large pore. Naudin (1850), Triana (1872), Krasser Chaetogastra.
(1893) and Pereira (1959–1961) all treated Purpurella as dis-
tinct from Tibouchina (or Pleroma), while Rambo (1958),
Wurdack (1962) and Souza (1986) all accepted T. sect. ■ MATERIALS AND METHODS
Purpurella.
Ule (1895) published Purpurella cleistoflora, but in the Taxon sampling. — To expand previous analyses of the
next year he modified the epithet of this species to P. cleisto- Melastomateae (Michelangeli & al., 2013), we increased the
petala (Ule, 1896a). Later, Ule (1908) considered that the sampling with 98 additional taxa, some of which were pro-
closed configuration of the petals in this species would be duced for a study of the Marcetieae (Rocha & al., 2016).
more than enough to segregate it from Tibouchina or Brachyo- Whenever possible, we tried to cover the morphological vari-
tum, which have distended or campanulate petals in the anthe- ation and geographical distribution for each genus, and speci-
sis, respectively. Ule (1908) then established the genus Itatiaia mens from all sections of Tibouchina were analyzed. Trees
Ule to accommodate I. cleistopetala. were rooted with Cambessedesia, following the results of pre-
Tibouchinopsis Markgr. was described by Markgraf vious analyses of the Melastomateae and the family as a whole
(1927) to accommodate T. glutinosa Markgr. Later, Brade (see Goldenberg & al., 2012; Michelangeli & al., 2013, 2014;
& Markgraf (1961) added a second species, T. mirabilis Brade Rocha & al., 2016).
& Markgr. Tibouchinopsis differs from Tibouchina by the DNA extraction and sequencing. — Total genomic
absence of trichomes, presence of a glutinous surface on DNA was isolated from silica-dried or herbarium material
leaves and branches and a glabrous ovary. using the DNAeasy plant mini kit from Qiagen
Wurdack (1953) published the latest revision of Brachyo- (Germantown, Maryland, U.S.A.), following the manufac-
tum, in which he recognized 44 species. With more recent turer’s protocol. Some samples were isolated with modifica-
descriptions, the genus has been expanded to include 56 spe- tions that have shown to improve DNA quality and yield in
cies. It should be noted that in this work, Wurdack designated the Melastomataceae and/or from herbarium specimens as
Chaetogastra longifolia (Vahl) DC. as the type of Chaetogas- previously described by Martin & al. (2008) and Michelangeli
tra. He designated this species as the type mostly because he & al. (2008, 2013). Phylogenetic analyses were based on three
considered it to be part of Tibouchina, and thus he would not molecular markers: the nuclear internal transcribed spacer
have to consider whether Chaetogastra was part of Brachyo- (nrITS) and the plastid intergenic spacers accD-psaI and
tum and the possible generic limits between these two genera. psbK-psbL, the same regions employed by Michelangeli
Obviously, while his intention to synonymize Chaetogastra & al. (2013). The nrITS has been used to elucidate phyloge-
under Tibouchina was clearly stated and appropriate in the netic relationships at the tribal and generic levels in the Melas-
interest of taxonomic stability, he did not foresee our current tomataceae (Fritsch & al., 2004; Michelangeli & al., 2004,
understanding of phylogenetic relationships between these 2008, 2013; Ionta & al., 2007; Bécquer-Granados & al.,
genera. 2008; Goldenberg & al., 2008; Martin & al., 2008; Reginato

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Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

& al., 2010; Kriebel & al., 2015), and it was amplified as (Rambaut & al., 2014), and runs were considered satisfactory
either one fragment or in two sections using the primers and with effective sample size (ESS) values greater than 200. The
conditions outlined by Michelangeli & al. (2004). The psbK- stable posterior distributions of the independent runs were
psbL spacer was amplified using primers designed by Kim- combined using LogCombiner v.2.5.0 (Rambaut & Drum-
Joong Kim (Korea University, South Korea) and published mond, 2018a), and a maximum clade credibility tree was sum-
by Reginato & al. (2010), and the accD-psaI spacer was marized with TreeAnnotator v.2.5.0 (Rambaut & Drummond,
amplified using primers designed by Shaw & al. (2005). The 2018b).
two plastid spacers have been informative for Melastomata- Nomenclature and taxonomy. — We built a database
ceae and have been employed in several studies (Reginato with all validly published names in the genera Brachyotum,
& al., 2010; Michelangeli & al., 2013; Penneys & Judd, Itatiaia, Microlepis, Pleroma, Purpurella, Svitramia, Tibou-
2013; Kriebel, 2016; Rocha & al., 2016), and we followed china and Tibouchinopsis. These genera were selected based
the same protocols. Cycle sequencing was performed with on the results of phylogenetic analyses. Heterotypic synonyms
the same forward and reverse primers used for amplification prior to Cogniaux (1891) were not listed for the species. The
through the High-Throughput Sequencing Service of the Uni- nomenclatural data assembled here are based on Index
versity of Washington. Sequence contigs were built with Kewensis and Gray Herbarium Card Index (both available at
Sequencher v.4.9 (Gene Codes, Ann Arbor, Michigan, http://www.ipni.org), The Plant List (http://www.theplantlist.
U.S.A.). Sequence alignment was performed on each locus org) and Tropicos (http://www.tropicos.org), with later addi-
individually with MAFFT v.7 (Katoh & al., 2005) as imple- tions of recently described species. Authors and original pub-
mented through Geneious v.9.1.7. (http://www.geneious. lications were checked for each species and corrected when
com, Kearse & al., 2012). necessary.
We also generated sequences for about 85% of the terminals All genera and species currently accepted in the core Melas-
for ca. 650 bp of the 3′ end of the nuclear external transcribed tomateae clade were morphologically analyzed from field-col-
spacer (nrETS). This locus has been used successfully in other lected specimens, herbarium collections or original descriptions.
Melastomataceae phylogenetic studies (Stone & Andreasen, All taxonomic and nomenclatural adjustments proposed
2010; Kriebel & al., 2015; Majure & al., 2015; Reginato here are in accordance with the following criteria: (a) to recog-
& Michelangeli, 2016). However, within the Melastomateae it nize only monophyletic and well-supported groups; and (b) to
seems that there are either paralogous copies or PCR chimeric recognize groups that are morphologically distinguished by
artifacts that could not be resolved at this time, and thus, these synapomorphies or by a combination of diagnostic morpho-
data were not used in the final analyses. logical characters, and that are also usually geographically
Phylogenetic analyses. — Dataset alignments and phy- structured. Based on the results of our phylogenetic analyses
logenetic trees are deposited in the Dryad Digital Repository and the premise that genera are supported monophyletic
(https://doi.org/10.5061/dryad.4m6s6). All newly obtained groups recognized by morphological characteristics, a new
DNA sequences were deposited in GenBank (Appendix 1). classification is proposed below for Tibouchina and allies.
DNA substitution models for Bayesian inference (BI) and The types of generic names and their taxonomic syno-
maximum-likelihood (ML) analyses were selected for each nyms were examined either directly, through online image
marker using jModeltest v.2.1.3 (Darriba & al., 2012). Sepa- databases (such as JSTOR Global Plants: http://plants.
rate phylogenetic analyses under ML were conducted for each jstor.org/; Herbarium Hamburgense, HBG: http://www.
of the three datasets (suppl. Figs. S1–S4). ML analyses were herbariumhamburgense.de/colldata_hbg.php), or by physical
performed in RAxML v.8.2.10 (Stamatakis, 2014) and run photographic images. Type specimens examined through
through the CIPRES Science Gateway (http://www.phylo. online resources are cited as “online image!”. Other types,
org/; Miller & al., 2010). Rapid bootstrapping (BS) was done mostly B material that is presumed lost during World War II
on the ML tree using RAxML at 1000 replicates to determine but seen as mounted photographs from F were described as
branch support. Once we ensured there was no topological “negative” with the corresponding original herbarium. Frag-
conflict among loci (BS threshold >70), all loci were com- ments when duly registered are considered by us to be dupli-
bined into a single matrix. ML was run on the combined cates of the type collection, as determined by the Shenzen
matrix with four partitions maintaining the same parameters Code in Art. 8 Ex. 8: “It [the fragment] is a duplicate, i.e. an
as above. isotype” (Turland & al. 2018).
BI analyses were performed in BEAST2 v.2.5.0
(Drummond & Bouckaert, 2014), as implemented in the
CIPRES portal. Each partition was assigned the GTR+Γ+I ■ RESULTS
model, and sequence evolution partitions were unlinked.
Rate variation across branches was set as uncorrelated and ML and BI analyses were highly congruent, with mostly
lognormally distributed, and with tree prior set to the Yule minor differences in derived nodes with low support values.
process. We ran three independent analyses of 60 million The only exception concerns the relationship between Bra-
generations each, sampling every 20,000 generations with a chyotum and Chaetogastra (see below). In the interest of clar-
20% burn-in. Convergence was assessed using Tracer v.1.6 ity and consistency, all results discussed are based on the ML

6
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

results unless specifically noted otherwise. Melastomateae ■ DISCUSSION


s.l. are recovered in two separate non-sister clades: one con-
taining the exclusively Neotropical Marcetieae and one con- Our results agree with previous phylogenetic analyses of
taining both New World and Old World Melastomateae the capsular-fruited Melastomataceae with the recognition of
(Fig. 2). The tribes Rhexieae and Microlicieae are both recov- four major clades: Marcetieae, Melastomateae, Microlicieae
ered as monophyletic with high support, and sister to each and Rhexieae (Michelangeli & al., 2013; Kriebel, 2016;
other with moderate support. The Rhexieae+Microlicieae Rocha & al., 2016; Veranso-Libalah & al., 2017). Although
clade is recovered as sister to core Melastomateae, albeit with these four clades are well supported, our increased taxon sam-
low support. pling did not provide higher support for the relationships
Loricalepis Brade is resolved as sister to all remaining among them.
Melastomateae. Pterolepis and Pterogastra (probably with Here we recover Loricalepis, a monotypic genus from
Tibouchina cubense) form a clade sister to a monophyletic white-sand savannas in northern Amazonia, and not previ-
Old World Melastomateae, and this larger clade is in turn sister ously sampled, as sister to all remaining Melastomateae.
to a large clade that includes the Neotropical genera Brachyo- Loricalepis is a poorly collected taxon with a character combi-
tum, Bucquetia, Castratella, Centradenia G.Don, Chaetolepis nation that had made it difficult to place in any other genus or
(DC.) Miq., Desmoscelis, Heterocentron Hook. & Arn., Ita- even unequivocally in a tribe. Michelangeli & al. (2013) had
tiaia, Microlepis, Monochaetum, Pilocosta Almeda & Whiffin, suggested that based on its small habit and tetramerous
Pterolepis, Pterogastra, Schwackaea Cogn., Svitramia, Tibou- flowers it may be placed in the Marcetia alliance. However,
china, and Tibouchinopsis, with Desmoscelis resolved as sister the cochleate seeds of “Tibouchinoid” type (Brade, 1938;
to the remaining species of this clade. Whiffin & Tomb, 1972), as well as the presence of a corona
Tibouchina is not recovered as monophyletic, but rather in of setae on the ovary apex (Brade, 1938) support its placement
four distinct clades. Although the relationships of these clades in the Melastomateae.
to each other and to the remaining members of the Melastoma- As shown by Michelangeli & al. (2013), Centradenia,
teae are weakly or moderately supported, each of these four Heterocentron, and Pilocosta are all part of a mostly Central
groups is strongly supported. All sampled members of Tibou- American clade. This group also includes Schwackaea, a
china sect. Tibouchina and T. sect. Barbigerae form a clade monotypic genus not sampled in that study, but that Kriebel
(from now on called Tibouchina s.str.) sister (with low sup- (2016) had already shown is part of this clade and sister to
port) to a clade that contains all members of T. sect. Pleroma, Pilocosta. Similarly, Bucquetia, Castratella, Chaetolepis and
T. sect. Involucrales and T. sect. Diplostegia as well as Ita- Monochaetum are resolved in a mostly Andean clade. These
tiaia, Microlepis, Svitramia and Tibouchinopsis (from now results are consistent with those of Michelangeli & al.
on called Pleroma). This Tibouchina s.str. + Pleroma clade (2013), so they will not be discussed further here.
is resolved as sister to a polytomy that contains three well to A new taxonomic framework for Tibouchina. — While
moderately supported clades: clade 1, composed of mostly Tibouchina is not resolved as monophyletic, all other genera
Central American taxa in Heterocentron (including Tibou- in the Tibouchina clade are recovered as monophyletic
china breedlovei), Centradenia, Schwackaea and Pilocosta; (or closely so, see below). Moreover, all clades that contain
clade 2, a mostly Andean clade composed of Bucquetia, Cas- species of Tibouchina are composed of species that belong to
tratella, Chaetolepis and Monochaetum; clade 3, composed of one or few sections of this genus, evidence that these clades
all members sampled of Tibouchina sect. Lepidotae, T. sect. are often also supported by morphological characters. This
Diotanthera, T. sect. Octomeris, T. sect. Simplicicaulis and means that a new taxonomic framework is needed for Neotrop-
T. sect. Pseudoterolepis, as well as all members of Brachyo- ical Melastomateae, one that is simultaneously consistent with
tum. In this last group, members of Tibouchina sect. Lepidotae a phylogenetic framework and based on diagnosable characters
(from now on Andesanthus) form a clade sister to the remain- or character combinations. Thus, the species traditionally
ing members of the clade (from now on Chaetogastra placed in Tibouchina (sensu Cogniaux, 1885, 1888, 1891) are
+ Brachyotum). In both ML and BI analyses, Brachyotum is here recognized in four different genera: Andesanthus, Chaeto-
not resolved as monophyletic, as some Andean species of gastra, Pleroma and Tibouchina. Alternative classifications
Tibouchina are intermixed within it. The main difference with additional or fewer genera were considered by us, but
between ML and BI is that in the ML analyses Chaetogastra we strongly feel that this is the most stable arrangement: It
is resolved as paraphyletic since the expanded Brachyotum allows for all the genera to be diagnosable and retains most
clade is derived from within it. In this case, the support for of the structural and biogeographical complexity of the group.
the expanded Brachyotum clade is high, but the support for Perhaps the most difficult decision was whether to merge Bra-
the previous two nodes is very low. Conversely, the BI ana- chyotum into Chaetogastra, or to recognize it as a separate
lyses recover Chaetogastra and the expanded Brachyotum genus. However, given the current sampling and phylogenetic
clades as sisters and reciprocally monophyletic. In this case results, we have chosen to recognize Brachyotum for the time
the support for the Chaetogastra + Brachyotum clade is low being as separate from Chaetogastra (see below).
as is the support for Chaetogastra, while the support for Bra- Tibouchina s.str. (as here narrowly defined) is composed
chyotum is moderate. of the species in T. sect. Tibouchina and T. sect. Barbigerae.

7
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

100 Cambessedesia hilariana


Cambessedesia espora
86
Marcetieae
100
75 Rhexieae
100 Microliceae
Loricalepis duckei
100 Pterolepis rotundifolia
100 Pterolepis parnassiifolia
100 Pterolepis stenophylla
72 Pterolepis glomerata
100 Pterogastra divaricata
96
Pterogastra minor
Pterolepis alpestris
94
94
Pterolepis repanda
95
Pterolepis polygonoides
Pterolepis trichotoma
Pterolepis sp.
100 Antherotoma phaeotricha
97 Guyonia ciliata
Tristemma mauritianum var. mauritianum
100 Tristemma littorale
83
99
Tristemma coronatum
Tristemma hirsutum
100 Dichaetanthera oblongifolia
73 Dichaetanthera africana
100 Heterotis decumbens
Heterotis rotundifolia
Dupineta multiflora
100 Antherotoma naudinii
Amphorocalyx rupestris
91 Amphorocalyx multiflorus
Rousseauxia andringitrensis
Dionycha bojeri
Rousseauxia minimifolia
Osbeckia stellata
// Osbeckia aspera var. courtallensis
76 Osbeckia australiana
Osbeckia nepalensis
Melastoma dodecandrum
100 87Melastoma malabathricum 2 (Melastoma affine)
Melastoma intermedium
79 Melastoma denticulatum
100 Melastoma malabathricum 3 (M. candidum)
Melastoma sanguineum
Melastoma malabathricum 1
Desmocelis villosa
Tibouchina aspera var. aspera

Tibouchina
100
Tibouchina aspera var. asperima
96 95
Tibouchina papyrus
90
Tibouchina aegopogon
98 Tibouchina llanorum
Tibouchina bipenicillata
99 Tibouchina striphnocalyx
Tibouchina fraterna
Pleroma raddianum (Tibouchina pulchra)
Pleroma aemulum 1 (T. valtherii)
Pleroma semidecandrum (T. semidecandra)
Pleroma wurdackianum (Svitramia wurdackiana)
95
79
Pleroma sp. 2 (Svitramia sp.)
97 Pleroma bandeirae (S. pulchra)
Pleroma gertii (S. hatschbachii)
Pleroma sp. 1 (Svitramia sp.)
Pleroma minus (S. minor)
Pleroma ursinum (T. ursina)
Pleroma kleinii (T. kleinii)
Pleroma dubia (T. dubia)
Pleroma ramboi (T. ramboi)
Pleroma echinatum (T. gardneriana)
Pleroma aemulum 2 (T. aemula)
Pleroma castellense (T. castellensis)
Pleroma trinervium
Pleroma melanocalyx (T. melanocalyx)
87 Pleroma oreophilum (T. oreophila)
Pleroma mirabile (Tibouchinopsis mirabilis)
84
Pleroma blanchetianum (T. blanchetiana)
96 Pleroma pereirae (T. pereira)
Pleroma velutinum (T. velutina)
Pleroma arboreum (T. arborea)
Pleroma candolleanum (T. candolleana)
Pleroma mutabile (T. mutabilis)
Pleroma nodosum (T. nodosa)
Pleroma foveolatum (T. foveolata)
Pleroma sellowianum (T. sellowiana)
Pleroma fothergillae (T. fothergillae)
Pleroma vimineum (T. corymbosa)
72
Pleroma benthamianum (T. benthamiana)
97 Pleroma axillare (T. axillaris)
Pleroma
Pleroma cristatum (T. cristata)
Pleroma urceolare (T. urceolaris)
Pleroma clavatum (T. clavata)
Pleroma cryptadenum (T. cryptadena)
Pleroma radula (T. radula)
Pleroma manicatum (T. manicata)
Pleroma heteromallum (T. heteromalla)
Pleroma clidemiodes (T. clidemioides)
Pleroma penduliflorum
Pleroma cucullatum
Pleroma marinanum
Pleroma trichopodum (T. trichopoda)
Pleroma barnebyanum (T. barnebyana)
Pleroma oleifolium (Microlepis oleifolia)
Pleroma itatiaiae (T. itatiaiae)
Pleroma martiale (T. martialis)
78 Pleroma salviifolium (T. salviaefolia)
Pleroma fissinervium (T. fissinervia)
Pleroma estrellense (T. estrellensis)
88
Pleroma granulosum (T. granulosa)
Pleroma boudettii (T. boudetii)
Pleroma microphyllum (T. microphylla)
Pleroma cleistoflorum (Itatiaia cleistopetala)
Pleroma cinereum (T. cinerea)
Pleroma laevicaule (T. laevicaulis)
Pleroma frigidulum (T. frigidula)
Pleroma angustifolium (T. angustifolia)
Pleroma martiale (T. lindeniana)
Pleroma stenocarpum (T. stenocarpa)
Pleroma hospitum (T. hospita)
Pleroma macrochiton (T. macrochiton)
Pleroma cardinale (T. cardinalis)
Pleroma arenarium (T. arenaria)
Pleroma martiusianum (T. martiusiana)

Fig. 2. Maximum likelihood phylogenetic cladogram of Melastomateae with representatives of Tibouchina and allies, and outgroups (Rhexieae,
Microlicieae, Cambessedesia) derived from the combined data matrix (nrITS, accD-psaI, psbK-psbL). Double slash indictes a branch shortened
to one-quarter.

8
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

These species are characterized by leaves and hypanthia cov- in Central America to Bolivia and northern Brazil, in white-
ered by scale-like trichomes (except in T. striphnocalyx and sand savannas and open areas. The second group is formed
T. catharinae, see below). The species of this clade can be by the morphologically distinct species pair of T. catharinae
informally divided into three subgroups based on morphology and T. striphnocalyx; they both have penicillate hairs on the
and geographical distribution. The first is comprised of spe- hypanthia, branched intercalycine emergences, and single
cies assigned to T. sect. Tibouchina, with the flower buds sub- smooth cochleate seeds. The last group is comprised by mem-
tended by 2 or 4 bracteoles, the calyx covered by scales, and bers of T. sect. Barbigerae. This section is characterized by a
glabrous stamen filaments. Cogniaux (1891) recognized two long tuft of hairs on the appendages of the connective, and
species in T. sect. Tibouchina, but only T. aspera is now the filament may have a few trichomes (Cogniaux, 1885). Its
accepted (Jørgensen & al., 2014). After the Cogniaux’s study, species occur in shrub savannas and forests in llanos and Cer-
five additional species were described, all endemic to the Ven- rado; 13 of them are found in central Brazil (Guimarães
ezuelan Guayana highlands, at 1800–2000 m. Tibouchina & Woodgyer, 2016), with 4 others in Costa Rica, Panama,
aspera is found beyond the Guayana Shield, also occurring Colombia, Venezuela and Bolivia at elevations up to 1600 m.

Heterocentron breedlovei (Tibouchina breedlovei)


100 Heterocentron elegans
100
100 Heterocentron subtriplinervium
Heterocentron muricatum
98
Centradenia grandifolia
Centradenia floribunda
97 Centradenia inaequilateralis clade 1
Schwackaea cupheoides
100
100
Pilocosta campanensis
Pilocosta nubicola
77 Pilocosta oerstedii
Pilocosta nana
98 Bucquetia vernicosa
83
Bucquetia glutinosa
Castratella piloselloides
100
Chaetolepis cufodontisii
100 98 Chaetolepis microphylla
Chaetolepis anisandra
Monochaetum polyneuron clade 2
94 Monochaetum discolor
88 Monochaetum vulcanicum
81
Monochaetum bonplandii
Monochaetum tenellum
98 Monochaetum lineatum
74 Monochaetum uribei
96
100 Monochaetum meridense
Monochaetum humboldtianum
100 Andesanthus wurdackii (T. wurdackii)
Andesanthus inopinatus (T. inopinata)
99
83 Andesanthus gleasonianus (T. gleasoniana) Andesanthus
Andesanthus gleasonianus (T. gleasoniana)
100 Chaetogastra geitneriana (T. geitneriana)
Chaetogastra longifolia (T. longifolia)
Chaetogastra cf. weberbaueri (T. weberbaueri)
Chaetogastra hieracioides 1 (T. hieracioides)
99 Chaetogastra chamaecistus (T. chamaecistus)
Chaetogastra inconstans (T. ornata)
Chaetogastra gracilis (T. gracilis)
71 Chaetogastra minor (T. minor)
96
Chaetogastra hieracioides 2 (T. hieracioides)
Chaetogastra clinopodifolia (T. clinopodifolia)
Chaetogastra cerastifolia (T. cerastifolia)
Chaetogastra sebastianopolitana (T. sebastianopolitana)
Chaetogastra

Chaetogastra saxosa (T. saxosa)


96 Chaetogastra dimorphylla 1 (T. dimorphylla)
88 Chaetogastra decora (T. decora)
90
Chaetogastra pleromoides (T. pleromoides)
Chaetogastra dimorphylla 2 (T. dimorphylla)
89 Chaetogastra mollis 1 (T. mollis)
Chaetogastra grossa (T. grossa)
Chaetogastra mariae (T. mariae)
95 Chaetogastra laxa (T. laxa)
Chaetogastra pendula (T. pendula)
100 Chaetogastra solmsii (T. solmsii)
Chaetogastra sp. (Tibouchina sp.)
Chaetogastra ciliaris (T. ciliaris)
Chaetogastra aff. mollis 2 (T. mollis) clade 3
Chaetogastra montana (T. alpestris)
94 Chaetogastra confertiflora (T. confertiflora)
99 Chaetogastra citrina (T. citrina)
Chaetogastra gayana 5 (T. gayana)
Chaetogastra cf. gayana 1 (T. gayana)
Chaetogastra gayana 4 (T. gayana)
98
Chaetogastra gayana 3 (T. gayana)
Chaetogastra gayana 2 (T. gayana)
Brachyotum ledifolium 2
92
Chaetogastra aff. scabriuscula (T. scabriuscula)
88 Chaetogastra naudiniana (T. naudiniana)
Chaetogastra latibracteolata (T. latibracteolata)
Brachyotum benthamianum
Brachyotum incrassatum
100 Brachyotum lindenii
Brachyotum ledifolium 2
Brachyotum fictum
Brachyotum

Brachyotum confertum
76
Brachyotum harlingii
Brachyotum fraternum
Tibouchina brevisepala
Brachyotum intermedium
Brachyotum rostratum 1
Brachyotum rostratum 2
Brachyotum angustifolium
Brachyotum radula
Brachyotum aff. maximowiczii
Brachyotum aff. ledifolium 1
Brachyotum cogniauxii
Brachyotum coronatum
Tibouchina octopetala
82 Brachyotum microdon
Tibouchina calycina
Tibouchina bicolor

0.04

Fig. 2. Continued.

9
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

All of these species will remain in Tibouchina under this new and with bilobed ventral appendages. Some species, mostly
scheme. from T. sect. Pseudoterolepis, do have dimorphic stamens
Pleroma includes species from eastern Brazil, in the Mata with a prolonged pedoconnective, but in this case at least
Atlântica and Cerrado biomes, rarely in Caatinga (although one of the whorls is always yellow. The oldest name available
few species occur in the Andes: T. cryptadena, T. lindeniana, for this group of species currently in Tibouchina is Chaetogas-
T. ochypetala and T. stenocarpa.), and contains species tra, and we will refer to it with this name herafter. While Bra-
assigned to T. sect. Diplostegia, T. sect. Involucrales and T. chyotum, with its tubular, nectar-producing flowers with
sect. Pleroma, as well as the genera Itatiaia, Microlepis, Svi- anthers with reduced pedoconnectives is easily diagnosable,
tramia and Tibouchinopsis. Although no character exclusively this genus is not resolved as monophyletic as some Andean
supports this group, the great majority of the species have species of Tibouchina are nested within it. Moreover, unfortu-
pink, lilac or purple anthers, often with a well-developed pedo- nately, the ML and BI results disagree on the relative position
connective and filaments covered with trichomes or glands, of the expanded Brachyotum clade, being recovered within
caducous calyx lobes during fruit development, pentamerous Chaetogastra in the ML analyses and as sister to Chaetogastra
flowers, and they lack scales on the leaves and hypanthia. in the BI analyses. It should be noted that in both cases, the
Microlepis had been segregated from Lasiandra (and later support for some of the basal nodes is rather low, allowing
Tibouchina) because of its filiform anther appendages for alternative topologies to be recovered. In fact, under some
(Miquel, 1840; Triana, 1872; Cogniaux, 1885). However, sampling schemes, ML analyses recovered a topology similar
the anther appendages seem to be similar to those present in to that found here in the BI analyses (Meyer, 2016). Therefore,
other species of Pleroma, but with an extreme level of devel- given the lack of support for either topology, and the fact that
opment. The species of Microlepis have already been trans- the BI analyses recovered Brachyotum and Chaetogastra as
ferred to Pleroma by Romero & Versiane (2014). Itatiaia separate and sister, we consider it more prudent to recognize
had been distinguished by its cleistogamous tetramerous Brachyotum as separate from Chaetogastra. It should be noted
flowers with anthers that lack a pedoconnective, but that have that a few species of Andean Tibouchina that lack tubular
a pair of long ventral appendages (Ule, 1895, 1896a,b). This corollas are currently resolved within Brachyotum. We have
genus has already been reviewed by da Silva & al. (2014), elected for the time being to recognize them as Tibouchina
and the appropriate transfer to Pleroma has been made. Tibou- incertae sedis, and not to transfer them to either Chaetogastra
chinopsis was segregated from Tibouchina mostly based on or Brachyotum for the same reasons explained above.
the glabrous ovary apex, but it also differs by its non-cochleate Two additional species currently recognized in Tibou-
seeds and persistent sepals. However, the stamen morphology is china are not resolved in any of these four groups
similar to that of most members of the Pleroma clade. Lastly, this (i.e., Andesanthus, Chaetogastra + Brachyotum, Pleroma or
clade also includes Svitramia. This is perhaps the group most Tibouchina). As already mentioned, T. breedlovei is resolved
difficult to fit into an expanded Pleroma because its anthers lack as sister to Heterocentron, and this placement is entirely con-
pedoconnectives, and it has a persistent calyx. However, it is sistent with its morphology. In fact, at the time of describing
restricted to eastern Brazil, and the anthers are usually pink or T. breedlovei, Wurdack (1967) already mentioned that it and
whitish, and not yellow. Pleroma is the oldest available name T. chiapensis may also be placed in Heterocentron, and the
for this clade, and indeed many of the species up until now rec- appropriate taxonomic change is made below. The second spe-
ognized in Tibouchina were described as Pleroma and later cies is T. cubense, which, although not included in this study,
transferred to Tibouchina by Cogniaux (1885, 1891). has been recovered in other analyses (Michelangeli & al.,
The species of Tibouchina sect. Lepidotae are all resolved unpub.) as sister to Pterogastra and nested within Pterolepis.
together as a monophyletic group. They are characterized by This species is characterized by anthers that lack a pedocon-
the presence of lepidote trichomes on the hypanthium. Addi- nective, but its placement has been problematic, as it has also
tionally, all members of this group have glabrous stamens, been placed in Arthrostemma, Chaetogastra, Pleroma and
usually isomorphic and with yellow anthers. Some species even Chaetolepis. However, T. cubense lacks the characteris-
have dimorphic stamens, but in that case the larger anthers tic anthers of both Pterolepis and Pterogastra, the intercaly-
are yellow, and the smaller anthers are pink or red. Most of cine emergences of Pterolepis, and the winged hypanthia of
the species in this clade are found in the Andes, with two of Pterogastra. At this point, it is perhaps most appropriate to
them extending into the mountains of Panama and Costa Rica. maintain it in Tibouchina as incertae sedis.
There is no available generic name for this clade, and we are The generic circumscriptions of Bucquetia, Castratella,
here naming it Andesanthus (see below). Centradenia, Chaetolepis, Monochaetum, Pilocosta, and
The fourth clade with species of Tibouchina contains all Schwackaea are not changed by our results. Two species are
species of Brachyotum, plus species from T. sect. Dio- transferred to Heterocentron, but given their morphology, this
tanthera, T. sect. Octomeris, T. sect. Pseudoterolepis and T. does not alter its diagnostic characteristics.
sect. Simplicicaulis. This clade is characterized by the absence Below we provide a key to the genera of the Tibouchina clade
of lepidote trichomes and anthers, at least in one set, yellow. of Melastomateae, along with lists of all species assigned to
Most of the species in this group have isomorphic or weakly Andesanthus, Brachyotum, Chaetogastra, Pleroma, and Tibou-
dimorphic stamens with the connective shortly prolonged china, including appropriate new combinations and new names.

10
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

Key to Tibouchina and related genera anthers pink or red; petal color commonly changing with
age .............................................................. Andesanthus
1. Stamens isomorphic and without pedoconnectives or 12. Anthers lilac or purple; petal color not changing with
well-developed appendages ............................................ 2 age .................................................................Tibouchina
1. Stamens almost always dimorphic and with well- 13. Stamens extremely dimorphic in color and shape; large
developed pedoconnectives, ventral appendages bilobed stamens with connective prolonged at least 2/3 the length
at least in the larger stamens ..........................................6 of the anther, ventrally with a pair of filiform appendages;
2. Petals never spreading, remaining in a pseudo-tube that small stamens with connective not prolonged and with
falls off as a unit after anthesis .................... Brachyotum short, thick ventral lobes ..............................Desmoscelis
2. Petals spreading, arranged perpendicular or inclined 13. Stamens similar in color and slightly dimorphic in shape;
to hypanthium and falling independently after large stamens with the connective prolonged no more
anthesis...........................................................................3 than 1/3 the length of the anther; ventral connective
3. Rosulate herbs, small (3–5 cm in height without the inflo- appendages usually present and similar in the two cycles
rescence); villous plants in all structures; leaf blade of stamens ....................................................................14
1.5–4 cm long ................................................Castratella 14. Anthers purple to lilac, rarely cream or white, with well-
3. Woody herbs, shrubs or small trees ...............................4 developed pedoconnectives, often filaments pubescent
4. Flower pentamerous ...........................................Pleroma and anther connectives with gland-tipped hairs; calyx
4. Flower tetramerous.........................................................5 caducous in fruit.................................................Pleroma
5. Stems and often the abaxial surface of the leaves with 14. Anthers at least partially yellow and often with reduced
barbellate or simple trichomes; petals yellow or pink, pedoconnectives, filament and anther connectives gla-
rarely magenta; plants sometimes glutinous; seeds brous; calyx persistent in fruit ....................Chaetogastra
cochleate....................................................... Chaetolepis
5. Stems and the abaxial surface of the leaves glabrous; Generic descriptions and species lists
petals magenta; plants glutinous; seeds straight, not
cochleate .........................................................Bucquetia Andesanthus P.J.F.Guim. & Michelang., gen. nov. ≡ Tibou-
6. Hypanthium quadrate or conspicuously winged or china sect. Lepidotae Cogn. in Martius, Fl. Bras. 14(3):
ribbed .............................................................................7 372. 1885 – Type: Andesanthus lepidotus (Bonpl.)
6. Hypanthium more or less terete, not conspicuously P.J.F.Guim. & Michelang. (≡ Rhexia lepidota Bonpl. ≡
angled, winged or ribbed................................................8 Tibouchina lepidota (Bonpl.) Baillon).
7. Subshrubs or perennial herbs; hypanthium square with 4 Species of Andesanthus can be recognized by their lepi-
prominent angles or ribs; flowers pedicellate ....Pilocosta dote indumentum; free bracts that envelop the floral buds; gla-
7. Annual herbs; hypanthium linear-oblong with 8 conspicu- brous stamens; yellow anthers on isomorphic stamens or
ous ribs; flowers subsessile ......................... Schwackaea larger anthers yellow and smaller anthers pink or red in dimor-
8. Leaves mostly penninerved, secondary veins diverging phic stamens. The indumentum consists of flattened scales
from the median vein alternately and irregularly similar to that of Tibouchina, but the stamens in Tibouchina
..................................................................Heterocentron are slightly dimorphic with purple anthers, glabrous or with
8. Leaves 3–7-nerved, all secondary veins originating at the hairs over the connective appendages and filaments.
base of the leaf blade or near it ......................................9 Trees or small trees 5–20 m at maturity or shrubs 1–3 m
9. Leaves obviously dimorphic in each pair, the base tall, with trichomes flattened scales of varying sizes and
oblique; seeds ellipsoid, pyriform, or clavi-form shapes, scabrous, trichomes bulla-based or with pustulate
..................................................................... Centradenia base. Leaves opposite, decussate, blades of a pair essentially
9. Leaves isomorphic or nearly so in each pair, the base sym- equal in size, symmetrical, petiolate, coriaceous, above (adax-
metrical; seeds cochleate, reniform or oblong-arched .... 10 ial foliar surface) flattened scales, scabrous bulla-based tri-
10. Anther connective with well-developed dorsal append- chomes or scabrous flattened and appressed trichomes,
ages, ventral appendages absent; seeds smooth or incon- below (abaxial foliar surface) scales, minute dendritic or
spicuously rugulose..................................Monochaetum smooth trichomes (papillae-like trichomes), minute glandular
10. Anther connective without dorsal appendages, ventral hairs, 3–7-nerved. Inflorescence an anthotelic terminal fron-
appendages present (often bilobed or as spurs); seeds dobracteose panicle or some modification of this structure in
tuberculated, ribbed or tuberculate-ribbed at least in the which the main and at least some lateral axes have nodes distal
anti-raphal side .............................................................11 to first pair of leaves on a branch axis; bracteoles 2, early
11. Hypanthium and stem internodes covered by scales, or rarely deciduous, commonly enveloping young floral buds. Flowers
hypanthia with prominent long-branched trichomes...........12 perigynous, pedicellate; hypanthium campanulate to urceo-
11. Hypanthium and stem internodes covered by simple late, mostly copiously pubescent externally, with 10 slight lon-
trichomes ......................................................................13 gitudinal vascular ribs. Calyx not calyptrate in bud, with
12. Anthers yellow, stamens isomorphic, or if stamens dimor- conspicuous lobes, persistent, lobes 5, oblong to triangular.
phic then with the larger anthers yellow and smaller Corolla patent; petals 5, free and spreading, pink, purple,

11
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

magenta, white, or initially magenta or rose-red but fading to 4. Andesanthus inopinatus (Wurdack) P.J.F.Guim. & Miche-
uniform lavender with age, essentially obovate, glabrous. Sta- lang., comb. nov. ≡ Tibouchina inopinata Wurdack in
mens 10, isomorphic, or weakly to strongly dimorphic. Phytologia 16(3): 169–170. 1967 – Holotype: COSTA
Anthers 2-celled, ventrally channeled, subulate, having ven- RICA. Cartago, on the wooded slopes of Cordillera de
trally inclined apical pores. Connectives glabrous, variously Talamanca, 12 miles southeast of Cartago, elev. 1600 m,
prolonged and modified ventrally at filament insertion into 15 Jul 1962, Webster, G.L.; Miller, K. & Miller, L. 12361
horizontal or upturned, bilobed glabrous ventral appendages. (US barcode 00120064!; isotypes: DUKE barcode
Filaments glabrous. Ovary superior, 5-celled, setose or bristly 10000288 [online image!], F barcode V0063972F [online
apically, axile placentation; style glabrous, hooked or image!], GH barcode 00073252 [online image!], LL bar-
incurved apically, typically declined to side of flower oppos- code 00372244 [online image!], MICH barcode 1111898
ing stamens. Mature capsule dry, semiwoody, loculicidal, [online image!], U barcode U 0004178 [online image!]).
not angled. Seeds numerous, cochleate to elongate-cochleate
and tuberculate with a terminal hilum. 5. Andesanthus lepidotus (Bonpl.) P.J.F.Guim. & Miche-
Etymology: This new genus is named for the Andes, the lang., comb. nov. ≡ Rhexia lepidota Bonpl. in Humboldt
region where the majority of the species occur. According to & Bonpland, Monogr. Melast. 2: 38, pl. 15. 1808 ≡ Chae-
Art. 62.2(c) of the ICN (Turland & al., 2018) names ending togastra lepidota (Bonpl.) DC., Prodr. 3: 134. 1828 ≡
in -anthus are treated as masculine in accordance with tradition. Pleroma lepidotum Triana in Trans. Linn. Soc. London,
Andesanthus has nine species and one variety with a Bot. 28(1): 44, t. 3. 1872 (“1871”) ≡ Tibouchina lepidota
mostly Andean distribution from western Venezuela in Mé- (Bonpl.) Baillon in Adansonia 12: 74. 1877 – Holotype:
rida to southcentral Peru in Pasco. Two extra-Andean species COLOMBIA or PERU. Bonpland, A.J.A. 3626
are restricted to floristically related areas in mountains of (P barcode P00708966!; isotype: P barcode P00136473!).
southern Central America in Costa Rica and Panama. The
center of diversity for Andesanthus is in Colombia, where 5a. Andesanthus lepidotus var. congestiflorus (Bonpl.) P.J.F.
six of the nine species are found. These species occupy the Guim. & Michelang., comb. nov. ≡ Tibouchina lepidota
cool montane forests, growing along road cuts or forest mar- var. congestiflora (Bonpl.) Cogn. in Bot. Jahrb. Syst. 8
gins (Todzia & Almeda, 1991). (1): 17. 1886 – Lectotype (designated by Todzia
& Almeda in Proc. Calif. Acad. Sci., ser. 4, 47(6): 194.
Species list 1991): ECUADOR. Pichincha: near S. Rorencio at Rio
Pilaton, 1800 m, Cordillera Central, 13 Jan 1881, Leh-
1. Andesanthus aristeguietae (Wurdack) P.J.F.Guim. & mann, F.C. 479a (BR barcode BR0000005223428!;
Michelang., comb. nov. ≡ Tibouchina aristeguietae isolectotype: B [probably destroyed; F negative
Wurdack in Bol. Soc. Venez. Ci. Nat. 20: 360. 1959 – F0BN016760!]).
Holotype: VENEZUELA. Mérida: from “arriba de Sto.
Domingo, alt. más o menos 2600 m”, Aug 1958, Ariste- 6. Andesanthus narinoensis (Wurdack) P.J.F.Guim.
guieta, L. 3263 (NY barcode 00245752; isotypes: US bar- & Michelang., comb. nov. ≡ Tibouchina narinoensis
codes 00120037! & 00946241!, VEN barcode 43276 Wurdack in Phytologia 35(3): 241–242. 1977 – Holo-
[online image!]). type: COLOMBIA. Dep. Nariño: between Ricaurte and
Diviso along the Pasto–Tumaco road, 11 Jun 1950, Espi-
2. Andesanthus elegantulus (Todzia & Almeda) P.J.F.Guim. nosa, R. 2954 (NY barcode 00245790!).
& Michelang., comb. nov. ≡ Tibouchina elegantula Todzia
& Almeda in Proc. Calif. Acad. Sci., ser. 4, 47(6): 187–189, 7. Andesanthus paleaceus (Triana ex Mart.) P.J.F.Guim.
fig. 1E–F, 8. 1991 – Holotype: COLOMBIA. Cauca: Parque & Michelang., comb. nov. ≡ Pleroma lepidotum var.
Nac. Munchique, km 50–55 along road above Uribe, paleaceum Triana in Trans. Linn. Soc. London, Bot.
1875–2256 m, 25 Apr 1979, Luteyn, J.L.; Lebrón-Luteyn, 28(1): 44. 1872 (“1871”) ≡ Tibouchina paleacea (Triana
M. & Morales, G. 7488 (NY barcode 00245763!; isotypes: ex Mart.) Cogn. in Martius, Fl. Bras. 14(3): 373–374.
AAU ImageID 3505 [online image!], CAS barcode 0004851 1885 – Lectotype (designated here): COLOMBIA.
[online image!], CM, COL barcodes COL000003424 Antioquia: Rio Negro, 2300 m, s.d., Triana, J.J. s.n.
& COL000003425 [online images!], MO barcode MO- (K barcode K000329134!; isolectotypes: BR barcode
313920 [online image!], US barcode 00345026!). BR0000005523597!, G-DC barcode G00318973 [online
image!], NY barcode 00245678!, P barcode
3. Andesanthus gleasonianus (Wurdack) P.J.F.Guim. & P00708681!).
Michelang., comb. nov. ≡ Tibouchina lepidota var. Other syntype: COLOMBIA. Nova Granada. Province
intermedia Gleason in Bull. Torrey Club 52: 328. 1925 Mariquita: in mont Quindiu, 1580 m, 1843, Linden,
≡ Tibouchina gleasoniana Wurdack in Phytologia 35(3): J.J. 1057 (BR barcode BR0000005523924!, F barcode
241. 1977 – Holotype: ECUADOR. Forest of the Andes, V0063927F [fragment] [online image!], MPU barcode
s.d., Jameson, W. 87 (NY barcode 00245779!). MPU013263 [online image!], P barcode P00708795!).

12
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

8. Andesanthus silvestris (Todzia & Almeda) P.J.F.Guim. off as a unit after anthesis; pollination is by hummingbirds.
& Michelang., comb. nov. ≡ Tibouchina silvestris Todzia The stamens, which are hidden inside the corolla tube, are iso-
& Almeda in Proc. Calif. Acad. Sci., ser. 4, 47(6): morphic with the connectives not prolonged or shortly pro-
200–202, fig. 5E–F, 6A–B, 11. 1991 – Holotype: longed with a bilobed ventral appendage. The leaves are
COLOMBIA. Valle: Anchicayá, 200 m, May 1969, generally of reduced size up to 4 cm long, with blades varying
Uribe, U.L. & Cortés, J.C. 6280 (COL barcode in texture from chartaceous to thick-coriaceous or thick-brit-
COL000003430 [online image!]; isotype: US barcode tle, providing another typical character.
00345027!).
Species list
9. Andesanthus wurdackii (Almeda & Todzia) P.J.F.Guim.
& Michelang., comb. nov. ≡ Tibouchina wurdackii 1. Brachyotum alpinum Cogn. in Candolle & Candolle,
Almeda & Todzia in Proc. Calif. Acad. Sci., ser. Monogr. Phan. 7: 167. 1891 – Lectotype (designated
4, 47(6): 202–204, fig. 6C–F, 12. 1991 – Holotype: PAN- here): ECUADOR. Prov. Chimborazo: Quito, mont.
AMA. Coclé: forested slopes above El Copé along aban- Chimborazo, at el. 12,000 ft., 1856, Jameson, W. 193
doned road leading to continental divide, 700–850 m, (G-DC barcode G00318699 [online image!]; isolecto-
08 38′N, 80 38′W, 24 Jan 1989, Almeda, F.; de types: BR, G barcodes G00353527 & G00353528 [online
Nevers, G. & McPherson, G. 6380 (CAS barcode images!], G-DC barcode G00318699 [online image!], K
0004864 [online image!]; isotypes: DUKE barcode barcode K000329226!, LE!, NY barcode 00221450!,
10000287 [online image!], MO barcode MO-129905 TCD barcode TCD0000747 [online image!], W).
[online image!], NY barcode 00022578!, PMA barcode Other syntypes: ECUADOR. 1860, Fraser s.n.
853 [online image!], TEX barcode 00372246 [online (F barcode V0063416F [fragment, F negative
image!], US barcode 00406144!). F0BN025862!] [online image!], G-DC barcodes
G00318712 & G00318684 [online images!], US barcode
Brachyotum (DC.) Triana in Bentham & Hooker, Gen. Pl. 1: 00120108 [online image!]).
743. 1867 ≡ Arthrostemma sect. Brachyotum DC., Prodr.
3: 136. 1828 – Type (designated by Wurdack in Mem. 2. Brachyotum andreanum Cogn. in Bull. Acad. Roy. Sci. Bel-
New York Bot. Gard. 8(4): 357. 1953): Brachyotum quin- gique, sér. 3, 14(7): 938–940. 1887 – Holotype: ECUA-
quenerve (Ruiz & Pav.) Triana (≡ Rhexia quinquenervis DOR. In Andibus centralibus aequatorensibus, alt.
Ruiz & Pav.). 3300 m, 1876, André, É.-F. 387 (K barcode K000329188!;
Shrubs or treelets covered with smooth to very shaggy tri- isotype: BR barcode BR0000005179725 [online image!]).
chomes, sometimes gland-tipped. Leaves opposite, decussate,
blades of a pair essentially equal in size, petiolate, membra- 3. Brachyotum angustifolium Wurdack in Mem. New York
naceous, chartaceous to coriaceous, variously pubescent to Bot. Gard. 8(4): 401–402. 1953 – Holotype: PERU.
nearly glabrous, or tuberculate, 3–5(–7)-nerved. Flowers pen- Dep. Amazonas: prov. Chachapoyas, Bajasan, 1835,
dulous, solitary or in 2–3-flowered dichasia, flowers are some- Mathews, A. 1253 (K barcode K000329236!; isotypes:
times aggregated into panicles or corymbs; bracteoles 1–3 E barcodes E00285771 & E00285772 [online images!]).
pairs, free, concave, variously pubescent, early deciduous or
persistent, smaller than the mature hypanthium or replaced 4. Brachyotum azuayense Wurdack in Phytologia 35(3):
by persistent undifferentiated leaves. Calyx lobes 4–5, persis- 242–243. 1977 – Holotype: ECUADOR. Prov. Azuay:
tent. Petals 4–5, dark purple to carmine or white with red mar- dry scrub near km 91 on Pan American Highway north
gins to greenish-white or yellowish, free but connivent and of Loja, 03 25′S, 79 10′W, elev. 2900 m, 5 May 1973,
imbricate in a cylindrical or campanulate tube, glabrous or Holm-Nielsen, L.; Jeppesen, S.; Lojtanant, B. & Oell-
in a few instances pubescent. Stamens 8–10, isomorphic. gaard, B. 5034 (US barcode 00120109 [online image!];
Anthers 2-celled, ventrally channeled, subulate, slightly arcu- isotypes: AAU ImageID 3490 [online image!], F barcode
ate and with lanceolate apical pore. Connectives glabrous, not V0093793F [online image!], GB barcode GB-0048125
or shortly prolonged, with bilobed glabrous ventral append- [online image!], MO barcode MO-288024 [online
ages. Filaments glabrous. Ovary superior, pubescent apically, image!], NY barcode 00221451!).
eglandular or with gland-tipped hairs, 4–5-celled with axile
placentation; style glabrous (except in B. campii), straight. 5. Brachyotum barbeyanum Cogn. in Candolle & Candolle,
Capsule dry, semiwoody, loculicidal. Seeds numerous, Monogr. Phan. 7: 158. 1891 – Holotype: PERU. [Dep.
cochleate, pitted with a terminal hilum. Amazonas]: Chachapoyas, 1862, Mathews, A. s.n. (G;
Brachyotum has 58 taxa (55 species, 2 varieties), which isotypes: BR, NY barcode 00221452!).
occur in páramo and subpáramo zones from Colombia to north-
ern Argentina; the species are mostly quite local distribution. 6. Brachyotum benthamianum Triana in Trans. Linn. Soc.
The genus is characterized by the presence of tubular London, Bot. 28(1): 48. 1872 (“1871”) – Lectotype (des-
drooping flowers with 4 to 5 petals that never spread and fall ignated by Wurdack in Mem. New York Bot. Gard. 8(4):

13
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

363. 1953): ECUADOR. s.d., Hartweg, K.T. 737 (designated here): ECUADOR. Prov. Loja: “entre Loxa
(K barcode K000329185!; isolectotypes: B [probably et Malacatos, à une élévation de 2000 mètres”, s.d., Bon-
destroyed; F negative F0BN016708!], BR barcode pland, A.J.A. & Humboldt, F.W.H.A. von 3378 (P barcode
BR0000005178940 [online image!], E barcode P00136399!; isolectotypes: COL barcode COL0000
E00285793 [online image!], F barcode V0063417F [frag- 03193 [online image!], P barcodes P00136400! &
ment] [online image!], G barcode G00353529, K barcode P00136401!).
K000329186!, LE!, NY barcode 00221454!, P barcodes = Brachyotum campylanthum Triana in Trans. Linn. Soc.
P02274342! & P02274343!, W). London, Bot. 28(1): 48. 1872 (“1871”) – Holotype:
ECUADOR. s.d., Jameson s.n. (K barcode K000
7. Brachyotum campanulare (Bonpl.) Triana in Trans. Linn. 329229!).
Soc. London, Bot. 28(1): 48. 1872 (“1871”) ≡ Rhexia
campanularis Bonpl. in Humboldt & Bonpland, Monogr. 12. Brachyotum coronatum (Triana) Wurdack in Phytologia
Melast. 2: 35, pl. 14. 1808 ≡ Arthrostemma campanulare 11(6): 379. 1965 ≡ Pleroma coronatum Triana in Trans.
(Bonpl.) DC., Prodr. 3: 136. 1828 ≡ Chaetogastra campa- Linn. Soc. Bot. 28(1): 41. 1872 (“1871”) ≡ Tibouchina
nularis (Bonpl.) Naudin in Ann. Sci. Nat., Bot., sér. coronata (Triana) Cogn. in Martius, Fl. Bras. 14(3):
3, 14(2): 130. 1850 – Holotype: PERU. “in Peruviae frigi- 307. 1885 – Holotype: PERU. [Dep. Amazonas]: Cha-
dis, juxta Loxam près de 2000 mètres”, s.d., Bonpland, chapoyas, 1840, Mathews, A. 3211 (K barcode
A.J.A. & Humboldt, F.W.H.A. von 3361 (P barcode K000329240!; isotypes: BR barcode BR000000517
P00136392!; isotypes: F barcode V0063943F [fragment] 8612 [fragment] [online image!], BM barcode BM00
[online image!], P barcode P00136393!). 0953910, G [F negative F0BN026104!], G-DC barcode
G00318885!, K barcode K000329241!).
8. Brachyotum campii Wurdack in Mem. New York Bot. = Brachyotum racemosum Cogn. in Bot. Jahrb. Syst. 42: 132.
Gard. 8(4): 364–365. 1953 – Holotype: ECUADOR. 1908 – Lectotype (designated by Wurdack in Mem.
Azuay-Oriente border, Páramo del Castillo and surround- New York Bot. Gard. 8(4): 369–370. 1953): PERU.
ing forested areas, crest of the eastern Cordillera on the Dep. Cajamarca: prov. Chota; in montibus ad occidentem
trail between Sevilla de Oro and Mendez, alt. a Huambos versus, 3100–3200 m a.s.l., 28 May 1904,
11,000–11,350 ft., 17 Dec 1944, Camp E-1629 Weberbauer, A. 4170 (BR barcode BR0000005184200
(NY barcode 00221456!). [online image!]; isolectotypes: B [probably destroyed; F
negative F0BN016714!], G barcode G00353530 [online
9. Brachyotum cernuum (Bonpl.) Triana in Trans. Linn. Soc. image!], MOL barcode 00003176 [online image!]).
London, Bot. 28(1): 48. 1872 (“1871”) ≡ Rhexia cernua
Bonpl. in Humboldt & Bonpland, Monogr. Melast. 2: 13. Brachyotum cutervoanum Wurdack in Phytologia 14(5):
32, pl. 13. 1808 ≡ Osbeckia cernua (Bonpl.) Spreng., 257–258. 1967 – Holotype: PERU. Dep. Cajamarca:
Syst. Veg. 2: 312. 1825 ≡ Chaetogastra cernua (Bonpl.) prov. Cutervo, La Pucarilla along the road between Socota
DC., Prodr. 3: 135. 1828 – Lectotype (designated here): and San Andrés, elev. 2420 m, 24 May 1965,
COLOMBIA. Dep. Cauca: “(Monte de Puracé), juxta Sagástegui, A. 5384 (US barcode 00120112 [online
urbem Popayan plus de 2000 mètres”, s.d., Bonpland, image!]; isotype: TRP).
A.J.A. & Humboldt, F.W.H.A. von s.n. (P barcode
P00136394!; isolectotypes: F barcode V0040203F [frag- 14. Brachyotum ecuadorense Wurdack in Mem. New York
ment] [online image!], P barcodes P00136395!, Bot. Gard. 8(4): 393–394. 1953 – Holotype: ECUADOR.
P00136396!, P00136397! & P00136398!, RB barcode Azuay-Oriente border, eastern cordillera between Oña
00541363!). and rio Yacuambi, crest, alt. 10,000–11,200 ft., 10–19
Sep 1945, Prieto, F. P-307 (NY barcode 00221457!; iso-
10. Brachyotum cogniauxii Wurdack in Mem. New York types: F barcode V0063418F [online image!], K barcode
Bot. Gard. 8(4): 392. 1953 – Holotype: PERU. Dep. K000329231!, US barcode 00120113 [online image!]).
Amazonas: “stony jalca, alt. 3500 m., above Colcamar”,
24–26 Jun 1948, Pennell, F.W. 15660 (PH barcode 15. Brachyotum fictum Wurdack in Mem. New York Bot.
00008041 [online image!]; isotype: USM barcode Gard. 8(4): 394. 1953 – Holotype: ECUADOR. Azuay-
USM000515 [online image!]). Oriente border, Páramo del Castillo and surrounding for-
ested areas, crest of the eastern cordillera on the trail
11. Brachyotum confertum (Bonpl.) Triana in Trans. Linn. between Sevilla de Oro and Mendez, alt. 11000–
Soc. London, Bot. 28(1): 48. 1872 (“1871”) ≡ Rhexia 11300 ft., 21 Aug 1945, Camp, W.H. E-4852 (NY bar-
conferta Bonpl. in Humboldt & Bonpland, Monogr. Me- code 00221458!; isotypes: F barcode V0063419F [online
last. 2: 53, pl. 20. 1808 ≡ Chaetogastra conferta image!], GH barcode 00071969 [online image!], K
(Bonpl.) DC., Prodr. 3: 135. 1828 ≡ Bolina conferta barcode K000329232!, US barcode 00120114 [online
(Bonpl.) Raf. in Sylva Tellur.: 101. 1838 – Lectotype image!]).

