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Systematics of the Trembleya sensu stricto clade of
Microlicia (Melastomataceae, Lavoisiereae)
Ricardo Pacico1,2, Frank Almeda2, Darin S. Penneys3, Karina Fidanza4
1Universidade Estadual de Maringá, Programa de Pós-Graduação em Biologia Comparada. Av. Colombo,
5790, 87020-900 Maringá, Paraná, Brazil 2California Academy of Sciences, Institute for Biodiversity Sci-
ence and Sustainability, Department of Botany, 55 Music Concourse Drive, Golden Gate Park, San Francisco,
California 94118-4503, USA 3Department of Biology and Marine Biology, University of North Carolina
Wilmington, Wilmington, North Carolina 28403, USA 4Departamento de Biologia, Universidade Estadual
de Maringá, Av. Colombo, 5790, Jardim Universitário, CEP 87020-900, Maringá, Paraná, Brazil
Corresponding author: Ricardo Pacico (ricardo_b9@hotmail.com)
Academic editor: Marcelo Reginato|Received 29 July 2022|Accepted 25 October 2022|Published 20 December 2022
Citation: Pacico R, Almeda F, Penneys DS, Fidanza K (2022) Systematics of the Trembleya sensu stricto clade of
Microlicia (Melastomataceae, Lavoisiereae). PhytoKeys 126: 1–101. https://doi.org/10.3897/phytokeys.126.91032
Abstract
A systematic monograph of the Trembleya s.s. clade is presented, a Brazilian endemic lineage of Melas-
tomataceae comprising 11 species and currently recognised as part of Microlicia s.l. (Melastomataceae).
First, we investigate phylogenetic relationships within Lavoisiereae using two nuclear markers and two
sampling datasets (102 and 134 terminals). en, we provide a systematic revision and new circumscrip-
tion of the Trembleya s.s. clade, including line drawings, photos of living specimens, leaves and oral parts,
distribution maps, a key to the 11 accepted species, comments on morphology, reproductive biology,
richness, endemism, biogeography and recommended conservation assessments. A nomenclatural update
of all taxa previously treated in Trembleya is also provided, including the designation of 45 lectotypes and
the proposal of 38 new synonyms.
Keywords
Brazil, Cadeia do Espinhaço, campo rupestre, Endemism, Minas Gerais
Introduction
Melastomataceae Juss. comprises about 5,880 species in 173 genera with about two-
thirds of this diversity restricted to the neotropics (Ulloa Ulloa et al. 2022). Brazil
is the most diverse country with 69 genera and about 1440 species and the family is
Copyright Ricardo Pacifico et al. This is an open access ar ticle distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
PhytoKeys 126: 1–101 (2022)
doi: 10.3897/phytokeys.126.91032
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Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
2
represented in all Brazilian biomes except for the Caatinga sensu stricto (Goldenberg
et al. 2020). e most comprehensive treatments of Brazilian Melastomataceae were
produced by Cogniaux (1883–1888, 1891) in “Flora Brasiliensis”, which were largely
based on tribal concepts proposed by Triana (1872). During the last century, the family
has been treated in many regional “Floras” and several genera and numerous new spe-
cies of Brazilian Melastomataceae were described (Goldenberg et al. 2012). Although
the work of Cogniaux (1883–1888) has long been the primary global reference for
the family, a new familial treatment for Brazil was recently published, including mor-
phological descriptions and keys to all Brazilian genera and species (Goldenberg et al.
2020), as part of the ambitious Brazilian Flora 2020 (BFG 2022).
As compared to the treatment in “Flora Brasiliensis”, the most recent infrafamil-
ial classications of Melastomataceae show signicant improvements. Based on DNA
sequence data, several new Neotropical tribes have been delimited, i.e. Henrietteeae
Penneys, Michelang., Judd & Almeda (Penneys et al. 2010), Marcetieae M.J.Rocha,
P.J.F.Guim. & Michelang. (Rocha et al. 2018), Pyramieae Naudin (as Cambessedesieae
Bochorny, Almeda, Michelang. & R.Goldenb.) (Bochorny et al. 2019), Rupestree-
ae Penneys & R.Goldenb., Stanmarkieae Penneys & Almeda (Penneys et al. 2022),
Trioleneae Bacci, Michelang. & R.Goldenb. (Bacci et al. 2019), Eriocnemeae Pen-
neys & Almeda and Lithobieae Penneys & Almeda (Penneys et al. 2020). Some new
genera have also proposed, for example, Rupestrea R.Goldenb., Almeda & Michelang.
(Goldenberg et al. 2015), Physeterostemon R.Goldenb. & Amorim (Goldenberg and
Amorim 2006), Brasilianthus Almeda & Michelang. (Almeda et al. 2016) and others
were relegated to synonymy (for example, see Penneys and Judd 2013 and Bochorny
et al. 2019); and there are still unplaced clades and genera that need to be tested for
monophyly (Penneys et al. 2022). Meanwhile, many new species are still being de-
scribed from little-studied genera and/or resulting from eldwork in areas that remain
poorly known or underexplored botanically.
With about 275–300 species, the near-endemic Brazilian tribe Lavoisiereae DC.,
which has long been known by the later tribal name Microlicieae Naudin, is one of the
richest clades of Neotropical Melastomataceae (Fritsch et al. 2004; Versiane et al. 2021;
Pacico and Almeda 2022). Currently, three genera are accepted, i.e. Rhynchanthera
DC. (15 spp.; Renner 1990), Poteranthera Bong. (5 spp.; Kriebel 2012; Almeda and
Pacico 2018) and Microlicia D.Don (ca. 250 spp.; Versiane et al. 2021; Pacico and
Almeda 2022). However, this tribe has a complex taxonomic history and its current ge-
neric delimitation has been drastically modied in light of molecular evidence (Fritsch
et al. 2004; Rocha et al. 2016; Versiane et al. 2021). Based on rpl16 and nrITS data,
Fritsch et al. (2004) found support for the restriction of Lavoisiereae to the six genera
previously enumerated by Almeda and Martins (2001). In turn, phylogenetic analyses
of Rocha et al. (2016) were decisive in assigning Poteranthera to the tribe where it
had been placed by Triana (1872) and several other early classications, thus resulting
in the recognition of seven genera in Lavoisiereae. Major taxonomic alterations were
proposed by Versiane et al. (2021), who included four traditionally recognised genera
(Chaetostoma DC., Lavoisiera DC., Stenodon Naudin, Trembleya DC.) in a broadly
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 3
circumscribed Microlicia s.l. Versiane et al. (2021) recovered little support for most
clades within Microlicia s.l., whereas some internal clades were strongly supported and
are recognisable morphologically, e.g. the “Trembleya s.s.” “Lavoisiera” and “Chaetosto-
ma” clades (Koschnitzke and Martins 2006; Martins and Almeda 2017; Pacico et al.
2019; Versiane et al. 2021). For a complete taxonomic history of the tribe, see Martins
and Almeda (2017: 32–34) and Pacico and Almeda (2022).
Trembleya s.s. is one of the clades currently recognised as part of Microlicia s.l. (Versi-
ane et al. 2021; Pacico and Almeda 2022). We here re-evaluate the status of this lineage
in a phylogenetic context and provide a systematic revision and new circumscription of
the Trembleya s.s. clade. Line drawings, photos of living specimens, leaves and oral parts,
distribution maps, a complete nomenclatural update, comments on morphology, repro-
ductive biology, richness, endemism, biogeography and conservation are also provided.
Historical background
Trembleya was named by Augustin Pyramus de Candolle (1778–1841) in honour of
the members of the family Trembley for their contributions to several elds of science
(Candolle 1828: 125). At that time, the genus was represented by six species, all of
which were described, based on the collections made by Carl Friedrich Philipp von
Martius in Brazil (Spix and Martius 1824). Candolle (1828: 114) placed Trembleya
along with other 13 genera in the tribe Rhexieae DC. is arrangement changed about
two decades later when Charles Victor Naudin (1818–1889) assigned this genus to the
tribe Lavoisiereae (as “Microliciales”) (Naudin 1849: 203). Trembleya was maintained
in the tribe Lavoisiereae (as “Microlicieae”) in subsequent treatments of Melastomata-
ceae like those by Triana (1872) and Cogniaux (1883–1888, 1891).
Trembleya was recognised as a heterogeneous group of species since it was rst
delimited by Candolle (1828), who recognised three morpho-groups. ree sections
were proposed, two of which were named in honour of members of the Trembley fam-
ily (Candolle 1828). e section Jacobia DC. was named after Jacob Trembley and the
section Abrahamia DC. was named after Abraham Trembley (Candolle 1828: 125–
126). e remaining section, composed of only Trembleya lycnithis DC. [= Microlicia
laniora (D.Don) Baill.], was named Erioleuca DC. in reference to the lanose appear-
ance of this species (Candolle 1828: 126). Candolle (1828b: 38) also claimed that
these three sections could one day either be brought together through more nuanced
intermediaries or separated into three distinct genera.
Chamisso (1834) followed Candolle’s infrageneric classication and described two
new species in the genus: Trembleya pithyoides Cham. in section Jacobia and Trembleya
calycina Cham. in sect. Abrahamia. Naudin (1849: 264) agreed with Candolle (1828)
on the polymorphic aspect of Trembleya, considering it a “genus habitu specierum omnino
heteromorphum” [genus in which the habit is highly heteromorphic amongst species].
Naudin (1849) reformulated the infra-generic classication of the genus under two
new sections, Trembleya sect. Verae Naudin and Trembleya sect. Heterogenae Naudin.
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
4
In “Flora Brasiliensis”, Cogniaux (1883–1888) mostly maintained the sections
proposed by Naudin (1849). Trembleya sect. Trembleya Cogn. contained the same spe-
cies of Trembleya attributed to sect. Verae, but included two taxa that were not available
for study in Paris by Naudin, Trembleya chamissoana Naudin and Trembleya rosmari-
noides Mart. & Schrank ex DC. In addition, Cogniaux (1883–1888) resurrected Can-
dolle’s monotypic section Erioleuca, which included only Trembleya laniora. e sec-
tion Heterogenae was also maintained by Cogniaux (1883–1888), who included three
additional species that were described after Naudin’s monographs, Trembleya selloana
Cogn., Trembleya warmingii Cogn. and Trembleya pradosiana Netto.
Renner (1990: 609) stated that Trembleya was “an assemblage of species diering
greatly in habit and pubescence”. Martins (1995) described another species, Trembleya
hatschbachii Wurdack & E.Martins. In the unpublished monograph of Trembleya by
Martins (1997), 18 species were recognised (several never validly published) and three
dubious taxa were noted. Martins (1997) did not recognise any of the infrageneric sec-
tions previously proposed in the genus.
Martins (1997) made a special eort to understand the supra-generic delimitation
of Trembleya. She emphasised that the number of ovary locules was the only feature
used by Cogniaux (1883–1888) to dierentiate this genus from Microlicia D.Don
(3-locular in Microlicia and 5-locular in Trembleya). In this context, a major objective
of Martins (1997) was to clarify the limits between Trembleya and Microlicia. Based on
comparative morphology, Martins (1997) proposed an expanded re-circumscription
of Trembleya, which was based on characters, such as inorescence structure, vegeta-
tive architecture and leaf venation patterns. However, Martins (1997) recognised that
none of these features could dierentiate Trembleya from Microlicia in an eective way
(Martins 1997). Besides, Martins (1997) mentioned taxa with overlapping features,
such as Microlicia pabstii Brade, that has leaves with evident secondary venation, a
feature attributed only to Trembleya (sensu Martins 1997) and isolated axillary owers,
a typical feature of Microlicia (Martins 1997).
Fritsch et al. (2004) conrmed the placement of Trembleya in Lavoisiereae using
molecular data. Subsequently, new species were described in Trembleya by Fidanza
et al. (2013), Pacico and Fidanza (2015) and Pacico et al. (2019). Additionally,
one species was transferred from Lavoisiera DC. to Trembleya by Almeda and Martins
(2001) [Trembleya elegans (Cogn.) Almeda & A.B.Martins], and another was trans-
ferred from Trembleya to Poteranthera by Almeda and Pacico (2018) [Poteranthera
warmingii (Cogn.) Almeda & R.B.Pacico]. Pacico et al. (2019) argued that Tremb-
leya was polyphyletic and a new circumscription was necessary in order to reduce this
genus to a monophyletic group of 11 species (Trembleya s.s.).
Based on an expanded phylogenetic analyses using ve molecular markers (atpF-
atpH, trnS-trnG, nrITS, nrETS and waxy) and 12 morphological characters for 113
taxa in Lavoisiereae, Versiane et al. (2021) transferred all species of Trembleya to the
enlarged Microlicia s.l. In order to accommodate these species in Microlicia, Versiane et
al. (2021) proposed 13 new combinations [M. altoparaisensis (R.B.Pacico, Fidanza &
Almeda) Versiane & R.Romero, M. calycina (Cham.) Versiane & R.Romero, M. chamis-
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 5
soana (Naudin) Versiane & R.Romero, M. inversa (Fidanza, A.B.Martins & Almeda)
Versiane & R.Romero, M. neopyrenaica (Naudin) Versiane & R.Romero, M. parviora
(D.Don) Versiane & R.Romero, M. pentagona (Naudin) Versiane & R.Romero, M.
phlogiformis (Mart. & Schrank ex DC.) Versiane & R.Romero, M. pithyoides (Cham.)
Versiane & R.Romero, M. pradosiana (Netto) Versiane & R.Romero, M. rosmari-
noides (Mart. & Schrank ex DC.) Versiane & R.Romero, M. thomazii (R.B.Pacico
& Fidanza) Versiane & R.Romero, M. tridentata (Naudin) Versiane & R.Romero)]
and three new names [M. acuminifolia Versiane & R.Romero, M. aviora Versiane &
R.Romero, M. serratifolia Versiane & R.Romero], whereas M. laniora (D.Don) Baill.
was reinstated. e analyses of Versiane et al. (2021) also supported the recognition
of Trembleya s.s. as a distinct clade, as previously suggested by Pacico et al. (2019).
Although recognisable morphologically, the Trembleya s.s. clade was transferred to Mi-
crolicia by Versiane et al. (2021) because the recognition of that clade as a distinct
genus would make Microlicia s.l. paraphyletic. is issue is also re-evaluated in the
present study.
Material and methods
Taxa sampling and DNA extraction
To investigate phylogenetic relationships in Lavoisiereae, we used two dierent datasets
including samples from the seven genera traditionally recognised in the tribe (Almeda
and Martins 2001; Fritsch et al. 2004). Dataset A comprised samples with complete
sequences (i.e. tting perfectly the alignment on both ends) of both nrITS and nrETS.
is dataset had 102 terminals, representing 99 species (ingroup, 91 species; eight
species are outgroups). Dataset B comprised samples with complete nrITS and nrETS
sequences, but also samples lacking sequences of one of these markers and/or partial
sequences, including 134 terminals representing a total of 131 species (ingroup, 127
species; four species are outgroups). Overall, taking the two datasets into considera-
tion, 247 sequences were generated for this study (125 nrITS plus 122 nrETS; total
of 131 species). Eight sequences were retrieved from GenBank and added to the align-
ment (seven nrETS plus one nrITS; total of seven species). For information on speci-
men vouchers and GenBank accessions, see Suppl. material 1.
Total genomic DNA was extracted from silica-dried leaves or from herbarium
specimens. DNA extraction was performed mainly using Qiagen DNeasy plant kits,
following the manufacturer’s recommendations (https://www.qiagen.com/) or modi-
ed to extend the incubation time in AP1 Buer and RNAase to 1 hour, followed by
1 hour of precipitation in ice. A few samples were extracted using the modied CTAB
(hexadecyltrimethylammonium bromide) protocol of Doyle and Doyle (1987).
Leaves were ground in a mixer mill, incubated in 1.2 ml CTAB and 5 µl of Proteinase
K for 60–180 minutes at 55 °C. DNA was precipitated overnight in isopropanol and
sodium acetate at -20 °C, centrifuged at 13,000 rpm for 20 minutes, washed twice
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
6
with 70% ethanol and dried at room temperature. e pellets with DNA were then
resuspended in 100µl of 1× TE (tris-EDTA buer, pH 8.5) and incubated at 55 °C
for about 30 minutes.
PCR and DNA sequencing
We amplied and sequenced the ribosomal markers nrETS and nrITS. In Melastomata-
ceae, the nrETS region has been used by Kriebel et al. (2015), Rocha et al. (2016), Bacci
et al. (2020) and Versiane et al. (2021). e nrITS is widely used in phylogenetic studies
of the family (e.g. Ionta et al. 2007; Penneys and Judd 2013; Reginato and Michelangeli
2016; Zhou et al. 2018). Information on the primers used in this study may be found
in the aforementioned literature. e PCR recipes were: (1) for nrETS, 1.5 µl of Buer
10×, 0.9 µl of MgCl2 25 uM, 0.3 µl of each primer (10 uM), 0.3 µl of dNTP (10 uM),
0.3 µl of Hotstart Ta q polymerase (1.25 uM) and 1 µl of template, reaching 15 µl of
nal reaction; (2) for nrITS, the recipe was similar to (1), but diered in including 1.2
µl of BSA (10 uM) and using Invitrogen Taq polymerase to improve the performance.
e thermocycler programmes were: (1) for nrETS initial denaturation at 95 °C for 2
minutes, 40 cycles of denaturation at 94 °C for 30 seconds, annealing at 58 °C for 30
seconds; extension at 72 °C for 1 minute and nal extension at 72°C for 7 minutes; (2)
for nrITS, initial denaturation at 94 °C for 5 minutes, 40 cycles of denaturation at 94
°C for 10 seconds; annealing at 61 °C for 45 seconds; extension at 72°C for 40 seconds
and nal extension at 72 °C for 10 minutes. e quality of amplications was veried
in agarose electrophoresis gel. e amplied fragments were cleaned using 5 µl of PCR
products (7 µl for weak bands) and 2 µl of diluted ExoSAP-IT. e PCR cleaning was
performed by incubating the samples at 37 °C for 30 minutes and 80 °C for 15 minutes.
Cycle sequencing was performed by using the same amplication primers. e thermo-
cycler programme used for cycle sequencing began at 96 °C for 60 sec, followed by 15
cycles of 10 sec at 96 °C, 5 sec at 50 °C and 75 sec at 60 °C; another 5 cycles of 10 sec at
96 °C, 5 sec at 50 °C, 90 sec at 60 °C; 5 sec at 50 °C and 120 sec at 60 °C. e Sanger
Sequencing was performed in an ABI 3130 Genetic Analyzer in the Center for Com-
parative Genomics at the California Academy of Sciences, San Francisco, CA. GenBank
accessions for all sequences used in this study are provided in Suppl. material 1.
Phylogenetic analyses
Contigs were assembled and edited using Geneious 11.0 (https://www.geneious.com).
e same software was employed to align (using Muscle algorithm), concatenate and
export the sequences as fasta les. Low-quality sequences were discarded. Final manual
edits on the alignment were performed using Mega 7 (Kumar et al. 2016). Maximum
Likelihood (ML) and Bayesian Inference (BI) phylogenetic analyses were performed
using the concatenated sequences of dataset A, whereas dataset B was only analysed
using ML. All analyses were run through the CIPRES online gateway (http://www.
phylo.org/; Miller et al. 2012) as four independent partitions (nrETS, ITS1, 5.8S and
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 7
ITS2). e ML analyses were performed using a fast Maximum Likelihood tree search
algorithm (RAxML, Stamatakis 2014) with the GTR+Γ+I model and 1,000 bootstrap
repetitions. e BI analysis was performed with MrBayes version v.3.2.6 using the
same model (GTR+Γ+I), which was employed for being the most parameter-rich (see
Abadi et al. 2019). BI analyses consisted of two independent runs, each one with one
cold and three heated simultaneous independent Markov chains. A total 15,000,000
generations were carried out, sampling one tree every 1000 generations and discarding
25% of the sampled trees as burn-in. A node was considered to be strongly supported if
it showed both a Bayesian Posterior Probability ≥ 0.95 and a Bootstrap Support ≥ 70%.
Taxonomic study
Specimens of all species of the Trembleya s.s. clade were examined. is study was car-
ried out using collections from the following herbaria: BM, BR, C, CAS, CESJ, ESA,
F, G, GH, HB, HUEM, INPA, K, L, M, MBM, MO, NY, P, R, RB, S, SPF, UEC, US
and W (acronyms follow iers 2022). Specimens of some of the mentioned herbaria
were examined only through online images (BM, C, CESJ, ESA, G, GH, INPA, L, M,
MO and S). All types and their synonyms were examined directly or through online
images available from databases, for example, JSTOR Global Plants (https://plants.
jstor.org/) and SpeciesLink (http://www.splink.org.br/). A list with all specimens cited
in this study is provided in Appendix 1.
Populations of all but two species (M. rosmarinoides and M. trembleyiformis) of the
Trembleya s.s. clade were sampled in the eld during several expeditions carried out by
the authors, mainly in the following regions of Brazil: Serra do Cipó, Serra do Cabral,
Serra do Bota, Serra de Grão Mogol, Serra do Caraça, Serra do Gandarela, Serra de
Ouro Preto and on the Diamantina Plateau (in Minas Gerais State), Chapada Dia-
mantina (Bahia State), Chapada dos Veadeiros and Serra dos Pirineus (Goiás State) and
Parque Estadual do Guartelá (Paraná State). All specimen identications were carefully
checked by the authors. We considered one specimen as a putative hybrid between M.
pentagona and M. laniora because it clearly intermixes characters between these two
species and was collected in an area where they occur sympatrically.
e morphological diagnosis was based on a comparative study of species in the
clade and all remaining taxa within Lavoisiereae. Species descriptions, lists of specimens
examined and circular histograms of phenology were prepared using the functions
“TabletoDescription”, “Collector List” and “PhenoHist” (respectively), available from
the package monographaR implemented in R (Reginato 2016). Herbarium specimens
were also the source of all distributional data used in this study. All coordinates obtained
from specimen labels were mapped to verify if they matched the locality described.
SEM images
Scanning Electron Microscope (SEM) photos were taken from herbarium specimens.
Sections of leaf blades (all species in the clade), anthers (four species), seeds (ve species)
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
8
and pollen grains (M. pithyoides) were axed using double-stick tape on pin stubs, sput-
ter-coated with gold and examined using a Hitachi SU3500 microscope with coupled
digital camera in the SEM Lab of the California Academy of Sciences, San Francisco, CA.
Anatomical studies
Dried leaves of M. altoparaisensis, M. chamissoana and M. aviora were obtained
from exsiccatae (housed in HUEM), rehydrated in sodium hydroxide (NaOH) 5%
(Anderson 1963, modied) and stored in 70% methyl alcohol (Johansen 1940). Epi-
dermal dissociation was performed either using acetic acid and hydrogen peroxide
(1:1) (Franklin 1945) or 10% nitric acid and 10% chromic acid (1:1) (Johansen 1940).
Epidermal sections were stained using Safranine and Astra blue (Bukatsch 1972) and
mounted in glycerine 50% (Kraus and Arduin 1997). Samples of the leaf blade were
embedded in historesin (2-hydroxyethylmethacrylate). Cross and longitudinal sections
of the leaf blade (7 µm) were obtained using a rotary microtome and stained with
0.05% toluidine blue, acetate buer at pH 4.7 (O’Brien et al. 1965, modied). Light
micrographs were taken using a Leica EZ4D digital camera in the Department of
Biology of the Universidade Estadual de Maringá.
Biogeography and mapping
Species richness and weight endemism analyses were run in R through the Dinamica
Ego platform (http://www.dinamicaego.com) using the tools “SR” and “WE” from the
toolkit BioDinamica 2.2 (Oliveira et al. 2019). In both cases, the distributional data were
saved as a csv le and analysed by employing a grid of hexagons in which the distances
amongst centroids were 1°. We delimited the main area of endemism for the Tremb-
leya s.s clade based on endemicity analyses implemented in the NDM/VNDM software
(Szumik et al. 2002; Szumik and Golobo 2004). A spreadsheet with the distributional
data was saved as a csv le and converted into a xyd le using the GeX online tool (http://
gex.mfuhlendorf.com/ (Santos and Fuhlendorf 2018). In NDM/VNDM, we used a grid
of 1° cells and searched for Areas of Endemism with 1000 repetitions. Only individual
areas of endemism with scores equal to or higher than 2 were considered and overlapping
subsets were kept only if they had at least 40% unique species. e consensus area of en-
demism was obtained using the loose consensus rule and a cut-o value of 40% (see Aa-
gesen et al. 2013). Coordinates of the consensus area were exported from NDM/VNDM
as a text le and converted into a shapele using DIVA-GIS (Hijmans et al. 2001). All
maps were prepared using QGIS 3.14 software (QGIS Development Team 2021).
Conservation assessments
Following the guidelines of criterion B of IUCN (2019), we provide informal con-
servation status recommendations for each species, based on georeferenced data
from herbarium material. GeoCAT (Bachman et al. 2011) was used to calculate Ex-
tent of Occurrence (EOO) and Area of Occupancy (AOO) for each species using a
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 9
user-dened cell width of 2 km. We are aware that the AOO is often inappropriately
used for conservation assessments, as most plant species are represented in herbaria
by only a few specimens, thereby underestimating the AOO (Lughadha et al. 2019).
For Lavoisiereae, our eld experience showed that most species in this tribe are of-
ten represented only by a few populations restricted to rocky outcrops on a shallow
layer of white quartzitic soil. ese populations usually occupy an area smaller than
4 km2; thus, we believe that the cell width of 2 km is appropriate for these analyses.
Results and discussion
Molecular phylogenetics
e aligned matrix of the combined nrETS+nrITS contains 1435 characters. In all
analyses, Lavoisiereae was recovered as monophyletic with strong statistical support,
in agreement with the analyses of Fritsch et al. (2004) and Versiane et al. (2021).
Rhynchanthera was recovered as the rst diverging lineage (as reported by Fritsch et
al. 2004 and Versiane et al. 2021), followed by Poteranthera and Microlicia s.l. (in
agreement with Versiane et al. 2021; Figs 1, 2). In this context, even with the inclu-
Figure 1. Maximum Likelihood (left) and Bayesian Inference (right) trees of Lavoisiereae, based on
analyses of nrITS+nrETS sequences using dataset A. Bootstrap support values ≥ 70% and Bayesian Poste-
rior Probability values ≥ 0.9 are given at each node.
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
10
sion of Poteranthera by Rocha et al. (2016), the Lavoisiereae can be distinguished
amongst other capsular-fruited Melastomataceae by its elongate to reniform seeds
with a reticulate-foveolate testa, stamens forming well-developed pedoconnectives
and anthers with rostrate thecae (Fritsch et al. 2004; Rocha et al. 2016; Pacico and
Almeda 2022).
Overall, species traditionally recognised in Trembleya (for example, by Cogniaux
1883–1888, 1891 and Martins 1997) were recovered in three distinct regions of the
Lavoisiereae tree. Using the dataset A, a clade composed exclusively of species of Trem -
bleya s.s. was recovered as sister to the rest of Microlicia s.l. in the ML tree, with a
Bootstrap Support of 70% (Fig. 1). However, this sister relationship was unresolved in
the BI tree (Fig. 1). Several clades within Microlicia s.l. had low topological resolution
in all analyses and especially using dataset B (Fig. 2).
e analyses of nrETS and nrITS resulted in incompletely resolved trees with sev-
eral weakly-supported clades, making it impossible to evaluate the monophyly of most
clades within Microlicia s.l. is was expected since these lineages have been recovered
with very short branch lengths in family-level phylogenetic analyses, based on the mul-
ti-loci approach that used a larger sampling of Melastomataceae (Penneys et al. 2022).
However, a clade that is diagnosable morphologically and is a good match to Trembleya
s.s. of Pacico et al. (2019) was identied. e recovery of this clade as sister to the re-
maining Microlicia s.l. in the ML tree, based on complete nrETS and nrITS sequences,
suggests that this group of species could be treated as a distinct genus. is taxonomic
decision was neither conrmed nor refuted by both the BI tree using dataset A and the
ML tree using dataset B. us, we here tentatively treat the Trembleya s.s. clade as part
of Microlicia. Future studies using enhanced taxonomic characters may provide a deni-
tive assessment of the limits between this clade and Microlicia s.l.
Figure 1. Continued.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 11
Figure 2. Maximum Likelihood tree of Lavoisiereae, based on analyses of nrITS+nrETS sequences using
dataset B. Bootstrap support values ≥ 70% at each node.
General characterisation
Habit and branching
Species of the Trembleya s.s. clade are woody and perennial erect shrubs, usually
0.5–2m tall, small trees 2.5–4 m tall (e.g. M. laniora and M. parviora) or caespi-
tose shrubs (e.g. M. pithyoides and M. rosmarinoides). Species can be highly variable
in habit, like M. laniora, which is typically a small tree (1.7–2.5 m tall), but may
also be a shrub up to 0.8 m tall with divaricate branching (Fig. 3). Both forms may
occur in the same population. e small trees of M. laniora stand out by having a
single trunk with corky bark. Variation in habit is also common in the wide-spread
M. parviora, generally from small shrubs ca. 0.5 m tall to small trees to 3–4 m
tall. On the Serra do Cipó (Minas Gerais), a “dwarf” specimen of M. chamis-
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
12
soana measuring about 10 cm tall when owering was collected once (Giulietti et
al CFSC12492). e branching pattern in the Trembleya s.s. clade is consistently
di-trichotomic, with quadrangular branchlets that are unwinged or have narrow
wings ca. 0.2 mm wide. e branchlets are typically glandular-punctate and may
be covered by other types of trichomes of variable type and distribution. Unlike
many taxa of the Lavoisiera clade of Microlicia s.l. (Martins and Almeda 2017), all
species in the Trembleya s.s. clade do not present conspicuous scars where the leaf
has fallen away.
Figure 2. (continued).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 13
Figure 3. Habit in the Trembleya s.s. clade of Microlicia A Celso de Paiva (ICMBio) taking a picture of
a small M. laniora tree B M. parviora, small tree C M. laniora with divaricate habit D M. chamissoana
with divaricate habit E M. rosmarinoides, a small, much-branched shrub F, G Saxicolous shrubs: F M. a-
viora G M. pentagona. Photos: A by O. Grae B by R. Penati C , E by L. Pedrosa D, F, G by R. Pacico.
