Alcantarea habitat and floral morphology, showing position of stamens. (a and b) Gneiss-granitic inselbergs inside the Atlantic rainforest domain. (a) Alcantarea geniculata (b) A. patriae (c and d) Quartzite rocky outcrops in rocky savanna-like vegetation along the Espinhaço range. (c) A. turgida. (d) A. duarteana. (e-i) Morphology of the stamens. (e and f) Spreading. (g and i) Presented in a bundle. (e) A. burle-marxii. (f) A. imperialis. (g) A. patriae. (h) A. heloisae. (i) A. tortuosa.

Alcantarea habitat and floral morphology, showing position of stamens. (a and b) Gneiss-granitic inselbergs inside the Atlantic rainforest domain. (a) Alcantarea geniculata (b) A. patriae (c and d) Quartzite rocky outcrops in rocky savanna-like vegetation along the Espinhaço range. (c) A. turgida. (d) A. duarteana. (e-i) Morphology of the stamens. (e and f) Spreading. (g and i) Presented in a bundle. (e) A. burle-marxii. (f) A. imperialis. (g) A. patriae. (h) A. heloisae. (i) A. tortuosa.

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The genus Alcantarea comprises near 30 species endemic to rocky outcrops from eastern Brazil. Most species are ornamental and several are threatened due to habitat loss and over collection. In this paper we examine the phylogenetics of Alcantarea and its relationship with the Brazilian members of Vriesea, a genus of which Alcantarea has been treate...

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... includes ornamental bromeliads characterized by their large size (sometimes reaching up to 5 m high when flowering, Fig. 1) and by well-developed tanks (phytotelma) and they are popular garden plants. A few species show a much-reduced habit and almost lack the ability to impound any water ( Benzing et al., 2000). Adaptation to life on inselbergs is a key feature of Alcantarea species (Martinelli, 1994). These adaptations include several anatomical ...
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... While many Alcantarea species present bat pollination syndrome characteristics (e.g. spreading stamens, nocturnal anthesis, abundant nectar production, cryptic coloration of the petals, Martinelli, 1994Martinelli, , 1997), other species present crepuscular or diurnal anthesis, smaller and bright yellow 2 perfumed flowers with bundled stamens (Fig. 1) and less developed septal nectaries that are suggestive of insect (sphingophily) or hummingbird pollination (Versieux, 2009;Vogel, 1969). Thus, the existence of a phylogenetic framework would be interesting to understand the evolution of these distinct floral morphologies and relate it to shifts in pollinators. Additionally, the ...
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... (Versieux, 2009;Vogel, 1969). Thus, the existence of a phylogenetic framework would be interesting to understand the evolution of these distinct floral morphologies and relate it to shifts in pollinators. Additionally, the species that grow on the quartzite stony grassland (campo rupestre) in the Espinhaço range of Bahia and Minas Gerais states (Fig. 1) have been hypothesized as derived from Atlantic forest mesic ancestors ( Benzing et al., 2000, p. 498) or, alternatively, have been proposed as the early divergent lineages that retained a relict inland distribution associated with other plesiomorphic character states, such as the rudimentary leaf rosettes (Leme, 1995, p. 23). ...
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... associated with other plesiomorphic character states, such as the rudimentary leaf rosettes (Leme, 1995, p. 23). Investigating these hypotheses related to the disjunctive occurrence between the Atlantic forest inselbergs versus the quartizitic outcrops of the savanna like habitats, as well as the morphological changes in Alcantarea flowers (Fig. 1), would be extremely useful as a baseline to address future questions and add evidences to discuss broader issues involving the evolution of eastern Brazil plants. Very little information is available in the literature regarding the appearance of the Espinhaço savanna-like habitats flora and their endemic plant species (e.g. Antonelli ...

