Article

New circumscription of Cryptanthus and new Cryptanthoid genera and subgenera (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular phylogeny

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  • Goethe-Universität Frankfurt am Main and Senckenberg Research Institute Frankfurt
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Abstract

The authors provide a new circumscription for genera of the “Cryptanthoid complex” of Bromeliaceae subfam. Bromelioideae, originally composed of Cryptanthus, Lapanthus, Orthophytum, and Sincoraea, on the basis of new or reevaluated ecological, geographical and morphological evidence, as well as molecular phylogenies. A new generic status is proposed for Cryptanthus subg. Hoplocryptanthus and two new genera, Forzzaea, and Rokautskyia, as well as four new subgenera in Orthophytum (Capixabanthus, Clavanthus, Krenakanthus, and Orthocryptanthus) are described to render taxonomic units monophyletic. The recognized taxa are well circumscribed by the combination of geographical range, ecology and morphological characters (sex distribution, leaf succulence, sepal and petal size and connation, petal appendages, pollen and stigma morphology, fruit size, calyx persistency, seed size and number per fruit). Field collected living specimens of 78 of the 81 species of Cryptanthus s.l., all species of Lapanthus, 58 of the 59 species of Orthophytum, and all species of Sincoraea were analysed in habitat and/or in cultivation, allowing the documentation and illustration of new and underutilized characters. The molecular analysis incorporated 91 accessions representing 33 species of Cryptanthus, all species (3) of Lapanthus, 42 species of Orthophytum, and 9 species of Sincoraea, including the type species for the first three genera and four outgroup taxa. The results suggest, that some morphological characters generally considered homoplasious for Bromelioideae, for the “Cryptanthoid complex”, are not homoplasious at least within the obtained, biogeographycally well delimited clades and their taxonomical utility is redeemed.

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... The species occur mainly in the Atlantic Forest phytogeographic domain; a few are found in the Caatinga (Maciel 2020). A recent phylogenetic study revealed relationships between Cryptanthus and allied genera, which resulted in a new circumscription for the genus with the transfer of some Cryptanthus species to new genera (Leme et al. 2017). Cryptanthus is characterized mainly by its terricolous or rupicolous habit, staminate and perfect (plants andromonoecious) flowers, sepals and petals partially connate, petals white or white with green or greenish apices and conduplicate-patent stigma (Ramírez-Morillo 1996, Leme et al. 2017). ...
... A recent phylogenetic study revealed relationships between Cryptanthus and allied genera, which resulted in a new circumscription for the genus with the transfer of some Cryptanthus species to new genera (Leme et al. 2017). Cryptanthus is characterized mainly by its terricolous or rupicolous habit, staminate and perfect (plants andromonoecious) flowers, sepals and petals partially connate, petals white or white with green or greenish apices and conduplicate-patent stigma (Ramírez-Morillo 1996, Leme et al. 2017). ...
... The last taxonomic revision of the genus was published by Ramírez-Morillo (1996) and included 21 species sensu Leme et al. (2017). After this revision, 37 new species were described for the genus sensu Leme et al. (2017), of which 26 occur in northeastern Brazil (Leme 1996, 1999, 2001, 2002a, 2002b, 2010, 2014, 2015, Leme & Siqueira-Filho 2001, Luther 1999a, 1999b, Leme & Siqueira-Filho 2007, Leme et al. 2008, 2010, 2014, 2020, Leme & Kollmann 2013, Braun & Brito 2019, Ferreira & Louzada 2020, Ferreira et al. 2021b, Souza & Leme 2021). ...
Article
Two new species of Cryptanthus from the Atlantic Forest of northeastern Brazil are described and illustrated. Cryptanthus pirambuensis occurs in Sergipe State and is compared to C. alagoanus, C. pickelii and C. ruthiae, from which it differs mainly by the number of flowers in the apical cluster of the inflorescence and the number of flowers in the lateral clusters of the inflorescence. Cryptanthus vinosibracteatus was discovered in Bahia State and is morphologically similar to C. arelli, but it can be differentiated by the absence of trichomes on the adaxial surface of the leaf blades, length and color of the floral bracts and sepals, and length and connation of the petals. Geographic distribution data, including a map, photographs and comments are also provided for the new species.
