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Species of Cambessedesieae. A, Huberia organensis. B, Huberia magdalenensis. C, Huberia insignis. D,
Huberia comosa. E, Huberia kollmannii. F, Huberia kollmannii. G, Huberia bradeana. H, Huberia bradeana. I, Huberia
espiritosantensis. J, Huberia espiritosantensis. K, Huberia sessilis. L, Huberia semiserrata. M, Merianthera burlemarxii.
N, Merianthera bullata. O, Merianthera bullata. P, Merianthera bullata. Q, Cambessedesia eichleri. R, Cambessedesia
membranacea S, Cambessedesia purpurata. T, Cambessedesia regnelliana. Photographs: A, B, I, P, Q, Thuane Bochorny; C,
E, F, G, H, J, K, L, N, O, S, Renato Goldenberg; D, André Fontana; M, Paulo Labiak; R, T, João M. do Carmo.

Species of Cambessedesieae. A, Huberia organensis. B, Huberia magdalenensis. C, Huberia insignis. D, Huberia comosa. E, Huberia kollmannii. F, Huberia kollmannii. G, Huberia bradeana. H, Huberia bradeana. I, Huberia espiritosantensis. J, Huberia espiritosantensis. K, Huberia sessilis. L, Huberia semiserrata. M, Merianthera burlemarxii. N, Merianthera bullata. O, Merianthera bullata. P, Merianthera bullata. Q, Cambessedesia eichleri. R, Cambessedesia membranacea S, Cambessedesia purpurata. T, Cambessedesia regnelliana. Photographs: A, B, I, P, Q, Thuane Bochorny; C, E, F, G, H, J, K, L, N, O, S, Renato Goldenberg; D, André Fontana; M, Paulo Labiak; R, T, João M. do Carmo.

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Systematic studies based on DNA sequences have shown that some traditional tribal delimitations in Melastomataceae remain unresolved, such as the ‘Merianthera and allies’ clade, an informal group which has not been formally assigned to a tribe. This clade includes Behuria, Cambessedesia, Dolichoura, Huberia and Merianthera and occurs mainly at high...

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Microlicieae are a monophyletic tribe comprising seven genera: Chaetostoma, Lavoisiera, Microlicia s.s., Poteranthera, Rhynchanthera, Stenodon and Trembleya. Microlicia s.s. includes 172 species predominantly distributed in the campo rupestre of Brazil. Its delimitation is complex because the generic boundaries, mostly with Lavoisiera and Trembleya...

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... & R.Goldenb. (Bochorny et al. 2019), is now assigned to Pyramieae Naudin, a tribal name with priority (Bochorny et al. 2022). Its four genera (Bisglaziovia Cogn., Cambessedesia DC., Huberia DC., and Merianthera Kuhlm.) with a total of 68 species are almost restricted to eastern Brazil, except for four species of Huberia occurring in the Peruvian and Ecuadorian Andes (Bochorny et al. 2022). ...
... All other species in the genus have either an open calyx dehiscing through regular lobes (Merianthera bullata, M. burlemarxii, M. eburnea, M. sipolisii, M. verrucosa) or the calyx is closed in bud and dehisces through irregular lobes in M. parvifolia and M. pulchra, but never through a circumscissile slit that releases a single calyptra. While absent from Merianthera and other genera in the tribe Pyramieae (Bochorny et al. 2019), this dehiscence mode can be found in at least five different tribes of Melastomataceae: Cyphostyleae Gleason, Henrietteeae, Merianieae Triana, Pyxidantheae Griseb., and in at least four unrelated clades of Miconieae DC. (Michelangeli et al.2011;Judd et al. 2022). ...
