Monograph |
Corresponding author: Ricardo Pacifico ( ricardo_b9@hotmail.com ) Academic editor: Marcelo Reginato
© 2022 Ricardo Pacifico, Frank Almeda, Darin S. Penneys, Karina Fidanza.
This is an open access article 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.
Citation:
Pacifico R, Almeda F, Penneys DS, Fidanza K (2022) Systematics of the Trembleya sensu stricto clade of Microlicia (Melastomataceae, Lavoisiereae). PhytoKeys 216: 1-101. https://doi.org/10.3897/phytokeys.216.91032
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A systematic monograph of the Trembleya s.s. clade is presented, a Brazilian endemic lineage of Melastomataceae 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). Then, we provide a systematic revision and new circumscription of the Trembleya s.s. clade, including line drawings, photos of living specimens, leaves and floral 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.
Brazil, Cadeia do Espinhaço, campo rupestre, Endemism, Minas Gerais
Melastomataceae Juss. comprises about 5,880 species in 173 genera with about two-thirds of this diversity restricted to the neotropics (
As compared to the treatment in “Flora Brasiliensis”, the most recent infrafamilial classifications of Melastomataceae show significant improvements. Based on DNA sequence data, several new Neotropical tribes have been delimited, i.e. Henrietteeae Penneys, Michelang., Judd & Almeda (
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 (
Trembleya s.s. is one of the clades currently recognised as part of Microlicia s.l. (
Trembleya was named by Augustin Pyramus de Candolle (1778–1841) in honour of the members of the family Trembley for their contributions to several fields of science (
Trembleya was recognised as a heterogeneous group of species since it was first delimited by
In “Flora Brasiliensis”,
Based on an expanded phylogenetic analyses using five molecular markers (atpF-atpH, trnS-trnG, nrITS, nrETS and waxy) and 12 morphological characters for 113 taxa in Lavoisiereae,
To investigate phylogenetic relationships in Lavoisiereae, we used two different datasets including samples from the seven genera traditionally recognised in the tribe (
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 modified to extend the incubation time in AP1 Buffer and RNAase to 1 hour, followed by 1 hour of precipitation in ice. A few samples were extracted using the modified CTAB (hexadecyltrimethylammonium bromide) protocol of
We amplified and sequenced the ribosomal markers nrETS and nrITS. In Melastomataceae, the nrETS region has been used by
Contigs were assembled and edited using Geneious 11.0 (https://www.geneious.com). The same software was employed to align (using Muscle algorithm), concatenate and export the sequences as fasta files. Low-quality sequences were discarded. Final manual edits on the alignment were performed using Mega 7 (
Specimens of all species of the Trembleya s.s. clade were examined. This study was carried 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
Populations of all but two species (M. rosmarinoides and M. trembleyiformis) of the Trembleya s.s. clade were sampled in the field 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 Diamantina (Bahia State), Chapada dos Veadeiros and Serra dos Pirineus (Goiás State) and Parque Estadual do Guartelá (Paraná State). All specimen identifications were carefully checked by the authors. We considered one specimen as a putative hybrid between M. pentagona and M. laniflora because it clearly intermixes characters between these two species and was collected in an area where they occur sympatrically.
The 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 (
Scanning Electron Microscope (SEM) photos were taken from herbarium specimens. Sections of leaf blades (all species in the clade), anthers (four species), seeds (five species) and pollen grains (M. pithyoides) were affixed using double-stick tape on pin stubs, sputter-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.
Dried leaves of M. altoparaisensis, M. chamissoana and M. flaviflora were obtained from exsiccatae (housed in HUEM), rehydrated in sodium hydroxide (NaOH) 5% (
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 (
Following the guidelines of criterion B of
The 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
Overall, species traditionally recognised in Trembleya (for example, by
The analyses of nrETS and nrITS resulted in incompletely resolved trees with several weakly-supported clades, making it impossible to evaluate the monophyly of most clades within Microlicia s.l. This was expected since these lineages have been recovered with very short branch lengths in family-level phylogenetic analyses, based on the multi-loci approach that used a larger sampling of Melastomataceae (
Species of the Trembleya s.s. clade are woody and perennial erect shrubs, usually 0.5–2 m tall, small trees 2.5–4 m tall (e.g. M. laniflora and M. parviflora) or caespitose shrubs (e.g. M. pithyoides and M. rosmarinoides). Species can be highly variable in habit, like M. laniflora, 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.
Habit in the Trembleya s.s. clade of Microlicia A Celso de Paiva (ICMBio) taking a picture of a small M. laniflora tree B M. parviflora, small tree C M. laniflora with divaricate habit D M. chamissoana with divaricate habit E M. rosmarinoides, a small, much-branched shrub F, G Saxicolous shrubs: F M. flaviflora G M. pentagona. Photos: A by O. Graeff B by R. Penati C, E by L. Pedrosa D, F, G by R. Pacifico.
Information on root and stem anatomy and development is limited to the study of
Microlicia parviflora stands out by having a spiralled aerenchymatous polyderm surrounding the root and stem (
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.
Leaves in the Trembleya s.s. clade of Microlicia in abaxial view A M. tridentata B M. altoparaisensis C–E M. pentagona F M. rosmarinoides G M. pithyoides H M. flaviflora I M. laniflora J M. parviflora 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 Pacifico 295 (CAS, HUEM, SPF) H Hatschbach et al. 52005 (S) I Pacifico 185 (HUEM, SPF) J Pacifico 191 (HUEM) K Romero & Nakajima 3593 (HUFU, K, UEC) L Pacifico 290 (CAS, SPF) M Romero et al. 8627 (HUEM, HUFU, RB).
Information on leaf anatomy is available for M. altoparaisensis (
Anatomical features of leaves in the Trembleya s.s. clade of Microlicia A–C Microlicia altoparaisensis: 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 flaviflora, 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: Stomata. All photos by A.A.O. Carmo. Voucher specimens: A–C Pacifico & Bressan 380 (CAS, HUEM, SPF) D, E Pacifico & Carmo 154 (HUEM, UEC) F Mello-Silva et al. 509 (SPF).