14
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

16. Brachyotum figueroae J.F.Macbr. in Publ. Field Mus. [both fragments] [online images!], G-DC barcode
Nat. Hist., Bot. Ser. 4(7): 173–174. 1929 – Holotype: G00318669 [online image!], GOET barcode
PERU. Dep. Ancash: “Catuc” 25 km east of Huarás, GOET007886 [online image!], K barcode K000
4 Oct 1922, Macbride, J.F. & Featherstone, W. 2504 329221!, LE!, NY barcode 00221462!, P barcodes
(F barcode V0040080F [online image!]; isotypes: G, P02274337! & P02274338!, S Nos. S05-3143
NY barcode 00221459!). & S09-13104 [online images!], W, TCD barcode
TCD0000749 [online image!]).
17. Brachyotum fraternum Wurdack in Mem. New York
Bot. Gard. 8(4): 375–376. 1953 – Holotype: ECUADOR. 21. Brachyotum harlingii Wurdack in Phytologia 29(2):
Azuay-Oriente border, Páramo del Castillo and surround- 140–141. 1974 – Holotype: ECUADOR. Prov. Morona-
ing forested areas, crest of the eastern cordillera on the Santiago: Campamento San Miguel along Sigsig-
trail between Sevilla de Oro and Mendez, alt. Gualaquiza road, elev. 3000 m, 8 Apr 1968,
11,000–11,300 ft., 21 Aug 1945, Camp, W.H. E-4871 Harling, G.; Storm, G. & Ström, B. 8038 (GB barcode
(NY barcode 00221460!; isotypes: F barcode GB-0048126 [online image!]).
V0063420F [online image!], G barcode G00353532
[online image!], GH barcode 00071970 [online image!], 22. Brachyotum huancavelicae Wurdack in Mem. New York
K barcode K000329199!, MO barcode MO-288052 Bot. Gard. 8(4): 381–382. 1953 – Holotype: PERU. Dep.
[online image!], P barcode P02274341!, US barcode Huancavelica: prov. Tayacaja, quebrada south of Salca-
00120115 [online image!], U barcode U 0003965 [online bamba in shrubwood, sandy loam, 8 Jan 1939, Stork
image!]). & Horton 10295 (F barcode V0040085F!; isotypes: G,
K barcode K000684620!, UC barcode UC656808 [online
18. Brachyotum gleasonii Wurdack in Mem. New York Bot. image!]).
Gard. 8(4): 370–371. 1953 – Holotype: ECUADOR.
Chimborazo-Cañar border (western escarpment), near 23. Brachyotum incrassatum E.Cotton in Nordic J. Bot.
Pimo, alt. 10,200–10,400 ft., 9 Jul 1945, Camp, W.H. 20(2): 185–187, fig. 4. 2000 – Holotype: ECUADOR.
E-4118 (NY barcode 00221461!; isotypes: F barcode Loja: 1 km S of Yangana, track to Sierra Toledo, km
V0063421F [online image!], G barcode G00353533 19.5, 04 23′S, 79 06′W, 3325 m, 14 Jan 1998, Lewis,
[online image!], GH barcode 00071971 [online image!], G.P. & Hughes, C.E. 3782 (AAU ImageID 3496 [online
K barcode K000329195 [online image!], MO barcode image!]; isotypes: K barcode K000329242 [online
MO-288050 [online image!], P barcode P02274340!, U image!], LOJA barcode 213311 [online image!], QCNE
barcode U 0003966 [online image!], UC barcode barcode QCNE340 [online image!]).
UC951481 [online image!], US barcode 00120116
[online image!]). 24. Brachyotum intermedium Wurdack in Mem. New York
Bot. Gard. 8(4): 367. 1953 – Holotype: PERU. Dep.
19. Brachyotum gracilescens Triana in Trans. Linn. Soc. Amazonas: Chachapoyas, s.d., Mathews, A. 3210
London, Bot. 28(1): 49. 1872 (“1871”) – Holotype: (K barcode K000329191!).
ECUADOR. Prov. Tungurahua: “in silvis montis Tungur-
ahua alt. 10,000 ped.”, Nov 1859, Spruce, R. 6084 25. Brachyotum jamesonii Triana in Trans. Linn. Soc.
(K barcode K000006134!; isotypes: B [probably London, Bot. 28(1): 49. 1872 (“1871”) – Holotype:
destroyed; F negative F0BN016710!], BR barcode ECUADOR. Southeastern, prov. Chimborazo near prov.
BR0000005179879 [online image!], C barcode Cañar border, in m. Azuay, loco Runa-rupashca, 12,000
C10014526 [online image!], E barcode E00285776 p., Aug 1859, Spruce, R. 6031 (K barcode
[online image!], G-DC barcode G00318689 [online K000006132!; isotypes: BM barcode BM000778236
image!], GH barcode 00071972 [online image!], GOET [online image!], BR barcode BR0000005177882 [online
barcode GOET007885 [online image!], K barcode image!], E barcode E00285774 [online image!], G-DC
K000006135!, LE!, MPU barcode MPU013842 [online barcode G00318668 [online image!], GH barcode
image!], P barcode P02274339!, S No. S05-3142 [online 00071973 [online image!], GOET barcode GOET007887
image!], TCD barcode TCD0005156 [online image!], W). [online image!], K barcode K000006133!, LE!, NY bar-
code 00221463!, P barcode P02274336!, TCD barcode
20. Brachyotum grisebachii Cogn. in Candolle & Candolle, TCD0000748 [online image!], W).
Monogr. Phan. 7: 153. 1891 – Lectotype (designated by
Wurdack in Mem. New York Bot. Gard. 8(4): 380. 26. Brachyotum johannesjulii E.Cotton in Nordic J. Bot.
1953): PERU. Dep. Puno: “prope Agapata”, 1854, Lech- 20(2): 187–189, fig. 6. 2000 – Holotype: ECUADOR.
ler, W. 1856 (BR barcode BR0000005178544 [online Loja: Parque Nacional Podocarpus, above Nudo de Caja-
image!]; isolectotypes: B [probably destroyed; F negative numa, Mirador, above Centro de Información, 04 05′S,
F0BN016711!], F barcodes V0040083F & V0040084F 79 10′W, 3000–3050 m, 15 May 1988, Ollgaard, B.

15
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

& al. 74157 (AAU ImageID 3508 [online image!]; iso- 29. Brachyotum longisepalum Wurdack in Phytologia
types: MO barcode MO-1515410 [online image!], LOJA, 11(6): 381–382. 1965 – Holotype: PERU. Dep. La Liber-
QCA barcode 58919 [online image!], QCNE barcode tad: Prov. Bolívar, jalca between Longotea and Bolí-
QCNE341 [online image!]). var, elev. 3600 m, 2 Jun 1960, Miranda, A.L. &
Sagástegui, A. 3338 (US barcode 00120117 [online
27. Brachyotum ledifolium (Desr.) Triana in Trans. Linn. image!]; isotypes: MO barcode MO-1704945 [online
Soc. London, Bot. 28(1): 48. 1872 (“1871”) ≡ Melastoma image!], TRP).
ledifolium Desr. in Lamarck, Encycl. 4: 48. 1796 ≡ Ple-
roma ledifolium (Desr.) DC., Prodr. 3: 151. 1828 – Lecto- 30. Brachyotum lutescens (Ruiz & Pav.) Triana in Trans.
type (designated here): ECUADOR. s.d., Jussieu, J. de Linn. Soc. London, Bot. 28(1): 48. 1872 (“1871”) ≡
s.n. (P barcode P02274335!). Rhexia lutescens Ruiz & Pav., Fl. Peruv. 3: 84. 1802 ≡
= Rhexia canescens Bonpl. in Humboldt & Bonpland, Arthrostemma lutescens (Ruiz & Pav.) DC., Prodr. 3:
Monogr. Melast. 2: 14–16, pl. 6. 1806 ≡ Chaetogastra 136. 1828 ≡ Alifana lutescens (Ruiz & Pav.) Raf. in Sylva
canescens (Bonpl.) DC., Prodr. 3: 134. 1828 ≡ Brachyo- Tellur.: 101. 1838 ≡ Chaetogastra lutescens (Ruiz & Pav.)
tum canescens (Bonpl.) Triana in Trans. Linn. Soc. Naudin in Ann. Sci. Nat., Bot., sér. 3, 14(2): 134. 1850 –
London, Bot. 28(1): 48. 1872 (“1871”) – Lectotype Lectotype (designated here): PERU. Dep. Huánuco: in
(designated here): COLOMBIA. Near Puracé, s.d., montibus Chaclla ad Piñapata et in Muña ad Tambo,
Bonpland, A.J.A. s.n. (COL barcode COL000003162 1787, Ruiz, H. & Pavón, J.A. s.n. (MA barcode MA
[online image!]; isolectotypes: F barcode V0040194F 813762 [online image!]; isolectotype: B [probably
[online image!], G, P barcodes P00136402!, destroyed; F negative F0BN016713!]).
P00136403! & P00136404!, VEN barcode 85194
[online image!]). 31. Brachyotum lycopodioides Triana in Trans. Linn. Soc.
London, Bot. 28(1): 49. 1872 (“1871”) – Lectotype (des-
28. Brachyotum lindenii Cogn. in Candolle & Candolle, ignated by Wurdack in Mem. New York Bot. Gard. 8(4):
Monogr. Phan. 7: 159. 1891 – Lectotype (designated by 396–397. 1953 ): PERU. Dep. Amazonas: prov. Chacha-
Wurdack in Mem. New York Bot. Gard. 8(4): 386. poyas ad Bajasan, 1835, Mathews, A. 1254 (K barcode
1953): COLOMBIA. Dep. Tolima: prov. Mariquita, ad K000329235!; isolectotype: E barcode E00285773
Tolima, “hauteur 2000 toises”, 1842, Linden, J.J. 925 [online image!]).
(G; isolectotypes: BR barcode BR0000005178537 = Brachyotum minimum Markgr. in Notizbl. Bot. Gart. Berlin-
[online image!], F barcodes V0063422F & V0063423F Dahlem 13: 459. 1937 – Type: PERU. Dep. Cajamarca:
[both fragments] [online images!], K barcode prov. Cutervo, Cutervo, May 1879 – Raimondi, A. 3505,
K000329224!, MPU barcode MPU013391 [online 3557 & 4898 (not located).
image!], P barcode P02274334!, W).
= Brachyotum rivetii Danguy & Cherm. in Bull. Mus. Hist. 32. Brachyotum lymphatum Wurdack in Mem. New York
Nat. (Paris) 28: 433. 1922 (“riveti”) – Lectotype (desig- Bot. Gard. 8(4): 395. 1953 – Holotype: COLOMBIA.
nated here): COLOMBIA. Province Mariquita: Forêts Dep. Cauca: Páramo de Guanacas, Cenual Andes of
du Quindiu, 3000 m, 1851, Triana s.n. (K barcode Popayan, alt. 3300–3600 m, s.d., Lebmann, F.C. 6025
K000329225!; isolectotype: P barcode P05202475!). (K barcode K000329233!).
Other syntypes: ECUADOR. El Pelado, Jan 1903, Rivet
342 (P barcode P02274333!); ECUADOR. Naes, Feb 33. Brachyotum markgrafii Wurdack in Mem. New York
1903, Rivet, C.G. 362 (P barcode P05202467!); ECUA- Bot. Gard. 8(4): 395–396. 1953 – Holotype: PERU.
DOR. Chemin d’El Angel à la Posta, 3500 m, Feb 1903, Dep. Huánuco: Cordillera of Pozuzo, 10–11,000 ft., Jul
Rivet, C.G. 407 (P barcode P05202469!). 1863, Pearce, R. s.n. (K barcode K000329234!).
= Brachyotum strigosum var. tolimensis Cuatrec. in Trab.
Mus. Nac. Ci. Nat., Ser. Bot. 33: 89. 1936 – Lectotype 34. Brachyotum maximowiczii Cogn. in Candolle & Can-
(designated here): COLOMBIA. Dep. Tolima: Andes, dolle, Monogr. Phan. 7: 154. 1891 – Holotype: PERU.
Cordillera Central, Vertient. merid. del monte Tolima, Dep. Amazonas: “in Peruvia ad Bajasan”, 1835,
subiendo al Nevado del Tolima, en Las Mesetas, malezas Mathews, A. 1257 (LE; isotypes: BR barcode BR00000
y bosque, alt. 3600 m, 13 May 1932, Cuatrecasas, J. 05183869 [fragment] [online image!], E barcodes
2834 (K barcode K000329222!; isolectotype: US barcode E00285779 & E00285780 [online images!], K barcode
00120125 [fragment] [online image!]). K000329194!).
Other syntypes: COLOMBIA. Dep. Tolima: Andes, Cor-
dillera Central, Entre ídem y el Páramo del Nevado, 34a. Brachyotum maximowiczii var. longifolium Wurdack in
14 May 1832, Cuatrecasas, J. 2835 (not located); Mem. New York Bot. Gard. 8(4): 369. 1953 – Holotype:
COLOMBIA. Dep. Tolima: Andes, Cordillera Central, PERU. Dep. Amazonas: Cerro de Fraijaco (Huaui-Huni)
15 May 1832, Cuatrecasas, J. 2836 (not located). northeast of Tambo Ventillas, in dry sandy soil, alt.

16
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

3000–3200 m, 7 Jul 1948, Pennell, F.W. 15857 (PH barcode lutescens Naudin in Ann. Sci. Nat., Bot., sér. 3, 14(2):
00008037 [online image!]; isotypes: PH barcode 00008036, 134. 1850 – Holotype: PERU. Dep. Huánuco [or more
USM barcode USM000516 [online image!]). probably Junín]: Palca, s.d., Dombey, J. s.n. (P barcode
P00136414!; isotypes: BM barcode BM001024973
35. Brachyotum microdon (Naudin) Triana in Trans. Linn. [online image!], F barcode V0040103F [fragment]
Soc. London, Bot. 28(1): 49. 1872 (“1871”) ≡ Chaetogas- [online image!], G, L barcode L 0008827 [online
tra microdon Naudin in Ann. Sci. Nat., Bot., sér. 3, 14(2): image!], P barcodes P00136415! [F negative
132–133. 1850 – Holotype: BOLIVIA. Dep. La Paz: near F0BN036134!], P00136416!, P00136417! & P001
“Carcuara Yungas” [this locality is perhaps Circuata], 36418!).
s.d., d’Orbigny 481 (P barcode P00136406!).
= Chaetogastra hermannioides Naudin in Ann. Sci. Nat., Bot., 39. Brachyotum nutans Gleason in Bull. Torrey Bot. Club
sér. 3, 14(2): 133. 1850 ≡ Brachyotum hermannioides 54(1): 25–26. 1927 – Holotype: PERU. Dep. Cuzco: Paso
(Naudin) Triana in Trans. Linn. Soc. London, Bot. 28(1): de Tres Cruces, 3 May 1925, Pennell, F.W. 13847
49. 1872 (“1871”) – Holotype: BOLIVIA. Prov. Tomina: (NY barcode 00221466!; isotypes: F barcode
s.d., Weddell, H.A. 3784 (P barcode P00136410!; isotypes: V0040102F [online image!], GH barcode 00071976
P barcodes P00136411! & P00136412!). [online image!], PH barcode 00008035 [online image!],
= Chaetogastra pentlandii Naudin in Ann. Sci. Nat., Bot., sér. S No. S05-3146 [online image!], US barcode 00120119
3, 14(2): 133. 1850 ≡ Brachyotum pentlandii (Naudin) [online image!]).
Triana in Trans. Linn. Soc. London, Bot. 28(1): 49.
1872 (“1871”) – Holotype: BOLIVIA. Cotagna, vallée 40. Brachyotum parvifolium Cogn. in Bot. Jahrb. Syst. 42:
chaude de l’Illimari, 1839, Pentland, J.B. s.n. (P barcode 132. 1908 – Lectotype (designated by Wurdack in
P00136407!; isotypes: F barcode V0063450F [fragment] Mem. New York Bot. Gard. 8(4): 366. 1953): Dep.
[online image!], P barcodes P00136408! & P00136409!). Amazonas: Tambo Ventillas apud Chachapoyas,
= Brachyotum setosum Gleason in Mem. New York Bot. elev. 2400–2500 m, Jul 1904, Weberbauer, A. 4406
Gard. 7: 314–315. 1927 – Holotype: BOLIVIA. 12 Jul (BR barcode BR0000005183876 [fragment] [online
1921, White, O.E. 151 (NY barcode 00221470!; isotypes: image!]; isolectotypes: B [probably destroyed; F negative
BKL barcode BKL00004024 [online image!], GH bar- F0BN016715!], G).
code 00071974 [online image!], K barcode
K000329219!, MICH barcode 1111797 [online image!], 41. Brachyotum quinquenerve (Ruiz & Pav.) Triana in Trans.
PH barcode 00008033 [online image!], US barcode Linn. Soc. London, Bot. 28(1): 48. 1872 (“1871”) ≡
00120124 [online image!]). Rhexia quinquenervis Ruiz & Pav., Fl. Peruv. 3:
83, t. 321, fig. b. 1802 ≡ Arthrostemma quinquenerve
36. Brachyotum multinervium Wurdack in Phytologia 11(6): (Ruiz & Pav.) DC., Prodr. 3: 136. 1828 ≡ Chaetogastra
379–380. 1965 – Holotype: PERU. Dep. Amazonas: prov. quinquenervis (Ruiz & Pav.) Naudin in Ann. Sci. Nat.,
Chachapoyas, slope seepage below km 415 of Balsas- Bot., sér. 3, 14(2): 130. 1850 – Lectotype (designated
Leimebamba road, middle eastern Calla-Calla slopes, elev. here): PERU. Habitat in Huassahuassi, Panao, Chaclla,
3100–3250 m, 11 Jul 1962, Wurdack, J.J. 1304 (US barcode et Muiña montibus, copiosè in Sancti Dominici et Lla-
00120118 [online image!]; isotypes: F barcode V0040105F mapaiñaui, collibus, s.d., Pavón, J.A. s.n. (BC barcode
[online image!], G barcode G00353534 [online image!], BC-873057 [online image!]; isolectotypes: B [probably
GH barcode 00071975 [online image!], K barcode destroyed; F negative F0BN016716!], MA barcodes
K000329243!, LIL barcode LIL001034 [online image!], MA 813772, MA 813773 & MA 813774 [online images!]).
MICH barcode 1111798 [online image!], MO barcode
MO-2272359 [online image!], NY barcode 00221464!, P 41a. Brachyotum quinquenerve var. pusillum Wurdack in
barcode P00136413!, S No. S05-3145 [online image!], U Mem. New York Bot. Gard. 8(4): 361. 1953 – Holotype:
barcode U 0003967 [online image!], UC barcode PERU. Dep. Amazonas: along Rio Sonche, west of Moli-
UC1289645 [online image!], USM). nopampa, dry sandy barren, 2400 m a.s.l., 8 Jul 1948,
Pennell, F.W. 15772 (PH barcode 00008034 [online
37. Brachyotum multituberculatum Wurdack in Mem. image!]; isotype: USM barcode USM000517 [online
New York Bot. Gard. 8(4): 394–395. 1953 – Holotype: image!]).
PERU. Dep. Amazonas: “La Jalca” between Chacha-
poyas and Moyobamba, alt. 2700–3300 m, 21 Jan 1930, 42. Brachyotum radula Triana in Trans. Linn. Soc. London,
Williams, L. 7587 (F barcode V0040104F [online Bot. 28(1): 48. 1872 (“1871”) – Holotype: PERU. Dep.
image!]; isotype: NY barcode 00221465!). Amazonas: Chachapoyas, s.d., Mathews, A. 3210
(K barcode K000329192!).
38. Brachyotum naudinii Triana in Trans. Linn. Soc. = Brachyotum asperum Cogn. in Bot. Jahrb. Syst. 42: 132.
London, Bot. 28(1): 48. 1872 (“1871”) ≡ Chaetogastra 1908 – Lectotype (designated by Burke & Michelangeli

17
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

in Phytotaxa 125(1): 11. 2013): PERU. Dep. Cajamarca: Azuay: N and NW of the Páramo del Castillo (ca. 6–8 km
supra Hacienda La Tahona apud Hualgayoc, elev. N-NE of Sevilla de Oro), 3048–3413.8 m, 31 Aug 1945,
3400–3700 m, 13 May 1904, Weberbauer, A. 4013 Camp, W.H. E-5161 (NY barcode 00221469!; isotypes:
(BR barcode BR0000005184217 [online image!]; iso- BM barcode BM000953911 [online image!], F barcode
lectotypes: B [probably destroyed; F negative V0063426F [online image!], G barcode G00353537
F0BN016707!], G, MOL). [online image!], GH barcode 00071978 [online image!],
K barcode K000329239!, MO barcode MO-288051
43. Brachyotum rosmarinifolium (Ruiz & Pav.) Triana in [online image!], P barcode P00136424!, U barcode
Trans. Linn. Soc. London, Bot. 28(1): 48. 1872 U 0003968 [online image!], UC barcode UC951482
(“1871”) ≡ Rhexia rosmarinifolia Ruiz & Pav. Fl. Peruv. [online image!], US barcode 00120123 [online image!],
3: 84. 1802 ≡ Arthrostemma rosmarinifolium (Ruiz VEN barcode 32934 [online image!]).
& Pav.) DC., Prodr. 3: 136. 1828 ≡ Chaetogastra rosmar-
inifolia (Ruiz & Pav.) Benth., Pl. Hartw.: 131. 1844 – Lec- 45. Brachyotum rotundifolium Cogn. in Bull. Acad. Roy.
totype (designated here): PERU. Dep. Huánuco and Sci. Belgique, sér. 3, 14(7): 937–938. 1887 – Holotype:
Junín: in Huanuci et Tarmae collibus, s.d., Ruiz, H. ECUADOR. Apud páramos, montium in Andibus Ecua-
& Pavón, J.A. s.n. (BC barcode BC-873059 [online dorensibus frequens, alt. 3200–3800 m, 1876, André,
image!]; isolectotypes: B [probably destroyed; F negative É.-F. 4501 (K barcode K000329190!; isotypes: BR bar-
F0BN016907!], MA barcodes MA 813775 & MA code BR0000005184194 [online image!], F barcode
813776 [online images!]). V0063424F [online image!], GH barcode 00071977
[online image!], K barcode K000329189!, NY barcode
44. Brachyotum rostratum (Naudin) Triana in Trans. Linn. 00221467!, P barcode P02274328!, US barcode
Soc. London, Bot. 28(1): 48. 1872 (“1871”) ≡ Chaetogas- 00120121 [online image!]).
tra rostrata Naudin in Ann. Sci. Nat., Bot., sér. 3, 14(2):
135–136. 1850 – Holotype: PERU. Probably in dept. 46. Brachyotum rugosum Wurdack in Mem. New York Bot.
Huánuco: Pasco, or Junín, s.d., Dombey, J. s.n. Gard. 8(4): 376. 1953 – Holotype: ECUADOR. Prov.
(P barcode P00136419!; isotypes: B [probably destroyed; Santiago-Zamora: trail between Pailas and El Pan, alt.
F negative F0BN016718!], BM, BR barcode 2255–3445 m, 10 Sep 1943, Steyermark, J.A. 54316
BR0000005184538 [online image!], F, G-DC barcode (F barcode V0063425F [online image!]; isotypes: NY
G00318688 [online image!], L barcode L 0008828 barcode 00221468!, US barcode 00120122 [online
[online image!], P barcodes P00136420!, P00136421! image!]).
& P00136422!, US).
= Chaetogastra microphylla Naudin in Ann. Sci. Nat., Bot., 47. Brachyotum russatum E.Cotton in Nordic J. Bot. 20(2):
sér. 3, 14(2): 136. 1850 ≡ Brachyotum microphyllum 187–188, fig. 5a–e. 2000 – Holotype: ECUADOR. Par-
(Naudin) Triana in Trans. Linn. Soc. 28(1): 49. 1872 que Nacional Podocarpus, Cerro Toledo, E of Yangana,
(“1871”) – Holotype: PERU. s.l., s.d., Bonpland, 04 24′S, 79 06′W, 3300–3450 m, 23 Feb 1985,
A.J.A. s.n. (P barcode P00136423!; isotype: F barcode Oellgaard, B.; Laegaard, S.; Thomsen, K.; Korning, J.
V0040106F [fragment] [online image!]). & Illum, T. 58025 (AAU ImageID 3507 [online image!];
= Brachyotum trianaei Cogn. in Candolle & Candolle, isotypes: GB barcode GB-0048127 [online image!], K
Monogr. Phan. 7: 167. 1891 – Holotype: Puno. “Tabina” barcode K000329244 [online image!], MO barcode
near Ayapata, s.d., Lechler, W. 2061 (BR barcode MO-149737 [online image!], LOJA, NY barcode
BR0000005184545 [online image!]; isotypes: B [proba- 00452035!, QCA barcode 59183 [online image!], US bar-
bly destroyed; F negative F0BN16719!], BR barcode code 00901924 [online image!]).
BR0000005183883 [online image!], F barcode
V0040100F [fragment] [online image!], G-DC barcode 48. Brachyotum sanguinolentum (Naudin) Triana in Trans.
G00318685 [online image!], K barcode K000329238!, Linn. Soc. London, Bot. 28(1): 48. 1872 (“1871”) ≡
NY barcode 00221471!, P, S No. S05-3148 [online Chaetogastra sanguinolenta Naudin in Ann. Sci. Nat.,
image!]). Bot., sér. 3, 14(2): 131–132. 1850 – Holotype: BOLIVIA.
= Brachyotum callosum J.F.Macbr. in Publ. Field Mus. Nat. Dep. La Paz: prov. de Larecaja et Caupolican (vallées
Hist., Bot. Ser. 4(7): 172–173. 1929 – Holotype: PERU. entre Tipoani et Apolobamba) à 2500 mètres, May
Huánuco: north of Huánuco, 16 Jun 1922, Macbride, 1847, Weddell, H.A. 4605 (P barcode P02274326!; iso-
J.F. & Featherstone, W. 2181 (F barcode V0040079F type: P barcode P02274327!).
[online image!]; isotypes: G, NY barcode 00221455!, S = Brachyotum floribundum Cogn. in Candolle & Candolle,
No. S05-3139 [online image!], US barcode 00120110 Monogr. Phan. 7: 155. 1891 – Lectotype (designated
[online image!]). here): PERU. Puno: “Tabina” near Ayapata, 1854,
= Brachyotum seorsum Wurdack in Mem. New York Bot. Lechler, W. 1857 (G-DC barcode G00318663 [online
Gard. 8(4): 402–403. 1953 – Holotype: ECUADOR. image!]; isolectotypes: B [probably destroyed; F negative

18
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

F0BN016709!], BR barcode BR0000005184521 [online Jahrb. Syst. 42: 133. 1908 – Lectotype (designated by
image!], F barcodes V0040081F & V0040082F [both Burke & Michelangeli in Phytotaxa 125(1): 11. 2013):
fragments] [online images!], GOET barcode GOET PERU. Dep. Junin: prov. Tarma, in montibus ad orientem
007884 [online image!], K barcode K000329220!, LE!, a Palca versus, in fruticeto denso secus rivum,
P barcodes P02274323! & P02274324!, S Nos. 3100–33200 m a.s.l., 12 Feb 1093, Weberbauer, A. 2475
S05-3140 & S09-13094 [online images!]). (BR barcode BR0000005226993 [online image!]; isolec-
= Brachyotum barbiferum Rusby in Phytologia 1: 69. 1934 – totype: B [probably destroyed; F negative F0BN
Holotype: BOLIVIA. [Department of La Paz]: Coco- 016807!]).
punco in the Cordillera Real, 10,000 ft., 24 Mar 1926,
Tate, G.H.H. 362 (NY barcode 00221453!; isotype: NY 55. Brachyotum weberbaueri Cogn. in Bot. Jahrb. Syst. 42:
barcode 00022556!). 133. 1908 – Lectotype (designated by Wurdack in
= Brachyotum pedicellatum Markgr. in Notizbl. Bot. Gart. Mem. New York Bot. Gard. 8(4): 373–374. 1953):
Berlin-Dahlem 13: 460. 1937 – Holotype: BOLIVIA. PERU. Dep. Amazonas: “Tambo Ventillas, ad orientem
Camata-Cuibaja, 3400 m, 3 Oct 1926, Troll, C. 2463 a Chachapoyas versus”, alt. 2400–2600 m, 28 Jul 1904,
(M barcode M-0165354 [online image!]). Weberbauer, A. 4405 (BR barcode BR0000005184224
[online image!]; isolectotypes: B [probably destroyed; F
49. Brachyotum sertulatum C.Ulloa in Anales Jard. Bot. negative F0BN016720!], F barcode V0040098F [frag-
Madrid 64(1): 70, fig. 1. 2007 – Holotype: COLOM- ment] [online image!], G, MOL barcode 00003175
BIA. Huila-Cauca: Macizo colombiano; Páramo de [online image!]).
Las Papas, colinas al SE de la laguna “La Magdalena”
sobre el cerro “La Carona” y “El Boquerón”, vertiente Bucquetia DC., Prodr. 3: 110. 1828 – Type: Bucquetia gluti-
del Magdalena, 3330–3520 m, 8 Sep 1958, Idrobo, nosa (L.f.) DC. (≡ Rhexia glutinosa Mutis ex L.f.).
Pinto & Bischler 3106 (COL; isotype: P barcode Shrubs and small trees, much branched. Plants
P00506904!). completely glabrous, except in some structures of the flower
or juveniles that tend to be puberulous; buds and young leaves
50. Brachyotum strigosum (L.f.) Triana in Trans. Linn. Soc. resinous. Leaves opposite, decussate, similar for the same
London, Bot. 28(1): 48. 1872 (“1871”) ≡ Melastoma node, symmetrical, petiolate, chartaceous; some lustrous,
strigosum L.f., Suppl. Pl.: 236. 1781 – Lectotype (desig- lower surface with tiny dark spots (puncticulate), obscurely
nated here): COLOMBIA. Dep. Cundinamarca: “Nova 3-nerved or shortly (ca. 1 mm) 3-plinerved. Inflorescence in
Granada”, probably in the vicinity of Bogotá, s.d., Mutis terminal few-flowered cymes with long stem, flowers short-
s.n. (LINN No. 559.6 [online image!]; isolectotype: pedicellate; bracteoles 2, enclosing the hypanthium. Flower
LINN No. 559.7 [online image!]). perigynous. Hypanthium campanulate, externally smooth or
with slight ribs (terete), glabrous. Calyx not calyptriform, with
51. Brachyotum trichocalyx Triana in Trans. Linn. Soc. conspicuous lobes, lobes 4, persistent, glabrous or with sparse
London, Bot. 28(1): 48. 1872 (“1871”) – Holotype: setae, broadly triangular. Corolla patent; petals 4, free and
ECUADOR. Prov. Loja: Quito, 1865, Jameson s.n. spreading, magenta or dark pink. Stamens 8, all fertile, iso-
(K barcode K000329230!). morphic. Anthers 2-celled, narrowly oblong, straight, ven-
trally smooth, with a minute apical pore dorsally oriented.
52. Brachyotum tyrianthinum J.F.Macbr. in Publ. Field Mus. Connectives glabrous, not prolonged, not or obscurely appen-
Nat. Hist., Bot. Ser. 4(7): 174–173. 1929 – Holotype: daged, typically declined to side of flower. Filaments thin, flat-
PERU. Dep. Huánuco: Mito, alt. 2750 m, Jul 1922, Mac- tened, glabrous. Ovary superior, 4-celled, strigose in the upper
bride, J.F. & Featherstone, W. 1438 (F barcode half; style filiform, glabrous; stigma punctiform, inconspicu-
V0040099F [online image!]; isotypes: G, K barcode ous. Mature capsule dry, semiwoody, 4-valved, no angle.
K000684621!, S No. S05-3147 [online image!], US bar- Seeds numerous, tiny, oblong-cuneate, essentially smooth
code 00120126 [online image!]). raphe inconspicuous.
Bucquetia has five taxa (three species, two varieties),
53. Brachyotum uribei Wurdack in Phytologia 29(2): which are restricted to the northern Andes, and found in areas
141–142. 1974 – Holotype: COLOMBIA. Dep. Cauca: of páramo and subpáramo of Venezuela (Táchira), northern
“monte Puracé camino hacia la laguna de San Rafael”, Colombia and southern Ecuador at 2300–4300 m.
elev. 3000 m, 7 Sep 1961, Uribe, L. 3847 (US barcode Bucquetia species are glabrous and glutinous shrubs with
00120127 [online image!]; isotype: COL barcode tetramerous flowers in which the magenta or dark pink color
COL000003194 [online image!], P barcode P00 of the petals contrasts with the yellow of the straight stamens.
136434!).
Centradenia G.Don, Gen. Hist. 2: 730, 765. 1832 – Type:
54. Brachyotum virescens (Cogn.) Wurdack in Phytologia Centradenia inaequilateralis (Schltdl. & Cham.) G.Don.
64(4): 296. 1988 ≡ Tibouchina virescens Cogn. in Bot. (≡ Rhexia inaequilateralis Schltdl. & Cham.).

19
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

Erect shrubs, subshrubs, or suffrutescent, (0.06–)0.2–2 Chaetogastra longifolia (Vahl) DC. (≡ Rhexia longifolia
(–3) m tall, perennial; puberulous, hispid or strigose on stems Vahl. ≡ Pleroma longifolium (Vahl.) Triana ≡ Tibouchina
and leaves with smooth, unbranched, multicellular trichomes; longifolia (Vahl.) Baill.).
glandular trichomes with multicellular stalks terminated by a = Oreocosmus Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(1):
globose cell. Leaves opposite, decussate, but apparently 37. 1849 – Type (designated here): Chaetogastra tor-
pseudo-alternate by partial or complete anisophylly; oblique tuosa (Bonpl.) DC. (≡ Rhexia tortuosa Bonpl. ≡ Oreocos-
or almost symmetric at the base; smaller leaves early absciss- mus tortuosus (Bonpl.) Naudin ≡ Tibouchina tortuosa
ing; short-petiolate or subsessile, glabrate to copiously pubes- (Bonpl.) Almeda).
cent, above and strigillose below, 3–7-plinerved. Inflorescence = Hephestionia Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(1):
terminal, basically a multiflowered dichasium, but frequently 36. 1850 – Type (designated by Pfeiffer in Nomencl. Bot.
becoming paniculate, racemose, or subfasciculate by reduc- 1(2): 1610. 1874): Chaetogastra strigosa (Rich.) DC. (=
tion; bracteoles 2. Flowers perigynous; hypanthium urceolate Chaetogastra inconstans (Vahl.) P.J.F.Guim. & Miche-
to campanulate; externally with 8 slight, longitudinal vascular lang.).
ribs, sparsely pubescent, trichomes simple, glandular stalked = Lasiandra sect. Simplicicaules Naudin in Ann. Sci. Nat.,
or sessile glandular and crystalline excrescences. Calyx lobes Bot., sér. 3, 13(3): 127. 1850, pro parte – Type (desig-
conspicuous, not calyptrate, lobes 4, small, persistent in fruit, nated here): Chaetogastra hieracioides DC. (≡ Lasian-
broadly deltoid to triangular-ovate or truncate. Corolla patent; dra hieracioides Naudin).
petals 4; free, spreading or connivent and somewhat cupulate; = Lasiandra sect. Macrodon Naudin in Ann. Sci. Nat., Bot.,
white or pink to magenta, glabrous. Stamens 8, all fertile, sér. 3, 13(3): 130. 1850 – Type (designated here):
strongly dimorphic to subequal, composed of 4 larger stamens Lasiandra sarmentosa Naudin (= Tibouchina laxa
in an antesepalous whorl and 4 smaller ones in an antepetalous (Desr.) Cogn.).
whorl; the largest connective conspicuously prolonged and = Lasiandra sect. Mavrocarpus Naudin in Ann. Sci. Nat.,
smaller with connective shortly prolongated. Anthers Bot., sér. 3, 13(3): 131. 1850 – Type (designated here):
2-celled, oblong to linear-oblong, straight, ventrally chan- Chaetogastra pleromoides (Naudin) P.J.F.Guim.
neled, with a small apical pore ventrally oriented. Connective & Michelang. (≡ Lasiandra pleromoides Naudin).
in the stamens larger geniculate and with two amorphous ven- = Purpurella Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3):
tral lobes at the point of articulation to the filament; in the 301. 1850, pro parte – Type (designated here): Purpu-
small stamens, connective prolonged little or no longer below rella muricata (Bonpl.) Naudin (= Tibouchina grossa
the thecae, however, presents a ventral basal appendix amor- (L.f.) Cogn.).
phous (sometimes articulated) and slightly lobed. Thin fila- = Micranthella Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(6):
ments compressed, glabrous. Ovary superior, 4-celled, 347. 1850 – Type (designated here): Chaetogastra clin-
setulose apically, completely enclosed by the hypanthium opodifolia DC. (≡ Micranthella clinopodifolia (DC.)
and calyx but not fused to them; axile placentation; style fili- Naudin ≡ Tibouchina clinopodifolia (DC.) Cogn.).
form, glabrous, straight or somewhat sigmoid and declinate; = Pleroma sect. Diotanthera Triana in Trans. Linn. Soc.
stigma inconspicuous. Mature capsule dry, semiwoody, loculi- London, Bot. 28(1): 45. 1872 (“1871”), pro parte – Type
cidal, with veins or ribs persisting on old fruits. Seeds numer- (designated here): Chaetogastra debilis Cham. (≡ Ple-
ous, tiny, narrowly pyriform to clavate or dolabriform, roma debile (Cham.) Triana).
echinate (often glandularly so), verrucose to muriculate or Shrubs, subshrubs, herbs or treelets covered with spreading-
papillate, raphe lateral conspicuous. pubescent trichomes or villose to setulose. Leaves opposite,
Six taxa are recognized in Centradenia (four species, two decussate, blades of a pair essentially equal in size, petiolate,
subspecies), which are restricted to Mexico, Central America membranaceous, chartaceous to coriaceous, variously pubes-
and northern South America. Their distribution extends from cent to nearly glabrous, or tuberculate 3–5(–7)-nerved. Inflo-
Hidalgo, Mexico, southeastward through Central America to rescence a terminal anthotelic frondobracteose panicle or
Panama, with one outlying species known from the Chocó some modification of this structure or few-flowered; bracte-
lowlands on the Pacific coast of Colombia (Almeda, 1977). oles 2, free, concave, variously pubescent, early deciduous
All species are restricted in habitat, occurring most commonly or persistent, smaller than the mature hypanthium. Flowers
on sheltered roadbanks, margins of streams and in disturbed hypogynous; hypanthium campanulate, mostly copiously
forests, or on moist cliffs in the spray of waterfalls between pubescent externally. Calyx lobes 4–5, persistent. Petals 4–5,
200 and 1900 m (Almeda, 1977). dark lilac to purple, rarely white or yellow or deep red, spread-
This genus is easily diagnosable by its pronounced aniso- ing or developing in an open-campanulate corolla and not
phylly, basally oblique larger leaves, and the pyriform, clavate, remaining in a tube, glabrous or in a few instances pubescent
or dolabriform seeds with a conspicuous lateral raphe on the adaxial side. Stamens 8 or 10, isomorphic, or weakly to
(Almeda, 1977). moderately or even strongly dimorphic. Anthers 2-celled, ven-
trally channeled, subulate, slightly arcuate and lanceolate, api-
Chaetogastra DC., Prodr. 3: 131. 1828 – Type (designated by cal pores ventrally inclined. Connectives variously prolonged,
Wurdack in Mem. New York Bot. Gard. 8(4): 343. 1953): bilobed ventrally, with glabrous appendages. Filaments

20
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

glabrous. Ovary superior, apical portion with hairs, 4–5-celled Cochabamba: “Im subalpinen Gebüsch an feuchten Stel-
with axile placentation; style glabrous or rarely moderately len bei Comarapa”, 2300 m, Apr 1911, Herzog,
short-setose with fine mostly eglandular hairs, hooked or T.C.J. 2014A (BR barcode BR0000005523931 [frag-
incurved apically, typically declined to side of flower oppos- ment]!; isolectotype: B [probably destroyed]).
ing stamens. Capsule dry, semiwoody, loculicidal. Seeds = Tibouchina brachyphylla Gleason in Phytologia 1(3): 136.
numerous, cochleate to elongate-cochleate and tuberculate 1935 – Holotype: BOLIVIA. Dep. Santa Cruz: Comarapa,
with a terminal hilum. Bergwälder, forest at Comarapa, alt. 2500–3000 m, 20 Oct
Chaetogastra has 117 species, which are distributed from 1928, Steinbach, J. 8332 (NY barcode 00245753!; isotypes:
Mexico, Central America and the West Indies through the Andes E barcode E00285782 [online image!], GH barcodes
(Venezuela, Colombia, Ecuador, Peru, Bolivia and Argentina) to 00073256 & 00073257 [online images!], GOET barcode
eastern Brazil, mostly in cloud forests above 1000 m elevation. GOET008065 [online image!], K barcode K000329165!,
Some herbaceous species of wide distribution, such as Chaeto- NY barcode 01795930!, PH barcode 00027260 [online
gastra gracilis, occupy open areas above 100 m from Colombia image!], S No. S-R-6110 [online image!], UC barcode
and Venezuela to Bolivia, northern Argentina, and central Brazil. UC394785 [online image!], US barcode 00120044!).
The center of diversity is in the Andes.
The genus is characterized by the presence of isomorphic 5. Chaetogastra anderssonii (Wurdack) P.J.F.Guim. & Miche-
or weakly to strongly dimorphic stamens with yellow anthers lang., comb. nov. ≡ Tibouchina anderssonii Wurdack in
and connective shortly prolonged and bilobed, ventrally with Phytologia 35(1): 1–2. 1976 – Holotype: ECUADOR. Prov.
appendages. Species with pronounced staminal dimorphism El Oro: between Piñas and El Placer along the Zaruma–
can have anthers of the larger whorl differently colored from Santa Rosa road, elev. 1100–1200 m, 6 May 1974, Harling,
the smaller whorl. G. & Andersson, L. 14363 (US barcode 00120036!; iso-
type: GB barcode GB-0048178 [online image!]).
Species list
6. Chaetogastra araneicalyx (de Santiago) P.J.F.Guim. &
1. Chaetogastra adenophora (Cogn.) P.J.F.Guim. & Miche- Michelang., comb. nov. ≡ Tibouchina araneicalyx de
lang., comb. nov. ≡ Tibouchina adenophora Cogn. in Santiago in Novon 14(2): 164–166, fig. 1. 2004 – Holo-
Bull. New York Bot. Gard. 4(14): 355. 1907 – Holotype: type: MEXICO. Guerrero: mun. Atoyac de Alvarez,
BOLIVIA. s.d., Bang, M. 2645 (NY barcode 00245748!; aproximadamente a 2 km al W de E1 Molote, 17 25′
isotypes: BR barcode BR0000005209453 [fragment]!, 06′′N, 100 10′41′′W, 1650 m, 5 Dec 2000, de Santiago
CORD barcode CORD 00003624 [online image!]). Gómez, J.R.; Lozada, L.; León, E. & Rojas, J. 870
(MEXU barcode MEXU 01138500 [online image!]; iso-
2. Chaetogastra alata P.J.F.Guim. & Michelang., nom. nov. ≡ types: CAS barcode 0026358 [online image!], ENCB,
Tibouchina stricta Wurdack in Phytologia 5(2): 56–57. FCME, K barcode K000328988!, MO barcode MO-
1954, non Chaetogastra stricta (Bonpl.) DC. 1828 – Holo- 1515292 [online image!]).
type: COLOMBIA. Dep. Cauca: “cabaceras del rio Palo,
quebrada del rio Lopez: Quebrada del Duende”, elev. 7. Chaetogastra arthrostemmoides (Cogn.) P.J.F.Guim.
3400–3450 m, 2 Dec 1944, Cuatrecasas, J. 18923 & Michelang., comb. nov. ≡ Tibouchina arthrostem-
(NY barcode 00245811!; isotypes: F barcode V0063999F moides Cogn. in Bull. Acad. Roy. Sci. Belgique, sér.
[online image!], MO barcode MO-2267315 [online 3, 14(7): 934. 1887 – Lectotype (designated here):
image!], P barcode P00708782!, US barcode 00120100!). ECUADOR. San Pablo Andium [pastensium] (Nova
Granate), 1270 m, s.d., André, É.-F. 3339 (K barcode
3. Chaetogastra almedae (Todzia) P.J.F.Guim. & Miche- K000329146!; isolectotype: BR barcode BR0000
lang., comb. nov. ≡ Tibouchina almedae Todzia in Britto- 005523948 [fragment]!).
nia 51(3): 255–259, fig. 1, 11. 1999 – Holotype: Other syntype: COLOMBIA. Altaquer in Cordillera me-
MEXICO. Sinaloa: mun. Concordia, dry slope with Pinus ridionalis Andium, 900 m, May 1876, André, E.-F. s.n.
and Quercus near crest of ridge on small dirt logging road (K barcode K000329147!).
near Loberas microwave station, 2000 m, 18 Oct 1983,
Breedlove, D.E. & Almeda, F. 58858 (CAS barcode 8. Chaetogastra asperifolia (Cogn.) P.J.F.Guim. & Miche-
0004734 [online image!]; isotypes: MEXU barcode lang., comb. nov. ≡ Tibouchina asperifolia Cogn. in
MEXU 00867943 [online image!], TEX barcode Bot. Jahrb. Syst. 42: 137. 1908 – Lectotype (designated
00375780 [online image!]). by Burke & Michelangeli in Phytotaxa 125(1): 12.
2013): PERU. Dep. Huánuco: prov. Huamalíes, in monti-
4. Chaetogastra montana P.J.F.Guim. & Michelang., nom. bus ad austro-occidentem a Monzon versus, in fruticeto
nov. ≡ Tibouchina alpestris Cogn. in Meded. Rijks-Herb. fruticibus et hic illic arboribus parvis foliis induratis
19: 78–79. 1913, non Chaetogastra alpestris (DC.) Mart. instructis composito, 2500–3000 m a.s.l., 12 Jul 1903,
1831 – Lectotype (designated here): BOLIVIA. Weberbauer, A. 3357 (BR barcode BR0000005209149!;

21
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

isolectotypes: B [probably destroyed; F negative Ann. Sci. Nat., Bot., sér. 3, 14: 122. 1850 ≡ Pterolepis cer-
F0BN016727!], MOL barcode 00003339 [online astifolia (Naudin) Triana in Trans. Linn. Soc. London,
image!]). Bot. 28(1): 40. 1872 (“1871”) ≡ Tibouchina cerastifolia
(Naudin) Cogn. in Martius, Fl. Bras. 14(3): 403–404.
9. Chaetogastra andreana (Cogn.) P.J.F.Guim. & Miche- 1885 – Holotype: BRAZIL. Minas Gerais: in sylva urbis
lang., comb. nov. ≡ Tibouchina andreana Cogn. in Bull. San João del Rey vicina, s.d., De Saint-Hilaire,
Acad. Roy. Sci. Belgique, sér. 3, 14(7): 936. 1887 – Lec- A.F.C.P. D566 (P barcode P00121003!; isotypes: P bar-
totype (designated here): COLOMBIA. Habitat at codes P00121004! & P00121005!).
Boquerón del Quindío, in Cordillera centrali Novae = Tibouchina cerastifolia var. major Cogn. in Martius,
Granatae, alt. 3450 m, 10 Mar 1876, André, É.-F. 2247 Fl. Bras. 14(3): 404. 1885, syn. nov. – Lectotype (desig-
part. (K barcode K000329164!; isolectotypes: BR bar- nated here): BRAZIL. São Paulo: in locis apricis prope
code BR0000005523610 [fragment]!, NY barcode Santos, alt. 1000 m, s.d., Mosén, C.W.H. 3267 (BR bar-
00245750!). code BR0000005224791!; isolectotype S No. S09-
13205!).
10. Chaetogastra aurea (Cogn.) P.J.F.Guim. & Michelang., = Tibouchina cerastifolia var. hirsuta Cogn. in Candolle
comb. nov. ≡ Tibouchina aurea Cogn. in Meded. & Candolle, Monogr. Phan. 7: 1177. 1891, syn. nov. –
Rijks-Herb. 19: 77. 1913 – Holotype: BOLIVIA. Lectotype (designated here): BRAZIL. Santa Catarina:
Cochabamba: “an kahlen Stellen im Bergwald über Serra do Mar prope São Bento, 11 Feb 1890,
Tablas”, 2400 m, May 1911, Herzog, T.C.J. 2157 Schwacke, A. 6916 (BR barcode BR0000005225132!;
(B [probably destroyed; F negative F0BN016729!]; iso- isolectotype: RB barcode 00541551!).
types: BR barcode BR0000005522286 [fragment]!,
JE barcode JE00004635 [online image!], S No. S-R- 15. Chaetogastra chamaecistus Sieber ex Griseb., Fl. Brit.
6108!, W!). W. I.: 267. 1860 ≡ Tibouchina chamaecistus (Sieber ex
Griseb.) Cogn. in Candolle & Candolle, Monogr. Phan.
11. Chaetogastra brittoniana (Cogn. ex Rusby) P.J.F.Guim. 7: 267. 1891 – Lectotype (designated here): MARTI-
& Michelang., comb. nov. ≡ Tibouchina brittoniana NICA. On the top of the Montagne Pêlée, Jun 1822, Sie-
Cogn. ex Rusby in Bull. Torrey Bot. Club 17(3): 55. ber, F.W. 297 (M barcode M-0165827!; isolectotypes: BR
1890 – Holotype: BOLIVIA. Yungas, 4000 ft., s.d., barcodes BR0000005530625 [fragment]! & BR00
Rusby, H.H. 2341 (NY barcode 00245754!; isotype: BR 00005530953!, F barcode V0063976F [fragment]!, K
barcode BR0000005530298 [fragment]!). barcodes K000328994! & K000328995!, LE [on 2
sheets]!, M barcode M-0165828!).
12. Chaetogastra campii (Wurdack) P.J.F.Guim. & Miche-
lang., comb. nov. ≡ Tibouchina campii Wurdack in 16. Chaetogastra chironioides Griseb., Fl. Brit. W. I.: 267.
Mem. New York Bot. Gard. 16: 1. 1967 – Holotype: 1864, pro parte ≡ Tibouchina chironioides (Griseb.)
ECUADOR. Prov. Morona-Santiago: between Tres Ran- Cogn. in Candolle & Candolle, Monogr. Phan. 7: 267.
chos and Chontal, eastern slope of the cordillera, valley 1891 – Lectotype (designated here): Dominica. s.d.,
of the ríos Negro and Chupianza (on the trail from Sevilla Imray, J. s.n. (NY barcode 00099392!; isolectotypes:
de Oro to Mendez), alt. 2700–5700 ft., 15 Dec 1944, E barcode E00531223 [online image!], GH barcode
Camp, W.H. E-1546 (US barcode 00120045!; isotypes: 00072043 [online image!], K barcode K000328996!).
F barcode V0063980F [online image!], NY barcode
00245755!). 17. Chaetogastra ciliaris (Vent.) DC., Prodr. 3: 132. 1828 ≡
Meriania ciliaris Vent. in Mém. Cl. Sci. Math. Inst. Natl.
13. Chaetogastra capitata (Naudin) P.J.F.Guim. & Miche- France 8: 14–15. 1807 ≡ Tibouchina ciliaris (Vent.)
lang., comb. nov. ≡ Micranthella capitata Naudin in Cogn. in Bull. Acad. Roy. Sci. Belgique, sér. 3, 14(7):
Ann. Sci. Nat., Bot., sér. 3, 13(6): 351. 1850 ≡ Pleroma 932. 1887 – Holotype: COLOMBIA. Santa Fé, s.d.,
capitatum (Naudin) Triana in Trans. Linn. Soc. London, Bonpland, A.J.A. s.n. (G barcode G00341509!
Bot. 28(1): 46. 1872 (“1871”) ≡ Tibouchina capitata [F negative F0BN026102!]; isotypes: G-DC barcode
(Naudin) Cogn. in Bull. Torrey Bot. Club 17(3): 54. G00310491 [fragment]!, P barcode P00121034!).
1890 – Holotype: BOLIVIA. [Dep. La Paz]: prov. de La-
recaja, Caupolican, vallées entre Tipoani et Apolobamba, 18. Chaetogastra cisplatensis (Cogn.) P.J.F.Guim. & Miche-
alt. 3000 m, Apr 1847, Weddell, H.A. 4584 (P barcode lang., comb. nov. ≡ Tibouchina cisplatensis Cogn. in
P00116995!; isotype: F barcode V0063876F [fragment] Martius, Fl. Bras. 14(3): 407–408. 1885 – Lectotype
[online image!]). (designated here): BRAZIL. In districtu Cisplatina, s.
d., Sellow, F.D. 2092 (BR barcode BR0000005224517!;
14. Chaetogastra cerastifolia (Naudin) P.J.F.Guim. & Miche- isolectotypes: B [probably destroyed; F negative
lang., comb. nov. ≡ Tetrameris cerastifolia Naudin in F0BN016734!], K barcode K000329100!, LE!).