... In all other species of Merianthera, the fruits apparently lack a dehiscence region, and "the seeds are dispersed after the hypanthium and ovary walls decay and open ("cápsula rompente" or "rupturing capsule" sensu Baumgratz 1985), and the seeds are released through the gaps left by the falling irregular hypanthium pieces" (Goldenberg et al. 2012). The same mechanism is found in some species of Huberia (also in the tribe Pyramieae; Bochorny et al. 2019Bochorny et al. , 2022, and in Allomaieta Gleason and Alloneuron Pilger (both in the tribe Cyphostyleae; Michelangeli et al. 2011). All these species in Merianthera, Huberia, Allomaieta and Alloneuron have an inferior ovary, and this dehiscence mechanism has been interpreted as a consequence of the ovary position (Goldenberg et al. 2012). ...
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Merianthera is a genus of flowering plants with up to now seven species occurring in eastern Brazil, in the states of Bahia, Espírito Santo, and Minas Gerais. It belongs to the tribe Pyramieae (Melastomataceae), and can be recognized by its shrubby or treelet habit with caducous leaves, 5-merous flowers with a strongly zygomorphic androecium, the latter with dimorphic stamens bearing complex dorsal connective appendages, as well as an inferior ovary and capsular fruits. We describe here a new species of Merianthera from two collections and a photographic record, all from the same locality, an inselberg in the Municipality of Jacinto, in northeastern Minas Gerais. Its candelabriform habit, with a fistulose stem, and solitary, axillary or cauliflorous flowers are only shared with M. burlemarxii. However, M. calyptrata R.Goldenb., Bochorny & Fraga sp. nov. has at least three characters that are absent in M. burlemarxii and all other species in the genus: the total absence of both a peduncle and bracteoles, the calyptrate calyx and the fruits developing from inferior ovaries and dehiscing through longitudinal slits. The first character appears to be unknown in other species in the family.
... The only succulent species is M. burlemarxii (Ziemmer & al. 2017). The genus was traditionally placed in tribe Merianieae (based on similarity in anther architecture), but in the molecular phylogeny of Goldenberg & al. (2012), Merianthera appears in the Cambessedia-Clade (formally published as tribe Cambessedesieae by Bochorny & al. 2019), but correctly Pyramieae for priority reasons, see Bochorny & al. (2022) and is placed as sister to the newly circumscribed Huberia De Candolle with 37 species (Bochorny & al. 2019)all of which are endemic to E Brazil except for 4 species from the Andes of Ecuador and Peru. U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: Urs.Eggli@zuerich.ch ...
... The only succulent species is M. burlemarxii (Ziemmer & al. 2017). The genus was traditionally placed in tribe Merianieae (based on similarity in anther architecture), but in the molecular phylogeny of Goldenberg & al. (2012), Merianthera appears in the Cambessedia-Clade (formally published as tribe Cambessedesieae by Bochorny & al. 2019), but correctly Pyramieae for priority reasons, see Bochorny & al. (2022) and is placed as sister to the newly circumscribed Huberia De Candolle with 37 species (Bochorny & al. 2019)all of which are endemic to E Brazil except for 4 species from the Andes of Ecuador and Peru. U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: Urs.Eggli@zuerich.ch ...
... & Amorim (Goldenberg and Amorim 2006), Brasilianthus Almeda & Michelang. (Almeda et al. 2016) and others were relegated to synonymy (for example, see Judd 2013 andBochorny et al. 2019); and there are still unplaced clades and genera that need to be tested for monophyly . Meanwhile, many new species are still being described from little-studied genera and/or resulting from fieldwork in areas that remain poorly known or underexplored botanically. ...
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A systematic monograph of the Trembleya s.s. clade is presented, a Brazilian endemic lineage of Melas-tomataceae comprising 11 species and currently recognised as part of Microlicia s.l. (Melastomataceae). First, we investigate phylogenetic relationships within Lavoisiereae using two nuclear markers and two sampling datasets (102 and 134 terminals). 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.
... & Amorim (Goldenberg and Amorim 2006), Brasilianthus Almeda & Michelang. (Almeda et al. 2016) and others were relegated to synonymy (for example, see Judd 2013 andBochorny et al. 2019); and there are still unplaced clades and genera that need to be tested for monophyly . Meanwhile, many new species are still being described from little-studied genera and/or resulting from fieldwork in areas that remain poorly known or underexplored botanically. ...