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
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, eglandular trichomes and stomata on the leaf abaxial surface D Microlicia flaviflora, stomata on the leaf abaxial surface E, F Microlicia laniflora: 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 Pacifico & Bressan 380 (CAS, HUEM, SPF) B Pacifico 290 (CAS, SPF) C Pacifico & Carmo 154 (HUEM, UEC) D Mello-Silva et al. 509 (SPF) E, F Almeda et al. 9179 (CAS, UEC).
SEM images of leaves in the Trembleya s.s. clade of Microlicia A Microlicia parviflora, 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 trichome on the abaxial surface F Microlicia tridentata, detail of a glandular trichome on the adaxial surface. All photos by R. Pacifico. Voucher specimens: A Almeda et al. 8483 (CAS, HUFU, NY) B, C Pacifico 295 (CAS, HUEM, SPF) E Romero & Nakajima 3593 (HUFU, UEC) F Pacifico 290 (CAS, HUEM, SPF).
Given the diversity of inflorescence types and the presence of many similar structures which are not guaranteed homologous, efforts have been made to establish natural systems for the flower-bearing parts of the Angiosperms (e.g.
Inflorescences in the Trembleya s.s. clade of Microlicia A M. altoparaisensis B M. chamissoana C M. laniflora D–F M. flaviflora G M. parviflora H M. tridentata. Photos: A by V. E. Bressan B by F.A.O. Silveira C by L. Pedrosa D, E by R. Pacifico F by A.V. Scatigna E by F. Almeda F by F.A. Michelangeli.
In the Trembleya s.s. clade, seven species have simple or compound dichasia (M. altoparaisensis, M. calycina, M. chamissoana, M. flaviflora, M. laniflora, M. parviflora and M. tridentata) and the remaining four have inflorescences reduced to solitary flowers at the apical region of the branches (M. pentagona, M. pithyoides, M. rosmarinoides and M. trembleyiformis). Floral pedicels are usually evident and measure 0.3–4 mm long. These are inconspicuous to 0.2 mm long only in M. altoparaisensis. Like some species of the Lavoisiera clade (
The hypanthium that envelops the superior ovaries is campanulate to urceolate and its length varies between 1.7 and 6.5 mm. The 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 usually referred to as the torus (
The calyx lobes that develop above the torus are united at the base and that union forms the calyx tube, which, in the Trembleya s.s. clade, varies in length between 0.1 and 0.7 mm. The 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. The indumentum of both calyx tube and calyx lobes is usually similar to that of the hypanthium. Differences in the shape, size and indumentum of the calyx lobes are generally useful diagnostic characters at the species level in this clade.
Flowers are generally 5-merous (Fig.
Diversity of flowers in the Trembleya s.s. clade of Microlicia A M. chamissoana B M. parviflora C M. flaviflora D M. laniflora 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.
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 pedoconnectives 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. The pedoconnectives of the antesepalous stamens are generally 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. The exception is M. altoparaisensis, which has subisomorphic stamens with all anthers yellow. The filaments are filiform, glabrous, measuring 1.5–6.3 mm long, usually similar in colour to the petals. The anthers are oblong (Fig.
SEM images of anthers and anther rostra in the Trembleya s.s. clade of Microlicia A Microlicia parviflora, anther of an antesepalous stamen B Microlicia tridentata, anther of antesepalous stamen C Microlicia flaviflora, anther rostrum of an antesepalous stamen D Microlicia flaviflora, anther rostrum of antepetalous stamen E Microlicia parviflora, anther rostrum of an antesepalous stamen F Microlicia parviflora, anther rostrum of an antepetalous stamen. Voucher specimens: A, E, F Pirani et al. CFSC 12361 (SPF) B Pacifico & Bressan 290 (CAS, HUEM) C, D Hatschbach et al. 54239 (CAS, INPA, MBM).
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 Pacifico & Bressan 290 (CAS, HUEM) E, F Pacifico 295 (CAS, HUEM, SPF).
Pollen grains in Melastomataceae are small, generally tricolporate, radially symmetrical and isopolar (
The 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. These differences in ovary locule number are additional characters that can be used for species recognition. The 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 intrusions. The style is filiform (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.
The brownish capsules are loculicidal measuring 2.3–8.0 mm in length, initially enveloped by the hypanthium and calyx that rupture and flake away tardily with age (Fig.
Capsules and infructescences in the Trembleya s.s. clade of Microlicia A Microlicia laniflora, infructescence B Microlicia parviflora, capsule enveloped by the hypanthium C–E Microlicia pentagona: C capsule enveloped by the hypanthium D infructescence E capsules enveloped by reddish mature hypanthia. Photos: A by F. Almeda B, C by F.A. Michelangeli E, F by R. Pacifico.
Seed micromorphological characters have been traditionally used in the tribal and generic classification of Melastomataceae (
SEM images of seeds in the Trembleya s.s. clade of Microlicia A Microlicia calycina B Microlicia laniflora C, D Microlicia parviflora E Microlicia pentagona F Microlicia rosmarinoides. All photos by R. Pacifico. Voucher specimens: A Pacifico & 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]).
Information on seed germination in the clade is limited to studies of M. parviflora and M. laniflora. According to
Flowers are hermaphroditic with a strongly dimorphic (rarely subisomorphic) androecium and poricidal anthers. Plants of this clade provide pollen as a primary resource to reward pollinators and apparently developed floral traits to protect the pollen, select pollinating agents and precisely deposit pollen on the body of the pollinator. These characteristics are usually understood as strategies to deal with the “pollen dilemma”, in which pollen grains fulfil two main functions, i.e. to transport the male gamete and to reward flower visitors with resource supply (
By having flowers with vibrant colours and diurnal anthesis, most species of Lavoisiereae seem to meet the requirements of the classic melittophily syndrome (
There is no evidence of apomixis in M. laniflora (
Populations of Microlicia parviflora from different localities may be distinct in phenological behaviour, with annual or episodic flowering and continuous or annual fruiting; this species is commonly reported from both typical campo rupestre and gallery forests (
Knowledge of secondary metabolites of Lavoisiereae is largely restricted to the work by
Information on histochemical tests is available only for M. parviflora and M. laniflora. Regarding the first, phenolic compounds were reported from the endodermis and parenchyma (
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 (
The Trembleya s.s. clade is restricted to Brazil (Fig.
Endemicity analyses detected one consensus area of endemism for the Trembleya s.s. clade, which resulted from the merging of four individual areas of endemism (Fig.
The 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.
