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Morphology of Neoregelia and its subgenera. A. Neoregelia carolinae. B. N. concentrica. C. N. guttata. D. N. lymaniana. E. N. mucugensis. F. N. tenebrosa. G. N. longisepala. H. N. tarapotoensis. I. N. myrmecophila. Photographs by Elton Leme. 

Morphology of Neoregelia and its subgenera. A. Neoregelia carolinae. B. N. concentrica. C. N. guttata. D. N. lymaniana. E. N. mucugensis. F. N. tenebrosa. G. N. longisepala. H. N. tarapotoensis. I. N. myrmecophila. Photographs by Elton Leme. 

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The ‘Nidularioid Complex’ is a group within the Bromelioideae usually characterized by inflorescences nested within water-impounding foliar rosettes. Currently, it comprises six genera: Canistrum, Canistropsis, Edmundoa, Neoregelia, Nidularium, and Wittrockia. While most of these genera occur in eastern Brazil, the distribution of Neoregelia is dis...

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... In the subfamily Bromelioideae, the so-called 'Nidularioid genera' Heller et al., 2015;Santos-Silva et al., 2017) comprise an assemblage of genera that mostly occupies humid forest habitats, typical of the Brazilian Atlantic Forest. Nidularioid genera include Canistropsis (Mez) Leme, Canistrum E.Morren, Edmundoa Leme, Nidularium Lem., Neoregelia L.B.Sm., and Wittrockia Lindm., which share a set of morphological traits, including a particular inflorescence morphology (Mez, 1934;Leme, 1997Leme, , 1998Leme, , 2000Nogueira et al., 2017;Santos-Silva et al., 2017). ...
... In the subfamily Bromelioideae, the so-called 'Nidularioid genera' Heller et al., 2015;Santos-Silva et al., 2017) comprise an assemblage of genera that mostly occupies humid forest habitats, typical of the Brazilian Atlantic Forest. Nidularioid genera include Canistropsis (Mez) Leme, Canistrum E.Morren, Edmundoa Leme, Nidularium Lem., Neoregelia L.B.Sm., and Wittrockia Lindm., which share a set of morphological traits, including a particular inflorescence morphology (Mez, 1934;Leme, 1997Leme, , 1998Leme, , 2000Nogueira et al., 2017;Santos-Silva et al., 2017). These inflorescences have a congested arrangement, with short internodes and large floral bracts that accumulate different amounts of water (Leme, 1998(Leme, , 2000(Leme, , 1997Nogueira et al., 2019Nogueira et al., , 2017. ...
... Although the inflorescence morphology seems to be characteristic to this group, its architecture is still poorly understood, and most of the descriptions on its basic morphology do not have a typology-based comparative approach (Mez, 1934;Smith and Downs, 1979;Leme, 1997Leme, , 1998Leme, , 2000. Moreover, the relationship among these genera is still not completely known, and the inflorescence arrangement could be a convergent morphological feature, instead of a synapomorphy of this group (Santos-Silva et al., 2017). ...
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In the subfamily Bromelioideae, the so-called Nidularioid genera share some morphological features, the most notable of which is the occurrence of congested inflorescences that accumulate different amounts of water. This study aims to describe and compare the inflorescence morphology in six Nidularioid genera and to estimate the “tank space” for water in each inflorescence. Inflorescences at different developmental stages were analyzed using light microscopy and described using a typology-based comparative approach. X-ray microtomography was used to obtain sections of young inflorescences, from which 3-dimensional images were built for exploring structural issues and estimating water impounding capacity. Inflorescences are simple racemes in N. johannis and heterothetic compound racemes in the other species. No adaxial prophylls were observed among Nidularioid genera, neither in floral axes nor in the inflorescence branches. The flowers are spirally arranged along the florescence axis although the polystichous condition is obscured by the dorsiventral compression of florescences. The development of Nidularioid compound inflorescences follows a bidirectional pattern since flower primordia differentiate acropetally and paraclade primordia develop basipetally. Bracts are used for water storage in species with a long scape, where such bracts are large and cover the entire paraclades. In species with a short floral scape, inflorescences remain submerged inside the vegetative phytotelmata, allowing these inflorescences to be always in contact with water. Despite the actual branching pattern is obscured because the inflorescence branches are closely condensed in the axils of water impounding bracts, all Nidularioid inflorescences constitute polytelic systems that follow the general pattern of Bromeliaceae. Although all Nidularioid inflorescences are associated with structures for water retention, they seem not to be synapomorphic. Thus, hypotheses of primary homology suggested by both the typological analysis of inflorescences and the survey of water-impounding structures are consistent with the non-monophyletic condition of the Nidularioid genera.