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14
Root and stem anatomy and development
Information on root and stem anatomy and development is limited to the study of So-
mavilla and Graciano-Ribeiro (2012) on M. parviora. e roots of M. parviora have
a uniseriate epidermis with outer periclinal walls and part of the anticlinal walls thick-
ened and the cuticle thinner than the outer periclinal wall. In more developed roots, the
epidermis is replaced by exodermis (Somavilla and Graciano-Ribeiro 2012). Likewise,
the stems of M. parviora have a uniseriate epidermis with outer periclinal walls slightly
thickened and covered by a cuticle (Somavilla and Graciano-Ribeiro 2012). e cortex
and the centre of the stems are separated by an endodermis layer and the central part
of the cortex has large intercellular spaces (Somavilla and Graciano-Ribeiro 2012). e
pericycle consists of 1–2 layers of parenchymatic cells surrounding the vascular system
and the pith is parenchymatic (Somavilla and Graciano-Ribeiro 2012).
Microlicia parviora stands out by having a spiralled aerenchymatous polyderm sur-
rounding the root and stem (Somavilla and Graciano-Ribeiro 2012). In this species, the
secondary growth of the stems and roots starts with the development of the vascular
cambium, as well as the phellogen from pericycle cells. Derivative cells from phellogen
then dierentiate externally into two cell types disposed in concentric layers and centrif-
ugally intercalated (Somavilla and Graciano-Ribeiro 2012). e one-layered phelloderm
is produced centripetally by the phellogen (Somavilla and Graciano-Ribeiro 2012).
Leaf morphology
Leaf shape is highly variable and, together with indumentum and venation, it is one
of the most taxonomically informative characters in the Trembleya s.s. clade (Fig. 4).
Leaves are opposite and decussate like most Melastomataceae, highly variable in size
along the branches and usually not imbricate. All species of the Trembleya s.s. clade
have petiole features that are shared in a consistent way only with Rhynchanthera
(see Renner 1990). e leaf blades vary from linear to oblong, lanceolate, narrowly-
elliptical, elliptical, ovate or rarely slightly obovate. Leaf consistency varies from
papyraceous to chartaceous or coriaceous. In 10 out of 11 species recognised in the
Trembleya s.s. clade, the leaf blades become strongly discoloured when dry. e blade
is usually plane throughout, although (when dry), the marginal region may become
slightly revolute (in three species) or conspicuously revolute in M. calycina. e leaf
blade is never keeled like it is in species of the Chaetostoma clade (Koschnitzke and
Martins 2006). e leaf margin is entire, slightly serrulate or entire along the basal
half and serrulate at the upper half; it lacks the stout glandular trichomes typical
of Poteranthera (Almeda and Pacico 2018). It is also not callose-thickened like in
most species of the Lavoisiera clade (Martins and Almeda 2017). e leaves always
have 1–3 primary basal veins and two or more pairs of secondary basal acrodromous
veins (see Carmo et al. 2019). In most species, the most external pair of acrodromal
veins is tenuous and positioned close to the leaf margin. Ten out of 11 species have
tertiary veins more or less evident on the abaxial leaf surface, which are arranged
nearly perpendicular to the mid-vein and may branch randomly or only apically.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 15
When mature, the leaf adaxial surface is always glandular-punctate to glabrescent.
e abaxial surface is always glandular-punctate and often covered with other types
of trichomes.
Leaf anatomy and indumentum
Information on leaf anatomy is available for M. altoparaisensis (Pacico et al.
2019), M. calycina (Paum 1897), M. chamissoana (Carmo et al. 2019, 2020), M.
laniora (Paum 1897; Silva et al. 2018), M. parviora (Paum 1897; Somavilla
and Graciano-Ribeiro 2011), M. aviora (Carmo et al. 2019, 2020), M. pithyoides
Figure 4. Leaves in the Trembleya s.s. clade of Microlicia in abaxial view A M. tridentata B M. altopara-
isensis C–E M. pentagona F M. rosmarinoides G M. pithyoides H M. aviora I M. laniora J M. parviora
K M. trembleyiformis L M. calycina M M. chamissoana. Voucher specimens: A Longhi-Wagner et al.
CFCR9184 (SPF, UEC, US) B Glaziou 21300 (F, P, S) C Barreto 10734 (BHCB) D Barreto 7025 (BHCB)
E Rapini et al. 296 (HUEM, SP, SPF) F Souza et al. 2584 (BHCB) G Pacico 295 (CAS, HUEM, SPF)
H Hatschbach et al. 52005 (S) I Pacico 185 (HUEM, SPF) J Pacico 191 (HUEM) K Romero & Naka-
jima 3593 (HUFU, K, UEC) L Pacico 290 (CAS, SPF) M Romero et al. 8627 (HUEM, HUFU, RB).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
16
(Paum 1897), M. rosmarinoides (Paum 1897) and M. tridentata (Paum 1897).
A plate with photos of leaf anatomical characters is presented in Fig. 5 (A–F).
Overall, the leaf epidermis is uniseriate with outer periclinal walls thicker than
the other walls. e adaxial epidermal cells have straight to slightly sinuous walls
in top view, while the abaxial epidermal cells have walls varying from straight to
sinuous (Fig. 5C). Microlicia altoparaisensis has amphistomatic leaves, while the
remaining four species have hypostomatic leaves. Cuticle striation was reported
for M. chamissoana and M. aviora. In cross-section, the petiole has an ovate or
arched shape, 3–5 bicollateral or amphicribal vascular bundles (Fig. 5E) and col-
lenchyma in the cortical region. e mesophyll varies from isobilateral (Fig. 5B) to
dorsiventral or mixed, always with palisade parenchyma as the cortical tissue and
the mid-rib projected. e shape of the mid-rib varies from arched to open arched,
with bicollateral or amphicribral vascular bundles (Fig. 5A) and parenchyma and/
or collenchyma at the cortical region. Sclereids and crystalliferous cells generally
occur in the petiole, mesophyll and mid-rib. In M. aviora and M. chamissoana,
the veinlets have imperfect areoles and terminal sclereids and the apical venation is
not ramied (g. 5C in Carmo et al. 2019). In the marginal region, the venation
is incomplete in M. aviora and arch-shaped in M. chamissoana (g. 5E in Carmo
et al. 2019).
All species of the Trembleya s.s. clade have some type of trichome on the leaves,
branchlets and abaxial leaf surfaces. Scanning Electronic Microscopy images showing
details of the leaf surface in the clade are presented in Figs 6–7. e common glandu-
lar-punctate indumentum consists of a set of glandular trichomes with a multicellular
head and a stalk that is too short to be seen with a typical stereomicroscope (Carmo
et al. 2019; Figs 5F, 6A, B, F, 7D–F). is type of glandular trichome was initially de-
scribed by Paum (1897), but only illustrated in detail by Mentink and Baas (1992; as
“bladder-like glandular hairs”). Similar trichomes were commonly referred to as “sessile
glands”, “sessile glandular trichomes” or “unforrowed sessile glands” (for example, by
Wurdack 1986) across the taxonomic literature. As these trichomes are not sessile, they
have been treated as typical glandular trichomes in anatomical studies (Reis et al. 2005;
Cassiano et al. 2010; Romero and Castro 2014; Silva et al. 2018; Carmo et al. 2019,
2020). In a more detailed investigation, Carmo et al. (2019) proposed new names for
trichomes and epidermal appendages, based on the structure of the stalk, head and
cuticle. Carmo et al. (2019) included two species of the Trembleya s.s. clade in their
study (M. chamissoana and M. aviora). Together with the anatomical descriptions
and gures provided by Somavilla and Graciano-Ribeiro (2011), Silva et al. (2018) and
Pacico et al. (2019), the following types of trichomes were reported for the Trembleya
s.s. clade:
1. Reduced glandular trichomes with apex consisting of 2–4 globular cells, dis-
tended cuticle and pluricellular stalk, in M. chamissoana and M. aviora (Carmo et
al. 2019);
2. Uniseriate elongated eglandular trichomes, narrow at the base and truncated
at the apex, with circular depressions in M. chamissoana (Carmo et al. 2019; Fig. 6C);
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 17
Figure 5. Anatomical features of leaves in the Trembleya s.s. clade of Microlicia A–C Microlicia altopara-
isensis: A cross section of the mid-vein, showing an amphicribral arch-shaped vascular bundle B cross
section of the leaf, showing the isobilateral mesophyll, the wavy epidermis and a glandular trichome in
a depression C Adaxial surface of the epidermis in frontal view, with stomata and glandular trichomes
D, E Microlicia chamissoana: D leaf stomatal crypt in cross section, with glandular trichomes E petiole
in cross section, showing an amphicribral vascular bundle F Microlicia aviora, cross section of the leaf
blade showing a reduced glandular trichome. AB: Abaxial surface; AD: Adaxial surface; CE: Common
epidermal cells; GT: Glandular trichome; PP: Palisade parenchyma; SP: Spongy parenchyma; ST: Sto-
mata. All photos by A.A.O. Carmo. Voucher specimens: A–C Pacico & Bressan 380 (CAS, HUEM,
SPF) D, E Pacico & Carmo 154 (HUEM, UEC) F Mello-Silva et al. 509 (SPF).
3. Short-stalked glandular trichomes with pluricellular clavate apex, with or
without a distended cuticle, in M. parviora (Somavilla and Graciano-Ribeiro 2011)
and M. altoparaisensis (Pacico et al. 2019; Fig. 6A);
4. Woolly eglandular trichomes in M. laniora (Silva et al. 2018; Fig. 6E).
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18
Inorescence
Given the diversity of inorescence types and the presence of many similar structures
which are not guaranteed homologous, eorts have been made to establish natural sys-
tems for the ower-bearing parts of the Angiosperms (e.g. Briggs and Johnson 1979;
Weberling 1988; Prusinkiewicz et al. 2007; Kircho and Claßen-Bockho 2013).
Figure 6. SEM images showing details of the leaf surface in the Trembleya s.s. clade of Microlicia
A Microlicia altoparaisensis, glandular trichomes and stomata on the leaf abaxial surface B Microlicia
calycina, glandular trichomes and stomata on the leaf abaxial surface C Microlicia chamissoana, eglandu-
lar trichomes and stomata on the leaf abaxial surface D Microlicia aviora, stomata on the leaf abaxial
surface E, F Microlicia laniora: E eglandular woolly trichomes on the leaf abaxial surface F glandular
trichomes on the leaf adaxial surface. ET: Eglandular trichomes; GT: Glandular trichomes; ST: Stomata.
Voucher specimens: A Pacico & Bressan 380 (CAS, HUEM, SPF) B Pacico 290 (CAS, SPF) C Pacico
& Carmo 154 (HUEM, UEC) D Mello-Silva et al. 509 (SPF) E, F Almeda et al. 9179 (CAS, UEC).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 19
Inorescences found in Lavoisiereae always have a terminal ower, which means that
they are determinate, i.e. monotelic sensu Weberling (1988) and anthotelic sensu Briggs
and Johnson (1979). Overall, inorescences in the Trembleya s.s. clade are similar in
structure to those of the Lavoisiera clade, which were described in detail by Martins
and Almeda (2017). In both clades, the inorescences always consist of a proliferation
or reduction, based on a 3-owered dichasial unit, with opposite branching like that of
sterile branches (Fig. 8). e dichasial unit consists of a terminal ower subtended by
Figure 7. SEM images of leaves in the Trembleya s.s. clade of Microlicia A Microlicia parviora, abaxial
surface in frontal view B–D Microlicia pithyoides: B abaxial surface in frontal view C pore on the leaf
abaxial surface D glandular trichome on the leaf adaxial surface E Microlicia trembleyiformis, glandular tri-
chome on the abaxial surface F Microlicia tridentata, detail of a glandular trichome on the adaxial surface.
All photos by R. Pacico. Voucher specimens: A Almeda et al. 8483 (CAS, HUFU, NY) B, C Pacico 295
(CAS, HUEM, SPF) E Romero & Nakajima 3593 (HUFU, UEC) F Pacico 290 (CAS, HUEM, SPF).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
20
Figure 8. Inorescences in the Trembleya s.s. clade of Microlicia A M. altoparaisensis B M. chamis-
soana C M. laniora D–F M. aviora G M. parviora H M. tridentata. Photos: A by V. E. Bressan
B by F.A.O. Silveira C by L. Pedrosa D, E by R. Pacico F by A.V. Scatigna E by F. Almeda F by
F.A. Michelangeli.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 21
a pair of modied leaves (bracts). Two lateral branches originate from the axils of the
bracts to produce lateral owers, which are, in turn, subtended by a pair of bracteoles.
e proliferation of this pattern produces biparous cymes in a compound inores-
cence. In some cases, only one lateral branch develops from the axil of a bract. is
pattern is associated with production of uniparous cymes at the distal portion of the
inorescence and is here reported for a few species.
In the Trembleya s.s. clade, seven species have simple or compound dichasia
(M. altoparaisensis, M. calycina, M. chamissoana, M. aviora, M. laniora, M. parvi-
ora and M. tridentata) and the remaining four have inorescences reduced to soli-
tary owers at the apical region of the branches (M. pentagona, M. pithyoides, M. ros-
marinoides and M. trembleyiformis). Floral pedicels are usually evident and measure
0.3–4 mm long. ese are inconspicuous to 0.2 mm long only in M. altoparaisensis.
Like some species of the Lavoisiera clade (Martins and Almeda 2017), a tendency of
inorescence reduction to congested clusters occurs in M. chamissoana and M. laniora
(Fig. 8B–C). e degree of inorescence development, as well as the shape, size and
venation of the bracteoles, are characters of taxonomic signicance at the species level.
Hypanthium and calyx
e hypanthium that envelops the superior ovaries is campanulate to urceolate and its
length varies between 1.7 and 6.5 mm. e external surface is light green, reddish or
golden and has (8)10(12) longitudinal vascular ribs that terminate apically in a circular
ring where the hypanthium and calyx lobes meet. In Melastomataceae, this region is usu-
ally referred to as the torus (Wurdack 1953; Almeda 1978; Martins and Almeda 2017).
In two species (M. tridentata and M. pentagona), the hypanthium could be described
as glabrous, although it is minutely granulose and covered by a glutinous substance. In
four species, the external surface is only glandular-punctate (M. altoparaisensis, M. caly-
cina, M. pithyoides and M. rosmarinoides), while, in the other three, eglandular or gland-
tipped trichomes usually occur associated with the glandular-punctate indumentum (M.
chamissoana, M. parviora and M. trembleyiformis). An atypical indumentum is found in
M. laniora, which has the external surface of the hypanthium totally hidden by a dense
lanose indumentum of eglandular trichomes and in M. parviora, which has a hypan-
thium that is usually pruinose, eventually sparsely to densely covered with gland-tipped
trichomes or rarely covered with what appear to be rigid hyaline eglandular trichomes.
e calyx lobes that develop above the torus are united at the base and that un-
ion forms the calyx tube, which, in the Trembleya s.s. clade, varies in length between
0.1 and 0.7 mm. e calyx lobes are either oblong, triangular, narrowly triangular or
subulate, with the apex acute or acuminate and between 0.7 and 9.7 mm long. e
indumentum of both calyx tube and calyx lobes is usually similar to that of the hypan-
thium. Dierences in the shape, size and indumentum of the calyx lobes are generally
useful diagnostic characters at the species level in this clade.
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Corolla
Flowers are generally 5-merous (Fig. 9) with petals convolute in bud. Rarely, one or
a few 4-merous or 6-merous owers are found in inorescences predominantly with
5-merous owers. e corolla is always radially symmetrical at anthesis. e petals
are free, attached to the torus alternating with calyx lobes and opposing the smaller
set of stamens. ey vary between 4.5 and 26.0 mm in length, exceeding 20 mm
only in M. laniora. e shape is obovate with the apex rounded or acute, varying
in colour from white (M. altoparaisensis, M. laniora) to magenta (M. calycina, M.
chamissoana, M. pentagona, M. pithyoides, M. trembleyiformis, M. tridentata) or yellow
Figure 9. Diversity of owers in the Trembleya s.s. clade of Microlicia A M. chamissoana B M. parviora
C M. aviora D M. laniora E, F M. pentagona G M. rosmarinoides H M. tridentata. Photos: A by
F.A.O. Silveira B and D–F by F. Almeda C by A.V. Scatigna G by L. Pedrosa H by F.A. Michelangeli.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 23
(M. aviora, M. rosmarinoides) (Fig. 9). In M. parviora, both magenta and white pet-
als occur in distinct populations. ese petals are more commonly described as white
with pink stains. A rare form of M. lanifora with pink-ushed petals was once reported
from Serra do Cipó, Minas Gerais. Both surfaces of the petals are generally glabrous
and the margin varies from entire to glandular-punctate (M. chamissoana) or ciliate
(M. laniora, sometimes in M. parviora).
Androecium
Flowers are diplostemonous and, therefore, generally have ten stamens, which are
organised in an antesepalous whorl of larger stamens and an antepetalous whorl of
smaller stamens. In all but one species of the clade (M. altoparaisensis), the pedocon-
nectives of the antepetalous stamens are much shorter than those of the antepetalous
stamens, forming a yellow ventral appendage that measures about 0.1(–0.5) mm and
is often inconspicuous. e pedoconnectives of the antesepalous stamens are gen-
erally well-developed, forming yellow ventral appendages (0.1–)0.7–3.0 mm long
that are usually emarginate at the apex. In the majority of species, the antesepalous
stamens have purple to vinaceous anthers in strong contrast to the yellow anthers of
the antepetalous whorl. e exception is M. altoparaisensis, which has subisomor-
phic stamens with all anthers yellow. e laments are liform, glabrous, measuring
1.5–6.3 mm long, usually similar in colour to the petals. e anthers are oblong
(Fig. 10A, B), 0.7–3.8 mm in length (excluding rostra), with a smooth external sur-
face and 2-celled tetrasporangiate thecae. Each anther tapers to a rostrum measuring
0.2–0.5(–0.7) mm long with a ventrally inclined apical pore (Figs 10C–F, 11A–D).
A remarkable dierence between the two sets of stamens is that the anther pores
of the antepetalous whorl are always narrower than those of the antesepalous set.
is distinction is usually only evident when anthers are examined with Scanning
Electron Microscopy (Figs 10C–F, 11A–D). Overall, in this clade, the androecium
is radially symmetrical at bud and bilaterally symmetrical at anthesis (see Fig. 9). For
notes on reproductive strategies, see the topic “Reproductive Biology, Pollination
and Phenology.”
Pollen
Pollen grains in Melastomataceae are small, generally tricolporate, radially symmetri-
cal and isopolar (Patel et al. 1984; Renner 1993). Apparently of little taxonomic
use, these characters have rarely been described in taxonomic studies of the family
(Almeda and Robinson 2011; Alvear and Almeda 2019). Based on SEM photos of
four species of the Trembleya s.s. clade taken during this study (T. altoparaisensis, M.
calcyina, M. chamissoana and M. pithyoides), the pollen grain structure of the clade is
apparently not informative taxonomically and matches the descriptions provided in
the above-cited references. SEM images of pollen grains of M. pithyoides are presented
in Fig. 11E, F.
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24
Gynoecium and capsule
e ovary is always superior, glabrous and completely enclosed by the hypanthium.
Ovaries are 5-locular in most species, 3(–4)-locular only in M. altoparaisensis and vary
from 3–5-locular only in M. trembleyiformis. ese dierences in ovary locule number
Figure 10. SEM images of anthers and anther rostra in the Trembleya s.s. clade of Microlicia A Microlicia
parviora, anther of an antesepalous stamen B Microlicia tridentata, anther of antesepalous stamen
C Microlicia aviora, anther rostrum of an antesepalous stamen D Microlicia aviora, anther rostrum
of antepetalous stamen E Microlicia parviora, anther rostrum of an antesepalous stamen F Microlicia par-
viora, anther rostrum of an antepetalous stamen. Voucher specimens: A , E , F Pirani et al. CFSC 12361
(SPF) B Pacico & Bressan 290 (CAS, HUEM) C, D Hatschbach et al. 54239 (CAS, INPA, MBM).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 25
are additional characters that can be used for species recognition. e ovaries are ovoid
to globose or cylindrical, measuring between 0.9 and 4.1 mm in length. Placentation
is axile with numerous tiny anatropous ovules attached to subpeltate placental intru-
sions. e style is liform (3–10 mm long) and incurved distally to sigmoid at anthesis,
glabrous, similar in colour to the petals and the stigma is punctiform to truncate.
Figure 11. SEM images of anther rostra and pollen grains in the Trembleya s.s. clade of Microlicia
A Microlicia pentagona, anther rostrum of an antesepalous stamen B Microlicia pentagona, anther rostrum
of an antepetalous stamen C Microlicia tridentata, anther rostrum of an antesepalous stamen D Microlicia
tridentata, anther rostrum of an antepetalous stamen E, F Microlicia pithyoides, pollen grains. Voucher
specimens: A , B Irwin et al. 20486 (CAS, MO, NY, US) C, D Pacico & Bressan 290 (CAS, HUEM)
E, F Pacico 295 (CAS, HUEM, SPF).
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26
e brownish capsules are loculicidal measuring 2.3–8.0 mm in length, initially envel-
oped by the hypanthium and calyx that rupture and ake away tardily with age (Fig. 12).
e shape of the capsule is ovoid to globose. e enveloping hypanthium is little or
markedly constricted at the apex (torus). e capsules dehisce from the apex to the base
(basipetal) into 3–5 valves with a deciduous columella. Both calyx tubes (0.2–3.1 mm
long) and calyx lobes (1.2–11.5 mm long) may become elongated in fruit. e apex of
the mature capsule never exceeds the torus, like it does in several species of Microlicia s.s.
In fruit, the calyx lobes become sharply thicker only in M. pentagona (Fig. 12C–E).
Seed structure and germination
Seed micromorphological characters have been traditionally used in the tribal and
generic classication of Melastomataceae (Cogniaux 1883–1888; Baumgratz 1983;
Fritsch et al. 2004; Ocampo et al. 2014; Goldenberg et al. 2015; Penneys et al. 2022).
In fact, one of the synapomorphies attributed to the Lavoisiereae is the presence of el-
lipsoid to reniform seeds with a reticulate-foveolate testa (Almeda and Martins 2001;
Fritsch et al. 2004; Rocha et al. 2016; Martins and Almeda 2017; Pacico and Almeda
Figure 12. Capsules and infructescences in the Trembleya s.s. clade of Microlicia A Microlicia laniora,
infructescence B Microlicia parviora, capsule enveloped by the hypanthium C–E Microlicia pentagona:
C capsule enveloped by the hypanthium D infructescence E capsules enveloped by reddish mature hypan-
thia. Photos: A by F. Almeda B , C by F.A. Michelangeli E , F by R. Pacico.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 27
2022). e use of seed morphology to recognise species from closely-related species
groups is limited in Melastomataceae, although some investigations have revealed
promising seed characters that could be used together with macromorphology for spe-
cies circumscriptions (Ocampo and Almeda 2013; Martins and Almeda 2017; Ocam-
po et al. 2022). In the Trembleya s.s. clade, the six species for which mature seeds were
available are similar in shape and testa sculpturing and typical of Lavoisiereae (Fig. 13).
Figure 13. SEM images of seeds in the Trembleya s.s. clade of Microlicia A Microlicia calycina B Micro-
licia laniora C, D Microlicia parviora E Microlicia pentagona F Microlicia rosmarinoides. All photos by
R. Pacico. Voucher specimens: A Pacico & Bressan 296 (CAS, HUEM, SPF) B Almeda et al. 7726
(CAS, UEC) C , D Almeda et al. 8483 (CAS, HUFU, NY) E Irwin et al. 20178 (CAS, MO, NY, UEC,
US) F Occhioni et al. s.n. (US [US001900109]).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
28
Overall, they are slightly curved to reniform with the raphal zone oblong, occupying
ca. 30–70% of the total length of the seed (total length = 0.2–0.8 mm) and the testa
cells are arranged in an aligned pattern.
Information on seed germination in the clade is limited to studies of M. parviora
and M. laniora. According to Silveira et al. (2012), seeds of these two species are
non-dormant. For M. laniora, the optimum temperature range for germination is
20–25°C, which matches the onset of the rainy season (Rodrigues and Silveira 2013).
Seeds of M. laniora are viable after 42 months of storage (Rodrigues and Silveira 2013)
and for being easily buried, they are likely easily incorporated into soil seed banks
(Rodrigues and Silveira 2013). According to the investigations by Giotto (2015), seeds
of M. parviora usually take 5–14 days to germinate and germination occurs more
quickly when the seeds are exposed to variations in temperature. Germination is higher
for seeds that are stored for more than a month after collection, and they continue to
be viable for at least 15 months after being collected (Giotto 2015).
Reproductive biology and pollination
Flowers are hermaphroditic with a strongly dimorphic (rarely subisomorphic) androe-
cium and poricidal anthers. Plants of this clade provide pollen as a primary resource
to reward pollinators and apparently developed oral traits to protect the pollen, select
pollinating agents and precisely deposit pollen on the body of the pollinator. ese
characteristics are usually understood as strategies to deal with the “pollen dilemma”,
in which pollen grains full two main functions, i.e. to transport the male gamete and
to reward ower visitors with resource supply (Westerkamp 1996; Luo et al. 2008).
In Melastomataceae, nectar production is apparently restricted to a few genera (see
Varassin et al. 2008) and has never been reported in Lavoisiereae. During anthesis, the
large anthers are generally positioned almost parallel to each other, serving as a “land-
ing platform”, while the smaller anthers are more erect and positioned to serve as the
feeding set; a clustering of the yellow anthers of the antepetalous stamens, together
with the yellow connective appendages of the antesepalous stamens, apparently acts as
a visual sign to pollinators in most species. As usual in Melastomataceae (Renner 1989;
Dellinger et al. 2022), the pollen is collected by bees that perform buzz pollination.
e bees position themselves over the androecium and perform rapid distortions of the
thoracic box and produce a characteristic buzzing sound (Renner 1989).
By having owers with vibrant colours and diurnal anthesis, most species of Lavoi-
siereae seem to meet the requirements of the classic melittophily syndrome (Faegri and
van Der Pijl 1979). e exception is M. laniora, whose white owers are the largest
in the clade and open at night and dusk (Soares and Morellato 2018). e reproduc-
tive success of M. laniora seems to be highly dependent upon large bees of the genus
Xylocopa that perform crepuscular foraging (Soares and Morellato 2018). Flowers of
M. laniora are self-incompatible and depend on cross-pollination; this behaviour sug-
gests that pollinator-mediated interbreeding is a strategy to promote genetic variability
of populations naturally isolated on rocky outcrops (Soares and Morellato 2018).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 29
ere is no evidence of apomixis in M. laniora (Soares and Morellato 2018).
e widely-distributed M. parviora has been reported as a facultative apomict (Silva
2001) or non-apomictic (Maia et al. 2016) and self-compatible (Silva 2001; Maia
et al. 2016). Still, Maia et al. (2016) reported reduced speed of pollen tube growth
in self-pollination treatments. Both staminal whorls have fertile pollen (Santos et al.
2012). Functional dimorphism of stamens in M. parviora is still possible because
germination for each type of pollen has not been tested (Santos et al. 2012). Natural
vegetative reproduction, probably from rhizomes, was reported by Campos (2005)
and Baumgratz et al. (2007). e occasional apomictic behaviour of M. parviora is
consistent with the general pattern found in Melastomaceae, a family in which apom-
ictic species generally have wider distributions than non-apomictic taxa (Santos et al.
2012; Caetano and Oliveira 2022). Silva et al. (2012) reported that M. parviora is
one of the native plant species most visited by the pollinating bee Eulaema negrita in
an agroecosystem in Uberlândia (Minas Gerais), indicating that this species performs
signicant ecosystem services in that region.
Populations of Microlicia parviora from dierent localities may be distinct in phe-
nological behaviour, with annual or episodic owering and continuous or annual fruit-
ing; this species is commonly reported from both typical campo rupestre and gallery
forests (Brito et al. 2017). Overall, a tendency to produce owers during the dry season
and seed dispersal at the beginning of the rainy season has been reported in phenologi-
cal studies for shrubby Melastomataceae from campo rupestre (Le Stradic 2012) and
especially in M. laniora (Rodrigues and Silveira 2013; Soares and Morellato 2018).
is tendency is believed to be an adaptation for eective pollination and seed dispersal
(Salazar et al. 2011; Silveira et al. 2012; Soares and Morellato 2018). Our histograms
of owering and fruiting periods, based on herbarium specimens (Fig. 14), agree with
what has been described for M. laniora and suggest a similar tendency for M. calycina,
M. chamissoana and M. aviora. ree other species (M. altoparaisensis, M. pithyoides
and M. rosmarinoides) could have a distinct phenological pattern, owering and fruiting
mainly during the wet season (Fig. 14A, H, I), although eld observations are neces-
sary to conrm this hypothesis. Microlicia pentagona and M. trembleyiformis apparently
produce owers and fruits more evenly throughout the year (Fig. 14G, J). Additional
phenological studies of this clade and the Lavoisiereae, in general, are desirable.
Chemical characters
Knowledge of secondary metabolites of Lavoisiereae is largely restricted to the work by
Bomm-Patrício et al. (2001), who provided leaf avonoid proles for 33 species of the
tribe, including four species from the Trembleya s.s. clade (M. chamissoana, M. laniora, M.
parviora and M. pentagona). ey reported 116 avonoids for the tribe with a predomi-
nance of avonol glycosides. Species of the Trembleya s.s. clade do not form a coherent
group with respect to a avonoid prole. A predominance of avonols in M. laniora and
M. parviora resembles Microlicia s.s., while 6-methoxylation found in M. chamissoana
and M. pentagona resembles species of the Lavoisiera clade (Bomm-Patrício et al. 2001).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
30
Figure 14. Circular histograms of phenology for species of the Trembleya s.s. clade of Microlicia. Infor-
mation on owering and fruiting periods were obtained from herbarium specimens.
Information on histochemical tests is available only for M. parviora and M. lani-
ora. Regarding the rst, phenolic compounds were reported from the endodermis and
parenchyma (Somavilla and Graciano-Ribeiro 2011), histochemical tests performed
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 31
on the exodermis cells of the roots were positive for suberin and negative for lignin
(Somavilla and Graciano-Ribeiro 2012), oil droplets were found in the mesophyl of
the leaves (Somavilla and Graciano-Ribeiro 2011), coumarins, steroids, triterpenes,
avonoids and tannins were also reported, with the following major compounds pre-
sent in the volatile oils: α-terpineol, α-pinene, β-pinene, sabinene, acetoxyeudesman-
4-α-ol and 2,4a-8,8-tetramethyldecahydrocyclopropanaphtalene (Farias et al. 2018).