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... pacifica, and considering that in angiosperms the inheritance of nDNA is biparental, whereas the inheritance of cpDNA is uniparental, mostly maternal, and highly conserved due to its slow molecular evolution and low mutation rates (Small et al. 2004, Robbins & Kelly 2023, it can be hypothesized that the origin of this taxon might be due to potential events of ancestral hybridization, introgression and/or incomplete lineage sorting (Barber et al. 2007, Kim & Donoghue 2008, Jabaily & Systma 2010. Incongruences between nDNA and cpDNA evidence has been constantly observed in Bromeliaceae, suggesting that phenomena such as hybridization, introgression and incomplete lineage sorting have been of importance to the diversification of bromeliads and are present in several groups within the family (e.g., Jabaily & Sytsma 2010, Schulte et al. 2010, Palma-Silva et al. 2011, Versieux et al. 2012, Krapp et al. 2014, Goetze et al. 2017, Mota et al. 2019, Ancona et al. 2022, being observed mostly in taxa resulting from a recent adaptative radiation in which the lineages experiment a process of diversification into various ecological niches in a short time period (Goetze et al. 2017). However, to determine whether these events influenced the evolutionary history of A. bracteata var. ...
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Background: Previous phylogenetic analyses suggested that Aechmea bracteata is not a monophyletic species, being each variety an individual lineage within the Aechmea bracteata complex. Hypothesis: A phylogenetic analysis based on molecular data and a morphological analysis will provide evidence to support the taxonomic recognition of A. bracteata var. pacifica as a distinct species. Studies species: A. bracteata var. bracteata, A. bracteata var. pacifica and related species. Study site and dates: Mexico to Northern South America; 17 populations from Southeastern and Western Mexico. Analyses were performed between 2021 and 2023. Methods: A linear morphometric analysis was conducted with 85 herbarium specimens testing 28 quantitative and three qualitative variables. Potential diagnostic traits were reviewed in living specimens. A phylogenetic analysis was performed with two nDNA markers (ETS and g3pdh) and one cpDNA marker (trnL-F). A total of 373 records were projected into biogeographical provinces of the Neotropics. Results: The morphometric analysis allowed to separate each variety into a group, also, nine of the evaluated traits resulted statistically significant through a univariate analysis. Five additional diagnostic traits from the inflorescence and leaves were recognized. According to nDNA both taxa present reciprocal monophyly, however, cpDNA groups A. bracteata var. pacifica with Central American species, showcasing a potential hybrid origin. Geographically, the Sierra Madre del Sur acts as a barrier between both taxa. Conclusions: Due to the gathered evidence is possible to recognize A. bracteata var. pacifica as an endemic species to Mexico, reestablishing the basionym A. laxiflora.
... & Schult.f., and Brocchinia Schult.f. (Bennett, 1991;Givnish et al., 1997;Barbará et al., 2009;Versieux et al., 2012;Hmeljevski et al., 2014). Semelparity is not ubiquitous in sky island Puya, with sympatric multiramet iteroparous Puya found in páramos. ...
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Phylogenomics enhances our understanding of plant radiations in the biodiverse Andes. Our study focuses on Puya , primarily Andean and a part of the Bromeliaceae family. Using a phylogenomic framework based on the Angiosperms353 probe set for 80 species, we explored Puya ′s phenotypic evolution and biogeography. Divergence time analyses and ancestral area estimations suggested that Puya originated in Central Coastal Chile around 9 million years ago (Ma). Subsequently, it dispersed to the dry valleys of the Central Andes and Puna regions between 5–8 Ma, leading to the emergence of major lineages. Key events in the last 2–4 million years include the recolonization of Chilean lowlands and dispersal to the northern Andes via Peru's Jalcas, facilitating passage through the Huancabamba depression. This event gave rise to the high‐elevation Northern Andes clade. Using phylogenetic comparative methods, we tested the hypothesis that adaptation to the Andes' island‐like high‐elevation ecosystems was facilitated by unique leaf and floral traits, life history, and inflorescence morphology. Our findings suggest correlations between inflorescence axis compression, protective bract overlap, and high‐elevation living, potentially preventing reproductive structure freezing. Semelparity evolved exclusively at high elevations, although its precise adaptive value remains uncertain. Our framework offers insights into Andean evolution, highlighting that lineages adapted to life in dry ecosystems can easily transition to high‐elevation biomes. It also underscores how the island‐like nature of high‐elevation ecosystems influences phenotypic evolution rates. Moreover, it opens avenues to explore genetic mechanisms underlying adaptation to extreme mountain conditions.