22
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

19. Chaetogastra cistoides Griseb. in Fl. Brit. W. I. 3: 267. isotypes: CAS barcode 0004850 [online image!], GBH
1860 ≡ Tibouchina cistoides (Griseb.) Cogn. in Candolle barcode GBH009952 [online image!], GH barcode
& Candolle, Monogr. Phan. 7: 266. 1891 – Lectotype 00073248 [online image!], K barcode K000328989!,
(designated here): Vincent. s.d., Guilding, L. s.n. NY barcodes 00581298! & 00581299!, RAS barcode
(K barcode K000328993!). RSA0004261 [online image!], US barcodes 00679974!
& 00680010!).
20. Chaetogastra citrina (Naudin) P.J.F.Guim. & Michelang.,
comb. nov. ≡ Micranthella citrina Naudin in Ann. Sci. Nat., 25. Chaetogastra cordeiroi F.S.Mey. & R.Goldenb. in Phyto-
Bot., sér. 3, 13(6): 350. 1850 ≡ Tibouchina citrina (Naudin) taxa 282(4): 240–244, fig. 2, 3. 2016 – Holotype: BRA-
Cogn. in Candolle & Candolle, Monogr. Phan. 7: 253. 1891 ZIL. Santa Catarina: Campo Alegre, Serra Quiriri, 7 Apr
– Holotype: BOLIVIA. Larecaja et Caupolican,vallées entre 2009, Cordeiro, J. & Barbosa, E. 3028 (MBM; iso-
Tipoani et Apolobamba, atl. 3000 m, May 1847, Weddell, type: ASE).
H.A. s.n. [4551] (P barcode P00121042!; isotypes: BR
barcode BR0000005520299 [fragment]!, P barcodes 26. Chaetogastra cornuta (Gleason) P.J.F.Guim. & Miche-
P00121043!, P00121044! & P00121045!). lang., comb. nov. ≡ Tibouchina cornuta Gleason in Bull.
Torrey Bot. Club 66(6): 415. 1939 – Holotype: BOLIVIA.
21. Chaetogastra clinopodifolia DC., Prodr. 3: 133. 1828 ≡ Dep. La Paz: prov. Sur Yungas, colaya, understory virgin
Micranthella clinopodifolia (DC.) Naudin in Ann. Sci. forest at Colaya, alt. 1870 m, 13 Dec 1935, Mexia,
Nat., Bot., sér. 3, 13(6): 348. 1850 ≡ Pleroma clinopodifo- Y.E.J. 4296 (GH barcode 00073258 [online image!]).
lium (DC.) Triana in Trans. Linn. Soc. London, Bot.
28(1): 46. 1872 (“1871”) ≡ Tibouchina clinopodifolia 27. Chaetogastra crassifolia F.S.Mey. & R.Goldenb. in Phy-
(DC.) Cogn. in Martius, Fl. Bras. 14(3): 411–412. 1885 totaxa 282(4): 244–249, fig. 4–6. 2016 – Holotype: BRA-
– Holotype: BRAZIL. Rio de Janeiro: in sylvis prope ZIL. Paraná: Campina Grande do Sul, Serra do Capivari,
S. Cruz ad Serram prov. Sebastianapolitanae, 1827, Mar- trilha que segue para o Capivari Grande, 2 Mar 2013,
tius, C.F.P. s.n. (M barcode M-0165831!; isotype: G-DC Meyer, F.S.; Schmitz, F. & Sade, G. 1620 (UEC; isotypes:
barcode G00318939!). MBM, NY, UPCB).
= Tibouchina clinopodifolia var. rurikiana Cham. in Linnaea
9(4): 450. 1835, syn. nov. – Lectotype (designated 28. Chaetogastra cristaensis F.S.Mey & R.Goldenb. in
here): BRAZIL. Santa Catarina, Dec 1815, Sellow, F. Phytotaxa 282(4): 249–253, fig. 7–9. 2016 – Holotype:
s.n. (K barcode K000329106!). BRAZIL. Santa Catarina: Garuva, Serra do Quiriri,
Pedra Cabeluda, Arredores do Monte Crista, 17 Mar
22. Chaetogastra confertiflora (Naudin) P.J.F.Guim. & 2013, Meyer, F.S. 1682 (UEC; isotypes: FURB,
Michelang., comb. nov. ≡ Micranthella confertiflora MBM, UPCB).
Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(6): 349–350.
1850 ≡ Tibouchina confertiflora (Naudin) Cogn. in 29. Chaetogastra debilis Cham. in Linnaea 9: 499. 1835 ≡
Candolle & Candolle, Monogr. Phan. 7: 253. 1891 – Pleroma debile (Cham.) Triana in Trans. Linn. Soc.
Holotype: BOLIVIA. Cupe-Yungas, s.d., D’Orbigny, London, Bot. 28(1): 45 8. 1872 (“1871”) ≡ Tibouchina
A.C.V.M.D. 415 (P barcode P00708854!). debilis (Cham.) Cogn. in Martius, Fl. Bras. 14(3):
401–402. 1885 – Lectotype (designated here): BRA-
23. Chaetogastra congestiflora (Todzia) P.J.F.Guim. & Miche- ZIL. s.d., Sellow, F. 4503 (F barcode V0063448F [online
lang., comb. nov. ≡ Tibouchina congestiflora Todzia in image!]; isolectotypes: B [probably destroyed; F negative
Brittonia 51(3): 259–262, fig. 3, 12. 1999 – Holotype: F0BN016741!], K barcodes K000329095 [no collector
MEXICO. Guerrero: Eastern slope of Cerro Alquitran, number]! & K000329096 [no collector number]!, P bar-
5–7 km (straight line distance) WSW of Mazatlan, code P00708983 [no collector number]!).
2450 m, 11 Feb 1970, Anderson, W.R. & Anderson, C. 5721
(US barcode 00679976!; isotypes: ENCB barcode 30. Chaetogastra decora (Gleason) P.J.F.Guim. & Miche-
ENCB008688 [online image!], MEXU barcode MEXU lang., comb. nov. ≡ Tibouchina decora Gleason in Bull.
00858980 [online image!], MICH barcode 1210031 [online Torrey Bot. Club 58(4): 218–219. 1931 – Holotype:
image!], TEX barcode 00374500 [online image!]). PERU. Dep. Ayacucho: Aina, between Huanta and Río
Apurimac, alt. 750–1000 m, 7 May 1929, Killip,
24. Chaetogastra connata (Gleason ex Todzia) P.J.F.Guim. E.P. & Smith A.C. 22700 (NY barcode 00245758!; iso-
& Michelang., comb. nov. ≡ Tibouchina connata Gleason types: K barcode K000329166!, US barcode 00120048!).
ex Todzia in Brittonia 51(3): 262, fig. 4, 13. 1999 – Holo-
type: MEXICO. Guerrero: Distr. Mina, Laguna- 31. Chaetogastra dimorphylla (Gleason) P.J.F.Guim. &
Tequeches, 1740 m, 12 Mar 1936, Hinton, G.B. 9952 Michelang., comb. nov. ≡ Tibouchina dimorphylla
(MEXU barcode MEXU 00100349 [online image!]; Gleason in Bull. Torrey Bot. Club 54(1): 27–28. 1927 –

23
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

Holotype: PERU. Dep. Cuzco: Cerro de Cusilluyoc, in a Jahrb. Syst. 50(2–3, Beibl. 111): 31. 1913 – Lectotype
thicket in a valley on the páramo above “Pillahuata”, alt. (designated here): Dep. Ayacucho: prov. Huanta, “ad
3000–3300 m, 3–6 May 1925, Pennell, F.W. 14109 viam a Tambo ultra Osno ad flum”, Apurimac, La Mar
(NY barcode 00245759!; isotypes: F barcode Locality between Tambo and the Apurimac, Yana-
V0040210F [online image!], GH barcode 00073259 monte alt. 2800 m, Jun 1910, Weberbauer, A. 5619 (BR
[online image!], PH barcode 00027248 [online image!], barcode BR0000005225200!; isolectotypes: F barcode
S No. S-R-6114!, US barcode 00120050!). V0040211F [online image!], GH barcode 00073261
[online image!], US barcode 00120056!).
32. Chaetogastra durangensis (Standl.) P.J.F.Guim. & = Tibouchina fulvipilis var. scrobiculata Cogn. in Bot. Jahrb.
Michelang., comb. nov. ≡ Tibouchina durangensis Syst. 50(2–3, Beibl. 111): 32. 1913, syn. nov. – Lecto-
Standl. in Contr. U.S. Natl. Herb. 23(4): 1054. 1924 – type (designated here): Dep. Ayacucho: prov. Huanta,
Holotype: MEXICO. San Ramón: Durango, 21 Apr–18 alt. 2600–2800 m, Jun 1910, Weberbauer, A. 5625
May 1906, Palmer, E. 163 (US barcode 00681573!; iso- (BR barcode BR0000005224555!; isolectotype: F bar-
types: GH barcode 00073249 [online image!], NY bar- code V0040212F [online image!], GH barcode
code 00581297!). 00073262 [online image!], US barcode 00120057!).

33. Chaetogastra echinata (Ruiz & Pav.) DC., Prodr. 3: 135. 38. Chaetogastra galeottiana (Naudin) P.J.F.Guim. & Miche-
1828 ≡ Rhexia echinata Ruiz & Pav., Fl. Peruv. 3: 85. lang., comb. nov. ≡ Oreocosmus galeottianus Naudin in
1802 ≡ Pleroma echinatum (DC.) Triana in Trans. Linn. Ann. Sci. Nat., Bot., sér. 3, 13(1): 38–39. 1850 ≡ Tibou-
Soc. London, Bot. 28(1): 47. 1872 (“1871”), nom. illeg., china galeottiana (Naudin) Cogn. in Candolle & Can-
non Gardner 1843 ≡ Tibouchina echinata (Ruiz & Pav.) dolle, Monogr. Phan. 7: 258. 1891 ≡ Tibouchina
Cogn. in Candolle & Candolle, Monogr. Phan. 7: 266. trianaei Cogn. in Candolle & Candolle, Monogr. Phan.
1891 – Lectotype (designated here): PERU. Muña, 7: 261. 1891, pro parte, nom. illeg. superfl. – Holotype:
s.d., Pavón s.n. (G! [F negative F0BN026105]; isolecto- Cordillera, Mountains E. of Oaxaca, 1840, Galeotti, H.
tyes: BR barcode BR0000005521005!, G-DC barcode 2935 (P barcode P00708796; isotypes: BR barcodes
G00310480 [fragment]!). BR0000005224937 & BR0000005225323, K barcode
K000328981!).
34. Chaetogastra erioclada (Triana) P.J.F.Guim. & Miche-
lang., comb. nov. ≡ Pleroma eriocladum Triana in Trans. 39. Chaetogastra gayana (Naudin) P.J.F.Guim. & Miche-
Linn. Soc. London, Bot. 28(1): 47. 1872 (“1871”) ≡ lang., comb. nov. ≡ Micranthella gayana Naudin in
Tibouchina erioclada (Triana) Cogn. in Candolle & Can- Ann. Sci. Nat., Bot., sér. 3, 13(6): 350. 1850 ≡ Tibouchina
dolle, Monogr. Phan. 7: 251. 1891 – Lectotype (desig- gayana (Naudin) Cogn. in Candolle & Candolle, Monogr.
nated here): COLOMBIA. Prov. Pasto: Tuquerres, alt. Phan. 7: 253. 1891 – Holotype: PERU. Cuzco, s.d.,
3000 m, May 1853, Triana, J.J. 3901 (COL barcode Gay, C. s.n. (P barcode P00708741!; isotype: BR barcode
COL000003426 [online image!]; isolectotypes: BM bar- BR0000005520220 [fragment]!).
code BM000953946!, BR barcode BR0000005521012!,
COL barcode COL000011608 [online image!], G-DC 40. Chaetogastra geitneriana Schltdl. in Linnaea 27(5): 523.
barcode G00319000!, K barcode K000329148!). 1854 ≡ Tibouchina geitneriana (Schltdl.) Cogn. in Can-
dolle & Candolle, Monogr. Phan. 7: 255. 1891 – Type:
35. Chaetogastra excoriata (Cogn.) P.J.F.Guim. & Miche- VENEZUELA. Geitner s.n. (W [not found]).
lang., comb. nov. ≡ Tibouchina excoriata Cogn. in Bull. = Tibouchina schumannii Cogn. in Candolle & Candolle,
New York Bot. Gard. 4(14): 356. 1907 – Holotype: Monogr. Phan. 7: 256. 1891 – Holotype: VENEZUELA.
BOLIVIA. “Epiphytisch an der Baumgrenze bei Tres Cru- Caracas ad Tovar, Dec 1848, Karsten, H. s.n.
ces”, 1500–1700 m, s.d., Bang, M. 1992 (NY barcode (B [probably destroyed; F negative F0BN016794!]; iso-
00245764!; isotypes: BR barcode BR0000005530632 type: BR barcode BR0000005225248!).
[fragment]!, US barcode 00120054!). = Tibouchina moritziana Klotzsch ex Cogn. in Candolle
& Candolle, Monogr. Phan. 7: 259. 1891 – Lectotype
36. Chaetogastra ferrariana (Cogn.) P.J.F.Guim. & Miche- (designated here): VENEZUELA [recorded as “Colum-
lang., comb. nov. ≡ Tibouchina ferrariana Cogn. in Can- bien”]. Mérida, 5 Nov 1844, Moritz, J.W.K. 937
dolle & Candolle, Monogr. Phan. 7: 1176. 1891 – (K barcode K000329154!; isolectoypes B [probably
Lectotype (designated here): MEXICO. Teziutlan, destroyed; F negative F0BN016769!], BR barcode
Nov 1884, Comisión Geográfica Exploradora 409 BR0000005225224!, K barcode K000329153!, NY
(BR barcode BR0000005224883!). barcode NY00245787!, P barcodes P00708723! &
P00708724!, US barcode 00604114!).
37. Chaetogastra fulvipilis (Cogn.) P.J.F.Guim. & Miche- Other syntypes: VENEZUELA. Caracas: 1854–5,
lang., comb. nov. ≡ Tibouchina fulvipilis Cogn. in Bot. Fendler, A. 404 part. (K barcode K000329158!);

24
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

VENEZUELA. s.d., Moritz, J.W.K. 291 (E barcode = Tibouchina gracilis var. australis Cogn. in Martius, Fl. Bras.
E00531231 [online image!], K barcodes K000329156! 14(3): 388. 1885, syn. nov. – Holotype: PARAGUAY.
& K000329157!, P barcodes P00708721! & P0070 Caaguazú, 16 Nov 1874, Balansa, B. 2566 (P barcode
8722!, US barcode 00604144!); VENEZUELA. s.d., P00708746!).
Karsten, H. s.n. (not located); VENEZUELA. Caracas, = Tibouchina gracilis var. chamissoana Cogn. in Martius,
s.d., Vargas, J.M. 147 (G-DC barcode G00318989!). Fl. Bras. 14(3): 388. 1885, pro parte, syn. nov. – Lecto-
= Tibouchina pseudomollis Gleason in Bull. Torrey Bot. Club type (designated here): BRAZIL. s.d., Sellow, F. 3657
56(8): 400. 1929 – Holotype: Guyana. Mt. Roraima, (BR barcode BR0000005520237!).
sandstone bouldres at Philipp Camp, alt. 5200–6000 ft., = Tibouchina gracilis var. longisetosa Cogn. in Martius,
7 Nov 1927, Tate, G.H.H. 275 (NY barcode 00245797!). Fl. Bras. 14(3): 388. 1885 – Lectotype (designated
= Tibouchina impressa Gleason in Fieldiana, Bot. 28(2): here): BRAZIL. s.d., Sellow, F. 1216 (K barcode
428–429. 1952 – Holotype: VENEZUELA. Anzoátegui: K000329073!).
on south-facing steep slopes, on rocky exposed shrubby = Tibouchina gracilis var. paucisetosa Cogn. in Martius,
portion of knife-like ridge above tree zone, Certo Peonía Fl. Bras. 14(3): 388. 1885, syn. nov. – Lectotype (desig-
(Cerro Los Pajaritos), above Santa Clara, alt. nated here): BRAZIL. Minas Gerais: 1842, Claussen, P.
1800–2000 m, 20 Mar 1945, Steyermark, J.A. 61616 102 (P barcode P05200653!).
(NY barcode 00245771!; isotypes: F barcode V0063985F Other syntypes: BRASIL. São Paulo: s.d., Sellow, F. D n
[online image!], VEN barcode 33981 [online image!]). 3204 (P barcode P00708742! [s.l., s.n.]); BRASIL. São
Paulo: s.d., Sellow, F. D n 3657 part. (BR barcode
41. Chaetogastra gracilis (Bonpl.) DC., Prodr. 3: 133. 1828 ≡ BR0000005224623!); BRAZIL. Rio Grande do Sul: s.d.,
Rhexia gracilis Bonpl. in Humboldt & Bonpland, Isabelle 6 (P barcode P05200617!).
Monogr. Melast. 2: 138, pl. 52. 1823 ≡ Lasiandra gracilis = Tibouchina gracilis var. grandiflora Cogn. in Bull. Herb.
(Bonpl.) Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): 159. Boissier, sér. 2(4): 1276. 1904, syn. nov. Type – PARA-
1850 ≡ Pleroma gracile (Bonpl.) A.Gray, U.S. Expl. GUAY. Campo pr. Cerro hu, Sep, Hassler, E. 977a (not
Exped., Phan. 15: 604. 1854 ≡ Tibouchina gracilis located).
(Bonpl.) Cogn. in Martius, Fl. Bras. 14(3): 386–389. = Tibouchina gracilis var. robusta Cogn. in Feddes Repert.
1885 – Lectotype (designated here): [illustration] Spec. Nov. Regni Veg. 7(4/6): 72. 1909, syn. nov. – Lec-
“RHEXIA gracilis” in Humboldt & Bonpland, Monogr. totype (designated here): PARAGUAY. Sierra de
Melast. 2: 138, pl. 52. 1823. Amambay, in campis altis ad Punta Porá, Dec,
= Chaetogastra fraterna DC., Prodr. 3: 133. 1828 ≡ Tibou- Hassler, E. 9883 (NY barcode 00245766!).
china gracilis var. fraterna (DC.) Cogn. in Martius,
Fl. Bras. 14(3): 387. 1885, syn. nov. – Lectotype (desig- 42. Chaetogastra grossa (L.f.) P.J.F.Guim. & Michelang.,
nated here): BRAZIL. Minas Gerais, 1833, comb. nov. ≡ Melastoma grossum L.f., Suppl. Pl.: 236.
Gaudichaud, C. s.n. (P barcode P005200634!). 1782 (“1781”) ≡ Purpurella grossa (L.f.) Triana in Trans.
= Chaetogastra hirsuta DC., Prodr. 3. 133. 1828 ≡ Pleroma Linn. Soc. London, Bot. 28(1): 48. 1872 (“1871”) ≡
gracile var. hirsuta Triana in Trans. Linn. Soc. London, Tibouchina grossa (L.f.) Cogn. in Martius, Fl. Bras.
Bot. 28(1): 45. 1872 (“1871”) ≡ Tibouchina gracilis var. 14(3): 297. 1885 – Holotype: COLOMBIA. Tortolito,
hirsuta (Triana) Cogn. in Martius, Fl. Bras. 14(3): 388. s.d., Mutis, J.C. 2730 (LINN No. 559.13 [online image!];
1885, syn. nov. – Lectotype (designated here): BRA- isotype: US barcode 00120453!).
ZIL. s.d., Raddi s.n. (G-DC barcode G00310488 [frag- = Rhexia muricata Bonpl. in Humboldt & Bonpland, Monogr.
ment, no collection data]!). Melast. 2: 1, pl. 1. 1806 ≡ Chaetogastra muricata
= Chaetogastra strigillosa Mart. ex DC., Prodr. 3: 133. 1828 (Bonpl.) DC., Prodr. 3: 134. 1828 ≡ Purpurella muricata
≡ Tibouchina gracilis var. strigillosa Cogn. in Martius, (Bonpl.) Naudin in Ann. Sci. Nat., Bot., sér. 3, 13: 301.
Fl. Bras. 14(3): 387. 1885, syn. nov. – Lectotype (desig- 1850 – Lectotype (designated here): COLOMBIA.
nated here): BRAZIL. Minas Gerais: “habitat in campis Andes du Pérou. Croît entre la ville de Popayan et le vol-
ad latera montinum prope Vam. S. Joannis Regiano”, can Puracé, à 3250 mètres de hauteur sur le niveau de le
s.d., Martius, C.F.P. von s.n. (M barcode M-0214978!; mer. “Crescit in frigidis montis Puracé prope Popayan,
isolectotype: G-DC barcode G00310486 [fragment]!). alt. 1600 m.; floret Decembri”, s.d., Bonpland, A.J.A.
= Tibouchina gracilis var. gracillima Cogn. in Martius, s.n. (P barcode P00136470!; isolectotypes: P barcodes
Fl. Bras. 14(3): 387. 1885, syn. nov. – Lectotype (desig- P00136471! & P00136472!, US barcodes 00120184!
nated here): BRAZIL. s.d., Sellow, F. s.n. (BR barcode & 01094678!).
BR000000552054!; isolectotype: HBG barode HBG- = Rhexia reticulata Bonpl. in Humboldt & Bonpland,
523024 [online image!]). Monogr. Melast. 2: 21, pl. 9. 1808 ≡ Chaetogastra reticu-
= Tibouchina gracilis var. vulgaris Cogn. in Martius, Fl. Bras. lata (Bonpl.) DC., Prodr. 3: 131. 1828 ≡ Purpurella reti-
14(3): 386. 1885, nom. illeg. superfl. [includes the type of culata (Bonpl.) Naudin in Ann. Sci. Nat., Bot., sér. 3, 13:
Rhexia gracilis Bonpl., Art. 6.8 & 32.3]. 302. 1850 ≡ Tibouchina reticulata (Bonpl.) Cogn. in

25
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

Martius, Fl. Bras. 14(3): 298. 1885 – Lectotype (desig- 45. Chaetogastra herincquiana (Cogn.) P.J.F.Guim. &
nated here): COLOMBIA. In montis Saraguru frigidis, Michelang., comb. nov. ≡ Tibouchina herincquiana
prope Loxam, alt. 1060 m; floret Septembri, s.d., Bon- Cogn. in Martius, Fl. Bras. 14(3): 399–400, t. 90, fig.
pland, A.J.A. s.n. (P barcode P00136465; isolectotypes: 3. 1885 – Holotype: BRAZIL. São Paulo: 1816, De
F barcode V0063950F [online image!]; P barcodes Saint-Hilaire, A.F.C.P. C2 1412 (P barcode P00708829!
P00136466! & P00136467!). [F negative F0BN036165!]; isotype: BR! barcode
BR0000005520909 [fragment]!).
43. Chaetogastra hassleri (Cogn.) P.J.F.Guim. & Michelang.,
comb. nov. ≡ Tibouchina hassleri Cogn. in Bull. Herb. 46. Chaetogastra herzogii (Cogn.) P.J.F.Guim. & Miche-
Boissier, sér. 2, 4: 1276–1277. 1904 – Lectotype (desig- lang., comb. nov. ≡ Tibouchina herzogii Cogn. in Meded.
nated here): PARAGUAY. In campo humido Ipe hu, Rijks-Herb. 19: 78. 1913 – Lectotype (designated here):
Sierra de Maracayú, Oct 1898, Hassler, E. 5196 BOLIVIA. An steinigen Stellen des “Meson” bei Samai-
(BR barcode BR0000005520565!; isolectotypes: B [prob- pata, 2100 m, Mar 1911, Herzog, T.C.J. 1765 (JE barcode
ably destroyed; F negative F0BN016750!], GH barcode JE00000961 [online image!]; isolectotypes: B [prob-
00073264 [online image!], K barcode K000329116!, ably destroyed; F negative F0BN016751!], BR
MPU barcode MPU013229 [online image!], NY barcode BR0000005520572 barcode [fragment]!, S No. S-R-
00245770!, P barcodes P00708956! & P00708957!, UC 6119 [online image!]).
barcode UC935016 [online image!]).
= Tibouchina guimaraensii Brade in Arch. Jard. Bot. Rio de 47. Chaetogastra hieracioides Schrank & Mart. ex. DC.,
Janeiro 14: 218–219, t. 5. 1956, syn. nov. – Holotype: Prodr. 3: 133. 1828 ≡ Lasiandra hieracioides (Schrank
BRAZIL. Paraná: Guarapuava, brejo no campo, 1000 m & Mart. ex. DC.) Naudin in Ann. Sci. Nat., Bot., sér.
a.s.l., 16 Feb 1949, Brade, A.C. 19555 (RB barcode 3, 13(3): 127. 1850 ≡ Pleroma hieracioides (Schrank
00662346!; isotype: RB barcode 00541560!, K barcode & Mart. ex. DC.) Triana in Trans. Linn. Soc. London,
K001089090!, NY barcode 01477584!, SP barcode Bot. 28(1): 45. 1872 (“1871”) ≡ Tibouchina hieracioides
SP002916 [online image!]). (Schrank & Mart. ex. DC.) Cogn. in Martius, Fl. Bras.
14(3): 389–390, t. 90, fig. 1. 1885 – Holotype: BRAZIL.
44. Chaetogastra herbacea (DC.) P.J.F.Guim. & Michelang., Minas Gerais: Serra de Itambé et S. de Montanha, campis
comb. nov. ≡ Arthrostemma herbaceum DC., Prodr. 3: altis, montium e saxo arenaris constantium prope vam de
137. 1828 ≡ Pterolepis herbacea (DC.) Triana in Trans. Campanha & alibi, soli duri incola, 1827, Martius, C.F.P.
Linn. Soc. London, Bot. 28(1): 39. 1872 (“1871”) ≡ von #Obs. 743 (M barcode M-0165838!; isotype: G-DC
Tibouchina herbacea (DC.) Cogn. in Martius, Fl. Bras. barcode G00310487!).
14(3): 408–409. 1885 – Holotype: BRAZIL. Sebastiana-
politanae et prope Minas Gerais: in sylvis caeduis locis 48. Chaetogastra hintonii (Gleason ex Todzia) P.J.F.Guim.
sicciusculis umbrosis prope Rio de Janeiro, Nov, Martius, & Michelang., comb. nov. ≡ Tibouchina hintonii Gleason
C.F.P. von 794 (M barcode M-0165836! [F negative ex Todzia in Brittonia 51(3): 264–267, fig. 5, 14. 1999 –
F0BN06351!]). Holotype: MEXICO. Guerrero: distr. Galeana, San Juan
= Arthrostemma hirsutissimum DC., Prodr. 3: 137. 1828 ≡ de las Flores, 700 m, 9 Dec 1937, Hinton, G.B. & al.
Tibouchina herbacea var. hirsutissima (DC.) Cogn. in 11015 (US barcode 00679975!; isotypes: BM barcode
Martius, Fl. Bras. 14(3): 409. 1885, syn. nov. – Type: BM000795740!, CAS barcode 0004852 [online image!],
BRAZIL. s.d., Raddi s.n. (not located). GBH barcode GBH011015 [online image!], GH barcode
= Tibouchina herbacea var. rosea Cogn. in Martius, Fl. Bras. 00073250 [online image!], K barcode K000328990!, NY
14(3): 409. 1885, syn. nov. – Lectotype (designated barcode 00581291!, RSA barcode RSA0004262 [online
here): PARAGUAY. Caaguazú, dans les prairies image!], US barcode 00680009!).
humides, 20 Mar 1876, Balansa, B. 2564 (P barcode
P00708694!). 49. Chaetogastra hutchisonii (Wurdack) P.J.F.Guim. &
Other syntype: PARAGUAY: Caaguazú, dans les marais, Michelang., comb. nov. ≡ Tibouchina hutchisonii
Apr 1876, Balansa, B. 2565 (P barcode P00708695!). Wurdack in Phytologia 14(5): 258–259. 1967 – Holo-
= Tibouchina herbacea f. grandifolia Cogn. in Bull. Herb. type: PERU. Dep. San Martín: wet steep banks at lower
Boissier, sér. 2, 4: 1277. 1904, syn. nov. – Lectotype Boquerón del Padre Abad, elev. 480 m, 25 Jul 1964,
(designated here): PARAGUAY. In campo Cordillera Hutchison, P.C.; Wright, J.K. & Straw, R.M. 6034
de Altos, Jul 1885, Hassler, E. 471 (K barcode (US barcode 00120062!; isotypes: UC, USM).
K000880905!).
= Tibouchina herbacea F. parvifolia Cogn. in Bull. Herb. 50. Chaetogastra incarum (Gleason) P.J.F.Guim. & Miche-
Boissier, sér. 2, 4: 1277. 1904, syn. nov. – Type PARA- lang., comb. nov. ≡ Tibouchina incarum Gleason in
GUAY. In palude pr. San Bernadino, Apr, Hassler, E. Amer. J. Bot. 26(8): 634. 1939 – Holotype: PERU. Near
4067 (not located). Cuzco, Machu Picchu, in shaded moist hedgerows, alt.

26
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

5000 ft., s.d., Balls, E.K. B-6817 (NY barcode 55. Chaetogastra lancifolia (Wurdack) P.J.F.Guim. & Miche-
00245772!; isotypes: CAS barcode 0004853 [online lang., comb. nov. ≡ Tibouchina lancifolia Wurdack in
image!], K barcode K000329168!, UC barcode Phytologia 5(2): 54–55. 1954 – Holotype: BOLIVIA.
UC683186 [online image!], US barcodes 00120063!, Cochabamba. Incachaca about 130 km, northeast of
00921433! & 00921434!). Cochabamba, elev. 1500 m, 20 Aug 1950, Brooke,
W.M.A. 6767 (NY barcode 00245777!; isotype: BM bar-
51. Chaetogastra inconstans (Vahl) P.J.F.Guim. & Miche- code BM000953942!).
lang., comb. nov. ≡ Rhexia inconstans Vahl, Eclog.
Amer. 1: 37. 1797 – Lectotype (designated here): GUA- 56. Chaetogastra latibracteolata (Paul G. Wilson) P.J.F.
DELOUPE. s.d., s.c. s.n. (LINN-HS No. 654.8 [online Guim. & Michelang., comb. nov. ≡ Tibouchina latibrac-
image!]). teolata Paul G. Wilson in Kew Bull. 13(1): 160–161.
= Rhexia strigosa Rich. in Actes Soc. Hist. Nat. Paris 1: 108. 1958 – Holotype: MEXICO. Chorrera: district of Temas-
1792 ≡ Chaetogastra strigosa (Rich.) DC., Prodr. 3. 134. caltepec; Chorrera, 26 Dec 1934, Hinton, G.B. & al. 7165
1828 ≡ Hephestionia strigosa (Rich.) Naudin in Ann. Sci. (K barcode K000328991!; isotypes: BM barcode
Nat., Bot., sér. 3, 13(1): 36–37. 1850 ≡ Tibouchina stri- BM000795747!, F barcode V0063973F [online image!],
gosa (Rich.) Cogn. in Candolle & Candolle, Monogr. GBH barcode GBH007165 [online image!], K barcode
Phan. 7: 266. 1891 – Holotype: Guadeloupe. s.d., Rich- K000328992!, MICH barcode 1210042 [online image!],
ard, L.C. s.n. (P barcode 00708783!; isotype: P barcode NY barcodes 00581283! & 00581284!, P barcode
00708784!). P00116890!, US barcode 00681568!).
= Melastoma ciliatum Desr. in Lamarck, Encycl. 4: 49. 1797,
nom. illeg., non Rich. 1792 – Holotype: Guadeloupe, s.d., 57. Chaetogastra latifolia (Naudin) P.J.F.Guim. & Miche-
Badier, Mr. de s.n. (P [not found]). lang., comb. nov. ≡ Micranthella latifolia Naudin in
= Osbeckia ornata Sw., Fl. Ind. Occid. 2: 647. 1798 – Lecto- Ann. Sci. Nat., Bot., sér. 3, 13(6): 351–352. 1850 ≡ Ple-
type (designated here): Guadeloupe. s.d., De Ponthieu, roma latifolium (Naudin) Triana in Trans. Linn. Soc.
H. s.n. (BM barcode BM000795772!). London, Bot. 28(1): 47. 1872 (“1871”) ≡ Tibouchina lati-
folia (Naudin) Britton in Bull. Torrey Bot. Club 17(3):
52. Chaetogastra kingii (Wurdack) P.J.F.Guim. & Miche- 54–55. 1890 – Holotype: BOLIVIA. Tipoani, alt.
lang., comb. nov. ≡ Tibouchina kingii Wurdack in Phyto- 2500 m, s.d., Weddell, H.A. 4535 (P barcode
logia 19(3): 191. 1969 – Holotype: COLOMBIA. Dep. P00708962!; isotypes: F barcode V0063878F [fragment]
Tolima: about 18 km west-southwest of Fresno, Cordil- [online image!], P barcode P00708963!).
lera Central, elev. 2150 m, 16–17 Jul 1965, King, R.M.;
Guevara, A.E. & Forero, E. 5999 (US barcode 58. Chaetogastra laxa (Desr.) P.J.F.Guim. & Michelang.,
00120065!; isotypes: COL barcode COL000003428 comb. nov. ≡ Melastoma laxum Desr. in Lamarck,
[online image!], F barcode V0063987F [online image!], Encycl. 4: 41. 1797 ≡ Tibouchina laxa (Desr.) Cogn. in
NY barcode 00245773!). Bull. Acad. Roy. Sci. Belgique, sér. 3, 14(7): 933. 1887
– Lectotype (designated here): PERU. s.d., Jussieu, J.
53. Chaetogastra kunhardtii (Gleason) P.J.F.Guim. & de s.n. (P barcode P00708964!; isolectotype: herbarium
Michelang., comb. nov. ≡ Tibouchina kunhardtii Glea- unknown [F negative F0BN036166!]).
son in Phytologia 3(5): 242–243. 1950 – Holotype: VEN- = Rhexia sarmentosa Bonpl. & Rich. in Humboldt & Bonpland,
EZUELA. Amazonas: on the summit of Cerro Sipapo, Monogr. Melast. 2: 23, pl. 10. 1806 ≡ Chaetogastra sarmen-
Cano, 15 Dec 1948, Maguire, B. & Politi, P. 27713 tosa (Bonpl.) DC., Prodr. 3: 134. 1828 ≡ Lasiandra sarmen-
(NY barcode 00245775!; isotypes: BM barcode tosa (Bonpl.) Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3):
BM000953943!, F barcode V0063988F [online image!], 130–131. 1850 – Holotype: PERU. Crescit in calidis Regni
MO barcode MO-313919 [online image!], RB barcode Quitensis, inter urbem Cuenca et pagum San Felipe, alt.
00541561!, US barcode 00120067!, VEN barcode 980–1400 hex, s.d., Bonpland, A.J.A. & Humboldt,
31436 [online image!]). F.W.H.A. von s.n. (P barcode P00136475!; isotypes: P bar-
codes P00708964! & P00708965!).
54. Chaetogastra laevis (Cogn.) P.J.F.Guim. & Michelang., = Tibouchina cymosa Cogn. in Bot. Jahrb. Syst. 42: 134–135.
comb. nov. ≡ Tibouchina laevis Cogn. in Bot. Jahrb. Syst. 1908 – Lectotype (designated by Burke & Michelangeli
42: 136. 1908 – Lectotype (designated by Burke & in Phytotaxa 125(1): 12. 2013): PERU. Dep. Cajamarca:
Michelangeli in Phytotaxa 125(1): 12. 2013): PERU. Ad Prov. Hualgayoc, infra San Miguel, in fruticeto ad rivum
viam a Sandia ad Chunchusmayo, inter Tambo Yunca- 2500–2600 m a.s.l., 5 May 1904, Weberbauer, A. 3911
coya et Tambo Cachicachi, in fruticetis, 1800–2200 m (BR barcode BR0000005224074!; isolectotype: B [prob-
a.s.l., 8 Jun 1902, Weberbauer, A. 1139 (BR barcode ably destroyed; F negative F0BN016740!]).
BR0000005224067; isolectotype: B [probably destroyed; = Tibouchina asperipilis S.F.Blake in Proc. Biol. Soc. Wash.
F negative F0BN016757!]). 35: 117. 1922 – Holotype: ECUADOR. Loja: open places

27
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

on hillsides, elev. 1400 ft., 8 Apr 1921, Popenoe, W. 1313 62. Chaetogastra longistyla (Cogn.) P.J.F.Guim. & Miche-
(US barcode 00120038!). lang., comb. nov. ≡ Pterolepis longistyla Cogn. in Mar-
tius, Fl. Bras. 14(3): 283–284, t. 65, fig. 2. 1885 ≡
59. Chaetogastra longifolia (Vahl.) DC., Prodr. 3: 132. 1828 Tibouchina longistyla (Cogn.) Renner in Nordic J. Bot.
≡ Rhexia longifolia Vahl., Eclog. Amer. 1: 39–40. 1797 ≡ 14(1): 102. 1994 – Lectotype (designated here): BRA-
Pleroma longifolium (Vahl.) Triana in Trans. Linn. Soc. ZIL. São Paulo: Cajuni, 8 Mar 1857, Regnell, A. III 15
London, Bot. 28(1): 45. 1872 (“1871”) ≡ Tibouchina (BR barcode BR0000005223442!; isolectotypes: S. No.
longifolia (Vahl.) Baill. in Adansonia 12: 74. 1877 – S-R-5379!, US barcode 01903133!).
Holotype: America meridionali, s.d., Rohr, J.P.B. von 15 Other syntype: BRAZIL. Minas Gerais: Furnas [near bor-
(C barcode C10015075 [online image!]). der to S. Paulo State], Mar 1826, Langsdorff, G.H. von
= Chaetogastra depauperata Naudin in Ann. Sci. Nat., Bot., 2073 (LE!).
sér. 3, 14(2): 129–130. 1850 ≡ Tibouchina longifolia
var. depauperata (Naudin) Kuntze, Revis. Gen. Pl. 63. Chaetogastra macvaughii (Todzia) P.J.F.Guim. & Miche-
1: 248. 1891, syn. nov. – Holotype: BOLIVIA. s.d., lang., comb. nov. ≡ Tibouchina macvaughii Todzia in
d’Orbigny, A.D. s.n. (P barcode P00708973!). Brittonia 51(3): 267–269, fig. 15. 1999 – Holotype:
= Tibouchina bourgaeana Cogn. ex Donn.Sm. in Bot. Gaz 16: MEXICO. Jalisco: steep mountains 20–22 km south of
4. 1891 – Lectotype (designated here): GUATEMALA. Talpa de Allende, in the headwaters of a west branch of
Dep. Yzabal: alt. 100 ft., Apr 1889, Donnell Smith, J. Rio de Talpa, 1200–1450 m, 28–30 Mar 1965,
1514 (GH barcode 00073247 [online image!]). McVaugh, R.; Baad, M.F.; Anderson, W.R. & Laskowski,
= Tibouchina aliena Brandegee in Univ. Calif. Publ. Bot. 6(4): C.W. 23292 (MICH barcode 1210032 [online image!];
58. 1914 – Holotype: MEXICO. Chiapas: Cerro del isotypes: ENCB barcode ENCB008690 [online image!],
Boquerón, s.d., Purpus, C.A. 6772 (UC barcode US barcode 00679972!).
UC172457 [online image!]; isotypes: F barcode
V0063968F [online image!], NY barcode 00581295!). 64. Chaetogastra mariae (Wurdack) P.J.F.Guim. & Miche-
= Tibouchina longifolia var. simulans in Publ. Field Colum- lang., comb. nov. ≡ Tibouchina mariae Wurdack in Phy-
bian Mus., Bot. Ser. 4(7): 174. 1929, syn. nov. – Holo- tologia 13(2): 65–67. 1966 – Holotype: PERU. Dep.
type: PERU. Dep. Junín: La Merced, 10–24 Aug 1923, Amazonas: prov. Chachapoyas, Quebrada Molina 5 km
Macbride, J.F. 5300 (F barcode V0040213F [online below Chachapoyas, elev. 2200 m, 30 May 1962,
image!]). Wurdack, J.J. 632 (US barcode 00120072!; isotypes: B
barcode B 10 0244602 [online image!], C barcode
60. Chaetogastra longipilosa (Cogn.) P.J.F.Guim. & Miche- C10015076 [online image!], F barcode V0040214F
lang., comb. nov. ≡ Tibouchina longipilosa Cogn. in Can- [online image!], GH barcode 00073266 [online image!],
dolle & Candolle, Monogr. Phan. 7: 1176. 1891 – LE!, LP barcode 004385 [online image!], MO barcode
Lectotype (designated here): BRAZIL. Santa Catarina: MO-313918 [online image!], NY barcode 00245782!,
“im Walde auf Sandstein bei Minas”, Feb 1890, Ule, TEX barcode 00372245 [online image!], UC barcode
E.H.G. 1446 (BR barcode BR0000005223435!; isolecto- UC1308370 [online image!], USM barcode
types: B [probably destroyed; F negative F0BN016763!], USM000525 [online image!], WAG barcode WAG0
HBG barcodes HBG522832 & HBG522833 [fragments] 002361 [online image!]).
[online images!], P barcode P00708702!, S No. S09-13202
[online image!], US barcode 00120071 [fragment]!). 65. Chaetogastra membranifolia (Cogn. & Rusby) P.J.F.
Guim. & Michelang., comb. nov. ≡ Tibouchina membra-
61. Chaetogastra longisepala (Cogn.) P.J.F.Guim. & Miche- nifolia Cogn. & Rusby in Bull. Torrey Bot. Club 23(1):
lang., comb. nov. ≡ Tibouchina longisepala Cogn. in 17. 1896 – Lectotype (designated here): BOLIVIA.
Candolle & Candolle, Monogr. Phan. 7: 259. 1891 – Lec- Jan 1895, Bang, M. 2857 (BR barcode BR0000
totype (designated here): GUATEMALA. Las Nubes, 005521166!; isolectotypes: MO barcode MO-345396
Nov 1877, Bernoulli & Cario 2872 (BR barcode [online image!], NY barcodes 00245783! & 00245784!,
BR0000005223329!; isolectotype: GOET barcode US barcode 00120074!).
GOET008066 [online image!]).
= Tibouchina spathulata Brandegee in Univ. Calif. Publ. Bot. 66. Chaetogastra minor (Cogn.) P.J.F.Guim. & Michelang.,
6(4): 58. 1914 ≡ Tibouchina longisepala var. spathulata comb. nov. ≡ Tibouchina minor Cogn. in Martius,
(Brandegee) L.O.Williams in Fieldiana, Bot. 29(10): Fl. Bras. 14(3): 390–391, t. 90, fig. 2. 1885 – Lectotype
581. 1963 – Holotype: MEXICO. Chiapas: Cerro del (designated here): BRAZIL. São Paulo: Campos da
Boquerón, Aug 1913, Purpus, C.A. 6771 (UC barcode Bocaina au Sobrado. 8 Jan 1876, Glaziou, A.F.M. 8369
UC172456!; isotypes: BM barcode BM000795746!, (P barcode P00708700!; isolectotypes: C barcode
MO barcode MO-277922 [online image!], NY barcode C10015078 [online image!], K barcode K000329084!, P
00581294!, S No. S09-15485 [online image!]). barcode P00708701!, R barcode R 000009658!).

28
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

67. Chaetogastra mollis (Bonpl.) DC., Prodr. 3. 134. 1828 ≡ nitida (Graham) Triana in Trans. Linn. Soc. London,
Rhexia mollis Bonpl. in Humboldt & Bonpland, Monogr. Bot. 28(1): 39. 1872 (“1871”) ≡ Tibouchina nitida
Melast. 2: 50, pl. 19. 1808 ≡ Tibouchina mollis (Bonpl.) (Graham) Cogn. in Martius, Fl. Bras. 14(3): 404–405.
Cogn. in Martius, Fl. Bras. 14(3): 349. 1885 – Holotype: 1885 – Type: not located. [In the protologue, the author
PERU. Inter montem Saraguru et urbem Loxae, alt. describes that “this plant was raised at Mr Neill’s garden,
1000–1200, s.d., Bonpland, A.J.A. & Humboldt, Canonmills, from seeds sent to him, in 1829, by Mr John
F.W.H.A. von s.n. (P barcode P00136477!; isotypes: F Tweedie” and we have an illustration of the cultivated
barcode V0063947F [fragment] [online image!], P bar- specimen, elaborated by Dr. Robert Kaye Greville, pub-
codes P00136478! & P00136479 [fragment]!). lished in Bot. Mag. 6: fig. 3142. 1832]. Neotype (desig-
= Tibouchina mollis var. glandulifera Wurdack in Phytologia 5 nated here): BRAZIL. Rio Grande do Sul, s.d.,
(2): 54 1954, syn. nov. – Holotype: COLOMBIA. Nariño Tweedie, J. 12 (MO barcode MO-2478495!).
Pasto, Volcán El Galeras, alt. about 3200–3500 m, 4 Jun
1946, Schultes, R.E. & Villarreal, M. 7978 (NY barcode 72. Chaetogastra obtusifolia (Cogn.) P.J.F.Guim. & Miche-
00245786!; isotypes: COL barcode COL000003427 lang., comb. nov. ≡ Tibouchina obtusifolia Cogn. in
[online image!], K barcode K000329142!). Bull. New York Bot. Gard. 4(14): 355. 1907 – Holo-
= Tibouchina mollis var. patenter-villosa Gilli in Feddes type: BOLIVIA. 1500–1700 m, s.d., Bang, M. 2646
Repert. 92: 683. 1981, syn. nov. – Holotype: ECUADOR. (NY barcode 00245792!; isotypes: BR barcode
Wald bei El Corazon, 1400 m, 28 Jun 1975, Gilli, A. BR0000005520985!, CORD barcode CORD 00003626
217 (W). [online image!]).

68. Chaetogastra monticola (Naudin) P.J.F.Guim. & Miche- 73. Chaetogastra oligantha (Gleason) P.J.F.Guim. & Miche-
lang., comb. nov. ≡ Oreocosmus monticola Naudin in lang., comb. nov. ≡ Tibouchina oligantha Gleason in
Ann. Sci. Nat., Bot., sér. 3, 13(1): 38. 1850 [illustration Phytologia 1(3): 134. 1935 – Holotype: BOLIVIA. Dep.
in Ann. Sci. Nat., Bot., sér. 3, 12(10): t. 14, fig. 6. Santa Cruz: Alto de Samaipata, berguchen El Fuerte,
1850] ≡ Tibouchina trianaei Cogn. in Candolle & Can- alt. 2200 m, 14 Oct 1928, Steinbach, J. 8262
dolle, Monogr. Phan. 7: 261. 1891, pro parte, nom. illeg. (NY barcode 00245793!; isotypes: E barcode
superfl. ≡ Tibouchina monticola (Naudin) Cogn. in E00285786 [online image!], GH barcode 00073268
Candolle & Candolle, Monogr. Phan. 7: 263. 1891 – [online image!], GOET barcodes GOET008067
Holotype: MEXICO. Cordillera de La Virgen 7–8000 ft – & GOET008068 [online images!], K barcode
Sierra Yavesia 6500–8000 ft, Nov–Apr 1840, Galeotti, H. K000329171!, MO barcode MO-2176113 [online
2931 (P barcode P00708706; isotypes: BR barcode image!], NY barcode 00659019!, PH barcode 00027261
BR0000005191017!, K barcode K000328982!, LE!, P [online image!], S No. S-R-6123 [online image!], UC
barcode P00708707!, herbarium unknown [F negative barcode UC394767 [online image!], US barcode
F0BN026116!]). 00120078!).

69. Chaetogastra nanifolia (Todzia) P.J.F.Guim. & Miche- 74. Chaetogastra orbignyana (Naudin) P.J.F.Guim. &
lang., comb. nov. ≡ Tibouchina nanifolia Todzia in Brit- Michelang., comb. nov. ≡ Micranthella orbignyana
tonia 51(3): 269–270, fig. 6, 16. 1999 – Holotype: Naudin in Ann. Sci. Nat., Bot., sér. 3, 13: 350–351.
MEXICO. México: 5 km al SW de Nanchititla, Teju- 1850 (“orbigniana”) ≡ Tibouchina orbignyana (Naudin)
pilco, 2000 m, 27 Dec 1972, Medrano, G. & al. 5019 Cogn. in Candolle & Candolle, Monogr. Phan. 7: 261.
(MEXU barcode MEXU 00158369 [online image!]; iso- 1891 – Holotype: BOLIVIA. Prope Cajapi-Yungas, s.d.,
types: CAS barcode 0004856 [online image!], MEXU d’Orbigny, A.C.V.M.D. 357 (P barcode P00708679!; iso-
barcode MEXU 00286070 [online image!]). types: BR barcode BR0000005527816 [fragment]!, F
barcode V0063879F [fragment] [online image!]).
70. Chaetogastra naudiniana Decne. in Rev. Hort. [Paris].
Ser. 3, 1: 86. 1847 ≡ Oreocosmus naudianus (Decne.) 75. Chaetogastra oroensis (Gleason) P.J.F.Guim. & Miche-
Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): 39. 1850 ≡ lang., comb. nov. ≡ Tibouchina oroensis Gleason in Bull.
Tibouchina naudiniana (Decne.) Cogn. in Candolle Torrey Bot. Club 57(1): 63–64. 1930 – Holotype: ECUA-
& Candolle, Monogr. Phan. 7: 264. 1891 – Lectotype DOR. Prov. Oro: between La Chorita and Portovelo, alt.
(designated here): MEXICO. Zacualpan, Mar 1843, 1000–2000 m, 28 Aug 1923, Hitchcock, A.S. 21203
Ghiesbreght, A.B. 38 (P barcode P00708713!; isolecto- (US barcode 00120079!; isotypes: GH barcode
type: P barcode P00708712!). 00073269 [online image!], NY barcode 00245794!).

71. Chaetogastra nitida (Graham) P.J.F.Guim. & Michelang., 76. Chaetogastra panicularis Naudin in Ann. Sci. Nat., Bot.,
comb. nov. ≡ Arthrostemma nitidum Graham in Edin- sér. 3, 14(2): 129. 1850 ≡ Tibouchina panicularis
burgh New Philos. J. 12: 186–187. 1831 ≡ Pterolepis (Naudin) Britton ex Cogn. in Bull. Torrey Bot. Club

29
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

17(3): 54. 1890 – Holotype: BOLIVIA. Prope Chulu- F0BN026112!], GH barcode 00073270 [online image!],
mani, s.d., d’Orbigny, A.C.V.M.D. 472 (P barcode K barcode K000329144!, P barcode P00708684!).
P00708672!).
82. Chaetogastra pentamera (Ule) P.J.F.Guim. & Miche-
77. Chaetogastra paratropica (Griseb.) P.J.F.Guim. & Miche- lang., comb. nov. ≡ Comolia pentamera Ule in Notizbl.
lang., comb. nov. ≡ Pleroma paratropicum Griseb. in Abh. Königl. Bot. Gart. Berlin 6(60): 352. 1915 ≡ Tibouchina
Königl. Ges. Wiss. Göttingen 19: 140–141. 1874 ≡ Tibou- pentamera (Ule) J.F.Macbr. in Publ. Field Mus. Nat.
china paratropica (Griseb.) Cogn. in Candolle & Candolle, Hist., Bot. Ser. 13(4/1): 290–291. 1941 – Lectotype (des-
Monogr. Phan. 7: 257. 1891 – Lectotype (designated ignated here): PERU. Loreto: Cerro de Escaler 1200 m,
here): ARGENTINA. Tucuman, Cuesta de Siambon, s.d., Ule, E.H.G. 6742 (HBG barcode HBG-522823
8 Mar 1872, Lorentz, P.G. 296 (CORD barcode CORD [online image!]; isolectotype B [probably destroyed; F
00005691 [online image!]; isolectotypes: B [probably negative F0BN016844!]).
destroyed; F negative F0BN016779!], BR barcode
BR0000005530328!, CORD barcode CORD 00005692 83. Chaetogastra pleromoides (Naudin) P.J.F.Guim. &
[online image!], G-DC barcode G00319182!). Michelang., comb. nov. ≡ Lasiandra pleromoides Nau-
din in Ann. Sci. Nat., Bot., sér. 3, 13(3): 131. 1850 ≡ Pler-
78. Chaetogastra parviflora (Cogn.) P.J.F.Guim. & Miche- oma maurocarpum Triana in Trans. Linn. Soc. London,
lang., comb. nov. ≡ Tibouchina parviflora Cogn. in Mar- Bot. 28(1): 47. 1872 (“1871”) ≡ Tibouchina maurocarpa
tius, Fl. Bras. 14(3): 410–411, t. 94. 1885 – Lectotype (Triana) Cogn. in Candolle & Candolle, Monogr. Phan. 7:
(designated here): BRAZIL. Goiás: in pascuis siccis ad 260. 1891, nom. illeg. superfl. ≡ Tibouchina pleromoides
Engenho do Abrantes, s.d., Pohl, J.B.E. 1279 (BR bar- (Naudin) Macbr. in Publ. Field Mus. Nat. Hist., Bot. Ser.
code BR0000005223800!; isolectotypes: B [probably 4(7): 174. 1929 – Holotype: PERU. Cuzco, s.d., Gay, C.,
destroyed; F negative F0BN016780!], BM barcode s.n. (P barcode P00708685!; istotypes: P barcodes
BM000795352!, BR barcodes BR0000005223459! P00708686! & P00708687!).
& BR0000005224135!, F barcode V0063993F [frag-
ment] [online image!], K barcode K000329105!, NY bar- 84. Chaetogastra pulcherrima (Gleason) P.J.F.Guim. & Miche-
codes 00245795! & 00245796!). lang., comb. nov. ≡ Tibouchina pulcherrima Gleason in
Bull. Torrey Bot. Club 54(1): 26–27. 1927 – Holotype:
79. Chaetogastra patens (Todzia) P.J.F.Guim. & Michelang., PERU. Dep. Cuzco: in forest, “Pillahuata”, Cerro de Cusil-
comb. nov. ≡ Tibouchina patens Todzia in Brittonia luyoc, alt. 2200–2400 m, 3–6 May 1925, Pennell 13971
51(3): 270–272, fig. 7, 17. 1999 – Holotype: MEXICO. (NY barcode 00245798!; isotypes: F barcode V0040215F
Guerrero: mun. Chilpancingo, 3 km al S de Omiltemi, [online image!], GH barcode 00073271 [online image!],
2200 m, 5 Dec 1966, Rzedowski, J. 23606 (MEXU bar- PH barcode 00027263 [online image!]).
code MEXU 00357703 [online image!]; isotypes: CAS
barcode 0004857 [online image!], ENCB barcode 85. Chaetogastra purpurascens (Cogn.) P.J.F.Guim. & Miche-
ENCB008691 [online image!]). lang., comb. nov. ≡ Tibouchina purpurascens Cogn. in Bull.
Torrey Bot. Club 17(3): 56–57. 1890 – Holotype: BOLIVIA.
80. Chaetogastra pauciflora P.J.F.Guim. & Michelang., Mapiri, 2500 ft., May 1886, Rusby, H.H. 2616 (NY barcode
nom. nov. ≡ Tibouchina lanceolata Cogn. in Bull. Tor- 00245799!; isotypes: BR barcode BR0000005531035 [frag-
rey Bot. Club 17(3): 55–56. 1890, non Chaetogastra ment]!, MICH barcode 1111897 [online image!], NDG bar-
lanceolata DC. 1828 – Holotype: BOLIVIA. Yungas, code NDG47044 [online image!]).
4000 ft., 1885, Rusby, H.H. 2343 (NY barcode 00
245776!; isotype: BR barcode BR0000005520244 86. Chaetogastra purpusii (Brandegee) P.J.F.Guim. &
[fragment]!). Michelang., comb. nov. ≡ Tibouchina purpusii Bran-
degee in Univ. Calif. Publ. Bot. 4(19): 378. 1913 – Holo-
81. Chaetogastra pendula (Cogn.) P.J.F.Guim. & Miche- type: MEXICO. Sierra Madre between Misantla and
lang., comb. nov. ≡ Tibouchina pendula Cogn. in Bot. Naolinco, Aug 1912, Purpus, C.A. 6104 (UC barcode
Jahrb. Syst. 8(1): 18. 1886 – Lectotype (designated UC146822 [online image!]; isotypes: BM barcode
here): ECUADOR. In silvis ad Ançua Calicali et S. Flor- BM000795745!, E barcode E00531224 [online image!],
encio prope Quito, alt. 1500–2000 m, 29 Nov 1880, Leh- F barcode V0063974F [online image!], G barcode
mann, F.C. 145 (BR barcode BR0000005530977!; G00368023 [online image!], GH barcode 00073254
isolectotypes: G!, LE!, US barcode 00120085!). [online image!], NY barcode 00581293!, P barcode
Other syntypes: ECUADOR. In declivitate occidentali P00708987!, US barcode 00681571!).
montis Pichincha, alt. 3000 m, s.d., Jameson, W. 388
(BM barcode BM000953945!, E barcode E00285794 87. Chaetogastra repens (Wurdack) P.J.F.Guim. & Miche-
[online image!], G-DC barcode G00318998! [F negative lang., comb. nov. ≡ Tibouchina repens Wurdack in

30
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

Phytologia 5(2): 55–56. 1954 – Holotype: PERU. Dep. 93. Chaetogastra rupestris (Cogn.) P.J.F.Guim. & Miche-
San Martin, Boquerón Pass, 92 km from Tingo Maria lang., comb. nov. ≡ Tibouchina rupestris Cogn. in Can-
on highway to Pucallpa, elev. 400 m, 16 Dec 1949–5 dolle & Candolle, Monogr. Phan. 7: 1176. 1891 –
Jan 1950, Allard, H.A. 22082 (US barcode 00120089!). Holotype: BRAZIL. Santa Catarina: inter rupibus ad
Campo do Serra Geral, zwischen Steinen auf dem Campos
88. Chaetogastra rhynchantherifolia (Cogn.) P.J.F.Guim. der Serra do Oratório, Jan 1890, Ule, E.H.G. 1447
& Michelang., comb. nov. ≡ Tibouchina rhynchantheri- (BR barcode BR0000005225316!; isotypes: B [probably
folia Cogn. in Bot. Jahrb. Syst. 42: 134. 1908 – Lectotype destroyed; F negative F0BN016790!, CORD barcode
(designated by Burke & Michelangeli in Phytotaxa 125 CORD 00003627 [online image!], HBG barcodes
(1): 12. 2013): PERU. Inter Sandia et Tambo Azalaya HBG523036, HBG523037, HBG523038 & HBG523039
in via a Sandia ad Chunchusmayo in fruticetis, [online images!], P barcode P00708678!, R barcode
1500–2000 m a.s.l., 5 Jun 1902, Weberbauer, A. 1062 R 000168428!, S No. S09-13188 [fragment] [online
(BR barcode BR0000005225958!; isolectotype: B [prob- image!]).
ably destroyed; F negative F0BN016785!]).
94. Chaetogastra rusbyi (Cogn.) P.J.F.Guim. & Michelang.,
89. Chaetogastra riograndensis F.S.Mey. in Phytotaxa 282 comb. nov. ≡ Tibouchina rusbyi Cogn. in Bull. Torrey
(4): 253–257, fig. 10–12. 2016 – Holotype: BRAZIL. Bot. Club 17(3): 54–55. 1890 – Holotype: BOLIVIA.
Rio Grande do Sul: São Francisco de Paula, Parque Esta- Unduavi, 8000 ft., Oct 1885, Rusby, H.H. 2339
dual do Rio Tainhas, 20 Apr 2014, Meyer, F.S.; Volet, (NY barcode 00245805!; isotype: BR barcode BR00
D.P. & Monge, M. 1996 (UPCB; isotypes: ICN, MBM). 00005530670!).