Alkaloids probably occur in M. parviora as well (Raauf 1996). In M. laniora, phe-
nolic compounds were reported from the leaf palisade parenchyma, collenchyma, vas-
cular bundle sheath (Silva et al. 2018), as well as saponins, triterpenes, tannins, avo-
noids (Ventura et al. 2007a) and a avanone (Ventura et al. 2007b).
Chromosome numbers
Only about 10% of the species of Melastomataceae are known cytologically, but some
patterns of chromosomal evolution are evident. Chromosome number stasis at the
diploid level and recurrent cycles of polyploidy and dysploidy are common (Almeda
2013; Almeda and Penneys 2022). e base number attributed to the family is x = 12
(Almeda 2013). Lavoisiereae is quite uniform with x = 12 and tetraploidy based on
that number, while Rhynchanthera diers in having x = 10 and tetraploidy on that base
number (Almeda and Penneys 2022). In the Trembleya s.s. clade, chromosome counts
are available only for two species. e diploid M. parviora has n = 12 and the tetra-
ploid M. pentagona has n = 24 (Almeda and Penneys 2022).
Geographic distribution, biogeography and endemism
e Trembleya s.s. clade is restricted to Brazil (Fig. 15). Both richness and weight en-
demism are concentrated in the Minas Gerais State (southern Cadeia do Espinhaço),
where 10 species occur (Fig. 16). e only two other States with more than one species
are Goiás (M. altoparaisensis and M. parviora) and Rio de Janeiro (M. parviora and
M. trembleyiformis). Microlicia parviora has the widest distribution in the clade and is
reported for the states of Bahia, Minas Gerais, Goiás, Distrito Federal, Espírito Santo,
Rio de Janeiro, São Paulo and Paraná.
Endemicity analyses detected one consensus area of endemism for the Tremb-
leya s.s. clade, which resulted from the merging of four individual areas of endemism
(Fig. 17). e consensus area consisted of ve cells and is a good match for the South-
ern Espinhaço biogeographic Province (ColliSilva et al. 2019), which is also a major
area of endemism for the Lavoisiereae (Pacico et al. 2020). e highest endemicity
score of the consensus area is 4.93 (at the southern region) and the lowest endemicity
score 3.30 (at the northern region). Microlicia calycina, M. chamissoana, M. laniora,
M. pentagona, M. rosmarinoides and M. tridentata were recovered as endemic species
and their endemicity scores are presented in Fig. 17.
e region with highest richness and weight endemism of the clade is the Iron
Quadrangle (Quadrilátero Ferrífero), located in the Municipalities of Barão dos Co-
cais, Belo Horizonte, Brumadinho, Caeté, Catas Altas, Congonhas, Itabirito, Mariana,
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
32
Figure 15. Geographic distribution of the Trembleya s.s. clade of Microlicia in Brazil.
Figure 16. Richness and weight endemism maps of the Trembleya s.s. clade of Microlicia A species rich-
ness map B species weight endemism map.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 33
Nova Lima, Ouro Branco, Ouro Preto, Raposos, Rio Acima, Sabará and Santa Bárbara
(Minas Gerais State). e Iron Quadrangle is one of the world’s largest mineral prov-
inces, consisting of rocks associated with the Archean and Paleoproterozoic periods
and includes mountain ranges, such as the Serra de Ouro Branco, Serra de Ouro Preto,
Serra da Moeda, Serra da Piedade, Serra do Gandarela, Sera do Garimpo, Serra do
Rola-Moça and Serra do Caraça (Ruchkys and Machado 2013). It has been recognised
as a distinct biogeographic district associated with campo rupestre in the Southern
Espinhaço Province (ColliSilva et al. 2019) and an area of endemism for the tribe
Lavoisiereae (Pacico et al. 2020). Eight species of the clade have been reported for
the Iron Quadrangle region: M. calycina, M. laniora, M. parviora, M. pentagona, M.
pithyoides, M. rosmarinoides, M. tridentata and M. trembleyiformis. Another relevant
endemism centre of the clade is the Serra do Cipó, where four narrow endemics can
be found (M. chamissoana, M. laniora, M. pentagona and M. tridentata). Only two
narrow endemics are not associated to the regions described above: Microlicia aviora,
which is restricted to northern Minas Gerais, in Grão Mogol, Botumirim and Rio
Pardo de Minas and M. altoparaisensis, restricted to Chapada dos Veadeiros, Goiás.
e richness and endemism of the Trembleya s.s. clade are concentrated in the
campo rupestre, a biodiversity-rich mosaic of mountaintop vegetation where about
15% of Brazilian vascular plant species occur in an area smaller than 1% of its ter-
ritory (Vasconcelos 2011; Alves et al. 2014; Fernandes 2016; Silveira et al. 2016).
Figure 17. e area of endemism of the Trembleya s.s. clade of Microlicia, based on Endemicity Analysis
A grid cells corresponding to the Southern Espinhaço area of endemism B the Southern Espinhaço area
of endemism manually drawn, based on distribution of species of Trembleya s.s. clade of Microlicia.
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
34
In Brazil, this azonal peinobiome is found mainly on the Cadeia do Espinhaço
(also referred to as “Espinhaço Range”), along the States of Bahia and Minas Ger-
ais and in the Brazilian Central Plateau in the State of Goiás (Giulietti et al. 1997;
Vasconcelos 2011; Alves et al. 2014). Its vegetation occurs mostly on mountains
from about 900–2000m, on lithosols associated with outcrops of Precambrian
quartzites, sandstones and ironstones (Giulietti et al. 1997; Vasconcelos 2011;
Alves et al. 2014; Silveira et al. 2016). All species of the Trembleya s.s. clade occur
on quartzitic soils (for example, at the Serra do Caraça). ree species were also
reported from ironstone outcrops known as cangas (M. laniora, M. rosmarinoides
and M. tridentata). Some species have non-overlapping elevational ranges, suggest-
ing an elevated specicity in their habitat preferences, for example, M. chamissoana
(1154–1462 m), M. rosmarinoides (1609–1807 m) and M. pithyoides (1827–2072
m). e most widely distributed species, M. parviora, also has the widest eleva-
tional range in the clade (560–2223 m), extending into the Cerrado Biome in
savannahs and gallery forests.
e traditional hypothesis for plant diversication in campo rupestre postulates
that repeated retraction-expansion events driven by Pleistocene climatic changes
would have worked as an evolutionary pump, creating a scenario where mountaintops
functioned as long-term refuges and maintained lineages despite climatic alterations
(Conceição et al. 2016). In this context, populations isolated on mountaintops would
have been susceptible to genetic drift and inbreeding depression, which would have fa-
voured speciation (Rapini et al. 2008). e evolutionary pump hypothesis has been re-
cently challenged by Rapini et al. (2020), who argue that the long-term fragmentation,
combined with recurrent extinctions and sporadic events of adaptive radiation, may
provide a better explanation for the current diversity and endemism in campo rupestre,
in accordance with the “escape-to-radiate model”. Another non-excluded hypothesis is
that mountains with campo rupestre are old climatically-buered infertile landscapes
(OCBILs) (Hopper 2009; Silveira et al. 2016; Zappi et al. 2017), consisting of both
a cradle of continuing diversication and a museum of ancient lineages (Silveira et al.
2016; Zappi et al. 2017; Vasconcelos et al. 2020).
In situ diversication apparently played an important role during the evolu-
tionary history of the Trembleya s.s. clade, along with several other plant lineages
(Fritsch et al. 2004; Pacico et al. 2020; Vasconcelos et al. 2020). Major diversica-
tion of Microlicia s.l. may have occurred during the Pliocene (Fritsch et al. 2004)
associated with the acquisition of xeromorphic features, for example, re-resistant
woody xylopodia (Simon et al. 2009), physiologically dormant seeds (Silveira et
al. 2012), petiole shortening, stomatal crypts (Fig. 5D), reduction in leaf blade
area, densication of palisade parenchyma, thickening of the outer periclinal wall of
the epidermal cells (Carmo et al. 2020) and aluminium accumulation mechanisms
(Jansen et al. 2002). ese are all possible innovations which could have favoured
a shift from mesophytic to xerophytic environments and the occupation of the
diverse microhabitats found in campo rupestre (Fritsch et al. 2004; Silveira et al.
2012; Carmo et al. 2020).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 35
Conservation
Overall, vascular plant lineages restricted to campo rupestre are highly vulnerable to
disturbances and their conservation deserves special attention (Conceição et al. 2016;
Silveira et al. 2016). Given their high phylogenetic conservatism, these lineages are not
expected to adapt quickly to new conditions and are usually unable to colonise new
habitats (Conceição et al. 2016; Silveira et al. 2016). Additionally, most of them would
be very unlikely to reach distant suitable areas because of their low dispersal capability
(Conceição et al. 2016; Silveira et al. 2016).
e historical stability of campo rupestre vegetation in the face of climatic changes
is still under debate (Antonelli et al. 2010; Bitencourt et al. 2016; Conceição et al.
2016; Rapini et al. 2020). According to the OCBIL theory, mountaintops with campo
rupestre have been buered for a long time from climatic alterations and rapid changes
could greatly reduce their biodiversity (Hopper 2009; Silveira et al. 2016). In global
warming scenarios, Bitencourt et al. (2016) estimated that the Espinhaço Range may
lose up to 56% of its plant diversity and 97% of microendemic species by 2080.
Most species of the Trembleya s.s. clade have narrow distributions limited to one or
a few mountains and several of them also have narrow elevational ranges. e Brazilian
Government already recognises three species of this clade as Endangered (EN), i.e. M.
calycina, M. chamissoanai and M. aviora and one as Critically Endangered (CR), M.
pithyoides (Brasília 2014). In this study, based on AOO and EOO values and criterion
B of IUCN (2019), our informal conservation status recommendations agree with
those assigned to these species by the Brazilian Government. We also recommend that
four additional species should be recognised as Endangered (EN), M. altoparaisensis,
M. pentagona, M. rosmarinoides and M. tridentata, M. laniora should be considered
as Vulnerable (VU) and M. parviora and M. trembleyiformis as Least Concern (LC).
Systematic treatment
Trembleya s.s. clade of Microlicia s.l.
Diagnosis. Perennial shrubs or treelets. Leaves petiolate, not imbricate, not keeled,
the adaxial surface glandular-punctate to glabrescent, venation basal acrodromous, im-
pressed on the adaxial surface and prominent on the abaxial surface, consisting of am-
phicribral or bicollateral vascular bundles. Flowers usually 5-merous, diplostemonous,
pedicellate, subtended by a pair of bracteoles. Hypanthia not fused to the ovary, lack-
ing a crown of trichomes at the apex. Stamens strongly dimorphic or subisomorphic,
anthers 2-celled, tetrasporagiate. Ovaries superior, (3–4–)5-locular. Capsules dehiscent
from the apex to the base, columella deciduous.
Description. Perennial erect shrubs or small trees (0.1–)0.3–4 m tall, woody, some-
times densely branched. Distal branches quadrangular, usually light green (when fresh)
and glutinous, glandular-punctate, sometimes granulose or pruinose, eventually covered
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
36
with eglandular or gland-tipped trichomes, internodes 0.1–4.5 cm long, angles unwinged
or narrowly winged, nodes thickened. Old branches terete, brownish and defoliating to-
wards the base. Leaves decussate, petiolate, not imbricate, not keeled, papyraceous, chart-
aceaous or coriaceous, usually discoloured when dry. Petioles 0.3–17 mm long. Blades
0.4–11.7 cm long, 0.05–5 cm wide, oblong, lanceolate, elliptic, narrowly elliptic, ovate
or linear, entire to slightly serrulate, sometimes entire along the basal half and serrulate
on the upper half, rarely ciliate, lacking support tissue on the leaf margin. Adaxial surface
green (when fresh), becoming pale green, pale brown, or darkened (when dry), glutinu-
ous, glandular-punctate to glabrescent, glandular trichomes (when present) appearing
sessile (i.e. on peduncles too short to be seen with a 40× magnication stereoscope).
Abaxial surface usually green (when fresh), becoming pale green (when dry), always light-
er than the adaxial surface, glandular-punctate to covered with eglandular or gland-tipped
trichomes, or totally concealed by a lanose indumentum. Venation composed of 1–7
basal acrodromous veins, mid-vein stout, lateral veins becoming faint towards the leaf
margin, impressed on the adaxial surface and prominent on the abaxial surface, consisting
of amphicribral or bicollateral vascular bundles, tertiaries usually evident, nearly perpen-
dicular to the mid-vein and branching towards the leaf margin. Inorescences simple or
compound dichasia, consisting of biparous cymes throughout or proximally biparous and
distally uniparous cymes or reduced to solitary owers on the apical region of the branch-
es. Inorescence bracts 0.7–5.0 cm long, 0.1–2.0 cm wide, petiolate, usually similar to
the principal leaves in shape and indumentum, 1–5-nerved from the base. Bracteoles
sessile or with petioles up to 6 mm long, blades 2.2–11 mm long, 0.5–5.5 mm wide, lin-
ear, elliptic, lanceolate, ovate, narrowly elliptic, oblong or oblanceolate, 1–3(–5)-nerved
from the base, entire to slightly serrulate, rarely ciliate, usually diering in shape, but
similar in indumentum to the principal leaves. Flowers (4–)5(–6)-merous, diplostemon-
ous, pedicellate, subtended by a pair of bracteoles. Hypanthia 1.7–6.5 mm long, 1.5–
5.2mm wide at the torus, campanulate to urceolate, not fused to the ovary, externally
glandular-punctate, sometimes covered with eglandular or gland-tipped trichomes, rarely
completely concealed by a lanose indumentum, lacking a crown of trichomes at the apex.
Calyx tubes 0.1–1.2 mm long, externally like the hypanthia. Calyx lobes 0.7–9.7mm
long, 0.4–3.2 mm wide at the base, oblong, triangular, narrowly triangular or subulate,
entire or rarely sparsely ciliate, apex acute to acuminate, rarely terminating in an apical
eglandular trichome, similar to the hypanthia in indumentum. Petals 4.5–26mm long,
2.4–15 mm wide, obovate, entire, eventually ciliate, white, magenta or yellow (when
fresh), apex acute, rounded, acuminate or emarginate, both surfaces glabrous or rarely
sparsely glandular-punctate on the adaxial surface. Stamens (8)10(12), strongly dimor-
phic or subisomorphic, glabrous throughout, laments linear, white, pink or yellow, pe-
doconnectives well-developed, anthers oblong, 2-celled (tetrasporangiate), rostrate, apical
pores circular and ventrally inclined. Larger (antesepalous) stamens (4–)5(–6), laments
1.5–6.3 mm long, pedoconnectives 1.3–7.3mm long, ventral appendages 0.1–3.0 mm
long, the apex usually emarginate to bilobate, thecae (excluding rostra) 0.8–3.8 mm long,
purple, red, vinaceous or rarely yellow, rostra 0.2–0.7 mm long, pores 0.1–0.3 mm wide.
Smaller (antepetalous) stamens (4–)5(–6), laments 1.5–5.4 mm long, pedoconnectives
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 37
0.2–1.9 mm long, ventral appendages usually up to 0.1 mm long, apex truncate to emar-
ginate, thecae (excluding rostra) 0.8–3.2 mm long, yellow to orange, rostra 0.2–0.6 mm
long, pores ca. 0.2 mm wide. Ovaries 0.9–4.1 mm long, 0.7–3.1 mm wide, ovoid, cy-
lindrical or globose, superior, (3–4–)5-locular, glabrous. Styles 3–10 mm long, liform,
sigmoid to incurved, white, pink or yellow, glabrous, stigmas punctiform. Capsules loc-
ulicidal, 2.3–8.0 mm long, 2.3–6.0 mm wide, ovoid or globose, the torus initially con-
stricted at the apex, dehiscent from the apex to the base, columella deciduous. Fruiting
calyx tubes 0.2–3.1mm long. Fruiting calyx lobes 1.2–11.5 mm long, rarely thickened.
Seeds 0.3–0.9 mm long, reniform, the testa foveolate-reticulate.
Nomenclatural notes
Based on collections housed in the herbarium at Paris (P), Martin and Cremers (2007)
selected types for Melastomataceae described by Charles Victor Naudin (1815–1889),
including six names for taxa that are part of the Trembleya s.s. clade (Microlicia calyci-
na, M. paniculata, M. pentagona, M. stenophylla, M. trembleyiformis and M. tridentata).
More than one specimen corresponding to the types cited by Martin and Cremers
(2007) were found at P for the six names. ese type citations were here treated as
rst-step lectotypications. us, second-step lectotypes were designated in these cases
following Article 9.17 of the Shenzhen Code (Turland et al. 2018). Overall, our no-
menclatural update of this clade includes the designation of 34 lectotypes for names at
specic and infraspecic ranks and 25 new synonyms.
Key to the species of Trembleya s.s. clade of Microlicia
1 Leaves 0.5–2 mm wide, 1-nerved from the base .............................................. 2
– Leaves 3–50 mm wide, 3–7-nerved from the base ........................................... 3
2 Leaf blades linear; petals magenta; larger (antesepalous) stamens with anthers
magenta to purple and smaller (antepetalous) stamens with yellow anthers .......
................................................................................................... M. pithyoides
– Leaf blades narrowly-lanceolate to narrowly-elliptic; petals yellow; all stamens
with anthers yellow to pale brown ........................................ M. rosmarinoides
3 Plants with a dense whitish lanose indumentum covering branchlets, leaf abaxial
surfaces and hypanthia; petals 19–26 mm long ............................. M. laniora
– Plants lacking a whitish lanose indumentum; petals 4.5–13 mm long ............. 4
4 Leaves with the abaxial surface entirely concealed by the indumentum, tertiary
venation “foveolate-like” (Fig. 4M) .........................................M. chamissoana
– Leaves with the abaxial surface exposed, tertiary venation (if present) not “fove-
olate-like”........................................................................................................5
5 Leaves 3-nerved from the base (including a tenuous inframarginal pair of
veins) ............................................................................................. M. calycina
– Leaves 5–7-nerved from the base (including a tenuous inframarginal pair of
veins) ..............................................................................................................6
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
38
6 Abaxial leaf venation with tertiaries absent or little evident (Fig. 4B–E) ..........7
– Abaxial leaf venation with tertiaries evident (Fig. 4A, H–M) ...........................8
7 Calyx lobes subulate, 6.2–8.5 mm long in ower, becoming stout and thick
in fruit (Fig. 12C–E); petals magenta; stamens strongly dimorphic (Minas
Gerais) ........................................................................................M. pentagona
– Calyx lobes oblong to triangular, 2.8–3.4 mm long in ower, tenuous in fruit;
petals white; stamens subisomorphic (Chapada dos Veadeiros, Goiás) ...............
........................................................................................... M. altoparaisensis
8 Leaf margins entire along the basal half, sharply serrulate on the upper half
(Fig. 4A), tertiary veins surrounding stout depressions on the abaxial leaf sur-
face; bracteoles 8.1–11.0 mm long .............................................. M. tridentata
– Leaf margins entire or slightly serrulate throughout, tertiary veins not surround-
ing stout depressions on the abaxial surface; bracteoles 2.2–6.1 mm long .......9
9 Calyx lobes 4–4.9 mm long; petals yellow; all anthers yellow to pale brown ......
.................................................................................................... M. aviora
– Calyx lobes 0.7–3 mm long; petals white, light pink or magenta; anthers of
larger (antesepalous) stamens pink to magenta .............................................. 10
10 Branchlets with narrow wings ca. 0.2 mm wide; owers solitary; calyx lobes
narrowly-triangular; ovaries 3–5-locular .............................M. trembleyiformis
– Branchlets unwinged; owers disposed in compound or simple dichasia; calyx
lobes triangular; ovaries 5-locular ................................................ M. parviora
1. Microlicia altoparaisensis (R.B.Pacico, Almeda & Fidanza) Versiane &
R.Romero, Bot. J. Linn. Soc. 197: 52. 2021.
Fig. 18
Trembleya altoparaisensis R.B.Pacico, Almeda & Fidanza, Phytotaxa 391(5): 291.
2019. basionym. Type: Brazil. Goiás, Alto Paraíso de Goiás, Parque Nacional
Chapada dos Veadeiros, próximo da cachoeira do Rio Preto, perto do povoado de
São Jorge, 6 February 1987, J.R. Pirani 1663, R.M. Harley, B.L. Stannard, A. Fur-
lan & C. Kameyama (holotype: SPF!; isotypes: HUEM!, K!, UEC!, US!).
Description. Erect shrubs 0.8–1.8 m tall. Branchlets quadrangular, glandular-punctate,
light green to golden (when fresh). Internodes 1.0–2.1 cm long, angles unwinged. Peti-
oles 2.0–5.5 mm long. Leaf blades 14–45 mm long, 3–9 mm wide, papyraceous (when
dry), oblong to lanceolate, both surfaces green (when fresh), pale green (when dry), con-
colorous (when dry), base attenuate, apex rounded to acute, margin at or slightly revo-
lute, slightly serrulate and glandular-punctate, 5-nerved from the base, one pair of acro-
dromous veins and one pair of tenuous veins close to the margin, tertiaries little or not
evident on the abaxial surface, adaxial surface densely glandular-punctate, abaxial surface
densely glandular-punctate. Inorescences compound dichasia consisting of proximally
biparous, distally uniparous cymes, not congested. Bracts (including petioles) 1.0–4.0 cm
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 39
long, 0.1–0.7 cm wide, 1-nerved, oblong to lanceolate, indumentum like that of the
principal leaves. Bracteoles (at anthesis) sessile or with petioles up to 0.1–0.8 mm long,
blades 5.0–6.7 mm long, 0.9–1.8 mm wide, oblong to oblanceolate, base acute, apex
rounded to emarginate, margin entire and glandular-punctate, 1-nerved, indumentum
like that of the principal leaves. Flowers (4–)5-merous, pedicels (at anthesis) inconspicu-
ous up to 0.2 mm long. Hypanthia (at anthesis) 2.5–5.5 mm long, 3.1–3.2 mm wide at
the torus, campanulate, light green to golden (when fresh), externally glandular-punc-
tate. Calyx tubes 0.1–0.3 mm long. Calyx lobes (at anthesis) 2.8–3.4 mm long, 3.1–3.2
mm wide at the base, oblong to triangular, apex acute, eventually terminating in an
eglandular trichome 0.1–0.4 mm long, margin entire, (when fresh) light green to golden,
externally glandular-punctate. Petals 8.1–10.2 mm long, 3.2–4.5 mm wide, white, obo-
vate, apex acuminate, margin entire and glabrous, both surfaces glabrous. Stamens (8)10,
subisomorphic. Larger (antesepalous) stamens (4–)5, laments 3.3–3.5 mm long, white,
pedoconnectives 1.3–1.5 mm long, white, appendages ca. 0.1 mm long, white, apex
emarginate, thecae (excluding rostra) 2.7–3.8 mm long, yellow, oblong, rostra 0.3–0.5
mm long, the circular pores ca. 0.2 mm wide. Smaller (antepetalous) stamens (4–)5,
laments 2.7–2.8 mm long, white, pedoconnectives 0.7–1.0 mm long, white, incon-
spicuous appendages ca. 0.1 mm long, white, apex truncate, thecae (excluding rostra)
2.5–3.2mm long, yellow, oblong, rostra 0.3–0.5 mm long, the circular pores ca. 0.2 mm
wide. Ovary 2.4–2.5 mm long, 1.4–1.6 mm wide, cylindrical, 3(–4)-locular. Style ca.
10 mm long, white. Capsules (at maturity) 4.5–5.3 mm long, 2.8–3.2 mm wide, ovoid,
initially enveloped by the hypanthium, torus constricted at the apex, fruiting calyx tubes
0.4–0.5 mm long, fruiting calyx lobes 3.2–3.4 mm long, not thickened. Seeds not seen.
Recognition and anities. Microlicia altoparaisensis can be recognised by its ob-
long-lanceolate leaf blades (1.4–4.5 × 0.3–0.9 cm), papyraceous, 3–5-nerved from the
base, glandular-punctate on both surfaces, tertiary veins not evident, inorescences
composed of proximally biparous and distally uniparous cymes, white petals, yellow
subisomorphic stamens and 3(–4)-locular ovaries. e subisomorphic androecium is
the most distinctive feature of M. altoparaisensis as it is unique in the clade. Microlicia
altoparaisensis is also the only species with amphistomatic leaves, attenuate thecae rostra
and a distribution restricted to Goiás (Pacico et al. 2019). Ovaries with 3–4 locules
occur only in the apparently distantly related M. trembleyiformis, which diers in hav-
ing leaves with tertiaries evident (vs. not evident), discoloured when dry (vs. concolor-
ous), solitary owers (vs. developed inorescences), petals magenta (vs. white) and
dimorphic stamens (vs. subisomorphic). Another possible relative is the widespread
M. parviora with a range that extends from Paraná to Bahia and Goiás and the only
representative of the clade that grows with M. altoparaisensis at the Chapada dos Vea-
deiros. Microlicia altoparaisensis diers from M. parviora by the leaves with tertiaries
not evident (vs. evident), concolorous when dry (vs. discoloured), amphistomatic (vs.
hipostomatic), inorescences with distally uniparous cymes (vs. biparous cymes), subi-
somorphic stamens (vs. dimorphic) and 5-locular ovaries (vs. 3–4-locular).
Distribution, habitat, and elevation range. Probably endemic to Chapada dos
Veadeiros in Alto Paraíso de Goiás Municipality, Goiás, Brazil (Fig. 19A). It occurs in
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
40
Figure 18. Microlicia altoparaisensis A habit B detail of the glandular-punctate indumentum on the
branches C leaf abaxial surface D bracteole adaxial surface E oral bud F ower in lateral view G petal
adaxial surface H antepetalous (left) and antesepalous (right) stamens. Drawn from Pirani et al. 1663
(UEC). Copyright Magnolia Press. Reproduced with permission from copyright holder.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 41
transitional formations between Cerrado and campo rupestre, on rocky quartzitic soils
partially exposed to sun, at elevations of 1043–1086 m.
Conservation. Microlicia altoparaisensis is known from less than 10 collections.
e EOO is 7.025 km2 and the AOO is 16 km2. Based on IUCN (2019) recommenda-
tions and criteria, we believe that this species should be classied as Endangered (EN):
B1ab(iii). e majority of the populations of M. altoparaisensis occur inside Parque
Nacional da Chapada dos Veadeiros, where this species is aorded some protection.
Notes. e collection Glaziou 21300 was listed under the name Trembleya debilis
by Glaziou (1908: 250). Following the proposal by Mansano and Pederneiras (2016),
the catalogue of collections by Glaziou (1908) was included in the List of Suppressed
Works for all taxonomic ranks in the Shenzhen Code (Turland et al. 2018). New
names at specied ranks included in publications listed as suppressed works are not
validly published according to Article 34 of the Code.
Specimens examined. Brazil. Goiás: Alto Paraíso de Goiás Municipality, Chapa-
da dos Veadeiros, Drummond et al. 321 (MBM, NY, RB), Klein et al. 2465 (HUFU,
UFG), Machado et al. 153 (HUFU), Meyer 1171 (UEC, UPCB), Pacico & Bressan
380 (CAS, HUEM, SPF), Pirani et al. 1663 (holotype: SPF; isotypes: HUEM, K,
UEC, US), Pirani et al. 1694 (K, SPF, UEC); Unknown municipality, Fazenda da Boa-
Vista, près Morro do Salto, Glaziou 21300 (F[photo], P, S).
Figure 19. Geographic distribution of species of the Trembleya s.s. clade of Microlicia A distributions
of M. altoparaisensis, M. calycina, M. chamissoana and M. aviora B distributions of M. laniora and
M. trembleyiformis.
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
42
2. Microlicia calycina (Cham.) Versiane & R.Romero, Bot. J. Linn. Soc. 197: 52. 2021.
Fig. 20
Trembleya calycina Cham., Linnaea 9(4): 430. 1835. basionym. Type: Brazil. “Bra-
silia, Itacolumi” [Minas Gerais, Ouro Preto], F. Sellow s.n. (lectotype, designated
here: K [K00530658]!; isolectotypes: BR [BR0000005227020]!, F [neg. 16634]!,
K [K00530659]!; image of lectotype is available at http://specimens.kew.org/her-
barium/K000530659).
Trembleya revoluta Naudin, Ann. Sci. Nat., Bot. Sér. 3, 2: 155. 1844. Type: Brazil.
“Minas Gerais, Capanema” [Minas Gerais, Santa Bárbara], 1841, P. Claussen 10
(lectotype, rst-step designated by Martin and Cremers (2007), second-step des-
ignated here: P [P00723384]!; isolectotypes: G [G00368001]!, P [P00723507]!;
image of lectotype is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/
p00723384).
Trembleya stenophylla Naudin, Ann. Sci. Nat., Bot. Sér. 3, 12: 265. 1849. syn. nov.
Type: Brazil. “Minas Gerais, Capanema” [Minas Gerais, Santa Bárbara], 1843,
P. Claussen 368 (lectotype, rst-step designated by Martin and Cremers (2007),
second-step designated here: P [P00723385]!; isolectotype: P [P00723386]!; im-
age of lectotype is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/
p00723385).
Description. Erect shrubs 0.65–1.5 m tall. Branchlets quadrangular, glandular-
punctate and sparsely covered with glandular trichomes 0.1–0.2 mm long, light green
(when fresh). Internodes 1.5–3.0 cm long, angles with narrow wings 0.2–0.4 mm
wide. Petioles 0.8–1.8 mm long. Leaf blades 10–26 mm long, 2–9 mm wide, charta-
ceous (when dry), elliptic to narrowly elliptic, both surfaces green (when fresh), adaxial
surface blackened and abaxial surface pale green (when dry), discoloured (when dry),
base attenuate, apex rounded to acute, margin revolute, entire along the basal half,
appearing entire to slightly serrulate on the upper half and minutely granulose and be-
coming glabrescent with age, 3-nerved from the base, one tenuous pair of acrodromal
veins, tertiaries evident on the abaxial surface, nearly perpendicular to the mid-vein,
little reticulate and branching apically, foveolate-like, adaxial surface sparsely glandu-
lar-punctate, appearing glabrous when dry, abaxial surface densely glandular-punctate.
