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A–D. Aechmea nigribracteata ( J.R. Maciel et al. 1950 ). A. Inflorescence, B. Longitudinal section of inflorescence, central axis and peduncle details, C. Floral bracts and flower details, D. Habit. (Photos: J.R. Maciel).
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Herein is described and illustrated a new Aechmea species. Aechmea nigribracteata grows in southern Bahia (Brazil), a region recognised as highly diverse in bromeliads. Aechmea nigribracteata belongs to Aechmea subg. Chevaliera and has completely dentate, blackish floral bracts and petals with two crenate-laciniate ligules.
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Citations
... Distribution-Trinidad and Tobago, French Guiana, and Suriname. In Trinidad, it is known from Aripo savannas, growing in savannas and surrounding forest at 30-50 m. Maciel et al. 2014;Gouda et al. 2023, continuously updated). In Trinidad and Tobago, nine species were reported by Baksh-Comeau et al. (2016). ...
... Thus, a more updated delimitation of this taxonomic complex includes A. multiflora, A. depressa, A. hostilis, A. nigribracteata J.R. Maciel, Louzada and M. Alves, A. prasinata G. Sousa and Wanderley, and A. saxicola. Aechmea gustavoi J.A. Siqueira and Leme was the last species assigned to the A. multiflora complex due to its morphological singularity (Leme and Siqueira-Filho 2001;Canela et al. 2003;Maciel et al. 2014;Sousa and Wanderley 2014). The monophyly of this group was confirmed recently in a molecular phylogenetic study of subgenus Chevaliera (Maciel et al. 2018). ...
... Most of the seven species (A. gustavoi, A. multiflora, A. nigribracteata, A. depressa, A. hostilis, A. prasinata, and A. saxicola; Fig. 1) gathered in this clade were formerly recognized as A. multiflora complex, except A. gustavoi (Leme and Siqueira-Filho 2001;Canela et al. 2003;Maciel et al. 2014;Sousa and Wanderley 2014). ...
... The first mention of any morphological relationship of A. gustavoi with species of the A. multiflora complex appears in Sousa (2004), who compared floral bracts of A. gustavoi and A. hostilis. Maciel et al. (2014) included the newly described A. nigribracteata in the A. multiflora complex, but did not relate the new species to A. gustavoi. Finally, Sousa and Wanderley (2014) related A. prasinata to A. hostilis and A. gustavoi. ...
... Thus, a more updated delimitation of this taxonomic complex includes A. multiflora, A. depressa, A. hostilis, A. nigribracteata J.R. Maciel, Louzada and M. Alves, A. prasinata G. Sousa and Wanderley, and A. saxicola. Aechmea gustavoi J.A. Siqueira and Leme was the last species assigned to the A. multiflora complex due to its morphological singularity (Leme and Siqueira-Filho 2001;Canela et al. 2003;Maciel et al. 2014;Sousa and Wanderley 2014). The monophyly of this group was confirmed recently in a molecular phylogenetic study of subgenus Chevaliera (Maciel et al. 2018). ...
... Most of the seven species (A. gustavoi, A. multiflora, A. nigribracteata, A. depressa, A. hostilis, A. prasinata, and A. saxicola; Fig. 1) gathered in this clade were formerly recognized as A. multiflora complex, except A. gustavoi (Leme and Siqueira-Filho 2001;Canela et al. 2003;Maciel et al. 2014;Sousa and Wanderley 2014). ...
... The first mention of any morphological relationship of A. gustavoi with species of the A. multiflora complex appears in Sousa (2004), who compared floral bracts of A. gustavoi and A. hostilis. Maciel et al. (2014) included the newly described A. nigribracteata in the A. multiflora complex, but did not relate the new species to A. gustavoi. Finally, Sousa and Wanderley (2014) related A. prasinata to A. hostilis and A. gustavoi. ...
... Until now, the widely accepted concept of Aechmea subgenus Chevaliera has been based on inflorescence and flower morphology, with c. 30 species sharing simple, strobiliform or rarely digitate, polystichous and often perennial inflorescences, coriaceous or ligneous floral bracts, free or connate sepals and reduced or absent petal appendages (Smith & Downs, 1979;Leme & Kollmann, 2009, 2011Leme et al., 2010Leme et al., , 2014Leme, 2010;Maciel, Louzada & Alves, 2014;Sousa & Wanderley, 2014). Additionally, A. subgenus Chevaliera is recognized as the only group in Bromelioideae that combines massive and strobiliform inflorescences with large rosettes forming dense clumps (Smith & Downs, 1979;Silva, 2003;Sousa, 2004;Maciel, Zizka & Alves, 2015). ...