The region with highest richness and weight endemism of the clade is the Iron Quadrangle (Quadrilátero Ferrífero), located in the Municipalities of Barão dos Cocais, Belo Horizonte, Brumadinho, Caeté, Catas Altas, Congonhas, Itabirito, Mariana, Nova Lima, Ouro Branco, Ouro Preto, Raposos, Rio Acima, Sabará and Santa Bárbara (Minas Gerais State). The Iron Quadrangle is one of the world’s largest mineral provinces, 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 (
The 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 territory (
The traditional hypothesis for plant diversification 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 (
In situ diversification apparently played an important role during the evolutionary history of the Trembleya s.s. clade, along with several other plant lineages (
Overall, vascular plant lineages restricted to campo rupestre are highly vulnerable to disturbances and their conservation deserves special attention (
The historical stability of campo rupestre vegetation in the face of climatic changes is still under debate (
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. The Brazilian Government already recognises three species of this clade as Endangered (EN), i.e. M. calycina, M. chamissoana and M. flaviflora and one as Critically Endangered (CR), M. pithyoides (
Perennial shrubs or treelets. Leaves petiolate, not imbricate, not keeled, the adaxial surface glandular-punctate to glabrescent, venation basal acrodromous, impressed on the adaxial surface and prominent on the abaxial surface, consisting of amphicribral or bicollateral vascular bundles. Flowers usually 5-merous, diplostemonous, pedicellate, subtended by a pair of bracteoles. Hypanthia not fused to the ovary, lacking 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.
Perennial erect shrubs or small trees (0.1–)0.3–4 m tall, woody, sometimes densely branched. Distal branches quadrangular, usually light green (when fresh) and glutinous, glandular-punctate, sometimes granulose or pruinose, eventually covered 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 towards the base. Leaves decussate, petiolate, not imbricate, not keeled, papyraceous, chartaceaous 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), glutinuous, glandular-punctate to glabrescent, glandular trichomes (when present) appearing sessile (i.e. on peduncles too short to be seen with a 40× magnification stereoscope). Abaxial surface usually green (when fresh), becoming pale green (when dry), always lighter 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 perpendicular to the mid-vein and branching towards the leaf margin. Inflorescences simple or compound dichasia, consisting of biparous cymes throughout or proximally biparous and distally uniparous cymes or reduced to solitary flowers on the apical region of the branches. Inflorescence 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, linear, elliptic, lanceolate, ovate, narrowly elliptic, oblong or oblanceolate, 1–3(–5)-nerved from the base, entire to slightly serrulate, rarely ciliate, usually differing in shape, but similar in indumentum to the principal leaves. Flowers (4–)5(–6)-merous, diplostemonous, pedicellate, subtended by a pair of bracteoles. Hypanthia 1.7–6.5 mm long, 1.5–5.2 mm 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 0.7–9.7 mm 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 4.5–26 mm 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 dimorphic or subisomorphic, glabrous throughout, filaments linear, white, pink or yellow, pedoconnectives well-developed, anthers oblong, 2-celled (tetrasporangiate), rostrate, apical pores circular and ventrally inclined. Larger (antesepalous) stamens (4–)5(–6), filaments 1.5–6.3 mm long, pedoconnectives 1.3–7.3 mm 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), filaments 1.5–5.4 mm long, pedoconnectives 0.2–1.9 mm long, ventral appendages usually up to 0.1 mm long, apex truncate to emarginate, 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, cylindrical or globose, superior, (3–4–)5-locular, glabrous. Styles 3–10 mm long, filiform, sigmoid to incurved, white, pink or yellow, glabrous, stigmas punctiform. Capsules loculicidal, 2.3–8.0 mm long, 2.3–6.0 mm wide, ovoid or globose, the torus initially constricted at the apex, dehiscent from the apex to the base, columella deciduous. Fruiting calyx tubes 0.2–3.1 mm long. Fruiting calyx lobes 1.2–11.5 mm long, rarely thickened. Seeds 0.3–0.9 mm long, reniform, the testa foveolate-reticulate.
Based on collections housed in the herbarium at Paris (P),
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. laniflora |
– | 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. |
M. chamissoana |
– | Leaves with the abaxial surface exposed, tertiary venation (if present) not “foveolate-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 |
6 | Abaxial leaf venation with tertiaries absent or little evident (Fig. |
7 |
– | Abaxial leaf venation with tertiaries evident (Fig. |
8 |
7 | Calyx lobes subulate, 6.2–8.5 mm long in flower, becoming stout and thick in fruit (Fig. |
M. pentagona |
– | Calyx lobes oblong to triangular, 2.8–3.4 mm long in flower, 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. |
M. tridentata |
– | Leaf margins entire or slightly serrulate throughout, tertiary veins not surrounding 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. flaviflora |
– | 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; flowers solitary; calyx lobes narrowly-triangular; ovaries 3–5-locular | M. trembleyiformis |
– | Branchlets unwinged; flowers disposed in compound or simple dichasia; calyx lobes triangular; ovaries 5-locular | M. parviflora |
Trembleya altoparaisensis R.B.Pacifico, 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. Furlan & C. Kameyama (holotype: SPF!; isotypes: HUEM!, K!, UEC!, US!).
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. Petioles 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), concolorous (when dry), base attenuate, apex rounded to acute, margin flat or slightly revolute, slightly serrulate and glandular-punctate, 5-nerved from the base, one pair of acrodromous 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. Inflorescences compound dichasia consisting of proximally biparous, distally uniparous cymes, not congested. Bracts (including petioles) 1.0–4.0 cm 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) inconspicuous 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-punctate. 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, obovate, apex acuminate, margin entire and glabrous, both surfaces glabrous. Stamens (8)10, subisomorphic. Larger (antesepalous) stamens (4–)5, filaments 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, filaments 2.7–2.8 mm long, white, pedoconnectives 0.7–1.0 mm long, white, inconspicuous appendages ca. 0.1 mm long, white, apex truncate, thecae (excluding rostra) 2.5–3.2 mm 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.
Microlicia altoparaisensis A habit B detail of the glandular-punctate indumentum on the branches C leaf abaxial surface D bracteole adaxial surface E floral bud F flower 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.
Microlicia altoparaisensis can be recognised by its oblong-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, inflorescences composed of proximally biparous and distally uniparous cymes, white petals, yellow subisomorphic stamens and 3(–4)-locular ovaries. The 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 (
Probably endemic to Chapada dos Veadeiros in Alto Paraíso de Goiás Municipality, Goiás, Brazil (Fig.