90. Chaetogastra rojasii (Cogn.) P.J.F.Guim. & Michelang., 95. Chaetogastra sandiensis (Wurdack) P.J.F.Guim.
comb. nov. ≡ Tibouchina rojasii Cogn. in Repert. Spec. & Michelang., comb. nov. ≡ Tibouchina sandiensis Wur-
Nov. Regni Veg. 7(4/6): 71–72. 1909 – Lectotype (desig- dack in Phytologia 14(5): 259–260. 1967 – Holotype:
nated here): PARAGUAY: Sierra de Amambay, in cam- PERU. Dep. Puno: low forest below Sandia, elev.
pis humidis ad Punta Porá, Dec 1907, Hassler, E. 9928 1800–1900 m, 13 May 1966, Ferreyra, R. 16677 (US
(NY barcode 00245804!; isolectotypes: B [probably barcode 00120091!; isotype: USM).
destroyed; F negative F0BN016787!], K barcode
K000329126!, P barcode P00708688!). 96. Chaetogastra saxicola (F.S.Mey., P.J.F.Guim. & R.Gold-
enb.) P.J.F.Guim. & Michelang., comb. nov. ≡ Tibou-
91. Chaetogastra roseotincta (Todzia) P.J.F.Guim. & Miche- china saxicola F.S.Mey., P.J.F.Guim. & R.Goldenb. in
lang., comb. nov. ≡ Tibouchina roseotincta Todzia in Hoehnea 36(1): 144–147, fig. 3. 2009 – Holotype: BRA-
Brittonia 51(3): 273–275, fig. 8, 18. 1999 – Holotype: ZIL. Paraná: Jaguariaíva, Parque Estadual do Cerrado,
MEXICO. Guerrero: mun. Atoyac de Alvarez, 23.5 km margens do Rio Jaguariaíva, 21 Apr 2007, Meyer,
al NE El Parafso, camino Atoyac-Puerto del Gallo, F.S. & Lisingen, L. von 325 (UPCB!; isotype: RB barcode
1700 m, 22 Nov 1983, Martínez, E. & Barrie, F. 5595 00669520!).
(MEXU barcode MEXU 00427279 [online image!]; iso-
type: CAS barcode 0004859 [online image!]). 97. Chaetogastra saxosa (Gleason) P.J.F.Guim. & Miche-
lang., comb. nov. ≡ Tibouchina saxosa Gleason in Phyto-
92. Chaetogastra rufipilis (Schltdl.) Walp. in Repert. Spec. logia 1(3): 135–136. 1935 – Holotype: PERU. Dep.
Nov. Regni Veg. Sonderbeih. E 2(1): 130. 1843 ≡ Rhexia Cusco: rocky forest at Pillahuata, Cerro de Cusilluyoc,
schiedeana var. macrantha Schltdl. & Cham. in Linnaea alt. 2200–2400 m, 3–5 May 1925, Pennell, F.W. 13992
5: 566. 1830 ≡ Rhexia rufipilis Schltdl. in Linnaea 13: (GH barcode 00073272 [online image!]; isotype: NY bar-
430–431. 1839 ≡ Tibouchina rufipilis (Schltdl.) Cogn. code 00245806!).
in Candolle & Candolle, Monogr. Phan. 7: 259. 1891 –
Holotype: MEXICO. Veracruz: in sylvis Jalapensibus, 98. Chaetogastra scabriuscula (Schltdl.) P.J.F.Guim.
monte Macultepec, San Andrés, Aug, Schiede, & Michelang., comb. nov. ≡ Rhexia scabriuscula Schltdl.
C.J.W. & Deppe, F. s.n. (B [probably destroyed; F nega- in Linnaea 13: 431. 1839 ≡ Pleroma scabriusculum
tive F0BN016789!]; isotypes: BM barcode BM000 (Schltdl.) Triana in Trans. Linn. Soc. 28(1): 46. 1872
901168!; BR barcode BR0000005530663!). (“1871”) ≡ Tibouchina scabriuscula (Schltdl.) Cogn. in
= Tibouchina rufipilis var. hirsuta Cogn. in Candolle & Can- Candolle & Candolle, Monogr. Phan. 7: 262. 1891 –
dolle, Monogr. Phan. 7: 259. 1891, syn. nov. – Lectotype Holotype: MEXICO. Schiede, C.J.W. s.n. (not located).
(designated here): MEXICO. Prope Jalapa, alt.
2100–2500 m, Aug 1884, Com. Geographico-explor. 99. Chaetogastra schiedeana (Schltdl. & Cham.) Walp. in
405 (BR barcode BR0000005530342!; isolectotype: BR Repert. Spec. Nov. Regni Veg. Sonderbeih. E 2(1): 130.
barcode BR0000005191437!). 1843 ≡ Rhexia schiedeana var. micrantha Schltdl.

31
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

& Cham. in Linnaea 5: 565. 1830 ≡ Tibouchina schie- Trans. Linn. Soc. London, Bot. 28(1): 40. 1872
deana (Schltdl. & Cham.) Cogn. in Candolle & Candolle, (“1871”) ≡ Tibouchina simplicicaulis (Naudin) Cogn. in
Monogr. Phan. 7: 261. 1891 – Lectotype (designated Martius, Fl. Bras. 14(3): 406–407. 1885 – Holotype:
here): MEXICO. In apricis Misantlae, Mar, Deppe, F. BRAZIL. Minas Gerais: “in pascuis uliginosis prope
& Schiede, C.J.W. 563 (K barcode K000328978!; isolec- Santo-Anjo, Braziliae australis, ad 23 –24 lineam aus-
totypes: BR barcode BR0000005190683!, HBG barcode tralem”, 1816, De Saint-Hilaire, A.F.C.P. B1-407
HBG-523035 [online image!], K barcode K000328979!). (P barcode P00708811!; isotype: P barcode P00708812!).

100. Chaetogastra sebastianopolitana (Raddi) P.J.F.Guim. 103. Chaetogastra sinaloensis (Todzia) P.J.F.Guim.
& Michelang., comb. nov. ≡ Rhexia sebastianopolitana & Michelang., comb. nov. ≡ Tibouchina sinaloensis Tod-
Raddi, Melast. Bras.: 8, t. 2, fig. 4. 1828 ≡ Tibouchina zia in Brittonia 51(3): 275–276, fig. 9, 19. 1999 – Holo-
sebastianopolitana (Raddi) Cogn. in Martius, Fl. Bras. type: MEXICO. Sinaloa: mun. Concordia, La Capilla
14(3): 409–410. 1885 – Holotype: BRAZIL. Rio de del Taxte on Mexican Hwy. 40, 1370 m, 18 Oct 1983,
Janeiro: Trovasi nella vicinanza di S. Sebastiano, altri- Breedlove, D.E. & Almeda, F. 58794 (CAS barcode
menti detta Rio de Janeiro, dove è comunissima (Raddi, 0004860 [online image!]; isotypes: MEXU, MICH, NY
op. cit.: 19), “Rhexia sebastianopolitana nob:” [scripsit: barcode 00346174!, TEX barcode 00374499 [online
G. Raddi on PI], s.d., Raddi s.n. (FI barcode FI005215 image!]).
[online image!]; isotype: FI005216 [online image!]).
= Arthrostemma macrantherum Miq. in Linnaea 22: 540. 104. Chaetogastra solmsii (Cogn.) P.J.F.Guim. & Miche-
1849 ≡ Tibouchina sebastianopolitana var. miqueliana lang., comb. nov. ≡ Tibouchina solmsii Cogn. in Candolle
Cogn. in Martius, Fl. Bras. 14(3): 410. 1885, syn. nov. – & Candolle, Monogr. Phan. 7: 250. 1891 – Lectotype
Lectotype (designated here): BRAZIL. Minas Gerais: (designated here): PERU. San Govan, Jul 1854,
Caldas, 9 Mar 1862, Regnell II 107 (BR barcode Lechler, W. 2179 (BR barcode BR0000005530724!; iso-
BR0000005531004 [date 12 Feb 1861]!; isolectotypes: lectotype: K barcode K000329143!).
BR barcode BR0000005530380 [no date]!, K barcode Other syntype: PERU, Tatanara, Aug 1854, Lechler, W.
K000329101 [date 9 Mar 1862]!, LE!, S No. S09-13135 2555 (GOET barcode GOET008069 [online image!]).
[no date] [online image!], U barcode U 0004183 [date
9 Mar 1862] [online image!], US barcodes 00120170! 105. Chaetogastra stenopetala (Cogn.) P.J.F.Guim. & Miche-
& 01906015!). lang., comb. nov. ≡ Tibouchina stenopetala Cogn. in
= Tibouchina sebastianopolitana var. hirsuta Cogn. in Mar- Candolle & Candolle, Monogr. Phan. 7: 262. 1891 –
tius, Fl. Bras. 14(3): 410. 1885, syn. nov. – Lectotype Lectotype (designated here): PERU. Prope San
(designated here): BRAZIL. Minas Gerais: Barbacena, Govan, Jul 1854, Lechler, W. & Hohenacker, R.F. 2271
26 Jan 1873, Glaziou, A.F.M. 8964 (BR barcode (BR barcode BR0000005225989!; isolectotypes: B
BR0000005531042!; isolectotypes: BR barcode [probably destroyed; F negative F0BN016798!], BR bar-
BR0000005531059!; K barcode K000329102!, P bar- code BR0000005225330!, G-DC barcode G00318990!,
code P00708802!). K barcode K000329161!, LE [on 3 sheets]!, M barcode
Other syntypes: BRAZIL. Rio de Janeiro: Alegria, 22 Feb M-0165862!, NY barcode 245809 [fragment]!, P bar-
1882, Glaziou, A.F.M. 13841 (K barcode K000880906!, codes P00708791 & P00708792!, S No. S-R-3521
P barcodes P00708800! & P00708801!). [online image!], TCD barcode TCD0000758 [online
image!]).
101. Chaetogastra sericea (de Santiago) P.J.F.Guim.
& Michelang., comb. nov. ≡ Tibouchina sericea de San- 106. Chaetogastra stenophylla (Cogn.) P.J.F.Guim. &
tiago in Brittonia 61(1): 51–54, fig. 1. 2009 – Holotype: Michelang., comb. nov. ≡ Tibouchina stenophylla
México. Guerrero: mun. Malinaltepec, Camino al cerro Cogn. in Bull. Torrey Bot. Club, 17(3): 56. 1890 – Lecto-
Borracho, 2000 m, 17 06′52.6′′N, 98 38′43.2′′W, type (designated here): BOLIVIA. Guanay, 2000 ft.,
22 Oct 2005, Lozada, L.; Santiago, R. & Rojas, J. 2909 May 1886, Rusby, H.H. 2323 (NY barcode 00245810!;
(FCME; isotypes: CAS barcode 0026357 [online isolectotypes: BR barcode BR0000005225668 [frag-
image!], ENCB, IEB, K barcode K000857990!, MEXU ment]!, CAS barcode 0004862 [online image!], E bar-
barcode MEXU 01262056 [online image!], MO barcode code E00531234 [online image!], GH barcode
MO-2195434 [online image!], NY barcode 01104836!, 00073273 [online image!], K barcode K000329159!,
UAMIZ, US barcode 01105775!, XAL). LE!, MICH barcode 1111896 [online image!], MO
barcode MO-313913 [online image!], P barcode
102. Chaetogastra simplicicaulis (Naudin) P.J.F.Guim. P00708793!, US barcodes 00120096! & 00120097!).
& Michelang., comb. nov. ≡ Tetrameris simplicicaulis
Naudin in Ann. Sci. Nat., Bot., sér. 3, 14(2): 122–123. 107. Chaetogastra tetrapetala (Cogn.) P.J.F.Guim. & Miche-
1850 ≡ Pterolepis simplicicaulis (Naudin) Triana in lang., comb. nov. ≡ Tibouchina tetrapetala Cogn. in Bull.

32
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

Torrey Bot. Club 276. 1896 – Lectotype (designated “Ayuruoca au Pico do Papagaio”, 22 Jan 1886, Glaziou,
here): BOLIVIA. Prov. Nor Yungas: Coroico, 10 Sep A.F.M. 16034 (BR barcode BR0000005225347!; iso-
1894, Bang, M. 2425 (BR barcode BR0000005226016!; lectotypes: BR barcode BR0000005225675!, C bar-
isolectotypes: E barcode E00285784!, F barcodes code C10015090 [online image!], K barcode
V0064001F & V0064002F [online images!], GH barcode K000329103!, P barcode P00708664!, R barcode
00073275 [online image!], K barcode K000329173!, R 000009669!).
LE!, MICH barcode 1111895 [online image!], MIN bar-
code 1002622 [online image!], MO barcode MO- 113. Chaetogastra venosa (Gleason) P.J.F.Guim. & Miche-
313912 [online image!], NY barcode 00245812!, PH bar- lang., comb. nov. ≡ Tibouchina venosa Gleason in Phyto-
code 00027265 [online image!], US barcodes 00120102!, logia 1(3): 134–135. 1935 – Holotype: BOLIVIA. Dep.
00120103! & 00921437!). Santa Cruz: Yungas de San Mateo Comarapa, alt.
2500 m, 24 Oct 1928, Steinbach, J. 8509 (K barcode
108. Chaetogastra thulia (Todzia) P.J.F.Guim. & Michelang., K000329174!; isotypes: E barcode E00285787 [online
comb. nov. ≡ Tibouchina thulia Todzia in Brittonia 51(3): image!], GH barcode 00073276 [online image!], NY bar-
276–278, fig. 10, 20. 1999 – Holotype: MEXICO. Sina- code 00245814!).
loa: mun. Badiraguato, Sierra Surutato, in the broad val-
ley of Ocurahui, 6000 ft., 2 Nov 1969, Breedlove, 114. Chaetogastra versicolor (Lindl.) P.J.F.Guim. & Miche-
D.E. & Kawahara, F.S. 16864 (US barcode 00679973!; lang., comb. nov. ≡ Rhexia versicolor Lindl. in Bot.
isotypes: CAS barcode 0004863 [online image!], MO Reg. 13: 1066, t. 1066. 1827 ≡ Tibouchina versicolor
barcode MO-277902 [online image!]). (Lindl.) Cogn. in Martius in Fl. Bras. 14(3): 405–406.
1885 – Lectotype (designated here): [illustration in]
109. Chaetogastra tortuosa (Bonpl.) DC., Prodr. 3: 132. 1828 Bot. Reg. 13: t. 1066. 1827.
≡ Rhexia tortuosa Bonpl. in Humboldt & Bonpland, = Tibouchina versicolor var. robusta Cogn. in Martius,
Monogr. Melast. 2: 17–18, pl. 7. 1808 ≡ Oreocosmus Fl. Bras. 14(3): 406. 1885, syn. nov. – Lectotype (desig-
tortuosus (Bonpl.) Naudin in Ann. Sci. Nat., Bot., sér. nated here): BRAZIL. Santa Catarina: s.d., Raben,
3, 13: 38. 1850 ≡ Tibouchina tortuosa (Bonpl.) Almeda F.C. 433 (BR barcode BR0000005530427!).
in Novon 15(5): 517–518. 2005 – Holotype: MEXICO. Other syntypes: BRAZIL. Santa Catarina: s.d., Sellow, F.
Guerrero: Taxco, “Crescit in aridis Regni Novae Hispa- s.n. (not located); idem, s.d., D’Urville, J.S.C.D. s.n. (not
niae, prope Tasco, ad altitudinem 2400 metrorum”, Bon- located); idem, s.d., Eschscholtz, J.F.G. von s.n. (not
pland, A.J.A. 3962 (P barcode P00708705!; isotypes: located); ARGENTINA. Buenos Aires: s.d., Fielding, E.
BR barcode BR0000005223756 [fragment]!, P barcodes s.n. (not located).
P00136463! & P00136464!). = Tibouchina meiodon Stapf in Bull. Misc. Inform. Kew 1895
= Pleroma mexicanum DC., Prodr. 3: 152. 1828 ≡ Melastoma (100–101): 104. 1895, syn. nov. – Holotype: BRAZIL.
mexicanum D.Don in Mem. Wern. Nat. Hist. Soc. 4(2): Cultivated at Kew from seeds believed to have been sent
290. 1823, nom. illeg., non Bonpl. 1816 ≡ Tibouchina by Glaziou, A.F.M. s.n. (K barcode K000329120!).
mexicana Cogn. in Candolle & Candolle, Monogr. Phan.
7: 258. 1891 – Type: MEXICO. Pavón, J.A. s.n. [Herbar- 115. Chaetogastra violacea (Cogn.) P.J.F.Guim. & Miche-
ium of Lambert, not located]. lang., comb. nov. ≡ Tibouchina violacea Cogn. in Can-
dolle & Candolle, Monogr. Phan. 7: 272. 1891 –
110. Chaetogastra triflora (Gleason) P.J.F.Guim. & Miche- Lectotype (designated here): PARAGUAY. Asuncion:
lang., comb. nov. ≡ Tibouchina triflora Gleason in Amer. “Loci humidis”, May 1874, Balansa, B. 1937
J. Bot. 19: 740. 1932 – Holotype: COLOMBIA. Dep. El (P barcode P00708946! [F negative F0BN036177!];
Cauca: “La Gallera”, Micay Valley, alt. 1800–1900 m, isolectotypes: BR BR0000005226986!, F barcode
1 Jul 1922, Killip, E.P. 7925 (NY barcode 00245813!; iso- V0064005F [fragment] [online image!], G!, K barcode
types: PH barcode 00027267!, US barcode 00120104!). K000329104!, P barcode P00708947!).

111. Chaetogastra trinitensis (Cogn.) P.J.F.Guim. & Miche- 116. Chaetogastra wasshausenii (Wurdack) P.J.F.Guim.
lang., comb. nov. ≡ Tibouchina trinitensis Cogn. in & Michelang., comb. nov. ≡ Tibouchina wasshausenii
Urban, Symb. Antill. 7(3): 309–310. 1912 – Holotype: Wurdack in Brittonia 40(1): 7–9, fig. 1. 1988 – Holotype:
TRINIDAD AND TOBAGO. Trinidad: s.d., Broadway, PERU. Puno: rocky outcrop edge in lowland rain forest
W.E. 3659 (not located). between San Juan del Oro and San Ignacio, Prov. Sandia,
elev. 1200 m, 7 Jun 1982, Wasshausen, D.C. & Salas, A.
112. Chaetogastra urbanii (Cogn.) P.J.F.Guim. & Miche- 1209 (US barcode 00048308!).
lang., comb. nov. ≡ Tibouchina urbanii Cogn. in Mar-
tius, Fl. Bras. 14(4): 602, t. 129, fig. 1. 1888 – 117. Chaetogastra weberbaueri (Cogn.) P.J.F.Guim. &
Lectotype (designated here): BRAZIL. Minas Gerais: Michelang., comb. nov. ≡ Tibouchina weberbaueri

33
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

Cogn. in Bot. Jahrb. Syst. 42: 136. 1908 – Lectotype (des- with a center of diversity in the Andes of Venezuela and
ignated by Burke & Michelangeli in Phytotaxa 125(1): Colombia with outlying species in the Guayana highlands of
12. 2013): PERU. Dep. Cuzco, in provincia Convencion, Venezuela. It is also found in Costa Rica in the Talamanca
supra Hacienda Idma apud Sta. Anna, in campo grami- mountains. It grows in temperate forests in subpáramos and
noso fruticibus intermixtis, 1700–1800 m.s.m., 24 Jun páramos on rocks and sandstone cliffs, on exposed slopes with
1905, Weberbauer, A. 5002 (BR barcode BR0000 shrubby vegetation, and along roadsides.
005226344!; isolectotypes: B [probably destroyed; F neg- Chaetolepis species are highly branched shrubs or suffru-
ative F0BN016808!], F barcode V0064006F [fragment] tescent herbs, with leaves strigose or hispid and rigid with an
[online image!]). ericoid appearance, small flowers with 4 petals yellow or pur-
ple and yellow stamen connective not or slightly prolonged at
Chaetolepis (DC.) Miq., Comm. Phytogr.: 72. 1840 ≡ Osbeckia the base, less then 1 mm long, without appendages.
sect. Chaetolepis DC., Prodr. 3: 140. 1828 – Type: Chaeto-
lepis microphylla (Bonpl.) Miq. (≡ Rhexia microphylla Heterocentron Hook. & Arn., Bot. Beechey Voy.: 290. 1838
Bonpl. ≡ Osbeckia microphylla (Bonpl.) DC.). – Type: Heterocentron mexicanum Hook. & Arn.
= Haplodesmium Naudin in Ann. Sci. Nat., Bot., sér. 3, 14(2): = Heeria Schltdl. in Linnaea 13: 432. 1839, nom. illeg., non
150, t. 5, fig. 4. 1850 – Type: Chaetolepis lindeniana Meisn. 1837 ≡ Schizocentron Meisn., Pl. Vasc. Gen. 2:
(Naudin) Triana (≡ Haplodesmium lindenianum Naudin). 355. 1843 – Type: Heterocentron elegans (Schltdl.)
= Trimeranthus H.Karst. in Linnaea 30: 159. 1859 – Type: Kuntze (≡ Heeria elegans Schltdl. ≡ Schizocentron ele-
Trimeranthus alpestris H.Karst. (= Chaetolepis lindeni- gans (Schltdl.) Rose).
ana (Naudin) Triana). Shrubs or subshrubs, erect to 1–3 m tall or procumbent
Erect annual herbs or compactly branched subshrubs, with with trailing branches to 0.5–1 m long, herbs suffrutescent
small leaves slightly overlapping or clustered towards the apex or prostrate cespitose, rooting at the nodes, stem indumentum
of the branches; indument varies from smooth eglandular tri- hirsute with smooth hairs or strigose. Leaves opposite, decus-
chomes to smooth glandular trichomes to barbellate trichomes sate, blades of a pair essentially equal in size, petiolate or
to scales on branches, leaves and floral structures. Leaves rarely subsessile, membranaceous to coriaceous, densely to
opposite, spreading or appressed, blades of a pair essentially sparsely hairy or glabrescent on both surfaces, penninerved
equal in size, symmetrical, conspicuously petiolate or sessile, or 3-nerved. Inflorescence in terminal bracteate dichasia, sim-
coriaceous or chartaceous; adaxial and abaxial leaf surfaces ple or composite, or solitary terminal flowers mostly on lateral
glabrous, sessile glands, punctate, or sparsely to densely cov- branches; bracteoles 2, persistent, smaller than the mature
ered with trichomes or scales, (1–)3–7-nerved. Inflorescence hypanthium. Flowers perigynous; pedicellate. Hypanthium
a very small corymb with 3–5 flowers, or flower solitary, campanulate to hemispheric or subglobose at maturity, vari-
terminal or in axils of upper leaves; flowers perigynous ously hairy to glabrous; calyx not calyptriform, with conspic-
or half-epigynous; sessile or short pedicellate. Hypanthium uous lobes, lobes 4, persistent in fruit, triangulate. Corolla
campanulate or urceolate, externally smooth or with slight lon- patent; petals 4, free and spreading, white, pink, purple or
gitudinal vascular ribs, glabrous or appressed pubescent. Calyx magenta, color remains unchanged with age, obovate to ovate,
not calyptriform, with lobes conspicuous, lobes 4, persistent to glabrous. Stamens 8, all fertile, strongly dimorphic, consisting
fruit, always with setose trichomes among lobes; deltoid to nar- of 4 large antesepalous stamens and 4 smaller antepetalous
rowly triangular, or lance-triangular. Corolla patent; petals ones. Anthers 2-celled, ventrally channeled, linear-oblong to
4, free and spreading, yellow, magenta or purple, color remains slightly subulate, opening by a solitary pore in the larger
unchanged with age, obovate, glabrous. Stamens 8, all fertile, anthers slightly inclined ventrally and in the smaller anthers
similar in size and shape. Anthers 2-celled, oblong-linear or truncated to slightly dorsally inclined. Connectives glabrous
oblong-subulate, linear to slightly arcuate, ventrally smooth or beset dorsally with two (rarely one) glandular hairs posi-
with a large apical pore oriented ventrally or dorsally. Connec- tioned above the filament insertion; larger stamens with con-
tives glabrous, not or slightly prolonged at the base (<1 mm nective prolonged below the thecae and extended beyond the
long), without dorsal or ventral appendages. Thin filaments filament insertion into an elongate bifid and glabrous append-
flattened, glabrous. Ovary superior or half-superior, 4-celled, age; smaller stamens with the connective just prolonged below
setulose apically, enclosed by the hypanthium thalamus but the thecae forming an inconspicuous appendage obtusely gla-
not fused to it or slightly fused at base, axile placentation; style brous. Filaments glabrous. Ovary superior, 4-celled, apically
glabrous, straight or slightly curved distally, stigma puncti- crowned by 4 triangular scales, rectangular or variously lobed,
form, inconspicuous. Mature capsule dry, semiwoody or scales glabrous or setose; style glabrous, declined to side of
papery, loculicidal, no angle, longitudinal vascular ribs con- flower, stigma punctiform, inconspicuous. Mature capsule
spicuous or sometimes cryptic upon drying. Seeds numerous dry, semiwoody, loculicidal, not angled. Seeds numerous
and tiny, cochleate, slightly to strongly curved, testa smooth and tiny, cochleate and tuberculate, hilum terminal.
or faveolate; raphe inconspicuous, hilum terminal. Heterocentron has 43 taxa (29 species, 14 varieties), con-
Chaetolepis is represented by 13 taxa (11 species, 2 varie- fined to Mexico and Central America, with the highest diver-
ties). This is a tropical, montane genus found above 2000 m, sity in Mexico (33 taxa). In Central America, most of the

34
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

species are distributed in the northwest. They are found mostly few to 10–15 flowers per inflorescence or solitary flowers.
in cloud forests, rain forests, and pine-oak forests, growing Flowers perigynous; pedicellate. Hypanthium campanulate
along forest margins and clearings, cliff faces, lake margins, or long-obconical (flared), externally smooth or with
rocky canyon walls, roadsides and disturbed sites from 8 slightly longitudinal vascular ribs, glabrous or pubescent,
750 to 2400 m (Almeda, 2009). trichomes simple or glandular. Calyx not calyptriform in
Heterocentron has tetramerous flowers, strongly dimor- bud, with conspicuous lobes, persistent or deciduous once
phic stamens in which the connective of the larger anther is opened the petals; lobes 4, ovate, lanceolate, narrowly tri-
long-produced below the thecae and appendaged; leaf vena- angular or linear obovate. Corolla patent; petals 4, free
tion is penninerved asymmetrical (rarely nerved from the and spreading, white, red-purple, pink, purple or lilac,
base), and the primary veins diverge from the median vein in color remains unchanged with age, obovate, glabrous. Sta-
a subopposite to irregularly alternate fashion. mens twice in number to petals (8) or less frequently in
The following species are transferred to Heterocentron: equal number (with 4 fertile and 4 staminodes), dimorphic.
Anthers linear subulate, straight or slightly curved, ven-
1. Heterocentron breedlovei (Wurdack) P.J.F.Guim. & Miche- trally smooth, with a small apical pore dorsally oriented.
lang., comb. nov. ≡ Tibouchina breedlovei Wurdack in Connectives glabrous, prolonged or not, with an conspicu-
Phytologia 14(5): 260–261. 1967 – Holotype: MEXICO. ous appendix dorsal. Filaments thin, usually flattened, gla-
Chiapas, mun. La Trinitaria, collected on a slope with brous. Ovary superior, 4-celled, globose or obovoid,
Pinus and Quercus at Lago de Montebello, 40 km E of apically setulose or pubescent; enclosed by the hypanthium
La Trinitaria, elev. 1600 m, 27 May 1965, Breedlove, but not fused to it; style filiform, glabrous, stigma incon-
D.E. 10060 (US barcode 00681569!; isotypes: CAS spicuous. Fruit capsule, usually elongated, completely
barcode 0004735 [online image!], DS, ENCB barcode enclosed by the hypanthium, which may be smooth or
ENCB008687 [online image!], F barcode V0063970F ribbed. Seeds numerous, tiny, cochleate, testa smooth or
[online image!], LL barcode 00372243 [online finely irregular, hilum terminal.
image!]). Monochaetum has over 50 species, which range from
Sinaloa, Mexico through Central America and the Andes to
2. Heterocentron chiapense (Wurdack) P.J.F.Guim. & Miche- Peru and the Venezuelan Coastal Cordillera, and are also pre-
lang., comb. nov. ≡ Tibouchina chiapensis Wurdack in sent in the Guayana highlands, usually at high elevations from
Phytologia 14(5): 261. 1967 – Holotype: MEXICO. Chia- 900 up to 3200 m (Almeda, 1974). They occur in cloud for-
pas: near Monserrate, 9 Mar 1952, MacDougall, T.B. s.n. ests, pine or pine-oak forests, moist thickets, volcanic crater
[collection no. series of E. Matuda 26064] (US No. margins, pastures, disturbed sites, trail margins, shaded road-
2364170!; isotypes: CAS barcode 0004849 [fragment] banks, boggy areas and rocky meadows.
[online image!], MEXU barcode MEXU 00015120 The genus is characterized by connectives below the
[online image!], NY barcode 00581296!). anther thecae with upturned or deflexed, brightly colored dor-
sal appendages (Almeda, 1974).
Monochaetum (DC.) Naudin in Ann. Sci. Nat., Bot., sér. 3, 4:
48–49, t. 2. 1845, nom. cons. ≡ Arthrostemma sect. Pilocosta Almeda & Whiffin in Syst. Bot. 5(3): 303. 1980 –
Monochaetum DC., Prodr. 3: 138. 1828 – Type: Mono- Type: Pilocosta oerstedii (Triana) Almeda & Whiffin (≡
chaetum calcaratum (DC.) Triana, typ. cons. (≡ Arthro- Pterolepis oerstedii Triana).
stemma calcaratum DC. ≡ Monochaetum candolleanum Subshrubs, erect or decumbent, 50 cm tall or prostrate matting
Naudin, nom. illeg. superfl.). or trailing herbs 30–50 cm long, pubescent to strigose in all
= Grischowia H.Karst., Auswahl Gew. Venez.: 15. 1848 – structures with trichomes simple, smooth or finely roughened.
Type: Monochaetum hirtum (H.Karst.) Triana ex Cogn. Leaves opposite, decussate or distichous, equal to somewhat
(≡ Grischowia hirta H.Karst.). unequal in each pair, symmetrical, petiolate, chartaceous;
= Loevigia H.Karst. & Triana in Triana, Nuev. Jen. Esp.: 22. pubescent to strigose above, puberulous below; 5–7-nerved
1854 – Type: Loevigia sericea H.Karst. & Triana (= or -plinerved. Inflorescence consisting of solitary, axillary
Monochaetum bonplandii (Kunth) Naudin ≡ Rhexia bon- and ebracteate flowers. Flowers perigynous; pedicellate.
plandii Kunth). Hypanthium pilose, sericeous, or setulose on the four promi-
= Roezlia Regel in Trudy Imp. S.-Peterburgsk. Bot. Sada 1: nent angles. Calyx not calyptriform, with conspicuous lobes,
98. 1871 – Type: Roezlia granadensis Regel. lobes 4, persistent to fruit, lance-triangular to narrowly subu-
Suffrutescent herbs or shrubs, 0.5–2 m tall, villous or late. Corolla patent; petals 4, free, spreading or connivent
pubescent appressed or laxly. Trichomes simple, dendritic and broadly campanulate, pink, fugacious, obovate, typically
or glandular. Leaves opposite, decussate, blades of a pair ciliate, glabrous. Stamens 8; all fertile; strongly dimorphic to
essentially equal in size, symmetrical, short petiolate, char- subequal, consisting of 4 large antesepalous stamens and
taceous, adaxial and abaxial leaf surfaces pubescent or less 4 smaller antepetalous ones. Anthers linear-subulate to
frequently glabrous, 3–9-nerved or -plinerved. Inflores- linear-oblong, straight or slightly curved, ventrally smooth
cence terminal, cymose, sub-corymbiform or paniculate, or channeled, with a small apical pore ventrally oriented.

35
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

Connectives glabrous, shortly prolonged at the base and Pleroma martiusianum (DC.) P.J.F.Guim. & Michelang.
slightly curved ventrally, with two truncated ventral lobules, (≡ Lasiandra martiusiana DC.).
glabrous. Filaments glabrous and complanate. Ovary superior, = Lasiandra sect. Brachypodae Naudin in Ann. Sci. Nat.,
4-celled; setulose apically; enclosed by the hypanthium but Bot., sér. 3, 13(3): 134. 1850 – Type (designated here):
not fused to it, axile placentation; style glabrous, linear, Pleroma martiale (Cham.) Triana (≡ Lasiandra martialis
straight or somewhat declinate; stigma punctiform, inconspic- Cham.).
uous. Mature capsule dry, semiwoody or coriaceous, loculici- = Lasiandra sect. Angustifoliae Naudin in Ann. Sci. Nat.,
dal, elongated with 4 angles, and 4 valves. Seeds numerous Bot., sér. 3, 13(3): 135. 1850 – Type (designated here):
and tiny, cochleate, conspicuously tuberculate, hilum Pleroma angustifolium (Naudin) Triana (≡ Lasiandra
terminal. angustifolia Naudin).
Five species are recognized in Pilocosta, distributed from = Lasiandra sect. Macrogastrae Naudin in Ann. Sci. Nat.,
Volcán Arenal in north-central Costa Rica south to the department Bot., sér. 3, 13(3): 136. 1850 – Type (designated here):
of Magdalena, Colombia (Almeda & Whiffin, 1981). The herba- Pleroma gaudichaudianum (DC.) A.Gray (≡ Lasiandra
ceous species grow along rivers or streams, and along seepage gaudichaudiana DC.).
areas, while the woody ones have a preference for well-drained, = Lasiandra sect. Mucoriferae Naudin in Ann. Sci. Nat., Bot.,
more open areas along roadsides, exposed gravelly banks, and sér. 3, 13(3): 139. 1850 – Type (designated here):
disturbed, often eroded slopes (Almeda & Whiffin, 1981). Lasiandra adenostemon Schrank ex DC. (= Pleroma het-
Pilocosta has solitary, axillary, ebracteate flowers, tetram- eromallum (D.Don) D.Don).
erous, and quadrangular, prominently ribbed hypanthia. Ptero- = Lasiandra sect. Sericophyllae Naudin in Ann. Sci. Nat.,
gastra also has a 4- or 5-winged hypanthium, but it is easily Bot., sér. 3, 13(3): 142. 1850 – Type (designated here):
distinguished by the presence of terminal, dichotomous, brac- Lasiandra longistyla Naudin (= Pleroma urceolare
teate cymes. Moreover, Pilocosta and Pterogastra are not (Schrank & Mart. ex DC.) Triana).
sympatric as the latter occurs in the savannas of South = Lasiandra sect. Involucrales Naudin in Ann. Sci. Nat., Bot.,
America. sér. 3, 13(3): 144. 1850 – Type (designated here): Ple-
roma canescens (D.Don) P.J.F.Guim. & Michelang. (≡
Pleroma D.Don in Mem. Wern. Nat. Hist. Soc. 4: Diplostegium canescens D.Don ≡ Lasiandra diploste-
283, 293–295. 1823 – Type (designated here): Pleroma gium Cham.).
heteromallum (D.Don) D.Don. (≡ Melastoma heteromal- = Pleroma sect. Involucrales Triana in Trans. Linn. Soc.
lum D.Don). London, Bot. 28(1): 40. 1872 (“1871”), pro parte – Type
= Diplostegium D.Don in Mem. Wern. Nat. Hist. Soc. 4: 296. (designated here): Pleroma thereminianum (DC.) Triana
1823 – Type: Pleroma canescens (D.Don) P.J.F.Guim. (≡ Lasiandra thereminiana DC.).
& Michelang. (≡ Diplostegium canescens D.Don.). = Itatiaia Ule in Bot. Jahrb. Syst. 42: 234. 1908 – Type: Ple-
= Lasiandra DC., Prodr. 3: 127. 1828 – Type (designated roma cleistoflorum (Ule) P.J.F.Guim. (≡ Purpurella cleis-
here): Pleroma frigidulum (Schrank & Mart. ex DC.) Tri- toflora Ule ≡ Itatiaia cleistopetala Ule).
ana (≡ Lasiandra frigidula Schrank & Mart. ex DC.). = Tibouchinopsis Markgr. in Notizbl. Bot. Gart. Berlin-
= Svitramia Cham. in Linnaea 9: 445. 1835 – Type: Pleroma Dahlem 10(91): 48. 1927 – Type: Pleroma glutinosum
bandeirae P.J.F.Guim. & Michelang. (≡ Svitramia pul- (Markgr.) P.J.F.Guim. & Michelang. (≡ Tibouchinopsis
chra Cham.). glutinosa Markgr.).
= Microlepis (DC.) Miq., Comm. Phytogr.: 71. 1840, nom. Shrubs, subshrubs or trees covered with smooth to very
cons. – Type: Pleroma oleifolium (DC.) R.Romero & Ver- shaggy trichomes, sometimes gland-tipped or sessile glutinous
siane (≡ Osbeckia oleifolia [“oleaefolia”] DC. ≡ Microle- glands. Leaves opposite or rarely verticillate, decussate, blades
pis oleifolia [“oleaefolia”] (DC.) Triana). of a pair essentially equal in size, petiolate or rarely sessile,
= Lasiandra sect. Urceolaria Naudin in Ann. Sci. Nat., Bot., essentially entire, papery to coriaceous, variously pubescent,
sér. 3, 13(3): 130. 1850 – Type (designated here): glutinous or nearly glabrous, 3–5(–11)-nerved. Inflorescence
Lasiandra urceolaris Naudin (= Tibouchina australis a terminal anthotelic frondobracteose panicle or some modifi-
(Triana) Cogn.). cation of this structure in which the main and at least some lat-
= Lasiandra sect. Dendroideae Naudin in Ann. Sci. Nat., eral axes have nodes distal to the first pair of leaves on a branch
Bot., sér. 3, 13(3): 131. 1850 – Type (designated here): axis, or solitary terminal flowers; bracteoles 2(–4–6), free, var-
Pleroma asperius (Cham.) Triana (≡ Lasiandra asperior iously pubescent or glabrous, concave or plane, rarely calyp-
Cham.). trate, persistent or deciduous, often enveloping young floral
= Lasiandra sect. Sphaerocarpus Naudin in Ann. Sci. Nat., buds. Flowers perigynous; hypanthium campanulate to urceo-
Bot., sér. 3, 13(3): 133. 1850 – Type (designated here): late, pubescent externally or uncommonly glabrous. Calyx
Lasiandra tetraptera Naudin (= Pleroma benthamianum lobes 5, deciduous or rarely persistent. Petals (4–)5, purple to
Gardner). lilac, rarely white, corolla patent or campanulate. Stamens
= Lasiandra sect. Martiales Naudin in Ann. Sci. Nat., Bot., (8–)10, weakly to strongly dimorphic or isomorphic. Anthers
sér. 3, 13(3): 133. 1850 – Type (designated here): 2-celled, ventrally channeled, subulate, ventrally with an

36
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

inclined apical pore, rarely with a large pore and short anthers. ca. 1300 m, 24 Apr 2002, Leoni, L.S. & Silva, A.E. 4961
Connectives glabrous, variously prolonged and modified ven- (RB barcode 01128398!; isotypes: HUFU!, RB barcodes
trally at filament insertion into horizontal or upturned bilobed 01128696! & 01131532!).
ventral appendages glabrous or pilose. Filaments pilose or gla-
brous. Ovary superior, (4–)5-celled, with axile placentation; 3. Pleroma aemulum (Schrank & Mart. ex DC.) Triana in Trans.
style glabrous or pilose, hooked or incurved apically, typically Linn. Soc. London, Bot. 28(1): 41. 1872 (“1871”) ≡ Lasian-
declined to side of flower opposing stamens. Capsule dry, dra aemula Schrank & Mart. ex DC., Prodr. 3: 128. 1828 ≡
semiwoody, loculicidal. Seeds numerous, cochleate to Tibouchina aemula (Schrank & Mart. ex DC.) Cogn. in
elongate-cochleate and tuberculate with a terminal hilum. Martius, Fl. Bras. 14(3): 412–413. 1885 – Holotype: BRA-
Pleroma has 157 species, which are distributed mostly ZIL. Minas Gerais: Serra aurifera prope Ouro Preto et alibi,
in eastern Brazil in Mata Atlântica and Cerrado biomes, s.d., Martius, C.F.P. von s.n. (M barcode M-0165826!; iso-
and rarely in the Caatinga. Most of them are species that type: G-DC barcode G00310199!).
are widely distributed in Brazil where they occur from the = Lasiandra hygrophila Mart. ex DC., Prodr. 3: 128. 1828 ≡
Atlantic Forest to the Cerrado. Only five species reach the Tibouchina hygrophila (Mart. ex DC.) Cogn. in Martius,
northwest and west of South America. Species of Pleroma Fl. Bras. 14(3): 413–414. 1885 – Holotype: BRAZIL.
are found in forests and forest margins, river banks, high- São Paulo: in sylvis udis prope civit São Paulo, 1827,
altitude grasslands, rocky outcrops, and shrubby restinga, Martius, C.F.P. von s.n. (M barcode M-0165842!
from sea level to 2650 m. [F negative F0BN06353!]; isotype: G-DC barcode
Species of this genus have well-developed pedoconnec- G00310198!).
tives, purple or pink anthers, and if the anther connectives = Tibouchina valtherii Cogn. in Martius, Fl. Bras. 14(3):
have trichomes, these are glandular. In this last group, the 317–418. 1885 – Lectotype (designated by Oliveira
calyx lobes are persistent in fruit. & al. in Brittonia 66(4): 356. 2014): BRAZIL. Minas Ger-
When describing Pleroma, D.Don used it as feminine ais: Mariana in sylvis, 1833, Vauthier, A.C. s.n. (P barcode
(e.g., Pleroma heteromalla). However, Pleroma is derived P00116913! [F negative F0BN036176!]; isolectotypes:
from Greek and ends in -ma, a Greek ending traditionally BR barcode BR0000005530755!, P barcode P00116912
used as neuter, and this was corrected by subsequent authors [probable]!).
and monographers that cited or described new species in = Tibouchina hirsuta Cogn. in Martius, Fl. Bras. 14(4): 601.
Pleroma (e.g., Candolle, 1828a,b; Gardner, 1843; Triana, 1888 – Lectotype (designated by Oliveira & al. in Britto-
1872). Following the Code of Nomenclature for algae, nia 66(4): 356. 2014): BRAZIL. Minas Gerais: Ouro
fungi, and plants (Turland & al., 2018), we have considered Preto, 8 Jan 1884, Glaziou, A.F.M. 14801 (P barcode
Don’s use of feminine endings as an error to be corrected P00708833!; isolectotypes: B [probably destroyed; F neg-
and have followed later authors and applied a neuter ending ative F0BN016753!], BR barcode BR0000005225286!,
when appropriate. C barcode C10015071 [online image!], K barcodes
K000329061! & K000329062!, P barcode P00708832!).
Species list Other syntypes: BRAZIL. Minas Gerais: Itacolumy
[Itacolomi], 14 Feb 1884, Glaziou, A.F.M. 14793
1. Pleroma ackermannii (Cogn.) P.J.F.Guim. & Michelang., (C barcode C10015070 [online image!], K barcode
comb. nov. ≡ Tibouchina ackermannii Cogn. in Martius, K000329063!, P barcode P00708831!, R barcode
Fl. Bras. 14(3): 365. 1885 – Holotype: BRAZIL. Minas R 000009685!).
Gerais, 1832, Ackermann, G. 159 (BR barcode = Tibouchina adamantinensis Brade in Arch. Jard. Bot. Rio de
BR0000005209422!; isotype: BR barcode BR0000 Janeiro 16: 9, t. 5. 1959 – Holotype: Minas Gerais: mun.
005209125 [fragment]!). Diamantina, Água Limpa, 22 May 1955, Pereira, E. 1450
= Tibouchina serrana P.J.F.Guim. & A.B.Martins in Revista (RB barcode 00541542!; isotype: NY barcode 01477592!).
Brasil. Bot. 20(1): 15–17, fig. 8–14. 1997, syn. nov. –
Holotype: BRAZIL. São Paulo: Brotas, 6 May 1992, Gui- 4. Pleroma alatum (Cogn.) P.J.F.Guim. & Michelang.,
marães, P.J.F. 114 (UEC!; isotypes: BM barcode comb. nov. ≡ Tibouchina alata Cogn. in Candolle & Can-
BM000799540!, F barcode V0044370F [online image!], dolle, Monogr. Phan. 7: 247. 1891 – Holotype: BRAZIL.
US barcode 00731238!). Rio de Janeiro: Nova Friburgo, s.d., Raben, F.C. s.n.
(C barcode C10015053 [online image!]; isotype: BR bar-
2. Pleroma ademarii (P.J.F.Guim., R.Romero & Leoni) code BR0000005523603 [fragment]!).
P.J.F.Guim. & Michelang., comb. nov. ≡ Tibouchina
ademarii P.J.F.Guim., R.Romero & Leoni in Pabstia 5. Pleroma amoenum (Herzog) P.J.F.Guim. & Michelang.,
13(3): 2, fig. A–F. 2002 – Holotype: BRAZIL. Minas comb. nov. ≡ Tibouchina amoena Herzog in Repert.
Gerais: Araponga, Parque Estadual da Serra do Briga- Spec. Nov. Regni Veg. 7(4/6): 64. 1909 – Syntypes:
deiro, área de de transição entre a floresta estacional semi- BOLIVIA. Prov. Chiquitos: An den Gipfelfelsen des
decidual e campos de altitude, em afloramentos rochosos, Cerro San Miserate (Serrania de Santiago), ca. 900 m,

37
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

May 1907, Herzog, T.C.J. 9 (B [probably destroyed; F Encycl. 4(1): 45. 1797, nom. illeg., non Sw. 1788 ≡ Me-
negative F0BN016725!], GH). lastoma clavatum Pers., Syn. Pl. 1: 476. 1805 ≡ Rhexia
holosericea Bonpl. in Humboldt & Bonpland, Monogr.
6. Pleroma andersregnellii P.J.F.Guim. & Michelang., nom. Melast. 2: 29–31, pl. 12. 1808 ≡ Lasiandra argentea
nov. ≡ Tibouchina regnellii Cogn. in Martius, Fl. Bras. (Desr.) DC., Prodr. 3: 131. 1828 ≡ Pleroma argenteum
14(3): 305–306. 1885, non Pleroma regnellii (Miq.) Tri- (Desr.) Gardner in London J. Bot. 1: 172. 1842 ≡ Tibou-
ana 1872 – Lectotype (designated here): BRAZIL. china clavata (Pers.) Wurdack in Phytologia 7(5): 233.
Minas Gerais. Serra de Caldas, 16 Apr 1870, Regnell, 1960 – Holotype: BRAZIL. Rio de Janeiro: de la mont
A.F. III 1763 (S No. S-R-6125 [online image!]; isolecto- das capuim, Jul 1767, Commerson, P. 224 (P barcode
types: BR barcode BR0000005520640 [fragment]!, S P00678364! [F negative F0BN038242!]; isotypes: P bar-
No. S09-13191 [online image!]). codes P00708848! & P00708849 [fragment]!).
= Melastoma holosericeum Sw., Observ. Bot.: 176. 1791,
7. Pleroma angustifolium (Naudin) Triana in Trans. Linn. nom. illeg., non L. 1753 ≡ Melastoma velutinum Willd.,
Soc. London, Bot. 28(1): 47. 1872 (“1871”) ≡ Lasiandra Sp. Pl. 2(1): 584. 1799 ≡ Tibouchina holosericea Baill.,
angustifolia Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): Hist. Pl. 7: 34. 1877 – Holotype: BRAZIL. Rio de
135–136. 1850 ≡ Tibouchina angustifolia (Naudin) Janeiro: 11 Dec 1768, Banks, J. & Solander, D.C. s.n.
Cogn. in Martius, Fl. Bras. 14(3): 396–397, t. 93, fig. 1. (BM barcode BM000020467!).
1885 – Holotype: BRAZIL. Minas Gerais: districtu Serro = Tibouchina paulensis Cogn. in Candolle & Candolle,
Frio, haud procul a vico Milho Verde, s.d., De Saint- Monogr. Phan. 7: 229. 1891 – Holotype: BRAZIL.
Hilaire, A.F.C.P. B2 no. 2126 (P barcode P00116918!; São Paulo: Mogy das Cruzes [Mogi das Cruzes],
isotypes: P barcodes P00116919! & P00116920!). 19 Apr 1889, Glaziou, A.F.M. 17523 (BR barcode
= Tibouchina angustifolia var. lanceolata Cogn. in Martius, BR0000005225699!; isotypes: B [probably destroyed; F
Fl. Bras. 14(3): 396. 1885, syn. nov. – Lectotype (desig- negative F0BN016781!], K barcode K000329048!, P bar-
nated here): BRAZIL. Minas Gerais: s.d., Riedel, L. code P00708683!).
1501 (LE!; isolectotypes: P barcodes P00116922!, = Tibouchina holosericea var. albipetala Handro in Arq. Bot.
P00116923! & P00116924!). Estado São Paulo, n.s., 3: 224. 1962 – Holotype: BRA-
ZIL. São Paulo: Santo André, Alto da Serra, estrada do
8. Pleroma apparicioi (Brade) P.J.F.Guim. & Michelang., Vergueiro, no campo à beira da estrada, 18 Apr 1958,
comb. nov. ≡ Tibouchina apparicioi Brade in Arch. Jard. Handro, O. 761 (SP barcode SP001229!; isotype SPF
Bot. Rio de Janeiro 14: 213–214, t. 1. 1956 – Holotype: barcode SPF00072053!).
BRAZIL. Espírito Santo: Itaguaçu, Alto Limoeiro,
800 m a.s.l., 14 May 1946, Brade, A.C.; Pereira, 12. Pleroma asperius (Cham.) Triana in Trans. Linn. Soc.
A.B. & Duarte, A.P. 18204 (RB barcode 00659612!; iso- London, Bot. 28(1): 44. 1872 (“1871”) ≡ Lasiandra asper-
types: NY barcodes 00169348! & 01477587!, RB bar- ior Cham. in Linnaea 9: 435. 1835 ≡ Tibouchina asperior
codes 00659641! & 00541544!). (Cham.) Cogn. in Martius, Fl. Bras. 14(3): 366–367, t. 87,
fig. 1. 1885 – Lectotype (designated here): BRAZIL.
9. Pleroma arboreum Gardner in London J. Bot. 2: 351. 1843 ≡ s.d., Sellow, F. 2105 (K barcode K000329059!; isolecto-
Lasiandra arborea (Gardner) Naudin in Ann. Sci. Nat., Bot., types: B [probably destroyed; F negative F0BN016728!],
sér. 3, 13(3): 159. 1850 ≡ Tibouchina arborea (Gardner) BR barcode BR0000005522613!).
Cogn. in Martius, Fl. Bras. 14(3): 299–300, t. 67. 1885 – Lec- = Tibouchina intermedia Cogn. in Martius, Fl. Bras. 14(3):
totype (designated here): BRAZIL. [Rio de Janeiro]: Organ 367–368. 1885 – Lectotype (designated here): BRA-
Mountains, 3–400 ft., forests, Mar 1837, Gardner, G. 411 ZIL. In Brasilia australi, loco haud designato, s.d.,
(BM barcode BM000795436!; isolectotypes: E barcodes Sellow, F. 2434 (W No. 0053864 [no collector num-
E00502356 & E00531232 [online images!], K barcode ber]!; isolectotypes: B [probably destroyed; F negative
K000328999!, P barcode P00116925!). F0BN016755!], P barcode P00116948 [no collector
number]!).
10. Pleroma arenarium (Cogn.) P.J.F.Guim. & Michelang.,
comb. nov. ≡ Tibouchina arenaria Cogn. in Martius, 13. Pleroma australe Triana in Trans. Linn. Soc. London,
Fl. Bras. 14(3): 329–330. 1885 – Holotype: BRAZIL. Bot. 28(1): 47. 1872 (“1871”) ≡ Lasiandra urceolaris
Minas Gerais: Congonhas do Campo, in locis arenariis Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): 130. 1850
humidisque, s.d., Vauthier, A.C. 51 (P barcode ≡ Tibouchina australis (Triana) Cogn. in Martius,
P00116935!; isotype: BR barcode BR0000005209484 Fl. Bras. 14(3): 392–393. 1885 – Holotype: BRAZIL. In
[fragment]!). Brasilia australi, loco haud designato, s.d., De Saint-
Hilaire, A.F.C.P. D-529 (P barcode P00116942!; iso-
11. Pleroma clavatum (Pers.) P.J.F.Guim. & Michelang., types: F barcode V0063926F [fragment] [online image!],
comb. nov. ≡ Melastoma argenteum Desr. in Lamarck, P barcodes P00708743! & P00116944!).

38
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

14. Pleroma axillare (Cogn.) P.J.F.Guim. & Michelang., Fl. Bras. 14(3): 391–392. 1885 – Lectotype (designated
comb. nov. ≡ Tibouchina axillaris Cogn. in Martius, here): BRAZIL. [Rio de Janeiro]: Organ Mountains, a
Fl. Bras. 14(3): 396, t. 92. 1885 – Lectotype (designated shrub, elev. 3000 ft., growing in moist shady places,
here): BRAZIL. Rio de Janeiro: Petrópolis, pris de la Mar 1837, Gardner, G. 410 (BM barcode
Cascatinha, 3 May 1880, Glaziou, A.F.M. 11971 BM000953965!; isolectotypes: E barcodes E00531240
(BR barcode BR0000005209392!; isolectotypes: BR bar- & E00531241 [online images!], K barcodes
code BR0000005209477!, C barcode C10015054 [online K000329085! & K000329086!, NY barcode 00245677!,
image!], F barcode V0063979F [fragment] [online P barcodes P00116953! & P00116954!, US barcode
image!], G barcode G00368029!, K barcode 00120042!).
K000329093!, LE!, P barcodes P00116945! & P001 = Lasiandra tetraptera Naudin in Ann. Sci. Nat., Bot., sér.
16946!, R barcode R000009661!). 3, 13(3): 133. 1850 – Holotype: BRAZIL. Rio de Janeiro:
Serra dos Órgãos, May 1839, Guillemin, J.B.A. 887
15. Pleroma bahiense (Wurdack) P.J.F.Guim. & Michelang., (P barcode P00116958!; isotypes: P barcodes
comb. nov. ≡ Tibouchina bahiensis Wurdack in Phytolo- P00116959! & P00116960!).
gia 45(4): 321–322. 1980 – Holotype: BRAZIL. Bahia: = Tibouchina benthamiana var. punicea Cogn. in Candolle
Porto Seguro, Parque Nacional do Monte Pascoal, Pico & Candolle, Monogr. Phan. 7: 247. 1891, syn. nov. –
do Monte Pascoal, 16 Jan 1973, Santos, T.S. 2316 Holotype: BRAZIL. Rio de Janeiro: Alto Macahé,
(CEPEC barcode 00009612!; isotypes: RB barcode 8 Apr 1888, Glaziou, A.F.M. 16788 (BR barcode
00541545!, US barcode 00120039!). BR0000005529131!; isotypes: BR barcode BR0000
= Tibouchina morii Wurdack in Phytologia 45(4): 322. 1980, 0055288061!, K barcode K000329088!, LE!, P barcodes
syn. nov. – Holotype: BRAZIL. Bahia: Itamaraju, P00116955!, P00116956! & P00116957, R barcode
fazenda Pau-brasil ca. 5 km northwest of Itamaraju, R 000009663!).
19 Sep 1978, Mori, S.; Mattos Silva, L.A. & Santos,
T.S. 10726 (CEPEC barcode 00014698!; isotypes: K bar- 19. Pleroma bergianum (Cogn.) P.J.F.Guim. & Michelang.,
code K000329121!, NY barcode 00245785!, RB barcode comb. nov. ≡ Tibouchina bergiana Cogn. in Martius,
00541569!, US barcode 00120076!). Fl. Bras. 14(3): 316, t. 71. 1885 – Holotype: BRAZIL.
Minas Gerais: Congonhas do Campo, 1843, Stephan
16. Pleroma bandeirae P.J.F.Guim. & Michelang., nom. nov. s.n. (BR barcode BR0000005209514!; isotype: BR bar-
≡ Svitramia pulchra Cham. in Linnaea 9: 446. 1835, non codes BR0000005209194 [fragment]! & BR00000052
Pleroma pulchrum Triana 1872 ≡ Svitramia canescens 09521!).
Walp., Repert. Bot. Syst. 2: 132. 1843, nom. superfl. –
Lectotype (designated here): BRAZIL. Minas Gerais: 20. Pleroma blanchetianum (Cogn.) P.J.F.Guim. & Miche-
Serra do Capivari, 10 May 1819, Sellow, F. s.n. lang., comb. nov. ≡ Tibouchina blanchetiana Cogn. in
(P barcode P02274322!; isolectotypes: K barcode Martius, Fl. Bras. 14(3): 351. 1885 – Lectotype (desig-
K000530433!, P barcode P02274321!). nated here): BRAZIL. Bahia: Jacobina, 1842, Blanchet,
This species is dedicated to Maria do Carmo Vaughan J.S. 3622 (BM barcode BM000020477!; isolectotypes:
Bandeira (1902–1992), Brazilian botanist, who worked for BR barcodes BR0000005224456! & BR0000005
about ten years at the Botanical Garden in Rio de Janeiro, 224784!, C barcode C10015055!, G barcode
Brazil. G00368026!, G-DC barcode G00319124!, K barcode
K000329042!, LE!, P barcodes P00116971!, P001
17. Pleroma barnebyanum (Wurdack) P.J.F.Guim. & Miche- 16972! & P00708726!, W!).
lang., comb. nov. ≡ Tibouchina barnebyana Wurdack in
Brittonia 33(3): 304–305, fig. 5. 1981 – Holotype: BRA- 21. Pleroma boraceiense (Brade) P.J.F.Guim. & Justino in
ZIL. Bahia: 2–3 km SW of Mucugê on road to Cascavel, Feddes Repert. 129: 235. 2018 ≡ Tibouchina boraceiensis
Serra do Sincorá, elev. 950 m, 17 Feb 1977, Harley, R.M.; Brade in Arq. Bot. Estado São Paulo 3: 210, t. 53,
Mayo, S.J.; Storr, R.M.; Santos, T.S. & Pinheiro, fig. 8–15. 1958 – Holotype: BRAZIL. [São Paulo]:
R.S. 18786 (CEPEC barcode 00019691!; isotypes: Santos, Serra do Mar, região de Boraceia, do rio Guara-
AAU ImageID 3475 [online image!], K barcodes tuba, 900 a.s.l., 15 Nov 1950, Kuhlmann, M. 2336
K000329149! & K000329150!, NY barcode 00169346!, (SP barcode SP001227!).
U barcode U 0004174 [online image!], US barcode
00120040!). 22. Pleroma boudetii (P.J.F.Guim. & R.Goldenb.) P.J.F.Guim.
& Michelang., comb. nov. ≡ Tibouchina boudetii P.J.F.
18. Pleroma benthamianum Gardner in Bot. Mag. 69: Guim. & R.Goldenb. in Kew Bull. 56: 989–993, fig. 1, 2.
t. 4007. 1843 ≡ Lasiandra benthamiana (Gardner) Nau- 2001 – Holotype: BRAZIL. Espírito Santo: Santa Teresa,
din in Ann. Sci. Nat., Bot., sér. 3, 13(3): 159. 1850 ≡ Vargem Alta, 28 Jan 1986, Boudet Fernandes, H.Q. 1809
Tibouchina benthamiana (Gardner) Cogn. in Martius, (MBML barcode MBML000764!; isotypes: UPCB!, US!).