... For this reason, recent research on the systematics of Tillandsioideae focused on resolving this issue either from a purely taxonomic approach (e.g. Bernal et al. 2015;Grant 1993Grant , 1995aKessous & Costa 2023) or a phylogenetic approach (Barfuss et al. 2005(Barfuss et al. , 2016Costa et al. 2014Costa et al. , 2015Gomes-da-Silva & Souza-Chies 2018;Machado et al. 2020;Versieux et al. 2012). Although these efforts provided clarity for Brazilian (e.g. ...
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A new species of Tillandsia from high elevation forests of the Eastern Cordillera of Colombia is described here: Tillandsia chingacensis. This species is unique because of its floral parts densely covered with a resin-like substance, and exceptionally long fruits. Although this species has distichous flowers and bears petal appendages, it was not placed in Vriesea because it shows morphological affinities to species of Tillandsia subg. Allardtia, suggesting a better placement of this taxon in Tillandsia. The discovery of this species is the result of recent explorations to a former war zone in the proximity of Bogotá (Colombia). This novelty demonstrates that further scientific expeditions to these areas recently opened for exploration will bring numerous taxonomic discoveries for Colombia. Finally, the description of this new species is complemented with notes on its geographical distribution, habitat, conservation status and a discussion of its taxonomic affinities.
... As recovered by Barfuss et al. (2016), tribes Catopsideae (Catopsis) and Glomeropitcairnieae (Glomeropitcairnia) conform the non-core Tillandsioideae clade, whereas the core Tillandsioideae clade is integrated by tribes Tillandsieae and Vrieseeae, and includes the genera Alcantarea (46 spp.), Barfussia (5 spp.), Cipuropsis (3 spp.), Goudaea (2 spp.), Gregbrownia (4 spp.), Guzmania (217 Barfuss et al. (2016), but formal taxonomic changes for these groups await for key taxa to be sampled (e.g., the Cipuropsis-Mezobromelia complex) or increased phylogenetic support to be achieved (e.g., Tillandsia biflora complex). Tillandsioideae phylogenetics, and in general in Bromeliaceae, has heavily relied on Sanger sequenced data, either from plastid, nuclear-ribosomal, or nuclear low copy loci that have been employed from deep to shallow taxonomic levels (Terry et al., 1997;Barfuss et al., 2005Barfuss et al., , 2016Granados Mendoza, 2008;Chew et al., 2010;Versieux et al., 2012;Donadío, 2013;Castello et al., 2016;Pinzón et al., 2016;Leme et al., 2017;Kessous et al., 2020;Donadío et al., in press). Recent studies have applied with great success Next Generation Sequencing (NGS) strategies to Tillandsioideae phylogenetics. ...
... Several other Sanger sequenced markers have been employed in previous molecular phylogenetic studies of the Tillandsioideae, including the ndhF and ycf 1 genes, the rpl32-trnL and trnK-rps16 IGS, the rps16 intron, and the trnK-matK-trnK, rpoB-trnC-petN, trnD-trnY-trnE-trnT, and trnH-rps19-psbA regions (Terry et al., 1997;Barfuss et al., 2005Barfuss et al., , 2016Granados Mendoza, 2008;Chew et al., 2010;Versieux et al., 2012;Donadío, 2013;Castello et al., 2016;Pinzón et al., 2016;Leme et al., 2017;Kessous et al., 2020;Donadío et al., in press). Among these markers, the ycf 1 gene far exceeds the PI of any other analyzed loci (Figure 2B and Supplementary Material 3). ...