Microlicia altoparaisensis is known from less than 10 collections. The EOO is 7.025 km2 and the AOO is 16 km2. Based on
The collection Glaziou 21300 was listed under the name Trembleya debilis by
Brazil. Goiás: Alto Paraíso de Goiás Municipality, Chapada dos Veadeiros, Drummond et al. 321 (MBM, NY, RB), Klein et al. 2465 (HUFU, UFG), Machado et al. 153 (HUFU), Meyer 1171 (UEC, UPCB), Pacifico & 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).
Trembleya calycina Cham., Linnaea 9(4): 430. 1835. basionym. Type: Brazil. “Brasilia, 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/herbarium/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, first-step designated by
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, first-step designated by
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, chartaceous (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 becoming 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 glandular-punctate, appearing glabrous when dry, abaxial surface densely glandular-punctate. Inflorescences simple dichasia or reduced to solitary flowers, 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, elliptic, 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, 1.9–2.2 mm wide at the torus, campanulate, light green or reddish (when fresh), externally glandular-punctate. Calyx tubes 0.4–0.7 mm long. Calyx lobes (at anthesis) 3.5–5.2 mm long, 0.5–0.7 mm wide at the base, subulate, apex acuminate, margin entire, (when 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, filaments 4.7–4.9 mm long, pink, pedoconnectives 5.9–6.2 mm long, pink, appendages 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, filaments 3.8–4.1 mm 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, oblong, rostra 0.4–0.6 mm long, the circular pores ca. 0.2 mm wide. Ovary 2.0–2.2 mm 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 hypanthium, 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.
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 flowers 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 species). Both species share the glandular-punctate indumentum, inflorescences reduced to solitary flowers (sometimes perfect dichasia only in M. calycina), subulate calyx lobes, magenta petals, and bicoloured anthers. Microlicia calycina differs 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). These two species may occur sympatrically in the seasonally dry grasslands of Parque Estadual do Itacolomi (
Endemic to central and southern Minas Gerais (Fig.
Microlicia calycina is known from about 20 collections. The EOO is 481 km2 and the AOO is 20 km2. This species is currently recognised as endangered (EN) by the Brazilian Government (
We agree with
Brazil. Minas Gerais: Caeté Municipality, Serra da Piedade, 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), Pacifico & 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), Pacifico & 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]).
Trembleya chamissoana Naudin, Ann. Sci. Nat., Bot. Sér. 3, 12: 270. 1849. basionym. Type: Brazil. “Brasilia, Itambé” [Minas Gerais, probably Santo Antônio do Itambé], 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).
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), elliptical, 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 flat 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 surface sparsely glandular-punctate, appearing glabrous when dry, abaxial surface densely glandular-punctate and covered with gland-tipped trichomes 0.2–0.4 mm long. Inflorescences simple or compound congested dichasia consisting of biparous cymes, or reduced to solitary flowers. 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 trichomes 0.2–0.4 mm long. Calyx tubes 0.9–1.2 mm long. Calyx lobes (at anthesis) 4.5–6.7 mm long, 1.9–2.1 mm 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 (antesepalous) stamens 5, filaments 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 rostra) 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, filaments 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 0.4–0.6 mm long, the circular pores ca. 0.2 mm wide. Ovary 2.2–2.8 mm 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 hypanthium, torus constricted at the apex, fruiting calyx tubes 1.5–1.7 mm long, fruiting calyx lobes 5.2–6 mm long, not thickened. Seeds ca. 0.5 mm long, reniform.
Microlicia chamissoana A habit B leaf abaxial surface C bracteole abaxial surface D floral bud E Flower in lateral view F flowering 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).
Largely restricted to the Serra do Cipó in central Minas Gerais (Fig.
Microlicia chamissoana is known from fewer than 20 collections. The EOO is 885.927 km2 and the AOO is 32 km2. This 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. This species is already recognised as endangered (EN) by the Brazilian Government (
Microlicia chamissoana may be recognised by its elliptic leaf blades with tertiaries densely reticulate and randomly branching, congested inflorescences or solitary flowers and narrowly triangular calyx lobes 4.5–5.7 mm long. It is probably more closely related to M. laniflora and M. pentagona, both of which may occur sympatrically with M. chamissoana at Serra do Cipó. Microlicia chamissoana differs from M. laniflora 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 hypanthia (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 differs 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 granulose), leaf blades with tertiaries densely reticulate and randomly branching (vs. parallel 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).
Major variation in M. chamissoana involves habit and degree of inflorescence development. This 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). This specimen was described as a herb, but it has woody branches. Most of the specimens examined have congested, many-flowered inflorescences (e.g. Barreto 6745), although in some of the inflorescences, these are reduced to solitary flowers (e.g. Pacifico & Carmo 154, Almeda et al. 8580).
Based on F. Sellow s.n.,
The type specimens of M. chamissoana, as cited by
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, Giulietti et al. CFSC12560 (SPF); Morro do Pilar Municipality, Serra do Cipó, Silveira 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), Pacifico & 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]).
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!; isotypes: BHCB!, C!, CAS!, CTES, ESA!, G!, HUFSJ, K!, MO!, RB!, S, SPF!, US!, VIC, VIES).
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 flat, 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. Inflorescences 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) 3.3–4.1 mm 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, filaments 3.4–4.0 mm long, yellow, pedoconnectives 3.7–4.0 mm long, yellow, appendages 1.0–1.2 mm long, yellow, apex truncate 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, filaments 2.7–2.9 mm long, yellow, pedoconnectives 1.2–1.4 mm long, yellow, short appendages ca. 0.5 mm long, yellow, apex truncate, thecae (excluding rostra) 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 thickened. Seeds ca. 0.6 mm long, reniform.
Microlicia flaviflora A habit B leaf abaxial surface C bracteole abaxial surface D floral bud E flower in lateral view F flowering 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).
Endemic to northern Minas Gerais (Fig.