39
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

23. Pleroma bracteolatum (J.G.Freitas, A.K.A.Santos & R.P. (P barcode P00116996!; isotype: G-DC barcode
Oliveira) P.J.F.Guim. & Michelang., comb. nov. ≡ Tibou- G00310481 [fragment]!).
china bracteolata J.G.Freitas, A.K.A.Santos & R.P.Oli- = Tibouchina valtherii var. minor Hoehne in Anexos Mem.
veira in Syst. Bot. 37(1): 190, fig. 1, 2E–F, 3A–B. 2012 Inst. Butantan, Secc. Bot. 1(5): 69. 1922, syn.nov. –
– Holotype: BRAZIL. Bahia: Lençóis, próximo ao rio Holotype: BRAZIL. Minas Gerais: Serra do Garimpo,
Lençóis, 30 Apr 2003, Silva-Pereira 55 (HUEFS barcode 13 Jan 1921, Hoehne, F.C. s.n. (SP barcode SP001237!).
HUEFS089925!).
28. Pleroma carvalhoi (Wurdack) P.J.F.Guim. & Michelang.,
24. Pleroma caissarum F.S.Mey. in Kew Bull. 69(3): 9527: comb. nov. ≡ Tibouchina carvalhoi Wurdack in Phytolo-
2–8, fig. 1, 2A–D. 2014 – Holotype: BRAZIL. Santa Cat- gia 53(2): 134–135. 1983 – Holotype: BRAZIL. Bahia:
arina, Município de Joinville, às margens da BR-101, Serra do Sincora ca. 15 km NW of Mucugê on road to
26 18′44.26′′S, 48 52′25.67′′W, 24 Feb 2013, Meyer, F. Guiné and Palmeiras, elev. 1300–1500 m, 26 Mar 1980,
& Schmitz 1578 (UPCB; isotype: K). Harley, R.M.; Bromley, G.L.; Carvalho, A.M. de
& Martinelli, G. 20993 (CEPEC barcode 00025558!; iso-
25. Pleroma candolleanum (Mart. ex DC.) Triana in Trans. types: K barcode K000329114!, US barcode 00120046!).
Linn. Soc. London, Bot. 28(1): 44, t. 3, fig. 31c. 1872
(“1871”) ≡ Lasiandra candolleana Mart. ex DC., Prodr. 29. Pleroma castellense (Brade) P.J.F.Guim. & Michelang.,
3: 129. 1828 ≡ Tibouchina candolleana (Mart. ex DC.) comb. nov. ≡ Tibouchina castellensis Brade in Arch. Jard.
Cogn. in Martius, Fl. Bras. 14(3): 339–340. 1885 – Holo- Bot. Rio de Janeiro 14: 215–216, t. 3. 1956 – Holotype:
type: BRAZIL. 1827, Martius, C.F.P. s.n. (G-DC barcode BRAZIL. Espírito Santo: Castelo, Forno Grande, 12 Aug
G00310191! [F negative F0BN07905!]). 1948, Brade, A.C. 19262 (RB barcode 00659644!; iso-
= Lasiandra setulosa Spring ex Mart. in Flora 22(1, Beibl.): types: RB barcodes 00541549! & 00659614!).
9. 1839 ≡ Pleroma setulosum Spring ex Triana in Trans.
Linn. Soc. London, Bot. 28(1): 44. 1872 (“1871”) ≡ 30. Pleroma cecilianum P.J.F.Guim. & Oliveira da Silva in
Tibouchina setulosa Spring ex Cogn. in Martius, Phytotaxa 205(1): 52–57, fig. 1, 2. 2015 – Holotype:
Fl. Bras. 14(3): 338–339, t. 80, fig. 1. 1885, syn. nov. – BRAZIL. Rio de Janeiro: Campos dos Goytacazes, Dis-
Lectotype (designated here): BRAZIL. Bahia. Cabula, trito de Ibitioca, fazenda Pedra Negra, sopé do morro de
Oct 1835, Luschnath, B. 20 (BR barcode BR00000 Itaoca (morro do Rato), elevation 40 m, 23 Mar 2001,
05224944!; isolectotype: LE!). fl., fr., Braga, J.M.A. 6648 (RB!; isotypes: K!, NY!,
R!, SP!).
26. Pleroma canescens (D.Don) P.J.F.Guim. & Michelang.,
comb. nov. ≡ Diplostegium canescens D.Don. in Mem. 31. Pleroma cinereum (Cogn.) P.J.F.Guim. & Michelang.,
Wern. Nat. Hist. Soc. 4: 296. 1823 ≡ Lasiandra diploste- comb. nov. ≡ Tibouchina cinerea Cogn. in Martius,
gium Cham. in Linnaea 9: 438. 1835, nom. illeg. superfl. Fl. Bras. 14(3): 416–417, t. 95, fig. 2. 1885 – Lectotype
≡ Pleroma diplostegium Triana in Trans. Linn. Soc. (designated here): BRAZIL. São Paulo: Campos da
London, Bot. 28(1): 40. 1872 (“1871”), nom. illeg. Bocaina, 11 Feb 1876, Glaziou, A.F.M 8367 (P barcode
superfl. ≡ Tibouchina canescens (D.Don) Cogn. in Mar- P00121035!; isolectotypes: B [probably destroyed; F neg-
tius, Fl. Bras. 14(3): 298–299, t. 66. 1885 – Lectotype ative F0BN016733!], BR barcode BR0000005225125!,
(designated here): BRAZIL. s.d., Sellow, F. s.n. C barcode C10015057!, K barcode K000329110!, LE!,
(K barcode K000328998!; isolectotypes: K barcode P barcode P00121036!, R barcode R 000009655!, S
K000328997!, P barcode P00116986!). No. S-R-6112 [fragment] [online image!]).
= Pleroma pohlianum O.Berg. ex Triana in Trans. Linn. Soc.
London, Bot. 28(1): 46–47. 1872 (“1871”) – Lectotype 32. Pleroma claussenii (Naudin) Triana in Trans. Linn. Soc.
(designated here): BRAZIL. Minas Gerais: Mundo Velho, London, Bot. 28(1): 41. 1872 (“1871”) ≡ Lasiandra
s.d., Pohl, J.B.E. D.1274-3489 (NY barcode 00941638!). claussenii Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3):
152–153. 1850 ≡ Tibouchina claussenii (Naudin) Cogn.
27. Pleroma cardinale (Bonpl.) Triana in Trans. Linn. Soc. in Martius, Fl. Bras. 14(3): 310–311. 1885 – Holotype:
London, Bot. 28(1): 44. 1872 (“1871”) ≡ Rhexia cardina- BRAZIL. Minas Gerais: 1841, Claussen, P.C.D. 42
lis Bonpl. in Humboldt & Bonpland, Monogr. Melast. 2: (P barcode P00121046!; isotypes: BR barcodes
96, pl. 37. 1818 ≡ Melastoma cardinale (Bonpl.) Spreng., BR0000005224845!, BR0000005264186! & BR000
Syst. Veg. 2: 298. 1825 ≡ Chaetogastra cardinalis 0005264254!, F barcodes V0063567F & V0063566F
(Bonpl.) DC., Prodr. 3: 134. 1828 ≡ Lasiandra cardinalis [both fragments] [online images!], G-DC barcode
(Bonpl.) Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): 132. G00319076!, M barcode M-0005213!, P barcode
1850 ≡ Tibouchina cardinalis (Bonpl.) Cogn. in Martius, P00121047!).
Fl. Bras. 14(3): 371–372. 1885 – Holotype: BRAZIL. = Tibouchina claussenii var. grandiflora Cogn. in Martius,
Minas Gerais: Gran-Pará, s.d., Bonpland, A.J.A. s.n. Fl. Bras. 14(3): 311. 1885, syn.nov. – Holotype:

40
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

BRAZIL. s.d., Ackermann, G. 964 (B [probably destroyed; barcode BR0000005224814 [fragment]!, C barcode
F negative F0BN016735!]). C10015058 [online image!], K barcodes K000329089!
= Tibouchina cogniauxii Glaz. ex Cogn. in Candolle & Can- & K000329090!, LE!, P barcode P00376767!, R barcode
dolle, Monogr. Phan. 7: 1175. 1891, syn. nov. – Holotype: R 000009662!).
BRAZIL. Minas Gerais: Morro de Pires ad Faria prope Other syntype: BRAZIL. Rio de Janeiro: Ayuruoca, au
Sabara, 24 Jan 1891, Glaziou, A.F.M. 18231 (P barcode sommet du Pico do Papagaio, 19 Apr 1878, Glaziou,
P00121049!; isotypes: BR barcode BR0000005224487 A.F.M. 9460 (K barcode 329090!, P barcode
[no collector number]!, R!, RB barcode 00541552!). P00376765!).

33. Pleroma cleistoflorum (Ule) P.J.F.Guim., Oliveira da 38. Pleroma costatocalyx F.S.Mey., L.Kollmann & R.Gold-
Silva & Michelang. in Phytotaxa 186(4): 225–226, fig. 1, enb. in Phytotaxa 282(3): 198–202, fig. 1, 2. 2016 –
2. 2014 ≡ Purpurella cleistoflora Ule in Ber. Deutsch. Holotype: BRAZIL. Espírito Santo: Mantenópolis (culti-
Bot. Ges. 13: 418, pl. 32. 1895 ≡ Itatiaia cleistopetala vated at Santa Teresa, at the property of Ludovic Koll-
Ule in Bot. Jahrb. Syst. 42: 235. 1908 – Lectotype (desig- mann), May 2015, (fl.), Kollmann, L. & Fontana,
nated by Silva & al. in Phytotaxa 186(4): 225. 2014): A.P. 12977 (MBML barcode MBML050000 [online
BRAZIL. Rio de Janeiro: Serra do Itatiaia [label], “habitat image!]; isotypes: NY, P, RB barcode 01128101!).
in campis ad Serram Itatiaiae, in altitudine 2100–2400
metrum” [protologue], Ule, E.H.G 3349 (HBG barcode 39. Pleroma crassirame (Cogn.) P.J.F.Guim. & Michelang.,
HBG-522987 [online image!]; isolectotypes: HBG bar- comb. nov. ≡ Tibouchina crassiramis Cogn. in Bot. Jahrb.
codes HBG522988 & HBG522989 [probably] [online Syst. 21(4): 446. 1896 – Lectotype (designated here):
images!]). BRAZIL. Goiás: Bäumchen Strauch in der Serra dos Pyr-
eneos, 1000–1100 m, Oct 1892, Ule, E.H.G. 2905 (HBG
34. Pleroma clidemioides Berg. ex Triana in Trans. Linn. barcode HBG-506568 [online image!]; isolectotypes: BR
Soc. London, Bot. 28(1): 43. 1872 (“1871”) ≡ Tibouchina barcode BR0000005520626 [fragment]!, R barcode
clidemioides (Berg. ex Triana) Cogn. in Martius, Fl. Bras. R 000186986 [fragment]!).
14(3): 398–399. 1885 – Lectotype (designated here):
BRAZIL. Espírito Santo: ad urbem Victoria, s.d., 40. Pleroma cristatum (Brade) P.J.F.Guim. & Michelang.,
Sellow, F. 432 (P barcode P00708747!; isolectotypes: B comb. nov. ≡ Tibouchina cristata Brade in Arq. Inst.
[probably destroyed; F negative F0BN016737!], K bar- Biol. Veg. 4(1): 76–77, fig. 8. 1938 – Holotype: BRA-
code K000329094!). ZIL. Rio de Janeiro: Frade de Macahé, Jul 1937, Brade,
A.C. 15865 (RB barcode 00659605!; isotypes: RB bar-
35. Pleroma collinum (Naudin) Triana in Trans. Linn. Soc. codes 00541553!, 00659637! & 00659636!).
London, Bot. 28(1): 42. 1872 (“1871”) ≡ Lasiandra col-
lina Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): 41. Pleroma cryptadenum (Gleason) P.J.F.Guim. & Miche-
148–149. 1850 ≡ Tibouchina collina (Naudin) Cogn. in lang., comb. nov. ≡ Tibouchina cryptadena Gleason in
Martius, Fl. Bras. 14(3): 318–319. 1885 – Holotype: Fieldiana, Bot. 28(2): 426–428. 1952 – Holotype: VENE-
BRAZIL. Minas Gerais: 1816–1821, De Saint-Hilaire, ZUELA. Mérida: between Beguilla and Mucuchachí, alt.
A.F.C.P., Catal. D no. 67 (P barcode P00121397!; iso- 1065–2430 m, 4 May 1944, Steyermark, J.A. 56307
types: P barcodes P00121398! & P00121399!). (F barcode V0063981F [online image!]; isotypes: NY bar-
code 00245757!, VEN barcode 26260 [online image!]).
36. Pleroma comosum (J.G.Freitas, A.K.A.Santos & R.P.
Oliveira) P.J.F.Guim. & Michelang., comb. nov. ≡ Tibou- 42. Pleroma cucullatum F.S.Mey., Fraga & R.Goldenb. in
china comosa J.G.Freitas, A.K.A.Santos & R.P.Oliveira Phytotaxa 348(4): 237–242, fig. 2, 3. 2018 – Holotype:
in Syst. Bot. 37(1): 190 –196, fig. 2A–D, 3C–D, BRAZIL. Espírito Santo: Nova Venécia, Área de Prote-
5. 2012 – Holotype: BRAZIL. Bahia: Mucugê, Serra do ção Ambiental–APA da Pedra do Elefante, 19 Feb
Esbarrancado, 16 Apr 2005, Conceição, A.A. 1273 2008, Kollmann, L.; Fraga, C.N. & Fontana, A.P. 10701
(HUEFS barcode HUEFS094535!). (UPCB; isotypes: CEPEC, RB).

37. Pleroma cordifolium (Cogn.) P.J.F.Guim. & Michelang., 43. Pleroma decemcostatum (Cogn.) P.J.F.Guim. & Miche-
comb. nov. ≡ Tibouchina cordifolia Cogn. in Martius, lang., comb. nov. ≡ Tibouchina decemcostata Cogn. in
Fl. Bras. 14(3): 393. 1885 – Lectotype (designated Martius, Fl. Bras. 14(3): 331–332, t. 78, fig. 1. 1885 – Lec-
here): BRAZIL. Rio de Janeiro: Engenho da Serra, totype (designated here): BRAZIL. Minas Gerais: s.d.,
Rodrigues Jardim à 1.900 metros environ altitude avec Ackermann, G. s.n. (BR barcode BR0000005224494!).
Netto, Schwacke & Rangel, 10 Apr 1879, Glaziou,
A.F.M. 10770 (P barcode P00376766!; isolectotypes: B 44. Pleroma dendroides (Naudin) Triana in Trans. Linn. Soc.
[probably destroyed; F negative F0BN016738!], BR London, Bot. 28(1): 44. 1872 (“1871”) ≡ Lasiandra

41
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

dendroides Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): = Tibouchina hieronymi Cogn. in Candolle & Candolle,
132. 1850 ≡ Tibouchina dendroides (Naudin) Cogn. in Monogr. Phan. 7: 1175. 1891 – Lectotype (designated
Martius, Fl. Bras. 14(3): 369–370. 1885 – Holotype: here): BRAZIL. Santa Catarina: campos de Boa Vista,
BRAZIL. s.d., Martius, C.F.P. von, Herb. Fl. Bras. Serra do Mar alt. 1000 m, Dec 1886, Ule, E.H.G. 577
961 (P barcode P00481670!; isotypes: B [probably (BR barcode BR0000005224821!; isolectotype: HBG
destroyed; F negative F0BN016742!], BR barcode barcode HBG-522834 [online image!]).
BR0000005520954!, G-DC barcode G00318922!, M Other syntypes: BRAZIL. Santa Catarina: an der Costa der
barcodes M-0165832! & M-0165833!). Serra Geral, Jan 1891, Ule, E.H.G. 1445 (B [probably
= Tibouchina rotundifolia Cogn. in Martius, Fl. Bras. 14(3): destroyed; F negative F0BN016752!, no collection data,
370–371. 1885, pro parte, syn. nov. – Lectotype (des- label partially photographed], BR barcode BR0000
ignated here): BRAZIL. Minas Gerais: 1840, 005225156!, HBG barcodes HBG-522835 & HBG-
Claussen, P. 296a (BR barcode BR0000005225965!). 522836 [online images!], P barcode P00708830!).

45. Pleroma discolor (Brade) P.J.F.Guim. & Michelang., 48. Pleroma dusenii (Cogn.) P.J.F.Guim. & Michelang.,
comb. nov. ≡ Tibouchina discolor Brade in Arq. Inst. comb. nov. ≡ Tibouchina dusenii Cogn. in Ark. Bot. 9:
Biol. Veg. 4(1): 75–76, fig. 6. 1938 – Holotype: BRA- 8. 1910 – Holotype: BRAZIL. Paraná: Serra do Mar, Mar-
ZIL. Rio de Janeiro: Santa Maria Magdalena, Serra da umbi, in Dickichten in einer Höhe von etwa 1000 m,
Furquilha 1400 m, 4 Mar 1935, Santos Lima, J. & Brade, 13 Feb 1904, Dusén, P.K.H. 3777 (BR barcode
A.C. 14268 (RB barcode 00541554!; isotypes: NY bar- BR0000005521333!; isotypes: RB barcode 00541557!,
codes 00245760! & 01477588!, RB barcode 00556708!, S No. S-R-9198!).
SP barcode SP002914!). = Tibouchina catharinensis Brade in Sellowia 12: 139, fig. 2,
= Tibouchina discolor var. alba Brade in Arq. Inst. Biol. Veg. 14–21. 1960 – Holotype: BRAZIL. Santa Catarina: Horto
4: 76. 1938, syn. nov. – Holotype: BRAZIL. Rio de Florestal do Instituto Nacional do Pinho: Ibirama, 700 m
Janeiro: Santa Maria Magdalena, Serra da Furquilha a.s.l., 26 Jan 1957, Klein, R.M. 2205 (HBR barcode
1500 m, 4 Mar 1935, Santos Lima, J. & Brade, HBR0027671 [online image!]; isotypes: M, NY barcode
A.C. 14269 (RB barcode 00541555!; isotypes: RB bar- 00658698!, US barcode 00120047!).
codes 00556708! & 00556709 [fragment]!).
49. Pleroma echinatum Gardner in London J. Bot. 2: 350.
46. Pleroma divaricatum (Cogn.) P.J.F.Guim. & Michelang., 1843 ≡ Lasiandra echinata (Gardner) Naudin in Ann.
comb. nov. ≡ Tibouchina divaricata Cogn. in Martius, Sci. Nat., Bot., sér. 3, 13(3): 159. 1850 ≡ Pleroma gard-
Fl. Bras. 14(3): 397–398. 1885 – Holotype: BRAZIL. nerianum Triana in Trans. Linn. Soc. London, Bot.
Minas Gerais: St-Hilaire, A.F.C.P., Catal. B’ no. 1446 28(1): 42. 1872 (“1871”), nom. illeg. superfl. ≡ Tibou-
(P barcode P00708656!; isotypes: BR barcode china gardneriana (Triana) Cogn. in Martius, Fl. Bras.
BR0000005530649 [fragment]!, F barcode V0063982F 14(3): 319, t. 72, fig. 1. 1885, nom. illeg. – Lectotype
[fragment]!, P barcodes P00708655! & P00708657!). (designated here): BRAZIL. Rio de Janeiro: Organ
Mountains at an elevation of nearly 6000 ft., May 1837,
47. Pleroma dubium (Cham) P.J.F.Guim. & Michelang., Gardner, G. 404 (K barcode K000329014!; isolecto-
comb. nov. ≡ Lasiandra dubia Cham. in Linnaea 9: types: BM barcode BM000953940!, BR barcode
437. 1835 ≡ Tibouchina dubia (Cham.) Cogn. in Martius, BR0000005224890 [fragment]!, K barcode K000
Fl. Bras. 14(3): 415. 1885 – Lectotype (designated by 329015!).
Guimarães & Oliveira in Martins & al., Fl. Fanerog. = Tibouchina alba Cogn. in Candolle & Candolle, Monogr.
Estado São Paulo 6: 134–135. 2009): BRAZIL. São Phan. 7: 210. 1891, syn. nov. – Lectotype (designated
Paulo: 1836, Sellow, F. 4211 (K barcode K000895005 here): BRAZIL. Rio de Janeiro: Alto Macahé, 6 Feb
[no collector number]!; isolectotypes: B [probably 1888, Glaziou, A.F.M. 16805 (BR barcode BR00
destroyed; F negative F0BN016743!], BR barcode 00005224913!; isolectotypes: B [probably destroyed; F
BR0000005263523!, P barcodes P00708984! & P007 negative F0BN016723!], BR barcode BR0000005
08985!, S No. S09-13203 [fragment]!). 224586!, C barcode C10015067!, F barcode V00
= Tibouchina glazioviana Cogn. in Martius, Fl. Bras. 14(3): 63977F!, GH barcode 00073255 [online image!], K bar-
325–326, t. 72, fig. 2. 1885, syn. nov. – Lectotype (desig- code K000329013!, LE!, NY barcode 00169347!, P bar-
nated by Guimarães & Oliveira in Martins & al., codes P00116914!, P00116915! & P00116916!, R
Fl. Fanerog. Estado São Paulo 6: 134–135. 2009): BRA- barcode R 000009286!).
ZIL. São Paulo: Campos da Bocaina, 10 Sep 1879,
Glaziou, A.F.M. 11954 (C barcode C10015068!; isolecto- 50. Pleroma edmundoi (Brade & Markgraf) P.J.F.Guim.
types: BR barcode BR0000005224593!, F barcode & Michelang., comb. nov. ≡ Pterolepis edmundoi Brade
V0063984F [fragment]!, K barcode K000329023!, LE!, & Markgraf in Arch. Jard. Bot. Rio de Janeiro 17:
P barcode P02441081!, R barcode R 000009713!). 45–46, fig. 2 11–19. 1959–1961 (= in Willdenowia 2

42
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

(5): 771. 1961) ≡ Tibouchina bradeana Renner in Nordic Fl. Bras. 14(3): 342–343, t. 81. 1885 – Holotype: BRA-
J. Bot. 14(1) 101. 1994 – Holotype: BRAZIL. Bahia: Sal- ZIL. Rio de Janeiro: Trovasi sulle montagne d’Estrella,
vador, Itapoan, Lagoa do Abaeté, 2 Sep 1956, Pereira, E. e fiorisce come la precedente, alla quale rassomiglia mol-
s.n. (HB; isotypes: RB barcode 00541535!, US barcode tissimo, s.d., Raddi, G. s.n. (PI; isotype: FI barcode
00118371!). FI005212 [online image!]).
Brade (1959–1961) published new Brazilian species of = Tibouchina scrobiculata Cogn. in Martius, Fl. Bras. 14(3):
Melastomataceae, manuscript submitted in December 1960, 341–342. 1885, syn. nov. – Lectotype (designated
but Brade & Markgraf (1961) also published many of these here): BRAZIL. Rio de Janeiro: in via ad Real fazenda
species plus others in another journal. In both articles, they de Santa Cruz, Feb–Mar [without year], Schott,
received the status of species nova. For this reason, both works H.W. & Pohl, J.B.E. 1176 [Kat. no. 3960] (BR barcode
are cited. BR0000005224524!; isolectotype: W No. 0053977!
[F negative F0BN032315!]).
51. Pleroma eichleri (Cogn.) P.J.F.Guim. & Michelang., Other syntype: BRAZIL. s.l., s.d., Sellow, F. 114
comb. nov. ≡ Tibouchina eichleri Cogn. in Martius, (B [probably destroyed]).
Fl. Bras. 14(3): 370. 1885 – Holotype: BRAZIL. São = Tibouchina scrobiculata var. membranifolia Cogn. in Mar-
Paulo: Bom Jardim, Campos da Bocaina, 5 Jan 1876, tius, Fl. Bras. 14(3): 342. 1885, syn. nov. – Lectotype
Glaziou, A.F.M. 8359 (P barcode P00121414!; isotype: (designated here): BRAZIL. Rio de Janeiro: in dumetis
B [probably destroyed; F negative F0BN016744!]). ad Mathias Ramos, s.d., Pohl, J.B.E. 3959 (W No.
0071708! [F negative F0BN032317!]; isolectotype: BR
52. Pleroma elegans Gardner in London J. Bot. 2: 350. 1843 ≡ BR0000005224524 [fragment]!).
Lasiandra elegans (Gardner) Naudin in Ann. Sci. Nat., Other syntype: BRAZIL. s.d., Sellow, F. s.n. (B [probably
Bot., sér. 3, 13(3): 159. 1850 ≡ Tibouchina elegans destroyed]).
(Gardner) Cogn. in Martius, Fl. Bras. 14(3): 323–324. 1885 = Tibouchina scrobiculata var. obtusifolia Cogn. in Martius,
– Lectotype (designated here): BRAZIL. Rio de Janeiro: Fl. Bras. 14(3): 342. 1885, syn. nov. – Lectotype (desig-
Organ Mountains, May 1837, Gardner, G. 405 (K barcode nated here): BRAZIL. Queiroz, s.d., Pohl, J.B.E. 70
K000329022!; isolectotype: BM barcode BM000953939!). (BR barcode BR0000005224852!; isolectotypes: W
= Lasiandra weddellii Naudin in Ann. Sci. Nat., Bot., sér. No. 2004-0008175! [F negative F0BN032316!]).
3, 13(3): 151. 1851 ≡ Tibouchina weddellii (Naudin)
Cogn. in Martius, Fl. Bras. 14(3): 311–312. 1885 ≡ Pler- 54. Pleroma ferricolum A.L.F.Oliveira, R.Romero & P.J.F.
oma weddellii (Naudin) Triana in Trans. Linn. Soc. Guim. in Brittonia 66(4): 354, fig. 1. 2014 – Holotype:
London, Bot. 28: 42. 1872 (“1871”), syn. nov. – Holo- BRAZIL. Minas Gerais: Nova Lima, Capão Xavier,
type: BRAZIL. Rio de Janeiro: in montibus Serra d’Es- 13 Mar 2001, Pimentel, M.; Madsen, V.; Barros, A.M.;
trella, Braziliae australis prope Bell Monte, 1844, Almeida, L. & Viana, M.C. 36 (BHCB!; isotypes:
Weddell, H.A. 757 (P barcode P00121402!; isotypes: HUFU!, MO!, RB!, US!).
B [probably destroyed; F negative F0BN026117!], BR
barcode BR0000005224500 [fragment]!, F barcode 55. Pleroma fissinervium (Schrank & Mart. ex DC.) Gardner in
V0063574F [fragment] [online image!], G-DC barcode London J. Bot. 2: 352. 1843 ≡ Lasiandra fissinervia Schrank
G00318950!, P barcode P00121403!). & Mart. ex DC., Prodr. 3: 130. 1828 ≡ Tibouchina fissinervia
= Tibouchina weddellii var. angustifolia Cogn. in Martius, (Schrank & Mart. ex DC.) Cogn. in Martius, Fl. Bras. 14(3):
Fl. Bras. 14(3): 312. 1885, syn. nov. – Lectotype (des- 343–344. 1885 – Holotype: BRAZIL. Minas Gerais: s.d.,
ignated here): BRAZIL. Rio de Janeiro: Serra da Martius, C.F.P. von s.n. (M barcode M-0165834!; isotype:
Estrella, 27 Jan 1875, Glaziou, A.F.M. 7611 (BR G-DC barcode G00310187 [fragment]!).
barcode BR0000005224838!; isolectotypes: C barcode
C10015060!, P barcodes P00121400! & P00121401!). 56. Pleroma floribundum (Cogn.) P.J.F.Guim. & Michelang.,
= Tibouchina weddellii var. parvifolia Cogn. in Candolle comb. nov. ≡ Tibouchina floribunda Cogn. in Martius,
& Candolle, Monogr. Phan. 7: 1175. 1891, syn. nov. – Fl. Bras. 14(3): 312–313. 1885 – Lectotype (designated
Lectotype (designated here): BRAZIL. Rio de Janeiro: here): BRAZIL. Rio de Janeiro: 28 Aug 1866, Glaziou,
Serra dos Órgãos, Dec 1888, Moura, J.T. 358 A.F.M. 587 (BR barcode BR0000005264162!; isolecto-
(BR barcode BR0000005225163!). types: BR barcode BR0000005263509!, C barcode
Other syntype: BRAZIL. Rio de Janeiro: Serra dos Órgãos, C10015062!, P barcode P00708978!).
Feb 1890, Moura, J.T. s.n. (RB barcode 00541587!). Other syntypes: BRAZIL. Rio de Janeiro: Nova Friburgo,
11 Aug 1881, Glaziou, A.F.M. 12685 (B [probably
53. Pleroma estrellense (Raddi) P.J.F.Guim. & Michelang., destroyed; F negative F0BN016746!], BR barcodes
comb. nov. ≡ Rhexia estrellensis Raddi in Mem. Mat. BR0000005224869! & BR0000005225194!, C barcode
Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis. 18(2): 388. C10015061!, K barcode K000329020!, LE!, P barcodes
1820 ≡ Tibouchina estrellensis (Raddi) Cogn. in Martius, P00708976! & P00708977!, R barcode R 000009292!).

43
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

= Tibouchina ovata Cogn. in Candolle & Candolle, Monogr. 59. Pleroma foveolatum (Naudin) Triana in Trans. Linn. Soc.
Phan. 7: 211. 1891, syn. nov. – Lectotype (designated London, Bot. 28(1): 41. 1872 (“1871”) ≡ Lasiandra
here): BRAZIL. Rio de Janeiro: Alto Macahé de foveolata Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3):
N. Friburgo, 19 Dec 1887, Glaziou, A.F.M. 16803 148. 1850 ≡ Tibouchina foveolata (Naudin) Cogn. in
(BR barcode BR0000005225187!; isolectotypes: B Martius, Fl. Bras. 14(3): 307–308. 1885 – Holotype:
[probably destroyed; F negative F0BN016778!], BR bar- BRAZIL. Minas Gerais: Serra Negra, s.d., De Saint-
code BR0000005224531!, C barcodes C10015064! Hilaire, A.F.C.P. B56 (P P00121410!; isotypes: BR
& C10015063!, K barcode K000329018!, LE!, P barcode barcode BR0000005520978 [fragment]!, F barcode
P00708676!, R barcode R 000009282!). V0063570F [fragment]!, P barcodes P00121411! &
P00121412!).
57. Pleroma formosum (Cogn.) P.J.F.Guim. & Michelang., = Tibouchina organensis Cogn. in Martius Fl. Bras. 14(3):
comb. nov. ≡ Tibouchina formosa Cogn. in Martius, 308–309. 1885 – Lectotype (designated here): BRA-
Fl. Bras. 14(3): 348–349, t. 83. 1885 – Lectotype (desig- ZIL. Rio de Janeiro: Serra dos Orgãos, s.d., Saldanha, J.
nated here): BRAZIL. Minas Gerais: Caldas, 18 Jan 7363 (K barcode K000329009!; isolectotypes: B [proba-
1870, Regnell, A.F. III 16 (BR barcode BR000 bly destroyed; F negative F0BN016777!], P barcode
0005224548!; isolectotypes: C barcodes C10015065! P00708680!).
& C10015066!, P barcode P00708979!, S Nos. S-R- = Tibouchina organensis var. silvestris Brade in Arq. Inst.
6116! & S08-20241!). Biol. Veg. 4(1): 72–73. 1938, syn. nov. – Holotype:
BRAZIL. Rio de Janeiro: Itatiaia km 12, Aug 1933,
58. Pleroma fothergillae (Schrank & Mart. ex DC.) Triana in Brade, A.C. 12682 (RB barcode 00541574!; isotype:
Trans. Linn. Soc. London, Bot. 28(1): 42. 1872 (“1871”) RB barcode 00556743!).
≡ Rhynchanthera fothergillae Schrank & Mart. ex DC., = Tibouchina camposportoi Brade in Arq. Inst. Biol. Veg. 4
Prodr. 3: 108. 1828 ≡ Lasiandra fothergillae (Schrank (1): 73–74, t. 3. 1938 (“campos-portoi”) – Holotype:
& Mart. ex DC.) Cham. in Linnaea 9: 437. 1835 ≡ Tibou- BRAZIL. São Paulo: Campos do Jordão, 20 Feb 1937,
china fothergillae (Schrank & Mart. ex DC.) Cogn. in Campos Porto, P. 3253 (RB barcode 00659604!; iso-
Martius, Fl. Bras. 14(3): 320–321, t. 73. 1885 – Holotype: types: NY barcodes 00245756! & 01477586!, RB bar-
BRAZIL. Minas Gerais?: sylvis locorum supra campos code 00541548!, SP barcode SP002921!).
prope S. Pauli Civitatem et alibi, s.d., Martius, C.F.P.
von 1026 (M barcode M-0165835!). 60. Pleroma fragae L.Kollmann & R.Goldenb. in Brittonia
= Lasiandra kunthiana DC., Prodr. 3: 128. 1828 ≡ Pleroma 68(1): 38–41, fig. 1, 2. 2016 – Holotype: BRAZIL. Espír-
fothergillii var. kunthiana (DC.) Triana in Trans. Linn. ito Santo: Colatina, Sapucaia, Alto Moacir, Prop. Laulau,
Soc. London, Bot. 28(1): 42. 1872 (“1871”) ≡ Tibouchina 300 m, 18 Apr 2008 (fl), Kollmann, L.; Fontana,
moricandiana var. kunthiana (DC.) Cogn. in Martius, A.P. & Brahim, K. 10918 (MBML barcode MBML
Fl. Bras. 14(3): 322. 1885, syn. nov. – Lectotype (desig- 048397; isotypes: CEPEC barcode 00120751 [online
nated here): BRAZIL. Rio de Janeiro: s.d., Langsdorff, image!], K, MO, NY barcode 02688306!, P, RB barcode
G.H. von s.n. (G-DC barcode G00310200!). 01043778!, UPCB).
= Pleroma moricandianum f. parvifolium Wawra, Itin. Princ.
S. Coburgi 1: 33. 1883 ≡ Tibouchina moricandiana var. 61. Pleroma francavillanum (Cogn.) P.J.F.Guim. & Miche-
parvifolia (Wawra) Cogn. in Martius, Fl. Bras. 14(3): lang., comb. nov. ≡ Tibouchina francavillana Cogn. in
322. 1885, syn. nov. – Holotype: BRAZIL. Rio de Martius, Fl. Bras. 14(3): 314–315. 1885 – Lectotype
Janeiro: Teresópolis, s.d., Wawra, H. II 352 (W No. (designated here): BRAZIL. Bahia: prope Ilheus, S.
1880-0000967 [online image!]). Domingo, Jan 1822, Riedel, L. 635 (LE!; isolectotype:
= Tibouchina fothergillae var. grandifolia Cogn. in Martius, B [probably destroyed; F negative F0BN016747
Fl. Bras. 14(3): 320. 1885, syn. nov. – Lectotype [no collector number]!).
(designated here): BRAZIL. Minas Gerais: Caldas, Other syntypes: BRAZIL. Bahia: prope S. Domingo, Feb
1 Jun 1874, Mosén, C.W.H. 1973 (BR barcode 1822, Riedel, L. 26 (LE!); idem, Serra da Jacobina, s.d.,
BR0000005530960!). Blanchet, J.S. s.n. [3078 in G] (BR BR0000005224876
Other syntypes: BRASIL. Minas Gerais: in sylvis monto- [fragment]!, G-DC barcode G00318961!).
sis ad Pedra Branca in Serra de Caldas: Regnell, A.F. I
157½ (U barcode U 0004176 [online image!]; BRASIL. 62. Pleroma frigidulum (Schrank & Mart. ex DC.) Triana in
s.l., s.d., Widgren J.F. s.n. (not located). Trans. Linn. Soc. London, Bot. 28(1): 42. 1872
= Tibouchina moricandiana var. vinacea Handro in Loefgre- (“1871”) ≡ Lasiandra frigidula Schrank & Mart. ex
nia 27: 3. 1968, syn. nov. – Holotype: Origin unknown, DC., Prodr. 3: 127. 1828 ≡ Tibouchina frigidula
probably BRAZIL. Cultivated in the Botanical Garden (Schrank & Mart. ex DC.) Cogn. in Martius, Fl. Bras.
of São Paulo, 20 Mar 1963, Handro, O. 1063 (SP barcode 14(3): 328–329, t. 76. 1885 – Holotype: BRAZIL. Minas
SP001231!; isotypes: HB, US barcode 00120075!). Gerais: campis alpestribus Serro Frio, s.d., Martius,

44
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

C.F.P. von s.n. (M barcode M-0005192!; isotype: G-DC Notizbl. Bot. Gart. Berlin-Dahlem 10(91): 48. 1927 –
barcode G00310206 [fragment]!). Holotype: BRAZIL. Bahia, 1914, Lützelburg 531
(M barcode M-0165864 [online image!]).
63. Pleroma gardneri (Naudin) P.J.F.Guim. & Michelang.,
comb. nov. ≡ Lasiandra gardneri Naudin in Ann. Sci. 67. Pleroma goldenbergii (F.S.Mey., P.J.F.Guim. & Kozera)
Nat., Bot., sér. 3, 13(3): 140. 1850 ≡ Tibouchina gardneri P.J.F.Guim. & Michelang., comb. nov. ≡ Tibouchina
(Naudin) Cogn. in Martius, Fl. Bras. 14(3): 334–335. goldenbergii F.S.Mey., P.J.F.Guim. & Kozera in Revista
1885 – Holotype: Tipo BRAZIL Ceará: Serra do Araripe, Brasil. Bot. 33: 266, fig. 1–10. 2010 – Holotype: BRA-
Sep 1838, Gardner, G. 1603 (P barcode P00708988!; iso- ZIL. Paraná: Balsa Nova, Ponte dos Arcos, 25 33′29′′S,
types: BM barcode BM000020628!, BR barcode 49 48′34′′W, 31 Jan 2006, Kozera, C. & Kozera,
BR0000005224562!, CAS barcode 0003133 [online O.P. 2922 (UPCB!; isotype: MBM!).
image!], F barcode V0063571F [fragment]!, G-DC bar-
codes GH00073263! & G00319123!, GH barcode 68. Pleroma granulosum (Desr.) D.Don in Mem. Wern. Nat.
00073263 [online image!], K barcodes K000294020! Hist. Soc. 4: 295. 1823 ≡ Melastoma granulosum Desr. in
& K000329032!, NY barcode 00228672!, P barcodes Lamarck, Encycl. 4: 44. 1797 ≡ Tibouchina granulosa
P00708989! & P00708990!, S No. S-R-6118!, W Nos. (Desr.) Cogn. in Martius, Fl. Bras. 14(3): 340–341.
0053865!, 1889-0008886! & 1889-0333554!). 1885 – Lectotype (designated here): BRAZIL. Rio de
Janeiro: a la moutague des Capucius, Jul 1767, Commer-
64. Pleroma gaudichaudianum (DC.) A.Gray, U.S. Expl. son s.n. (P barcode P00708953!; isolectotypes: C!, P bar-
Exped., Phan.: 603. 1854 ≡ Lasiandra gaudichaudiana codes P00708952 & P00708954!).
DC., Prodr. 3: 128. 1828 ≡ Tibouchina gaudichaudiana
(DC.) Baill. in Hist. Pl. 7: 7, fig. 9. 1879 – Holotype: 69. Pleroma guartelaense F.S.Mey. & R.Goldenb. in Kew
BRAZIL. Rio de Janeiro: 1824, Gaudichaud & Kunth Bull. 69(3): 9527, fig. 2E–H, 3. 2017 – Holotype: BRA-
s.n. (B [probably destroyed; F negative F0BN016748!]). ZIL. Mun. Tibagi, Parque Estadual do Guartelá, trilha
Neotype (designated here): BRAZIL. Rio de Janeiro: para o mirante, 24 33′40.67′′S, 50 15′18.82′′W, 29 Nov
Rio de Janeiro, entre Gavea et Tijuca, 18 Jan 1864, Gla- 2008, Meyer, F. & Jardim 844 (UPCB; isotype: RB bar-
ziou, A.F.M. 592 (P barcode P05200668!; isoneotypes: code 00669844!).
BR!, US barcode 01904044!).
= Lasiandra moricandiana DC., Prodr. 3: 128. 1828 ≡ Ple- 70. Pleroma hatschbachii (Wurdack) P.J.F.Guim. & Michelang.,
roma moricandianum (DC.) Triana in Trans. Linn. Soc. comb. nov. ≡ Tibouchina hatschbachii Wurdack in Pap.
London, Bot. 28(1): 42, t. 2. 1872 (“1871”) ≡ Tibouchina Avulsos Herb. Hatschbach 4: 1–2, fig. 1. 1963 – Type: BRA-
moricandiana (DC.) Baill. in Adansonia 12: 75. 1877 – ZIL. Paraná: Piraí do Sul, Serra das Furnas, 26 Feb 1957,
Holotype: BRAZIL. Without locality, s.d., Herb. Mori- Hatschbach, G. 3624 (US barcode 00120060!; isotypes:
cand s.n. (G-DC barcode G00368034!). MBM barcode MBM049749!, US barcode 00946240!).
= Tibouchina reichardtiana Cogn. in Martius, Fl. Bras. 14(3):
361–362, t. 85. 1885 – Lectotype (designated here): 71. Pleroma heteromallum (D.Don) D.Don in Mem. Wern.
BRAZIL. Rio de Janeiro: S. Cristovão et Coral Falza, Nat. Hist. Soc. 4: 295. 1823 ≡ Melastoma heteromallum
s.d., Pohl, J.B.E. & Schott, H.W. 3957 (W No. 0053861!; D.Don in Bot. Reg. 644, t. 644. 1822 ≡ Lasiandra heter-
isolectotype: BR barcode BR0000005530335 [frag- amalla (D.Don) Naudin in Ann. Sci. Nat., Bot., sér.
ment]!, NY barcode 00245801!). 3, 13(3): 158. 1850 ≡ Tibouchina heteromalla (D.Don)
Cogn. in Martius, Fl. Bras. 14(3): 336–337. 1885 – Lec-
65. Pleroma gertii P.J.F.Guim. & Michelang., nom. nov. ≡ Svi- totype (designated here): [illustration in:] Bot. Reg.:
tramia hatschbachii Wurdack in Bol. Mus. Bot. Munic. 10: t. 644. 1822.
1–2, fig. 1. 1973, non Pleroma hatschbachii (Wurdack) = Lasiandra adenostemon Schrank ex DC., Prodr. 3: 130.
P.J.F.Guim. & Michelang. – Holotype: BRAZIL. Minas 1828 ≡ Pleroma adenostemon (Schrank ex DC.) Triana
Gerais: São Tomé das Letras, rocha do alto do morro, Feb in Trans. Linn. Soc. London, Bot. 28(1): 43. 1872
1973, Hatschbach, G. & Ahumada, Z. 31280 (US barcode (“1871”) ≡ Tibouchina adenostemon (Schrank ex DC.)
00120234!; isotypes: C barcode C10015049 [online image!], Cogn. in Martius, Fl. Bras. 14(3): 332–333. 1885, syn.
CTES barcode CTES0014016 [online image!], F barcode nov. – Holotype: BRAZIL. Sebastianapolitanae et S.
V0063965F [online image!], K barcode K000530436!, LP Pauli, s.d., Martius, C.F.P. von s.n. (M barcode M-
barcode 004384 [online image!], NY barcode 00720504!, 0165825! [F negative F0BN06346!]; isotype: G-DC bar-
RB barcode 00541540!, UC barcode UC1386107 [online code G00310501!).
image!], VEN barcode 196581 [online image!]). = Pleroma multiflorum Gardner in London J. Bot. 2: 351.
1843 ≡ Lasiandra multiflora (Gardner) Naudin in Ann.
66. Pleroma glutinosum (Markgr.) P.J.F.Guim. & Miche- Sci. Nat., Bot., sér. 3, 13(3): 159. 1850 ≡ Tibouchina mul-
lang., comb. nov. ≡ Tibouchinopsis glutinosa Markgr. in tiflora (Gardner) Cogn. in Martius, Fl. Bras. 14(3):

45
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

333–334. 1885, syn. nov. – Holotype: BRAZIL. Rio de (designated here): BRAZIL. Rio de Janeiro, s.d.,
Janeiro: Organ Mountains 3000 ft., Mar 1837, Gardner, G. 398 (BM barcode BM000795368!; isolecto-
Gardner, G. 409 (BM barcode BM000020626!; isotypes: types: E barcodes E00531225 & E00531243 [online
BM barcode BM000020627!, E barcodes E00531226 images!], G-DC!, K barcode K000329111!, NY barcodes
& E00531244 [online images!], GH barcode 00073267 00245673! & 00245674!, P barcodes P00708692!
[online image!], K barcode K000329031!, P barcodes & P00708693!, TCD barcode TCD0005090 [online
P00708710! & P00708711!, herbarium unknown image!], US barcode 00120035!).
[F negative F0BN026109!]). = Tibouchina hospita var. minor Cogn. in Martius, Fl. Bras.
= Tibouchina grandifolia Cogn. in Martius, Fl. Bras. 14(3): 14(3): 418. 1885, syn. nov. – Lectotype (designated
335–336, t. 79. 1885 – Lectotype (designated here): here): BRAZIL. Minas Gerais: Itatiaia, 4 Jun 1871, Gla-
BRAZIL. Rio de Janeiro: s.d., Riedel, L. s.n. (BR barcode ziou, A.F.M. 4805 (P barcode P00708834!; isolectotype:
BR0000005225293!; isolectotypes: BR barcode BR0 P barcode P00708835!).
000005225170!, C barcode C10015069!, M barcode M- Other syntypes: BRAZIL. Rio de Janeiro, 22 Jan 1873,
0165837!). Glaziou, A.F.M. 6531 (BR barcode BR0000005224579!,
= Tibouchina grandifolia var. obtusifolia Cogn. in Martius, K barcode K000329112!, R barcode R 000009657!).
Fl. Bras. 14(3): 336. 1885, syn. nov. – Lectotype (desig- = Tibouchina hospita var. australis Cogn. in Candolle & Can-
nated here): BRAZIL. Rio de Janeiro: 8 Apr 1868, Gla- dolle, Monogr. Phan. 7: 1177. 1891, syn. nov. – Lecto-
ziou, A.F.M. 2569 (BR barcode BR0000005224951!; type (designated here): BRAZIL. Santa Catarina: Serra
isolectotype P barcode P00708950!). Geral, Jan 1890, Ule, E.H.G. 1443 (BR barcode
= Tibouchina magdalenensis Brade in Arq. Inst. Biol. Veg. 4 BR0000005225620!; isolectotype: BR barcode
(1): 74–75, fig. 5. 1938, syn. nov. – Holotype: BRAZIL. BR0000005295562!, CORD barcode CORD 00003625
Rio de Janeiro: Santa Maria Magdalena, Pedra Dubois [online image!], HBG barcodes HBG-523029 & HBG-
1200 m, 28 Feb 1934, Santos Lima, J. & Brade, 523030 [online images!]).
A.C. 13237 (RB barcode 00662326!; isotypes: RB bar-
code 00541566!, SP barcode SP002919!). 74. Pleroma integerrimum (R.Romero & A.B.Martins) P.J.F.
Guim. & Michelang., comb. nov. ≡ Svitramia integer-
72. Pleroma hirsutissimum (Cogn.) P.J.F.Guim. & Miche- rima R.Romero & A.B.Martins in Kew Bull. 58(3):
lang., comb. nov. ≡ Tibouchina hirsutissima Cogn. in 403–406, fig. 1. 2003 – Holotype: Minas Gerais: Delfi-
Martius, Fl. Bras. 14(3): 356. 1885 – Lectotype (desig- nópolis, estrada para a fazenda do Antônio Lopes, Serra
nated here): BRAZIL. Rio de Janeiro, cabo Frio, s.d., Preta, 23 May 1996, Nakajima, J. & Romero, R. 1742
s.c. s.n. (LE!; isolectotype: BR barcode BR0000 (HUFU; isotypes: BHCB barcode BHCB000312!, K bar-
005225149 [fragment]!). code K000530435!, R barcode R 000212033!, UEC, US
= Tibouchina prostrata Ule in Notizbl. Königl. Bot. Gart. Ber- barcode 00810591!).
lin 6(60): 350. 1915, syn nov. – Holotype: BRAZIL. Rio
de Janeiro: in der Strandgegend von Cabo Frio, Oct 1899, 75. Pleroma itatiaiae (Wawra) P.J.F.Guim. & Michelang.,
Ule, E.H.G. 4727 (B [probably destroyed; F negative comb. nov. ≡ Purpurella itatiaiae Wawra in Oesterr. Bot.
F0BN016783!]; isotypes: HBG barcode HBG-523041 Z. 32: 38. 1882 ≡ Tibouchina itatiaiae (Wawra) Cogn. in
[online image!], R barcode R 000185946!). Martius, Fl. Bras. 14(3): 414–415, t. 93, fig. 2. 1885 –
Holotype: Minas Gerais: Hügel um die Facenda, 1879,
73. Pleroma hospitum (Schrank & Mart. ex DC.) Triana in Wawra II, H. 416 (W No. 1880-0000974 [online image!];
Trans. Linn. Soc. London, Bot. 28(1): 42. 1872 isotype: F barcode V0063940F [fragment] [online image!]).
(“1871”) ≡ Lasiandra hospita Schrank & Mart. ex DC.,
Prodr. 3: 129. 1828 ≡ Tibouchina hospita (Schrank 76. Pleroma kleinii (Wurdack) P.J.F.Guim. & Michelang.,
& Mart. ex DC.) Cogn. in Martius, Fl. Bras. 14(3): comb. nov. ≡ Tibouchina kleinii Wurdack in Sellowia
417–318. 1885 ≡ Purpurella hospita (Schrank & Mart. 14: 124, fig. 1G–J. 1962 – Holotype: BRAZIL. Santa
ex DC.) Krasser in Engler & Prantl, Nat. Pflanzenfam. 3 Catarina: Lajes, ruderal 1 km east of Indios, alt.
(7): 148. 1893 – Holotype: BRAZIL. Minas Gerais: in 800–900 m, 11 Feb 1957, Smith, L.B. & Klein,
montibus frigidis Serro Frio praecipue in Serra de Mon- R.M. 11249 (US barcode 00120066!; isotypes: B barcode
tanha et in monte Itambé, 1827, Martius, C.F.P. von s.n. B 10 0244604 [online image!], GH barcode 00073265
(M barcode M-0165839! [F negative F0BN06352!]; iso- [online image!], HBR, K barcode K000329117!, NY bar-
type: G-DC barcode G00310196 [fragment]!). code 00245774!, R barcode R 000136934!, S No. S-R-
= Pleroma albiflorum Gardner in London J. Bot. 2: 347. 1843 6120 [online image!], UC barcode UC1243598 [online
≡ Purpurella albiflora (Gardner) Triana in Trans. Linn. image!]).
Soc. London, Bot. 28(1): 48. 1872 (“1871”) ≡ Tibouchina
hospita var. albiflora (Gardner) Cogn. in Martius, 77. Pleroma kollmannianum F.S.Mey. & R.Goldenb. in Phy-
Fl. Bras. 14(3): 418. 1885, syn. nov. – Lectotype totaxa 282(3): 202–205, fig. 4–5. 2016 – Holotype:

46
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

BRAZIL. Espírito Santo: Santa Teresa, Pedra da Onça, lhotskyana C.Presl, Symb. Bot. 2: 2, t. 52. 1834 ≡ Chae-
cultivated at Santa Teresa, in the Museu de Biologia togastra lhotskyana (C.Presl.) Steud. in Flora 27(2):
Mello Leitão, 13 Apr 2016, (fl., fr.), Meyer, 720. 1844 ≡ Tibouchina lhotskyana (C.Presl) Cogn. in
F.S. & Kollmann, L. 2219 (UPCB; isotypes: MBM, Martius, Fl. Bras. 14(3): 357–358. 1885 – Lectotype
MBML barcode MBML051020 [online image!], NY). (designated here): BRAZIL. Bahia, s.d., Lhotzky, J. s.n.
(G-DC barcode G00319091!; isolectotypes: B [proba-
78. Pleroma kuhlmannii (Brade) P.J.F.Guim. & Michelang., bly destroyed; F negative F0BN016761!], P barcode
comb. nov. ≡ Tibouchina kuhlmannii Brade in Arq. Bot. P00708697!).
Estado São Paulo 3: 210, t. 53, fig. 1–7. 1958 – Holotype:
BRAZIL. São Paulo: Santos, Serra do Mar, região de bora- 83. Pleroma lilacinum (Cogn.) P.J.F.Guim. & Michelang.,
ceia, cabeceira do Rio Guaratuba, 15 Nov 1950, comb. nov. ≡ Tibouchina lilacina Cogn. in Candolle
Kuhlmann, M. & Kühn, E. 2332 (SP barcode SP001230!; & Candolle, Monogr. Phan. 7: 212. 1891 – Lectotype
isotype: SPF barcode SPF00072030 [online image!]). (designated here): BRAZIL. Rio de Janeiro: Mauá,
16 Jan 1887, Schwacke, C.A.W 5433 (RB!; isolectotype:
79. Pleroma laevicaule (Cogn. ex Wurdack) P.J.F.Guim. R barcode R 000114955!).
& Michelang., comb. nov. ≡ Tibouchina laevicaulis
Cogn. ex Wurdack in Phytologia 29(2): 139–140. 1974 84. Pleroma limae (Brade) P.J.F.Guim. & Michelang., comb.
– Holotype: BRAZIL. Distrito Federal: Chapada da nov. ≡ Tibouchina limae Brade in Arq. Inst. Biol. Veg. 4
Contagem, ca. 20 km E of Brasília, elev. 700–1000 m, (1): 72–73, fig. 2. 1938 – Holotype: BRAZIL. Rio de
17 Aug 1964, Irwin, H.S. & Soderstrom, T.R. 5247 Janeiro: Santa Maria Madalena, Águas Paradas, 800 m
(NY barcode 00019389!; isotype: US barcode a.s.l., 5 Mar 1935, Santos Lima, J. & Brade, A.C. 14267
00120068!). (RB barcode 00541562!; isotype: RB barcode
00556707!).
80. Pleroma langsdorffianum (Bonpl.) Triana in Trans. Linn.
Soc. London, Bot. 28(1): 43. 1872 (“1871”) ≡ Rhexia 85. Pleroma lithophilum (Wurdack) P.J.F.Guim. & Miche-
langsdorffiana Bonpl. in Humboldt & Bonpland, lang., comb. nov. ≡ Tibouchina lithophila Wurdack in
Monogr. Melast. 2: 135, pl. 51. 1821 ≡ Osbeckia langs- Brittonia 33(3): 305. 1981 – Holotype: BRAZIL Bahia:
dorffiana (Bonpl.) Spreng., Syst. Veg. 2: 312. 1825 ≡ Milagres Morro de Couro ou Morro de São Cristovão,
Lasiandra langsdorffiana (Bonpl.) DC., Prodr. 3: 130. 12 54′S, 39 52′W, alt. 500–600 m, 6 Mar 1977, Har-
1828 ≡ Tibouchina langsdorffiana (Bonpl.) Baill., Hist. ley, R.M.; Mayo, S.J.; Storr, R.M.; Santos, T.S. & Pin-
Pl. 7: 34. 1879 – Holotype: BRAZIL. Rio de Janeiro, heiro, R.S. 19431 (CEPEC barcode 00019719!;
1820, Langsdorff, G.H. von s.n. (B [probably destroyed; isotypes: AAU ImageID 3478 [online image!], K bar-
F negative F0BN016758!]). Neotype (designated here): code K000329119!, NY barcode 00245781!, U barcode
BRAZIL. Rio de Janeiro, Corcovado a Lagoa de Freitas U 0004179 [online image!], US barcode 00120069!).
19 Dec 1886, Glaziou, A.F.M. 15991 (P barcode
P05315843!; isoneotypes: C!, F!, G!, K barcode 86. Pleroma luetzelburgii (Markgr.) P.J.F.Guim. & Miche-
K000329045!, P barcode P05315844!, R!, RB barcode lang., comb. nov. ≡ Tibouchina luetzelburgii Markgr. in
00231443!). Notizbl. Bot. Gart. Berlin-Dahlem 10(91): 49. 1927 –
= Tibouchina langsdorffiana var. chartacea Cogn. in Martius, Holotype: BRAZIL. Bahia, 1913, Lützelburg, P. von
Fl. Bras. 14(3): 352–353. 1885, syn. nov. – Lectotype 445 (M barcode M-0165844!).
(designated here): BRAZIL. Rio de Janeiro: Laranjeiras,
27 Dec 1868, Glaziou, A.F.M. 2993 (C barcode 87. Pleroma lutzii (Brade) P.J.F.Guim. & Michelang., comb.
C10015074!; isolectotypes: BR barcode BR0000005 nov. ≡ Tibouchina lutzii Brade in Mem. Inst. Oswaldo
223749!, P barcodes P00708698! & P00708699!, R bar- Cruz 53: 353–355, fig. 1–6. 1955 – Holotype: BRAZIL.
code R 000009693!). São Paulo: Serra da Bocaina. 1700 m a.s.l., 21 Apr
1951, Brade, A.C. 20675 (RB barcode 00541565!; iso-
81. Pleroma leopoldinense L.Kollmann & R.Goldenb. in types barcodes RB 00556705! & 00556706!).
Brittonia 68(1): 41–43, fig. 2, 4. 2016 – Holotype: BRA-
ZIL. Espírito Santo: Santa Leopoldina, estrada para o Rio 88. Pleroma macrochiton (Mart. ex DC.) Triana in Trans.
das Farinhas, 14 Apr 2015, Goldenberg, R.; Kollmann, L. Linn. Soc. London, Bot. 28(1): 42. 1872 (“1871”) ≡
& Filgueiras, T. 2128 (MBML barcode MBML048572 Lasiandra macrochiton DC., Prodr. 3: 129. 1828 ≡ Tibou-
[online image!]; isotypes: NY, RB barcode china macrochiton (Mart. ex DC.) Cogn. in Martius,
01043800!, UPCB). Fl. Bras. 14(3): 313–314. 1885 – Holotype: BRAZIL. s.l.,
1827, Martius, C.F.P. s.n. (G-DC barcode G00310195!
82. Pleroma lhotskyanum (C.Presl.) Triana in Trans. Linn. [F negative F0BN07907!]; isotype: E barcode
Soc. London, Bot. 28(1): 43. 1872 (“1871”) ≡ Lasiandra E00285785 [online image!], G barcode G00310194!).