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... In this line of thought, based on several reports of environmental effects on the composition of epicuticular waxes, (Hull et al., 1979;Shepherd et al., 1995;Medina et al., 2004;Domínguez et al., 2011;Ortúñez and Cano-Ruiz, 2013;Rajčević et al., 2014;Menzel et al., 2017;Xue et al., 2017;Sharma et al., 2018) as well as on the close link established between the characteristics of epicuticular waxes of many taxa and taxonomic, ecological or evolutive issues, (Li and Christophel, Frontiers in Plant Science | www.frontiersin.org 2000; Versieux et al., 2012;Li et al., 2016;Mitić et al., 2016;Harris et al., 2017;Klimko et al., 2018;Lindelof et al., 2020;Weber and Schwark, 2020;Faria et al., 2021) we propose in this work an unprecedented strategy to contribute to the study of vicariant phenomena in plant science. This new strategy involves a comprehensive physical-chemical analysis of the surface of plant leaves that strengthens the more usual phylogenetic, biogeographic and ecological approaches. ...
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Classically, vicariant phenomena have been essentially identified on the basis of biogeographical and ecological data. Here, we report unequivocal evidences that demonstrate that a physical–chemical characterization of the epicuticular waxes of the surface of plant leaves represents a very powerful strategy to get rich insight into vicariant events. We found vicariant similarity between Cercis siliquastrum L. (family Fabaceae, subfamily Cercidoideae) and Ceratonia siliqua L. (family Fabaceae, subfamily Caesalpinoideae). Both taxa converge in the Mediterranean basin (C. siliquastrum on the north and C. siliqua across the south), in similar habitats (sclerophyll communities of maquis) and climatic profiles. These species are the current representation of their subfamilies in the Mediterranean basin, where they overlap. Because of this biogeographic and ecological similarity, the environmental pattern of both taxa was found to be very significant. The physical–chemical analysis performed on the epicuticular waxes of C. siliquastrum and C. siliqua leaves provided relevant data that confirm the functional proximity between them. A striking resemblance was found in the epicuticular waxes of the abaxial surfaces of C. siliquastrum and C. siliqua leaves in terms of the dominant chemical compounds (1-triacontanol (C30) and 1-octacosanol (C28), respectively), morphology (intricate network of randomly organized nanometer-thick and micrometer-long plates), wettability (superhydrophobic character, with water contact angle values of 167.5 ± 0.5° and 162 ± 3°, respectively), and optical properties (in both species the light reflectance/absorptance of the abaxial surface is significantly higher/lower than that of the adaxial surface, but the overall trend in reflectance is qualitatively similar). These results enable us to include for the first time C. siliqua in the vicariant process exhibited by C. canadensis L., C. griffithii L., and C. siliquastrum.
... The floral morphology of Vrieseinae is highly diverse. The corolla shape is tubular, campanulate, or spiralescent and drooping; the stamens have different positions related to the throat of the corolla (included or exserted) and can form bundles resulting in different types of zygomorphy of the flower; the ovary is superior to half-inferior, and seven of the numerous stigma types known for bromeliads are present (Versieux et al. 2012;Costa et al. 2014; Barfuss et al. 2016). ...
... Besides their common occurrence in rocky environments, Stigmatodon share the batpollination syndrome with most species of V. sect. Xiphion, and with some species of Alcantarea (Versieux et al. 2012). Additionally, Stigmatodon and V. sect. ...
... Besides being currently used to recognize genera in the Tillandsioideae subfamily (Mez 1896;Smith and Downs 1977;Versieux et al. 2010Versieux et al. , 2012Gomes-da-Silva and Costa 2013;Costa et al. 2014;Barfuss et al. 2016;Leme et al. 2017a), such characters are especially important for delimitation of our study group, including the two anatomical ones (Couto 2017). The lifestyle (character i) is an important character used in the delimitation of the genera in Tillandsioideae (Barfuss et al. 2016), especially in Vrieseinae , and it is important for understanding the adaptive radiations in the different ecosystems (character ii) of the Brazilian shield by Vrieseinae genera. ...