Inorescences simple dichasia or reduced to solitary owers, not congested. Bracts
(including petioles) 1.0–1.3 cm long, 0.3–0.4 cm wide, 3-nerved, elliptic to narrowly
elliptic, indumentum like that of the principal leaves. Bracteoles (at anthesis) sessile
or with petioles up to 1.0 mm long, blades 4.0–5.0 mm long, 1.3–1.8 mm wide, el-
liptic, base attenuate, apex rounded to acute, margin entire and glandular-punctate,
1-nerved, indumentum like that of the principal leaves. Flowers 5-merous, pedicels (at
anthesis) 0.5–0.7 mm long. Hypanthia (at anthesis) 2.6–3.5 mm long, 1.9–2.2mm
wide at the torus, campanulate, light green or reddish (when fresh), externally glan-
dular-punctate. Calyx tubes 0.4–0.7 mm long. Calyx lobes (at anthesis) 3.5–5.2 mm
long, 0.5–0.7mm wide at the base, subulate, apex acuminate, margin entire, (when
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 43
Figure 20. Microlicia calycina A habit B leaf abaxial surface C bracteole abaxial surface D oral
bud E ower in lateral view F owering hypanthium G petal adaxial surface H antesepalous stamen
I antepetalous stamen J gynoecium K capsule. Drawn from Barreto 9019 (UEC).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
44
fresh) light green or reddish, externally glandular-punctate. Petals 7.8–10 mm long,
5.0–6.2 mm wide, magenta, obovate, apex acuminate, margin entire and glabrous,
both surfaces glabrous. Stamens 10, strongly dimorphic. Larger (antesepalous) stamens
5, laments 4.7–4.9mm long, pink, pedoconnectives 5.9–6.2 mm long, pink, append-
ages 1.4–1.6 mm long, yellow, apex truncate to slightly emarginate, thecae (excluding
rostra) 2.3–2.6 mm long, purple, oblong, rostra 0.4–0.7 mm long, the circular pores
ca. 0.2 mm wide. Smaller (antepetalous) stamens 5, laments 3.8–4.1mm long, pink,
pedoconnectives 0.6–0.8 mm long, yellow, inconspicuous appendages ca. 0.1 mm
long, yellow, apex truncate, thecae (excluding rostra) 2.1–2.3 mm long, yellow, ob-
long, rostra 0.4–0.6 mm long, the circular pores ca. 0.2 mm wide. Ovary 2.0–2.2mm
long, 1.9–2.1 mm wide, globose, 5-locular. Style ca. 6.5 mm long, pink. Capsules (at
maturity) 2.3–2.7 mm long, 2.3–2.7 mm wide, globose, initially enveloped by the hy-
panthium, torus constricted at the apex, fruiting calyx tubes 0.7–0.8 mm long, fruiting
calyx lobes 5.5–6.0 mm long, not thickened. Seeds ca. 0.8 mm long, reniform.
Recognition and anities. Microlicia calycina may be recognised by its elliptic
to narrowly elliptic leaf blades with revolute margins, 3-nerved from the base, simple
dichasia or solitary owers and subulate calyx lobes 3.5–5.2 mm long. In morphology,
M. calycina resembles narrow-leaved forms of M. pentagona (see notes under this spe-
cies). Both species share the glandular-punctate indumentum, inorescences reduced
to solitary owers (sometimes perfect dichasia only in M. calycina), subulate calyx
lobes, magenta petals, and bicoloured anthers. Microlicia calycina diers by the leaf
blades that are 3-nerved from the base (vs. 5-nerved) and calyx lobes 3.5–5.2 mm long
(vs. 6.2–8.5 mm long) that become thick in fruit (vs. tenuous). ese two species may
occur sympatrically in the seasonally dry grasslands of Parque Estadual do Itacolomi
(Rolim 2011) and at Serra do Caraça. Microlicia parviora is also morphologically
similar to M. calycina, which may be distinguished by leaves 3-nerved from the base
(vs. 5-nerved) and subulate calyx lobes (vs. triangular). Additionally, M. parviora is
distinct in having openly ramied inorescences that are sometimes reduced to sim-
ple dichasia. In turn, the inorescences of M. calycina consist of simple dichasia that
are frequently reduced to solitary owers. e distributions of Microlicia calycina and
M. parviora overlap in Caeté, Ouro Preto and Catas Altas, where sympatry is likely to
occur. Microlicia calycina is also similar to M. pithyoides (see notes under this species).
Distribution, habitat and elevation range. Endemic to central and southern Mi-
nas Gerais (Fig. 19A), at Serra do Caraça, Serra do Itacolomi and Serra da Piedade.
It occurs on quartzitic campo rupestre exposed to full sun at elevations between 1692
and 1920 m.
Conservation. Microlicia calycina is known from about 20 collections. e EOO
is 481 km2 and the AOO is 20 km2. is species is currently recognised as endangered
(EN) by the Brazilian Government (Brasília 2014). Based on IUCN (2019) recom-
mendations and criteria, we recommend a similar assessment: (EN): B1ab(iii). e
conservation units of Parque Estadual do Itacolomi and RPPN Serra do Caraça (Natu-
ral Heritage Private Reserve) are of major importance for the long-term conservation
of M. calycina.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 45
Notes. We agree with Cogniaux (1883–1888) and treat Trembleya revoluta as a
synonym of M. calycina. e type of Trembleya revoluta (P. Claussen 10) diers from
typical M. calycina collections only by its revolute leaf margins, a feature that appears
to be an artefact of drying. Likewise, we consider Trembleya stenophylla a narrow-leaved
form of M. calycina. Specimens of M. calycina with leaf blades conspicuously revo-
lute have also been confused with M. pithyoides. For a comparison of M. calycina and
M. pithyoides, see the notes under the latter.
Specimens examined. Brazil. Minas Gerais: Caeté Municipality, Serra da Pie-
dade, Grandi et al. 6593 (BHCB, HUFU); Catas Altas Municipality, Serra do Caraça,
Castro et al. 283 (HUFU), Oliveira & Giacomin 47 (BHCB), Oliveira & Giacomin
84 (BHCB), Oliveira et al. 480 (BHCB), Pacico & Bressan 296 (CAS, HUEM, SPF);
Ouro Preto Municipality, Serra do Itacolomi, Barreto 9019 (BHCB, ESA, FUEL,
HUFU, SP, SPF, UEC, UPCB), Damazio s.n. (RB [48391]), Glaziou 14745 (P),
Glaziou 18232 (K, P, R), Pacico & Bressan 291 (CAS, HUEM, SPF), Peron 220 (RB),
Peron 268 (RB), Peron 269 (RB), Riedel s.n. (K [K00530657], NY [NY00941982],
W [18890019737]), Rolim 366 (HUFU, NY, RB, VIC), Schwacke 9368 (RB, W);
Unknown municipality, Claussen 10 (P [P00723384, P00723507]), “capanema”,
Claussen 368 (P [P00723385, P00723386]), Sellow s.n. (lectotype: K [K00530659];
isolectotypes: BR [BR0000005227020], F [neg. 16634], K [K00530658]).
3. Microlicia chamissoana (Naudin) Versiane & R.Romero, Bot. J. Linn. Soc. 197:
53. 2021.
Fig. 21
Trembleya chamissoana Naudin, Ann. Sci. Nat., Bot. Sér. 3, 12: 270. 1849. basionym.
Type: Brazil. “Brasilia, Itambé” [Minas Gerais, probably Santo Antônio do Itam-
bé], F. Sellow s.n. [b. 1171 c. 1156] (lectotype, designated here: K [K00530656]!;
isolectotype: P [P00723508]!; image of lectotype is available at http://specimens.
kew.org/herbarium/K000530656).
Description. Erect shrubs (0.1–)0.3–1.5 m tall. Branchlets quadrangular, glandular-
punctate and covered with gland-tipped trichomes 0.2–0.4 mm long, light green
(when fresh). Internodes 0.3–1.5 cm long, angles unwinged. Petioles 1.0–4.9 mm
long. Leaf blades 10–28 mm long, 4–18 mm wide, chartaceous (when dry), ellipti-
cal, both surfaces green (when fresh), adaxial surface blackened and abaxial surface
pale green (when dry), discoloured (when dry), base attenuate, apex obtuse to acute,
margin at or slightly revolute, entire throughout or slightly serrulate on the upper half
and glandular-punctate, 7-nerved from the base, two pairs of acrodromous veins and
one tenuous pair of veins close to the margin, tertiaries evident on the abaxial surface,
nearly perpendicular to the mid-vein, reticulate and randomly branching, adaxial sur-
face sparsely glandular-punctate, appearing glabrous when dry, abaxial surface dense-
ly glandular-punctate and covered with gland-tipped trichomes 0.2–0.4 mm long.
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
46
Figure 21. Microlicia chamissoana A habit B leaf abaxial surface C bracteole abaxial surface D oral
bud E Flower in lateral view F owering hypanthium G detail of the indumentum of the hypanthium
H petal adaxial surface I gynoecium J antepetalous (behind) and antesepalous (in front) stamens. Drawn
from Barreto 6745 (SPF, UEC).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 47
Inorescences simple or compound congested dichasia consisting of biparous cymes,
or reduced to solitary owers. Bracts (including petioles) 0.8–1.0 cm long, 0.6–0.7 cm
wide, 5-nerved, elliptical, indumentum like that of the principal leaves. Bracteoles (at
anthesis) with petioles 1.6–2.0 mm long, blades 4.2–4.9 mm long, 1.4–2.1 mm wide,
narrowly elliptic, base attenuate, apex acuminate, margin entire along the basal half,
sparsely serrulate on the upper half, 3–5-nerved, indumentum like that of the principal
leaves. Flowers 5-merous, pedicels (at anthesis) 2–4 mm long. Hypanthia (at anthesis)
2.5–3.7 mm long, 1.9–2.1 mm wide at the torus, campanulate, reddish (when fresh),
externally glandular-punctate and sparsely to densely covered with gland-tipped tri-
chomes 0.2–0.4 mm long. Calyx tubes 0.9–1.2 mm long. Calyx lobes (at anthesis)
4.5–6.7 mm long, 1.9–2.1mm wide at the base, narrowly triangular, apex acuminate,
margin entire and sparsely ciliate with gland-tipped trichomes 0.2–0.4 mm long, (when
fresh) reddish, externally glandular-punctate and covered with gland-tipped trichomes
0.2–0.4 mm long. Petals 11.5–13 mm long, 5.8–6.2 mm wide, magenta, obovate, apex
acuminate, margin entire and glandular-punctate, adaxial surface sparsely glandular-
punctate, abaxial surface glabrous. Stamens 10, strongly dimorphic. Larger (antese-
palous) stamens 5, laments 5.2–6.0 mm long, pink, pedoconnectives 5.5–6.5 mm
long, pink, appendages 1.2–1.8 mm long, yellow, apex bilobate, thecae (excluding ros-
tra) 1.7–2.6 mm long, purple, oblong, rostra 0.3–0.6 mm long, the circular pores ca.
0.3 mm wide. Smaller (antepetalous) stamens 5, laments 4.7–5.0 mm long, pink,
pedoconnectives 1.0–1.2 mm long, yellow, inconspicuous appendages ca. 0.1 mm
long, yellow, apex truncate, thecae (excluding rostra) 1.5–2.0 mm long, yellow, oblong,
rostra 0.4–0.6mm long, the circular pores ca. 0.2 mm wide. Ovary 2.2–2.8mm long,
1.6–1.8 mm wide, cylindrical, 5-locular. Style ca. 6.3 mm long, pink. Capsules (at
maturity) 2.5–3.2 mm long, 2.3–3.0 mm wide, ovoid, initially enveloped by the hyp-
anthium, torus constricted at the apex, fruiting calyx tubes 1.5–1.7mm long, fruiting
calyx lobes 5.2–6 mm long, not thickened. Seeds ca. 0.5 mm long, reniform.
Distribution, habitat and elevation range. Largely restricted to the Serra do Cipó
in central Minas Gerais (Fig. 19A), but extending to the Serra dos Alves and probably
to Pico do Itambé. Most of the recent collections were made at Serra do Cipó; only
one collection studied came from Serra dos Alves, Souza & Miranda 1639 (BHCB).
Considering Sellow’s itinerary in Brazil (see Rego et al. 2013), the type series was prob-
ably collected at the Pico do Itambé region. It occurs on quartzitic campo rupestre
exposed to full sun at elevations between 1154 and 1462 m.
Conservation. Microlicia chamissoana is known from fewer than 20 collections.
e EOO is 885.927 km2 and the AOO is 32 km2. is species was collected on
Pico do Itambé only in the 19th century and the local extinction of that population
is a possibility. Several populations found more recently are protected in the Parque
Nacional da Serra do Cipó. is species is already recognised as endangered (EN) by
the Brazilian Government (Brasília 2014). Our study based on IUCN (2019) recom-
mendations and criteria reached a similar conclusion regarding its conservation status
(EN): B1ab(iii).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
48
Recognition and anities. Microlicia chamissoana may be recognised by its el-
liptic leaf blades with tertiaries densely reticulate and randomly branching, congested
inorescences or solitary owers and narrowly triangular calyx lobes 4.5–5.7 mm long.
It is probably more closely related to M. laniora and M. pentagona, both of which
may occur sympatrically with M. chamissoana at Serra do Cipó. Microlicia chamissoana
diers from M. laniora by the shorter height (0.1–)0.5–0.8 m tall (vs. 0.5–3.5 m
tall), absence of lanose indumentum on branchlets, abaxial leaf surfaces and hypan-
thia (vs. present), leaves with shorter petioles 1.0–4.9 mm long (vs. 6–11 mm long),
blades with tertaries densely reticulate and randomly branching (vs. little reticulate and
branching apically), bracteoles with apices acuminate (vs. rounded), shorter hypanthia
2.5–3.7 mm long (vs. 5.0–6.5 mm long), shorter calyx lobes 4.5–6.7 mm long (vs.
7.9–9.7 mm long) and petals magenta (vs. white) 11.5–13.0 mm long (vs. 19.0–26.0).
In turn, Microlicia chamissoana diers from M. pentagona by the branchlets, abaxial
surfaces of the leaves and hypanthia that are densely glandular-punctate and covered
with gland-tipped trichomes (vs. appearing glabrous, vernicose and minutely granu-
lose), leaf blades with tertiaries densely reticulate and randomly branching (vs. paral-
lel or little reticulate and branching apically) and calyx lobes tenuous (vs. thickened)
5.2–6.0 mm long (vs. 6.5–11.0 mm long).
Notes. Major variation in M. chamissoana involves habit and degree of inores-
cence development. is species is usually a shrub about 1 m tall, although an atypical
specimen from Serra do Cipó is about 10 cm tall (A.M. Giulietti et al. CFSC12492).
is specimen was described as a herb, but it has woody branches. Most of the speci-
mens examined have congested, many-owered inorescences (e.g. Barreto 6745), al-
though in some of the inorescences, these are reduced to solitary owers (e.g. Pacico
& Carmo 154, Almeda et al. 8580).
Based on F. Sellow s.n., Chamisso (1834: 396–397) provided a detailed descrip-
tion for M. chamissoana under Microlicia sp., indicating his uncertainty of its generic
position, especially because of its 5-valvate capsules. Naudin (1849: 270) proposed the
epithet chamissoana for this species mentioning the description of Chamisso (1834:
396–397). Even preceded by a short description that is not diagnostic (5-valvate cap-
sules), the name Trembleya chamissoana Naudin is still valid because Naudin (1849)
made reference to Chamisso’s decription.
e type specimens of M. chamissoana, as cited by Cogniaux (1883–1888), have
two collection numbers indicated on each of their labels (F. Sellow 1171 and 1156).
Both collection numbers were cited by Cogniaux in “Flora brasiliensis”. At least one
set of Sellow’s duplicates of M. chamissoana was housed at B and probably destroyed
during World War II. As the Sellow duplicate at K is in good shape, it is here designated
as the lectotype for this species.
Specimens examined. Brazil. Minas Gerais: “Itambé” [probably Santo Antônio
do Itambé Municipality], Sellow s.n. (lectotype: K [K00530656]; isolectotype: P
[P00723508]); Conceição do Mato Dentro Municipality, Serra do Cipó, Kameyama et
al. CFSC10403 (SPF), Sheperd & Kirzenzaft 10214 (SP); Itabira Municipality, Serra
dos Alves, Souza & Miranda 1639 (BHCB); Jaboticatubas Municipality, Serra do Cipó,
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 49
Giulietti et al. CFSC12560 (SPF); Morro do Pilar Municipality, Serra do Cipó, Silvei-
ra s.n. (HUFU [56533]); Santana do Riacho Municipality, Serra do Cipó, Almeda et
al. 8580 (CAS, HUEM, UEC), Escaramai et al. 52 (SPF), Giulietti et al. CFSC12492
(HUEM, SPF), Pacico & Carmo 154 (HUEM), Pena & Viana 417 (SPF), Rocha
694 (BHCB), Romero et al. 8627 (HUEM, HUFU, RB), Semir CFSC5607 (SP);
Unknown municipality, Serra do Cipó, Barreto 6745 (BHCB, HUFU, NY, SP, SPF,
UEC, UPCB), Damazio 2026 (RB), Sena s.n. (W [W19110004181]).
4. Microlicia aviora Versiane & R.Romero, Bot. J. Linn. Soc. 197: 53. 2021.
Fig. 22
Trembleya hatschbachii Wurdack & E.Martins, Bol. Bot. Univ. São Paulo 14: 40. 1995.
original name. Type: Brazil. Minas Gerais, Grão Mogol, Rio das Mortes, 15 May
1988, G. Hatschbach, M. Hatschbach & O. Ribas 52005 (holotype: MBM!; iso-
types: BHCB!, C!, CAS!, CTES, ESA!, G!, HUFSJ, K!, MO!, RB!, S, SPF!, US!,
VIC, VIES).
Description. Erect shrubs 0.8–2.5 m tall. Branchlets quadrangular, appearing glabrous,
vernicose and minutely granulose, light green (when fresh). Internodes 0.6–3.0 cm
long, angles with narrow wings 0.2–0.4 mm wide. Petioles 3.9–17 mm long. Leaf
blades 37–90 mm long, 17–50 mm wide, coriaceous (when dry), elliptic to slightly
ovate, both surfaces green (when fresh), adaxial surface blackened and abaxial surface
pale brown (when dry), discoloured (when dry), base cuneate to attenuate, apex acute,
margin at, entire and minutely granulose and becoming glabrescent with age, 5-nerved
from the base, one pair of acrodromous veins and one pair of tenuous veins close to the
margin, tertiaries evident on the abaxial surface, nearly perpendicular to acute to the
mid-vein, little reticulate and branching apically, adaxial surface glabrous to minutely
granulose, vernicose, abaxial surface glabrous to minutely granulose. Inorescences
compound dichasia consisting of biparous cymes, not congested. Bracts (including
petioles) 3.2–5.0 cm long, 1.4–2.0 cm wide, 5-nerved, elliptical, appearing glabrous,
vernicose. Bracteoles (at anthesis) with petioles 1.6–1.9 mm long, blades 3.5–6.0 mm
long, 1.3–1.9 mm wide, narrowly elliptic, base attenuate, apex acute, margin entire,
1–3-nerved, indumentum appearing glabrous, vernicose. Flowers 5-merous, pedicels
(at anthesis) 1.8–2.2 mm long. Hypanthia (at anthesis) 3.3–4.1mm long, 3.0–3.2 mm
wide at the torus, campanulate, light green (when fresh), externally glabrous, minutely
granulose, vernicose. Calyx tubes inconspicuous, 0.1–0.2 mm long. Calyx lobes (at
anthesis) 4.0–4.9 mm long, 1.3–1.9 mm wide at the base, narrowly triangular, apex
acute, margin entire, (when fresh) light green, externally glabrous, minutely granulose,
vernicose. Petals 6.0–8.8 mm long, 5.2–7 mm wide, yellow, obovate, apex rounded,
margin entire and glabrous, both surfaces glabrous. Stamens 10, strongly dimorphic.
Larger (antesepalous) stamens 5, laments 3.4–4.0 mm long, yellow, pedoconnec-
tives 3.7–4.0 mm long, yellow, appendages 1.0–1.2 mm long, yellow, apex truncate
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
50
to slightly emarginate, thecae (excluding rostra) 1.4–1.6 mm long, brownish, oblong,
rostra 0.3–0.4 mm long, the circular pores ca. 0.2 mm wide. Smaller (antepetalous)
stamens 5, laments 2.7–2.9 mm long, yellow, pedoconnectives 1.2–1.4 mm long,
Figure 22. Microlicia aviora A habit B leaf abaxial surface C bracteole abaxial surface D oral bud
E ower in lateral view F owering hypanthium G petal adaxial surface H antesepalous (left) and antepetalous
(right) stamens I gynoecium J capsule enveloped by the hypathium. Drawn from Meireles et al. 1124 (UEC).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 51
yellow, short appendages ca. 0.5 mm long, yellow, apex truncate, thecae (excluding ros-
tra) 1.4–1.6 mm long, yellow-brownish, oblong, rostra 0.3–0.5 mm long, the circular
pores ca. 0.2 mm wide. Ovary 3.5–4.1 mm long, 2.9–3.1 mm wide, globose, 5-locular.
Style 4–4.2 mm long, yellow. Capsules (at maturity) 3.4–3.6 mm long, 3.5–4.2 mm
wide, globose, initially enveloped by the hypanthium, torus constricted at the apex,
fruiting calyx tubes 0.2–0.4 mm long, fruiting calyx lobes 3.7–4.0 mm long, not thick-
ened. Seeds ca. 0.6 mm long, reniform.
Distribution, habitat and elevation range. Endemic to northern Minas Gerais
(Fig. 19A), at Serra de Grão Mogol, Serra de Botumirim and Serra Nova. It occurs on
quartzitic campo rupestre exposed to full sun at elevations between 760 and 1243 m.
e distribution of M. aviora is a good match to the Grão Mogol biogeographic
district (Colli-Silva et al. 2019).
Conservation. is species is known from about 20 collections. e EOO is
468,668 km2 and the AOO is 32 km2. Most of the populations of M. aviora oc-
cur within the following conservation units: Parque Estadual de Grão Mogol, Parque
Estadual de Botumirim and Parque Estadual da Serra Nova, where this species is af-
forded protection. e Brazilian Government assigned a conservation status of Endan-
gered (EN) to this species (Brasília 2014). Based on IUCN (2019) recommendations
and criteria, we concur with that conclusion (EN): B1ab(iii).
Recognition and anities. Microlicia aviora may be recognised by its leaves
and hypanthia that appear to be glabrous, but are vernicose and minutely granulose,
leaf blades 3.7–9.0 cm long, elliptic to slightly ovate, compound dichasia and yel-
low petals. In overall vegetative morphology, M. aviora resembles M. tridentata. In
turn, its yellow petals, staminal laments and styles are shared only with M. rosma-
rinoides. Microlicia aviora diers from M. tridentata by its leaves that have entire
margins throughout (vs. serrulate along the upper half), abaxial surfaces appearing
glabrous (vs. glandular-punctate), shorter bracteoles with blades 3.5–6.0 mm long (vs.
8.1–11.0 mm long) and apex acute (vs. rounded to obtuse) and yellow petals (vs. ma-
genta or rarely white) that are 6.0–8.8 mm long (vs. 11.5–13.0 mm long). Microlicia
aviora diers from M. rosmarinoides by its taller habit 0.8–2.0 m tall (vs. 0.3–0.6m
tall), branchlets, abaxial surfaces of the leaves and hypanthia appearing glabrous (vs.
glandular-punctate), leaf blades 3.7–9.0 cm long (vs. 0.4–1.0 cm long) that are el-
liptic to slightly ovate (vs. linear to lanceolate) and have 5 basal acrodromous veins
(vs. 1-nerved from the base), compound dichasia (vs. solitary owers), longer calyx
lobes 4.0–4.9 mm long (vs. 2.2–2.8 mm long) and longer petals 6.0–8.8 mm long (vs.
5.0–5.3 mm long).
Specimens examined. Brazil. Minas Gerais: Botumirim Municipality, Estrada
para o Rio do Peixe, Forzza et al. 4897 (NY, RB, SPF), Serra da Canastra, Mello-Silva et
al. 509 (HUEM, SPF, UEC), Nakajima et al. 4764 (HUFU), Scatigna & Galvão 376
(UEC); Grão Mogol Municipality, Serra de Grão Mogol, Bidá et al. CFCR11951 (SPF,
US), Cerati et al. 246 (K, SP, UEC), Furlan et al. CFCR771 (SPF, UEC, US), Hatsch-
bach & Hatschbach 52005 (holotype: MBM; isotypes: BHCB, C, CAS, CTES, ESA,
G, HUFSJ, K, MBM, MO, S, SPF, US, UPCB, VIC, VIES), Hatschbach 41337 (ESA,
FLOR, HCF, HUEFS, MBM, NY, RB, SPF, UPCB, US), Hatschbach et al. 54239 (CAS,
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
52
INPA, MBM), Hatschbach et al. 68067 (MBM), Hensold et al. CFCR3525 (SPF, US),
Kral et al. 72723 (SP, SPF), Leitão Filho et al. 7893 (MBM, UEC), Meireles et al. 1124
(CAS, HUEM, UEC), Oliveira et al. CFCR12997 (SPF, US), Pacico & Simoes 353
(CAS, HUEM), Pacico 565 (CAS, HUEM, RB); Pirani & Mello-Silva CFCR10814
(HUEM, SPF, UEC, US), Zappi et al. CFCR9903 (SPF, UEC); Rio Pardo de Minas
Municipality, Serra Nova, Araújo et al. 2043 (BHCB), Rocha et al. 497 (BHCB, NY).
5. Microlicia laniora (D.Don) Baill., Adansonia 12: 95. 1877.
Fig. 23
Melastoma laniora D.Don, Mem. Wern. Nat. Hist. Soc. 4: 292. 1823. basionym. Type:
Brazil. “in Brazilia”, F. Sellow s.n. (lectotype, designated here: K [K00957812]!;
probable isolectotypes: K [K00957818]!, P [P005317063]!, P [P005317064]!, P
[P005317070]!; image of lectotype is available at http://specimens.kew.org/her-
barium/K000957812).
Trembleya lychnitis Schrank & Mart. ex DC., Prod. 3: 126. 1828. Type: Brazil. “In
Brasiliae lapidosis apricis ad latera montium prov. Min. gener.” [Minas Gerais],
C.F.P. Martius s.n. (lectotype, designated here: G [G00368004]!; isolectotype: M
[M0165875]!).
Trembleya laniora (D.Don) Cogn. in Martius et al., Fl. Bras. 14(3): 130. 1883.Type:
Based on Melastoma laniora D.Don.
Trembleya laniora var. acutifolia Cogn. in Martius et al., Fl. Bras. 14(3): 131. 1883.
syn. nov. Type: Brazil. “Brasilia meridionalis, ad Serra da S. Antonio” [Minas Ge-
rais], 1878, F. Sellow 1727 (lectotype, designated here: K [K00530655]!; isolecto-
type: P [P00723419]!; image of lectotype available at http://specimens.kew.org/
herbarium/K000530655).
Trembleya laniora var. genuina Cogn. in Martius et al., Fl. Bras. 14(3): 130. 1883.
syn. nov. Type: Brazil. “Minas Geraes” [Minas Gerais], 1840, P. Claussen
332A (lectotype, designated here: BR [BR0000005228225]!, isolectotype: BR
[BR0000005227891]!).
Trembleya laniora var. grandifolia Cogn. in Martius et al. Fl. Bras. 14(3):131. 1883.
syn. nov. Type: Brazil. “ad Pico d’Itabira et ad Caxoeira do Campo” [Minas Ger-
ais, Itabira and Cachoeira do Campo], C.F.P. Martius 930 (lectotype, designated
here: P [P005317085]!; isolectotypes: BM [BM00516949]!, G [G00368005]!,
G [G00318589]!, GH [GH00053137]!, L [L00056323]!, L [L00056324]!, M
[M0165876]!, M [M0165877]!, M [M0165878]!, P [P005317086]!, S [S09-
12961]!; image of lectotype is available at http://coldb.mnhn.fr/catalognumber/
mnhn/p/p05317085).
Trembleya laniora var. intermedia Cogn. in Martius et al., Fl. Bras. 14(3):130. 1883.
syn. nov. Type: Brazil. “In prov. Minas Geraës loco haud indicato” [Minas Gerais],
G. Gardner 4601 (lectotype, designated here: BM [BM00525899]!; isolectotypes:
G [G00318586]!, GH [GH00053136]!, NY [NY00245856]!, P [P005317081]!,
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 53
R [R000168426]! US [US00623967]!; image of isolectotype at P is available at
http://coldb.mnhn.fr/catalognumber/mnhn/p/p05317081).
Description. Erect shrubs or treelets 0.5–3.5 m tall. Branchlet surfaces concealed by a
lanose indumentum of eglandular trichomes 0.1–0.5 mm long, whitish (when fresh).
Internodes 0.7–4.5 cm long, angles unwinged. Petioles 6–11 mm long. Leaf blades
20–39 mm long, 9–25 mm wide, coriaceous (when dry), ovate, elliptic or narrowly
elliptic, adaxial surface green and partially covered by a thin layer of whitish indu-
mentum, abaxial surface totally concealed by the white lanose indumentum (when
fresh), adaxial surface blackened, abaxial surface hidden by the white to pale brown
lanose indumentum (when dry), discoloured (when dry), base cuneate to rounded,
apex acute to rounded, margin at, entire and minutely granulose and becoming gla-
brescent with age, 5-nerved from the base, one pair of acrodromous veins and one
pair of tenuous veins close to the margin, tertiaries evident on the abaxial surface,
nearly perpendicular to the mid-vein, little reticulate and branching apically, adaxi-
al surface glandular-punctate, usually pruinose and becoming glabrescent with age,
abaxial surface densely covered with lanose eglandular trichomes 0.1–0.5 mm long.