... In a recent study by Silva et al. (2018) the anatomy of vegetative organs, inflorescence axis and pedicel of the Neoregelia bahiana complex (Bromelioideae) were studied, showing that leaf also provides taxonomic characters. The results observed here, as well as those by Sajo et al. (1998), Santos-Silva et al. (2017, and Silva et al. (2018) show the relevance of leaf anatomical characters in the delimitation of morphologically related species that form complexes in the subfamily Bromelioideae. Hence, a broader leaf morpho-anatomical study is being conducted (Cavalcante et al. in prep.) ...
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We here establish the Hohenbergia capitata complex composed of three species endemic to the Atlantic Forest, in the State of Bahia, Northeastern Brazil. When compared with other Hohenbergia species endemic to the Atlantic Forest, the complex is recognized by the small size of the plants (shorter than 1 m tall when flowering), lanceolate leaf blades, inflorescence with main axis short and congested branches, concentrated on the apical portion of the inflorescence (creating a capitulate shape), primary branches short pedunculate (short stipes), basal primary bracts sub-orbicular, large flowers (over 3 cm long) with spatulate petals with a cuspidate apex. Within this complex, we describe H. nidularioides sp. nov., a critically endangered species from the southern coastal region of Bahia, only known from a small fragment of Restinga forest in Una municipality. This species is very similar to H. capitata but differs by the nidular aspect of the inflorescence, which is only short-pedunculate and more or less hidden inside the rosette. In addition to the taxonomic treatment, we provide illustrations, the geographic distribution, taxonomic comments, and anatomical comparison of the species in the H. capitata complex.
... Neoregelia ibitipocensis (Leme) Leme, the species on which this work focuses, is an endemic and rare bromeliad from the Brazilian Atlantic Forest. Neoregelia stands out as the genus with the largest species number within the 'nidularioid complex', which is characterised by congested inflorescences with short peduncles that develop within the water tank formed by the rosette of leaves (Santos- Silva et al. 2017). The populations of N. ibitipocensis have restricted distribution to a few portions of cloud forests in the Mantiqueira Mountain Range, south-east Brazil (Leme 2000). ...
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Herbivory in plant reproductive tissues can severely reduce the fitness of the species, especially for those threatened with extinction. We investigated aspects of the reproductive ecology of Neoregelia ibitipocensis (Leme) Leme, an endangered species restricted to a few cloud forests sites in the Brazilian Atlantic Forest. Data on breeding systems, fertility, and viability of its populations are provided, as well as alternatives for development of strategies for ex situ conservation. Hand pollination experiments revealed that N. ibitipocensis is partially self-incompatible. Fruit and seeds were affected by intense herbivory caused by Tipulidae (Diptera) larvae. The predation reached 100% of the analysed infructescences during the first year of the study. We observed the production of parthenocarpic fruit both through controlled pollination experiments and under natural conditions. The potential for self-pollination and autogamy probably compensate for the reduced ability of the species to attract pollinators due to its small floral display. In the same way, parthenocarpy seems to represent a strategy to enhance the fitness of N. ibitipocensis, mitigating the effects of the predation. We developed an efficient protocol of in vitro multiplication, which represents a viable alternative for ex situ conservation of the species.
... 2; 3), traditional morphological characters plus new or underutilized traits, and molecular phylogeny provides a consistent pattern and thus proved to be useful to propose a new status for Hylaeaicum as a distinct genus, well differentiated from the genera of the "Nidularioid complex", where it has been traditionally placed in. An integrative approach with the combination of multiple traits (i.e., morphological, palynological, biogeographic, ecological, and molecular) and the critical assessment of the diagnostic characters (e.g., inflorescence structure, flower pedicels, sepal concrescence, petal appendages, stigma and seed types) as proposed by Leme (1997Leme ( , 1998Leme ( , 2000, Santos-Silva et al. (2017), and Leme et al. (2017) for generic delimitation in bromelioids, was applied here to revise Hylaeaicum. The thorough consideration of several morphological characters is the principal goal of this study. ...