This species is known from about 20 collections. The EOO is 468,668 km2 and the AOO is 32 km2. Most of the populations of M. flaviflora occur within the following conservation units: Parque Estadual de Grão Mogol, Parque Estadual de Botumirim and Parque Estadual da Serra Nova, where this species is afforded protection. The Brazilian Government assigned a conservation status of Endangered (EN) to this species (
Microlicia flaviflora 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 yellow petals. In overall vegetative morphology, M. flaviflora resembles M. tridentata. In turn, its yellow petals, staminal filaments and styles are shared only with M. rosmarinoides. Microlicia flaviflora differs 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. magenta or rarely white) that are 6.0–8.8 mm long (vs. 11.5–13.0 mm long). Microlicia flaviflora differs from M. rosmarinoides by its taller habit 0.8–2.0 m tall (vs. 0.3–0.6 m 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 elliptic to slightly ovate (vs. linear to lanceolate) and have 5 basal acrodromous veins (vs. 1-nerved from the base), compound dichasia (vs. solitary flowers), 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).
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), Hatschbach & 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, 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), Pacifico & Simoes 353 (CAS, HUEM), Pacifico 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).
Melastoma laniflora 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/herbarium/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 laniflora (D.Don) Cogn. in Martius et al., Fl. Bras. 14(3): 130. 1883.Type: Based on Melastoma laniflora D.Don.
Trembleya laniflora 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 Gerais], 1878, F. Sellow 1727 (lectotype, designated here: K [K00530655]!; isolectotype: P [P00723419]!; image of lectotype available at http://specimens.kew.org/herbarium/K000530655).
Trembleya laniflora 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 laniflora 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 Gerais, 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 laniflora 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]!, R [R000168426]! US [US00623967]!; image of isolectotype at P is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p05317081).
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 indumentum, 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 flat, 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 the mid-vein, little reticulate and branching apically, adaxial surface glandular-punctate, usually pruinose and becoming glabrescent with age, abaxial surface densely covered with lanose eglandular trichomes 0.1–0.5 mm long. Inflorescences simple dichasia or reduced to solitary flowers, 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 attenuate, 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, 4.5–5.2 mm wide at the torus, campanulate to urceolate, surface hidden by the whitish lanose indumentum composed of eglandular trichomes 0.1–0.5 mm long. Calyx tubes 1.0–2.0 mm 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 eglandular trichomes 0.1–0.5 mm long. Petals 19–26 mm long, 10–15 mm wide, white, rarely with pink stains at the apical region, obovate, apex emarginate, margin entire and ciliate with eglandular trichomes 0.1–0.4 mm long at the apical region, both surfaces glabrous. Stamens 10(–12), strongly dimorphic. Larger (antesepalous) stamens 5(–6), filaments 5.2–6.3 mm long, white, pedoconnectives 6.1–7.3 mm long, white to light yellow, appendages 1.9–3.0 mm long, yellow, apex emarginate, thecae (excluding rostra) 2.3–2.7 mm long, vinaceous, oblong, rostra 0.4–0.6 mm long, the circular pores ca. 0.2 mm wide. Smaller (antepetalous) stamens 5(6), filaments 4.0–5.4 mm long, white, pedoconnectives 0.8–1.1 mm long, yellow, inconspicuous appendages ca. 0.1 mm 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 6.5–8.0 mm 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 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.
Microlicia laniflora A habit B leaf abaxial surface C bracteole abaxial surface D flower in lateral view E flowering 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).
Endemic to central and southern Minas Gerais (Fig.
Microlicia laniflora has a symbiotic relationship with arbuscular mycorrhizae (
Leaf and stem extracts from Microlicia laniflora have antimicrobial activity against Staphylococcus aureus and Micrococcus luteus (
This 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. The 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. laniflora. The EOO is 19,113.356 km2 and the AOO is 248 km2. Based on
Microlicia laniflora may be recognised by its branches, abaxial surfaces of the leaves and hypanthia that are densely covered by a lanose indumentum, white petals (rarely flushed 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 laniflora differs from M. pentagona by the branches, abaxial foliar surfaces and hypanthia densely covered by the lanose indumentum (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 flushed with pink (vs. magenta). For additional comparisons, see comments under M. chamissoana.
The four varieties proposed by
According to
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), Brumadinho Municipality, Serra do Rola-Moça, Carmo 4819 (BHCB), Serra da Calçada, Martens 7 (SPF), Martens 383 (SPF), Retiro das Pedras, Carvalho s.n. (HUFU [39992]), Stehmann & Morais 2650 (BHCB); Caeté Municipality, Serra do Gandarela, 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, “Diamantina 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, Serra 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), Delfini 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), Forzza 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), Farinaccio 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), Mattos & Rizzini 107 (RB, US), Mattos & Rizzini 480 (RB), Monge et al. 384 (UEC), Ordones et al. 1856 (BHZB, HUFU), Pacifico 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), Claussen 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 isolectotypes: K [K00957818], P [P005317063, P005317064, P005317070]), Vauthier s.n. (P [P005317061]).
(M. laniflora × M. pentagona). Brazil. Minas Gerais: Santana do Riacho Municipality [“Santa Luzia”], Serra do Cipó, Barreto 7026 (BHCB).
Meriania parviflora 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]!; isolectotypes: G [G00310210]!, M [M0165883]!).
Trembleya triflora 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).
Trembleya paniculata
Naudin, Ann. Sci. Nat., Bot. Sér. 3, 2: 154. 1844. Type: Brazil. “In campis circa Juruoca in prov. Minas Geraës” [Minas Gerais], 1816–1821, [catal. D, n° 462] A. Saint-Hilaire s.n (lectotype, first-step designated by
Trembleya parviflora (D.Don) Cogn. in Martius et al., Fl. Bras. 14(3): 127. 1883. Type: Based on Meriania parviflora D.Don.
Trembleya parviflora 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 parviflora subsp. triflora (Mart. & Schrank ex DC.) Cogn. in Martius et al., Fl. Bras. 14(3): 129. 1883. syn. nov. Type: Based on Trembleya triflora Mart. & Schrank ex DC.
Trembleya parviflora 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/catalognumber/mnhn/p/p05317041).
Trembleya parviflora 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/catalognumber/mnhn/p/p00723407).
Trembleya parviflora 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 isolectotype at P is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p05317106).
Trembleya parviflora 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-online 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 parviflora 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]!; isolectotypes: 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).
Trembleya parviflora var. multiflora 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 isolectotype at P is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p05317099).