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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.
... Some studies in the Atlantic Forest also recovered conserved climatic niches for lineages as Leandra Raddi s.s. (Reginato, 2014), Cambessedesieae (Bochorny et al., 2019) and Bertolonia . Some plant lineages (mainly herbaceous) can show these same patterns in locations distant from each other, for example, regions of the Andes (Tovar et al., 2020) and southeastern Australia (Morgan and Venn, 2017). ...
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The asymmetric pattern in species richness is a notable feature across different lineages and geographic regions. While some lineages have high richness, diversity and wide distribution, others have the opposite. Despite low rates of diversification, the latter might also be phylogenetically isolated. Lineages that accumulate these characteristics are known as "depauperons'' and explaining their existence and persistence through time is still a challenge. The plant family Melastomataceae contains both megadiverse lineages (such as the tribe Miconieae, with around 1900 species) and groups with few species (such as the tribes Eriocnemeae, Lithobieae, and Rupestreeae with 7, 1 and 2 species, respectively). These three clades are restricted to eastern Brazil, where they have been seldom studied. The lack of information about their basic biology as well as which processes determine their distribution have not been previously studied. Here we integrated metrics of dispersal ability, species distribution models (SDMs) and natural history data compilation in order to uncover common patterns shared by these depauperons in Melastomataceae and raise conservation concerns. For all nine species we estimated the dispersal ability and generated SDMs in different time-periods (past, present and future). Dispersal ability was associated with predicted distribution models under future scenarios to evaluate shifts and/or retractions in suitable areas. In addition, we compared the climatic tolerances of the depauperons with their megadiverse sister tribes via climatic envelopes. Overall, our results indicate limited dispersal ability, dependency on water for dispersal, and restricted niche as common characteristics for all species in the deupauperon tribes Eriocnemeae, Lithobieae and Rupestreeae. Our analyses also show that the climatic niche spaces of the depauperons are limited and totally included within the niche space of its sister tribes. Based on our findings, the level of threat in these groups can be potentiated by rapid climate change, mainly due to their inability to spread over long distances, restricted niches and increased habitat fragmentation. We suggest that future conservational actions prioritize these unique taxa in Melastomataceae, especially if a phylogenetic diversity perspective is taken into account.
... 3a-c). In addition, closely related groups may exhibit contrasting character states(Penneys et al., chapter "A New Melastomataceae Classification Informed by Molecular Phylogenetics and Morphology"), even within tribes such as Eriocnemeae (Eriocnema-superior, Physeterostemon-inferior,Penneys et al. 2020) and Pyramieae (Cambessedesiasuperior, Huberia and Merianthera-inferior,Bochorny et al. 2019). The range of variation in ovary position can be larger or smaller according to the clade considered (see ...
Chapter
The floral diversity of Melastomataceae is stunning and clearly expressed in sepal and stamen structure and in the position of the ovary. Comparative developmental studies are effective in order to understand these variations because they reveal the often-enigmatic origin of the structures. The diverse calyx structure originates from variations in the degree of union between the sepals. The contort corolla aestivation, widespread in the family, is influenced by the floral architecture. Stamen size and shape depend on the space available in the floral bud after the growth of the perigynous hypanthium that may cause the delay in stamen emergence and flexion. Dimorphic stamens originate from differences in their developmental time and position. Prolonged connectives and most of their appendages are formed late during floral development. Ontogeny also explains the decrease or increase in organ number. The intercalary meristems can promote the formation of a hypanthium associated with the gynoecium, and their extension is responsible for the gradual variation in ovary position. These meristems also act on the development of a perigynous hypanthium. Thus, intercalary meristems play an important role for floral diversification in Melastomataceae. The potential of comparative floral development is wide and is illustrated here through several examples in this family.