Inorescences simple dichasia or reduced to solitary owers, usually congested. Bracts
(including petioles) 1–1.9 cm long, 0.3–0.8 cm wide, 3-nerved, ovate, elliptical or
narrowly elliptic, indumentum like that of the major leaves. Bracteoles (at anthesis)
with petioles 4–6 mm long and blades eventually linear and rudimentary, blades (when
well-developed) 2.2–4.4 mm long, 1.5–2 mm wide, linear to elliptic, base attenu-
ate, apex rounded, margin entire, 1-nerved, indumentum like that of the principal
leaves. Flowers 5(–6)-merous, pedicels (at anthesis) 0.7–2.0 mm long. Hypanthia (at
anthesis) 5.0–6.5 mm long, 4.5–5.2mm wide at the torus, campanulate to urceo-
late, surface hidden by the whitish lanose indumentum composed of eglandular tri-
chomes 0.1–0.5 mm long. Calyx tubes 1.0–2.0mm long. Calyx lobes (at anthesis)
7.9–9.7 mm long, 2.0–2.7 mm wide at the base, subulate, apex acute, margin entire,
(when fresh) surface hidden by the whitish lanose indumentum composed of eglan-
dular trichomes 0.1–0.5 mm long. Petals 19–26mm long, 10–15 mm wide, white,
rarely with pink stains at the apical region, obovate, apex emarginate, margin entire
and ciliate with eglandular trichomes 0.1–0.4mm long at the apical region, both sur-
faces glabrous. Stamens 10(–12), strongly dimorphic. Larger (antesepalous) stamens
5(–6), laments 5.2–6.3 mm long, white, pedoconnectives 6.1–7.3 mm long, white
to light yellow, appendages 1.9–3.0mm long, yellow, apex emarginate, thecae (exclud-
ing rostra) 2.3–2.7mm long, vinaceous, oblong, rostra 0.4–0.6 mm long, the circular
pores ca. 0.2mm wide. Smaller (antepetalous) stamens 5(6), laments 4.0–5.4mm
long, white, pedoconnectives 0.8–1.1mm long, yellow, inconspicuous appendages ca.
0.1mm long, yellow, apex emarginate, thecae (excluding rostra) 2.4–2.6 mm long,
yellow, oblong, rostra 0.3–0.6 mm long, the circular pores ca. 0.2 mm wide. Ovary
2.5–4.0 mm long, 1.7–2.0 mm wide, globose to cyclindrical, 5-locular. Style 6.5–
8.0mm long, white. Capsules (at maturity) 5.0–8.0 mm long, 5.0–6.0 mm wide,
globose, initially enveloped by the hypanthium, torus constricted at the apex, fruiting
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
54
Figure 23. Microlicia laniora A habit B leaf abaxial surface C bracteole abaxial surface D ower in
lateral view E owering hypanthium F petal adaxial surface G detail of indumentum on the apex of the
petal H antepetalous (behind) and antesepalous (in front) stamens I gynoecium J ovary in cross-section
K capsule enveloped by the hypanthium. Drawn from Almeda et al. 7726 (UEC) and Almeda et al.
9197 (UEC).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 55
calyx tubes 2.0–3.1 mm long, fruiting calyx lobes 9.0–11.5 mm long, not thickened.
Seeds ca. 0.8 mm long, reniform.
Distribution, habitat and elevation range. Endemic to central and southern Mi-
nas Gerais (Fig. 19B), mainly at Serra de Ouro Branco, Serra do Itacolomi, Serra do
Cipó, Serra da Piedade, Serra da Moeda, Serra do Caraça, Serra do Gandarela, Serra de
Lavras Novas, Serra dos Alves, Serra de Itabirito, Serra do Rola-Moça, Serra do Cur-
ral, Serra de Taquaril, Serra do Garimpo, Serra do Belo Vale, Diamantina Plateau and
mountains in the Municipalities of Brumadinho and Mariana. It occurs on quartzitic or
ferrugineous campo rupestre exposed to full sun at elevations between 868 and 1540 m.
Ecology. Microlicia laniora has a symbiotic relationship with arbuscular mycor-
rhizae (Abrahão et al. 2019). is is likely related to phosphorus and nitrogen acquisi-
tion strategies (Abrahão et al. 2019). is species easily colonises degraded campo rup-
estre areas and has been recommended for ecological restoration (Amaral et al. 2013).
According to Pereira (2011), its leaves develop increased asymmetry when damaged
by insects. Vasconcelos et al. (2001) reported the use of M. laniora for nesting by the
endemic hummingbird, Hyacinth Visorbearer (Augastes scutatus), in campo rupestre at
the Serra do Cipó.
Uses. Leaf and stem extracts from Microlicia laniora have antimicrobial activity
against Staphylococcus aureus and Micrococcus luteus (Cota et al. 2002; Ventura et al. 2007a).
Conservation. is species is known from more than 100 herbarium specimens
making it one of the better sampled species in the Trembleya s.s. clade. However, this
sampling is geographically biased towards the surroundings of the MG-010 highway,
where more than a half of these collections were made. e MG-010 highway is the
main road crossing the mountains at south-eastern Serra do Cipó and provides easy
access to large populations of M. laniora. e EOO is 19,113.356 km2 and the AOO
is 248 km2. Based on IUCN (2019) recommendations and criteria, we suspect that
this species would be classied as Vulnerable (VU): B2ab(iii). Several populations of
M. laniora occur within the following conservation units: Parque Estadual Serra do
Ouro Branco, Parque Estadual do Biribiri, Parque Estadual de Itacolomi, Monumento
Natural Estadual Serra da Piedade, Monumento Natural Estadual Serra da Moeda,
Parque Estadual da Serra do Rola-Moça, Parque Nacional da Serra do Gandarela,
Parque Nacional da Serra do Cipó and RPPN Serra do Caraça (Natural Heritage Pri-
vate Reserve), where M. laniora is aorded protection.
Recognition and anities. Microlicia laniora may be recognised by its branch-
es, abaxial surfaces of the leaves and hypanthia that are densely covered by a lanose
indumentum, white petals (rarely ushed with pink) that are 19.0–26.0 mm long
and subulate calyx lobes 7.9–9.7 mm long. It appears to be most closely related to
M. pentagona and M. chamissoana. Microlicia laniora diers from M. pentagona by
the branches, abaxial foliar surfaces and hypanthia densely covered by the lanose indu-
mentum (vs. appearing glabrous, vernicose and minutely granulose), leaves with longer
petioles 6.0–11.0 mm long (vs. 0.4–2.5 mm long) and margins entire throughout (vs.
serrulate on the upper half), longer hypanthia 5.0–6.5 mm long (vs. 2.5–3.5 mm long)
and longer petals 19.0–26.0 mm long (vs. 11.8–13.8 mm long), that are white, rarely
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
56
ushed with pink (vs. magenta). For additional comparisons, see comments under
M. chamissoana.
Notes. e four varieties proposed by Cogniaux (1883–1888) were based mainly
on dierences in petiole length, leaf size and leaf shape. An examination of Cogniuax’s
varieties and many additional collections showed these purported dierences to be in-
consistent with much size overlap in the characters he used to delimit his infraspecic
taxa. e ovate leaf shape that Cogniuax (1883–1888) attributed only to var. genuina is
present in most of the individuals examined, along with a great deal of variation in leaf
and petioles sizes. Cavalcanti et al. CFSC10628 (UEC), for example, has ovate leaves
(a feature attributed only to var. genuina) and well-developed petioles (a feature at-
tributed to the varieties intermedia, grandifolia and acutifolia). is specimen could be
identied as var. intermedia since its leaf measurements match those of the protologue
(3–4 cm long). However, when comparing the Cavalcanti specimen with the type of
var. intermedia (Gardner 4601), it is clear that this variety has lanceolate leaves that are
very distinct from the ovate leaves of Cavalcanti et al. CFSC10628. Characters, such
as leaf shape and size, also vary along branches of a single individual and, thus, do not
constitute good diagnostic features to delimit varieties of this species.
According to Cogniaux (1883–1888), some of Glaziou’s collections of M. laniora
(his numbers 14740 and 14741) were made “in prov. Rio de Janeiro” [Rio de Janeiro
State]. On the other hand, Glaziou (1908: 251) asserts that his number 14740 came
from “Serra do Caraça, au Morro do Incionado” and the number 14741 came from
“Campo de São Sebastião, prés Ouro Preto”, both sites located in Minas Gerais State.
e labels of Glaziou 14740 and 14741 from P had annotations of collections’ sites
similar to those shown in Glaziou (1908), while a duplicate of Glaziou 14740 from K
has a label that says “14741 the same” and the location site is cited as “Environs de Rio
de Janeiro et Ouro Preto”. Due to these contradictions, the cited collections were not
used for the geographical distribution summary of M. laniora. e discordant data
on Glaziou’s collections of Melastomataceae have been noted by Wurdack (1970) for
other collections of Melastomataceae, such as Clidemia, Leandra, Macairea, Miconia,
Rhynchanthera, Tibouchina and Tococa. is is the rst report of dubious geographic
information for a species of Microlicia. We found similar contradictory information
on labels of Glaziou 14746 which is discussed in the comments under M. pithyoides.
Specimens examined. Brazil. Minas Gerais: Barão de Cocais Municipality, Serra
do Garimpo, Hensold 778 (SPF, US), Semir et al. 28809 (UEC), Souza 1606 (BHCB);
Belo Horizonte Municipality, Serra do Taquaril – Serra do Curral, Barreto 6769 (SP),
Ducke s.n. (RB [241970]), Ferreira 5544 (HUFU), Roth s.n. (BHCB, CESJ, CTES,
ESA, R, RB, SP, UB, UPCB), Morro do Chapéu, Brandão 28574 (HUFU), Bru-
madinho Municipality, Serra do Rola-Moça, Carmo 4819 (BHCB), Serra da Calça-
da, Martens 7 (SPF), Martens 383 (SPF), Retiro das Pedras, Carvalho s.n. (HUFU
[39992]), Stehmann & Morais 2650 (BHCB); Caeté Municipality, Serra do Gandare-
la, Damazio 1025 (RB); Catas Altas Municipality, Serra do Caraça, Vasconcelos s.n.
(SPF [145870, 145871]); Conceição do Mato Dentro Municipality, Serra do Cipó,
Macedo 3758 (S), Martinelli & Tavora 2583 (RB); Diamantina Municipality, “Dia-
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 57
mantina Plateau”, Araújo et al. 323 (HUFU, RB, UEC), Brade 13735 (NY, RB, US),
Franco et al. 1270 (HUFU), Hatschbach et al. 27400 (K, MBM, UPCB), Hatschbach
et al. 68138 (MBM, UPCB), Leitão et al. 17281 (RB, UEC), Lima et al. 49 (SPF),
Maguire et al. 49140 (NY), Marques et al. 274 (HUFU), Mello et al. 61 (HUFU),
Vauthier 37 (P); Itabira Municipality, Martius 930 (GH, K, NY, P), Torres s.n. (RB
[241965]); Itabirito Municipality, Serra do Itabirito, Irwin 19974 (NY), Irwin et al.
19974 (K, NY, US), Krieger 10641 (CESJ, ESA, HUFU, MBM), Lima et al. 1443
(RB), Teixeira s.n. (BHCB [21783], HUFU [19443]); Jaboticatubas Municipality, Ser-
ra do Cipó, Goldenberg & Silveira 1573 (UPCB); Mariana Municipality, Collo et al.
s.n. (SPF [62806]), Messias et al. 1922 (OUPR, RB); Nova Lima Municipality, Serra
do Curral, Nakajima & Romero 3040 (HUFU), Pereira & Pabst 3107 (RB), Sampaio
7194 (BHCB), Williams & Assis 6352 (GH, NY); Ouro Branco Municipality, Serra
de Outro Branco, Almeda et al. 7726 (CAS, UEC), Almeda et al. 8395 (CAS, UEC),
Alves & Almeida-Lafetá 5573 (R), Alves et al. 6925 (R), Araújo et al. 334 (ESA, RB),
Arbo et al. 3906 (CTES, MBM, RB, SPF, UB), Delni et al. 78 (ESA, HUFU, RB),
Forzza et al. 993 (SPF), Nakajima et al. 4547 (HUFU), Paula et al. 136 (HUFU,
VIC), Paula et al. 295 (HUFU, VIC), Paula et al. 8 (HUFU, VIC), Pereira & Pabst
2944 (RB), Pirani et al. CFCR11211 (SPF), Rocha et al. 602 (BHCB, NY), Saraiva
et al. 85 (OUPR, RB); Ouro Preto Municipality, Barreto & Viégas s.n. (IAC [6389]),
Damazio 1540 (NY, US), Ferreira et al. 433 (HUFU), Fontana et al. 2288 (RB), For-
zza et al. 6344 (OUPR, RB, UPCB), Forzza et al. 6359 (NY, OUPR, RB, UPCB),
Groppo & Ulwin 676 (SPF), Lima et al. 1296 (RB), Meireles et al 1362 (HUEM,
UEC), Messias et al. 2151 (OUPR, RB), Rolim et al. 329 (HUFU, RB, VIC), Rolim
et al. 386 (RB, VIC), Rolim et al. 61 (HUFU, VIC), Teixeira s.n. (SPF [114173]),
Valente et al. 2574 (RB, VIC); Raposos Municipality, Tameirão Neto & Mansur 4874
(BHCB); Rio Acima Municipality [Gandarela], Emygdio 3314 (NY), Emygdio 3353
(NY); Sabará Municipality, Barreto 6771 (BHCB), Barreto 6772 (NY); Santa Bárbara
Municipality, Serra do Caraça, Almeda et al. 7753 (CAS, UEC), Almeda et al. 8862
(CAS, UEC), Arbo et al. 4030 (SPF), Barreto 706 (NY, UEC), Fraga et al. 3333 (NY,
RB), Fraga et al. 3341 (NY, RB), Marcondes-Ferreira et al. 281 (SPF), Pirani & Yano
696 (CAS, SPF), Pirani et al. 696 (SP, SPF), Rocha et al. 672 (BHCB, RB), Romero et
al. 5307 (CAS, UEC), Tales et al. 17 (BHCB, HUFU), Temponi & Vasconcelos s.n.
(BHCB [36249], HUFU [19313]), TSMG & Tales 81 (BHCB, HUFU), Valente et
al. 1229 (HUFU, VIC), Valente et al. 1230 (HUFU, VIC), Valente et al. 535 (HUFU,
VIC); Santana do Riacho Municipality, Serra do Cipó, Almeda et al. 8549 (CAS,
UEC), Almeda et al. 9179 (CAS, UEC), Alves et al. 2109 (SPF), Andrade et al. 374
(BHZB, HUFU), Antar et al. 1662 (SPF), Borges et al. 153 (HUEFS, K, NY, SPF),
Bruniera et al. 37 (HUFU, SPF), Castro et al. s.n. (HUEM [24842], HUFU [2254]),
Ceccantini et al. 3914 (SPF), Chuckr et al. s.n. (HUEM [24776]), Cordeiro et al.
CFSC10504 (SPF), Escaramai et al. 65 (SPF), Faria & Mazucato 105 (SPF), Farinac-
cio et al. 36 (MBM, RB), Fernandes et al. 1468 (BHZB, HUFU), Forero et al. 7700
(SPF), Forero et al. 7831 (SPF), Giulietti et al. CFSC12564 (UEC), Kral et al. 72997
(CEN, SP), Kubo et al. 109 (SPF), Kubo et al. 125 (SPF), Maguire et al. 49016 (NY),
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
58
Mattos & Rizzini 107 (RB, US), Mattos & Rizzini 480 (RB), Monge et al. 384 (UEC),
Ordones et al. 1856 (BHZB, HUFU), Pacico 185 (HUEM, SPF), Pena & Viana 365
(SPF), Pereira & Pabst 152 (HUEM), Pirani et al. 5082 (NY, SPF), Pires & Braga s.n.
(CESJ [21520]), Reginato et al. 1401 (NY, UPCB), Sakuragui & Souza 38 (ESA),
Salatino et al. 14 (NY, SPF), Salatino et al. 18 (NY, SPF), Souza et al. 11565 (ESA,
RB), Souza et al. 25181 (ESA), Stehmann & Morais 2354 (SPF), Verdi et al. 6501
(RB), Zappi et al. CFSC9353 (SPF); Santana do Riacho Municipality [“Conceição
do Mato Dentro”], Serra do Cipó, Marquete et al. 3817 (RB); Santana do Riacho
Municipality [“Jaboticatubas”], Serra do Cipó, Joly CFSC2405 (SPF), Joly CFSC82
(SPF), Mantovani 99 (SP, SPF), Semir CFSC2005 (SP, SPF), Semir CFSC4133 (SPF),
Semir CFSC5001 (SPF), Semir CFSC5068 (SPF); Serro, Semir et al CFCR230 (SPF);
Unknown municipality, Bunge s.n. (P [P005317069]), Claussen 1 (BR, CAS), Claus-
sen 1645 (P), Claussen 332A (BR), Claussen 554 (P), Claussen 640 (BR), Claussen
923 (CAS), Claussen s.n. (K [K00957804, K00957810, K00957811, K00957816],
NY [NY00941988], P [P005317066, P005317067, P005317071, P005317072]),
Gardner 4601 (BM, G, GH, K, NY, P, R, US), Glaziou 11953 (K), Glaziou 14740
(K, P), Glaziou 14741 (K, P), Glaziou s.n. (P [P005317096]), Gounelle s.n. (P
[P005317089]), Lund 135 (C), Martius 930 (BM, G, GH, L, M, P, S), Martius s.n. (P
[P005317088]), Netto s.n. (BR [BR0000005520688]), Raben 428 (S), Riedel s.n. (K
[K00957817], P [P005317084]), Saint-Hilaire 216 (P), Sellow 1446 (BR, US), Sellow
1727 (P [P00723419]), Sellow s.n. (lectotype: K [K00957812]; probable isolecto-
types: K [K00957818], P [P005317063, P005317064, P005317070]), Vauthier s.n.
(P [P005317061]).
Putative hybrid (M. laniora × M. pentagona). Brazil. Minas Gerais: Santana
do Riacho Municipality [“Santa Luzia”], Serra do Cipó, Barreto 7026 (BHCB).
6. Microlicia parviora (D.Don) Versiane & R.Romero, Bot. J. Linn. Soc. 197:
54. 2021.
Fig. 24
Meriania parviora D.Don, Mem. Wern. Nat. Hist. Soc. 4: 323. 1823. basionym.
Type: Brazil. F. Sellow s.n. (lectotype, designated here: G [G00396696]!; probable
isolectotype: P [P05317745]!).
Trembleya heterostemon Mart. & Schrank ex DC., Prod. 3: 126. 1828. syn. nov. Type:
Brazil. “In Brasiliae subalpinis ad fontes in prov. Minarum Generalium” [Minas
Gerais], C.F.P. Martius 961 (lectotype, designated here: M [M0165884]!; isolecto-
types: G [G00310210]!, M [M0165883]!).
Trembleya triora Mart. & Schrank ex DC., Prod. 3: 126. 1828. syn. nov. Type: Brazil.
“In sylvis caeduis prope Villam-Riceam prov. Minarum generalium” [Minas Gerais,
Ouro Preto], 1827, C.F.P. Martius s.n. (lectotype, designated here: M [M0165882]!;
isolectotypes: G [G00310209]!, P [P00723390]!; image of isolectotype at P is
available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p00723390).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 59
Trembleya paniculata Naudin, Ann. Sci. Nat., Bot. Sér. 3, 2: 154. 1844. Type: Bra-
zil. “In campis circa Juruoca in prov. Minas Geraës” [Minas Gerais], 1816–1821,
[catal. D, n° 462] A. Saint-Hilaire s.n (lectotype, rst-step designated by Martin
and Cremers (2007), second-step designated here: P [P00723414]!; isolectotype:
P [P00723415]!; image of lectotype is available at http://coldb.mnhn.fr/catalog-
number/mnhn/p/p00723414).
Trembleya parviora (D.Don) Cogn. in Martius et al., Fl. Bras. 14(3): 127. 1883. Type:
Based on Meriania parviora D.Don.
Trembleya parviora subsp. heterostemon (Mart. & Schrank ex DC.) Cogn. in Martius
et al., Fl. Bras. 14(3): 128. 1883. syn. nov. Type: Based on Trembleya heterostemon
Mart. & Schrank ex DC.
Trembleya parviora subsp. triora (Mart. & Schrank ex DC.) Cogn. in Martius et al.,
Fl. Bras. 14(3): 129. 1883. syn. nov. Type: Based on Trembleya triora Mart. &
Schrank ex DC.
Trembleya parviora var. angustifolia Cogn. in Martius et al., Fl. Bras. 14(3): 128.
1883. syn. nov. Type: Brazil. “In prov. Rio de Janeiro ad Serra dos Orgâos” [Rio
de Janeiro], 1838, G. Gardner 379 (lectotype, designated here: P [P005317041]!;
isolectotypes: BR [BR0000005520404]!, NY [NY00245857-online image]!, US
[US00623960]!; image of lectotype is available at http://coldb.mnhn.fr/catalog-
number/mnhn/p/p05317041).
Trembleya parviora var. denticulata Cogn. in Martius et al., Fl. Bras. 14(3): 129. 1883.
syn. nov. Type: Brazil. “In prov. S. Paulo ad Paitura” [São Paulo], 1846, Prates
s.n. (lectotype, designated here: P [P00723407]!; isolectotypes: P [P00723406]!,
P [P00723408]!; image of lectotype is available at http://coldb.mnhn.fr/catalog-
number/mnhn/p/p00723407).
Trembleya parviora var. farinacea Cogn. in Martius et al., Fl. Bras. 14(3): 128. 1883.
syn. nov. Type: Brazil. “In campo sicco apricot vel umbroso ad Caldas prov. Minas
Geraës” [Minas Gerais, Caldas], H. Mosen 1971 (lectotype, designated here: R
[R000166846]!; isolectotypes: C [C10015104]!, P [P005317106]!; image of isolecto-
type at P is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p05317106).
Trembleya parviora var. latifolia Cogn. in Martius et al., Fl. Bras. 14(3): 128. 1883.
syn. nov. Type: Brazil. “In prov. Rio de Janeiro ad Serra dos Orgâos”, 1838, G.
Gardner 380 (lectotype, designated here: P [P00723401]!; isolectotypes: BR
[BR0000005225798]!, F [F0064040F]!, G [G00359408]!, G [G00368014-on-
line image]!, NY [NY00217747]!, NY [NY00245859]!, P [P00723402]!, S [S05-
3227]]!, US [US00623963]!; image of lectotype is available at http://coldb.mnhn.
fr/catalognumber/mnhn/p/p00723401).
Trembleya parviora var. martii Cogn. in Martius et al., Fl. Bras. 14(3): 129. 1883. syn.
nov. Type: Brazil. “In prov. Minas Geraës ad Serra do Ouro Preto” [Minas Gerais,
Ouro Preto], C.F.P. Martius 931 (lectotype, designated here: P [P005317103]!; isolec-
totypes: BM, G [G00368009]!, G [G00318575]!, M [M0165879]!, M [M0165880]!,
MO [MO-2267366]!, P [P005317750]!, P [P005317751]!; image of lectotype is
available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p05317103).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
60
Trembleya parviora var. multiora Cogn. in Martius et al., Fl. Bras. 14(3): 129.
1883. syn. nov. Type: Brazil. “Ayuruoca, Minas Gerais’”, 17 April 1878, A.F.M.
Glaziou 9454 (lectotype, designated here: R [R000009197]!; isolectotypes: BR
[BR0000005226733]!, BR [BR0000005226405]!, BR [BR0000005227068]!, C
[C10015105-online image]!, G [G00368008]!, G [G00318573-online image]!, S
[S-0912956-online image]!, P [P005317099]!, P [P005317057]!; image of isolecto-
type at P is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p05317099).
Trembleya parviora var. parvifolia Cogn. in Martius et al., Fl. Bras. 14(3): 129. 1883.
syn. nov. Type: Brazil. “In prov. Minas Geraës ad Rio das Pedras” [Minas Gerais],
F. Sellow 1154 (lectotype, designated here: US [US00623966-online image]!; im-
age of lectotype is available at http://n2t.net/ark:/65665/392812a8c-782b-431c-
b2fc-3644dcfad16e).
Trembleya parviora var. selloana Cogn. in Martius et al., Fl. Bras. 14(3): 128. 1883.
syn. nov. Type: Brazil. “In prov. Minas Gerais” [Minas Gerais], F. Sellow 5278 (lec-
totype, designated here: P [P00723410]!; image of lecotype is available at http://
coldb.mnhn.fr/catalognumber/mnhn/p/p00723410).
Trembleya parviora var. tomentosa Cogn. in Martius et al., Fl. Bras. 14(3): 128). syn.
nov. Type: Brazil. “In prov. Rio de Janeiro ad Serra dos Orgâos” [Rio de Janeiro],
J.B.A. Guillemin 946 (lectotype, designated here: P [P005317767]!; isolectotypes:
G [G00368013]!, P [P00723409]!; image of lectotype is available at http://coldb.
mnhn.fr/catalognumber/mnhn/p/p05317767).
Trembleya parviora var. triora (Mart. & Schrank ex DC.) Cogn. in Martius et al.,
Fl. Bras. 14(3): 129. 1883. syn. nov. Type: Based on Trembleya triora Mart. &
Schrank ex DC.
Trembleya parviora var. valtheri Cogn. in Martius et al., Fl. Bras. 14(3): 128.
1883. syn. nov. Type: Brazil. “In prov. Minas Geraës” [Minas Gerais], 1833,
M. Vauthier 43 (lectotype, designated here: P [P00723403]!; isolectotypes: BR
[BR0000005226375]!, G [G00368012]!, G [G00318569]!, P [P00723404]!, P
[P00723405]!; image of lectotype is available at http://coldb.mnhn.fr/catalog-
number/mnhn/p/p00723405).
Trembleya parviora var. vulgaris Cogn. in Martius et al., Fl. Bras. 14(3): 128. 1883. syn.
nov. Type: Brazil. “In prov. Minas Geraës” [Minas Gerais], 1840, G. Gardner 4602
(lectotype, designated here: R [R000168427]!, isolectotypes: NY [NY00245860]!,
NY [NY00245861]!, US; image of isolectotype at NY is available at http://sweet-
gum.nybg.org/science/vh/specimen_details.php?irn=535234).
Trembleya parviora var. warmingii Cogn. in Martius et al., Fl. Bras. 14(3): 128. 1883.
syn. nov. Type: Brazil. “In prov. Minas Gerais ad Lagoa Santa” [Minas Gerais, Lagoa
Santa], E. Warming s.n. (lectotype, designated here: P [P005317116]!; isolectotypes:
BR [BR0000005520732]!; C [C10015107-online image]!; image of lectotype is
available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p05317116).
Trembleya parviora var. widgrenii Cogn. in Martius et al., Fl. Bras. 14(3): 128. 1883. syn.
nov. Type: Brazil. “In prov. Rio de Janeiro” [Rio de Janeiro], Widgren s.n. (lectotype,
designated here: P [P005317115]!; isolectotypes: BR [BR0000005227044]!, PH
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 61
[PH00027744]!, S [S-053231]!; image of lectotype is available at http://coldb.
mnhn.fr/catalognumber/mnhn/p/p05317115).
Trembleya parviora var. heterophylla Cogn. in de Candolle & de Candolle [A.D.C.
& C.DC.], Monogr. Phan. 7: 75. 1891. syn. nov. Type: Brazil. “In prov. Rio
de Janeiro, Nova Friburgo” [Rio de Janeiro, Nova Friburgo], 31 July 1877,
A.F.M. Glaziou 16778 (lectotype, designated here: R [R000009196]!; isolecto-
types: C [C10015106]!, G [G00368010]!, G [G00368011]!, L [L0056325]! P
[P00723411]!, P [P00723412]!, P [P00723413]!; image of isolectotype at P is
available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p00723411).
Description. Erect shrubs or treelets 0.7–4.0 m tall. Branchlets quadrangular, always
glandular-punctate and usually pruinose-granulose, eventually sparsely to densely cov-
ered with gland-tipped trichomes 0.1–0.9 mm long, rarely covered with rigid hyaline
eglandular trichomes 0.1–0.5 mm long, light green (when fresh). Internodes 0.6–3.8cm
long, angles unwinged. Petioles 1–15 mm long. Leaf blades 12–117 mm long, 3–38 mm
wide, papyraceous (when dry), elliptic to narrowly elliptic or lanceolate, rarely obovate,
adaxial surface green and partially covered by a thin layer of whitish indumentum, abaxial
surface totally concealed by the white lanose indumentum (when fresh), adaxial surface
blackened and abaxial surface pale green to pale brown (when dry), discoloured (when
dry), base attenuate, apex acute to obtuse, margin at or slightly revolute, entire to slight-
ly serrulate and glabrescent, granulose or granular-punctate or ciliate with gland-tipped
trichomes 0.1–0.9 mm long, 5-nerved from the base, one pair of acrodromous veins and
one pair of tenuous veins close to the margin, tertiaries evident on the abaxial surface,
nearly perpendicular to the mid-vein, reticulate and randomly branching, adaxial surface
glandular-punctate, usually pruinose and becoming glabrescent with age, abaxial sur-
face glandular-punctate and usually pruinose, eventually sparsely to densely covered with
gland-tipped or eglanduar trichomes 0.1–0.9 mm long. Inorescences simple or com-
pound dichasia consisting of with biparous cymes, not congested. Bracts (including peti-
oles) 0.7–1.6 cm long, 0.1–0.6 cm wide, 3-nerved, elliptic to narrowly elliptic, indumen-
tum like that of the principal leaves. Bracteoles (at anthesis) with petioles 0.9–1.9 mm
long, blades 2.2–3.8 mm long, 0.5–1.0 mm wide, narrowly elliptic to oblanceolate,
base attenuate, apex acute to obtuse, margin entire, 1–3-nerved, indumentum like that
of the principal leaves. Flowers 5(–6)-merous, pedicels (at anthesis) 1.1–2.0 mm long.