... These species occur in Central America, the Andes, Amazon and Atlantic Forest, displaying a disjunct distribution between northern and eastern South America. The Atlantic Forest along the east coast of Brazil is the centre of diversity for this species group (Smith & Downs, 1979;Canela et al., 2003;Silva, 2003;Sousa, 2004;Leme & Kollmann, 2009, 2011Leme, 2010;Leme et al., 2010Leme et al., , 2014Maciel et al., 2014Maciel et al., , 2015Sousa & Wanderley, 2014). Silva (2003) provided an improved concept of A. subgenus Chevaliera as given by Smith & Downs (1979). ...
... This point of view is similar to that of Mez (1896) Mez and A. veitchii Baker] and kept the Atlantic species as the separate genus Chevaliera. The results of Silva (2003) become more relevant from the perspective of geographical conservatism in phylogenetics of Bromelioideae, which means that species with morphological affinities and sharing geographical patterns tend to be monophyletic (Sass & Specht, 2010 (Leme & Siqueira-Filho, 2001;Leme & Silva, 2002;Canela et al., 2003;Leme & Kollmann, 2009, 2011Leme, 2010;Maciel et al., 2014). ...
A better understanding of the phylogenetics of Aechmea subgenus Chevaliera is hampered by the limited taxonomic sampling in molecular studies of Bromelioideae. To test the monophyly of Aechmea subgenus Chevaliera, we conducted a molecular phylogenetic study using three DNA regions (ETS, matK and phyC) and a reconstruction of ancestral states of morphological characters. Our sampling included 22 of the up to 30 species recorded in the subgenus. Aechmea subgenus Chevaliera turned out to be clearly polyphyletic, with Amazonian species previously included in A. subgenus Chevaliera being found in separate clades of the phylogenetic tree for Bromelioideae. Two distinct groups comprise the majority of the Atlantic species up to now grouped in Chevaliera. The first of these, referred to as the A. multiflora group, includes seven species related to A. multiflora. The second group includes 11 species, referred to as the A. sphaerocephala group. Morphological analyses show that the A. multiflora and A. sphaerocephala groups are convergent in plant size, growth form and inflorescence morphology. The groups can morphologically be best differentiated based on the margin of the floral bracts and petal colour.
... Until now, the widely accepted concept of Aechmea subgenus Chevaliera has been based on inflorescence and flower morphology, with c. 30 species sharing simple, strobiliform or rarely digitate, polystichous and often perennial inflorescences, coriaceous or ligneous floral bracts, free or connate sepals and reduced or absent petal appendages (Smith & Downs, 1979;Leme & Kollmann, 2009, 2011Leme et al., 2010Leme et al., , 2014Leme, 2010;Maciel, Louzada & Alves, 2014;Sousa & Wanderley, 2014). Additionally, A. subgenus Chevaliera is recognized as the only group in Bromelioideae that combines massive and strobiliform inflorescences with large rosettes forming dense clumps (Smith & Downs, 1979;Silva, 2003;Sousa, 2004;Maciel, Zizka & Alves, 2015). ...
... These species occur in Central America, the Andes, Amazon and Atlantic Forest, displaying a disjunct distribution between northern and eastern South America. The Atlantic Forest along the east coast of Brazil is the centre of diversity for this species group (Smith & Downs, 1979;Canela et al., 2003;Silva, 2003;Sousa, 2004;Leme & Kollmann, 2009, 2011Leme, 2010;Leme et al., 2010Leme et al., , 2014Maciel et al., 2014Maciel et al., , 2015Sousa & Wanderley, 2014). Silva (2003) provided an improved concept of A. subgenus Chevaliera as given by Smith & Downs (1979). ...
... This point of view is similar to that of Mez (1896) Mez and A. veitchii Baker] and kept the Atlantic species as the separate genus Chevaliera. The results of Silva (2003) become more relevant from the perspective of geographical conservatism in phylogenetics of Bromelioideae, which means that species with morphological affinities and sharing geographical patterns tend to be monophyletic (Sass & Specht, 2010 (Leme & Siqueira-Filho, 2001;Leme & Silva, 2002;Canela et al., 2003;Leme & Kollmann, 2009, 2011Leme, 2010;Maciel et al., 2014). ...