Trembleya parviflora 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]!; image of lectotype is available at http://n2t.net/ark:/65665/392812a8c-782b-431c-b2fc-3644dcfad16e).
Trembleya parviflora 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 (lectotype, designated here: P [P00723410]!; image of lecotype is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p00723410).
Trembleya parviflora 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 parviflora var. triflora (Mart. & Schrank ex DC.) Cogn. in Martius et al., Fl. Bras. 14(3): 129. 1883. syn. nov. Type: Based on Trembleya triflora Mart. & Schrank ex DC.
Trembleya parviflora 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/catalognumber/mnhn/p/p00723405).
Trembleya parviflora 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://sweetgum.nybg.org/science/vh/specimen_details.php?irn=535234).
Trembleya parviflora 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 parviflora 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 [PH00027744]!, S [S-053231]!; image of lectotype is available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p05317115).
Trembleya parviflora 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]!; isolectotypes: 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).
Erect shrubs or treelets 0.7–4.0 m tall. Branchlets quadrangular, always glandular-punctate and usually pruinose-granulose, 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, light green (when fresh). Internodes 0.6–3.8 cm 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 flat or slightly revolute, entire to slightly 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 surface glandular-punctate and usually pruinose, eventually sparsely to densely covered with gland-tipped or eglanduar trichomes 0.1–0.9 mm long. Inflorescences simple or compound dichasia consisting of with biparous cymes, not congested. Bracts (including petioles) 0.7–1.6 cm long, 0.1–0.6 cm wide, 3-nerved, elliptic to narrowly elliptic, indumentum 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, 1.5–2.2 mm 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 hypanthia. Petals 4.5–8.1 mm long, 3.0–4.9 mm wide, white, usually flushed 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 0.1–0.4 mm long at the apical region, both surfaces glabrous. Stamens 10(–12), strongly dimorphic. Larger (antesepalous) stamens 5(–6), filaments 3.5–4.0 mm long, white or pink, pedoconnectives 3.0–4.0 mm 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), filaments 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 0.2–0.4 mm 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.
Microlicia parviflora A habit B leaf abaxial surface C bracteole abaxial surface D flower in lateral view E flowering 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).
(one specimen selected for each municipality of occurrence). 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); Buritizeiro, 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); Delfinópolis, Romero & Nakajima 3432 (HUFU); Diamantina, Hatschbach & Pelanda 28014 (MBM); Espera Feliz, Foster & Leoni 64 (ESA); Estrela do Sul, Costa et al. 60 (HUFU); Extrema, Yamamoto 1549 (UEC); Gouveia, Hatschbach 27277 (MBM); Grão Mogol, Cavalcanti 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, Delfini 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, Pacifico 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, Pacifico 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 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); Itirapina, 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]).
Endemic to Brazil in the States of Bahia, Minas Gerais, Goiás, Distrito Federal, Espírito Santo, Rio de Janeiro, São Paulo and Paraná (Fig.
Microlicia parviflora is the most widely distributed species in the clade and the most studied from an ecological perspective.
This is the most abundant species in the Trembleya s.s. clade, with more than 1000 collections currently housed in herbaria. The EOO is 1,152,078.308 km2 and the AOO is 1,640 km2. Populations of M. parviflora are found in all conservation units that protect the remaining species of the clade. As it occurs in large and dense populations, some have claimed that M. parviflora has the potential to become an invasive species (e.g.
Microlicia parviflora 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-flowered inflorescences, triangular 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. parviflora in leaf shape and venation and stamen morphology. Microlicia parviflora differs from M. pentagona by its leaves with tertiaries evident (vs. little evident), moderately reticulate and randomly branching (vs. parallel and/or branching apically), inflorescences (vs. solitary flowers) 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. parviflora differs from M. trembleyiformis by the unwinged branchlets (vs. with narrow wings ca. 0.2 mm wide), flowers disposed in inflorescences (vs. flowers solitary), triangular calyx lobes (vs. narrowly-triangular) and consistently 5-locular ovaries (vs. 3–5-locular). For a comparison between M. parviflora and M. altoparaisensis, see notes under the latter species.
The morphological variation found in M. parviflora is mainly related to leaf blade shape (varying from elliptic to ovate and lanceolate), indumentum on branches 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 inflorescence development (three- to many-flowered). The petals are usually white flushed with pink at the base, although some populations have entirely white petals, for example, in Serra de Carrancas (
The 15 varieties of M. parviflora proposed by
Like Microlicia cataphracta (Mart. & Schrank ex DC.) Versiane & R.Romero [= Lavoisiera imbricata (Thunb.) DC.], M. parviflora fits all the six criteria enumerated by
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, first-step designated by
Erect shrubs or treelets 0.5–1.5 m tall. Branchlets quadrangular, appearing 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 elliptic 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 flat, 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, vernicose. Inflorescences reduced to solitary flowers 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) 2.5–3.5 mm 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, 1.8–2.2 mm 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, filaments 4.0–5.0 mm long, pink, pedoconnectives 5.3–5.8 mm long, pink, appendages 1.2–1.6 mm long, yellow, apex emarginate to bilobate, thecae (excluding rostra) 1.2–2.0 mm long, vinaceous, oblong, rostra 0.5–0.7 mm long, the circular pores ca. 0.2 mm wide. Smaller (antepetalous) stamens (4)5, filaments 3.9–4.4 mm long, pink, pedoconnectives 1.0–1.6 mm long, yellow, inconspicuous appendages ca. 0.1 mm long, yellow, apex emarginate, 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) 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.
Microlicia pentagona A habit B leaf abaxial surface C bracteole abaxial surface D floral bud E flower in lateral view F flowering 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).
Endemic to Minas Gerais State (Fig.
Microlicia pentagona is known from about 70 collections. The 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
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, inflorescences reduced to solitary terminal flowers and calyx lobes 6.2–8.5 mm long that become thickened in fruit. The elongate calyx lobes of M. pentagona are comparable in length only to those of M. laniflora (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. laniflora and M. parviflora. For comparisons, see comments under these species.
Microlicia pentagona is remarkably variable in leaf shape and size. The 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 variation 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
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), Sobral 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]), Michelangeli et al. 1580 (NY, UPCB), Pacifico & Bressan 294 (HUEM, SPF), Peron 256 (RB), Peron 260 (RB), Rezende 507 (HUFU), Rolim et al. 353 (UPCB, VIC), Rolim 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), Romero 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), Pacifico & Bressan 531 (CAS, HUEM, RB). Romero & Nakajima 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), Hatschbach 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]).