... Stamen characteristics have always drawn attention in the family and served as an important background for traditional taxonomic studies (Cogniaux 1891;Jacques-Félix 1981, 1995Triana 1872). Significant progress in the systematics of Melastomataceae (see Michelangeli et al. 2020) has revealed that some groups recognized in these traditional studies are not monophyletic, in addition to disclosing the emergence of "unexpected" clades and the proposal of new tribes (e.g., Bacci et al. 2019;Bochorny et al. 2019;Penneys et al. 2010Penneys et al. , 2020Rocha et al. 2016;Veranso-Libalah et al. 2017). This suggests that the stamen morphology of Melastomataceae is subject to a high degree of homoplasy (Gavrutenko et al. 2020;Goldenberg et al. 2008;Rocha et al. 2016). ...
... Inferior ovary species tend to have fewer stamen elaborations. Moreover, in species with inferior to semi-inferior ovaries, when the stamen shows a thickened connective or appendages, these tend to be dorsal, as is the case, e.g., of Henrietteeae (Bellucia; Penneys et al. 2010), Pyramieae (Huberia, Merianthera; Bochorny et al. 2019) and Pyxidantheae (Blakea, Chalybea; Penneys and Judd 2011). One hypothesis, not yet tested, is that in Melastomataceae the flowers with superior ovaries have a proportionately longer perigynous hypanthium than those of the inferior ovary. ...
Article
The androecium of Melastomataceae presents notable modifications in its merosity, morphology between whorls and in prolonged connectives and appendages. We carried out a comparative study of six Melastomataceae species to shed light on the developmental processes that originate such stamen diversity. The development of stamens was studied using scanning electron microscopy and histological observations. The stamens of all species studied have a curved shape because they emerge on a plane displaced by the perigynous hypanthium. They are the last flower organs to initiate and therefore their growth is inwards and towards the floral center. Despite the temporal inversion between carpels and stamens in Melastomataceae, the androecium maintains the centripetal pattern of development, the antepetalous stamens emerging after antesepalous stamens. The isomerous androecium can be the result of abortion of the antepetalous stamens, whereas heterostemony seems to be caused by differences in position and the stamen development time. Pedoconnectives and ventral appendages originate from the basal expansion of the anther late in floral development. The delay in stamen development may be a consequence of their dependence on the formation of a previous space so that they can grow. Most of the stamen diversity is explained by the formation of the connectives and their appendages. The formation of a basal-ventral anther prolongation, which culminates in the development of the pedoconnective, does not differ from other types of sectorial growth of the connective, which form shorter structures.
... Seed appendages are rare in Melastomataceae, but several species of Miconia (formerly in Leandra section Tshudya) have short appendages (Martin et al. 2008;Martin and Michelangeli 2009). Otherwise, lateral wings are present in some species of Huberia, and most prominently in Acanthella (Whiffin and Tomb 1972;Baumgratz 1988Baumgratz , 2004Bochorny et al. 2019). ...
Chapter
The Cyphostyleae are an exclusively neotropical and small tribe, yet poorly studied and collected. It includes 25 species in four genera from low to mid-elevation in the Andes of Colombia, Ecuador, and Peru. Even though the group is not species-rich, some or all of its species present several characters otherwise rare in the family: leaves with pinnate venation, haplostemonous flowers, calyptrate calyces, incurved style, bi-colored petals, capsular fruits derived from inferior ovaries, and seeds with long appendages. We review the history of the group, its systematics, and taxonomy, and discuss the prevalence of these unusual characters across the family.
... Of the 544 species of the family that occur in the domain, 386 are endemic (Goldenberg et al. 2020a) and vary widely in overall morphology (i.e., lineages with both capsular and berry fruits, Reginato et al. 2020) and habitats, or are restricted to specific AF regions. For example, Physeterostemon is distributed only in the Bahian portion of the domain, or Huberia, usually occurs on mountaintops in southern AF (Bochorny et al. 2019;Goldenberg and Amorim 2006). There are also widespread groups throughout the AF and its different physiognomies, such as the Leandra clade and Pleroma. ...