... The stigma type (character viii) is applied here following Barfuss et al. (2016), who presented a synthesis of all stigma types present in Tillandsioids. Although neglected for many years, the stigma type contributed significantly to improve the circumscription of historically problematic genera in Bromeliaceae (Brown and Gilmartin 1984; Gomes-da-Silva and Souza-Chies 2017; Leme et al. 2017aLeme et al. , 2017bGomes-da-Silva et al. 2019) and the understanding of the evolution of this character in Vrieseinae, composed of four types of stigma, becomes relevant. ...
... However, in the combined BI tree, the tribe Vrieseeae was not recovered as monophyletic, with the subtribe Cipurosidinae emerging as sister to the tribe Tillandsieae. The nuclear gene PHYC showed high polymorphism, and showed better resolution when compared to the other markers, in agreement with other studies of Bromeliaceae (Jabaily and Sytsma 2010;Krapp et al. 2014;Louzada et al. 2014;Leme et al. 2017b;Maciel et al. 2018;Matuszak-Renger et al. 2018), especially of the subfamily Tillandsioideae (Barfuss et al. 2016;Leme et al. 2017a). Recently, Loiseau et al. (2021) argued that ancestral hybridization between Vriesea s.s. and Tillandsieae genera could be responsible for both hypotheses of relationships of Cipuropsidinae, depending on whether the plastidial or nuclear data is considered. ...
... The species of Stigmatodon s.s. are characterized by the presence of a tubo-laciniate stigma, which is also synapomorphic for the genus (Barfuss et al. 2016; Gomes-da-Silva and Souza-Chies 2017; Leme et al. 2017a); while the conduplicate types (patent and erect) are synapomorphic for Alcantarea (Leme 2007(Leme , 2009; Barfuss et al. 2016;Leme et al. 2017a) (Fig. 4D). The stigma type has proven to be taxonomically powerful when used in combination with other morphological, ecological and geographical characters (Barfuss et al. 2016;Leme et al. 2017aLeme et al. , 2017b, and contributed significantly to improve the circumscription of historically problematic genera in Bromeliaceae as a whole (Leme et al. 2017b;Gomesda-Silva et al. 2019). ...
... The "Cryptanthoid" genera share ecological, geographical, and morphological characteristics such as endemic occurrence in southeastern and northeastern Brazil, terrestrial or saxicolous habit, leaf rosettes without waterholding capacity, sessile flowers usually arranged in subsessile fascicles, and fruits without noticeable mucilaginous substance (Leme et al. 2013, Leme 2015, Leme et al. 2017b). Its current circumscription was proposed in a recent monograph (Leme et al. 2017b) in which an integrative approach was applied, combining data of biogeography, morphology (traditionally used characters plus new or underutilized traits) together with a molecular phylogeny, to forward new generic and subgeneric delimitations of monophyletic groups. ...
... The "Cryptanthoid" genera share ecological, geographical, and morphological characteristics such as endemic occurrence in southeastern and northeastern Brazil, terrestrial or saxicolous habit, leaf rosettes without waterholding capacity, sessile flowers usually arranged in subsessile fascicles, and fruits without noticeable mucilaginous substance (Leme et al. 2013, Leme 2015, Leme et al. 2017b). Its current circumscription was proposed in a recent monograph (Leme et al. 2017b) in which an integrative approach was applied, combining data of biogeography, morphology (traditionally used characters plus new or underutilized traits) together with a molecular phylogeny, to forward new generic and subgeneric delimitations of monophyletic groups. The data at hand and the monophyly of the "Cryptanthoid complex" allowed to assign generic and subgeneric rank to the several monophyletic groups, which were well-circumscribed biogeographically, morphologically and genetically. ...
... When measures provided in the protologue were not confirmed in the studied material, values are indicated inside parentheses. The descriptive terminology follows Smith & Downs (1974, 1977, 1979, with adaptations suggested by Scharf & Gouda (2008), and definitions used by Barfuss et al. (2016) and Leme et al. (2017b) to avoid confusion resulting from an inconsistent application by previous authors (e.g. definition of lifestyle, habit, petal appendages, stigma types). ...