47
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

= Tibouchina macrochiton var. angustifolia Cogn. in Martius, 92. Pleroma martiusianum (DC.) P.J.F.Guim. & Michelang.,
Fl. Bras. 14(3): 314. 1885, syn. nov. – Lectotype (desig- comb. nov. ≡ Lasiandra martiusiana DC., Prodr. 3: 127.
nated here): BRAZIL. s.l., s.d., Wied-Neuwied, M.A.P. zu 1828 ≡ Tibouchina martiusiana (DC.) Cogn. in Martius,
s.n. (BR barcode BR0000006587543!; isolectotypes: BR Fl. Bras. 14(3): 327–328. 1885 – Holotype: BRAZIL.
barcodes BR0000005629374! & BR0000006588076!). Minas Gerais: Itambé, “in campis altis 800–900 [m]
= Tibouchina macrochiton var. uniflora Cogn. in Martius, supra mare”, s.d., Martius, C.F.P. von s.n. (M barcode M-
Fl. Bras. 14(3): 314. 1885, syn. nov. – Holotype: BRA- 0005193! [F negative F0BN06354!]; isotype: G-DC
ZIL. Bahia, 7 Aug 1839, Luschnath, B. 1566 (LE!). barcode G00310204!).

89. Pleroma manicatum (Cogn.) P.J.F.Guim. & Michelang., 93. Pleroma marumbiense (Wurdack) P.J.F.Guim. & Miche-
comb. nov. ≡ Tibouchina manicata Cogn. in Candolle lang., comb. nov. ≡ Tibouchina marumbiensis Wurdack
& Candolle, Monogr. Phan. 7: 220. 1891 – Holotype: in Phytologia 55(3): 137–138. 1984 – Holotype:
BRAZIL. Minas Gerais: Serra do Caparaó, Jun 1888, BRAZIL. Paraná: Morretes, in cloud forest on Serra Mar-
Moura, J. 259 (BR barcode BR0000005282074!; iso- umbi, elev. 1300–1400 m, 27 Feb 1970, Hatschbach, G.
type: B [probably destroyed; F negative F0BN016766!]). 23934 (MBM barcode MBM013221!; isotypes: HBG
barcode HBG-522831 [online image!], US barcode
90. Pleroma marinanum P.J.F.Guim. & Fraga in Phytotaxa 166 00120073!).
(1): 78–81, fig. 1, 2B–E. 2014 – Holotype: BRAZIL. Espírito
Santo: mun. Pancas, Serra do Mutum Preto, elev. 750 m, 7 Mar 94. Pleroma maximilianum (DC.) Triana in Trans. Linn. Soc.
2003, Fraga, C.N.; Boudet Fernandes, H.Q. & Vervloet, London, Bot. 28(1): 43. 1872 (“1871”) ≡ Lasiandra max-
R.R. 962 (MBML barcode MBML021011!; isotypes: imiliana DC., Prodr. 3: 128. 1828 ≡ Tibouchina maximili-
CEPEC, K, NY barcode 00879073!, RB, UPCB). ana (DC.) Baill., Hist. Pl. 7: 34. 1877 – Holotype:
BRAZIL. 1815, Wied-Neuwied, M.A.P. zu s.n. (M bar-
91. Pleroma martiale (Cham.) Triana in Trans. Linn. Soc. code M-0165848!; isotypes: BR barcodes BR000000
London, Bot. 28(1): 44. 1872 (“1871”) ≡ Lasiandra mar- 6587871 & BR0000006588533!, GOET barcode GOET
tialis Cham. in Linnaea 9: 433. 1835 ≡ Tibouchina mar- 000776 [online image!]).
tialis (Cham.) Cogn. in Martius, Fl. Bras. 14(3):
346–347, t. 80, fig. 2. 1885 – Lectotype (designated 95. Pleroma melanocalyx (R.Romero, P.J.F.Guim. & Leoni)
here): BRAZIL. s.d., Sellow, F. 5124 (K barcode P.J.F.Guim. & Michelang., comb. nov. ≡ Tibouchina
K000329039!; isolectotypes: E barcode E00531227 melanocalyx R.Romero, P.J.F.Guim. & Leoni in Novon
[no collector number] [online image!], HBG barcode 19(2): 219–221, fig. 1. 2009 – Holotype: BRAZIL.
HBG-508544 [no collector number] [online image!], M Minas Gerais: Parque Estadual da Serra do Brigadeiro,
barcode M-0165846 [no collector number]!). Fervedouro, 7 Jul 1995, Leoni, L.S. 2995 (RB barcode
= Pleroma corymbosum Benth., Pl. Hartw.: 181. 1845 ≡ 01128401!; isotypes: HUFU barcode HUFU00067943!,
Lasiandra corymbosa (Benth.) Naudin in Ann. Sci. MO, RB barcode 01128698!).
Nat., Bot., sér. 3, 13(3): 159. 1850 ≡ Tibouchina lindeni-
ana Cogn. in Martius, Fl. Bras. 14(3): 347. 1885, non 96. Pleroma mellobarretoi (Brade) P.J.F.Guim. & Miche-
Tibouchina corymbosa (Raddi) Cogn. 1885, syn. nov. – lang., comb. nov. ≡ Tibouchina mellobarretoi Brade in
Lectotype (designated here): COLOMBIA. Socorro, Arq. Inst. Biol. Veg. 4(1): 76, fig. 7. 1938 (“mello-barre-
Nov 1842, Linden, J.J. 753 (BR barcode BR000 toi”) – Holotype: BRAZIL. Minas Gerais: Serra do Cipó,
0005223411!; isolectotypes: BM barcodes BM0000 15 Apr 1935, Mello Barreto & Brade, A.C. 1168
20553! & BM000020555!, F!, G [on 2 sheets]!, K bar- (RB barcode 00662327!; isotypes: RB barcodes
code K000329131!, MPU barcode MPU013405 [online 00541568! & 00662433!).
image!], P barcode P00708696!).
= Tibouchina pauciflora Cogn. in Martius, Fl. Bras. 14(3): 97. Pleroma michelangii P.J.F.Guim. & J.G.Freitas in Syst.
348. 1885, syn. nov. – Lectotype (designated here): Bot. 40(2): 553–554, fig. 1, 2A,E,I,M. 2015 – Holotype:
BRAZIL. Minas Gerais: Caldas, 27 Feb 1862, Regnell, BRAZIL. Bahia: Barra da Estiva, Serra do Sincorá,
A.F. III 17 (BR barcode BR0000005223787!; isolecto- 13 31′19′′S, 4112′11′′W, 980 m, 7 Mar 1996, Sali-
types: C barcodes C10015077! [F negative F0BN mena-Pires, F.R.; Lima, R.; Roque, N.; Conceição, A.
021208!] & C10015079!, F barcode V0063994F [online & Nobre, P.H. 2140 (ALCB!).
image!], P barcode P00708682!, S Nos. S08-20260!
& S-R-612!, W No. 1927-0001185!). 98. Pleroma microphyllum (Cogn.) P.J.F.Guim. & Miche-
= Tibouchina martialis var. ovata Cogn. in Candolle & Can- lang., comb. nov. ≡ Tibouchina microphylla Cogn. in
dolle, Monogr. Phan. 7: 225. 1891, syn. nov. – Holotype: Candolle & Candolle, Monogr. Phan. 7: 277. 1891 – Lec-
BRAZIL. Minas Gerais: Caraça, Feb. 1885, totype (designated here): BRAZIL. Minas Gerais, Serra
Mendonça, R. 22 (S!). do Caparaó, 8 Feb 1890, Schwacke, C.A.W. 6722

48
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

(P barcode P00708822!; isolectotype: P barcode P00708823 103. Pleroma mosenii (Cogn.) P.J.F.Guim. & Michelang.,
[fragment]!). comb. nov. ≡ Tibouchina mosenii Cogn. in Martius,
Other syntypes: BRAZIL. Minas Gerais: Serra do Caparaó, Fl. Bras. 14(3): 393–394, t. 91. 1885 – Lectotype (desig-
s.d., Moura, J.T. de 186 (B [probably destroyed; F negative nated here): BRAZIL. Minas Gerais: Caldas, in fissuris
F0BN016767!], S No. S09-13201 [fragment] [online rupium apricarum infra cacumen montis Pedra Branca
image!]); BRAZIL. Minas Gerais: Serra do Caparaó, ad Serra de Caldas, 1 Jun 1874, Mosén, C.W.H. 1972
13 Mar 1889, Kromer, E. 6472 (P barcode P00708824!); (S No. S-R-6122!; isolectotypes: BR barcode BR
BRAZIL. Minas Gerais: Serra do Caparaó, in campis eleva- 0000005223466!, P barcode P00708825!, S No.
tis, 2 Jul 1888, Schwacke, C.A.W. 6163 (P barcode S09-13194!).
P05228373!). Other syntypes: BRAZIL. Minas Gerais: Caldas, 18 Apr
1847, Regnell, A.F. III 1527 (B [probably destroyed;
99. Pleroma minus (R.Romero & A.B.Martins) P.J.F.Guim. F negative F0BN016771!], BR barcodes BR000
& Michelang., comb. nov. ≡ Svitramia minor R.Romero 0005281985!, BR0000005282401! & BR000000528
& A.B.Martins in Kew Bull. 58(3): 406–408, fig. 2. 2003 – 2739!, C barcode C10015080!, F barcodes V0093865F!
Holotype: BRAZIL. Minas Gerais: mun. São Roque de Minas, & V0093866F!, NY barcode 00245788!, P barcodes
Parque Nacional da Serra da Canastra, Morro após o córrego P00708825! & P00708826!, R barcode R 000142032!,
dos Passageiros, 21 Nov 1996, Nakajima, J. & Romero, R. U barcodes U 0004181 & U 0004182).
2130 (HUFU; isotypes: K barcode K000530434!, NY barcode = Tibouchina mosenii var. jordanensis Brade in Arq. Inst.
00688091!, R barcode R 000212035!, SP barcode Biol. Veg. 4(1): 77. 1938, syn. nov. – Holotype: BRA-
SP001223!, UEC, US barcode 00810592!). ZIL. São Paulo: Campo do Jordão, Apr 1937,
Lanstyák, L. s.n. (RB barcode 00541570!; isotypes: NY
100. Pleroma minutiflorum (Cogn.) P.J.F.Guim. & Miche- barcode 00245789!, RB barcode 00556737!).
lang., comb. nov. ≡ Tibouchina minutiflora Cogn. in Mar- = Tibouchina mosenii var. ciliato-alata Brade in Arq. Inst.
tius, Fl. Bras. 14(3): 415–416, t. 95, fig. 1. 1885 ≡ Biol. Veg. 4(1): 77. 1938, syn. nov. – Holotype: BRAZIL.
Purpurella minutiflora (Cogn.) Krasser in Engler & Prantl, Rio de Janeiro: Itatiaia, Maromba 1000 a.s.l., 22 May
Nat. Pflanzenfam. 3(7): 148. 1893 – Lectotype (designated 1935, Brade, A.C. 14615 (RB barcode 00541599!; iso-
here): BRAZIL. São Paulo [Rio de Janeiro in BR]: Campos types: NY barcode 02350106!, RB barcode 00541606!).
da Bocaina, Sobrado, 8 Jan 1876, Glaziou, A.F.M. 8366
(BR barcode BR0000005224081!; isolectotypes: B [proba- 104. Pleroma mourae (Cogn.) P.J.F.Guim. & Michelang.,
bly destroyed; F negative F0BN016768!], F barcode comb. nov. ≡ Tibouchina mouraei Cogn. in Candolle
V0063991F [fragment]!, K barcode K000329109!, P & Candolle, Monogr. Phan. 7: 236. 1891 – Lectotype
barcodes P00708836! & P00708837!, R barcode (designated here): BRAZIL. Minas Gerais: Serra do
R000009665!, S No. S09-13200 [online image!]). Caparaó, Jun 1888, Moura, J.T.de 185 (BR barcode
BR0000005520787!; isolectotypes: B [probably de-
101. Pleroma mirabile (Brade & Markgr.) P.J.F.Guim. stroyed; F negative F0BN016770!]).
& Michelang., comb. nov. ≡ Tibouchinopsis mirabilis
Brade & Markgr. in Arch. Jard. Bot. Rio de Janeiro 17: 105. Pleroma mutabile (Vell.) Triana in Trans. Linn. Soc.
45–46, fig. 2: 11–19. 1959–61 (= in Willdenowia 2(5): London, Bot. 28(1): 40. 1872 (“1871”) ≡ Melastoma muta-
773–774. 1961) – Holotype: BRAZIL. Bahia: Morro do bile Vell., Fl. Flumin.: 181. 1829 (“1825”) ≡ Lasiandra mu-
Chapéu, 1000 m, 11 Sep 1956, Pereira, E. 2143 (HB; iso- tabilis (Vell.) Riedel ex Naudin in Ann. Sci. Nat., Bot., sér.
types: RB barcodes 00541588! & 00556744!, US barcode 3, 13(3): 145. 1850 ≡ Tibouchina mutabilis (Vell.) Cogn.
00120107!). in Martius, Fl. Bras. 14(3): 300–301. 1885 – Lectotype
Brade (1959–1961) published new Brazilian species of (designated here): [illustration] “MELASTOMA MUTA-
Melastomataceae, manuscript submitted in December 1960, BILIS” in Vellozo, Fl. Flumin. Icones 4: t. 130. 1831.
but Brade & Markgraf (1961) also published many of these = Tibouchina mutabilis var. alba Mattos in Loefgrenia 85:
species plus others in another journal. In both articles, they [2]. 1984, syn. nov. – Holotype: BRAZIL. São Paulo:
received the status of species nova. For this reason, both works Cubatão, junto à rodovia dos Imigrantes, Apr 1974,
are cited. Mattos, J. 26282 (HAS No. 87549).

102. Pleroma molle (Cham.) Triana in Trans. Linn. Soc. 106. Pleroma noblickii (Wurdack) P.J.F.Guim. & Michelang.,
London, Bot. 28(1): 43. 1872 (“1871”) ≡ Lasiandra mol- comb. nov. ≡ Tibouchina noblickii Wurdack in Kew
lis Cham. in Linnaea 9: 444. 1835 ≡ Tibouchina chamis- Bull. 50(4): 823–824. 1995 – Holotype: BRAZIL. Bahia:
soana Cogn. in Martius, Fl. Bras. 14(3): 349–350. 1885 Serra de S. José, Feira de Santana, 12 15′S, 38 58′W,
– Lectotype (designated here): BRAZIL. s.d., 10 May 1984, Noblick, L.R. 3173 (HUEFS barcode
Sellow, F. 18 (P barcode P00121040!; isolectotype: B HUEFS003329!; isotypes: CEPEC barcode 00035279!,
[probably destroyed; F negative F0BN016732!]). RB barcode 00541571!, US barcode 00478964!).

49
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

107. Pleroma nodosum (Wurdack) P.J.F.Guim. & Michelang., (BR barcode BR0000005223473!, C barcode C100
comb. nov. ≡ Tibouchina nodosa Wurdack in Los Angeles 15082!, F barcode V0063992F [fragment]!, K barcode
County Mus. Contr. Sci. 28: 9, fig. 3. 1959 – Holotype: K000329017!, P barcodes P00708673! & P00708674!).
BRAZIL. Goiás: Region of the Chapada dos Veadeiros,
sandstone outcrop 7 km south of Veadeiros 47 30′S, 112. Pleroma pauloalvinii (Vinha) P.J.F.Guim. & Miche-
14 30′W, 24 Apr 1956, Dawson, E.Y. 14596 (H; isotype: lang., comb. nov. ≡ Tibouchina pauloalvinii Vinha in
NY barcode 00245791!, R barcode R 000171760!, RSA Bradea 1(30): 320, fig. 1. 1973 (“paulo-alvinii”) – Holo-
barcode RSA0004263 [fragment] [online image!]). type: BRAZIL. Bahia: Itacaré/Ubaitaba, margem da
estrada, 17 Apr 1970, Santos, T.S. 733 (CEPEC barcode
108. Pleroma ochypetalum (Ruiz & Pav.) D.Don in Mem. 00005622!; isotype: US barcode 00120084!).
Wern. Nat. Hist. Soc. 4: 295. 1823 ≡ Rhexia ochypetala
Ruiz & Pav., Fl. Peruv. 3: 86, t. 321, fig. a. 1802 ≡ Lasian- 113. Pleroma penduliflorum Fraga & P.J.F.Guim. in Phyto-
dra ochypetala (Ruiz & Pav.) DC., Prodr. 3: 128. 1828 ≡ taxa 166(1): 81–83, fig. 2F–I, 4. 2014 – Holotype:
Tibouchina ochypetala (Ruiz & Pav.) Baillon in Adanso- BRAZIL. Espírito Santo: mun. Pancas, Serra do Mutum
nia 7: 75. 1877 – Lectotype (designated here): PERU. Preto, 750 m, 7 Mar 1999, Fraga, C.N.; Fernandes,
s.d., Pavón, J. s.n. (BC barcode BC-872998 [online H.Q.B. & Vervloet, R.R. 965 (MBML barcode
image!]; isolectotypes: MA barcodes MA 813969 MBML021014!; isotypes: NY barcode 00879071!,
& MA 813970 [online images!]). RB, UPCB).
= Tibouchina tessmannii Markgr. in Notizbl. Bot. Gart. Berlin-
Dahlem 9: 1139. 1927 – Lectotype (designated here): 114. Pleroma pereirae (Brade & Markgr.) P.J.F.Guim.
PERU. Marañón, 15 Aug 1924, Tessmann, G. 3788 & Michelang., comb. nov. ≡ Tibouchina pereirae Brade
(NY barcode 00169344!; isolectotypes: B [probably & Markgr. in Arch. Jard. Bot. Rio de Janeiro 17: 46–46,
destroyed; F negative F0BN016800!], F barcode fig. 2: 1–10. 1959–61 (= in Willdenowia 2(5): 772–773.
V0064000F [fragment] [online image!], G!, S No. S-R- 1961) – Holotype: BRAZIL. Bahia: entre Lençóis e Ita-
6127 [online image!]). beraba, 15 Sep 1956, Pereira, E. 2062 (HB; isotype: RB
barcode 00541576!).
109. Pleroma oleifolium (DC.) R.Romero & Versiane in Brade (1959–1961) published new Brazilian species of
Novon 23(2): 218–223, fig. 1. 2014 ≡ Osbeckia oleifolia Melastomataceae, manuscript submitted in December 1960,
DC., Prodr. 3: 139. 1828 (“oleaefolia”) ≡ Microlepis olei- but Brade & Markgraf (1961) also published many of these
folia (DC.) Triana in Trans. Linn. Soc. London, Bot. species plus others in another journal. In both articles, they
28(1): 48. 1872 (“1871”) (“oleaefolia”) – Holotype: received the status of species nova. For this reason, both works
BRAZIL. São Paulo: in campis siccis elatioribus in valle are cited.
flum. Paraiba et prope Lorena, s.d., Martius, C.F.P. von
s.n. (M barcode M-0165680!). 115. Pleroma petiolatum (R.Romero & A.B.Martins) P.J.F.
Guim. & Michelang., comb. nov. ≡ Svitramia petiolata
110. Pleroma oreophilum (Wurdack) P.J.F.Guim. & Miche- R.Romero & A.B.Martins in Kew Bull. 58(3): 408–410,
lang., comb. nov. ≡ Tibouchina oreophila Wurdack in fig. 3. 2003 – Holotype: BRAZIL. Minas Gerais: Estrada
Brittonia 33(3): 305–308, fig. 1–4. 1981 – Holotype: Furnas-Capitão, c. de 15 km após a ponte da represa de Fur-
BRAZIL. Bahia: among rocks at summit of Pico das nas, beira da estrada, afloramentos rochosos, lado direito da
Almas ca. 25 km WNW of the vila do Rio da Contas, estrada, 13 Feb 1998, Romero, R; Semir, J.; Goldenberg, R.
alt. 1600–1850 m, 19 Mar 1977, Harley, R.M.; Mayo, & Dutilh, J.H.A. 5131 (UEC; isotype: HUFU).
S.J.; Ston, R.M.; Santos, T.S. & Pinheiro, R.S. 19695
(CEPEC barcode 00019728!; isotypes: K barcode 116. Pleroma pilosum (Cogn.) P.J.F.Guim. & Michelang.,
K000329124!, US barcode 00120080!). comb. nov. ≡ Tibouchina pilosa Cogn. in Martius in
Fl. Bras. 14(4): 600. 1888 – Holotype: BRAZIL. São
111. Pleroma pallidum (Cogn.) P.J.F.Guim. & Michelang., Paulo: Ipanema dans le bois, 22 Mar 1884, Glaziou,
comb. nov. ≡ Tibouchina pallida Cogn. in Martius, A.F.M. 15977 (P barcode P00121413!; isotypes: B [prob-
Fl. Bras. 14(3): 316–317, t. 70, fig. 2. 1885 – Lectotype ably destroyed; F negative F0BN016782!], BR barcode
(designated here): BRAZIL. Rio de Janeiro: Rio de BR0000005530656 [fragment]!, C barcode C10015085!).
Janeiro, Pico do Andarahy Grande ou Tijuca, 25 Jul = Tibouchina pilosa var. glandulosa Wurdack in Sellowia 14:
1866., Glaziou, A.F.M. 1090 (BR barcode BR0000 117. 1962, syn. nov. – Holotype: BRAZIL. Santa Cata-
005224128!; isolectotypes: C barcode C10015083!, P rina: an Uferabhängen bei Blumenau, Jan 1888, Ule,
barcodes P00708675! & P04888221!, R barcodes E.H.G. 669 (US barcode 00120086!).
R 000009283! & R 000009283a!).
Other syntypes: BRAZIL. Rio de Janeiro: Rio de Janeiro, 117. Pleroma quartzophilum (Brade) P.J.F.Guim. & Miche-
Pico de Tijuca, 3 Apr 1875, Glaziou, A.F.M. 8356 lang., comb. nov. ≡ Tibouchina quartzophila Brade in

50
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

Arch.. Jard. Bot. Rio de Janeiro 14: 219–220, t. 5. 1956 – K001089085!, NY barcodes 00245780! & 1477595!,
Holotype: BRAZIL. Espírito Santo: Cachoeiro do Itape- RB barcodes 00541564! & 00662437!, SP barcode
mirim, Vargem Alta, morro do Sal, alt. 600 m, 6 May SP002915!).
1949, Brade, A.C. 19767 (RB barcode 662347!; isotypes:
K barcode K001089089!, NY barcodes 00245800! 120. Pleroma ramboi (Brade) P.J.F.Guim. & Michelang.,
& 01477590!, RB barcodes 00541578! & 00662439!, comb. nov. ≡ Tibouchina ramboi Brade in Sellowia 8:
SP barcode SP002920!). 367–368, fig. 1. 1957 – Holotype: BRAZIL. Rio Grande
do Sul: Serra da rocinha pr. Bon Jesus in silvula nebulari,
118. Pleroma raddianum (DC.) Gardner in London J. Bot. 2: 3 Feb 1953, Rambo, B. s.n. (PACA No. 53871!; isotypes:
351. 1843 ≡ Rhexia uniflora Raddi in Mem. Mat. Fis. HB [fragment], RB barcode 00541579!).
Soc. Ital. Sci. Modena, Pt. Mem. Fis. 18(2): 388. 1820,
nom. illeg., non Vahl 1791 ≡ Lasiandra raddiana DC., 121. Pleroma redivivum O.Berg ex Triana in Trans. Linn.
Prodr. 3: 129. 1828 ≡ Tibouchina raddiana (DC.) Cogn. Soc. London, Bot. 28(1): 43. 1872 (“1871”) ≡ Tibou-
in Martius, Fl. Bras. 14(3): 303–304. 1885 – Holotype: china rediviva (O.Berg ex Triana) Cogn. in Martius,
BRAZIL. Rio de Janeiro, s.d., Raddi, G. s.n. (PI; isotype: Fl. Bras. 14(3): 399. 1885 – Holotype: BRAZIL. Minas
G barcode G00310190!). Gerais: Serra de Caraza [Caraça], s.d., Sellow, F. 1329
= Lasiandra pulchra Cham. in Linnaea 9: 439. 1835 ≡ Tibou- (B [probably destroyed; F negative F0BN016784!]).
china pulchra (Cham.) Cogn. in Martius, Fl. Bras. 14(3):
301–302, t. 68. 1885, syn. nov. – Lectotype (designated 122. Pleroma regelianum (Cogn.) P.J.F.Guim. & Michelang.,
here): BRAZIL. s.d., Sellow, F. s.n. (M barcode M- comb. nov. ≡ Tibouchina regeliana Cogn. in Martius,
0165851!; isolectotypes: K barcodes K000329000! Fl. Bras. 14(3): 356–357. 1885 – Lectotype (designated
& K000684628!, US barcode 00730886!). here): Cultivatum, s.l. s.d, s.c. s.n. (BR barcode
= Tibouchina saldanhaei Cogn. in Martius, Fl. Bras. 14(3): BR0000005224906!; isolectotype LE!).
302–303, t. 69, fig. 1. 1885, syn. nov. – Lectotype
(designated here): BRAZIL. Rio de Janeiro, Serra 123. Pleroma reitzii (Brade) P.J.F.Guim. & Michelang.,
d’Estrella, 27 Jan 1878, Glaziou, A.F.M. 7610 (BR comb. nov. ≡ Tibouchina reitzii Brade in Sellowia 12:
barcode BR0000005530373!; isolectotypes: BR bar- 137–138, fig. 2: 1–7. 1960 – Holotype: BRAZIL. Santa
code BR0000005530700!, C barcodes C10015086! Catarina: Joinville, estrada Dona Francisca, 600 m a.s.l.,
& C10015087!, K barcode K000329003!, LE!, P bar- 18 Dec 1957, Reitz, P.R. & Klein, R.M. 5705 (HB No.
codes P00708813!, P00708814! & P00708815!). 10704!; isotypes: B barcode B 10 0295489!, HBR bar-
Other syntypes: BRAZIL. Rio de Janeiro: Organ Moun- code HBR0027600!, MBM barcode MBM002133!, NY
tains, Apr 1837, Gardner, G. 408 (BM!, G!, K!); BRA- barcode 00558377!, US barcode 00120087!).
ZIL. Rio de Janeiro: Serra dos Orgãos entre Garapão e = Tibouchina reitzii var. alpestris Brade in Sellowia 12:
Ventania, 13 Jan 1883, Saldanha, J. 6805 (P barcode 138–139, fig. 2: 8–13. 1960, syn. nov. – Holotype: BRA-
P00708816!, R barcode R 000182445!). ZIL. Santa Catarina: Campo Alegre, Morro Iquererim,
= Tibouchina petroniana Saldanha & Cogn., Bouquet Mélast. 1300 m a.s.l., 5 Feb 1958, Reitz, P.R. & Klein, R.M.
Brés.: t. 2, fig. 1. 1887, syn. nov. – Lectotype (desig- 6484 (HB; isotypes: B barcode B 10 0295489!, HBR bar-
nated here): BRAZIL. Rio de Janeiro: alto da Serra de code HBR0027537!, NY barcode 00169343!, US bar-
Nova Friburgo, 18 Mar 1887, Glaziou, A.F.M. 15997 code 00120088!).
(BR barcode BR0000005223480!; isolectotypes: BR bar-
codes BR0000005223503! & BR0000005223831!, C 124. Pleroma riedelianum (Cogn.) P.J.F.Guim. & Miche-
barcode C10015084!, F barcode V0063995F [fragment]!, lang., comb. nov. ≡ Tibouchina riedeliana Cogn. in Mar-
G [on 2 sheets]!, K barcode K000329001!, LE!, P bar- tius, Fl. Bras. 14(3): 368–369, t. 87, fig. 2. 1885 –
codes P00708667!, P00708668! & P00708669!, R bar- Lectotype (designated here): BRAZIL. São Paulo:
code R 000009302!, RB barcode 00541577!). prope Mogi, Nov 1833, Riedel, L. 1426 (BR barcode
BR0000005225309!; isolectotypes: B [probably
119. Pleroma radula (Markgr.) P.J.F.Guim. & Michelang., destroyed; F negative F0BN016786!], K barcode
comb. nov. ≡ Tibouchina radula Markgr. in Notizbl. K000329060!, LE!, M barcode M-0165853!, NY barcode
Bot. Gart. Berlin-Dahlem 10(91): 49. 1927 – Holotype: 00245802!, P barcodes P00708840!, P00708841!
BRAZIL. Espírito Santo: Leopoldina, May 1917, & P00708842!, W!).
Luetzelburg, P. von 13348 (M barcode M-0165852!).
= Tibouchina limoeirensis Brade in Arch. Jard. Bot. Rio de 125. Pleroma rigidulum (Naudin) P.J.F.Guim. & Michelang.,
Janeiro 14: 214–215, t. 2. 1956, syn. nov. – Holotype: comb. nov. ≡ Lasiandra rigidula Naudin in Ann. Sci.
BRAZIL. Espírito Santo: Itaguassú, Alto Limoeiro, Nat., Bot., sér. 3, 13(3): 150–151. 1850 ≡ Tibouchina rigi-
10 May 1946, Brade, A.C.; Barbosa, A. & Duarte, dula (Naudin) Wurdack in Phytologia 21(2): 116. 1971 –
A. 18057 (RB barcode 00662342!; isotypes: K barcode Holotype: BRAZIL. Minas Gerais: in monticulo dicto

51
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

Morro-Pilado prope urbem Villa do Principe, 1816– na Chapada, perto rio Manso, Apr 1911, Hoehne,
1821, De Saint-Hilaire, A.F.C.P. B1.996 (P barcode F.C. [Comissão Rondon] 2391 (R barcode R 000168429!).
P00046774!; isotype: P barcode P00708839!).
131. Pleroma salviifolium (Cham.) Triana in Trans. Linn.
126. Pleroma riparium (Markgr.) P.J.F.Guim. & Michelang., Soc. London, Bot. 28(1): 44. 1872 (“1871”) ≡ Lasiandra
comb. nov. ≡ Tibouchina riparia Markgr. in Notizbl. Bot. salviaefolia Cham. in Linnaea 9: 441. 1835 ≡ Tibouchina
Gart. Berlin-Dahlem 10(91): 50. 1927 – Lectotype (des- salviaefolia (Cham.) Cogn. in Martius, Fl. Bras. 14(3):
ignated here): BRAZIL. Bahia: Rio de Contas, Flußufer, 364–365. 1885 – Lectotype (designated here): BRA-
Jul 1914, Luetzelburg, P. 505 (M barcode M-0165855!; ZIL. Inter Vitoria et Bahia, s.d., Sellow, F. 128
isolectotype: M barcode M-0165854!). (BR barcode BR0000005225972!; isolectotypes: B
[probably destroyed; F negative F0BN016791!], K bar-
127. Pleroma robustum (Cogn.) P.J.F.Guim. & Michelang., code K000329056!, P barcode P00708818!).
comb. nov. ≡ Tibouchina robusta Cogn. in Martius, = Lasiandra sulcata Naudin in Ann. Sci. Nat., Bot., sér.
Fl. Bras. 14(3): 394–395. 1885 – Lectotype (desig- 3, 13(3): 137. 1850 ≡ Pleroma sulcatum (Naudin) Triana
nated here): BRAZIL. Goiás: in sylvis montosis as in Trans. Linn. Soc. London, Bot. 28(1): 44. 1872
Morro do Manoel Gomez prope Villa Boa, s.d., Pohl, (“1871”) ≡ Tibouchina sulcata (Naudin) Cogn. in Mar-
J.B.E. 1628 (M barcode M-0165856!; isolectotypes: tius, Fl. Bras. 14(3): 337–338, t. 78, fig. 2. 1885, syn.
BR barcode BR0000005225637 [fragment]!, F barcode nov. – Holotype: BRAZIL. Bahia, s.d., Blanchet,
V0063996F [fragment]!, NY barcode 00245803!, W!, J.S. 1697 (P barcode P00708775; isotypes: BM barcode
herbarium unknown [F negative F0BN032313 or BM000020473!, BR barcode BR0000005225682 [frag-
32343, illegible!]). ment]!, F!, G barcode G00368037!, G-DC barcode
G00319122!, LE!, P barcode P00708774!).
128. Pleroma rubrobracteatum (R.Romero & P.J.F.Guim.)
P.J.F.Guim. & Michelang., comb. nov. ≡ Tibouchina 132. Pleroma setosociliatum (Cogn.) F.S.Mey. & F.B.Matos
rubrobracteata R.Romero & P.J.F.Guim. in Novon in Phytotaxa 305(3): 202, fig. 2. 2017 ≡ Tibouchina seto-
15(2): 210–212, fig. 1. 2005 – Holotype: BRAZIL. sociliata Cogn. in Ark. Bot. 9: 9. 1910 (“setoso-ciliata”) –
Minas Gerais: São Roque de Minas. Par. Nac. Serra da Holotype: BRAZIL. Paraná: Marumbi, Serra do Mar, in
Canastra, estrada para o sítio João Domingos, Serra da einer Höhe von etwa 1000 m, 13 Feb 1904, Dusén,
Canastra, afloramentos rochosos, 20 00′–20 39′S, P.K.H. 3614A (BR barcode BR0000005225651!;
46 15′–47 00′W, 26 May 1996, Nakajima, J.N. & isotypes: B [probably destroyed; F negative
Romero, R. 1807 (HUFU!; isotypes: K barcode F0BN016795!], S No. S-R-9197!).
K000343212!, MO barcode MO-1515228!, RB barcodes
00557100! & 00557463!, UEC, US barcode 00956377!). 133. Pleroma scaberrimum Triana in Trans. Linn. Soc.
London, Bot. 28(1): 47. 1872 (“1871”) ≡ Tibouchina sca-
129. Pleroma rubrum J.G.Freitas in Phytotaxa 288(3): berrima (Triana) Cogn. in Martius, Fl. Bras. 14(3):
250–255, fig. 1, 2. 2016 – Holotype: BRAZIL. Bahia: 400–401. 1885 – Lectotype (designated here): BRA-
Rio de Contas, vertente leste do Campo do Queiroz, ZIL. Prov. St. Paul and Rio, South BRAZIL, 1861–2,
13 32′S, 41 59′W, 1500 m, 24 Dec 1988, fl., fr., Harley, Weir, J. 246 (K barcode K000329097!).
R.M. 27358 (HUEFS; isotypes: CEPEC barcode = Tibouchina nobilis Rech. in Denkschr. Kaiserl. Akad.
00047370 [online image!], K, MBM barcode Wiss., Wien. Math.-Naturwiss. Kl. 79: 254. 1908 – Holo-
MBM302116 [online image!], NY barcodes 00558327! type: BRAZIL. São Paulo: in silvis Alto da Serra,
& 00687343!, P barcode P05287214!, SPF barcode 900 m a.s.l., kleiner Baum, Blüten im Leben schön vio-
SPF00178641 [online image!], US barcode 01906444 lett, May 1901, Wettstein, R. & Schiffner, V.F. 196
[online image!]). (W No. 1907-0005462 [online image!]; isolectotypes: B
[probably destroyed; F negative F0BN016774!], WU
130. Pleroma rupicola (Hoehne) P.J.F.Guim. & Michelang., Nos. 0034180, 0034181, 0034182 & 0034183 [online
comb. nov. ≡ Tibouchina rupicola Hoehne in Anexos images!]).
Mem. Inst. Butantan, Secc. Bot. 1(5): 85–86, t. 12, fig. 1.
1922 – Lectotype (designated here): BRAZIL. Mato 134. Pleroma schenckii (Cogn.) P.J.F.Guim. & Michelang.,
Grosso: Casa da Pedra, na Chapada, Apr 1911, Hoehne, comb. nov. ≡ Tibouchina schenckii Cogn. in Candolle
F.C. [Comissão Rondon] 2390 (SP barcode SP001234!; & Candolle, Monogr. Phan. 7: 207. 1891 – Lectotype
isolectotype: S No. S09-13187!). (designated here): BRAZIL. Serra do Picú, entre as pro-
Other syntypes: BRAZIL. Mato Grosso: Casa da Pedra, víncias de Rio de Janeiro, São Paulo e Minas Gerais,
na Chapada, perto Rio Manso, Apr 1911, Hoehne, Nov–Dec 1886, Schenck, H. 1487 (BR barcode
F.C. [Comissão Rondon] 2389 (R barcode BR0000005224616!; isolectotype: BR barcode
R 000007194!); BRAZIL. Mato Grosso: Casa da Pedra, BR0000005225262!, RB barcode 00541582!).

52
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

135. Pleroma schwackei (Cogn.) P.J.F.Guim. & Michelang., 137. Pleroma semidecandrum (Schrank & Mart. ex DC.) Tri-
comb. nov. ≡ Tibouchina schwackei Cogn. in Candolle ana in Trans. Linn. Soc. London, Bot. 28(1): 41. 1872
& Candolle, Monogr. Phan. 7: 206. 1891 – Lectotype (“1871”) ≡ Lasiandra semidecandra Schrank & Mart. ex
(designated here): BRAZIL. Rio de Janeiro: Alto DC., Prodr. 3: 129. 1828 ≡ Tibouchina semidecandra
Macahé, Friburgo, 19 Dec 1887, Glaziou, A.F.M. 16802 (Schrank & Mart. ex DC.) Cogn. in Martius, Fl. Bras.
(BR barcode BR0000005282821!; isolectotypes: B 14(3): 309–310. 1885 – Holotype: BRAZIL. Minas Gerais:
[probably destroyed; F negative F0BN016793!], BR bar- campestribus montium altiorum prope Villan Riccam et
codes BR0000005289097! & BR0000005289233!, C Villan Principis, s.d., Martius, C.F.P. von 766 (M barcode
barcodes C10015088! & C10015089!, F barcode M-0005194!; isotype G-DC barcode G00310192!).
V0063997F!, G!, K barcode K000329019!, LE!, P bar-
codes P00708658!, P00708659! & P00708660!, R bar- 138. Pleroma stellipile (Wurdack) P.J.F.Guim. & Michelang.,
code R 000009293!, RB barcodes 00541583! & comb. nov. ≡ Tibouchina stellipilis Wurdack in Phytolo-
00541585!). gia 29(2): 138–139. 1974 – Holotype: BRAZIL. Maran-
hão: Morro do Chapéu, Carolina, foot of sandstone
136. Pleroma sellowianum (Cham.) P.J.F.Guim. & Miche- cliffs, 29 Mar 1967, Prance, G.T. 4806 (US barcode
lang., comb. nov. ≡ Lasiandra sellowiana Cham. in Lin- 00120095!; isotype: CAS barcode 0004861 [online
naea 9: 440. 1835 ≡ Tibouchina sellowiana (Cham.) image!], K barcode K000329127!, NY barcode
Cogn. in Martius, Fl. Bras. 14(3): 304–305, t. 69, fig. 2. 00019390!).
1885 – Lectotype (designated here): BRAZIL. s.d.,
Sellow, F. 1145 (US barcode 00730913!; isolectotypes: 139. Pleroma stenocarpum (Schrank & Mart. ex DC.) Triana
F barcode V0063573F [fragment]!, K barcode in Trans. Linn. Soc. London, Bot. 28(1): 44. 1872
K000329008!, US barcode 00730887!, W No. 0071711!). (“1871”) ≡ Lasiandra stenocarpa Schrank & Mart. ex
= Tibouchina sellowiana var. brevifolia Cogn. in Martius, DC., Prodr. 3: 130. 1828 ≡ Tibouchina stenocarpa
Fl. Bras. 14(3): 305. 1885, syn. nov. – Lectotype (desig- (Schrank & Mart. ex DC.) Cogn. in Martius, Fl. Bras.
nated here): BRAZIL. São Paulo [or Rio de Janeiro], 14(3): 344–345, t. 82. 1885 – Holotype: BRAZIL. Bahia:
19 Apr 1881 (or 1882), Glaziou, A.F.M. 12692 in arenariis saxis prope Sincorâ et in graniticis prope
(BR barcode BR0000005520312!; isolectotypes: C!, Maracás, s.d., Martius, C.F.P. von s.n. (M barcode M-
G barcode G00368038!, K barcode K000329005!, LE!, 0165861!; isotype: G-DC barcode G00310503!).
P barcodes P00708805!, P00708806! & P00708807!, = Tibouchina stenocarpa var. boliviensis Cogn. in Martius,
R barcode R 000009295!). Fl. Bras. 14(3): 345. 1885, syn. nov. – Holotype:
Note: Glaziou transcribed different dates and locations for BOLIVIA. From 1500 to 2000 miles in the interior; lat.
this collection. 15–18 South, s.d., Bridges 257 (LE!; isotype: LE!).
= Tibouchina imperatoris Saldanha & Cogn., Bouquet Mél- = Tibouchina stenocarpa var. latifolia Cogn. in Martius,
ast. Brés.: t. 2, fig. 2. 1887, syn. nov. – Holotype: BRA- Fl. Bras. 14(3): 345. 1885, syn. nov. – Lectotype (desig-
ZIL. Rio de Janeiro: Serra do Alto Macahé, 18 Aug nated here): BRAZIL. Minas Gerais: Caldas, 1861,
1887, Glaziou, A.F.M. 15982 (P barcode P00121407!; Regnell, A.F. I 159 (R barcode R 000168431!; isolecto-
isotypes: BR barcodes BR0000005223336! & type: C!, LE!,).
BR0000005223695!, C barcode C10015073!, P barcode Other syntypes: BRAZIL. Minas Gerais: Lagoa Santa,
P00121408!, R barcode R 000009297!). 1834, Warming, E. 2238/2 (C!); BRAZIL. Minas Gerais:
= Tibouchina imperatoris var. parvifolia Saldanha & Cogn., Lagoa Santa, 1835, Warming, E. 311 (C!); BRAZIL. São
Bouquet Mélast. Brés.: t. 2, fig. 2. 1887, syn. nov. – Lec- Paulo: prope Itú, s.d., Lund s.n. (BR barcode
totype (designated here): BRAZIL. Minas Gerais: BR0000005530403 [fragment]!); BRAZIL. São Paulo:
Ayuruoca, le bois, 22 Jan 1886, Glaziou, A.F.M. 15983 in campis siccis apricis as Mugy prov. S. Paulo, s.d.,
(BR barcode BR0000005224029!; isolectotypes: B Mosén, C.W.H. 1291 (S No. S09-13109!); BRAZIL.
[probably destroyed; F negative F0BN016754!], C bar- Minas Gerais, s.d., Claussen, P. s.n. (not located).
code C10015072!, K barcode K000329004!, P barcode = Tibouchina stenocarpa var. longifolia Cogn. in Martius,
P00121409!, R barcode R 000009298!, RB barcode Fl. Bras. 14(3): 345. 1885, syn. nov. – Lectotype (desig-
00231199!). nated here): BRAZIL. Minas Gerais: Caldas, 20 Apr
= Tibouchina ulaei Cogn. in Candolle & Candolle, Monogr. 1874, Mosén, C.W.H. 1292 (S No. S-R-6126!; isolecto-
Phan. 7: 202. 1891, syn. nov. – Lectotype (designated types: R barcode R 000168430!, S No. S09-13114!).
here): BRAZIL. Santa Catarina: am Ufer des Tuberão Other syntypes: BRAZIL. São Paulo, s.d., De Saint-
bei Minas, Apr 1889, Ule, E.H.G. 1126 (BR barcode Hilaire, A.F.C.P. D709 (P barcodes P00708789!
BR0000005226009!; isolectotypes: B [probably & P00708790!).
destroyed; F negative F0BN016803!], BR barcode = Tibouchina caldensis Cogn. in Martius, Fl. Bras. 14(3):
BR0000005225361!, HBG barcode HBG-523033 306. 1885, syn. nov. – Holotype: BRAZIL. Minas Ger-
[online image!]). ais: Caldas, 8 May 1875, Regnell, A.F. III 1764 (S No.

53
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

S-R-6111!; isotype: BR barcode BR0000005224463 a.s.l., 29 Jun 1935, Brade, A.C. 14906 (RB barcode
[fragment]!). 00659607!; isotypes: NY barcode 00245751!, RB bar-
= Tibouchina granulosa var. angustifolia Britton in Bull. Tor- codes 00541543! & 00659638!, SP barcode SP002922!).
rey Bot. Club 17(3): 54. 1890, syn. nov. – Holotype:
BOLIVIA. Yungas, 6000 ft., s.d., Rusby, H.H. 2245 146. Pleroma tomentulosum (Wurdack) P.J.F.Guim. &
(NY barcode 00245769!). Michelang., comb. nov. ≡ Tibouchina tomentulosa
Wurdack in Kew Bull. 50(4): 824–825. 1995 – Holotype:
140. Pleroma stipulaceum (Vinha) P.J.F.Guim. & Miche- BRAZIL. Bahia: Santa Terezinha, Serra da Pioneira –
lang., comb. nov. ≡ Tibouchina stipulacea Vinha in Bra- 0.3 km de P. Branca, alt. 800 m, 12 45′S, 39 32′W,
dea 1(30): 321. 1973 – Holotype: BRAZIL. Bahia: rod. 16 May 1983, Noblick, L.R. & al. 3233 (HUEFS barcode
Itaimbé/Itapebí, 15 May 1970, Santos, T.S. 800 (CEPEC HUEFS003389!; isotype: NY barcode 02286702!, P bar-
barcode 00005783!; isotype: US barcode 00120099!). code P01167438!, RB barcode 00883978!, US barcode
00478965!).
141. Pleroma subglabrum (Wurdack) P.J.F.Guim. & Miche-
lang., comb. nov. ≡ Tibouchina subglabra Wurdack in 147. Pleroma trichopodum DC., Prodr. 3: 151. 1828 ≡
Phytologia 53(2): 133–134. 1983 – Holotype: BRAZIL. Lasiandra trichopoda (DC.) Naudin in Ann. Sci. Nat.,
Bahia: Morro do Chapéu, rio do Ferro Doido 19.5 km Bot., sér. 3, 13(3): 159. 1850 ≡ Tibouchina trichopoda
SE of Morro do Chapéu on the Ba 052 highway to Novo (DC.) Baill. in Adansonia 12: 75. 1877 – Lectotype
Mundo, alt. ca. 900 m, 11 38′S, 41 02′W, 13 May 1980, (designated here): BRAZIL. Rio de Janeiro: 1881,
Harley, R.M. 22895 (CEPEC barcode 00029661!; isotype: Langsdorff, G.H. von s.n. (P barcode P00708788!
CEPEC barcode 00040490!, K, US barcode 00120099!). [F negative F0BN036175!]; isolectotype: P barcode
P00708787!).
142. Pleroma subsessile F.S.Mey. & L.Kollmann in Phyto- = Lasiandra tibouchinoides Schrank & Mart. ex DC., Prodr.
taxa 282(3): 205–208, fig. 6–7. 2016 – Holotype: BRA- 3: 128. 1828 ≡ Tibouchina trichopoda var. tibouchinoides
ZIL. Espírito Santo: Afonso Cláudio, Alto Três Pontões Cogn. in Martius, Fl. Bras. 14(3): 325. 1885, syn. nov. –
(Pico da Antena de Rádio Amador), 16 Aug 1990, Boudet Holotype: BRAZIL. São Paulo: campis inter Villa de Lor-
Fernandes, H.Q. & al. 2999 (MBML barcode eno et Guaratingueta, s.d., Martius, C.F.P. von s.n.
MBML006043 [online image!]; isotypes: RB, UPCB). (M barcode M-0005199!; isotype: G-DC barcode
G00310197!).
143. Pleroma taperoense (Wurdack) P.J.F.Guim. & Miche- = Lasiandra multiceps Naudin in Ann. Sci. Nat., Bot., sér.
lang., comb. nov. ≡ Tibouchina taperoensis Wurdack in 3, 13(3): 149. 1850 ≡ Tibouchina multiceps (Naudin)
Phytologia 64(4): 295–296. 1988 – Holotype: BRAZIL. Cogn. in Martius, Fl. Bras. 14(3): 326–327. 1885, syn.
Bahia: Taperoá, ramal de fazendas a W de Taperoá a nov. – Holotype: BRAZIL. São Paulo: 1833,
4–7 km da cidade, 8 Dec 1980, Carvalho, A.M.; Mattos Gaudichaud, C. 815 (P barcode P00708709!).
Silva, L.A. & Hage, J.L. 354 (CEPEC barcode = Tibouchina multiceps var. parvifolia Cogn. in Martius,
00022902!; isotypes: HUEFS barcode HUEFS003754!, Fl. Bras. 14(3): 327. 1885, syn. nov. – Lectotype (desig-
MBM, US barcodes 00048299! & 01013508!). nated here): BRAZIL. s.d., Sellow, F. 1146 (K barcode
K000329024!).
144. Pleroma tedescoi (Meirelles, L.Kollmann & R.Gold- Other syntype: BRAZIL. s.d., Sellow, F. 1722 (not located).
enb.) P.J.F.Guim. & Michelang., comb. nov. ≡ Tibou-
china tedescoi Meirelles, L.Kollmann & R.Goldenb. in 148. Pleroma trinervium P.J.F.Guim. in Syst. Bot. 40(2):
Kew Bull. 67(3): 461–464, fig. 1, 2. 2012 – Holotype: 554–559, fig. 2D,H,L,P, 4. 2015 – Holotype: BRAZIL.
BRAZIL. Espírito Santo: mun. Castelo, Parque Estadual Minas Gerais: Serra do Funil au Rio Paranaua, 29 Apr
do Forno Grande, Forninho, 21 Jan 2009, 1892, Glaziou, A.F.M. 19301 (P barcode P00708663!;
Goldenberg, R.; Fraga, C.N. & Forzza, R. 1290 (RB bar- isotypes: B [probably destroyed; F negative F0BN
code 00549493!). 016801!], K!, R!).