Hypanthia (at anthesis) 1.9–2.8 mm long, 1.5–2.2mm wide at the torus, campanulate
to urceolate, light green, sometimes with reddish stains (when fresh), externally always
glandular-punctate and usually pruinose, eventually sparsely to densely covered with
gland-tipped trichomes 0.1–0.9 mm long, rarely covered with rigid hyaline eglandular
trichomes 0.1–0.5 mm long. Calyx tubes 0.2–0.7 mm long. Calyx lobes (at anthesis)
0.7–2.5(–3.0) mm long, 0.9–1.5 mm wide at the base, triangular, apex acute, acuminate
or apiculate, margin entire, (when fresh) light green or reddish, externally like the hyp-
anthia. Petals 4.5–8.1mm long, 3.0–4.9mm wide, white, usually ushed with pink at
the base and around the veins, rarely entirely pink, obovate, apex emarginate, rounded
or acute, margin entire, glabrous or ciliate with eglandular or gland-tipped trichomes
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
62
Figure 24. Microlicia parviora A habit B leaf abaxial surface C bracteole abaxial surface D ower in
lateral view E owering hypanthium and pedicel F petal adaxial surface G detail of indumentum on the
apex of the petal H antepetalous (left) and antesepalous (right) stamens I gynoecium J ovary in cross-
section K capsule enveloped by the hypathium. Drawn from Souza-Buturi 328 (UEC) and Matsumoto
et al. 428 (UEC).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 63
0.1–0.4 mm long at the apical region, both surfaces glabrous. Stamens 10(–12), strongly
dimorphic. Larger (antesepalous) stamens 5(–6), laments 3.5–4.0 mm long, white or
pink, pedoconnectives 3.0–4.0mm long, white or pink, appendages 0.7–1.5 mm long,
yellow, apex emarginate to bilobate, thecae (excluding rostra) 0.8–1.3 mm long, red to
vinaceous, oblong, rostra 0.2–0.4 mm long, the circular pores 0.1–0.2 mm wide. Smaller
(antepetalous) stamens 5(6), laments 2.0–3.1 mm long, white or pink, pedoconnectives
0.2–0.5 mm long, white or pink, inconspicuous appendages ca. 0.1 mm long, yellow
or pink, apex emarginate, thecae (excluding rostra) 0.8–1.3 mm long, yellow, oblong,
rostra 0.2–0.4mm long, the circular pores ca. 0.2 mm wide. Ovary 1.8–2.2 mm long,
1.6–1.8 mm wide, globose, 5-locular. Style 3.0–3.6 mm long, white or pink. Capsules
(at maturity) 2.9–5.0 mm long, 2.5–4.0 mm wide, globose, initially enveloped by the
hypanthium, torus constricted at the apex, fruiting calyx tubes 0.2–1.0 mm long, fruiting
calyx lobes 1.2–2.9(–3.5) mm long, not thickened. Seeds 0.3–0.6 mm long, reniform.
Representative specimens (one specimen selected for each municipality of oc-
currence). Bahia: Abaíra, Ganev 879 (HUEFS, NY, SPF); Andaraí, Orlandi et al. 778
(MBM); Lençóis, Carvalho 1086 (CEPEC, RB, SPF); Miguel Calmon, Guedes et al.
12092 (ALCB); Morro do Chapéu, Hage et al. 2317 (CEPEC, MBM, RB); Mucugê,
Roque 2869 (ALCB); Palmeiras, Bautista 1347 (CEPEC); Piatã, Ganev 965 (HUEFS,
SPF); Ribeirão do Largo, Carvalho 6986 (CEPEC, NY); Rio de Contas, Carvalho
et al. 6651 (CEPEC); Seabra, Irwin et al. 31141 (NY); Utinga, Samento & Bautista
859 (RB); Wenceslau Guimarães, Goldenberg & Michelangeli 2093 (UPCB). Distrito
Federal: Brasília, Azevedo et al. 675 (CAS); Taguatinga, Irwin et al. 8149 (SP). Espírito
Santo: Domingos Martins, Pereira 309 (CEPEC, SP); Dores do Rio Preto, Monge et
al. 2595 (UEC); Fundão, Kollmann 231 (RB, UPCB); Itaguaçu, Brade et al. 18205
(RB); Iúna, Fontana et al. 7678 (UPCB); Marechal Floriano, Hatschbach et al. 74974
(FURB, HCF, MBM); Santa Rita de Jacutinga, Krieger 8857 (MBM); Santa Teresa,
Martinelli et al. 10932 (RB). Goiás: Alto Paraiso de Goiás, Marquete et al. 2332 (RB,
US); Cocalzinho de Goiás, Versiane et al. 305 (HUFU, RB); Corumbá de Goiás, Irwin
et al. 34521 (NY); Cristalina, Monteiro 78 (RB, SPF, UPCB); Pirenópolis, Delprete
9205 (RB). Minas Gerais: Aiuruoca, Glaziou 9454 (P); Alagoa, Guiamarães 333 (RB);
Alto Caparaó, Hatschbach & Guimarães 55455 (MBM); Baependi, Souza et al. 1013
(CESJ, MBM); Barão de Cocais, Fontana 2309 (RB, UPCB); Barbacena, Barreto 4621
(SP); Barroso, Assis et al. 520 (MBM); Belo Horizonte, Vidal s.n. (CEN [46402]); Boa
Esperança, Silva s.n. (UPCB [52391]); Bom Jardim de Minas, Krieger et al. 24423
(SPF); Brumadinho, Martens 524 (SPF); Buenópolis, Davis et al. 2298 (UEC); Bur-
itizeiro, Hatschbach et al. 75995 (MBM); Caeté, Paula & Grandi s.n. (BHCB [7956],
MBM [178326]); Carandaí, Costa 451 (RB); Carangola, Leoni 1 (SPF); Carmésia,
Stehmann 2532 (ESA); Carrancas, Sobral et al. 14085 (RB); Catas Altas, Oliveira et
al. 508 (BHCB, RB); Conceição do Ibitipoca, Oliveira 25197 (CESJ, RB); Conceição
do Mato Dentro, Guarçoni & Sartori 1367 (HUFU); Coromandel, Brandão 14509
(HUFU); Cristália, Hatschbach 41498 (MBM, US); Delnópolis, Romero & Na-
kajima 3432 (HUFU); Diamantina, Hatschbach & Pelanda 28014 (MBM); Espera
Feliz, Foster & Leoni 64 (ESA); Estrela do Sul, Costa et al. 60 (HUFU); Extrema,
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
64
Yamamoto 1549 (UEC); Gouveia, Hatschbach 27277 (MBM); Grão Mogol, Caval-
canti CFCR8314 (SPF); Itabira, Faria et al. 1332 (HUFU); Itabirito, Brandão 22298
(HUFU); Itamonte, Batista & Naves 405 (UEC); João Pinheiro, Heringer 8544/736
(UB, US); Joaquim Felício, Cavalcanti et al. CFCR8035 (SPF, UEC); Lagoa Santa,
Warming s.n. (BR [BR0000005520732], C [C10015107], P [P005317116]); Lavras,
Avezum & Almeida 10 (SPF); Lima Duarte, Heluey & Castro 108 (RB); Manhuaçu,
Hatschbach & Silva 49393 (HUEM, MBM); Mariana, Lombardi 4045 (BHCB, ESA);
Moeda, Silva & Grandi 6627 (HUFU); Nova Lima, Williams & Assis 7274 (SP); Ouro
Branco, Delni et al. 97 (ESA, RB); Ouro Preto, Colletta 161 (SPF), Martius 931
(BM, G, M, MO, P); Paracatu, Bovini & Barros 3235 (RB); Passa Quatro, Meireles et
al. 1766 (RB, UEC); Patrocínio, Farah et al. 580 (ESA, HUEM); Perdizes, Mendes &
Araújo 970 (SPF); Piuhmhi, Emygdio 3613 (NY, R); Poços de Caldas, Oliveira 1013
(US); Prados, Sobral et al. 12797 (UPCB); Presidente Soares, Hatschbach et al. 55434
(MBM); Rio Acima [Gandarela], Emygdio 3312 (NY, R); Rio Pardo de Minas, Sevilha
et al. 7075 (CEN); Rio Preto, Barros & Feteira 1625 (RB); Rio Vermelho, Mello-Silva
et al. CFCR7836 (SPF); Rosário de Limeira, Marcolino 222 (RB); Sabará, Barreto 6759
(BHCB, SP); Sacramento, Romero et al. 2155 (MBM); Santa Bárbara, Barreto 6757
(BHCB, SP); Santa Bárbara do Monte Verde, Pivari 15 (MBM); Santana de Pirapama,
Zappi et al. 2504 (SPF); Santana do Riacho, Pacico 191 (HUEM); Santos Dumont,
Mello-Silva et al. 1217 (SPF); São Gonçalo do Rio Preto, Foresto et al. 59 (SPF); São
Gonçalo do Sapucaí, Hatschbach 26964 (MBM); São João Batista da Glória, Kinoshita
et al. 43767 (HUEM); São João del Rei, Barreto 4654 (SP, US); São Roque de Minas,
Pacico 413 (CAS, HUEM); São Tomé das Letras, Valente & Azevedo 57 (RB); Serro,
Almeda et al. 9076 (CAS, UEC); Tapira, Salgado 167 (RB); Tiradentes, Alves 600 (R);
Tombos, Fraga & Saddi 1786 (CEPEC, RB); Uberlândia, Romero & Nakajima 3021
(HUFU); Unknown municipality in Minas Gerais State, Gardner 4602 (NY, R, US),
Martius 961 (G, M), Mosen 1971 (C, P, R), Saint-Hilaire s.n. [D462] (P [P00723414],
P [P00723415]), Sellow 1154 (US), Sellow 5278 (P), Vauthier 43 [part] (BR, G, P),
Widgren 967 (BR). Paraná: Adrianópolis, Camargo et al. 109 (UPCB); Antonina,
Hatschbach & Guimarães 56167 (MBM); Araponga, Caiafa & Umbelino 172 (UPCB,
VIC); Arapoti, Hatschbach 6908 (MBM); Balsa Nova, Hatschbach et al. 42967
(MBM); Bocaiúva do Sul, Ribas et al. 6769 (MBM, UPCB); Campina Grande do Sul,
Brotto & Vieira 1921 (MBM); Colombo, Hatschbach 647 (MBM, RB); Jaguariaíva,
Ribas et al. 8528 (MBM); Ortigueira, Silva et al. 6478 (MBMB, UNOP); Palmeira,
Hatschbach 2775 (MBM); Piraí do Sul, Goldenberg et al. 1652 (NY, UPCB); Ponta
Grossa, Silva & Koch 7610 (MBM); Rio Branco do Sul, Silva & Abe 2310 (MBM);
Sengés, Hatschbach 39954 (MBM); Tibagi, Kirizawa 3665 (SP); Tunas do Paraná,
Brotto 2600 (MBM, RB); Ventania, Estevan et al. 617 (UPCB). Rio de Janeiro: Bom
Jardim, Hottz et al. 302 (MBM, RB); Duque de Caxias, Lima et al. 8445 (RB); Itatiaia,
Baumgratz et al. 1127 (MBM, SPF); Miguel Pereira, Wängler & Ferreira 1352 (RB);
Nova Friburgo, Forzza et al. 3427 (RB); Petrópolis, Vieira & Yamamoto 26186 (FUEL,
RB, UEC); Resende, Eiten & Eiten 7305 (NY, P, RB, SP, US); Rio Claro, Moutinho
et al. 76 (RB); Santa Maria Madalena, Lima 400 (US); Teresópolis, Duarte & Brade
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 65
1155 (RB); Unknown municipality in Rio de Janeiro state, Gardner 379 (BR, NY, P,
US), Gardner 380 (BR, F, G, NY, P, S, US), Guillemin 921 (P), Guillemin 946 (G,
P), Widgren s.n. (BR [BR0000005227044], PH-27744, S-53231). São Paulo: Apiaí,
Souza et al. 6098 (ESA, RB); Bananal, Martinelli 19464 (RB); Bom Sucesso de Itararé,
Aguiar 102 (MBM); Brotas, Queiroz 2808 (CEPEC); Campos do Jordão, N. da Cruz
135 (CAS, MBM); Cunha, Mamede et al. 666 (RB, SP); Eldorado, Pastore et al. 685
(RB); Iporanga, Souza et al. 5934 (SPF); Itapeva, Baitello et al. 2136 (UPCB); Itirapi-
na, Tannus 760 (HUFU); Lavrinhas, Caddah et al. 636 (UPCB); Mogi Mirim, Hoehne
20521 (NY, SP); Pedregulho, Polisel et al. 176 (UPCB); Pindamonhangaba, Nicolau
et al. 2212 (SP); Piquete, Gonçalves et al. 172 (RB); Rio Claro, Loefgren 557 (BHCB,
SP); Santo André, Kirizawa et al. 2132 (SP); São Bernardo do Campo, Kuhlmann 4381
(SP); São Carlos, Eiten et al. 3019 (SP, US); São José do Barreiro, Handro 790 (NY);
São José dos Campos, Mimura 479 (SP, US); São Paulo, Beraldo & França 85 (SPF);
Ubatuba, Souza et al. 1108 (UPCB); Unknown municipality in São Paulo State, Prates
s.n. (P [P00723407], P [P00723406, P [P00723408]). Unknown state: Bunbury s.n.
(BR [BR0000005520374]), Glaziou 12704 (BR), Glaziou 16679 (BR), Glaziou 16778
(C, L, P), Glaziou 2579 (BR, P), Glaziou 8680 (BR, P), Glaziou 9454 (BR, C, G, P, R,
S), Martius 989 (BR), Raben 409 (BR), Riedel s.n. (BR [BR0000005520367]), Sellow
s.n. (lectotype: G [G00396696], probable isolectotype: P [P05317745]).
Distribution, habitat and elevation range. Endemic to Brazil in the States of
Bahia, Minas Gerais, Goiás, Distrito Federal, Espírito Santo, Rio de Janeiro, São Paulo
and Paraná (Fig. 25A). Trembleya parviora is common in gallery forests surround-
ing campo rupestre, Cerrado, campo sujo, campo limpo. campos de altitude, Veredas
(palm swamps), and margins of roads throughout Cerrado and Atlantic forest frag-
ments, usually exposed to full sun, at elevations between 560 and 2223 m.
Ecology. Microlicia parviora is the most widely distributed species in the clade
and the most studied from an ecological perspective. Giotto (2015) investigated plant
occupation in palm swamps with very dense populations of M. parviora, where seed-
lings of this species corresponded to 78% of the total seed bank. Silva (2003) re-
ported a population density of 1.47 individuals per square metre in these areas. is
elevated local dominance of M. parviora negatively aects the overall plant species
richness of these regions and is apparently favoured by reduction in humidity (Giotto
2015), although M. parviora occurs with large populations in both wet and dry areas
(Melo 2013). is species is strongly recommended for ecological restoration (Amaral
et al. 2013) because it readily colonises degraded campo rupestre. Leaf extracts of
M. parviora proved to have allelopathic properties, inhibiting root and shoot growth
of Sesamum indicum (Borghetti et al. 2005). According to Pereira (2011), leaves of
M. parviora develop increased asymmetry when damaged by insects.
Conservation. is is the most abundant species in the Trembleya s.s. clade, with
more than 1000 collections currently housed in herbaria. e EOO is 1,152,078.308
km2 and the AOO is 1,640 km2. Populations of M. parviora are found in all conser-
vation units that protect the remaining species of the clade. As it occurs in large and
dense populations, some have claimed that M. parviora has the potential to become
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
66
an invasive species (e.g. Silva 2003). We are not aware of records of this species occupy-
ing regions outside its natural distributional range. Based on the IUCN (2019) recom-
mendations and criteria, we recommend that a conservation status of Least Concern
(LC) be assigned to this species.
Recognition and anities. Microlicia parviora can be distinguished from its
congeners by its leaves that are 5-nerved from the base, the abaxial surface glandular-
punctate, sometimes covered with an indumentum of pedicellate trichomes (never
dense enough to conceal the epidermis), three to many-owered inorescences, trian-
gular calyx lobes 0.7–2.5(–3) mm long, petals white or pink and dimorphic stamens.
Microlicia pentagona and M. trembleyiformis are somewhat similar to M. parviora in
leaf shape and venation and stamen morphology. Microlicia parviora diers from
M. pentagona by its leaves with tertiaries evident (vs. little evident), moderately reticu-
late and randomly branching (vs. parallel and/or branching apically), inorescences
(vs. solitary owers) and triangular calyx lobes 0.7–2.5(–3) mm long (vs. subulate and
6.2–8.5 mm long) that are tenuous in fruit (vs. thickened). In turn, M. parviora diers
from M. trembleyiformis by the unwinged branchlets (vs. with narrow wings ca. 0.2 mm
wide), owers disposed in inorescences (vs. owers solitary), triangular calyx lobes (vs.
narrowly-triangular) and consistently 5-locular ovaries (vs. 3–5-locular). For a com-
parison between M. parviora and M. altoparaisensis, see notes under the latter species.
Notes. e morphological variation found in M. parviora is mainly related to leaf
blade shape (varying from elliptic to ovate and lanceolate), indumentum on branches
Figure 25. Geographic distribution of species of the Trembleya s.s. clade of Microlicia A distributions
of M. parviora and M. tridentata B distributions of M. pentagona, M. pithyoides and M. rosmarinoides.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 67
and abaxial leaf surface (always glandular-punctate, usually pruinose, eventually
sparsely to densely covered with gland-tipped or eglandular trichomes 0.1–0.9 mm
long) and inorescence development (three- to many-owered). e petals are usually
white ushed with pink at the base, although some populations have entirely white
petals, for example, in Serra de Carrancas (Matsumoto and Martins 2005) and Serra
Negra (Justino et al. 2016) in Minas Gerais. Based on descriptions of M. parviora
in local oras, magenta-owered and white-owered variants of this species occur to-
gether in Poços de Caldas and Paque Estadual do Ibitipoca, both in Minas Gerais. In
the Serra da Canastra, Romero (2000: 283) recognised a variant of M. parviora as a
distinct entity (Trembleya sp.) characterised by tertiaries more evident, trichomes on
the abaxial leaf surface with 3–4-lobed heads, inorescence internodes 7–9 mm long
and calyx lobes 2.5–3 mm long. Several other modal extremes were designated as types
of infraspecic taxa by Cogniaux (1883–1888, 1891), for example, the specimen Prates
s.n. (P [P00723407]; type of var. denticulata) has more conspicuous serrations on the
leaf margin, whereas Guillemin 946 (P; type of var. tomentosa) has a denser indumen-
tum on the branchlets. However, none of the mentioned variations is correlated with
other characteristics in a meaningful way.
e 15 varieties of M. parviora proposed by Cogniaux (1883–1888, 1891) were
based exclusively on dierences in leaf blade size, shape and apex and degree of ino-
rescence development. Based on a comprehensive sampling and analysis of representa-
tive collections from throughout the range of the species, it is clear that Cogniaux’s
infraspecic taxonomy is articial and untenable. In fact, many of the specimens
examined had leaf sizes that do not t into any of the infraspecic taxa proposed
by Cogniaux (1883–1888, 1891). For example, the specimen Kinoshita et al. 10/19
(UEC) has leaves 1.5–8 cm long and could t all varieties proposed, based on this fea-
ture. e features that Cogniaux (1883–1888) used to circumscribe the two subspecies
are also imprecise and not diagnostic. After analysing several representative collections
of M. parviora, we did not nd any specimen with clearly tetragonal and totally
glabrous branches, both purported diagnostic features of M. parviora subsp. triora.
us, given the highly polymorphic nature of M. parviora, we do not recognise any
infraspecic taxa in this species.
Like Microlicia cataphracta (Mart. & Schrank ex DC.) Versiane & R.Romero [=
Lavoisiera imbricata (unb.) DC.], M. parviora ts all the six criteria enumerated
by Cronk (1998) to be considered as an ochlospecies (see Martins and Almeda 2017).
Besides sharing outstanding levels of morphological variation, M. cataphracta and
M. parviora also have similar distributional ranges and usually occur in large popula-
tions. e reproductive biology, population characteristics, chemical composition and
other ecological aspects of M. parviora are summarised in other sections of this paper.
Martin and Cremers (2007) examined a supposed holotype of Trembleya paniculata
at P. However, we found two duplicates of the same collection used by Naudin (1844)
to describe Trembleya paniculata (Saint-Hilaire s.n., P [P00723414], P [P00723415]).
us, we designated one of these sheets as the lectotype (P [P00723414]) and the other
as an isolectotype (P [P00723415]) for Trembleya paniculata.
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
68
7. Microlicia pentagona (Naudin) Versiane & R.Romero, Bot. J. Linn. Soc. 197:
54. 2021.
Fig. 26
Trembleya pentagona Naudin, Ann. Sci. Nat., Bot. Sér. 3, 2: 154. 1844. basionym.
Type: Brazil. “In montibus vulgo Serra d’Ouro Branco, provincia Minas-Geraes”
[Minas Gerais, Ouro Branco], A. Saint-Hilaire s.n. (lectotype, rst-step designated
by Martin and Cremers (2007), second-step designated here: P [P00723399]!;
isolectotype: P [P00723398]!; image of lectotype is available at http://coldb.mnhn.
fr/catalognumber/mnhn/p/p00723399).
Description. Erect shrubs or treelets 0.5–1.5 m tall. Branchlets quadrangular, appear-
ing glabrous, vernicose and minutely granulose, vinaceous (when fresh). Internodes
0.4–2.5 cm long, angles unwinged. Petioles 1.8–2.4 mm long. Leaf blades 12–40 mm
long, 3–15 mm wide, chartaceous to coriaceous (when dry), elliptic, ovate, narrowly el-
liptic or linear, both surfaces green (when fresh), adaxial surface blackened and abaxial
surface pale green to pale brown (when dry), discoloured (when dry), base cuneate to
attenuate, apex rounded to acute, margin at, entire along the basal half, serrulate on
the upper half and minutely granulose and becoming glabrescent with age, 5-nerved
from the base, one pair of acrodromous veins and one pair of tenuous veins close to
the margin, tertiaries little evident on the abaxial surface, nearly perpendicular to the
mid-vein, parallel or little reticulate and branching apically, adaxial surface glabrous to
minutely granulose, vernicose, abaxial surface glabrous to minutely granulose, vernic-
ose. Inorescences reduced to solitary owers apically on the branches. Bracts absent.
Bracteoles (at anthesis) with petioles 1.8–3.0 mm long, blades 3.0–6.5 mm long, 1.1–
3.6 mm wide, elliptic, ovate or lanceolate, base attenuate to cuneate, apex acute to
obtuse, margin entire, 3-nerved, indumentum like that of the principal leaves. Flowers
(4–)5-merous, pedicels (at anthesis) 0.8–2.0 mm long. Hypanthia (at anthesis) 2.5–
3.5mm long, 2.7–3.0 mm wide at the torus, campanulate to urceolate, light green or
reddish (when fresh), externally glabrous, minutely granulose, vernicose. Calyx tubes
0.2–0.4 mm long. Calyx lobes (at anthesis) 6.2–8.5 mm long, 1.8–2.2mm wide at the
base, subulate, apex acute, margin entire, (when fresh) light green or reddish, externally
like the hypanthia. Petals 11.8–13.8 mm long, 7.1–9.5 mm wide, magenta, obovate,
apex shortly acuminate, margin entire and glabrous, both surfaces glabrous. Stamens
(8)10, strongly dimorphic. Larger (antesepalous) stamens (4)5, laments 4.0–5.0mm
long, pink, pedoconnectives 5.3–5.8 mm long, pink, appendages 1.2–1.6mm long,
yellow, apex emarginate to bilobate, thecae (excluding rostra) 1.2–2.0mm long, vina-
ceous, oblong, rostra 0.5–0.7 mm long, the circular pores ca. 0.2 mm wide. Smaller
(antepetalous) stamens (4)5, laments 3.9–4.4 mm long, pink, pedoconnectives 1.0–
1.6 mm long, yellow, inconspicuous appendages ca. 0.1mm long, yellow, apex emar-
ginate, thecae (excluding rostra) 1.6–2.0 mm long, yellow, oblong, rostra 0.5–0.6 mm
long, the circular pores ca. 0.2 mm wide. Ovary 2.5–2.7 mm long, 2.4–2.6 mm
wide, globose, 5-locular. Style 6.0–6.6 mm long, magenta. Capsules (at maturity)
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 69
Figure 26. Microlicia pentagona A habit B leaf abaxial surface C bracteole abaxial surface D oral
bud E ower in lateral view F owering hypanthium G petal adaxial surface H antesepalous (left) and
antepetalous (right) stamens I gynoecium J capsule enveloped by the hypanthium. Drawn from Sazima
CFSC4259 (UEC) and Almeda et al. 9203 (UEC).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
70
4.5–5.5 mm long, 5.0–6.0 mm wide, globose, initially enveloped by the hypanthium,
torus constricted at the apex, fruiting calyx tubes 0.2–0.4 mm long, fruiting calyx lobes
6.5–11.0 mm long, stout, thickened. Seeds 0.5–0.7 mm long, reniform.
Distribution, habitat and elevation range. Endemic to Minas Gerais State
(Fig. 25B) in Serra do Caraça, Serra do Cipó, Serra de Ouro Preto, Serra de Ouro
Branco, Serra do Itabirito, Mariana and Barão de Cocais. It occurs on quartzitic and
ferruginous campo rupestre, exposed to full sun, at elevations of about 1000–1830 m.
Conservation. Microlicia pentagona is known from about 70 collections. e EOO
is 2,667.623 km2 and the AOO is 56 km2. Several populations are protected within the
following conservation units: Parque Estadual do Itacolomi, Parque Estadual Serra do
Intendente, Parque Estadual Serra do Ouro Branco, Parque Nacional da Serra do Cipó
and RPPN Serra do Caraça (Natural Heritage Private Reserve). Following the IUCN
(2019) recommendations and criteria, we recommend an Endangered (EN): B1ab(iii)
status for this species.
Recognition and anities. Microlicia pentagona can be recognised by its branches,
leaves and hypanthia that are glandular-punctate and vernicose, leaves 5-nerved from
the base with one pair of acrodromous veins and one pair of tenuous veins close to the
margin, inorescences reduced to solitary terminal owers and calyx lobes 6.2–8.5 mm
long that become thickened in fruit. e elongate calyx lobes of M. pentagona are com-
parable in length only to those of M. laniora (7.9–9.7 mm long) and are consistently
longer than those of all remaining congeners. In fruit, the thickened calyx lobes of
M. pentagona are unique in the clade. Overall, M. pentagona is morphologically similar
to M. calycina, M. chamissoana, M. laniora and M. parviora. For comparisons, see
comments under these species.
Notes. Microlicia pentagona is remarkably variable in leaf shape and size. e leaf
blades vary from almost linear (e.g. Irwin et al. 20537, Barreto 10734) to narrowly elliptic
(e.g. Barreto 7025, Joly et al. CFSC 3196) and elliptic (e.g. Silva 1043). Examples of vari-
ation in leaf size are Almeda et al. 9203 (1.7–2.1 cm long) and Silva 1043 (2.3–3.1 cm
long). Some modal extremes were informally recognised as distinct taxa by Mello Barreto
on herbaria labels. As no character varies in a meaningful way, we agree with Martins
(1997) and treat all these extremes within the variation here attributed to M. pentagona.
Specimens examined. Brazil. Minas Gerais: Barão de Cocais Municipality,
Brandão 20838 (HUFU); Catas Altas Municipality, Serra do Caraça, Hatschbach et al.
s.n. (HUFU [25601]), Irwin et al. 29213 (NY), Oliveira et al. 478 (BHCB), Oliveira
et al. 480 (BHCB, RB), Oliveira et al. 518 (BHCB), Silva et al. 1043 (HUFU), So-
bral et al. 14508 (HUFSJ, RB); Santa Bárbara Municipality [“Capanema”], Claussen
296 (W), Vainio 33501 (US); Jaboticatubas Municipality [“Caeté”], Serra do Cipó,
Magalhães 2278 (BHCB, US); Mariana Municipality, Pivari et al. 2512 (BHCB);
Ouro Branco Municipality, Saint-Hilaire s.n. (lectotype: P [P00723399]; isolectotype:
P [P00723398]); Ouro Preto Municipality, Antônio-Silva et al. 315 (HUFU, OUPR),
Bortoluzzi et al. 678 (RB, VIC), Damazio s.n. (RB [48352], NY [NY00942037]), Mi-
chelangeli et al. 1580 (NY, UPCB), Pacico & Bressan 294 (HUEM, SPF), Peron 256
(RB), Peron 260 (RB), Rezende 507 (HUFU), Rolim et al. 353 (UPCB, VIC), Rolim
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 71
et al. 361 (NY, RB, VIC), Rolim et al. 370 (HUFU, NY, RB, VIC), Schwacke 10814
(W), Schwacke 9325 (RB, W); Santa Bárbara Municipality, Serra do Caraça, Almeda et
al. 7751 (CAS, HUFU, UEC), Damazio s.n. (RB [48392]), Ordones et al. 213 (BHZB,
HUFU), Pirani & Yano 692 (RB, SP, SPF), Rapini et al. 296 (HUEM, SP, SPF), Rome-
ro et al. 5303 (CAS, UEC); Santana do Riacho Municipality, Serra do Cipó, Almeda
et al. 8555 (CAS, UEC), Almeda et al. 8911 (CAS, UEC), Almeda et al. 9203 (CAS,
HUEM, UEC), Barreto 1182 (RB), Contro & Marques 21 (HUEM, HUFU), Duarte
2004 (FLOR, MBM, RB), Fernandes s.n. (HUFU [56535]), Irwin et al. 20178 (CAS,
MO, NY, UEC, US), Irwin et al. 20486 (CAS, MO, NY, US), Irwin et al. 20537
(CAS, NY, UEC, US), Pacico & Bressan 531 (CAS, HUEM, RB). Romero & Na-
kajima 5998 (HUEM, HUFU, UEC), Semir CFSC5608 (SP, SPF), Semir & Sazima
CFSC3395 (UEC); Santana do Riacho Municipality [“Conceição do Mato Dentro”],
Serra do Cipó, Barreto 7025 (BHCB, HB, UEC); Santana do Riacho Municipality
[“Jaboticatubas”], Serra do Cipó, Giulietti & Menezes 4017 (SP, SPF, UEC), Hatsch-
bach et al. 28759 (MBM, SPF, US), Hatschbach et al. 29958 (MBM), Joly & Semir
CFSC2957 (UEC), Joly & Semir CFSC3196 (SPF, UEC), Sazima CFSC4259 (RB,
SP, UEC), Semir & Joly CFSC3743 (UEC); Santana do Riacho Municipality [“Santa
Luzia”], Serra do Cipó, Barreto 10734 (HB, UEC), Barreto 7023 (BHCB), Barreto
7024 (UEC), Brade 14756 (R), Duarte 2692 (R, RB, US); Unknown municipality
in Minas Gerais State, Casaretto s.n. (G [G00318565]), Claussen 1617 (P), Claussen
1637 (P), Claussen 22 (P, US), Claussen 350 (W), Claussen s.n. (P [P005317034], P
[P005317035]), Glaziou 14742 (K, NY, P, R), Ule 2543 (US), Schüch s.n. (W [70510]).
8. Microlicia pithyoides (Cham.) Versiane & R.Romero, Bot. J. Linn. Soc. 197:
54. 2021.
Fig. 27
Trembleya pithyoides Cham., Linnaea 9(4): 428. 1835. basionym. Type: Brazil.