... When the taxonomical revision of the "Nidularioid complex" s.l. was concluded proposing new circumscriptions for Canistrum, Nidularium, and Wittrockia, as well as the new genera Canistropsis and Edmundoa, and two new subgenera for Neoregelia (Leme 1997(Leme , 1998(Leme , 2000 it became clear that Hylaeaicum was neither related to Neoregelia nor any other genus of the complex. It confirms the particular idea of Ramírez (1991Ramírez ( , 2000 that Hylaeaicum is only apparently related to Neoregelia due to homoplasious characters, especially the nidular inflorescence, but it is rather associated with Aechmea (see also Santos-Silva et al. 2017), which reinforced the common sense of the taxonomists on the distinctness of the group (e.g., H. Luther, pers. comm.). ...
... 1), it is evident that the Hylaeaicum species are a monophyletic group forming a strongly supported clade. The same applies to the remaining species of the "Nidularioid complex" s.str (clade W) However, both clades are located on different major branches of the phylogenetic tree (clade Y vs. clade O) and clearly do not form a monophyletic group similarly as previously shown by Silvestro et al. (2014), Santos-Silva et al. (2017), andHeller (2018). Thus, the phylogenetic analyses support Hylaeaicum as a separate genus. ...
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Generic status for the Amazonian Hylaeaicum is proposed within the Aechmea alliance, excluding it from the “Nidularioid complex” in general and from Neoregelia in particular. The monophyly of this new genus is supported by molecular phylogenetic analyses. The taxonomic circumscription of Hylaeaicum is based on the combination of geographical range and morphological characters, such as clonal growth, inflorescence structure, petal and corolla conformation, petal appendages, ovary, ovule, stigma, pollen, fruit, and seed, as well as seed anatomy, thoroughly documented and illustrated from field-collected specimens that flowered in cultivation in the Rio de Janeiro Botanical Garden, the Marie Selby Botanical Gardens, and in Refúgio dos Gravatás. The presence of seeds with long bicaudate appendages on both chalazal and micropylar ends is reported for the first time in Bromelioideae and considered an important character to distinguish Hylaeaicum from the related genera in the Aechmea alliance. In order to support the morphological distinctness of Hylaeaicum, the most varied and complete documentation of stigmata, fruits, and seeds of Bromelioideae is also presented for the first time, covering 24 genera and 17 subgenera. Fourteen new combinations, including 12 species and two varieties, are proposed.
... (Figure 1). The bromeliads were identified using reference taxonomic works of literature and monographs done by Santos-Silva et al. (2017), Parkhurst (2000, as well as Leme and Marigo (1993). The water samples were collected from leaf cavities of individual bromeliads (Neoregelia carolinae (Beer) L.B. Sm. "Devroe," Neoregelia cruenta (R. Graham) L.B. Sm. "Selecta," and Neoregelia compacta (Mez) L.B. Sm.) using a 50 mL syringe with a plastic hose ( Figure 2). ...
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The aquatic microcosms in plant axils of bromeliads are unique and favorable habitat for diverse kinds of new and rare species of microalgae. This study presents the first taxonomic survey in the Philippines to focus on the algal community existing in phytotelmata of Neoregelia spp. L.B. Smith (Bromeliaceae) from florist wholesalers in Laguna. A total of 20 microalgal taxa were described and taxonomically identified-of which six species belong to Cyanophyceae, five species both for Bacillariophyceae and Chlorophyceae, two species for Trebouxiophyceae, and one species each for Euglenophyceae and Zygnematophyceae. The study reports an additional of eleven species (Chlorella vulgaris, Dictyochloropsis splendida, Stigeoclonium tenue, Chlorococcum infusionum, Pinnularia rumrichiae, Cymbella affinis, Chroococcus schizodermaticus, Pseudanabaena minima, Oscillatoria limosa, Oscillatoria tenuis, and Euglena agilis) as new records in the global inventory list of phytotelm microalgae and 14 new taxa as additional records of microalgae in phytotelm microhabitat of Asia. Also, the occurrence of a microalga, Dictyochloropsis splendida Geitler, is described for the first time in the Philippines. Keys and diagnostic descriptions as well as photomicrographs are presented to differentiate the species of phytotelm microalgae. The survey provided important taxonomic records and information on the species composition of phytotelm microalgae of bromeliad tanks found in the Philippines.
... Neoregelia is a genus of bromeliads that have the highest diversity of species compared to other genera. Neoregelia has a characteristic arrangement of dense leaves and forms a bowl like a rose (Santos-Silva et al. 2017). Bromeliads are the most popular ornamental for plant enthusiasts and are found in Indonesia (Trubus 2008). ...