145. Pleroma thereminianum (DC.) Triana in Trans. Linn. 149. Pleroma tuberosum Gardner ex Triana in Trans. Linn.
Soc. London, Bot. 28(1): 41. 1872 (“1871”) ≡ Lasiandra Soc. London, Bot. 28(1): 46. 1872 (“1871”) ≡ Tibouchina
thereminiana DC., Prodr. 3: 129. 1828 ≡ Tibouchina tuberosa (Gardner ex Triana) Cogn. in Martius, Fl. Bras.
thereminiana (DC.) Cogn. in Martius, Fl. Bras. 14(3): 14(3): 395–396. 1885 – Holotype: BRAZIL. Goiás: in
310. 1885 – Holotype: BRAZIL. 1819, Theremin, K.W. clefts of dry rocks Serra de Natividade, 1840,
s.n. (G-DC barcode G00310193!). Gardner, G. 3738 (BM barcode BM000953964!; iso-
= Tibouchina angraensis Brade in Arq. Inst. Biol. Veg. 4(1): types: B [probably destroyed; F negative F0BN0
74; pl. 4. 1938, syn. nov. – Holotype: BRAZIL. Rio de 16802!], BR barcode BR0000005530731!, E barcode
Janeiro: Angra dos Reis, Serra do Mar, Jussural, 300 m E00531242 [online image!], G-DC barcode G00319063

54
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

[online image!], K barcodes K000329091! & K0003 Martius, Fl. Bras. 14(3): 350–351. 1885 – Holotype:
29092!, P barcodes P00708772! & P00708773!). BRAZIL. Bahia, 1844, Blanchet, J.S. 291 (P barcode
2274607!; isotypes: B [probably destroyed; F negative
150. Pleroma urceolare (Schrank & Mart. ex DC.) Triana F0BN016806!], BM barcode BM000020541!, BR bar-
in Trans. Linn. Soc. London, Bot. 28(1): 42. 1872 code BR0000005226979!, G!, W No. 0053860!).
(“1871”) ≡ Osbeckia urceolaris Schrank & Mart. ex
DC., Prodr. 3: 139. 1828 ≡ Tibouchina urceolaris 154. Pleroma villosissimum Mart. ex Triana in Trans. Linn.
(Schrank & Mart. ex DC.) Cogn. in Martius, Fl. Bras. Soc. London, Bot. 28(1): 43. 1872 (“1871”) ≡ Lasiandra
14(3): 355, t. 84, fig. 1. 1885 – Holotype: BRAZIL. holosericea Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3):
[Rio de Janeiro]: Rio-Janeiro circa urbem, s.d., Martius, 139. 1850 ≡ Tibouchina villosissima (Mart. ex. Triana)
C.F.P. von s.n. (M barcode M-0165860!). Cogn. in Martius, Fl. Bras. 14(3): 330–331, t. 77. 1885
= Tibouchina litoralis Ule in Notizbl. Königl. Bot. Gart. Ber- – Holotype: BRAZIL. Minas Gerais: 1838, Claussen, P.
lin 6(60): 351. 1915, syn. nov. – Holotype: BRAZIL. Rio 556 (P barcode P00708945!; isotypes: BR!, P barcode
de Janeiro: in Sümpfen bei Cabo Frio, Oct 1899, Ule, P00708944!).
E.H.G. 4728 (B [probably destroyed; F negative = Tibouchina villosissima var. longifolia Cogn. in Martius,
F0BN016762!]; isotypes:, HBG barcode HBG-523023, Fl. Bras. 14(3): 331. 1885, syn. nov. – Lectotype (desig-
R barcode R 000114953!). nated here): BRAZIL. Goiás: Catalão, Aug 1834, Riedel,
L. 1543 (LE!).
151. Pleroma ursinum (Cham.) Triana in Trans. Linn. Soc. Other syntypes (excluded because original material is
London, Bot. 28(1): 43. 1872 (“1871”) ≡ Lasiandra attributed to two different varieties): BRAZIL. Minas
ursina Cham. in Linnaea 9. 443. 1835 ≡ Tibouchina Gerais. (fl), Claussen, P. 556 (BR!, P barcodes
ursina (Cham.) Cogn. in Martius, Fl. Bras. 14(3): P00708944! & P00708945!).
351–352. 1885 – Lectotype (designated here): BRA- = Tibouchina villosissima var. parvifolia Cogn. in Martius,
ZIL. s.d., Sellow, F. 5407 (K barcode K000329043!; iso- Fl. Bras. 14(3): 331. 1885, syn. nov. – Lectotype (desig-
lectotype: K barcode K000329044 [no collector nated here): BRAZIL. Minas Gerais: 1840, Claussen, P.
number]!). 1067 (BR barcode BR0000005226658!; isolectotype: G!).
Other syntype: BRAZIL. s.d., Sellow, F. 659 (B [probably = Tibouchina villosissima var. petiolaris Cogn. in Martius,
destroyed; F negative F0BN016804!], US barcode Fl. Bras. 14(3): 331. 1885, syn. nov. – Lectotype (desig-
01882587!). nated here): BRAZIL. Minas Gerais, 1817, Langsdorff,
G.H. von 2159 (LE!; isolectotype: LE!).
152. Pleroma urvilleanum (DC.) P.J.F.Guim. & Michelang.,
comb. nov. ≡ Lasiandra urvilleana DC., Prodr. 3: 130. 155. Pleroma vimineum (D.Don) D.Don in Mem. Wern. Nat.
1828 ≡ Tibouchina urvilleana (DC.) Cogn. in Martius, Hist. Soc. 4: 296. 1823 ≡ Rhexia viminea D.Don. in Bot.
Fl. Bras. 14(3): 358–359, t. 84, fig. 2. 1885 – Lectotype Reg. 8: 664. 1822 ≡ Lasiandra viminea (D.Don) Naudin
(designated here): BRAZIL. Santa Catarina: 1825, in Ann. Sci. Nat., Bot., sér. 3, 13(3): 159. 1850 ≡ Tibou-
D’Urville, J.S.C.D. s.n. (W No. 1889-0015758!; isolecto- china viminea (D.Don) Cogn. in Martius, Fl. Bras.
type: B [probably destroyed; F negative F0BN016805!], P 14(3): 359–360. 1885 – Lectotype (designated here):
barcodes P00708937! & P00708938!). [illustration in] Bot. Reg.: 8. t. 664. 1822.
= Tibouchina paulistana Hoehne in Anexos Mem. Inst. = Rhexia corymbosa Raddi in Mem. Mat. Fis. Soc. Ital. Sci.
Butantan, Secc. Bot. 1: 70. 1922 – Holotype: BRAZIL. Modena, Pt. Mem. Fis. 20: 124, t. 2, fig. 1. 1828 ≡ Tibou-
São Paulo: São Francisco dos Campos, 29 Dec 1896, china corymbosa (Raddi) Cogn. in Martius, Fl. Bras.
Löfgren, A. [CGGSP] 3433 (SP barcode SP001232!; iso- 14(3): 363–364, t. 86, fig. 1. 1885, syn. nov. – Holotype:
type: US barcode 00120083 [fragment]!). BRAZIL. Rio de Janeiro: s.d., Raddi, G. s.n. (PI; isotype:
= Tibouchina urceolaris var. papillosa Hoehne in Anexos FI barcode FI005211 [online image!]).
Mem. Inst. Butantan, Secc. Bot. 1: 79. 1922 – Holotype:
BRAZIL. São Paulo: Campinas, s.d., Novaes, C. s.n. 156. Pleroma virgatum Gardner in London J. Bot. 2: 350.
(SP barcode SP001236!). 1843 ≡ Lasiandra virgata (Gardner) Naudin in Ann. Sci.
= Tibouchina urvilleana var. glandulifera Wurdack in Sellowia Nat., Bot., sér. 3, 13(3): 159. 1850 ≡ Tibouchina virgata
14: 122. 1962, syn. nov. – Holotype: BRAZIL. Santa (Gardner) Cogn. in Martius, Fl. Bras. 14(3): 365–366.
Catarina: Itajaí, Cunhas, 29 Nov 1954, Klein, R.M. 882 1885 – Lectotype (designated here): BRAZIL. Rio de
(US barcode 00120105!; isotype: M barcode M0165859!). Janeiro: Organ mountain, elev. 4500 ft., May 1837,
Gardner, G. 403 (BM barcode BM000020625!; isolecto-
153. Pleroma velutinum (Naudin) Triana in Trans. Linn. type: K barcode K000329058!, NY barcode 00245682!).
Soc. London, Bot. 28(1): 42. 1872 (“1871”) ≡ Lasiandra = Tibouchina nervulosa Cogn. in Martius, Fl. Bras. 14(3):
velutina Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): 315, t. 70. 1885, syn. nov. – Lectotype (designated
144. 1850 ≡ Tibouchina velutina (Naudin) Cogn. in here): BRAZIL. Rio de Janeiro: Frade, Serra dos Orgãos,

55
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

4 Apr 1870, Glaziou, A.F.M. 3974 (BR barcode minute setae, the remaining surface sparingly hairy with
BR0000005226023!; isolectotypes: C barcode C1001 simple or glandular trichomes. Calyx not calyptriform, with
5091!, K barcode K000329011!, P barcodes P00708 4 short lobes, persistent in fruit, linear-subulate. Corolla
719! & P00708720!, R barcode R 000009289!). patent; petals 4, free, spreading, purplish-pink or white. Sta-
= Tibouchina nervulosa var. glabrata Cogn. in Candolle mens 8, dimorphic, consisting of 4 large antesepalous sta-
& Candolle, Monogr. Phan. 7: 209. 1891, syn. nov. – mens fertile and 4 atrophied antepetalous ones unfertile.
Lectotype (designated here): BRAZIL. Rio de Janeiro: Anthers in the large stamens ventrally channeled, oblong
Alto Macahé, 21 Apr 1888, Glaziou, A.F.M. 16794 part. to oblong-subulate, slightly arcuate and not lanceolate. Con-
(BR barcode BR0000005226665!; isolectotypes: C bar- nective in the large stamens prolonged conspicuously in the
code C10015092!, G [on 3 sheets]!, K barcode basal part, slightly arched, with two ventral lobes at the
K000329011!, LE!, P barcode P00708718!). point of articulation with the filament; in the atrophied sta-
= Tibouchina nervulosa var. ovata Cogn. in Candolle & Can- mens, connective is shortly prolonged and has two small
dolle, Monogr. Phan. 7: 209. 1891, syn. nov. – Lectotype ventral lobes, with a small apical pore directed ventrally.
(designated here): BRAZIL. Rio de Janeiro: Alto Filaments thin, terete, glabrous. Ovary superior, 4-locular,
Macahé, 8 Apr 1888, Glaziou, A.F.M. 16794 part. glabrous or with few trichomes at the apex; axile placenta-
(BR barcode BR0000005226337!; isolectotypes: BR bar- tion; style clavate, apically incurved, stigma broadly trun-
code BR0000005226641!, K barcode K000329010!, MO cate, glabrous. Mature capsule dry, semiwoody, 8-winged,
barcode MO-100754164!, P barcodes P00708714!, loculicidal. Seeds numerous and tiny, cochleate and tuber-
P00708715! & P00708716!, R barcode R 000009288!). culate, hilum terminal.
= Tibouchina aspericaulis Cogn. in Candolle & Candolle, A monospecific genus, only represented by Schwackaea
Monogr. Phan. 7: 209. 1891, syn. nov. – Lectotype (desig- cupheoides, which occurs from southern Mexico (Chiapas)
nated here): BRAZIL. Rio de Janeiro: mun. Petrópolis, throughout Central America, chiefly on the Pacific slope, to
14 Mar 1888, Glaziou, A.F.M. 16795 (BR barcode northern Colombia. It grows in humid places in savannas,
BR0000005225378!; isolectotypes: B [probably destroyed; along ditches, in dried-out river beds, open fields and planta-
F negative F0BN016726!]), BR barcode BR000000 tions, and between cliffs near the sea; from sea level to
5226030!, C barcodes C10015093! & C10015094!, G 2000 m (Renner, 1994; Kriebel, 2016).
[on 2 sheets]!, K barcode K000329012!, P barcode Schwackaea is easily differentiated from other Melasto-
P00708727!). mateae by the combination of annual habit and 8-winged
hypanthia that closely envelop the capsules.
157. Pleroma wurdackianum (R.Romero & A.B.Martins)
P.J.F.Guim. & Michelang., comb. nov. ≡ Svitramia wur- Tibouchina Aubl., Hist. Pl. Guiane 1: 445–446, t. 177. 1775 ≡
dackiana R.Romero & A.B.Martins in Kew Bull. 58(3): Savastania Scop., Intr. Hist. Nat.: 213. 1777 ≡ Lasiandra
410–412, fig. 4. 2003 – Holotype: BRAZIL. Minas Ger- sect. Tibouchina Naudin in Ann. Sci. Nat., Bot., sér.
ais: São Roque de Minas, Parque Nacional da Serra da 3, 13(3): 153. 1850 ≡ Pleroma sect. Tibouchina
Canastra, morro próximo à sede administrativa, 20 15′ (Naudin) Triana in Trans. Linn. Soc. London, Bot.
22′′S, 46 25′20′′W, 1352 m, 9 May 1995, Romero, R. 28(1): 45. 1872 (“1871”), pro parte ≡ Tibouchina sect.
& al. 2162 (HUFU; isotypes: K barcode K000530432!, Tibouchina (Naudin) Cogn. in Martius, Fl. Bras. 14(3):
R barcode R 000212036!, UEC, US barcode 00810593!). 374. 1885 – Type: Tibouchina aspera Aubl.
= Lasiandra sect. Barbigerae Naudin in Ann. Sci. Nat., Bot.,
Schwackaea Cogn. in Candolle & Candolle, Monogr. Phan. sér. 3, 13(3): 153. 1850 ≡ Tibouchina sect. Barbigerae
7: 182–183. 1891 – Type: Schwackaea cupheoides (Naudin) Cogn. in Martius, Fl. Bras. 14(3): 376. 1885 –
(Benth.) Cogn. (≡ Heeria cupheoides Benth. ≡ Pterogas- Type (designated here): Tibouchina aegopogon
tra cupheoides (Benth.) Seem.). (Naudin) Cogn. (≡ Lasiandra aegopogon Naudin).
Erect annual herb, 15–40(–50) cm tall; trichotomously Shrubs, subshrubs or treelets, strigose in all structures
branched, the branchlets quadrangular, setulose on the with trichomes that are compressed into flattened scales of
nodes and sometimes along the angles, trichomes short varying sizes and shapes. Leaves opposite, decussate or verti-
and simple except for reproductive structures where they cillate, blades of a pair essentially equal in size, symmetrical,
are setose and in the hypanthium where they are rarely petiolate or rarely sessile, coriaceous, covered with scales or
small glandular, otherwise glabrous. Leaves opposite, strigose, tomentose or glandular, (1–)3–5(–7)-nerved. Inflo-
decussate, symmetrical, blades of a pair essentially equal rescence a terminal anthotelic frondobracteose panicle or
in size, petiolate, papyraceous or membranaceous, moder- some modification with reduction of branches and the
ately setulose or glabrous on both surfaces; 3-nerved. Inflo- agglomeration of dichasia in glomerules; bracteoles 2 or
rescence consisting of solitary flowers borne in the upper 4, persistent, smaller than the mature hypanthium; free or less
leaf axils or ramifications; ebracteate. Flowers perigynous; commonly, the inner bracteoles united into a cuplike involu-
subsessile. Hypanthium narrowly oblong, with 8 thick lon- cre. Flowers perigynous; pedicellate. Hypanthium campanu-
gitudinal wings, the wings dorsally tuberculate and with late, the sepals and bracts mostly covered with flattened

56
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

scales externally, rarely densely covered with villous and copi- BM000953966 [without collector number]!, K barcode
ous white trichomes up to 6 mm. Calyx not calyptriform, with K000329072!, NY barcode 00581288!).
conspicuous lobes, lobes 5, persistent to fruit, triangular. Other syntype: BRAZIL. Minas Gerais: in pascuis siccis
Corolla patent; petals 5, free and spreading, purple, lilac, color aridis glareosis montosis ad Serra de Ipicons, s.d., Pohl,
remains unchanged with age, obovate, glabrous. Stamens 10; J.B.E. 643 (not located).
all fertile; isomorphic, or weakly dimorphic, consisting of
5 large antesepalous stamens and 5 smaller antepetalous ones. 2. Tibouchina albescens Cogn. ex P.J.F.Guim., A.L.F.Oliveira
Anthers with 2-celled ventrally channeled, subulate, slightly & R.Romero in Syst. Bot. 40(4): 1003–1005, fig. 1A–C, 2,
arcuate and lanceolate apical pores ventrally inclined. Con- 4A–E. 2015 – Holotype: BRAZIL. Goiás: fazenda dos Vea-
nectives variously prolonged and modified ventrally at fila- deiros, s.d., Glaziou, A.F.M. 21376 (P barcode P00708771!;
ment insertion into horizontal or upturned bilobed ventral isotypes: B [probably destroyed; F negative F0BN016724!],
appendages glabrous or with a cluster of long filamentous BR barcode BR0000005209156!, K).
hairs. Filaments glabrous or with sparse long filamentous
hairs. Ovary superior, 5-celled; setulose apically; enclosed 3. Tibouchina araguaiensis P.J.F.Guim. in Novon 23(1):
by the hypanthium but not fused to it; axile placentation; style 44–45, fig. 2. 2014 – Holotype: BRAZIL. Tocantins:
glabrous, hooked or incurved apically, typically declined to Pium, Parque Nac. Araguaia, ponta da ilha, Rio Javaés,
side of flower opposing stamens; stigma punctiform, incon- 195 m a.s.l., 09 51′43′′S, 50 10′35′′W, 28 Mar 1999,
spicuous. Mature capsule dry, semiwoody, loculicidal, not Mendonça, R.C.; Silva, M.A.; Cardoso E.; Oliveira,
angled. Seeds numerous and tiny or rarely a single seed N.R. & Souza, N.G. 4083 (IBGE; isotype: FLOR).
unusually large, cochleate to elongate-cochleate and tubercu-
late, rarely smooth, hilum terminal. 4. Tibouchina aspera Aubl., Hist. Pl. Guiane 1: 446–448,
Currently there are 35 taxa recognized in Tibouchina: fig. 177. 1775 ≡ Melastoma aromaticum Vahl., Eclog.
30 species, 4 varieties and 1 subspecies. Tibouchina occurs Amer. 1: 41. 1796, nom. illeg. superfl. ≡ Rhexia aspera
from Central America (Belize, Honduras, Nicaragua, Costa (Aubl.) Willd., Sp. Pl. 2: 304. 1798 ≡ Melastoma tibou-
Rica, Panama) to Amazonian Bolivia and Brazil, including china Desr. in Lamarck, Encycl. 4: 49. 1798, nom. illeg.
Venezuela and all the Guianas. In Brazil it occurs in northern superfl. – Lectotype (designated here): French Guiana.
and central-western regions, with a southern boundary in the Habitat in locis aridis Aroura prope praedium de Bertier,
Cerrado of Minas Gerais and Mato Grosso do Sul States. It s.d., Aublet, J.B.C.F. s.n. (BM barcode BM000953948!;
inhabits lowland savannas (up to 1200 m) and Cerrado in isolectotypes: MPU barcode MPU013398 [online
grassland, open areas, forest edges, stream margins and rocky image!]).
slopes and outcrops. = Tibouchina belizensis Lundell in Amer. Midl. Naturalist
Tibouchina has leaves and hypanthia covered with scale- 29(2): 483–484. 1943 – Holotype: HONDURAS. Toledo
like trichomes except T. catharinae and T. striphnocalyx, which District, Cow pen under oak trees in pine ridge, 17 Sep
have penicillate hairs on the hypanthia. The presence of a lepi- 1942, Gentle, P.H. 4155 (MICH barcode 1111899
dote indumentum is shared with Andesanthus, but the stamens [online image!]; isotypes: A barcode 00073246 [online
in Tibouchina are slightly dimorphic with purple anthers and a image!], F barcode V0063969F!, GH, NY barcode
developed pedoconnective in both whorls. In Andesanthus the 00245746!, LL barcodes LL00208466 & LL00372242
stamens are isomorphic with yellow anthers or if dimorphic [online images!], US barcode 00120041!).
then with larger anthers yellow and the smaller ones pink or = Tibouchina brachyanthera Pittier in J. Wash. Acad. Sci.
red. These genera are also easily separated based on the habitat, 13(17): 386. 1923 – Holotype: VENEZUELA. Toro-
with Andesanthus found in cool montane forests. coco: Trujillo, 1100 m, in sunny spots, 11 Jan 1922,
Jahn, A. 755 (US barcode 00120043!).
1. Tibouchina aegopogon (Naudin) Cogn. in Martius,
Fl. Bras. 14(3): 383–384. 1885 ≡ Lasiandra aegopogon 4a. Tibouchina aspera var. asperrima Cogn. in Martius,
Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): 153–154. Fl. Bras. 14(3): 375. 1885 – Lectotype (designated
1850 ≡ Pleroma aegopogon (Naudin) Triana in Trans. here): SURINAME. s.d., Hostmann, F.W.R. & Kappler, A.
Linn. Soc. London, Bot. 28(1): 45. 1872 (“1871”) – Holo- 989 (K barcode K000329064!; isolectotypes: LE!, P
type: BRAZIL. Minas Gerais: 1844, Weddell, H.A. 1911 barcodes P00708728! & P00708729!, S No. S09-13208
(P barcode P00708770!). [online image!]).
Other syntypes: GUYANA. Berbice, s.d., Schomburgk,
1a. Tibouchina aegopogon var. angustifolia Cogn. in Mar- R.H. 252 part. (BR [fragment]!, P barcodes P00708730!
tius, Fl. Bras. 14(3): 384. 1885 – Lectotype (designated & P00708731!, TCD); “Savannis Parae” [location uncer-
here): BRAZIL. Goiás: in pascuis siccis arridis glareosis tain], s.d., Splitgerber, F.L. s.n. (MPU barcode
montosis! Montium Serra de Ipicons, s.d., Pohl, MPU013397 [online image!]); SURINAME. “Pl. Berlyn,
J.B.E. 1191 (W No. 0053869!; isolectotypes: B [probably Para”, s.d., Wullschlaegel, H.R. 177 (BR barcode
destroyed; F negative F0BN016722!], BM barcode BR0000005523290!).

57
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

4b. Tibouchina aspera var. poeppigii Cogn. in Martius, elev. 1850–1900 m, 15 Jan 1954, Maguire, B.; Wurdack,
Fl. Bras. 14(3): 375. 1885 – Lectotype (designated J.J. & Bunting, G.S. 37294 (NY barcode 00245761!; iso-
here): BRAZIL. Pará: in arenosis campestribus ad Cam- types: G barcode G00368017!, K barcode K000329167!,
pinas de Colares, s.d., Poeppig, E.F. 3014 (BR barcode RB barcode 00541556!, S No. S-R-6115 [online image!],
BR0000005523955!). US barcode 00120051!, VEN barcode 42247 [online
image!]).
5. Tibouchina barbigera (Naudin) Baill. in Adansonia 12:
75. 1877 ≡ Lasiandra barbigera Naudin in Ann. Sci. 11. Tibouchina duidae Gleason in Fieldiana, Bot. 28(2): 428.
Nat., Bot., sér. 3, 13(3): 155. 1850 ≡ Pleroma barbigerum 1952 – Holotype: VENEZUELA. Amazonas: summit of
(Naudin) Triana in Trans. Linn. Soc. London, Bot. 28(1): Cerro Duida, alt. 1700–1980 m, 1 Sep 1944, Steyermark,
45. 1872 (“1871”) – Holotype: BRAZIL. 1844, Weddell, J.A. 58203 (F barcode V0063983F!; isotypes: GH
H.A. 2601 (P barcode P00708766!; isotypes: F barcode barcode 00073260 [online image!], NY barcode
V0047549F [fragment]!, P barcode P00708767!). 00245762!, US barcode 00120052!).

6. Tibouchina bipenicillata (Naudin) Cogn. in Martius, 12. Tibouchina edmundoi Brade in Arch. Jard. Bot. Rio de
Fl. Bras. 14(3): 385–386. 1885 ≡ Lasiandra bipenicillata Janeiro 16: 10–11, t. 4. 1959 – Holotype: BRAZIL. Pará:
Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): 156. 1850 ≡ Serra do Cachimbo, 20 Sep 1955, Pereira, E. 1878
Pleroma bipenicillatum (Naudin) Triana in Trans. Linn. (RB barcode 00541558!; isotype: US barcode 00680002
Soc. London, Bot. 28(1): 45. 1872 (“1871”) – Lectotype [fragment]!).
(designated here): VENEZUELA. Mérida: San Cristo-
bal, 600 m, Oct 1842, Linden, J.J. 699 (P barcode 13. Tibouchina exasperata (Naudin) Cogn. in Martius,
P00481684!; isolectotypes: BR!, FI barcode FI004722 Fl. Bras. 14(3): 377–378. 1885 ≡ Lasiandra exasperata
[online image!], G barcode G00368025!, GENT barcode Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(13): 154–155.
BR0000005209200 [online image!], K barcodes 1850 – Holotype: BRAZIL. Goiás: montibus Perineos,
K000329138!, K000329139! & K000329140!, LE!, P s.d., St.-Hilaire A.F.C.P. de C1 697 (P barcode
barcode P00481685!, W!). P00708690!; isotypes: F barcode V0063569F [frag-
Other syntypes: VENEZUELA. Mérida: haud procul ab ment]!, P barcodes P05228409! & P00708691!).
oppidulo San Cristobal, 400–1000 m, 1846, Funck,
N. & Schlim, L.J. 1275 (BR barcode BR00000052 14. Tibouchina fraterna N.E.Br. in Trans. Linn. Soc.
09538!, G barcode G00368024!, LE!, US barcode London, Bot. 6(1): 28. 1901 – Holotype: BRITISH GUI-
00680003 [fragment]!). ANA. Upper slopes Roraima: Mount Roraima Expedi-
tion. [Fladas superiores del Cerro Roraima], 1894,
7. Tibouchina bruniana P.J.F.Guim. in Novon 23(1): 42–44, Quelch, J.J. & McConnell, F. 17 (K barcode K000
fig. 1. 2014 – Holotype: BRAZIL. Goiás: Niquelândia, 329115!).
próximo ao povoado de Macedo, ca. 7 km da Mina de
Níquel Tocantins, do lado esquerdo passando por baixo 14a. Tibouchina fraterna subsp. paruana Wurdack in Mem.
do teleférico, 14 18′59′′S, 48 23′34′′W, 20 Sep 1996, New York Bot. Gard. 10(1): 101. 1958 – Holotype: VEN-
Silva, M.A.; Fonseca, M.L.; Lopes, E.C. & Ferreira, EZUELA. Amazonas: Serrania Paru, frequent in cumbre
C.C.S. 3172 (IBGE!; isotype: UB!). along west rim just north of camp caño head, elev.
2000 m, 31 Jan 1951, Cowan, R.S. & Wurdack, J.J. 31060
8. Tibouchina catharinae Pittier in Bol. Soc. Venez. Ci. Nat. (NY barcode 00245765!; isotypes: S No. S-R-6117!, US
11: 18. 1947 – Holotype: VENEZUELA. Amazonas: Cerro barcode 00120055!, VEN barcode 42267 [online image!]).
Paraque, 1600 m, Alto Orinoco, Feb 1946, Phelps, K. 17
(VEN barcode 26269 [online image!]). 15. Tibouchina huberi Wurdack in Phytologia 69(5):
318–319. 1990 – Holotype: VENEZUELA. Amazonas:
9. Tibouchina cujabensis Cogn. in Martius, Fl. Bras. 14(3): Atures, on northwest part of Serra de Maigualida in headwa-
382–383, t. 88, fig. 2. 1885 – Lectotype (designated ters of Río Iguana (affluent of Río Asisa), 05 43′N, 65 19′
here): BRAZIL. Mato Grosso: in locis subhumidis W, elev. 1720 m, 25 Mar 1988, Huber, O. 12686 (VEN
prope Cujaba, Feb 1827, Riedel, L. 850 (LE No. 4153!; barcode 357992 [online image!]; isotypes: M barcodes
isolectotypes: BR barcode BR0000005209507!, LE M-0165840! & M-0165841!, US barcode 00323870!).
No. 4154!).
16. Tibouchina johnwurdackiana Todzia in BioLlania
10. Tibouchina dissitiflora Wurdack in Mem. New York Bot. (Ed. Espec.) 6: 537, fig. 1. 1997 – Holotype: BRAZIL.
Gard. 10(1): 101. 1958 – Holotype: VENEZUELA. Ama- Goiás: Sandstone summit of Serra Dourada, 20 km SE
zonas: Cerro de la Neblina, Río Yatua, frequent in west of Goiás Velho, 800 m, 18 Jan 1966, lrwin, H.S.; Souza,
escarpment savanna 4–8 km southwest of cumbre camp, R. & Santos, R.R. 11711 (UB barcode 0037523!;

58
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

isotypes: CAS barcode 0004854 [online image!], K bar- K000329071!, herbarium unknown [F negative
code K000276345!, NY barcode 00689107!, MO No. F0BN021207!]).
04840126 [barcode MO-1515257]!, SPF barcode SPF
00184547!, TEX barcode 00208185 [online image!], 23. Tibouchina pogonanthera (Naudin) Cogn. in Martius,
US barcodes 00679971! & 00955905!). Fl. Bras. 14(3): 381–382. 1885 ≡ Lasiandra pogonanthera
Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3): 155. 1850 ≡
17. Tibouchina karstenii Cogn. in Martius, Fl. Bras. 14(3): Pleroma pogonantherum (Naudin) Triana in Trans. Linn.
381. 1885 – Lectotype (designated here): COLOMBIA. Soc. London, Bot. 28(1): 45. 1872 (“1871”) – Holotype:
Bogotá, Guetame i Susumuco, s.d., Karsten, H. s.n. BRAZIL. s.l., s.d., Bonpland, A.J.A. s.n. (P barcode
(W No. 2002-0013116 [online image!]; isolectotype: LE!). P00708769!; isotype: P barcode P00708768!).
Other syntype: COLOMBIA. Meta: Saint Martin, Llano de
la Quebradita, Rio Meta, s.d., Karsten, H. s.n. (LE!). 23a. Tibouchina pogonanthera var. latifolia Cogn. in Mar-
tius, Fl. Bras. 14(3): 382. 1885 – Lectotype (designated
18. Tibouchina llanorum Wurdack in Mem. New York Bot. here): BRAZIL. Goiás: s.d., Pohl, J.B.E. s.n. (BR barcode
Gard. 10(5): 142. 1964 – Holotype: VENEZUELA. BR0000005209491!).
Cojedes: Sabanas between San Carlos and Acarigua, Other syntypes: BRAZIL. “Pl. Bras. Occidental”, s.d., Tam-
Cojedes, 15 Sep 1940, Chardon-Palacios, C.E. 223 berlik s.n. (BR barcode BR0000005530311 [fragment]!).
(US barcode 00120070!).
24. Tibouchina rosanae P.J.F.Guim. & Woodgyer in Phyto-
19. Tibouchina mathaei Cogn. in Martius, Fl. Bras. 14(3): taxa 246(2): 146–150, fig. 1 & 2. 2016 – Holotype: BRA-
380–381. 1885 ≡ Lasiandra lepidota Naudin in Ann. Sci. ZIL. Mato Grosso: Santa Cruz do Xingu, Parque Estadual
Nat., Bot., sér. 3, 13(3): 155. 1850 ≡ Pleroma barbigerum do Xingu, limite norte do parque, 09 45′22′′S, 52 38′06′′
var. lepidotum Triana in Trans. Linn. Soc. London, Bot. W, 289 m, 10 Mar 2011, (fl.), Soares, C.R.A.; Zappi,
28(1): 45. 1872 (“1871”), pro parte – Holotype: PERU. D.C.; Frisby, S.; Milliken, W.; Forte, A.F.; Silva,
Prov. Moyobamba: 1836, Mathews, A. 1267 (P barcode D.R. & Lindolfo, L.F. 3217 (RB barcode 00621965!; iso-
P00708661!; isotypes: E barcode E00531233 [online types: K!, UPCB).
image!], F barcodes V0063989F & V0063990F [both frag-
ments] [online images!], K barcodes K000329135! 25. Tibouchina sickii Brade in Arch. Jard. Bot. Rio de Janeiro
& K000329136!, NY barcode 00581285!). 14: 217–218, t. 4. 1956 – Holotype: BRAZIL. Mato
Grosso: Xavantina, Rio das Mortes, Fev 1947, Sick,
20. Tibouchina melastomoides (Naudin) Cogn. in Martius, H. B283 (RB barcode 00541586!; isotypes: RB barcode
Fl. Bras. 14(3): 377, t. 88, fig. 1. 1885 ≡ Lasiandra me- 00556741!, US barcode 00679970 [fragment]!).
lastomoides Naudin in Ann. Sci. Nat., Bot., sér. 3, 13(3):
154. 1850 ≡ Pleroma melastomoides (Naudin) Triana in 26. Tibouchina sipapoana Gleason in Phytologia 3:
Trans. Linn. Soc. London, Bot. 28(1): 45. 1872 (“1871”) 240–241. 1950 – Holotype: VENEZUELA. Amazonas:
– Holotype: BRAZIL. Goiás: Arrayas foot of Serra de Cerro Sipapo (Paraque), SE slopes North Park I, 12 Dec
Santa Brida, Feb 1841, Gardner, G. 3740 (P barcode 1948, Maguire, B. & Politi, L. 27658 (NY barcode
P00708819!; isotypes: E barcode E00531238 [online 00245807!; isotypes: BM barcode BM000953941!, F
image!], FI barcode FI004721 [online image!], G-DC barcode V0063998F!, US barcode 00120093!, VEN bar-
barcode G00319183!, K barcode K000329067!, P bar- code 31744 [online image!]).
codes P00708820! & P00708821!, herbarium unknown
[F negative F0BN032311!]). 27. Tibouchina spruceana Cogn. in Martius, Fl. Bras. 14(3):
376, t. 86, fig. 2. 1885 – Lectotype (designated here):
21. Tibouchina nigricans Cogn. ex P.J.F.Guim., A.L.F. BRAZIL. Vicinibus Barra prov. Rio negro, Campo de Jana-
Oliveira & R.Romero in Syst. Bot. 40(4): 1005– narí, Jan 1851, Spruce, R. 1279 (K barcode K000329065!;
1010, fig. 1G–I, 4F–H, 5. 2015 – Holotype: BRAZIL. isolectotypes: B [probably destroyed; F negative
Goiás: Cocalzinho de Goiás, Serra dos Pireneus, F0BN016796!], E barcode E00285783 [online image!], G-
13 Dec 2012, Versiane, A.F.A. & Pacheco, R.A. 457 DC barcode G00319104!, LE!, NY barcode 00245808!,
(HUFU!; isotypes: K! barcode K000329067!, RB!, UFG!). TCD barcode TCD0005085 [online image!]).
Other syntypes: BRAZIL. Maypurés, prov. Alto Amazo-
22. Tibouchina papyrus (Pohl) Toledo in Arq. Bot. Estado nas, Jun 1854, Spruce, R. 3649 (BR barcode BR000
São Paulo 3: 30. 1952 ≡ Lasiandra papyrus Pohl, Reise 0005225279!, K barcode K000329066!, P barcode
Brasil. 1: 397. 1832 – Lectotype (designated here): P00708777!).
BRAZIL. Goiás: Aldeya São José de Mossamedes, in
montibus Serra d’Ourada, s.d., Pohl, J.B.E. 1273 28. Tibouchina steyermarkii Wurdack in Mem. New York
(BM barcode BM000953967!; isolectotypes: K barcode Bot. Gard. 23: 873, fig. 11. 1972 – Holotype:

59
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

VENEZUELA. Bolívar: vicinity of campsite, Cerro Jáua, 2. Tibouchina bicolor (Naudin) Cogn. in Candolle & Can-
cumbre de la porción Central-Occidental de la Meseta de dolle, Monogr. Phan. 7: 265. 1891 ≡ Chaetogastra
Jáua, 04 45′N, 64 26′W, 36 millas nauticas o 60 km nor- bicolor Naudin in Ann. Sci. Nat., Bot., sér. 3, 14(2):
oeste de la misión de Campamento Sanidad del Río 128, t. 4., fig. 3. 1850 – Holotype: BOLIVIA. s.d.,
Canaracuni, alt. 1922–2100 m, 22–27 Mar 1967, Steyer- d’Orbigny 418 (P barcode P00116967!).
mark, J.A. 97854 (US barcode 00120098!; isotypes: K
barcode K000329172!, P barcode P00708794!, VEN bar- 3. Tibouchina brevisepala Cogn. in Bot. Jahrb. Syst. 42: 135.
code 71154 [online image!]). 1908 – Lectotype (designated by Burke & Michelangeli
in Phytotaxa 125(1): 12. 2013): PERU. Ad viam a Cuzco
29. Tibouchina striphnocalyx (DC.) Pittier in Pittier & al., ad Sta. Anna, infra Yanamanche, locis apertis frutice-
Cat. Fl. Venez. 2: 249. 1947 ≡ Osbeckia striphnocalyx torum, 3100–3200 m a.s.l., 17 Jun 1905, Weberbauer, A.
DC., Prodr. 3: 140. 1828 ≡ Chaetogastra striphnocalyx 4965 (BR barcode BR0000005225118 [online image!];
(DC.) Mart., Nov. Gen. Sp. Pl. 3(2): 101, t. 246. 1831≡ isolectotypes: B [probably destroyed; F negative F0BN01
Pleroma striphnocalyx (DC.) Triana in Trans. Linn. 6730!], G barcode G00368030 [online image!]).
Soc. London, Bot. 28(1): 41. 1872 (“1871”) ≡ Pterolepis
striphnocalyx (DC.) Cogn. in Martius, Fl. Bras. 14(3): 4. Tibouchina calycina Cogn. in Bot. Jahrb. Syst. 42: 135.
286–287. 1885 ≡ Tibouchina striphnocalyx (DC.) Glea- 1908 – Lectotype (designated by Burke & Michelangeli
son in Phytologia 3: 241. 1950, nom. illeg. superfl. – in Phytotaxa 125(1): 12. 2013): PERU. Sandia: in fruti-
Holotype: BRAZIL. Amazonas: Rio Negro, 1827, Mar- ceto, ca. 2400 m a.s.l., 3 Apr 1902, Weberbauer, A. 650
tius, C.F.P. von s.n. (M barcode M-0165863!; isotype: (BR barcode BR0000005224807 [online image!]; isolec-
G-DC barcode G00310455!, herbarium unknown totype: B [probably destroyed; F negative F0BN
[F negative F0BN06345!]). 016731!]).
= Pterolepis striphnocalyx var. grandifolia Cogn. in Martius,
Fl. Bras. 14(3): 287. 1885 – Holotype: VENEZUELA. 4a. Tibouchina calycina var. parvifolia Cogn. in Bot. Jahrb.
Amazonas: prope San Carlos ad Rio Negro, 1853–4, Syst. 42: 135. 1908 – Lectotype (designated by Burke
Spruce, R. 3132 (BR barcode BR0000005225354!; & Michelangeli in Phytotaxa 125(1): 12. 2013): PERU.
isotypes: BM, BR barcode BR0000005225996 [frag- Dep. Cuzco: prov. Convencion, supra Hacienda Idma
ment]!, E barcode E00531245 [online image!], G bar- apud Sta. Anna, in fruticeto fruticibus et hic illic arbo-
codes G00368027! & G00368028!, G-DC barcode ribus parvis foliis induratis instructis composito,
G00319181 [online image!], GH barcode 00073164 2300–2400 m a.s.l., 25 Jun 1905, Weberbauer, A. 5008
[online image!], GOET barcode GOET008070 [online (BR barcode BR0000005224470 [online image!]).
image!], K barcodes K000006140! & K000006139!,
LE, NY barcode 00245700!, P barcode P00708785!, 5. Tibouchina octopetala Cogn. ex Britton in Bull. Torrey
S No. S-R-5384 [online image!], TCD barcode Bot. Club 17(3): 57. 1890 – Holotype: BOLIVIA. Near
TCD0000757 [online image!], W). Yungas, 4000 ft., 1885, Rusby, H.H. 2332 (NY barcode
= Tibouchina yavitensis Pittier in Bol. Soc. Venez. Ci. Nat. 00169345!; isotype: BR barcode BR0000005527489
11: 19. 1947 – Holotype: VENEZUELA. Amazonas: [fragment] [online image!]).
Yavita, 129 m, Alto Orinoco, 22 Jan 1942, Williams,
L.W. 13891a (VEN barcode 26267 [online image!]).
■ AUTHOR CONTRIBUTIONS
30. Tibouchina verticillaris Cogn. in Martius, Fl. Bras. 14(3):
379–380, t. 89. 1885 – Lectotype (designated here): PJFG and FAM designed the research. All authors collected
BRAZIL. Goiás: in campis siccis glascosis ad S. Izidro, samples in the field and conducted laboratory work. FAM and KS
performed data analysis with input from PJFG. PJFG, FAM and KS
Villa Boa et Coralinho, s.d., Pohl, J.B.E. 1192 integrated and interpreted the results. PJFG and FAM wrote the
(NY barcode 00581287!; isolectotypes: F barcode paper. — PJFG, https://orcid.org/0000-0001-5266-4552; FAM,
V0064004F [fragment]!, K barcode K000329070!). https://orcid.org/0000-0001-7348-143X; KS, https://orcid.org/0000-
0001-8780-0120; JRdSG, https://orcid.org/0000-0002-9384-2455
Incertae sedis

1. Arthrostemma cubense A.Rich., Hist. Phys. Cuba, ■ ACKNOWLEDGMENTS


Pl. Vasc.: 550. 1846 (“1845”) ≡ Chaetogastra cubensis
(A.Rich.) Griseb., Cat. Pl. Cub.: 103. 1866 ≡ Tibouchina The authors thank curators and staff for assistance during visits to
cubensis (A.Rich.) M.Gómez in Anales Soc. Esp. Hist. ALCB, BHCB, BM, CEPEC, CVRD, EFC, G, G-DC, HUEFS,
INPA, K, LE, M, MBML, P, NY, R, RB, SJRP, SP, SPF, UB, UEC
Nat. 23: 68. 1894 – Holotype: CUBA. Macurigy et in and US and also for specimen loans from B, BR, C, F, FLOR, FUEL,
insula Pinorum, s.d., De La Sagra, R. s.n. (P barcode HRB, HRCB, IAC, ICN, IPA, MO, S, SPF TEPB, UFG, VIC and
P00507408!). W. The Brazilian conservation body ICMBio is thanked for granting

60
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

the scientific collection permits. We wish to thank M. Alford, brasiliensis, vol. 14(4). Lipsiae [Leipzig]: apud Frid. Fleischer,
F. Almeda, J.F. Baumgratz, I. Capote, R. Goldenberg, M. Hopkins, 1886–1888. https://doi.org/10.5962/bhl.title.454
W. Judd, L. Kelly, L. Kollmann, A. Lobão, C.V. Martin, H. Mendoza, Cogniaux, A.C. 1891. Melastomaceae. Pp. 1–1256 in: Candolle, A. de
R. Naczi, M. Nee, K. Redden, A.K dos Santos, K. Steiner and & Candolle, C. de (eds.), Monographiae phanerogamarum, vol. 7.
K. Wurdack for providing leaf and/or herbarium material. This research Parisiis [Paris]: sumptibus G. Masson. https://doi.org/10.5962/
was supported by grant from CAPES and CNPq, Brazil to PJFG bhl.title.45961
(Scholarship Process Number BEX 2128/09-8; CNPq Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. 2012. jModel-
563541/2010-5). Partial support was also provided by the Cullman Pro- Test 2: More models, new heuristics and parallel computing.
gram for Molecular Systematics and the structural botany laboratory at Nature, Meth. 9: 772. https://doi.org/10.1038/nmeth.2109
The New York Botanical Garden and by the National Science Founda- Desrousseaux, M. 1797. Melastome. Pp. 31–57 in: Lamarck, B. de
tion (DEB-1343612) to FAM. (ed.), Encyclopédie méthodique: Botanique, vol. 4. Paris: chez
H. Agasse. https://doi.org/10.5962/bhl.title.824
Don, D. 1823. An illustration of the natural family of plants called Melas-
tomataceae. Mem. Wern. Nat. Hist. Soc. 4(2): 276–329.
■ LITERATURE CITED Drummond, A.J. & Bouckaert, R.R. 2014. Bayesian evolutionary
analysis with BEAST 2. Cambridge University Press. http://
Almeda, F. 1974. Monochaetum talamancense (Melastomataceae): A beast2.cs.auckland.ac.nz/
new species endemic to Costa Rica. Brittonia 26: 70–73. https:// Fraga, C.N. & Guimarães, P.J.F. 2014. Two new species of Pleroma
doi.org/10.2307/2805922 (Melastomataceae) from Espírito Santo, Brazil. Phytotaxa 166:
Almeda, F. 1977. Systematics of the Neotropical genus Centradenia 77–84. https://doi.org/10.11646/phytotaxa.166.1.5
(Melastomataceae). J. Arnold Arbor. 58: 73–108. https://doi.org/ Fritsch, P.W., Almeda, F., Renner, S.S., Martins, A.B & Cruz, B.C.
10.5962/bhl.part.29236 2004. Phylogeny and circumscription of the near-endemic Brazil-
Almeda, F. 2009. Melastomataceae. Pp. 164–337 in: Davidse, G., ian tribe Microlicieae (Melastomataceae). Amer. J. Bot. 91:
Sousa-Sânchez, M., Knapp, S. & Chiang, F. (eds.), Flora Meso- 1105–1114. https://doi.org/10.3732/ajb.91.7.1105
americana, vol. 4(1). México: Universidad Nacional Autónoma Gardner, G. 1843. Contributions toward a flora of Brazil. Part II. Plants
de México. from the Organ Mountains. London J. Bot. 2: 239–355.
Almeda, F. & Whiffin, T. 1981 [“1980”]. Pilocosta, a new genus of Goldenberg, R. & Kollmann, L.J.C. 2016. Two new species of Ple-
tropical American Melastomataceae. Syst. Bot. 5: 294–311. roma (Melastomataceae) from Espírito Santo, Brazil. Brittonia
https://doi.org/10.2307/2418374 68: 37–45. https://doi.org/10.1007/s12228-015-9390-4
Almeda, F., Michelangeli, F.A. & Viana, P.L. 2016. Brasilianthus Goldenberg, R., Penneys, D.S., Almeda, F., Judd, W.S. &
(Melastomataceae), a new monotypic genus endemic to ironstone Michelangeli, F.A. 2008 Phylogeny of Miconia (Melastoma-
outcrops in the Brazilian Amazon. Phytotaxa 273: 269–282. taceae): Patterns of stamen diversification in a megadiverse neo-
https://doi.org/10.11646/phytotaxa.273.4.3 tropical genus. Int. J. Pl. Sci. 24: 315–327.
Aublet, J.B.C.F. 1775. Histoire des plantes de la Guiane françoise, Goldenberg, R., Fraga, C.N., Fontana, A.P. & Nicolas, A.N. 2012.
vol 1. London & Paris: Pierre-François Didot. https://doi.org/10. Taxonomy and phylogeny of Merianthera (Melastomataceae).
5962/bhl.title.64646 Taxon 61: 1040–1056. https://doi.org/10.1002/tax.615010
Baillon, H.E. (ed.) 1877. Mélastomacées. Pp. 1–65 in: Histoire des Goldenberg, R., Almeda, F., Sosa, K., Ribeiro, R.C. & Miche-
plantes, vol. 7. Paris: Librairie Hachette, 1880. langeli, F.A. 2015. Rupestrea: A new Brazilian genus of Melasto-
Bécquer-Granados, E.R., Neubig, K.M., Judd, W.S., Miche- mataceae, with anomalous seeds and dry indehiscent fruits. Syst.
langeli, F.A., Abbott, J.R. & Penneys, D.S. 2008. Preliminary Bot. 40: 561–571. https://doi.org/10.1600/036364415X688862
molecular phylogenetic studies in Pachyanthus (Miconieae, Mel- Goyena, M.R. 1909. Flora nicaragüense, vol. 1. Managua: Compañia
astomataceae). Bot. Rev. (Lancaster) 74: 37–52. https://doi.org/ tipográfica internacional.
10.1007/s12229-008-9010-z Guimarães, P.J.F. 1992. Tibouchina sect. Pleroma (D. Don) Cogniaux
Brade, A.C. 1938. Melastomataceae Novae II. Arq. Inst. Biol. Veg. 4 no estado de São Paulo. M.Sc. thesis, Universidade Estadual de
(1): 71–84. Campinas, Campinas, Brazil.
Brade, A.C. 1959–1961. Melastomatáceas novas do Estado da Bahia. Guimarães, P.J.F. 1997. Estudos taxonômicos de Tibouchina sect.
Arch. Jard. Bot. Rio de Janeiro 17: 43–50. Pleroma (D. Don) Cogniaux (Melastomataceae). Ph.D. thesis,
Brade, A.C. & Markgraf, F. 1961. Neue Melastomataceen aus Brasi- Universidade Estadual de Campinas, Campinas, Brazil. https://
lien. Willdenowia 2(5): 769–774. doi.org/10.1590/S0100-84041997000100002
Candolle, A.P. de 1828a. Prodromus systematis naturalis regni veget- Guimarães, P.J.F. & Freitas, J.G. 2015. Two new species of Pleroma
abilis, vol. 3. Parisiis [Paris]: sumptibus sociorum Treuttel & (Melastomataceae) from Brazil. Syst. Bot. 40: 553–560. https://
Würtz. https://doi.org/10.5962/bhl.title.286 doi.org/10.1600/036364415X688736
Candolle, A.P. de 1828b. Mémoires sur la famille des Mélastomacées. Guimarães, P.J.F. & Oliveira, C.M.S. 2009. Tibouchina Aubl.
Paris: chez Treuttel & Würtz. http://books.google.com/books?id= Pp. 127–149 in: Martins, S.E., Wanderley, M.G.L., Sheperd,
hz8AAAAAQAAJ G.J., Giulietti, A.M. & Melhem, T.S. (eds.), Flora fanerogâmica
Chamisso, A. 1835 [“1834”]. De plantis in expeditione speculatoria do estado de São Paulo, vol. 6. São Paulo: FAPESP.
Romanzoffiana et in herbariis Regiis Berolinensibus observatis – Guimarães, P.J.F. & Oliveira da Silva, M.F. 2015. A new species of
Melastomaceae Americanae. Linnaea 9: 368–402. Pleroma (Melastomataceae, Melastomeae) from southeastern Brazil.
Clausing, G. & Renner, S.S. 2001. Molecular phylogenetics of Mela- Phytotaxa 205: 51–58. https://doi.org/10.11646/phytotaxa.205.1.4
stomataceae and Memecylaceae: Implications for character evolu- Guimarães, P.J.F. & Woodgyer, E.M. 2016. A new and unusual
tion. Amer. J. Bot. 88: 486–498. https://doi.org/10.2307/2657114 amphistomatic species of Tibouchina (Melastomeae, Melastoma-
Cogniaux, A.C. 1885. Melastomataceae. Tribus II. Tibouchineae. taceae) from Brazil. Phytotaxa 246: 145–151. https://doi.org/10.
Pp. 205–480 in: Martius, C.F.P. de & Eichler A.G. (eds.), Flora 11646/phytotaxa.246.2.6
brasiliensis, vol. 14(3). Lipsiae [Leipzig]: apud Frid. Fleischer, Ionta, M.G., Judd, W.S., Williams, N.H. & Whitten, W.M. 2007.
1883–1885. https://doi.org/10.5962/bhl.title.454 Phylogenetic relationships in Rhexia (Melastomataceae): Evi-
Cogniaux, A.C. 1888. Melastomataceae. Addenda et Emendanda. dence from DNA sequence data and morphology. Int. J. Pl. Sci.
Pp. 596–604 in: Martius, C.F.P. de & Eichler A.G. (eds.), Flora 168: 1055–1066. https://doi.org/10.1086/518837

61
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

Jørgensen, P.M., Nee, M. & Beck, S.G. (eds.) 2014. Catálogo de las (Melastomataceae) in the Caribbean. Bot. Rev. (Lancaster) 74:
plantas vasculares de Bolivia. Monographs in Systematic Botany 53–77. https://doi.org/10.1007/s12229-008-9004-x
127. St. Louis: Missouri Botanical Garden Press. Michelangeli, F.A., Guimarães, P.J.F., Penneys, D.S., Almeda, F. &
Jussieu, A.L. de 1789. Genera plantarum. Parisiis [Paris]: apud Kriebel, R. 2013. Phylogenetic relationships and distribution of
viduam Herissant et Theophilum Barrois. https://doi.org/10. new world Melastomeae (Melastomataceae). Bot. J. Linn. Soc.
5962/bhl.title.284 171: 38–60. https://doi.org/10.1111/j.1095-8339.2012.01295.x
Katoh S. 2013. MAFFT multiple sequence alignment software version Michelangeli, F.A., Ulloa, C.U. & Sosa, K. 2014. Quipuanthus, a new
7: improvements in performance and usability. Molec. Biol. Evol. genus of Melastomataceae from the foothills of the Andes in
30: 772–780. Ecuador and Peru. Syst. Bot. 39: 533–540. https://doi.org/10.
Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., 1600/036364414X680924
Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Miller, M.A., Pfeiffer, W. & Schwartz, T. 2010. Creating the CIPRES
Thierer, T., Ashton, B., Meintjes, P. & Drummond, A. 2012. Science Gateway for inference of large phylogenetic trees.
Geneious Basic: An integrated and extendable desktop software Pp. 45–52 in: Proceedings of the Gateway Computing Environ-
platform for the organization and analysis of sequence data. Bioin- ments Workshop (GCE), New Orleans, Louisiana, 14 Nov 2010.
formatics 28: 1647–1649. Piscataway: IEEE. https://doi.org/10.1109/GCE.2010.5676129
Krasser, F. 1893. Melastomataceae. Pp. 130–199 in: Engler, A. & Miquel, F.A.G. 1840. Commentarii phytographici, quibus varia rei
Prantl, K. (eds.), Die natürlichen Pflanzenfamilien, vol. 3(7). herbariae capita illustrantur. Lugduni Batavorum [Leiden]: apud
Leipzig: Engelmann. S. et J. Luchtmans. https://doi.org/10.5962/bhl.title.13717
Kriebel, R. 2016. Phylogenetic placement of the monotypic genus Miquel, F.A.G. 1849. Melastomataceae. Pp. 537–545 in: Plantae
Schwackaea (Melastomeae: Melastomataceae) and the evolution Regnellianae. Linnaea 22: 511–583.
of its unique fruit. Int. J. Pl. Sci. 177(5): 440–448. https://doi. Naudin, C.V. 1849. Melastomacearum monographicae descriptiones.
org/10.1086/685689 Ann. Sci. Nat., Bot., sér. 3, 13: 25–39, 126–159.
Kriebel, R., Michelangeli, F.A. & Kelly, L.M. 2015. Discovery of Naudin, C.V. 1850. Melastomacearum monographicae descriptiones.
unusual anatomical and continuous characters in the evolutionary Ann. Sci. Nat., Bot., sér. 3, 14: 53–76, 118–165.
history of Conostegia (Miconieae: Melastomataceae). Molec. Oliveira, A.L.F., Romero, R. & Guimarães, P.J.F. 2014. A new Bra-
Phylogen. Evol. 82: 289–313. https://doi.org/10.1016/j.ympev. zilian species and some synonyms in Pleroma (Melastomataceae).
2014.09.021 Brittonia 66: 353–357. https://doi.org/10.1007/s12228-014-
Majure, L.C., Neubig, K.M., Skean, J.D., Jr., Bécquer, E.R. & 9345-1
Judd, W.S. 2015. Evolution of the Sandpaper clade (Miconia, Oliveira da Silva, M.F., Guimarães, P.J.F. & Michelangeli, F.A.
Miconieae, Melastomataceae). Int. J. Pl. Sci. 176: 607–626. 2014. Nomenclatural and taxonomic novelties in the tribe Mela-
https://doi.org/10.1086/682148 stomeae (Melastomataceae). Phytotaxa 186: 222–228. https://
Markgraf, F. 1927. Plantae Luetzelburgianae Brasilienses VII. doi.org/10.11646/phytotaxa.186.4.6
Notizbl. Bot. Gart. Berlin-Dahlem 10(91): 43–54. https://doi.org/ Penneys, D.S. & Judd, W.S. 2013. Combined molecular and morpho-
10.2307/3994816 logical phylogenetic analyses of the Blakeeae (Melastomataceae).
Martin, C.V., Little, D.P., Goldenberg, R. & Michelangeli, F.A. Int. J. Pl. Sci. 174: 802–817. https://doi.org/10.1086/670011
2008. A phylogenetic evaluation of Leandra (Miconieae, Melasto- Pereira, E. 1959–1961. Contribuição ao conhecimento das Melasto-
mataceae): A polyphyletic genus where the seeds tell the story, not mataceae brasileiras. Arch. Jard. Bot. Rio de Janeiro 17: 125–169.
the petals. Cladistics 24: 315–327. https://doi.org/10.1111/j.1096- Rambaut, A. & Drummond, A.J. 2018a. LogCombiner, version
0031.2007.00185.x 2.5.0. Part of BEAST 2 package, available at http://beast2.cs.
Martius, C.F.P. von 1829. Nova genera et species plantarum brasilien- auckland.ac.nz/
sium, vol. 3. Monachii [Munich]: typis Caroli Wolf. https://doi. Rambaut, A. & Drummond, A.J. 2018b. Tree Annotator, version
org/10.5962/bhl.title.450 2.5.0. Part of BEAST 2 package, available at http://beast2.cs.
Meyer, F.S. 2016. Estudos sistemáticos no clado de Chaetogastra auckland.ac.nz/
DC. e gêneros aliados (Melastomataceae: Melastomeae). Ph.D. Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. 2014.
thesis, UNICAMP, Campinas, Brazil. Tracer, version 1.6. Institute of Evolutionary Biology, University
Meyer, F.S. & Goldenberg, R. 2014. Two new species of Pleroma of Edinburgh. Available at http://beast.bio.ed.ac.uk/
(Melastomataceae: Melastomeae) from Brazil. Kew Bull. 69: Rambo, B. 1958. Geografía das Melastomatáceas Riograndenses. Sel-
9527. https://doi.org/10.1007/S12225-014-9527-8 lowia 10(9): 147–167.
Meyer, F.S. & Goldenberg, R. 2016. Four new species of Chaetogastra Reginato, M. & Michelangeli, F.A. 2016. Untangling the phylogeny
(Melastomeae, Melastomataceae) from southern Brazil. Phytotaxa of Leandra s.str. (Melastomataceae, Miconieae). Molec. Phylogen.
282: 239–258. https://doi.org/10.11646/phytotaxa.282.4.1 Evol. 96: 17–32. https://doi.org/10.1016/j.ympev.2015.11.015
Meyer, F.S. & Matos, F.B. 2017. The recognition of Tibouchina seto- Reginato, M., Michelangeli, F.A. & Goldenberg, R. 2010. Phylogeny
sociliata Cogn. (Melastomataceae, Melastomeae) and its transfer of Pleiochiton A. Gray (Melastomataceae, Miconieae): Total evi-
to Pleroma D. Don. Phytotaxa 305: 201–208. https://doi.org/10. dence. Bot. J. Linn. Soc. 162: 423–434. https://doi.org/10.1111/j.
11646/phytotaxa.305.3.7 1095-8339.2009.01022.x
Meyer, F.S., Goldenberg, R. & Kollmann, L.J.C. 2016. Three new Renner, S.S. 1989. Systematic studies in the Melastomataceae: Bel-
species of Pleroma (Melastomataceae) from Inselbergs of Espírito lucia, Loreya, and Macairea. Mem. New York Bot. Gard. 50:
Santo, Brazil. Phytotaxa 282: 197–210. https://doi.org/10.11646/ 1–112.
phytotaxa.282.3.3 Renner, S.S. 1993. Phylogeny and classification of the Melastomata-
Michelangeli, F.A., Penneys, D.S, Giza, J., Soltis, D., Hils, M.H. & ceae and Memecylaceae. Nordic J. Bot 13: 519–540. https://doi.
Skean, J.D. 2004. A preliminary phylogeny of the tribe Mico- org/10.1111/j.1756-1051.1993.tb00096.x
nieae (Melastomataceae) based on nrITS sequence data and its Renner, S.S. 1994. Revisions of Pterogastra and Schwackaea (Me-
implications on inflorescence position. Taxon 53: 279–290. lastomataceae: Melastomeae). Nordic J. Bot. 14: 65–71. https://
https://doi.org/10.2307/4135608 doi.org/10.1111/j.1756-1051.1994.tb00574.x
Michelangeli, F.A., Judd, W.S., Penneys, D.S., Dan Skean, J., Jr., Rocha, M.J.R., Batista, J.A.N., Guimarães, P.J.F. & Miche-
Bécquer-Granados, E.R., Goldenberg, R. & Martin, C.V. 2008. langeli, F.A. 2016. Phylogenetic relationships in the Marcetia alli-
Multiple events of dispersal and radiation of the tribe Miconieae ance (Melastomeae, Melastomataceae) and implications for

62
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

generic circumscription. Bot. J. Linn. Soc. 181: 585–609. https:// Triana, J.J. 1872 [“1871”]. Les Mélastomacées. Trans. Linn. Soc.
doi.org/10.1111/boj.12429 London, Bot. 28: 1–188.
Rocha, M.J.R., Guimarães, P.J.F., Michelangeli, F.A. & Batista, Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W.,
J.A.N. 2018. Taxonomy of Marcetieae: A new neotropical tribe Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.-H.,
of Melastomataceae. Int. J. Pl. Sci. 179: 50–74. https://doi.org/ Li, D.-Z., Marhold, K., May, T.W., McNeill, J., Monro, A.M.,
10.1086/694932 Prado, J., Price, M.J. & Smith, G.F. (eds.) 2018. International
Romero, R. & Martins, A.B. 2003. Four new species of Svitramia Code of Nomenclature for algae, fungi, and plants (Shenzhen
Cham. (Melastomataceae, Melastomeae) from Minas Gerais, Bra- Code) adopted by the Nineteenth International Botanical Congress
zil. Kew Bull. 58: 403–413. https://doi.org/10.2307/4120623 Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashütten:
Romero, R. & Versiane, A.F.A. 2014. Taxonomic novelty and typifi- Koeltz Botanical Books. https://doi.org/10.12705/Code.2018
cations in Microlepis (Melastomataceae). Novon 23: 217–223. Ule, E.H.G. 1895. Ueber die Blütheneinrichtungen von Purpurella
https://doi.org/10.3417/2011077 cleistoflora, einer neuen Melastomacee. Ber. Deutsch. Bot. Ges.
Scopoli, J.A. 1777. Introductio ad historiam naturalem. Pragae [Prague]: 13: 415–420, pl. 32.
apud Wolfgangum Gerle. https://doi.org/10.5962/bhl.title.10827 Ule, E.H.G. 1896a. Berichtigung. Ber. Deutsch. Bot. Ges. 14: 163.
Shaw, J., Lickey, E.B., Beck, J.T., Farmer, S.B., Liu, W., Miller, J., Ule, E.H.G. 1896b. Weiteres zur Blütheneinrichtung von Purpurella
Siripun, K.C., Winder, C.T., Schilling, E.E. & Small, R.L. cleistopetala und Verwandten. Ber. Deutsch. Bot. Ges. 14:
2005. The tortoise and the hare II: Relative utility of 21 noncoding 169–178, pl. 13, fig. 1–6.
chloroplast DNA sequences for phylogenetic analysis. Amer. Ule, E.H.G. 1908. Beiträge zur Flora von Bahia. I. - 9. Melastomata-
J. Bot. 92: 142–166. ceae. Bot. Jahrb. Syst. 42: 232–236.
Souza, M.L.D.R. 1984. Estudos taxonômicos do gênero Tibouchina Ventenat, E.P. 1807. Notice sur les plantes qui seront publiées dans les
Aubl. (Melastomataceae) no Rio Grande do Sul. M.Sc. thesis, cinq dernières livraisons de l’ouvrage intitulé Choix de plantes.
Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil. Mém. Cl. Sci. Math. Inst. Natl. France 8: 1–20.
https://doi.org/10.3732/ajb.92.1.142 Veranso-Libalah, M.C., Stone, R.D., Fongod, A.G.N., Couvreur,
Souza, M.L.D.R. 1986. A taxonomic study of the genus Tibouchina T.L.P. & Kadereit, G. 2017. Phylogeny and systematics of Afri-
Aubl. (Melastomataceae) in Rio Grande do Sul, Brazil. Insula can Melastomateae (Melastomataceae). Taxon 66: 584–614.
16: 3–109. https://doi.org/10.12705/663.5
Stamatakis, A. 2014. RAxML version 8: A tool for phylogenetic anal- Whiffin, T. & Tomb, A.S. 1972. The systematic significance of seed
ysis and post-analysis of large phylogenies. Bioinformatics 30: morphology in the neotropical capsular-fruited Melastomataceae.
1312–1313. https://doi.org/10.1093/bioinformatics/btu033 Amer. J. Bot. 59: 411–422. https://doi.org/10.1002/j.1537-2197.
Stone, R.D. & Andreasen, K. 2010. The Afro-Madagascan genus 1972.tb10112.x
Warneckea (Melastomataceae): Molecular systematics and Wurdack, J.J. 1953. A revision of the genus Brachyotum
revised infrageneric classification. Taxon 59: 83–92. https://doi. (Tibouchineae - Melastomataceae). Mem. New York Bot. Gard.
org/10.1002/tax.591009 8: 343–407.
Todzia, C.A. & Almeda, F. 1991. A revision of Tibouchina Wurdack, J.J. 1962. Melastomataceae of Santa Catarina. Sellowia 14:
section Lepidotae (Melastomataceae: Tibouchineae). Proc. Calif. 109–217.
Acad. Sci. 47: 175–206. Wurdack, J.J. 1967. Certamen Melastomataceae XI. Phytologia 14:
Triana, J.J. 1865. Dispositio Melastomacearum. Bull. Congr. Int. Bot. 257–274.
Hort. Amsterdam, 1865: 457–461. https://doi.org/10.1111/j.1096- Wurdack, J.J. 1973. Uma nova Melastomataceae de Minas Gerais.
3642.1871.tb00222.x Bol. Mus. Bot. Munic. 10: 1–3.