Trembleya pithyoides Cham., Linnaea 9(4): 428. 1835. basionym. Type: Brazil. “Caraça” [Minas Gerais, Serra do Caraça], 20 December 1830, F. Sellow 1316 (lectotype, 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).
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 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), discoloured (when dry), base cuneate, apex rounded to acute, margin flat, entire and glandular-punctate, 1-nerved from the base, tertiaries not evident on the abaxial surface, adaxial surface glandular-punctate, abaxial surface glandular-punctate. Inflorescences reduced to solitary flowers apically on the branches. Bracts absent. Bracteoles (at anthesis) with petioles 0.6–0.9 mm long, blades 3.0–3.9 mm long, 0.3–0.5 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) 0.3–0.6 mm long. Hypanthia (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, 0.4–0.7 mm wide at the base, subulate, 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 acuminate, margin entire and glabrous, adaxial surface glabrous or sparsely glandular-punctate, abaxial surface glabrous. Stamens 10, strongly dimorphic. Larger (antesepalous) stamens 5, filaments 2.6–2.9 mm long, pink, pedoconnectives 2.9–3.2 mm 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 0.2–0.4 mm long, the circular pores ca. 0.2 mm wide. Smaller (antepetalous) stamens 5, filaments 1.9–2.1 mm long, pink, pedoconnectives 0.6–0.9 mm long, yellow, inconspicuous appendages ca. 0.1 mm long, yellow, apex emarginate, thecae (excluding rostra) 1.4–1.6 mm 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 hypanthium, 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 0.7–0.8 mm long, reniform.
Microlicia pithyoides A habit B detail of nodes and internodes C leaf abaxial surface D bracteole abaxial surface E floral bud F flower 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 Pacifico 295 (CAS).
Brazil. Minas Gerais: Catas Altas Municipality, Serra do Caraça, Oliveira 382 (BHCB, RB), Pacifico 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]).
Known only from Minas Gerais State (Fig.
Microlicia pithyoides species is the least collected species of the Trembleya s.s. clade and apparently has the narrowest distribution. Less than 10 collections 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 available for M. pithyoides refer to the same population, we were unable to calculate its EOO. The AOO is 4 km2. The only population known of this species occurs inside a private protected area, the RPPN Serra do Caraça (Natural Heritage Private Reserve). The type material was probably collected at the Serra de Capanema in Santa Bárbara 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 vegetation of Serra do Caraça is largely intact and affords a good measure of protection for the populations of M. pithyoides. This species is considered Critically Endangered by the Brazilian Government (
Microlicia pithyoides can be recognised by its narrow leaves that are 0.5–1.4 mm wide, 1-nerved from the base, inflorescences reduced to solitary flowers, 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 flowers with subulate calyx lobes and the overall shape of its stamens. Microlicia pithyoides differs 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 flowers (sometimes simple dichasia only in M. calycina), magenta petals and stamens with bicoloured anthers. Microlicia pithyoides may be differentiated 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).
Recent collections of M. pithyoides (Oliveira 382, Pacifico 295) have leaves with blades that are 8–12 mm long. These measurements bridge those given in the protologues of varieties pithyoides (8–10 mm long) and major (12–20 mm long). We consider these size differences to represent a continuum and here relegate var. major to synonymy.
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]!).
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 narrow wings 0.2–0.3 mm wide. Petioles 0.5–1.2 mm long. Leaf blades 4–10 mm 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 flat, entire and glandular-punctate, 1-nerved from the base, tertiaries little evident on the abaxial surface, adaxial surface glandular-punctate, abaxial surface glandular-punctate. Inflorescences reduced to solitary flowers on apical branches. Bracts absent. Bracteoles (at anthesis) with petioles 0.5–1.0 mm long, blades 2.5–3.3 mm 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) 2.2–3.5 mm long, 1.5–2.0 mm wide at the torus, campanulate to urceolate, light green (when fresh), externally glandular-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, filaments 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, yellow to orange, oblong, rostra 0.2–0.3 mm long, the circular pores ca. 0.2 mm wide. Smaller (antepetalous) stamens 5, filaments 1.5–2.0 mm long, yellow, pedoconnectives 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 0.3–0.6 mm long, fruiting calyx lobes 2.5–3.2 mm long, not thickened. Seeds 0.5–0.9 mm long, reniform.
Microlicia rosmarinoides A habit B detail of nodes and internodes C leaf abaxial surface D bracteole abaxial surface E floral bud F flower 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]).
Restricted to central Minas Gerais State (Fig.
Microlicia rosmarinoides is a little-collected species, known from about 15 specimens. The 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
Microlicia rosmarinoides can be readily recognised amongst congeners by its narrow leaves (1–1.9 mm wide) and flowers with yellow petals. The 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. flaviflora. For comparisons, see notes under M. flaviflora and M. pithyoides.
Brazil. Minas Gerais: Barão de Cocais, Mina do Baú, Souza et al. 2584 (BHCB); Belo Vale, Occhioni et al. s.n. (RFA, US [US001900109]); Itabirito, Serra de Capanema, Carmo 192 (BHCB), Carmo 377 (BHCB); Ouro Preto municipality, 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 Municipality, 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).
Brazil. “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, first-step designated by
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 flat, slighly serrulate and ciliate with eglandular trichomes 0.1–0.4 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, 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. Inflorescences reduced to solitary flowers 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-punctate 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, narrowly 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. Larger (antesepalous) stamens 5, filaments 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 circular pores ca. 0.2 mm wide. Smaller (antepetalous) stamens 5, filaments 1.5–2.0 mm 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.
Microlicia trembleyiformis A habit B detail of nodes and internodes C leaf abaxial surface D leaf adaxial surface E bracteole abaxial surface F floral bud G flower in lateral view H petal adaxial surface I antesepalous stamen J antepetalous stamen K gynoecium L capsule enveloped by the hypanthium. Drawn from Duarte 2767 (US).
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.