“Caraça” [Minas Gerais, Serra do Caraça], 20 December 1830, F. Sellow 1316 (lec-
totype, designated here: K [K00530665]!; isolectotypes: F-BN016638-photo]!, P
[P00723396]!, P [P00723397]!; image of isolectotype at P is available at http://
coldb.mnhn.fr/catalognumber/mnhn/p/p00723396).
Trembleya pithyoides var. major Cogn. in Martius et al., Fl. Bras. 14(4): 594. 1888. syn.
nov. Type: Brazil. “Minas, Serra de Capanema” [Minas Gerais, Santa Bárbara], 21
February 1884, A.F.M. Glaziou 14746 (lectotype, designated here: P [P00723395]!;
isolectotypes: BR [BR0000005223930]!, BR [BR0000005520169]!, C
[C10015114-online image]!, C [C10015113-online image]!, K [K00530666]!, P
[P00723394]!, P [P00723393]!; image of lectotype is available at http://coldb.
mnhn.fr/catalognumber/mnhn/p/p00723395).
Description. Erect, densely-branched shrubs 0.3–0.6 m tall. Branchlets quadrangular,
glandular-punctate, vinaceous (when fresh). Internodes 0.2–0.5 cm long, angles with
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
72
narrow wings ca. 0.2 mm wide. Petioles 0.7–1.5 mm long. Leaf blades 4–15 mm long,
0.5–1.4 mm wide, chartaceous (when dry), linear, both surfaces green (when fresh),
adaxial surface blackened and abaxial surface pale green to pale brown (when dry), dis-
coloured (when dry), base cuneate, apex rounded to acute, margin at, entire and glan-
dular-punctate, 1-nerved from the base, tertiaries not evident on the abaxial surface,
adaxial surface glandular-punctate, abaxial surface glandular-punctate. Inorescences re-
duced to solitary owers apically on the branches. Bracts absent. Bracteoles (at anthesis)
with petioles 0.6–0.9 mm long, blades 3.0–3.9 mm long, 0.3–0.5mm wide, lanceolate,
base cuneate, apex acute to obtuse, margin entire, 1-nerved, indumentum like that of
the principal leaves. Flowers 5-merous, pedicels (at anthesis) 0.3–0.6 mm long. Hypan-
thia (at anthesis) 1.7–2.0 mm long, 1.9–2.1 mm wide at the torus, campanulate, light
green or reddish (when fresh), externally glandular-punctate. Calyx tubes 0.3–0.4 mm
long. Calyx lobes (at anthesis) 2.5–3.0 mm long, 0.4–0.7mm wide at the base, subu-
late, apex acute, margin entire, (when fresh) light green or reddish externally like the
hypanthia. Petals 5.0–5.7 mm long, 3.7–4.2 mm wide, magenta, obovate, apex acumi-
nate, margin entire and glabrous, adaxial surface glabrous or sparsely glandular-punc-
tate, abaxial surface glabrous. Stamens 10, strongly dimorphic. Larger (antesepalous)
stamens 5, laments 2.6–2.9 mm long, pink, pedoconnectives 2.9–3.2mm long, pink,
appendages 0.7–0.9 mm long, yellow, apex emarginate to bilobate, thecae (excluding
rostra) 1.8–2.0 mm long, vinaceous, oblong, rostra 0.2–0.4mm long, the circular pores
ca. 0.2 mm wide. Smaller (antepetalous) stamens 5, laments 1.9–2.1mm long, pink,
pedoconnectives 0.6–0.9 mm long, yellow, inconspicuous appendages ca. 0.1 mm long,
yellow, apex emarginate, thecae (excluding rostra) 1.4–1.6mm long, yellow, oblong,
rostra 0.2–0.3 mm long, the circular pores ca. 0.2 mm wide. Ovary 1.3–1.5 mm long,
1.0–1.2 mm wide, globose, (4–)5-locular. Style 4.0–4.5 mm long, pink. Capsules (at
maturity) 2.7–3.1 mm long, 2.9–3.3 mm wide, globose, initially enveloped by the hy-
panthium, torus constricted at the apex, fruiting calyx tubes 0.4–0.6 mm long, fruiting
calyx lobes 3.2–3.7 mm long, not thickened. Seeds 0.7–0.8mm long, reniform.
Specimens examined. Brazil. Minas Gerais: Catas Altas Municipality, Serra
do Caraça, Oliveira 382 (BHCB, RB), Pacico 295 (CAS, HUEM, SPF); Unknown
municipality in Minas Gerais State, Serra do Caraça. Glaziou 14746 (BR, C, K, P,
R), Glaziou 19239 (K, P), Sellow 1316 (lectotype: K [K00530665; isolectotypes: F-
BN016638-photo, P [P00723396], P [P00723397]), Sellow s.n. (K [K00957781, K
[K00957783]), Weddell s.n. (P [P005317975]).
Distribution, habitat and elevation range. Known only from Minas Gerais State
(Fig. 25B), where it is probably endemic to the Serra do Caraça. Microlicia pithyoides
grows on quartzitic campo rupestre exposed to full sun at elevations between 1827
and 2072 m. It is the only species in the clade that reaches the highest peak in Cadeia
do Espinhaço, the Pico do Sol in Catas Altas Municipality (elev. 2,072 m) (personal
observation by R. Pacico).
Conservation. Microlicia pithyoides species is the least collected species of the
Trembleya s.s. clade and apparently has the narrowest distribution. Less than 10 col-
lections of this species are housed in herbaria. It had not been collected for more than
a hundred years until it was re-discovered in 2009 (Oliveira 382). As all coordinates
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 73
available for M. pithyoides refer to the same population, we were unable to calculate its
EOO. e AOO is 4 km2. e only population known of this species occurs inside a
private protected area, the RPPN Serra do Caraça (Natural Heritage Private Reserve).
e type material was probably collected at the Serra de Capanema in Santa Bárbara
Figure 27. Microlicia pithyoides A habit B detail of nodes and internodes C leaf abaxial surface
D bracteole abaxial surface E oral bud F ower in lateral view G petal adaxial surface H antesepalous
stamen I antepetalous stamen J capsule enveloped by the hypanthium K capsule in cross section. Drawn
from Pacico 295 (CAS).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
74
Municipality. Currently, this locality is also part of a private property. We are not aware
of recent collections of M. pithyoides from the Serra de Capanema. Overall, the vegeta-
tion of Serra do Caraça is largely intact and aords a good measure of protection for
the populations of M. pithyoides. is species is considered Critically Endangered by
the Brazilian Government (Brasília 2014). Based on criterion B of the IUCN (2019),
we concur with this assessment: (CR): B1ab(iii).
Recognition and anities. Microlicia pithyoides can be recognised by its narrow
leaves that are 0.5–1.4 mm wide, 1-nerved from the base, inorescences reduced to
solitary owers, magenta petals and stamens with bicoloured anthers. In morphology, it
is most like M. rosmarinoides, the only congener with leaves that are 1-nerved from the
base. Microlicia rosmarinoides also shares with M. pithyoides the solitary owers with sub-
ulate calyx lobes and the overall shape of its stamens. Microlicia pithyoides diers from
M. rosmarinoides by the leaves with the mid-vein thickened (vs. not thickened), magenta
petals (vs. yellow) and stamens with anthers vinaceous and yellow (vs. all anthers yellow
to orange). Both species occur in central Minas Gerais State, but their distributions do
not overlap; M. rosmarinoides has never been collected on the Serra do Caraça.
Microlicia calycina is also morphologically similar. It shares with M. pithyoides
the narrow leaves, solitary owers (sometimes simple dichasia only in M. calycina),
magenta petals and stamens with bicoloured anthers. Microlicia pithyoides may be dif-
ferentiated by its leaf blades (oblong to lanceolate) 0.5–1.4 mm wide (vs. 2–9 mm
wide), 1-nerved from the base (vs. 3-nerved) with tertiaries not evident (vs. evident)
and shorter calyx lobes 2.5–3.0 mm long (vs. 3.5–4.2 mm long). Both M. calycina and
M. pithyoides occur sympatrically on the Serra do Caraça, but only M. pithyoides occurs
on the highest peak in that mountain range. In fact, M. pithyoides apparently prefers
slightly higher elevations since it has only been collected between 1827 and 2072 m
(vs. 1692–1920 m for M. calycina).
Notes. Recent collections of M. pithyoides (Oliveira 382, Pacico 295) have leaves
with blades that are 8–12 mm long. ese measurements bridge those given in the proto-
logues of varieties pithyoides (8–10 mm long) and major (12–20 mm long). We consider
these size dierences to represent a continuum and here relegate var. major to synonymy.
9. Microlicia rosmarinoides (Mart. & Schrank ex DC.) Versiane & R.Romero,
Bot. J. Linn. Soc. 197: 54. 2021.
Fig. 28
Trembleya rosmarinoides Mart. & Schrank ex DC., Prodr. 3: 125. 1828. basionym.
Type: Brazil. ”Habitat in summo Monte de V. Rica et in Itacolumi 5000 ped. alt.
Provinciae Min. Gen.” [Minas Gerais, Ouro Preto], C.F.P. Martius 808 (lectotype,
designated here: M [M0165886]!; isolectotype: G [G00310213-online image]!).
Description. Erect, densely branched shrubs 0.3–0.6 m tall. Branchlets quadrangular,
glandular-punctate, light green (when fresh). Internodes 0.1–0.4 cm long, angles with
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 75
Figure 28. Microlicia rosmarinoides A habit B detail of nodes and internodes C leaf abaxial surface
D bracteole abaxial surface E oral bud F ower in lateral view G petal adaxial surface H antesepalous
stamen I antepetalous stamen J capsule enveloped by the hypanthium K capsule in cross section L seed in
lateral view. Drawn from Occhioni et al. s.n. (US [US001900109]).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
76
narrow wings 0.2–0.3 mm wide. Petioles 0.5–1.2 mm long. Leaf blades 4–10mm long,
1.0–1.9 mm wide, chartaceous (when dry), narrowly-lanceolate to narrowly-elliptic,
both surfaces green (when fresh), adaxial surface blackened and abaxial surface pale
green to pale brown (when dry), discoloured (when dry), base cuneate, apex rounded
to acute, margin at, entire and glandular-punctate, 1-nerved from the base, tertiaries
little evident on the abaxial surface, adaxial surface glandular-punctate, abaxial surface
glandular-punctate. Inorescences reduced to solitary owers on apical branches. Bracts
absent. Bracteoles (at anthesis) with petioles 0.5–1.0 mm long, blades 2.5–3.3mm
long, 0.5–0.9 mm wide, lanceolate, base cuneate, apex acute to obtuse, margin entire,
1-nerved, indumentum like that of the principal leaves. Flowers 5-merous, pedicels (at
anthesis) 0.4–0.6 mm long. Hypanthia (at anthesis) 2.2–3.5mm long, 1.5–2.0 mm
wide at the torus, campanulate to urceolate, light green (when fresh), externally glandu-
lar-punctate. Calyx tubes 0.3–0.5 mm long. Calyx lobes (at anthesis) 2.2–2.8 mm long,
0.5–0.8 mm wide at the base, subulate, apex acute, margin entire, (when fresh) light
green, externally like the hypanthia. Petals 5.0–5.3 mm long, 2.4–3.0 mm wide, yellow,
obovate, apex acute or rounded, margin entire and glabrous, both surfaces glabrous.
Stamens 10, strongly dimorphic. Larger (antesepalous) stamens 5, laments 2.2–3.0 mm
long, yellow, pedoconnectives 3.2–3.8 mm long, yellow, appendages 0.8–1.0 mm long,
yellow, apex emarginate to bilobate, thecae (excluding rostra) 1.4–1.7 mm long, yel-
low to orange, oblong, rostra 0.2–0.3 mm long, the circular pores ca. 0.2 mm wide.
Smaller (antepetalous) stamens 5, laments 1.5–2.0 mm long, yellow, pedoconnec-
tives 0.8–1.0 mm long, yellow, inconspicuous appendages ca. 0.1 mm long, yellow,
apex emarginate, thecae (excluding rostra) 1.4–1.6 mm long, yellow to orange, oblong,
rostra 0.2–0.3 mm long, the circular pores ca. 0.2 mm wide. Ovary 0.9–1.1 mm long,
0.7–0.9 mm wide, ovoid, 5-locular. Style ca. 3 mm long, yellow. Capsules (at maturity)
2.5–3.5 mm long, 2.0–3.0 mm wide, globose, initially enveloped by the hypanthium,
torus constricted at the apex, fruiting calyx tubes 0.3–0.6mm long, fruiting calyx lobes
2.5–3.2 mm long, not thickened. Seeds 0.5–0.9 mm long, reniform.
Distribution, habitat, and elevation range. Restricted to central Minas Gerais
State (Fig. 25B), where it is known only from Serra do Gandarela, Serra de Capanema,
Serra de Ouro Preto, Serra de Itabirito, Belo Vale and Barão de Cocais. Microlicia
rosmarinoides occurs in campo rupestre and canga, on ferruginous or quartzitic soils,
exposed to full sun, at elevations between 1609 and 1807 m.
Conservation. Microlicia rosmarinoides is a little-collected species, known from
about 15 specimens. e EOO is 298.172 km2 and the AOO is 28 km2. Populations
of M. rosmarinoides are protected in the Parque Nacional da Serra do Gandarela and
probably in Parque Estadual do Itacolomi. Following IUCN (2019) criteria, we rec-
ommend an assessment of Endangered (EN): B1ab(iii) for this species.
Recognition and anities. Microlicia rosmarinoides can be readily recognised
amongst congeners by its narrow leaves (1–1.9 mm wide) and owers with yellow
petals. e narrow leaves of M. rosmarinoides are more similar to those of the closely
related M. pithyoides, while the yellow petals are shared only with the distantly related
M. aviora. For comparisons, see notes under M. aviora and M. pithyoides.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 77
Specimens examined. Brazil. Minas Gerais: Barão de Cocais, Mina do Baú, Souza
et al. 2584 (BHCB); Belo Vale, Occhioni et al. s.n. (RFA, US [US001900109]); Itabiri-
to, Serra de Capanema, Carmo 192 (BHCB), Carmo 377 (BHCB); Ouro Preto mu-
nicipality, Serra do Itacolomi, Martius 808 (lectotype: M [M0165886]; isolectotype:
G [G00310213]), Pedrosa 110 (HUFU, OUPR), Schwacke 9184 (BHCB, RB, US,
W); Rio Acima Municipality, Serra do Gandarela, Carmo 2262 (BHCB), Versiane &
Castello 677 (HUFU, UEC), Vidal s.n. (BHCB [191426]); Santa Bárbara Municipal-
ity, Serra do Gandarela, Vidal s.n. (BHCB [181346]); Unknown municipality in Minas
Gerais State, Glaziou 14747 (IAN, K, RB, US), Glaziou 19242 (K), Ule 2528 (US).
10. Microlicia trembleyiformis Naudin, Ann. Sci. Nat., Bot. Sér. 3, 3: 172. 1845.
Fig. 29
Type. B. “Minas Geraes, in campis circa urbem Villa Ricca frequens” [Minas
Gerais, Ouro Preto], 1816–1821, [catal. B1, n° 160] A. Saint-Hilaire s.n. (lectotype,
rst-step designated by Martin and Cremers (2007), second-step designated here: P
[P002297746]!; isolectotype: F [F0360366]!; image of lectotype is available at http://
coldb.mnhn.fr/catalognumber/mnhn/p/p02297746).
Description. Erect shrubs 0.5–1.0 m tall. Branchlets quadrangular, glandular-
punctate and sparsely covered with eglandular trichomes 0.1–0.5, light green (when
fresh). Internodes 0.2–1.1 cm long, angles with narrow wings ca. 0.2 mm wide. Petioles
0.3–1.4 mm long. Leaf blades 4–25 mm long, 1.5–12 mm wide, papyraceous (when
dry), ovate or elliptic, both surfaces green (when fresh), adaxial surface blackened and
abaxial surface pale green to pale brown (when dry), discoloured (when dry), base
rounded or attenuate, apex acute or obtuse, margin at, slighly serrulate and ciliate
with eglandular trichomes 0.1–0.4 mm long, 5-nerved from the base, one pair of acro-
dromous veins and one pair of tenuous veins close to the margin, tertiaries evident on
the abaxial surface, nearly perpendicular to the mid-vein, little reticulate and branching
apically, adaxial surface glandular-punctate, abaxial surface densely glandular-punctate
and sparsely covered with eglandular trichomes 0.1–0.5 mm long around the veins.
Inorescences reduced to solitary owers on the aplical region of the branches. Bracts
absent. Bracteoles (at anthesis) with petioles 0.3–0.5 mm long, blades 2.4–6.1 mm
long, 1.0–3.2 mm wide, ovate or elliptic, base cuneate, apex acute to obtuse, margin
slightly serrulate and ciliate with eglandular trichomes 0.1–0.4 mm long, 3-nerved,
indumentum like that of the principal leaves. Flowers 5-merous, pedicels (at anthesis)
0.7–1.9 mm long. Hypanthia (at anthesis) 2.2–2.6 mm long, 1.8–2.2 mm wide at the
torus, campanulate to urceolate, light green (when fresh), externally glandular-punc-
tate and sparsely covered with eglandular trichomes 0.1–0.5. Calyx tubes 0.2–0.4 mm
long. Calyx lobes (at anthesis) 1.5–2.1 mm long, 0.5–0.7 mm wide at the base, nar-
rowly triangular, apex acute, margin entire, (when fresh) light green, externally like the
hypanthia. Petals 6.1–7.9 mm long, 3.0–3.5 mm wide, magenta, obovate, apex acute,
margin entire and glabrous, both surfaces glabrous. Stamens 10, strongly dimorphic.
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
78
Larger (antesepalous) stamens 5, laments 1.5–3.0 mm long, pink, pedoconnectives
1.9–2.9 mm long, pink, appendages 1.0–1.5 mm long, yellow, apex bilobate, thecae
(excluding rostra) 1.0–1.5 mm long, purple, oblong, rostra 0.2–0.3 mm long, the cir-
cular pores ca. 0.2 mm wide. Smaller (antepetalous) stamens 5, laments 1.5–2.0 mm
Figure 29. Microlicia trembleyiformis A habit B detail of nodes and internodes C leaf abaxial surface
D leaf adaxial surface E bracteole abaxial surface F oral bud G ower in lateral view H petal adaxial sur-
face I antesepalous stamen J antepetalous stamen K gynoecium L capsule enveloped by the hypanthium.
Drawn from Duarte 2767 (US).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 79
long, pink, pedoconnectives 0.8–1.2 mm long, yellow, inconspicuous appendages ca.
0.1 mm long, yellow, apex emarginate, thecae (excluding rostra) 0.7–1.1 mm long,
yellow, oblong, rostra 0.2–0.3 mm long, the circular pores ca. 0.2 mm wide. Ovary
1.8–2.0 mm long, 1.9–2.1 mm wide, globose, 3–5-locular. Style 3.7–4.2 mm long,
pink. Capsules (at maturity) 2.5–3.4 mm long, 2.4–3.0 mm wide, globose, initially
enveloped by the hypanthium, torus constricted at the apex, fruiting calyx tubes 0.3–
0.4 mm long, fruiting calyx lobes 2.4–2.9 mm long, not thickened. Seeds not seen.
Distribution, habitat and elevation range. Microlicia trembleyiformis is known from
quartzitic campo rupestre, Cerrado and veredas (palm swamps) at Ouro Preto, Serra da
Canastra, Uberlândia and Patrocínio (in Minas Gerais State) and campos de altitude in
Serra Negra (Itatiaia), Rio de Janeiro State (Fig. 19B). On Serra da Canastra, it was collect-
ed on sandy soils near streams, exposed to full sun, at elevations between 786 and 1300 m.
Collections from Ouro Preto and Serra Negra lack additional habitat information.
Conservation. is species is known from about 10 specimens. It is a little-col-
lected species that, however, has a comparatively wide distributional range. e EOO
is 92,214.048 km2, a value that would indicate a Least Concern conservation status
if criterion B of IUCN (2019) was applied. However, the AOO of 24 km2 matches a
status of Endangered in accordance with the same criterion. Both EOO and AOO re-
quirements have to be fullled for the correct use of criterion B. us, we recommend
a status of Least Concern (LC) for M. trembleyiformis. We suspect that this species oc-
curs in low population densities. Some populations of M. trembleyiformis are protected
in the Parque Nacional da Serra da Canastra.
Recognition and anities. Microlicia trembleyiformis can be recognised by its el-
liptic to ovate leaves that are 5-nerved from the base and solitary owers with narrow-
ly-triangular calyx lobes 1.5–2.1 mm long. In morphology, it is closest to M. parviora
and more distantly to M. altoparaisensis (see notes under these species for compari-
sons). Microlicia pentagona is the only congener that shares with M. trembleyiformis
both the leaves 5-nerved from the base and solitary owers. Microlicia trembleyiformis
diers in having leaves that are papyraceous when dry (vs. chartaceous to coriaceous in
M. pentagona), the margin is ciliate with eglandular trichomes 0.1–0.4 mm long (vs.
glabrous to minutely granulose) and the shorter calyx lobes are 1.5–2.1 mm long (vs.
6.2–8.5 mm long) and tenuous in fruit (vs. thickened). Amongst the compared spe-
cies, only M. parviora occurs in Ouro Preto, Serra da Canastra and Serra Negra, where
sympatry with M. trembleyiformis is possible.
Notes. According to Naudin (1845), this species was frequent in Ouro Preto Mu-
nicipality, although we are not aware of any recent collections from that region. Naudin
(1845) justied the placement of M. trembleyiformis in Microlicia because the type had
3-locular ovaries. Recent collections of M. trembleyiformis have 3–5-locular ovaries (e.g.
Porto 2834). In the Parque Nacional da Serra da Canastra, M. trembleyiformis is known
from only a few individuals that consistently have 3-locular ovaries (Romero 2000).
Specimens examined. Brazil. Minas Gerais: Capitólio Municipality, estrada
para Cachoeira Fecho da Serra, Romero et al. 7550 (HUFU, US); Ouro Preto Munici-
pality, [catal. B1, n° 160] Saint-Hilaire s.n. (lectotype: P [P002297746]!; isolectotype:
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
80
F [F0360366]); São Roque de Minas Municipality, Serra da Canastra, Nakajima et
al. 1476 (HUFU, US), Romero & Nakajima 3593 (HUFU, K, UEC), Romero 4157
(BHCB, F, HUFU), Santos 411 (HUFU, K); Serra do Salitre Municipality, Serra de
Catiara, Duarte 2767 (BHCB, US); Uberlândia Municipality, Clube Caça e Pesca
Itororó, Romero et al. 8689 (HUFU), Romero et. al. 8694 (HUFU); unknown mu-
nicipality in Minas Gerais State, Glaziou 19221 (K, P). Rio de Janeiro: Itatiaia Mu-
nicipality, Serra Negra, Porto 2834 (NY, RB, US).
11. Microlicia tridentata (Naudin) Versiane & R.Romero, Bot. J. Linn. Soc. 197:
55. 2021.
Fig. 30
Trembleya tridentata Naudin, Ann. Sci. Nat., Bot. Sér. 3, 2: 154: 1844. basionym. Type:
Brazil. “In montibus Serra de San Jose, provinciae Minas Geraes” [Minas Gerais,
Serra de São José], 1816–1821, [catal. B2, n° 2397] A. Saint-Hilaire s.n. (lectotype,
rst-step designated by Martin and Cremers (2007), second-step designated here: P
[P00723392]!; isolectotypes: P [P00723391]!, P [P00723506]!; image of lectotype
is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p00723392).
Description. Erect shrubs or treelets 1.0–1.8 m tall. Branchlets quadrangular, appear-
ing glabrous, vernicose and minutely granulose, light green (when fresh). Internodes
0.3–1.5 cm long, angles with narrow wings 0.2–0.4 mm wide. Petioles 1.0–4.9 mm long.
Leaf blades 20–49 mm long, 12–24 mm wide, coriaceous (when dry), elliptic, ovate, or
rarely narrowly elliptic, both surfaces green (when fresh), adaxial surface blackened and
abaxial surface pale green to pale brown (when dry), discoloured (when dry), base cune-
ate or attenuate, apex rounded to emarginate, margin at, entire along the basal half, ser-
rulate on the upper half and glandular-punctate, 5-nerved from the base, one pair of ac-
rodromous veins and one pair of tenuous veins close to the margin, tertiaries evident on
the abaxial surface, nearly perpendicular to the mid-vein, reticulate, randomly branching
and surrounding stout depressions on the abaxial leaf surface, adaxial surface glandular-
punctate, abaxial surface glandular-punctate. Inorescences simple dichasia or reduced to
solitary owers, not congested. Bracts (including petioles) 1.0–1.6 cm long, 0.4–0.8 cm
wide, 3- to inconspicuously 5-nerved, elliptic, indumentum like that of the principal
leaves. Bracteoles (at anthesis) with petioles 2.5–5.0 mm long, blades 8.1–11.0 mm long,
4.0–5.5 mm wide, elliptic, base attenuate, apex rounded to obtuse, margin sparsely serru-
late, 3-nerved, indumentum like that of the principal leaves. Flowers 5-merous, pedicels
(at anthesis) 1.3–1.6 mm long. Hypanthia (at anthesis) 3.5–3.9 mm long, 2.3–2.7 mm
wide at the torus, campanulate to urceolate, light green (when fresh), externally appear-
ing glabrous, vernicose and minutely granulose. Calyx tubes 0.3–0.4 mm long. Calyx
lobes (at anthesis) 4.1–4.9mm long, 0.8–1.1 mm wide at the base, subulate, apex acute,
margin entire, (when fresh) light green or reddish, externally like the hypanthia. Petals
11.5–13.0mm long, 6.0–7.0 mm wide, magenta or rarely white, obovate, apex acute
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 81
Figure 30. Microlicia tridentata A habit B leaf abaxial surface C bracteole abaxial surface D oral bud
E ower in lateral view F owering hypanthium G petal adaxial surface H antesepalous (behind) and an-
tepetalous (in front) stamens I gynoecium J capsule enveloped by the hypanthium. Drawn from Hensold
et al. CFCR2880 (UEC) and Souza et al. 8015 (UEC).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
82
or rounded, margin entire and glabrous, both surfaces glabrous. Stamens 10, strongly
dimorphic. Larger (antesepalous) stamens 5, laments 4.9–5.6 mm long, pink or rarely
white, pedoconnectives 6.0–6.5 mm long, pink or rarely white with the apical region
ushed with pink, appendages 1.7–1.9 mm long, yellow, apex emarginate or truncate,
thecae (excluding rostra) 1.7–1.9 mm long, purple, oblong, rostra 0.3–0.5 mm long, the
circular pores ca. 0.2 mm wide. Smaller (antepetalous) stamens 5, laments 4.0–4.4mm
long, pink or rarely white, pedoconnectives 1.5–1.9 mm long, pink or rarely white with
the apical region ushed with pink, inconspicuous appendages ca. 0.2 mm long, yel-
low, apex emarginate or truncate, thecae (excluding rostra) 1.4–1.7 mm long, yellow,
oblong, rostra 0.3–0.5 mm long, the circular pores ca. 0.2 mm wide. Ovary 2.2–2.4mm
long, 2.1–2.3 mm wide, globose, 5-locular. Style 6.1–6.5 mm long, pink or rarely white.
Capsules (at maturity) 3.0–4.0 mm long, 2.7–3.6 mm wide, globose, initially enveloped
by the hypanthium, torus constricted at the apex, fruiting calyx tubes 0.4–0.5mm long,
fruiting calyx lobes 4.5–5.4 mm long, not thickened. Seeds not seen.
Distribution, habitat and elevation range. Microlicia tridentata is endemic to
Minas Gerais State (Fig. 25A) at Serra de Ouro Branco, Serra de Ouro Preto, Serra de
São José, Serra do Gandarela, Serra do Cipó, Serra do Lenheiro, Serra do Caraça and
Barão de Cocais. It grows on quartzitic or ferruginous campo rupestre, exposed to full
sun, at elevations between 1110 and 1614 m.
Conservation. Microlicia tridentata is known from about 40 collections. e
EOO is 4,613.081 km2 and the AOO is 24 km2. Following IUCN (2019) recommen-
dations, we recommend Endangered (EN): B1ab(iii) status for this species. Popula-
tions of M. tridentata are protected in Parque Estadual do Itacolomi, Parque Estadual
Serra do Ouro Branco and RPPN Serra do Caraça (Natural Heritage Private Reserve).
Recognition and anities. Microlicia tridentata can be recognised by its leaves
that are serrulate along the upper half, 5-nerved from the base, abaxially conspicuously
glandular-punctate, with tertiary veins reticulate and randomly branching and owers
with subulate calyx lobes 4.1–4.9 mm long. It shares with M. pentagona the shrubby
to treelet habit and the serrulate leaves along the upper half that are 5-nerved from the
base. Microlicia tridentata is readily dierentiated by the abaxial surfaces of the leaves
that are more conspicuously glandular-punctate, with tertiares reticulate and randomly
branching (vs. parallel or little reticulate and branching apically) and shorter calyx
lobes 4.1–4.9 mm long (vs. 6.2–8.5 mm long) that are tenuous in fruit (vs. thickened).
Another close relative is M. parviora, from which M. tridentata diers by the leaves
that are not pruinose (vs. usually pruinose), inorescences composed of simple dicha-
sia or reduced to solitary owers (vs. compound or simple dichasia), bracteoles longer
8.1–11.0 mm long (vs. 2.2–3.8 mm long) and longer calyx lobes 4.1–4.9 mm long [vs.
0.7–2.5(–3.0)] that are subulate (vs. triangular). e distributions of M. parviora and
M. pentagona overlap with M. tridentata on several mountain ranges in Minas Gerais
(e.g. Serra de Ouro Branco, Serra de Ouro Preto, Serra do Cipó).
Notes. Major variation in M. tridentata involves inorescence development (simple
dichasia or solitary owers) and the petals typically magenta (Fig. 8H), rarely white
(Fig. 9H), which may be rounded or acute at the apex.
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 83
Specimens examined. Brazil. Minas Gerais: Barão de Cocais Municipality,
Irwin et al. 29029 (IAN, NY, US), Irwin et al. 29079 (K, MO, NY, P, RB); Catas
Altas Municipality, Serra do Caraça, Castro et al. 281 (HUFU); Ouro Branco Mu-
nicipality, Serra de Ouro Branco, Pacico & Bressan 290 (CAS, HUEM), Souza et
al. 8015 (ESA, SPF, UEC); Ouro Preto Municipality, Serra do Itacolomi, Badini
& Ferreira 9774 (HUFU), Ferreira & Helena 7844 (ESA, HUFU), Giulietti et al.