Chapter
Melastomataceae are the fifth richest plant family in the Atlantic Forest with 544 species belonging to few phylogenetic lineages and present considerable variation in habitat, ecological and reproductive characteristics, and morphology. Some of these lineages are prevailingly centered in the Atlantic Forest, being either endemic (Bertolonieae, Eriocnema, Physeterostemon) and/or highly diverse [Leandra clade (Miconia s.l.), Chaenanthera clade (Miconia s.l.), Discolores clade (Miconia s.l.), Pleroma and Pyramieae] in this biome. In this chapter, we summarize the diversification of Melastomataceae in the Atlantic Forest. Based on the phylogenetic hypothesis generated for the family by Penneys et al. (chapter “A New Melastomataceae Classification Informed by Molecular Phylogenetics and Morphology”), we recovered at least 22 different Melastomataceae lineages currently distributed in the Atlantic Forest, nine of which are highly diverse in the domain. The colonization and diversification events that led to the current distribution of the family in the Atlantic Forest would have occurred during three distinct periods (Miocene, Pliocene and Pleistocene). Moreover, diversification in the AF is heterogeneous, meaning that some AF-centered lineages show higher or lower diversification rates when compared with those outside AF. Future studies on a smaller scale, taking into account putative effects of traits or more specific habitats and regions within and surrounding the AF, should be carried out to explore this diversification heterogeneity.
... Species in the genus Huberia DC. (including Behuria Cham. and Dolichoura Brade) have anthers with dorsal connective appendages that are simple, descending and vary from straight to coiled in shape (Goldenberg et al., 2012;Bochorny et al., 2019). In most species they are short and have the same yellow colour as the thecae (see Baumgratz, 1997;Tavares, 2005;Goldenberg et al., 2016). ...
... Currently the suggestion that stamen appendages are a primitive character in Melastomataceae is not considered because it contradicts the distribution of the character amongst taxa in relation to their phylogenetic relationships (Clausing and Renner, 2001). Indeed, stamen appendages in Melastomataceae have appeared in different tribes and are variable in position, shape, colour, and vascularization (Renner, 1993;Clausing and Renner, 2001;Goldenberg et al., 2012;Michelangeli et al., 2013;Bochorny et al., 2019). This raises the hypothesis that these appendages may have evolved several times in the family. ...
... Stamen appendages in Huberia were present ancestrally, probably simple and linear, as in most species of the genus, and probably evolved into long and coiled structures in only two species (i.e., H. espiritosantensis and H. bradeana; Bochorny et al., 2019). Our data suggest that the small and inconspicuous stamen appendages in H. insignis are probably vestigial and play no role in the pollination system of these species. ...
Article
In most species of Melastomataceae pollen is the only reward for pollinators. Pollen is locked within poricidal anthers and is only legitimately removed by bees that specifically buzz the anthers. Stamen appendages in many species of plants are morphological adaptations of the androecium for pollination, and are evident in several species of Melastomataceae. In an experimental study, anthers with large, coiled appendages released more pollen during artificial buzzing in flowers of the genus Huberia than anthers without appendages. Other aspects of reproductive biology and pollination ecology in Huberia are poorly known, including the relationship between size and shape of widely varying stamen appendages with pollen release. In the Brazilian Atlantic Forest, we studied the breeding system and pollinator assemblage of the endemic Huberia insignis, and we experimentally tested how stamen appendages are associated with pollination. With hand-pollination experiments, morphoanatomical analysis, observations of the pollinator assemblage, using flowers with and without stamen appendages, we tested pollinator preferences and the effect of appendage presence on pollen removal rates. Huberia insignis is self-compatible and requires pollinators, and bees that buzz-pollinate are the only pollinators, typically two species of Bombus (ca. 80% of visits). The presence of stamen appendages of H. insignis had no effects on attracting pollinators and pollen removal. Anthers and stamen appendages have neither secretory structures nor specialized biomechanical tissues. We conclude that stamen appendages play no role in the pollination of H. insignis. We suggest that these typically small and inconspicuous stamen appendages are vestigial.