This species is known from about 10 specimens. It is a little-collected species that, however, has a comparatively wide distributional range. The EOO is 92,214.048 km2, a value that would indicate a Least Concern conservation status if criterion B of
Microlicia trembleyiformis can be recognised by its elliptic to ovate leaves that are 5-nerved from the base and solitary flowers with narrowly-triangular calyx lobes 1.5–2.1 mm long. In morphology, it is closest to M. parviflora and more distantly to M. altoparaisensis (see notes under these species for comparisons). Microlicia pentagona is the only congener that shares with M. trembleyiformis both the leaves 5-nerved from the base and solitary flowers. Microlicia trembleyiformis differs 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 species, only M. parviflora occurs in Ouro Preto, Serra da Canastra and Serra Negra, where sympatry with M. trembleyiformis is possible.
According to
Brazil. Minas Gerais: Capitólio Municipality, estrada para Cachoeira Fecho da Serra, Romero et al. 7550 (HUFU, US); Ouro Preto Municipality, [catal. B1, n° 160] Saint-Hilaire s.n. (lectotype: P [P002297746]!; isolectotype: 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 municipality in Minas Gerais State, Glaziou 19221 (K, P). Rio de Janeiro: Itatiaia Municipality, Serra Negra, Porto 2834 (NY, RB, US).
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, first-step designated by
Erect shrubs or treelets 1.0–1.8 m tall. Branchlets quadrangular, appearing 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 cuneate or attenuate, apex rounded to emarginate, margin flat, entire along the basal half, serrulate on the upper half and glandular-punctate, 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, randomly branching and surrounding stout depressions on the abaxial leaf surface, adaxial surface glandular-punctate, abaxial surface glandular-punctate. Inflorescences simple dichasia or reduced to solitary flowers, 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 serrulate, 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 appearing glabrous, vernicose and minutely granulose. Calyx tubes 0.3–0.4 mm long. Calyx lobes (at anthesis) 4.1–4.9 mm 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 11.5–13.0 mm long, 6.0–7.0 mm wide, magenta or rarely white, obovate, apex acute or rounded, margin entire and glabrous, both surfaces glabrous. Stamens 10, strongly dimorphic. Larger (antesepalous) stamens 5, filaments 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 flushed 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, filaments 4.0–4.4 mm long, pink or rarely white, pedoconnectives 1.5–1.9 mm long, pink or rarely white with the apical region flushed with pink, inconspicuous appendages ca. 0.2 mm long, yellow, 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 2.2–2.4 mm 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 0.4–0.5 mm long, fruiting calyx lobes 4.5–5.4 mm long, not thickened. Seeds not seen.
Microlicia tridentata A habit B leaf abaxial surface C bracteole abaxial surface D floral bud E flower in lateral view F flowering hypanthium G petal adaxial surface H antesepalous (behind) and antepetalous (in front) stamens I gynoecium J capsule enveloped by the hypanthium. Drawn from Hensold et al. CFCR2880 (UEC) and Souza et al. 8015 (UEC).
Microlicia tridentata is endemic to Minas Gerais State (Fig.
Microlicia tridentata is known from about 40 collections. The EOO is 4,613.081 km2 and the AOO is 24 km2. Following
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 flowers 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 differentiated 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. parviflora, from which M. tridentata differs by the leaves that are not pruinose (vs. usually pruinose), inflorescences composed of simple dichasia or reduced to solitary flowers (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). The distributions of M. parviflora 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).
Major variation in M. tridentata involves inflorescence development (simple dichasia or solitary flowers) and the petals typically magenta (Fig.
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 Municipality, Serra de Ouro Branco, Pacifico & 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), Michelangeli 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, Pacifico & Bressan 303 (CAS, HUEM), Versiane & Devides 682 (UEC); Santa Bárbara Municipality, Serra do Caraça, Ordones 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 Municipality, 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, Claussen 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).
1. Trembleya acuminata R.B.Pacifico & 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.Pacifico & 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.
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 September 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, first-step designated by
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. (lectotype, first-step designated by
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, designated here: BR [BR0000005520855]!; isolectotype: BR [BR0000005520190]!).
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 image]!, 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/catalognumber/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 (lectotype, 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, designated 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 flumen 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!).
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 December 2003, K. Fidanza 108 (holotype: UEC!; isotype: CAS!).
Replaced with: Microlicia serratifolia Versiane & R.Romero.
12. Trembleya thomazii R.B.Pacifico & 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.Pacifico & 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.Pacifico.
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 Griff. [probably = Melastoma L.]
6. Trembleya rosea Sessé & Moc. [probably = Coreopsis grandiflora Hogg ex Sweet (Asteraceae)]
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 locating the type of Meriania parviflora; 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 fieldwork activities; André V. Scatigna, Fabian A. Michelangeli, Fernando A.O. Silveira, Luciano Pedrosa, Orlando Graeff, Rodrigo Penati, and Vania E. Bressan for allowing us to use their photos of living specimens; Renato Goldenberg, an anonymous reviewer and the handling editor (Marcelo Reginato) for their comments and suggestions on an earlier version of this monograph. RP thanks the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for a Ph.D. degree grant and the M. Stanley Rundel 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. This research was supported in part by U.S. National Science Foundation grant DEB-1146409.
List of species
1. Microlicia altoparaisensis (R.B.Pacifico, Almeda & Fidanza) Versiane & R.Romero
2. Microlicia calycina (Cham.) Versiane & R.Romero
3. Microlicia chamissoana (Naudin) Versiane & R.Romero
4. Microlicia flaviflora Versiane & R.Romero
5. Microlicia laniflora (D.Don) Baill.
6. Microlicia parviflora (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). Andrade 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. laniflora × M. pentagona); 706 (5); 9019 (2). Barreto & Viégas s.n. (5). Barros & 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). Cavalcanti 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). Delfini 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 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). Ferreira 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). Kameyama 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). Leoni 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). Macedo 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); 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). Pacifico 185 (5); 191 (6); 295 (8); 413 (6); 565 (4). Pacifico & Bressan 290 (11); 291 (2); 294 (7); 296 (2); 303 (11); 380 (1); 531 (7); 565 (4). Pacifico & Carmo 154 (3). Pacifico & 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). Santos 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). Versiane 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).
List of species sampled in this study, their respective Genbank accession codes (for nrITS and nrETS), and specimen vouchers
Data type: table (word document)
Explanation note: List of species sampled in this study, their respective Genbank accession codes (for nrITS and nrETS), and specimen vouchers. Codes highlighted in bold refer to sequences previously obtained from GenBank.