CFCR13780 (K, MO, SPF), Longhi-Wagner et al. CFCR9184 (SPF, UEC, US),
Magalhães 1160 (IAN, US), Michelangeli et al. 1586 (NY, RB, UPCB), Michel-
angeli et al. 1595 (NY, UPCB), Pedrosa 120 (OUPR), Rolim et al. 326 (HUFU,
VIC), Rolim et al. 327 (UPCB, VIC), Rolim et al. 328 (HUFU, VIC), Rolim et al.
394 (HUFU, NY, RB, VIC), Roschel et al. s.n. (OUPR [1506], RB [RB01301048]),
Souza et al. 8047 (ESA, SORO, SPF), Souza et al. 8061 (ESA, RB, SPF, UPCB); Rio
Acima Municipality, Serra do Gandarela, Pacico & Bressan 303 (CAS, HUEM),
Versiane & Devides 682 (UEC); Santa Bárbara Municipality, Serra do Caraça, Or-
dones et al. 84 (BHZB, HUFU), Pereira & Pabst 2619 (RB, US); Santana do Riacho
Municipality, Serra do Cipó, Duarte 8157 (G, LE, RB, US); São João del-Rei Mu-
nicipality, Schwacke 10135 (BHCB, RB, W), Glaziou 17513 (P, R, US), Serra do
Lenheiro, Glaziou 16781 (K, NY, P, US); Tiradentes Municipality, Serra de São José,
Alves 595 (US), Alves 7359 (R), Rutter 128 (R), Rutter 150 (R), Rutter 161 (R),
Rutter 186 (R), Rutter 191 (R); Unknown municipality in Minas Gerais State, Claus-
sen 1631 (P), Glaziou 14743 (K, P, US), Glaziou 17573 (K), Mendonça 557 (US),
Riedel s.n. (K [K009597799, W-18800001419, W-18890019740]), Saint-Hilaire
s.n. [catal. B2, n° 2397] (lectotype: P [P00723392]; isolectotypes: P [P00723391], P
[P00723506]), Sellow s.n. (US [US00292635]); Serra da Conceição (Serra B. Vista),
Hensold et al. CFCR2880 (SPF, US).
A nomenclatural review on the remaining taxa previously treated in Trembleya
1. Trembleya acuminata R.B.Pacico & Fidanza, Phytotaxa 238(2): 164. 2015.
Type: Brazil. Minas Gerais, Joaquim Felício, Serra do Cabral, estrada Joaquim Felício–
Várzea da Palma, ca. 24 km de Joaquim Felício, 10 July 2001, V.C. Souza et al. 25654
(holotype: MBM!; isotypes: BHCB!, ESA!, SPF!).
Replaced with: Microlicia acuminifolia Versiane & R.Romero.
2. Trembleya agrestis Mart. & Schrank ex DC., Prodr. 3: 126. 1828. Type: Brazil.
“In prov. Minas Geraes” [Minas Gerais], C.F.P. Martius s.n. (holotype: M [M0165682]!,
isotype [fragment]: BR [BR0000005521203]!).
Basionym of: Microlicia agrestis (Mart. & Schrank ex DC.) Cogn.
3. Trembleya botaensis R.B.Pacico & Fidanza, Phytotaxa 238(2): 167. 2015.
Type: Brazil. Minas Gerais, Guaraciama, Cadeia do Espinhaço, Serra do Bota, 19
March 2005, E. Guarçoni 903 & M. A. Sartori (holotype: MBM!).
Synonym of: Microlicia curralensis Brade.
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
84
4. Trembleya elegans (Cogn.) Almeda & A.B.Martins, Novon 11(1): 6. 2001. Type:
Based on Lavoisiera elegans Cogn.
Replaced with: Microlicia speciosa Versiane & R.Romero.
5. Trembleya inversa Fidanza, A.B.Martins & Almeda, Brittonia 65(3): 281. 2013.
Type: Brazil. Minas Gerais, Joaquim Felício, Serra do Cabral, Armazém da Laje, 9 Sep-
tember 2003, K. Fidanza & R. Belinello 12 (holotype: UEC!; isotype: CAS!).
Basionym of: Microlicia inversa (Fidanza, A.B.Martins & Almeda) Versiane &
R.Romero.
6. Trembleya neopyrenaica Naudin, Ann. Sci. Nat., Bot. Sér. 3, 2: 154. 1844.
Type: Brazil. “In montibus Pyreneos prov. Goyaz” [Goiás, Serra dos Pireneus], 1816,
A. Saint-Hilaire C1-694 (lectotype, rst-step designated by Martin and Cremers
(2007), second-step designated here: P [P00723416]!; isolectotypes: P [P00723417]!,
P [P00723418]!, F [fragment] [F0064039F-online image]!; image of lectotype is avail-
able at http://coldb.mnhn.fr/catalognumber/mnhn/p/p00723416).
Basionym of: Microlicia neopyrenaica (Naudin) Versiane & R.Romero.
7. Trembleya phlogiformis Mart. & Schrank ex DC., Prodr. 3: 126. 1828. Type:
Brazil. “In Brasiliae campis prov. S.-Pauli” [São Paulo State], Martius s.n. (holotype: M
[M0165885]!, probable isotype: G [G00310212]!).
Basionym of: Microlicia phlogiformis (Mart. & Schrank ex DC.) Versiane & R.Romero.
Melastoma pumilum Vell., Fl. Flumin. 179. 1829. Type: Brazil. “Habitat campis apricis
mediterraneis prope Predium Boavista inter gramina” (lectotype, designated here:
Original illustration published in Vellozo, Fl. Flumin. Icones 3: t. 151. 1831; the
original parchment is in the manuscript section of the Biblioteca Nacional, Rio de
Janeiro; a copy of the illustration is deposited at G [G00368052]!).
Trembleya phlogiformis var. glabra Cham., Linnaea 9: 429. 1835. syn. nov. Type: Brazil.
“In Brasilia australi loco haud indicato”, Sellow 2387 (lectotype, designated here: S
[S09-12942-online image]!; probable isolectotype: K [K000530643]!).
Trembleya stachyoides Naudin, Ann. Sci. Nat., Bot. Sér. 3, 2: 154. 1844. syn. nov.
Type: Brazil. “In Brasilia australi, praecipue circa Tocoropa”, Laruotte s.n. (lecto-
type, rst-step designated by Martin and Cremers (2007), second-step designated
here: P [P00723388]!; isolectotype: P [P00723389]!; image of lectotype is avail-
able at http://coldb.mnhn.fr/catalognumber/mnhn/p/p00723388).
Trembleya phlogiformis var. cuneifolia Cogn. in Martius et al., Fl. Bras. 14(3): 132. 1883.
syn. nov. Type: Brazil. “In prov. Minas Geraës ad Congonhas do Campo” [Minas
Gerais], Stephan s.n. (lectotype, designated here: BR [BR0000005224203]!).
Trembleya phlogiformis var. genuina Cogn. in Martius et al., Fl. Bras. 14(3): 132. 1883.
syn. nov. Type: Based on Trembleya phlogiformis Mart. & Schrank ex DC.
Trembleya phlogiformis var. latifolia Cogn. in Martius et al., Fl. Bras. 14(3): 132. 1883.
syn. nov. Type: Brazil. “In Brasilia loco haud indicato”, Raben 781 (lectotype, des-
ignated here: BR [BR0000005520855]!; isolectotype: BR [BR0000005520190]!).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 85
Trembleya phlogiformis var. microlicioides Cogn. in Martius et al., Fl. Bras. 14(4): 594.
1888. syn. nov. Type: Brazil. In prov. Rio de Janeiro, Glaziou 16046 (lectotype,
designated here: R [R000009186]!; isolectotypes: BR [BR0000005223596]!, BR
[BR0000005520503]!, BR [BR0000005520176]!, C [C10015111-online im-
age]!, G [G00368051]!, P [P00723400]!; image of isolectotype at P is available at
http://coldb.mnhn.fr/catalognumber/mnhn/p/p00723400).
Trembleya phlogiformis var. parvifolia Cogn. in Martius et al., Fl. Bras. 14(3): 132. 1883.
syn. nov. Type: Brazil. “In paludibus prope Lorena prov. S. Paulo”, Riedel 1418
(lectotype, designated here: P [P05317991]!; isolectotypes: K [K000530649]!, P
[P05317995]!; image of lectotype is available at http://coldb.mnhn.fr/catalog-
number/mnhn/p/p05317991).
Trembleya phlogiformis var. quinquenervia Cogn. in Martius et al., Fl. Bras. 14(3):
132. 1883. syn. nov. Type: Brazil. “In prov. Goyaz” [Goiás], Gardner 4147 (lec-
totype, designated here: BM [BM000516951]!; isolectotypes P [P05317981]!, P
[P05317982]!; image of lectotype is available at http://plants.jstor.org and http://
data.nhm.ac.uk/object/255f17a9-b371-4dd3-8eb0-9f0eafbd2e76).
Trembleya phlogiformis var. ramosissima Cogn. in Martius et al., Fl. Bras. 14(3): 132. 1883.
syn. nov. Type: Brazil. “In prov. Minas Geraës”, Regnell I. 152 part (lectotype, desig-
nated here: R [R000166805]!; isolectotypes: P [P05318036]!, S [S09-12952-online
image]!; images of the lectotype and an isolectotype are available at http://plants.
jstor.org and http://coldb.mnhn.fr/catalognumber/mnhn/p/p05318036).
Trembleya phlogiformis var. stachyoides (Naudin) Cogn. in Martius et al., Fl. Bras. 14(3):
132. 1883. syn. nov. Type: Based on Trembleya stachyoides Naudin.
Trembleya phlogiformis var. villosa Cogn. in Martius et al., Fl. Bras. 14(3): 132. 1883.
syn. nov. Type: Brazil. “In campis siccis ad Registo Velho”, Pohl & Schüch 230
(lectotype, designated here: BR [BR0000005223541]!).
Trembleya selloana Cogn. in Martius et al., Fl. Bras. 14(3): 133. 1883. syn. nov. Type:
Based on Trembleya phlogiformis var. glabra Cham.
8. Trembleya pityrophylla (Mart. ex DC.) Triana, Trans. Linn. Soc. London 28(1):
24. 1872. Type: Based on Osbeckia pityrophylla Mart. ex DC.
Synonym of: Cambessedesia pityrophylla (Mart. ex DC.) A.B.Martins.
9. Trembleya pradosiana Netto, Ann. Sci. Nat., Bot. Sér. 5, 3: 378. 1865. Type:
Brazil. “Habitat in campis ad umen Rio das Velhas, prope vivum Trahiras, in parte
centrali provinciae Minas Geraes” [Minas Gerais], May 1862, Netto s.n. (holotype: P
[P00723509]!; image of holotype is available at http://coldb.mnhn.fr/catalognumber/
mnhn/p/p00723509).
Basionym of: Microlicia pradosiana (Netto) Versiane & R.Romero.
Trembleya rubra Fidanza, A.B.Martins & Almeda, Brittonia 65: 286. 2013. syn.
nov. Type: Brazil. Minas Gerais, Joaquim Felício, Serra do Cabral, ca. 5 km S do
Armazém da Laje, 4 December 2003, K. Fidanza & R. Belinello 112 (holotype:
UEC!; isotype: CAS!).
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
86
10. Trembleya purpurascens Fidanza, A.B.Martins & Almeda, Brittonia 65(3):
284. 2013. Type: Brazil. Minas Gerais, Joaquim Felício, Serra do Cabral, estrada
Joaquim Felício-Armazém, 4 May 2003, K. Fidanza & R. Belinello 56 (holotype:
UEC!; isotype: CAS!).
Synonym of: Microlicia curralensis Brade.
11. Trembleya serrulata Fidanza, A.B.Martins & Almeda, Brittonia 65(3): 288.
2013. Type: Brazil. Minas Gerais, Joaquim Felício, Serra do Cabral, Pedreira, 7 De-
cember 2003, K. Fidanza 108 (holotype: UEC!; isotype: CAS!).
Replaced with: Microlicia serratifolia Versiane & R.Romero.
12. Trembleya thomazii R.B.Pacico & Fidanza, Phytotaxa 238(2): 171. 2015.
Type: Brazil. Minas Gerais, Guaraciama, Cadeia do Espinhaço, Serra do Bota, 20
March 2005, E. Guarçoni 929 & M. A. Sartori (holotype: MBM!)
Basionym of: Microlicia thomazii (R.B.Pacico & Fidanza) Versiane & R.Romero.
13. Trembleya warmingii Cogn. in Martius et al., Fl. Bras. 14(3): 133. 1883. Type:
Brazil. “Ad Lagoa Santa” [Minas Gerais, Lagoa Santa], 9 February 1866, E. Warming
2306 (holotype: C!).
Basionym of: Poteranthera warmingii (Cogn.) Almeda & R.B.Pacico.
Excluded and/or dubious names
1. Trembleya capitata Cogn. [nom. inval.]
2. Trembleya canescens Schnizl. [probably = Pleroma D.Don]
3. Trembleya debilis Glaz. [nom. inval.]
4. Trembleya santae-luziae Glaz. [nom. inval.]
5. Trembleya rhynanthera Gri. [probably = Melastoma L.]
6. Trembleya rosea Sessé & Moc. [probably = Coreopsis grandiora Hogg ex Sweet
(Asteraceae)]
Acknowledgements
We thank: Boni Cruz and Heritiana Ranarivelo for their assistance with lab work in
the Center for Comparative Genomics at the California Academy of Sciences; Sandra
Knapp, Laurent Gautier, Elizabeth Woodgyer and Nicholas Hind for help in locat-
ing the type of Meriania parviora; BHCB, CAS, HUEM, UEC, MBM, R, RB, SP
and SPF, for herbarium-related logistical support; Mateus Reck for his help with the
BI analysis; Alan Chou and Klei Souza for the line drawings; Zhi-Qiang Zhang for
granting permission to reproduce the illustration of Microlicia altoparaisensis; V.E.
Bressan, R. Gabani and P.V. Simões for their assistance during eldwork activities;
André V. Scatigna, Fabian A. Michelangeli, Fernando A.O. Silveira, Luciano Pedrosa,
Orlando Grae, Rodrigo Penati, and Vania E. Bressan for allowing us to use their
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 87
photos of living specimens; Renato Goldenberg, an anonymous reviewer and the han-
dling editor (Marcelo Reginato) for their comments and suggestions on an earlier
version of this monograph. RP thanks the Coordenação de Aperfeiçoamento de Pes-
soal de Nível Superior (CAPES) for a Ph.D. degree grant and the M. Stanley Run-
del Charitable Trust (U.S.A.) and the Lakeside Foundation (U.S.A.) for grants that
supported his training and research visits to the California Academy of Sciences in
2017 and 2019–2020. is research was supported in part by U.S. National Science
Foundation grant DEB-1146409.
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98
Appendix 1
List of species
1. Microlicia altoparaisensis (R.B.Pacico, Almeda & Fidanza) Versiane & R.Romero
2. Microlicia calycina (Cham.) Versiane & R.Romero
3. Microlicia chamissoana (Naudin) Versiane & R.Romero
4. Microlicia aviora Versiane & R.Romero
5. Microlicia laniora (D.Don) Baill.
6. Microlicia parviora (D.Don) Versiane & R.Romero
7. Microlicia pentagona (Naudin) Versiane & R.Romero
8. Microlicia pithyoides (Cham.) Versiane & R.Romero
9. Microlicia rosmarinoides (Mart. & Schrank ex DC.) Versiane & R.Romero
10. Microlicia trembleyiformis Naudin
11. Microlicia tridentata (Naudin) Versiane & R.Romero
Specimens examined
Aguiar 102 (6). Almeda et al. 7726 (5); 7751 (7); 7753 (5); 8395 (5); 8549 (5); 8555
(7); 8580 (3); 8862 (5); 8911 (7); 9076 (6); 9179 (5); 9203 (7). Alves 595 (11); 600
(6); 7359 (11). Alves & Almeida-Lafetá 5573 (5). Alves et al. 2109 (5); 6925 (5). An-
drade et al. 374 (5). Antar et al. 1662 (5). Antônio-Silva et al. 315 (7). Araújo et al.
2043 (4); 323 (5); 334 (5). Arbo et al. 3906 (5); 4030 (5). Assis et al. 520 (6). Avezum
6017 & Almeida 10 (6). Azevedo et al. 675 (6). Badini & Ferreira 9774 (11). Baitello
et al. 2136 (6). Barreto 10734 (7); 1182 (7); 4621 (6); 4654 (6); 6745 (3); 6757 (6);
6759 (6); 6769 (5); 6771 (5); 6772 (5); 7023 (7); 7024 (7); 7025 (7); 7026 (putative
hybrid M. laniora × M. pentagona); 706 (5); 9019 (2). Barreto & Viégas s.n. (5). Bar-
ros & Feteira 1625 (6). Batista & Naves 405 (6). Baumgratz et al. 1127 (6). Bautista
1347 (6). Beraldo & França 85 (6). Bidá et al. CFCR11951 (4). Borges et al. 153 (5).
Bortoluzzi et al. 678 (7). Bovini & Barros 3235 (6). Brade 13735 (5); 14756 (7). Brade
et al. 18205 (6). Brandão 14509 (6); 20838 (7); 22298 (6); 28574 (5). Brotto 2600
(6). Brotto & Vieira 1921 (6). Bruniera et al. 37 (5). Bunbury s.n. (6). Bunge s.n. (5).
Caddah et al. 636 (6). Caiafa & Umbelino 172 (6). Camargo et al. 109 (6). Carmo 192
(9); 2262 (9); 377 (9); 4819 (5). Carvalho 1086 (6); 6986 (6); s.n. (5). Carvalho et al.
6651 (6). Casaretto s.n. (7). Castro et al. 281 (11). Castro et al. 283 (2); s.n. (5). Cav-
alcanti CFCR8314 (6). Cavalcanti et al. CFCR8035 (6). Ceccantini et al. 3914 (5).
Cerati et al. 246 (4). Chuckr et al. s.n. (5). Claussen 1 (5); 1617 (7); 1631 (11); 1637
(7); 1645 (5); 22 (7); 296 (7); 332A (5); 350 (7); 554 (5); 640 (5); 923 (5); s.n. (5,7).
Claussen 10 (2); 368 (2). Colletta 161 (6). Collo et al. s.n. (5). Contro & Marques 21
(7). Cordeiro et al. CFSC10504 (5). Costa 451 (6). Costa et al. 60 (6). Damazio 1025
(5); 1540 (5); 2026 (3); s.n. (277). Davis et al. 2298 (6). Delni et al. 78 (5); 97 (6).
Delprete 9205 (6). Drummond et al. 321 (1). Duarte 2004 (7); 2692 (7); 2767 (10);
8157 (11). Duarte & Brade 1155 (6). Ducke s.n. (5). Eiten & Eiten 7305 (6). Eiten et
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 99
al. 3019 (6). Emygdio 3312 (6); 3314 (5); 3353 (5); 3613 (6). Escaramai et al. 52 (3);
65 (5). Estevan et al. 617 (6). Farah et al. 580 (6). Faria & Mazucato 105 (5). Faria et
al. 1332 (6). Farinaccio et al. 36 (5). Fernandes s.n. (7). Fernandes et al. 1468 (5). Fer-
reira 5544 (5). Ferreira & Helena 7844 (11). Ferreira et al. 433 (5). Fontana 2309 (6).
Fontana et al. 2288 (5); 7678 (6). Forero et al. 7700 (5); 7831 (5). Foresto et al. 59 (6).
Forzza et al. 3427 (6); 4897 (4); 6344 (5); 6359 (5); 993 (5). Foster & Leoni 64 (6).
Fraga & Saddi 1786 (6). Fraga et al. 3333 (5); 3341 (5). Franco et al. 1270 (5). Furlan
et al. CFCR771 (4). Ganev 879 (6); 965 (6). Gardner 379 (6); 380 (6); 4601 (5); 4602
(6). Giulietti & Menezes 4017 (7). Giulietti et al. CFCR13780 (11); CFSC12492 (3);
CFSC12560 (3); CFSC12564 (5). Glaziou 11953 (5); 12704 (6); 14740 (5); 14741
(5); 14742 (7); 14743 (11); 14745 (2); 14746 (8); 14747 (9); 16679 (6); 16778 (6);
16781 (11); 17513 (11); 17573 (11); 18232 (2); 19221 (10); 19239 (8); 19242 (9);
21300 (1); 2579 (6); 8680 (6); 9454 (6); s.n. (5). Goldenberg & Michelangeli 2093
(6). Goldenberg & Silveira 1573 (5). Goldenberg et al. 1652 (6). Gonçalves et al. 172
(6). Gounelle s.n. (5). Grandi et al. 6593 (2). Groppo & Ulwin 676 (5). Guarçoni &
Sartori 1367 (6). Guedes et al. 12092 (6). Guiamarães 333 (6). Guillemin 921 (6); 946
(6). Hage et al. 2317 (6). Handro 790 (6). Hatschbach 27277 (6); 2775 (6); 39954
(6); 41337 (4); 647 (6); 6908 (6). Hatschbach 26964 (6); 41498 (6). Hatschbach &
Guimarães 55455 (6); 56167 (6). Hatschbach & Hatschbach 52005 (4). Hatschbach
& Pelanda 28014 (6). Hatschbach & Silva 49393 (6). Hatschbach et al. 27400 (5);
28759 (7); 29958 (7); 42967 (6); 54239 (4); 55434 (6); 68067 (4); 68138 (5); 74974
(6); 75995 (6); s.n. (7). Heluey & Castro 108 (6). Hensold 778 (5). Hensold et al.
CFCR2880 (11); CFCR3525 (4). Heringer 8544/736 (6). Hoehne 20521 (66). Hottz
et al. 302 (6). Irwin 19974 (5). Irwin et al. 19974 (5); 20178 (7); 20486 (7); 20537
(7); 29029 (11); 29079 (11); 29213 (7); 31141 (6); 34521 (6); 8149 (6). Joly
CFSC2405 (5); CFSC82 (5). Joly & Semir CFSC2957 (7); CFSC3196 (7). Kamey-
ama et al. CFSC10403 (3). Kinoshita et al. 43767 (6). Kirizawa 3665 (6). Kirizawa et
al. 2132 (6). Klein et al. 2465 (1). Kollmann 231 (6). Kral et al. 72723 (4). Kral et al.
72997 (5). Krieger 10641 (5); 8857 (6). Krieger et al. 24423 (6). Kubo et al. 109 (5);
125 (5). Kuhlmann 4381 (6). Leitão et al. 17281 (5). Leitão Filho et al. 7893 (4). Le-
oni 1 (6). Lima 400 (6). Lima et al. 1296 (5); 1443 (5); 49 (5); 8445 (6). Loefgren 557
(6). Lombardi 4045 (6). Longhi-Wagner et al. CFCR9184 (11). Lund 135 (5). Mac-
edo 3758 (5). Machado et al. 153 (1). Magalhães 1160 (11); 2278 (7). Maguire et al.
49016 (5); 49140 (5). Mamede et al. 666 (6). Mantovani 99 (5). Marcolino 222 (6).
Marcondes-Ferreira et al. 281 (5). Marques et al. 274 (5). Marquete et al. 2332 (6);
3817 (5). Martens 383 (5); 524 (6); 7 (5). Martinelli 19464 (6). Martinelli & Tavora
2583 (5). Martinelli et al. 10932 (6). Martius 930 (5); 931 (6); 961 (6); 989 (6); s.n.
(59). Mattos & Rizzini 107 (5); 480 (5). Meireles et al 1362 (5). Meireles et al. 1124
(4); 1766 (6). Mello-Silva et al. 1217 (6); 509 (4); CFCR7836 (6). Mello et al. 61 (5).
Mendes & Araújo 970 (6). Mendonça 557 (11). Messias et al. 1922 (5); 2151 (5).
Meyer 1171 (1). Michelangeli et al. 1580 (7); 1586 (11); 1595 (11). Mimura 479 (6).
Monge et al. 2595 (6); 384 (5). Monteiro 78 (6). Mosen 1971 (6). Moutinho et al. 76
(6). N. da Cruz 135 (6). Nakajima & Romero 3040 (5). Nakajima et al. 1476 (10);
Ricardo Pacico et al. / PhytoKeys 126: 1–101 (2022)
100
4547 (5); 4764 (4). Netto s.n. (5). Nicolau et al. 2212 (6). Occhioni et al. s.n. (9).
Oliveira 1013 (6); 25197 (6); 382 (8). Oliveira & Giacomin 47 (2); 84 (2). Oliveira et
al. 478 (7); 480 (2); 508 (6); 518 (7); CFCR12997 (4). Ordones et al. 1856 (5); 213
(7); 84 (11). Orlandi et al. 778 (6). Pacico 185 (5); 191 (6); 295 (8); 413 (6); 565 (4).
Pacico & Bressan 290 (11); 291 (2); 294 (7); 296 (2); 303 (11); 380 (1); 531 (7); 565
(4). Pacico & Carmo 154 (3). Pacico & Simoes 353 (4). Pastore et al. 685 (6). Paula
& Grandi s.n. (6). Paula et al. 136 (5); 295 (5); 8 (5). Pedrosa 110 (9); 120 (11). Pena
& Viana 365 (5); 417 (3). Pereira 309 (6). Pereira & Pabst 152 (5); 2944 (5); 3107 (5).
Pereira & Pabstii 2619 (11). Peron 220 (2); 256 (7); 260 (7); 268 (2); 269 (2). Pirani
& Mello-Silva CFCR10814 (4). Pirani & Yano 692 (7); 696 (5). Pirani et al. 1663 (1);
1694 (1); 5082 (5); 696 (5); CFCR11211 (5). Pires & Braga s.n. (5). Pivari 15 (6).
Pivari et al. 2512 (7). Polisel et al. 176 (6). Porto 2834 (10). Prates s.n. (6). Queiroz
2808 (6). Raben 409 (6); 428 (5). Rapini et al. 296 (7). Reginato et al. 1401 (5).
Rezende 507 (7). Ribas et al. 6769 (6); 8528 (6). Riedel s.n. (255611). Rocha 694 (3).
Rocha et al. 497 (4); 602 (5); 672 (5). Rolim 366 (2). Rolim et al. 326 (11); 327 (11);
328 (11); 329 (5); 353 (7); 361 (7); 370 (7); 386 (5); 394 (11); 61 (5). Romero 4157
(10). Romero & Nakajima 3021 (6); 3432 (6); 3593 (10); 5998 (7). Romero et al.
2155 (6); 5303 (7); 5307 (5); 7550 (10); 8627 (3); 8694 (10); 8689 (10). Roque 2869
(6). Roschel et al. s.n. (11). Roth s.n. (5). Rutter 128 (11); 150 (11); 161 (11); 186
(11); 191 (11). Saint-Hilaire 160 [catal. B] (10); 216 (5); 2399 [catal. B2] (11); s.n.
(7). Saint-Hilaire s.n. [catal. D, n° 462] (6). Sakuragui & Souza 38 (5). Salatino et al.
14 (5); 18 (5). Salgado 167 (6). Samento & Bautista 859 (6). Sampaio 7194 (5). San-
tos 411 (10). Saraiva et al. 85 (5). Sazima CFSC4259 (7). Scatigna & Galvão 376 (4).
Schüch s.n. (7). Schwacke 10135 (11); 10814 (7); 9184 (9); 9325 (7); 9368 (2). Sellow
1154 (6); 1316 (8); 1446 (5); 1727 (5); 260 (3); 5278 (6); s.n. (3, 5). Semir CFSC2005
(5); CFSC4133 (5); CFSC5001 (5); CFSC5068 (5); CFSC5607 (3). Semir CFSC5608
(7). Semir & Joly CFSC3743 (7). Semir & Sazima CFSC3395 (7). Semir et al
CFCR230 (5). Semir et al. 28809 (5). Sena s.n. (3). Sevilha et al. 7075 (6). Sheperd &
Kirzenzaft 10214 (3). Silva s.n. (6). Silva & Abe 2310 (6). Silva & Grandi 6627 (6).
Silva & Koch 7610 (6). Silva et al. 1043 (7); 6478 (6). Silveira s.n. (3). Sobral et al.
12797 (6); 14085 (6); 14508 (7). Souza 1606 (5). Souza & Miranda 1639 (3). Souza
et al. 1013 (6); 1108 (6); 11565 (5); 25181 (5); 2584 (9); 5934 (6); 6098 (6); 8015
(11); 8047 (11); 8061 (11). Stehmann 2532 (6). Stehmann & Morais 2354 (5); 2650
(5). Tales et al. 17 (5). Tameirao Neto & Mansur 4874 (5). Tannus 760 (6). Teixeira
s.n. (5); s.n. (5). Temponi & Vasconcelos s.n. (5). Torres s.n. (5). TSMG & Tales 81
(5). Ule 2528 (9); 2543 (7). Vainio 33501 (7). Valente & Azevedo 57 (6). Valente et
al. 1229 (5); 1230 (5); 2574 (5); 535 (5). Vasconcelos s.n. (55). Vauthier 37 (5); 43
[part] (6); s.n. (5). Verdi et al. 6501 (5). Versiane & Devides 677 (9); 682 (11). Versi-
ane et al. 305 (6). Vidal s.n. (9). Vieira & Yamamoto 26186 (6). Wängler & Ferreira
1352 (6). Warming s.n. (6). Weddell s.n. (8). Widgren 967 (6); s.n. (6). Williams &
Assis 6352 (5); 7274 (6). Yamamoto 1549 (6). Zappi et al. 2504 (6); CFCR9903 (4);
CFSC9353 (5).
Systematics of the Trembleya s.s. clade of Microlicia (Melastomataceae) 101
Supplementary material 1
List of species sampled in this study, their respective Genbank accession codes (for
nrITS and nrETS), and specimen vouchers
Authors: Ricardo Pacico, Frank Almeda, Darin S. Penneys, Karina Fidanza
Data type: table (word document)
Explanation note: List of species sampled in this study, their respective Genbank acces-
sion codes (for nrITS and nrETS), and specimen vouchers. Codes highlighted in
bold refer to sequences previously obtained from GenBank.
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/phytokeys.126.91032.suppl1
Available via license: CC BY 4.0
Content may be subject to copyright.