Article
The genus Stigmatodon occurs in vertical and bare granite slopes, typical of the inselbergs of the Brazilian Atlantic Forest. Here, we present the first broad phylogenetic analysis focused on Stigmatodon , sampling a total of 83 terminals, including 16 of the 20 species of the genus and the morphologically similar species of Vriesea . We conducted a phylogenetic analysis using two plastid markers ( matK and rps16-trnK ) and the nuclear gene PHYC to infer phylogenetic relationships and reconstruct ancestral states for ecological and morphological characters. Our results suggest the monophyly of Stigmatodon as originally circumscribed is only possible with the inclusion of morphologically and ecologically similar Vriesea species. In addition, the morphological and anatomical traits led us to propose a new circumscription for the genus, combining eight species of Vriesea to Stigmatodon as S. andaraiensis , S. freicanecanus , S. lancifolius , S. limae , S. oliganthus , S. pseudoliganthus , S. vellozicolus , and S. zonatus . The stomata positioned above the ordinary epidermal cells, the adaxial water-storage parenchyma with axially elongated cells, the stamens positioned in two groups of three on each side of the corolla, and the tubo-laciniate stigma are exclusive to Stigmatodon in its new circumscription. These new morphological and phylogenetic results constitute a relevant contribution to the taxonomy and evolution of Bromeliaceae, one of the most diverse and ecologically important families of flowering plants of the Neotropics.
... Aechmea (Goetze et al., 2017), Alcantarea (Versieux et al., 2012;Lexer et al., 2016), Fosterella (Paule et al., 2017), Pitcairnia (Palma-Silva et al., 2011;Mota et al., 2020), Puya (Jabaily and Sytsma, 2010;Schulte et al., 2010), Tillandsia (Gardner, 1984;Gonçalves and de Azevêdo-Gonçalves, 2009), and Vriesea (Matos et al., 2016;Zanella et al., 2016;Neri et al., 2018). Despite this wealth of evidence for a strong hybridization potential in bromeliads, population genetic studies generally have found low levels of interspecific gene flow in extant populations of bromeliads. ...
... Bromeliaceae are known for their low rate of molecular evolution compared to other Poales (Gaut et al., 1992;Smith and Donoghue, 2008), resulting in a lack of resolution in specieslevel phylogenies reconstructed from a small number of plastid and sometimes nuclear markers (e.g., Sass and Specht, 2010;Versieux et al., 2012;Goetze et al., 2016). In this study, we aimed at circumventing this limitation by using genome skimming, a method which allows the obtention of a large amount of DNA from different genomic compartments without the need for upstream marker development. ...
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The tropics hold at least an order of magnitude greater plant diversity than the temperate zone, yet the reasons for this difference are still subject to debate. Much of tropical plant diversity is in highly speciose genera and understanding the drivers of such high species richness will help solve the tropical diversity enigma. Hybridization has recently been shown to underlie many adaptive radiations, but its role in the evolution of speciose tropical plant genera has received little attention. Here, we address this topic in the hyperdiverse Bromeliaceae genus Vriesea using genome skimming data covering the three genomic compartments. We find evidence for hybridization in ca. 11% of the species in our dataset, both within the genus and between Vriesea and other genera, which is commensurate with hybridization underlying the hyperdiversity of Vriesea, and potentially other genera in Tillandsioideae. While additional genomic research will be needed to further clarify the contribution of hybridization to the rapid diversification of Vriesea, our study provides an important first data point suggesting its importance to the evolution of tropical plant diversity.
... However, demographic analyses usually assume panmictic populations, leading to misinterpretations about demographic changes in highly structured populations, as in V. oligantha (Heller, Chikhi & Siegismund, 2013). Another caveat of demographic changes is the scarcity of polymorphisms in our genetic markers, a common and intrinsic issue found in Bromeliaceae (Versieux et al., 2012). In fact, diversification events throughout the Quaternary are also congruent with other extant lineages from distinct Neotropical montane formations, as in the Espinhaço itself (e.g., Barres et al., 2019;Bonatelli et al., 2014;Chaves et al, 2019;Nascimento et al., 2018; the páramos (e.g., Hughes & Atchison, 2015;Madriñán, Cortés & Richardson, 2013) and the pantepuis (e.g., Rull et al, 2020;Salerno et al., 2012), reinforcing the importance of climate changes throughout the whole Pleistocene in the diversification of the montane biota in the Neotropics. ...