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Ikhsan M, Hadi UK, Soviana S. 2020. Diversity and distribution bromeliads plants as breeding habitat for mosquito larvae (Diptera: Culicidae) in Bogor, Indonesia. Biodiversitas 21: 3494-3498. Bromeliads leaf structure can hold water and has potential as the breeding ground for mosquitoes. Information regarding the potential of bromeliads as a habitat for mosquito larvae in Indonesia is limited. This study aims to identify the diversity of bromeliads and their potential as breeding habitat for mosquito larvae in West Bogor Sub-district, Bogor City, West Java, Indonesia. Determination of the sample using purposive sampling method. Larvae collections were carried out on bromeliads that were outside the building in December 2019 to Maret 2020. The collection used a 50 ml syringe that was modified using a small pipe. Total bromeliads collected were 117 consisted of Neoregelia, Cryptanthus, dan Alcantarea from 50 observation points. The percentage of mosquito larvae presence from bromeliads was 74.4%. Total larvae collected were 859 larvae consisting of Aedes albopictus, Aedes aegypti, Aedes spp., Armigeres subalbatus, and Culex quinquefasciatus. The diversity of mosquito larvae found in the bromeliads were low (H'=0.16), while the dominant species of mosquito larvae were Ae. albopictus (61.55%) and confirmed as the second vector of Dengue Fever in Indonesia. The results of this study indicate that bromeliads have potential as a breeding site for mosquito larvae in West Bogor Sub-district, Bogor City.
... are rare species that are endemic to a few cloud forests in the Mantiqueira Mountain Range in south-eastern Brazil. These species have fleshy indehiscent fruits, produce monocarpic ramets and belong to the nidularioid complex, characterized by inflorescences with a short peduncle, located in the water tank formed by the rosette of leaves (Leme, 2000;Santos-Silva et al., 2017). Neoregelia ibitipocensis is predominantly terrestrial and occasionally found as an epiphyte. ...
... This type of pollination mechanism, involving corolla opening by bees, has not been described previously in Bromeliaceae, although N. leucophoea (Baker) L.B.Sm. and N. lymaniana R.Braga & Sucre bear flowers with a similar corolla appearance at anthesis (Costa & Wendt, 2007;Santos-Silva et al., 2017). ...
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Although plant demography, genetics and reproduction are interrelated processes, few studies on rare tropical plants have attempted to integrate them. We used an integrated approach to study two endangered Neoregelia spp. from the Brazilian Atlantic Forest. The floral phenotypes of N. ibitipocensis and N. oligantha are similar, and both species are visited and pollinated only by bumblebees. Flowers of N. ibitipocensis secrete copious nectar, which bees can access only by forcing the corolla to open, whereas flowers of N. oligantha are nectarless. Neoregelia ibitipocensis is self-incompatible and pollen-limited, whereas N. oligantha is self-compatible and sets fruits spontaneously. Population growth rates indicate an increasing population size for both species, although to a lesser extent for N. oligantha. Genetic diversity as a whole was moderate; Bayesian analysis indicated a spatially structured cluster for N. ibitipocensis populations, and genetic diversity was scattered across the distribution of N. oligantha. Despite these differences, vegetative reproduction via clonal growth was the main component of population structuring in both species. Limited seed production in N. ibitipocensis and the risk of genetic drift in N. oligantha seem to be the main threats to their persistence.
... are rare species that are endemic to a few cloud forests in the Mantiqueira Mountain Range in south-eastern Brazil. These species have fleshy indehiscent fruits, produce monocarpic ramets and belong to the nidularioid complex, characterized by inflorescences with a short peduncle, located in the water tank formed by the rosette of leaves (Leme, 2000;Santos-Silva et al., 2017). Neoregelia ibitipocensis is predominantly terrestrial and occasionally found as an epiphyte. ...
... This type of pollination mechanism, involving corolla opening by bees, has not been described previously in Bromeliaceae, although N. leucophoea (Baker) L.B.Sm. and N. lymaniana R.Braga & Sucre bear flowers with a similar corolla appearance at anthesis (Costa & Wendt, 2007;Santos-Silva et al., 2017). ...