A better understanding of the phylogenetics of Aechmea subgenus Chevaliera is hampered by the limited taxonomic sampling in molecular studies of Bromelioideae. To test the monophyly of Aechmea subgenus Chevaliera, we conducted a molecular phylogenetic study using three DNA regions (ETS, matK and phyC) and a reconstruction of ancestral states of morphological characters. Our sampling included 22 of the up to 30 species recorded in the subgenus. Aechmea subgenus Chevaliera turned out to be clearly polyphyletic, with Amazonian species previously included in A. subgenus Chevaliera being found in separate clades of the phylogenetic tree for Bromelioideae. Two distinct groups comprise the majority of the Atlantic species up to now grouped in Chevaliera. The first of these, referred to as the A. multiflora group, includes seven species related to A. multiflora. The second group includes 11 species, referred to as the A. sphaerocephala group. Morphological analyses show that the A. multiflora and A. sphaerocephala groups are convergent in plant size, growth form and inflorescence morphology. The groups can morphologically be best differentiated based on the margin of the floral bracts and petal colour. ADDITIONAL KEYWORDS: Bromelioideae-evolution-floral bracts-flower-inflorescence-plant size-taxonomy.
... Historically, the central Atlantic Forest has been poorly collected when compared to the southern region, especially in medium and high elevation areas . Several new species of vascular plants have been described in the last eight years from the Atlantic Forest, with the highest rate of discoveries in the central portion of the biome: Acanthaceae (Profice and Leitman 2013); Bromeliaceae (Amorim and Leme 2009;Costa et al. 2012;Versieux et al. 2012;Maciel et al. 2014;Sousa and Wanderley 2014); Clusiaceae (Marinho et al. 2015(Marinho et al. , 2016a; Chrysobalanaceae (Asprino and Amorim 2016); Cyperaceae (Mayedo and Thomas 2016); Dichapetalaceae ; Erythroxylaceae (Araújo et al. 2015); Gentianaceae (Grant and Trunz 2011); Hypericaceae (Marinho et al. 2016b); Leguminosae (Cardoso et al. 2009(Cardoso et al. , 2017; Malpighiaceae Almeida et al. 2016); Moraceae (Machado et al. 2013); Myrtaceae (Sobral and Mazine 2010;Sobral et al. 2016;Tuler et al. 2017); Ochnaceae (Fraga and Saavedra 2014); Orchidaceae (Fraga et al. 2015); Piperaceae (Monteiro et al. 2016); Pteridaceae (Prado and Hirai 2013); Thelypteridaceae (Matos et al. 2010). Some of these were new genera [e.g. ...
Information regarding conservation actions and species distribution are based on specimens collected and deposited in scientific collections and taxonomic studies. In the tropics, these data might be biased for several reasons. Even in well-explored regions, such as the Atlantic Forest, there is still a lot to learn regarding distribution and evolution of genera and species. Intense sampling effort during the last 15 yr in the central Atlantic Forest has led to the description of six new species of Bertolonia (Melastomataceae). Bertolonia angustipetala, B. cuspidata, B. linearifolia, B. reginatoi, and B. vitoriana also occurs in Minas Gerais, near the border between both states. These and other recent discoveries in Bertolonia shift the diversity center of the genus from the southern to the central region of the Atlantic Forest, and provide a new context to understand morphological and biogeographical variation within the genus.
... Chevaliera (A. saxicola, A. multiflora, A. sphaerocephala) to be monophyletic and well supported in an otherwise mostly unsupported cladogram. According to Maciel et al. (2014) the subgenus Chevaliera comprises 30 species. ...
... The phylogeny presented here adds the species Aechmea perforata and A. gustavoi to this complex. Further potential members of this group are A. nigribracteata (Maciel et al., 2014) which is morphologically close to A. depressa, and A. prasinata which is closely related to A. hostilis (de Sousa and Wanderley, 2015). The position of A. perforata in the Aechmea subgen. ...