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The exceptional species-richness associated with mountains worldwide is linked to the fragmented topography of these areas, responsible for constantly isolating populations during periods of climatic fluctuations. Consequently, endemism and spatial turnover in mountains are very high and few species are widespread among entire mountain ranges, precluding population-level studies that help understanding how macroevolutionary patterns were shaped. Here, we used the bromeliad Vriesea oligantha, a species endemic to, but widespread in, one of the most species-rich ancient montane areas in the globe, the Espinhaco Range, to test how environmental changes over time may have acted on the evolutionary history of this taxon, contributing to understanding how montane macroevolutionary patterns were shaped. Through analyses of plastidial and nuclear DNA of V. oligantha, we dated its origin and intraspecific diversification, and estimated the genetic diversity, structure and migration rates among populations. Using climatic and geographic variables, we modeled suitable areas for the present and the past, estimating corridors between isolated populations. We also used demographic analyses to estimate ancient population dynamics of V. oligantha. Finally, we tested whether climatic variables or geographical distance explain the observed population structure. The origin and intraspecific diversification of V. oligantha are related to early climatic oscillations during the Plio-Pleistocene. This species has a high population structure due to its low pollen and seed dispersibility. The analysis of species distribution modeling estimated corridors between populations in the past, whereas the structure of V. oligantha results from both models of isolation by distance and isolation by environment. The phylogeographic patterns of Vriesea oligantha reflect previously recognized spatial and temporal macroevolutionary patterns in the Espinhaco Range, providing insights into how microevolutionary processes may have given rise to this astonishing mountain biodiversity.
... Many studies have described that species of Bromelioideae possess an inferior ovary, species of Tillandsioideae have a superior ovary, and species of Glomeropitcairnia Mez and Alcantarea (E. Morren ex Mez) Harms have a half-inferior ovary (Smith and Downs, 1977;Coffani-Nunes et al., 2010;Versieux et al., 2012;Barfuss et al., 2016). The remaining subfamilies are polymorphic with respect to this character and many intermediate levels of the perianth insertions have been observed between the two previous types cited ( Barfuss et al., 2005;Benzing, 2000). ...
... Investigating the gene expression that underlies the formation of petal appendages in bromeliads is an open field of study. Previous studies have revealed little variation in genetic markers that have traditionally been used to construct phylogenetic histories of Bromeliaceae (Terry, Brown & Olmstead, 1997, Givnish et al., 2011, Versieux et al., 2012. Phylogenetic hypotheses indicate that petal appendages appeared several times as an evolutionary adaptation, influenced by pollinators (Schulte & Zizka, 2008;Aguirre-Santoro et al., 2016). ...
Article
Aechmea (Bromeliaceae) is a large genus with controversial systematics and distinct flower shapes and pollinators. We explored floral anatomy and development in two Aechmea spp. belonging to different subgenera to contribute useful information on reproductive biology and taxonomy. We examined floral buds using scanning electron and light microscopy to characterize the development of septal nectaries, petal appendages, ovules, stamens and carpels. In A. gamosepala, we confirmed that the petal appendages develop late, whereas in A. correia-araujoi they develop earlier during floral development. Petal appendage formation included positional changes, possibly affecting floral attributes and visitation by insects, rather than vertebrates. Nectar is released through three basal orifices distally on the ovary, and here we document the link between the nectary region, through discrete canals, upward to the conduplicate lobes of the wet stigma. Improved understanding of the floral development and morphology of Aechmea may help to explain the existence of polymorphic flowers in this genus and may have implications for studies on interactions with pollinators and systematics.