Article
Full-text available
Although plant demography, genetics and reproduction are inter-related processes, few studies on rare tropical plants have attempted to integrate them. We used an integrated approach to study two endangered Neoregelia spp. from the Brazilian Atlantic Forest. The floral phenotypes of N. ibitipocensis and N. oligantha are similar, and both species are visited and pollinated only by bumblebees. Flowers of N. ibitipocensis secrete copious nectar, which bees can access only by forcing the corolla to open, whereas flowers of N. oligantha are nectarless. Neoregelia ibitipocensis is self-incompatible and pollen-limited, whereas N. oligantha is self-compatible and sets fruits spontaneously. Population growth rates indicate an increasing population size for both species, although to a lesser extent for N. oligantha. Genetic diversity as a whole was moderate; Bayesian analysis indicated a spatially structured cluster for N. ibitipocensis populations, and genetic diversity was scattered across the distribution of N. oligantha. Despite these differences, vegetative reproduction via clonal growth was the main component of population structuring in both species. Limited seed production in N. ibitipocensis and the risk of genetic drift in N. oligantha seem to be the main threats to their persistence.
... The corresponding sequences present in our Agt1 dataset cluster with identity values above 98% and no non-"Portea/Gravisia complex" members are present within this cluster. Another six clusters represent groups recently assigned to the "Cryptanthoid complex" [23], five are members of the "Nidularioid complex" [30,31] and one comprises only taxa of the recently studied genus Ananas [25]. Among the remaining Bromelioideae Agt1 sequences we found seven clusters that consist only of members from either one of the following genera: Wittmackia, Billbergia, Araeococcus, Hohenbergia, Lymania or Neoregelia. ...
... Among the remaining Bromelioideae Agt1 sequences we found seven clusters that consist only of members from either one of the following genera: Wittmackia, Billbergia, Araeococcus, Hohenbergia, Lymania or Neoregelia. All of these genera are currently considered as being monophyletic [29][30][31][32][33][34][35][36][37][38]. ...
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Background The angiosperm family Bromeliaceae comprises over 3.500 species characterized by exceptionally high morphological and ecological diversity, but a very low genetic variation. In many genera, plants are vegetatively very similar which makes determination of non flowering bromeliads difficult. This is particularly problematic with living collections where plants are often cultivated over decades without flowering. DNA barcoding is therefore a very promising approach to provide reliable and convenient assistance in species determination. However, the observed low genetic variation of canonical barcoding markers in bromeliads causes problems. Result In this study the low-copy nuclear gene Agt1 is identified as a novel DNA barcoding marker suitable for molecular identification of closely related bromeliad species. Combining a comparatively slowly evolving exon sequence with an adjacent, genetically highly variable intron, correctly matching MegaBLAST based species identification rate was found to be approximately double the highest rate yet reported for bromeliads using other barcode markers. Conclusion In the present work, we characterize Agt1 as a novel plant DNA barcoding marker to be used for barcoding of bromeliads, a plant group with low genetic variation. Moreover, we provide a comprehensive marker sequence dataset for further use in the bromeliad research community.
... In the present analysis, anatomical characters ( Fig. 4) were demonstrated to be synapomorphies and, therefore, contributed significantly to producing a tree with higher resolution. In Bromeliaceae, few phylogenetic studies include anatomical data, but when anatomical characters are included they contribute to tree resolution (e.g., Gomes-da-Silva, Vargens, Arruda, & Costa, 2012;Santos-Silva et al., 2017;Saraiva, Mantovani, & Forzza, 2015). 8. Dyckia and Encholirium were published in the same work. ...
Article
The genera Deuterocohnia, Dyckia, and Encholirium are recognized as the xeric clade Pitcairnioideae. More than 220 xeric species of this group occur in South America in the Andean region, the Chacoan subregion, and the South American Transition Zone. The species recently and rapidly diversified, and such phylogenetic relationships are difficult. This clade is well-supported in phylogenetic analyses, but despite the monophyly of this group of xeric species, the analyses have demonstrated the paraphyly or polyphyly of genera in the xeric clade. However, most of these studies suffer with partial resolution, limited taxon sampling or incorrect homology statements. To investigate this group, we conducted the most comprehensive phylogenetic analysis of the xeric clade to date using parsimony criteria with morphological and molecular datasets, including plastid and nuclear genomes concatenated for the first time, to establish a suitable classification. A total of 133 accessions were included that corresponded to 109 taxa. The dataset included 70 morphological characters and �6 kb of sequence data from the plastid and nuclear genomes. Phylogenetic analyses were conducted using equal and implied weighting. The analysis resulted in a well-supported cladistic hypothesis and demonstrated the three genera as currently circumscribed are not supported as monophyletic. Based on the results we propose a new classification that reflects the evolutionary relationships, a new circumscription for Dyckia is proposed that includes the species belonging to Deuterocohnia and Encholirium. For this circumscription, 41 combinations were made for Dyckia.