Phylogenetic trouble unleashed
The first part of my thesis deals with a comprehensive phylogeny of the Bromelioideae subfamily. The family Bromeliaceae is subdivided into eight subfamilies, one of them is the Bromelioideae. Phylogenetic relationships among the Bromelioideae are still poorly understood and many of the extant genera are suspected to be not monophyletic. Especially Aechmea, the largest and most polymorphic genus constitutes many questions and the genus was used as a depot for taxonomically problematic species. The phylogenetic study presented here is the most comprehensive one so far, covering about half of the known species (434 of 965, Table 1) of Bromelioideae. The phylogeny was generated using plastid (atpB-rbcL, matK, rps16, ycf1_1, ycf1_6) and nuclear (AGT1_exon, ETS, G3PDH, PHYC, RPB2) genetic markers. The markers were analysed individually as well as combined using maximum likelihood and Bayesian analysis. The comparison of plastid vs. nuclear data revealed significant differences which were discussed in detail and hypothesised to indicate hybridisation in certain lineages. Nevertheless, the combination of both datasets increased the overall resolution of the phylogeny and was used to discuss the results in the light of previous studies. The entire phylogeny was divided into 32 groups for discussion. These groups represent potential genera or starting points for further studies in order to reorganise the polyphyletic genera of Bromelioideae into monophyletic lineages. Many extant genera of the eu-Bromelioideae were found to be not monophyletic. Monophyly was observed for the genera Acanthostachys, Billbergia, Cryptanthus, Disteganthus, Hoplocrypanthus, Lapanthus, Orthocryptanthus, Orthophytum, Rokautskyia, Ronnbergia, Sincoraea, Wittmackia and the monotypic ones (Deinacanthon, Eduandrea, Fascicularia, Hohenbergiopsis, Pseudananas). The genus concept proposed by Smith and Downs (1979) is therefore rejected, as well as the taxonomic utility of petal appendages, which were mainly used to delimit genera. In summary, this study and recent studies highlighted other morphological characters (e.g. pollen morphology, stigma type) as much more informative. However, no single character should be used to delimit genera and combinations of relevant characters are required. Even the petal appendages can pose a taxonomical important character at certain taxonomic level.
The combination of biogeography and phylogeny revealed that species of some groups which co- occur in a biome or region are also phylogenetically closely related. These groups were not recognised before because the misinterpretation of homoplastic characters led to wrong taxonomical conclusion. For example, the recent re-organisation of the Cryptanthoid group and the re-establishment of Wittmackia with the former Hohenbergia subgen. Wittmackiopsis species highlighted, among other characters, the importance of biogeography. Another case is the subgenus Neoregelia subgen. Hylaeaicum which is geographically and phylogenetically separated from the Nidularioid group and therefore has to be excluded.
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The large phylogeny presented here gives evidence for multiple invasions of the Brazilian biomes (Amazon Forest, Atlantic Forest, Cerrado, Caatinga) as well as of Central America and the Greater Antilles. It is important to note that the phylogeny is lacking resolution in the deeper nodes. Confident assumptions are therefore hindered and the historical biogeography of Bromelioideae remains cryptic. Anyway, the Atlantic Forest is nowadays the diversity hotspot of the core Bromelioideae and critically endangered. Extensive conservation efforts are required to protect the diverse flora, including the bromeliads.
The genetic markers used so far in bromeliad phylogenies provided only limited variation resulting in often unresolved complexes. The search for additional suitable genetic markers in bromelioid phylogenies yielded the nuclear marker AGT1. The amplified fragment consists of one well conserved exon region as well as a highly variable intron. The intron was too variable for aligning it across the entire bromelioid set. On the other hand, the intron provides relevant information for inferring phylogenies of closely related species groups (e.g. in Ananas, Cryptanthoid group). Furthermore, AGT1 is proposed as a genetic barcode in Bromelioideae because it poses much more information then the commonly used ones (e.g. matK).
Does size matter?
The second part of this thesis deals with the genome size evolution within the family Bromeliaceae. Samples from seven subfamilies were screened with the emphasis on the subfamily Bromelioideae. The data were combined with data from literature and the observed patterns were discussed in relation to known phenomena (e.g. correlations to environment and life form). In the second sub-chapter I have chosen the species Tillandsia usneoides to study the intraspecific genome size variation in combination with morphology and biogeography. Genome size and base composition were measured using the flow cytometry technique.