Appendix 1. Voucher information.


Species name, collector and number (herbarium code), GenBank accessions numbers for nrITS, accD-psaI, psbK-psbL. Missing sequence data is indicated by a
dash (–). DNA sequences newly generated for this study are marked with an asterisk (*).
Acanthella sprucei Hook.f., Brazil, Diaz, W., 4538 (NY), JQ730036, JQ730247, JQ730456; Aciotis acuminifolia (Mart. ex DC.) Triana, French Guiana, Martin,
C.V., 487 (NY), JQ730037, JQ730248, JQ730457; Aciotis annua (Mart. ex DC.) Triana, Brazil, Rocha, M.J.R., 304 (BH), KU501052, KU501161, –; Aciotis
circaeifolia (Bonpl.) Triana, Brazil, Caddah, M.K., 621 (NY), JQ730038, JQ730249, JQ730458; Aciotis indecora1 (Bonpl.) Triana, Guyana, Michelangeli,
F.A., 2456 (NY), KU501056*, KU501163*, –; Aciotis indecora2 (Bonpl.) Triana, French Guyana, Martin, C.V., 411 (NY), JQ730039, JQ730250,
JQ730459; Aciotis paludosa (Mart. ex DC.) Triana, Brazil, Guimarães, P.J.F., 317 (RB), JQ730040, JQ730251, JQ730460; Aciotis polystachya1 (Bonpl.) Tri-
ana, Brazil, Caddah, M.K., 581 (INPA), –, KF407949*, MH463415*; Aciotis polystachya2 (Bonpl.) Triana, Brazil, Rocha, M.J.R., 858 (BHCB), KU501053, –,
–; Aciotis purpurascens1 (Aublet) Triana, Guyana, Michelangeli, F.A., 2454 (NY), KU501055*, KU501164*, –; Aciotis purpurascens2 (Aublet) Triana, French
Guiana, Martin, C.V., 422 (NY), JQ730041, JQ730252, JQ730461; Aciotis rubricaulis1 (Mart. ex DC.) Triana, Brazil, Goldenberg, R., 850 (NY), JQ730042,
JQ730253, JQ730462; Aciotis rubricaulis2 (Mart. ex DC.) Triana, Guyana, Michelangeli, F.A., 2452 (NY), KU501057*, KU501165*, –; Acisanthera alsinae-
folia (Mart. & Schrank ex DC.) Triana, Brazil, Goldenberg, R., 826 (NY), JQ730043, JQ730254, JQ730463; Acisanthera ayangannae (Wurdack) M.J.Rocha
& P.J.F.Guim., Guyana, Hoffman, B., 2943 (US), KU501063*, KU501171*, –; Acisanthera erecta J.St.-Hil., Venezuela, Michelangeli, F.A., 826 (BH),
JQ730045, JQ730256, JQ730465; Acisanthera paraguayensis (Hook.f.) Cogn., Paraguay, Krapovickas, A., 45640 (NY), KU501058*, –, –; Acisanthera uni-
flora (Vahl) Gleason, Colombia, Pedraza, P., 2281 (NY), KU501062*, KU501169*, –; Acisanthera variabilis (Mart. ex DC.), Brazil, Irwin, H.S., 19593 (NY),
–, MH569713*, –; Amphorocalyx multiflorus Baker, Madagascar, Almeda, F., 8669 (CAS), JQ730046, JQ730257, JQ730466; Amphorocalyx rupestris H.Per-
rier, Madagascar, Almeda, F., 8723 (CAS), JQ730047, JQ730258, JQ730467; Andesanthus gleasonianus (Wurdack) P.J.F.Guim. & Michelang., Ecuador,
Jaramillo, J., 7985 (NY), JQ730189, –, JQ730609; Andesanthus inopinatus (Wurdack) P.J.F.Guim. & Michelang., Costa Rica, Kriebel, R., 5342 (NY),
JQ730196, JQ730404, JQ730616; Andesanthus lepidotus (Bonpl.) P.J.F.Guim. & Michelang., Colombia, Morales, M.E., s.n. (UPTC), JQ730201,
JQ730409, JQ730621; Andesanthus wurdackii (Almeda & Todzia) P.J.F.Guim. & Michelang., Panama, Penneys, D.S., 1749 (FLAS), JQ730240,
JQ730449, JQ730661; Antherotoma naudinii Hook.f., Madagascar, Almeda, F., 8624 (CAS), JQ730048, JQ730259, –; Antherotoma cf. phaeotricha
(Hochst.) Jacq.-Fél., South Africa, Steiner, K.E., 3561 (NBG), JQ730078, JQ730289, JQ730496; Appendicularia pullei (Gleason) M.J.R.Rocha & P.J.F.Guim.,
French Guiana, Martin, C.V., 460 (NY), JQ730081*, JQ730294*, JQ730500*; Appendicularia thymifolia DC., French Guiana, Martin, C.V., 441 (NY),

63
Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

Appendix 1. Continued.
JQ730049, JQ730260, JQ730468; Arthrostemma ciliatum1 Pav. ex D.Don, Puerto Rico, Skean, J.D., 3806 (FLAS), AY460429, –, –; Arthrostemma ciliatum2
Pav. ex D.Don, Jamaica, Judd, W.S., 8344 (FLAS), MH430227, –, –; Arthrostemma primaevum Almeda, Mexico, Penneys, D.S., 1429 (FLAS), JQ730050,
JQ730261, JQ730469; Brachyotum alpinum Cogn., Ecuador, Ulloa Ulloa, M.C., 2084 (MO), MH430228*, MH569714*, MH463416*; Brachyotum angusti-
folium Wurdack, Peru, Michelangeli, F.A., 1734 (USM), KF463029*, KF407954*, KF462993*; Brachyotum barbeyanum Cogn., Peru, Michelangeli, F.A.,
1717 (NY), MH430229*, –, MH463417*; Brachyotum benthamianum Triana, Ecuador, Alford, M., 3083 (BH), JQ730051, JQ730262, JQ730470; Brachyo-
tum cogniauxii Wurdack, Peru, Michelangeli, F.A., 1736 (USM), KF463030*, KF407955*, KF462994*; Brachyotum confertum (Bonpl.) Triana, Ecuador,
Penneys, D.S., 1603 (FLAS), JQ730052, JQ730263, JQ730471; Brachyotum coronatum (Triana) Wurdack, Peru, Michelangeli, F.A., 1687 (USM),
KF463027*, KF407952*, –; Brachyotum fictum Wurdack, Ecuador, Penneys, D.S., 1594 (FLAS), JQ730053, JQ730264, JQ730472; Brachyotum fraternum
Wurdack, Ecuador, Penneys, D.S., 1604 (FLAS), JQ730054, JQ730265, JQ730473; Brachyotum grisebachii Cogn., Peru, Michelangeli, F.A., 1856 (NY),
MH430230*, MH569715*, MH463418*; Brachyotum harlingii Wurdack, Ecuador, Penneys, D.S., 1593 (FLAS), JQ730055, JQ730266, JQ730474; Brachyo-
tum huancavelicae Wurdack, Peru, Michelangeli, F.A., 1861 (NY), MH430231*, MH569716*, MH463419*; Brachyotum incrassatum E.Cotton, Ecuador,
Clark, J.L., 8896 (NY), JQ730056, JQ730267, JQ730475; Brachyotum intermedium Wurdack, Peru, Michelangeli, F.A., 1685 (USM), KF463026*, –, –; Bra-
chyotum aff. ledifolium1 (Desr.) Triana, Peru, Michelangeli, F.A., 1886 (USM), KF463025*, KF407951*, KF462991*; Brachyotum ledifolium2 (Desr.) Triana,
Ecuador, Penneys, D.S., 1554 (FLAS), JQ730057, JQ730268, JQ730476; Brachyotum lindenii Cogn., Ecuador, Penneys, D.S., 1555 (FLAS), JQ730058,
JQ730269, JQ730477; Brachyotum aff. maximowiczii Cogn., Peru, Michelangeli, F.A., 1738 (USM), KF463032*, –, –; Brachyotum microdon (Naudin) Tri-
ana, Bolivia, Nee, M., 55327 (NY), JQ730059, JQ730270, JQ730478; Brachyotum nutans Gleason, Peru, Michelangeli, F.A., 1887 (NY), MH430232*,
MH569717*, MH463420*; Brachyotum quinquenerve (Ruiz & Pav.) Triana, Peru, Michelangeli, F.A., 1863 (NY), MH430233*, MH569718*,
MH463421*; Brachyotum radula Triana, Peru, Michelangeli, F.A., 1737 (USM), KF463031*, –, –; Brachyotum rostratum1 (Naudin) Triana, Peru, Michelan-
geli, F.A., 1691 (USM), KF463028*, KF407953*, KF462992*; Brachyotum rostratum2 (Naudin) Triana, Ecuador, Penneys, D.S., 1586 (FLAS), JQ730060,
JQ730271, JQ730479; Brachyotum sp., Peru, Michelangeli, F.A., 1633 (NY), MH430234*, MH569719*, MH463422*; Brachyotum strigosum (L.f.) Triana,
Colombia, Gonzalez, F., s.n. (COL), MH430235*, MH569720*, MH463423*; Bucquetia glutinosa (L.f.) DC., Colombia, Alvear, M., 1398 (CAS), JQ730061,
JQ730272, JQ730480; Bucquetia vernicosa Gleason, Colombia, Almeda, F., 10559 (COL, CAS), –, KF407956*, –; Cambessedesia espora (A.St.-Hil. ex
Bonpl.) DC., Brazil, Guimarães, P.J.F., 397 (RB), JQ730062, JQ730273, JQ730481; Cambessedesia hilariana (Kunth) DC., Brazil, Guimarães, P.J.F., 405
(RB), JQ730063, JQ730274, JQ730482; Castratella piloselloides (Bonpl.) Naud., Colombia, Kelly, L., s.n. (CAS), JQ730064, JQ730275, JQ730483; Centra-
denia floribunda Planchon, Guatemala, Kriebel, R., 5557 (NY), –, –, KF462995*; Centradenia grandifolia (Sehlechtd.) Endl., Costa Rica, Penneys, D.S., 1544
(FLAS), JQ730065, JQ730276, –; Centradenia inaequilateralis (Schlechtd. & Cham.) G.Don., Costa Rica, Michelangeli, F.A., 838-a (NY), JQ730066,
JQ730277, JQ730484; Chaetogastra cerastifolia (Naudin) P.J.F.Guim. & Michelang., Brazil, Varassin, I.G., 96 (UPCB), JQ730167, –, JQ730588; Chaetogas-
tra chamaecistus Sieber ex Griseb., Martinique, Penneys, D.S., 1276 (FLAS), JQ730168, JQ730378, JQ730589; Chaetogastra ciliaris (Vent.) DC., Costa Rica,
Kriebel, R., 5461 (NY), JQ730169, –, JQ730590; Chaetogastra citrina (Naudin) P.J.F.Guim. & Michelang., Bolivia, Nee, M., 55308 (NY), JQ730171,
JQ730380, JQ730592; Chaetogastra clinopodifolia DC., Brazil, Goldenberg, R., 774 (NY), JQ730174, JQ730383, JQ730594; Chaetogastra confertiflora
(Naudin) P.J.F.Guim. & Michelang., Bolivia, Nee, M., 55377 (NY), JQ730175, JQ730384, JQ730595; Chaetogastra congestiflora (Todzia) P.J.F.Guim.
& Michelang., Mexico, Rojas, J., 480 (MEXU), MH430238*, –, MH463428*; Chaetogastra cf. decora (Gleason) P.J.F.Guim. & Michelang., Peru, Michelan-
geli, F.A., 1985 (USM), KF463041*, KF407974*, KF463014*; Chaetogastra dimorphylla1 (Gleason) P.J.F.Guim. & Michelang., Peru, Michelangeli, F.A.,
1892 (NY, USM), KF463044*, MH569727*, KF463017*; Chaetogastra dimorphylla2 (Gleason) P.J.F.Guim. & Michelang., Peru, Michelangeli, F.A., 1983
(USM, NY), KF463045*, MH569728*, KF463018*; Chaetogastra echinata (Ruiz & Pav.) DC., Peru, Michelangeli, F.A., 2672 (USM), –, MH569729*,
MH463429*; Chaetogastra cf. gayana1 (Naudin) P.J.F.Guim. & Michelang., Bolivia, Nee, M., 55287 (NY), MH430237*, MH569724*, MH463427*; Chae-
togastra gayana2 (Naudin) P.J.F.Guim. & Michelang., Peru, Michelangeli, F.A., 1804 (USM), KF463049*, –, –; Chaetogastra gayana3 (Naudin) P.J.F.Guim.
& Michelang., Peru, Michelangeli, F.A., 1910 (USM), KF463038*, KF407971*, KF463013*; Chaetogastra gayana4 (Naudin) P.J.F.Guim. & Michelang.,
Bolivia, Nee, M., 55380 (NY), JQ730187, JQ730396, JQ730607; Chaetogastra geitneriana Schltdl., Venezuela, Capote, I., 852 (NY), JQ730188,
JQ730397, JQ730608; Chaetogastra gracilis (Bonpl.) DC., Brazil, Guimarães, P.J.F., 336 (RB), JQ730190, JQ730398, JQ730610; Chaetogastra grossa
(L.f.) P.J.F.Guim. & Michelang., unknown, Zabala, J.C., 1 (NY), JQ730192, JQ730400, JQ730612; Chaetogastra hieracioides Schrank & Mart. ex. DC., Bra-
zil, Almeda, F., 8380 (CAS), JQ730194, JQ730402, JQ730614; Chaetogastra inconstans (Vahl) P.J.F.Guim. & Michelang., Dominica, Penneys, D.S., 1298
(FLAS), JQ730218, JQ730426, JQ730638; Chaetogastra latibracteolata (Paul G.Wilson) P.J.F.Guim. & Michelang., Mexico, Cowan, 4810 (NY),
MH430240, MH569730, MH463430; Chaetogastra laxa (Desr.) P.J.F.Guim. & Michelang., Ecuador, Harling, G., 23570 (NY), JQ730200, JQ730408,
JQ730620; Chaetogastra longifolia (Vahl.) DC., Venezuela, Michelangeli, F.A., 816 (BH), JQ730204, JQ730412, JQ730624; Chaetogastra mariae
(Wurdack) P.J.F.Guim. & Michelang., Peru, Michelangeli, F.A., 1702 (NY), KF463046, MH569732, KF463019; Chaetogastra minor (Cogn.) P.J.F.Guim.
& Michelang., Brazil, Guimarães, P.J.F., 393 (RB), JQ730211, JQ730419, JQ730631; Chaetogastra mollis1 (Bonpl.) DC., Ecuador, Penneys, D.S., 1895
(FLAS), JQ730212, JQ730420, JQ730632; Chaetogastra aff. mollis2 (Bonpl.) DC., Peru, Michelangeli, F.A., 1881 (USM, NY), KF463050*, MH569722*,
KF463022*; Chaetogastra montana P.J.F.Guim. & Michelang., Bolivia, Nee, M., 55379 (NY), JQ730150, JQ730361, JQ730573; Chaetogastra naudiniana
Decne., Cultivated NY (NY), JQ730214, JQ730422, JQ730634; Chaetogastra pendula (Cogn.) P.J.F.Guim. & Michelang., Ecuador, Penneys, D.S., 1893
(FLAS), JQ730220, JQ730428, JQ730640; Chaetogastra pleromoides (Naudin) P.J.F.Guim. & Michelang., Peru, Michelangeli, F.A., 1859 (USM),
KF463047*, MH569733*, KF463020*; Chaetogastra saxosa (Gleason) P.J.F.Guim. & Michelang., Peru, Michelangeli, F.A., 1803 (NY), KF463048*,
MH569735*, KF463021*; Chaetogastra aff. scabriuscula (Schltdl.) P.J.F.Guim. & Michelang., Mexico, de Santiago, J.R., 1803 (MEXU), –, MH569723*,
MH463426*; Chaetogastra sebastianopolitana (Raddi) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 330 (RB), JQ730226, JQ730434, JQ730646;
Chaetogastra solmsii (Cogn.) P.J.F.Guim. & Michelang., Peru, Michelangeli, F.A., 1929 (USM), KF463042*, MH569737*, KF463015*; Chaetogastra sp.,
Peru, Michelangeli, F.A., 1935 (USM), KF463051*, MH569738*, KF463023*; Chaetogastra cf. weberbaueri (Cogn.) P.J.F.Guim. & Michelang., Peru, Miche-
langeli, F.A., 1877 (USM), KF463043*, MH569725*, KF463016*; Chaetolepis anisandra Naudin, Colombia, Almeda, F., 10152 (CAS), –, –, KF462996;
Chaetolepis cufodontisii Standl., Costa Rica, Michelangeli, F.A., 1160 (NY), JQ730067, JQ730278, JQ730485; Chaetolepis microphylla (Bonpl.) Miquel, Bra-
zil, Michelangeli, F.A., 1268 (NY), JQ730068, JQ730279, JQ730486; Chaetostoma armatum (Spreng.) Cogn., Brazil, Guimarães, P.J.F., 396 (RB), JQ730069,
JQ730280, JQ730487; Comolia leptophylla (Bonpl.) Naudin, Venezuela, Wurdack, J.J., 39943 (US), KU501072, –, –; Comolia microphylla Benth., Guyana,
Redden, K.M., 1454 (NY), JQ730070, JQ730281, JQ730488; Comolia ovalifolia (DC.) Triana, Brazil, Rocha, M.J.R., 719 (BHCB), KU501077, –, –; Comolia
smithii Wurdack, Guyana, Jasen-Jacobs, M.J., 4461 (US), KU501075*, –, –; Comolia villosa (Aubl.) Triana, Brazil, Goldenberg, R., 1621 (UPCB), –,
KF407957*, KF462997*; Comoliopsis montana (Gleason) M.J.R.Rocha & P.J.F.Guim., Venezuela, Huber, O., 8851 (US), KU501078*, –, –; Desmoscelis vil-
losa (Aubl.) Naud., Venezuela, Michelangeli, F.A., 827 (BH), JQ730073, JQ730284, JQ730491; Dichaetanthera africana Jacq.-Fél., Gabon, Smith, S., 1885
(US), JQ730074, JQ730285, JQ730492; Dichaetanthera oblongifolia Baker, Madagascar, Almeda, F., 7926a (CAS), JQ730075, JQ730286, JQ730493; Dicra-
nanthera hedyotidea C.Presl., Guyana, Wurdack, K.J., 4145 (NY), JQ730044, JQ730255, JQ730464; Dionycha bojeri Naudin, Madagascar, Almeda, F., 8626
(CAS), JQ730076, JQ730287, JQ730494; Dupineta multiflora (Sm.) Raf., Gabon, Smith, S., 1886 (US), JQ730077, JQ730288, JQ730495; Ernestia
adenotricha L. Uribe, Colombia, Uribe, L., 3662 (NY), KU501083*, KU501186*, –; Ernestia confertiflora Wurdack, French Guyana, Penneys, D.S., 1913
(FLAS), JQ730079, JQ730292, JQ730498; Ernestia maguirei Wurdack, Venezuela, Michelangeli, F.A., 707 (NY), KU501079, KU501182, –; Ernestia quad-
riseta Berg. ex Triana, Peru, Rimachi, M., 11642 (NY), –, KU501187*, –; Ernestia tenella (Bonpl.) DC., Venezuela, Michelangeli, F.A., 386 (BH), JQ730082,
JQ730295, JQ730501; Fritzschia edmundoi (Brade) M.J.R.Rocha & P.J.F.Guim., Brazil, Rocha, M.J.R., 975 (BHCB), –, KU501176, –; Fritzschia erecta
Cham., Brazil, Guimarães, P.J.F., 406 (RB), JQ730083, JQ730296, JQ730502; Fritzschia furnensis R.Romero & M.J.R.Rocha, Brazil, Rocha, M.J.R., 984

64
TAXON 00 (00) • 1–66 Guimarães & al. • Systematics of Tibouchina and allies

Appendix 1. Continued.
(BHCB), KU501069, –, –; Fritzschia lanceiflora (DC.) M.J.R.Rocha & P.J.F.Guim., Brazil, Rocha, M.J.R., 905 (BHCB), KU501068, KU501177, –; Fritzschia
rupestris R.Pacifico, Almeda & D.Nunes, Brazil, Mota, N., 2694 (BHCB), KU501070, KU501179, –; Fritzschia sertularia (Schrank & Mart. ex DC.) M.J.R.
Rocha & P.J.F.Guim., Brazil, Almeda, F., 7724 (CAS), JQ730071, JQ730282, JQ730489; Fritzschia sessilis (Spreng.) M.J.R.Rocha & P.J.F.Guim., Brazil,
Rocha, M.J.R.531 (BHCB), –, KU501178, –; Guyonia ciliata Hook.f., Cameroon, Mambo, P., 248 (NY), JQ730084, –, JQ730505; Heterocentron breedlovei
(Wurdack) P.J.F.Guim. & Michelang., Mexico, Kelly, L., 1361 (CAS), JQ730163, JQ730374, JQ730584; Heterocentron elegans (Schltdl.) Kuntze, Cultivated
NY (NY), JQ730085, JQ730299, JQ730506; Heterocentron muricatum Gleason, Mexico, Penneys, D.S., 1949 (FLAS), JQ730086, JQ730300, JQ730507; Het-
erocentron subtriplinervium (Link & Otto) A.Br. & Bouche, Mexico, Penneys, D.S., 1430 (FLAS), JQ730087, JQ730301, JQ730508; Heterotis decumbens
(P.Beauv.) Jacq.-Fél., Gabon, Smith, S., 1705 (US), JQ730088, JQ730302, JQ730509; Heterotis rotundifolia (Sm.) Jacq.-Fél., Dominica, Penneys, D.S., 1304
(FLAS), JQ730089, JQ730290, –; Lavoisiera caryophyllea A.St.-Hil. ex Naudin, Brazil, Goldenberg, R., 1570 (UPCB), –, –, KF462999; Lavoisiera conferti-
flora Naudin, Brazil, Goldenberg, R., s.n. (UPCB), –, KF407959, KF463000; Lavoisiera imbricata (Thunb.) DC., Brazil, Goldenberg, R., 820 (UPCB),
KF463034, KF407960, KF463001; Lavoisiera mucorifera Mart. & Schrank ex DC., Brazil, Guimarães, P.J.F., 345 (RB), KF463033, KF407958,
KF462998; Lavoisiera pulchella Chamisso, Brazil, Goldenberg, R., 814 (NY), JQ730093, JQ730306, JQ730513; Leiostegia vernicosa Benth., Guyana, Wur-
dack, K.J., 4181 (NY), JQ730072, JQ730283, JQ730490; Loricalepis duckei Brade, Brazil, Hopkins, M., 1904 (INPA), –, KF407961, –; Macairea lasiophylla
(Benth. Wurdack, Brazil, Rocha, M.J.R., 873 (BHCB), KU501081, –, –; Macairea multinervia Benth., Brazil, Rocha, M.J.R., 876 (BHCB), KU501082,
KU501185, –; Macairea pachyphylla Benth., Guyana, Redden, K.M., 3869 (NY), JQ730094, –, –; Macairea radula (Bonpl.) DC., Brazil, Lima, J., 715
(UPCB), JQ730095, JQ730307, JQ730514; Macairea thyrsiflora DC., Guyana, Wurdack, K.J., 4153 (NY), JQ730096, JQ730308, JQ730515; Marcetia acerosa
DC., Brazil, Santos, A.K.A., 681 (UFB), JQ730097, –, JQ730516; Marcetia alba Ule, Brazil, Goldenberg, R., 2085 (UPCB), KU501095, KU501200, –; Mar-
cetia bahiensis (Brade & Markgr.) Wurdack, Brazil, Rocha, M.J.R. 308 (BHCB), KU501086, KU501190, –; Marcetia candolleana A.K.A.Santos & A.B.Mar-
tins, Brazil, Bunger, M.O., 615 (BHCB), KU501085, KU501189, –; Marcetia canescens Naud., Brazil, Rocha, M.J.R., 295 (BHCB), KU501089, KU501193, –;
Marcetia eimeriana A.B.Martins & Woodgyer, Brazil, Santos, A.K.A., 832 (UFB), JQ730098, JQ730309, JQ730517; Marcetia ericoides (Spreng.) O.Berg ex
Cogn., Brazil, Santos, A.K.A., 532 (UFB), JQ730099, JQ730310, JQ730518; Marcetia formosa Wurdack, Brazil, Rocha, M.J.R., 345 (BHCB), KU501090,
KU501195, –; Marcetia grandiflora Markgr., Brazil, Rocha, M.J.R., 319 (BHCB), KU501091, KU501196, –; Marcetia harleyi Wurdack, Brazil, Santos,
A.K.A., 558 (UFB), JQ730100, –, JQ730519; Marcetia latifolia Naud., Brazil, Santos, A.K.A., 366 (UFB), JQ730101, –, JQ730520; Marcetia macrophylla
Wurdack, Brazil, Rocha, M.J.R., 297 (BHCB), KU501084, KU501188, –; Marcetia mucugensis Wurdack, Brazil, Rocha, M.J.R., 285 (BHCB), KU501088,
KU501192, –; Marcetia taxifolia (A.St.-Hil.) DC., Venezuela, Michelangeli, F.A., 680 (BH), JQ730102, JQ730311, JQ730521; Marcetia vellutina Markgr.,
Brazil, Rocha, M.J.R., 293 (BHCB), KU501087, KU501191, –; Marcetia viscida Wurdack, Brazil, Rocha, M.J.R., 334 (BHCB), KU501094, KU501199, –;
Melastoma denticulatum Labill., New Caledonia, Simmons, M.P., 1842 (BH), JQ730104, JQ730313, JQ730523; Melastoma dodecandrum Lour., [unknown;
from GenBank], GQ265883, –, –; Melastoma intermedium Dunn, [unknown; from GenBank], GQ265879, –, –; Melastoma malabathricum1 L., China, Pen-
neys, D.S., 1998 (CAS), JQ730105, JQ730314, JQ730524; Melastoma malabathricum2 L., [unknown; from GenBank], GQ265878, –, –; Melastoma malaba-
thricum3 L., Taiwan, Chen, C.H., 6443 (TAIF), JQ730103, JQ730312, JQ730522; Melastoma sanguineum Sims, China, Penneys, D.S., 2000 (CAS),
JQ730106, JQ730315, JQ730525; Microlicia cordata (Spreng.) Cham., Brazil, Michelangeli, F.A., 1578 (UPCB), –, KF407962, KF463002; Microlicia crenu-
lata (DC.) Mart., Brazil, Michelangeli, F.A., 1581 (UPCB), –, –, KF463003; Microlicia fulva (Spreng.) Cham., Brazil, Michelangeli, F.A., 1576 (NY),
KU501105, KU501205, MH463424; Microlicia glazioviana Cogn., Brazil, Michelangeli, F.A., 1572 (UPCB), –, KF407964, KF463005; Microlicia graveolens
DC., Brazil, Michelangeli, F.A., 1593 (UPCB), –, –, KF463006; Microlicia isophylla DC., Brazil, Michelangeli, F.A., 1564 (UPCB), –, KF407965, KF463007;
Monochaetum bonplandii (Kunth) Naudin, Venezuela, Michelangeli, F.A., 1239 (NY), JQ730108, JQ730320, JQ730530; Monochaetum discolor H.Karst. ex
Triana, Venezuela, Capote, I., 811 (NY), JQ730109, JQ730321, JQ730531; Monochaetum humboldtianum (Kunth & Bouche) Kunth ex Walpers, Venezuela,
Michelangeli, F.A., 796 (BH), JQ730110, JQ730322, JQ730532; Monochaetum lineatum (D.Don) Naudin, Peru, Michelangeli, F.A., 1700 (USM), KF463035*,
–, –; Monochaetum meridense (Klotzsch ex H.Karst.) Naudin, Venezuela, Capote, I., 862 (NY), JQ730111, JQ730323, JQ730533; Monochaetum polyneuron
Triana, Venezuela, Michelangeli, F.A., 1267 (NY), JQ730112, JQ730324, JQ730534; Monochaetum tenellum Naudin, Cultivated Strybing Arboretum,
AY460432, –, –; Monochaetum uribei Wurdack, Colombia, Mendoza, H., 17593 (FMB), JQ730113, JQ730325, JQ730535; Monochaetum vulcanicum Cogn.,
Venezuela, Michelangeli, F.A., 718 (BH), JQ730114, JQ730326, JQ730536; Nepsera aquatica (Aubl.) Naud., Porto Rico, Struwe, L., 1158 (NY), JQ730115,
JQ730327, JQ730537; Noterophila bivalvis (Aubl.) Kriebel & M.J.R.Rocha, Brazil, Rocha, M.J.R., 871 (BHCB), KU501064, KU501172, –; Noterophila cras-
sipes (Naudin) Kriebel & M.J.R.Rocha, Brazil, Rocha, M.J.R., 833 (BHCB), KU501065, –, –; Noterophila genliseoides (Hoehne) Kriebel & M.J.R.Rocha, Bra-
zil, Rocha, M.J.R., 942 (BHCB), KU501066, KU501173, –; Noterophila limnobios (DC.) Mart., Brazil, Moreira, S.N., 312 (BHCB), –, KU501174, –; Osbeckia
aspera Blume, Sri Lanka, Klackenberg, 376 (NY), JQ730117, –, –; Osbeckia australiana Naudin, Australia, Brennan, K.G., 7008 (NY), JQ730116, JQ730328,
JQ730538; Osbeckia nepalensis Hook., China, Penneys, D.S., 1986 (CAS), JQ730118, JQ730329, JQ730539; Osbeckia stellata Buch.-Ham. ex Ker Gawl.,
China, Penneys, D.S., 1969 (CAS), JQ730119, JQ730330, JQ730540; Pachyloma huberioides (Naud.) Triana, Brazil, Caddah, M.K., 558 (NY), JQ730120,
JQ730331, JQ730541; Pilocosta campanensis (Almeda & Whiffin) Almeda, Panama, Penneys, D.S., 1647 (FLAS), JQ730122, JQ730333, JQ730543; Pilocosta
nana (Standl.) Almeda & Whiffin, Ecuador, Moran, R.C., 6928 (NY), JQ730123, JQ730334, JQ730544; Pilocosta nubicola Almeda, Costa Rica, Penneys, D.S.,
1775 (FLAS), JQ730124, JQ730335, JQ730545; Pilocosta oerstedii (Triana) Almeda & Whiffin, Costa Rica, Kriebel, R., 5412 (NY), JQ730125, JQ730336,
JQ730546; Pleroma aemulum1 (Schrank & Mart. ex DC.) Triana, Brazil, Almeda, F., 8443 (CAS), JQ730238, JQ730447, JQ730659; Pleroma aemulum2
(Schrank & Mart. ex DC.) Triana, Brazil, Guimarães, P.J.F., 313 (RB), JQ730149, JQ730360, JQ730572; Pleroma angustifolium (Naudin) Triana, Brazil,
Almeda, F., 9568 (CAS), JQ730151, JQ730362, JQ730574; Pleroma arboreum Gardner, Brazil, Guimarães, P.J.F., 324 (RB), JQ730152, JQ730363,
JQ730575; Pleroma arenarium (Cogn.) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 352 (RB), JQ730153, JQ730364, JQ730576; Pleroma axillare
(Cogn.) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 338 (RB), JQ730156, JQ730367, JQ730579; Pleroma bandeirae P.J.F.Guim. & Michelang., Bra-
zil, Guimarães, P.J.F., 388 (RB), JQ730145, JQ730356, JQ730567; Pleroma barnebyanum (Wurdack) P.J.F.Guim. & Michelang., Brazil, Almeda, F., 8262
(CAS), JQ730157, JQ730368, –; Pleroma benthamianum Gardner, Brazil, Almeda, F., 9759 (CAS), JQ730158, JQ730369, JQ730580; Pleroma blanchetia-
num (Cogn.) P.J.F.Guim. & Michelang., Brazil, Santos, A.K.A., 332 (UFB), JQ730161, JQ730372, –; Pleroma boudetii (P.J.F.Guim. & R.Goldenb.) P.J.F.
Guim. & Michelang., Brazil, Kollman & Goldenberg, 8563 (NY), JQ730162, JQ730373, JQ730583; Pleroma candolleanum (Mart. ex DC.) Triana, Brazil,
Lima, J., 711 (UPCB), JQ730164, JQ730375, JQ730585; Pleroma cardinale (Bonpl.) Triana, Brazil, Guimarães, P.J.F., 407 (RB), JQ730165, JQ730376,
JQ730586; Pleroma castellense (Brade) P.J.F.Guim. & Michelang., Brazil, Goldenberg, R., 1071 (NY), JQ730166, JQ730377, JQ730587; Pleroma cinereum
(Cogn.) P.J.F.Guim. & Michelang., Brazil, Almeda, F., 9843 (CAS), JQ730170, JQ730379, JQ730591; Pleroma clavatum (Pers.) P.J.F.Guim. & Michelang.,
Brazil, Santos, A.K.A., 696 (UFB), JQ730172, JQ730381, JQ730593; Pleroma cleistoflorum (Ule) P.J.F.Guim., Oliveira da Silva & Michelang., Brazil, Baum-
gratz, J.F.A, 1138b (R), JQ730090, JQ730303, JQ730510; Pleroma clidemioides Berg. ex Triana, Brazil, Goldenberg, R., 1439 (NY), JQ730173, JQ730382, –;
Pleroma cristatum (Brade) P.J.F.Guim. & Michelang., Brazil, Baumgratz, J.F.A, 819 (R), JQ730177, JQ730386, JQ730597; Pleroma cryptadenum (Gleason)
P.J.F.Guim. & Michelang., Venezuela, Michelangeli, F.A., 708 (BH), JQ730178, JQ730387, JQ730598; Pleroma cucullatum F.S.Mey., Fraga & R.Goldenb.,
Brazil, Goldenberg, R., 1143 (NY), JQ730230, JQ730438, JQ730650; Pleroma dubium (Cham) P.J.F.Guim. & Michelang., Brazil, Varassin, I.G., 101 (UPCB),
JQ730179, JQ730388, JQ730599; Pleroma echinatum Gardner, Brazil, Guimarães, P.J.F., 305 (RB), JQ730186, JQ730395, JQ730606; Pleroma estrellense
(Raddi) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 340 (RB), JQ730180, JQ730389, JQ730600; Pleroma fissinervium (Schrank & Mart. ex DC.)
Gardner, Brazil, Guimarães, P.J.F., 351 (RB), JQ730181, JQ730390, JQ730601; Pleroma fothergillae (Schrank & Mart. ex DC.) Triana, Brazil, Guimarães,
P.J.F., 314 (RB), JQ730182, JQ730391, JQ730602; Pleroma foveolatum (Naudin) Triana, Brazil, Guimarães, P.J.F., 350 (RB), JQ730183, JQ730392,
JQ730603; Pleroma frigidulum (Schrank & Mart. ex DC.) Triana, Brazil, Guimarães, P.J.F., 394 (RB), JQ730185, JQ730394, JQ730605; Pleroma gertii

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Guimarães & al. • Systematics of Tibouchina and allies TAXON 00 (00) • 1–66

Appendix 1. Continued.
P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 391 (RB), JQ730143, JQ730354, JQ730565; Pleroma granulosum (Desr.) D.Don, Brazil, Guimarães,
P.J.F., 378 (RB), JQ730191, JQ730399, JQ730611; Pleroma heteromallum (D.Don) D.Don, Brazil, Guimarães, P.J.F., 339 (RB), JQ730193, JQ730401,
JQ730613; Pleroma hospitum (Schrank & Mart. ex DC.) Triana, Brazil, Baumgratz, J.F.A, 1008 (R), JQ730195, JQ730403, JQ730615; Pleroma itatiaiae
(Wawra) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 310 (RB), JQ730197, JQ730405, JQ730617; Pleroma kleinii (Wurdack) P.J.F.Guim. & Miche-
lang., Brazil, Almeda, F., 9868 (CAS), JQ730198, JQ730406, JQ730618; Pleroma laevicaule (Cogn. ex Wurdack) P.J.F.Guim. & Michelang., Brazil, Guimar-
ães, P.J.F., 412 (RB), JQ730199, JQ730407, JQ730619; Pleroma macrochiton (Mart. ex DC.) Triana, Brazil, Guimarães, P.J.F., 320 (RB), JQ730205,
JQ730413, JQ730625; Pleroma manicatum (Cogn.) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 328 (RB), JQ730206, JQ730414, JQ730626; Ple-
roma marinanum P.J.F.Guim. & Fraga, Brazil, Guimarães, P.J.F., 377 (RB), JQ730232, JQ730440, JQ730652; Pleroma martiale1 (Cham.) Triana, Brazil,
Goldenberg, R., 821 (NY), JQ730207, JQ730415, JQ730627; Pleroma martiale2 (Cham.) Triana, Venezuela, Michelangeli, F.A., 1269 (NY), JQ730202,
JQ730410, JQ730622; Pleroma martiusianum (DC.) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 335 (RB), JQ730208, JQ730416, JQ730628; Ple-
roma melanocalyx (R.Romero, P.J.F.Guim. & Leoni) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 341 (RB), JQ730209, JQ730417, JQ730629; Ple-
roma microphylla (Cogn.) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 344 (RB), JQ730210, JQ730418, JQ730630; Pleroma minus (R.Romero
& A.B.Martins) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 390 (RB), JQ730144, JQ730355, JQ730566; Pleroma mirabile (Brade & Markgr.)
P.J.F.Guim. & Michelang., Brazil, Santos, A.K.A., 338 (UFB), JQ730241, JQ730450, JQ730662; Pleroma mutabile (Vell.) Triana, Brazil, Lobão, A, 1450
(RB), JQ730213, JQ730421, JQ730633; Pleroma nodosum (Wurdack) P.J.F.Guim. & Michelang., Brazil, Santos, A.K.A., 1049 (UFB), JQ730215,
JQ730423, JQ730635; Pleroma oleifolium (DC.) R.Romero & Versiane, Brazil, Goldenberg, R., 1036 (NY), JQ730107, JQ730319, JQ730529; Pleroma oreo-
philum (Wurdack) P.J.F.Guim. & Michelang., Brazil, Santos, A.K.A., 829 (UFB), JQ730217, JQ730425, JQ730637; Pleroma penduliflorum Fraga & P.J.F.
Guim., Brazil, Goldenberg, R., 1150 (NY), JQ730231, JQ730439, JQ730651; Pleroma pereirae (Brade & Markgr.) P.J.F.Guim. & Michelang., Brazil, Santos,
A.K.A., 302 (UFB), JQ730221, JQ730429, JQ730641; Pleroma raddianum (DC.) Gardner, Brazil, Baumgratz, J.F.A, 1068 (R), JQ730222, JQ730430,
JQ730642; Pleroma radula (Markgr.) P.J.F.Guim. & Michelang., Brazil, Goldenberg, R., 1281 (NY), JQ730223, JQ730431, JQ730643; Pleroma ramboi
(Brade) P.J.F.Guim. & Michelang., Brazil, Almeda, F., 9889 (CAS), JQ730224, JQ730432, JQ730644; Pleroma salviifolium (Cham.) Triana, Brazil, Guimar-
ães, P.J.F., 322 (RB), JQ730225, JQ730433, JQ730645; Pleroma sellowianum (Cham.) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 331 (RB),
JQ730227, JQ730435, JQ730647; Pleroma semidecandrum (Schrank & Mart. ex DC.) Triana, Brazil, Goldenberg, R., 765 (NY), JQ730228, JQ730436,
JQ730648; Pleroma sp.1, Brazil, Guimarães, P.J.F., 402 (RB), JQ730146, JQ730357, JQ730568; Pleroma sp.2, Brazil, Guimarães, P.J.F., 400 (RB),
JQ730147, JQ730358, JQ730569; Pleroma stenocarpum (Schrank & Mart. ex DC.) Triana, Bolivia, Nee, M., 55386 (NY), JQ730233, JQ730441,
JQ730653; Pleroma trichopodum DC., Brazil, Lobão, A, 1437 (RB), JQ730235, JQ730443, JQ730655; Pleroma trinervium P.J.F.Guim., Brazil,
Almeda, F., 9682 (CAS), –, JQ730444, JQ730656; Pleroma urceolare (Schrank & Mart. ex DC.) Triana, Brazil, Guimarães, P.J.F., 321 (RB), JQ730236,
JQ730445, JQ730657; Pleroma ursinum (Cham.) Triana, Brazil, Almeda, F., 9859 (CAS), JQ730237, JQ730446, JQ730658; Pleroma velutinum (Naudin) Tri-
ana, Brazil, Santos, A.K.A., 349 (UFB), JQ730239, JQ730448, JQ730660; Pleroma vimineum (D.Don) D.Don, Brazil, Guimarães, P.J.F., 327 (RB), JQ730176,
JQ730385, JQ730596; Pleroma wurdackianum (R.Romero & A.B.Martins) P.J.F.Guim. & Michelang., Brazil, Guimarães, P.J.F., 395 (RB), JQ730148,
JQ730359, JQ730570; Pseudoernestia glandulosa (Gleason) M.J.R.Rocha & P.J.F.Guim., French Guiana, Martin, C.V., 471 (NY), JQ730080, JQ730293,
JQ730499; Pterogastra divaricata (Bonpl.) Naudin, Venezuela, Michelangeli, F.A., 540 (BH), JQ730126, JQ730337, JQ730547; Pterogastra minor Naudin,
Venezuela, Romero, G.A., 1682 (CAS), JQ730127, JQ730338, JQ730548; Pterolepis alpestris (DC.) Triana, Brazil, Almeda, F., 8469 (CAS), JQ730128,
JQ730339, JQ730549; Pterolepis glomerata (Rottb.) Miq., French Guiana, Martin, C.V., 419 (NY), JQ730129, JQ730340, JQ730550; Pterolepis parnassiifolia
(DC.) Triana, Brazil, Almeda, F., 8306 (CAS), JQ730130, JQ730341, JQ730551; Pterolepis polygonoides (DC.) Triana, Brazil, Goldenberg, R., 1463 (UPCB),
–, –, KF463008; Pterolepis repanda (DC.) Triana, Brazil, Almeda, F., 7731 (CAS), JQ730131, JQ730342, JQ730552; Pterolepis rotundifolia Wurdack, Brazil,
Almeda, F., 8290 (CAS), JQ730132, JQ730343, JQ730553; Pterolepis sp., Bolivia, Nee, M., 55383 (NY), JQ730133, JQ730344, JQ730554; Pterolepis steno-
phylla Gleason, Peru, Michelangeli, F.A., 2609 (NY), –, MH569721, MH463425; Pterolepis trichotoma (Rottb.) Cogn., Venezuela, Michelangeli, F.A., 823
(VEN), –, KF407966, KF463009; Rhexia alifanus Walt., United States of America, Abbott, J.R., 14892 (FLAS), DQ985623, –, –; Rhexia aristosa Britton,
United States of America, Naczi, R.F.C., 12065 (NY), JQ730134, JQ730345, JQ730555; Rhexia cubensis Griseb., unknown, Bostick, S., 388-1 (NCU),
DQ985627, –, –; Rhexia lutea Walt., United States of America, Abbott, J.R., 14882 (FLAS), DQ985628, –, –; Rhexia mariana L., United States of America,
Longbottom, W.D., 11730 (NY), JQ730135, –, JQ730556; Rhexia nashii Small, United States of America, Ionta, G.M., 398 (FLAS), DQ985662, –, –; Rhexia
nuttallii C.W. James, United States of America, Abbott, J.R., 14883 (FLAS), DQ985634, –, –; Rhexia parviflora Chapm., United States of America, Ionta, G.M.,
395 (FLAS), DQ985636, –, –; Rhexia petiolata Walt., United States of America, Ionta, G.M., 147 (FLAS), DQ985637, –, –; Rhexia salicifolia Kral & Bostick,
United States of America, Ionta, G.M., 407 (FLAS), DQ985639, –, –; Rhexia virginica L., United States of America, Michelangeli, F.A., 1448 (NY), JQ730136,
JQ730346, JQ730557; Rhynchanthera bracteata Triana, Bolivia, Zenteno, F., 8801 (NY), JQ730137, JQ730347, JQ730558; Rhynchanthera grandiflora
(Aubl.) DC., Venezuela, Michelangeli, F.A., 541 (BH), JQ730138, JQ730348, JQ730559; Rhynchanthera serrulata (L.C.Rich.) DC., Venezuela, Michelangeli,
F.A., 828 (BH), AY460435, JQ730349, JQ730560; Rostranthera tetraptera (Cogn.) M.J.R.Rocha & P.J.F.Guim., Brazil, Rocha, M.J.R., 757 (BHCB),
KU501067, KU501007, –; Rousseauxia andringitrensis (H.Perrier) Jacq.-Fél., Madagascar, Almeda, F., 9390 (CAS), JQ730139, JQ730350, JQ730561; Rous-
seauxia minimifolia (Jum. & H.Perrier) Jacq.-Fél., Madagascar, Almeda, F., 8704 (CAS), JQ730140, JQ730351, JQ730562; Sandemania hoehnei (Cogn.)
Wurdack, Brazil, Goldenberg, R., 1007 (NY), JQ730141, JQ730352, JQ730563; Schwackaea cupheoides Cogn., Costa Rica, Kriebel, R., 5819 (INB), –,
KF407967, KF463010; Siphanthera arenaria (DC.) Cogn., Brazil, Mota, N., 2656 (BHCB), KU501099, KU501203, –; Siphanthera cordata Pohl ex DC., Bra-
zil, Rocha, M.J.R., 964 (BHCB), KU501102, KU501208, –; Siphanthera cordifolia (Benth.) Gleason, Brazil, Rocha, M.J.R., 882 (BHCB), KU501096,
KU501201, –; Siphanthera dawsonii Wurdack, Brazil, Versiane, A.F., 641 (HUFU), KU501103, KU501207, –; Siphanthera fasciculata (Gleason) Almeda
& O.R.Rob., Peru, Wurdack, J.J., 1959 (NY), KU501097, –, –; Siphanthera foliosa1 (Naudin) Wurdack, Brazil, Goldenberg, R., 1611 (UPCB), –,
KF407968, –; Siphanthera foliosa2 (Naudin) Wurdack, Brazil, Rocha, M.J.R., 728 (BHCB), MH430236, –, –; Siphanthera gracillima (Naudin) Wurdack, Bra-
zil, Rocha, M.J.R., 941 (BHCB), KU501101, KU501206, –; Siphanthera hostmanii Cogn., Guyana, Wurdack, K.J., 4142 (NY), JQ730142, JQ730353,
JQ730564; Siphanthera paludosa (DC.) Cogn., Brazil, Rocha, M.J.R., 621 (BHCB), KU501100, KU501204, –; Siphanthera subtilis Pohl ex DC., Brazil, Mor-
eira, S.N., 878 (BHCB), KU501104, KU501209, –; Tibouchina aegopogon (Naudin) Cogn., Brazil, Almeda, F., 9523 (CAS), –, –, JQ730571; Tibouchina
aspera Aubl., French Guiana, Martin, C.V., 409 (NY), JQ730154, JQ730365, JQ730577; Tibouchina aspera Aubl., French Guiana, Martin, C.V., 423 (NY),
JQ730155, JQ730366, JQ730578; Tibouchina bicolor (Naudin) Cogn., Bolivia, Nee, M., 55307 (NY), JQ730159, JQ730370, JQ730581; Tibouchina bipeni-
cillata (Naudin) Cogn., Venezuela, Michelangeli, F.A., 735 (BH), JQ730160, JQ730371, JQ730582; Tibouchina brevisepala Cogn., Peru, Michelangeli, F.A.,
1958 (NY, USM), KF463039, KF407972, –; Tibouchina calycina Cogn., Peru, Michelangeli, F.A., 1883 (NY, USM), KF463040, KF407973, –; Tibouchina
fraterna N.E.Br., Venezuela, Michelangeli, F.A., 512 (BH), JQ730184, JQ730393, JQ730604; Tibouchina llanorum Wurdack, Venezuela, Michelangeli,
F.A., 736 (BH), JQ730203, JQ730411, JQ730623; Tibouchina octopetala Cogn. ex Britton, Bolivia, Solomon, J.C., 14934 (CAS), JQ730216, JQ730424,
JQ730636; Tibouchina papyrus (Pohl) Toledo, Brazil, Guimarães, P.J.F., 380 (RB), JQ730219, JQ730427, JQ730639; Tibouchina striphnocalyx (DC.) Glea-
son, Venezuela, Michelangeli, F.A., 379 (BH), JQ730234, JQ730442, JQ730654; Trembleya laniflora (D.Don) Cogn., Brazil, Almeda, F., 8549 (CAS),
AY553744, –, –; Trembleya parviflora (D.Don) Cogn., Brazil, Goldenberg, R., 824 (NY), JQ730242, JQ730451, JQ730663; Trembleya pentagona Naudin,
Brazil, Almeda, F., 8555 (CAS), AY553745, –, –; Tristemma coronatum Benth., Ghana, Schmidt, H.H., 3386 (CAS), JQ730243, JQ730452, JQ730664; Tri-
stemma hirtum P.Beauv., Gabon, Smith, S., 1879 (US), JQ730244, JQ730453, JQ730665; Tristemma littorale Benth., Gabon, Smith, S., 1725 (US),
JQ730245, JQ730454, JQ730666; Tristemma mauritianum J.F.Gmel., Madagascar, Almeda, F., 8021 (CAS), JQ730246, JQ730455, JQ730667.

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