Bromeliaceae comprises mostly diploid species with predominantly 50 small chromosomes (2n), small genome sizes (0.59-4.11 pg) and normal GC content (36.46-42.21 %) compared to other families. Polyploidy was observed so far in the subfamilies Bromelioideae, Tillandsioideae and Pitcairnioideae. Triploids, tetraploids and potential hexaploids were identified. The genera show significant differences in holoploid genome size and base composition throughout the entire family. GC content is weakly positively correlated with genome size. Significant intraspecific genome size variation has been observed, including polyploidization, but no endopolyploidy and no variation in dioecious species. Within the subfamily Bromelioideae, the observed genome size between the early diverging lineages and the core Bromelioideae supports this division. The differences are due to a higher proportion of polyploids in the early diverging lineages and a significant higher
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GC content in the core Bromelioideae. Both groups differ in their life strategies and occupy principally different habitats with corresponding morphological adaptations. Hence, the early diverging lineages are predominantly terrestrial and xeromorphic. In contrast, the prevailing epiphytic core Bromelioideae are characterised by a tank habit and mostly adapted to more humid environments. Across the family and the subfamily Bromelioideae in particular, significant genome size differences between the different life forms have been observed, but no correlation to biomes within Brazil.
Tillandsia usneoides is the most widely distributed species of the family Bromeliaceae. It ranges from the southeastern United States to Argentina and Chile. Tillandsia usneoides grows epiphytic and is dispersed by seeds as well as by fragments of the plant. Within the species striking morphological differences can be observed as far as size characters are concerned. Morphotypes have shown to be stable in cultivation while growing under the same conditions. In order to investigate possible reasons for the variation the relative genome size of 75 specimens covering the whole distribution range was measured and combined with morphological, distribution and climatic data. Significant variation in the relative genome size corresponded to the morphological differences and reflected the north-south distribution gradient. Genome size and morphotypes showed a positive correlation, as well as with the mean temperature of the driest and coldest quarter and the minimal temperature of the coldest month.
... Aechmea prasinata and A. hostilis are registered to the state of Espírito Santo and A. gustavoi to the states of Pernambuco and Bahia (Leme & Siqueira-Filho 2001). Others species of Aechmea of the subgenus Chevaliera were recently described, occurring in the state of Espírito Santo and Bahia, however they are very distinctive of the new species manly by the long peduncled inflorescence (Leme et al. 2010, Leme & Kollmann 2011, Leme et al. 2014, Maciel et al. 2014). Besides Canela et al. (2003) as considered A. hostilis as synonym of A. saxicola L.B. Sm. this position is not accepted in the present work, considering two distinctive species. ...
Aechmea prasinata G. Sousa & Wand., a new species from southeast Brazil, is described and illustrated. The species belongs to the subg. Chevaliera and shows morphological similarities with Aechmea hostilis E. Pereira and Aechmea gustavoi J. A. Siqueira & Leme, in the short-peduncled and capituliform inflorescence, but differs in the oval to oblong floral bracts, the acuminate sepals and the green colour of all perianth structures. (© 2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)
Epiphytes make up roughly 10% of all vascular plant species globally and play important functional roles, especially in tropical forests. However, to date, there is no comprehensive list of vascular epiphyte species. Here, we present EpiList 1.0, the first global list of vascular epiphytes based on standardized definitions and taxonomy. We include obligate epiphytes, facultative epiphytes, and hemiepiphytes, as the latter share the vulnerable epiphytic stage as juveniles. Based on 978 references, the checklist includes >31,000 species of 79 plant families. Species names were standardized against World Flora Online for seed plants and against the World Ferns database for lycophytes and ferns. In cases of species missing from these databases, we used other databases (mostly World Checklist of Selected Plant Families). For all species, author names and IDs for World Flora Online entries are provided to facilitate the alignment with other plant databases, and to avoid ambiguities. EpiList 1.0 will be a rich source for synthetic studies in ecology, biogeography, and evolutionary biology as it offers, for the first time, a species‐level overview over all currently known vascular epiphytes. At the same time, the list represents work in progress: species descriptions of epiphytic taxa are ongoing and published life form information in floristic inventories and trait and distribution databases is often incomplete and sometimes even wrong. Since the epiphytic growth blends into soil‐rooted growth and vice versa, the inclusion or exclusion of particular species in the current list will sometimes be contentious. Thus, initiating a well‐founded discussion was one of the motivations for compiling this database; our list represents 31,311 hypotheses on the life form of plant species, and we welcome feedback on possible omission or erroneous inclusions. We release these data into the public domain under a Creative Commons Zero license waiver. When you use the data in your publication, we request that you cite this data paper. If EpiList 1.0 is a major part of the data analyzed in your study, you may consider inviting the EpiList 1.0 core team as collaborators.