Article
The continuous refinement of the morphological, anatomical, and molecular studies on the genera of the “Cryptanthoid complex”, mainly considering the recently discovered outlier species, gave rise to new interpretations of species relationships. It reinforced the monophyly of the groups, and allowed the recognition of the new monotypic genus Siqueiranthus based on a morphologically unique endemic species from northeastern Brazil on the brink of extinction, as well as generic status for Krenakanthus and Orthocryptanthus previously conceived as subgenera of Orthophytum, with readjustments in the positioning of some species. An integrative approach with the combination of multiple traits and the critical assessment of the diagnostic characters proved to be useful for the establishment of the new monophyletic genera. A new unusual species of Orthophytum is also described and illustrated, and its phylogenetic relationships are discussed.
... updated). The morphological circumscription of the genus was narrowed in the revision proposed by Leme et al. (2017), in order to encompass only andromonoecious plants with compound or rarely pseudosimple, sessile, and shortly corymbose inflorescences (fertile part) with inconspicuously stipitate basal/outer flower fascicles, with flowers bearing unappendaged, usually white, rarely green or greenish petals, which are sublinear-lanceolate to narrowly spathulate, arcuate to recurved at anthesis and distinctly exposing the stamens. All Cryptanthus species have sulcate and spherical pollen, with reticulate exine. ...
... The morphological descriptions and illustrations are based on living fertile material using a stereomicroscope before herborization. The descriptive terminology follows Smith & Downs (1979), with adaptations suggested by Scharf & Gouda (2008), and definitions proposed by Leme et al. (2017) for the genera of the "Cryptanthoid complex". Photographs were taken as indicated in the captions. ...
... Latin abbreviations used in nomenclature follow the Shenzhen Code (Turland et al. 2018) andHawksworth (2010). The affinity/distinctness of the species was established based on the most relevant characters selected by Leme et al. (2017). ...
Article
The authors describe and illustrate four new species of Cryptanthus: C. flesherii, C. lutandensis, C. santosii, and C. solidadeanus. The affinities and distinctness of the new species are supported by the combination of geographical distribution, ecological preference, and morphological characters regarded as the most useful to distinguish taxa in "Cryptanthoid complex". All described species were recovered as critically endangered. A distribution map as well as taxonomic comments are presented.
... 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. ...
... Biogeography. The relevance of biogeography in detecting evolutionary lineages is evident in Bromeliaceae (see e.g., Leme et al. 2017). According to Sass & Specht (2010), the distinct pattern of geographical distribution is important because the apparently homoplastic morphological characters used to characterize species, subgenera or genera may be useful taxonomically when biogeography is also considered. ...
... For the first time, they named a "capituliform" type for species with simple, compact inflorescences. In the review of the genera of the "Cryptanthoid complex", Leme et al. (2017) adopted Mez's (1981Mez's ( , 1934Mez's ( -1935 and most of Louzada's (2012) inflorescence classification for Orthophytum Beer (1854: 347), to recognize the sessile and compound, usually compact, and shortly corymbose inflorescence type principally observed in the group, with subsessile or shortly stipitate, more or less complanate and flabellate, sometimes pulvinate fascicles, which, not considering the non-impounding leaf rosettes and the foliaceous primary bracts of these, is an inflorescence model similar to Hylaeaicum. ...
Article
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.
... The late ontogenetic development of petal appendages as well as the repeated loss and/or gain without affecting the function of other floral organs (Brown and Terry, 1992;Schulte and Zizka, 2008) illustrate their unreliability as a taxonomical character. However, to delimitate related species groups they might be useful tools in combination with other characters (Brown and Terry, 1998;Barfuss et al., 2005;Louzada et al., 2014;Leme et al., 2017). According to Schulte and Zizka (2008) the absence of petal appendages is plesiomorphic for Bromelioideae and most of the early diverging lineages are lacking these, while as the eu-Bromelioideae exhibit a heterogeneous picture. ...
... shape, length, width, intertwining, twisting, papillae). Indeed, further stigma types were described by Leme and Brown (2004), Leme (2007aLeme ( , 2009, Barfuss et al. (2016) and Leme et al. (2017). The new studies of Barfuss et al. (2016) and Leme et al. (2017) considered the stigma types as highly specific and valuable diagnostic character. ...
... Indeed, further stigma types were described by Leme and Brown (2004), Leme (2007aLeme ( , 2009, Barfuss et al. (2016) and Leme et al. (2017). The new studies of Barfuss et al. (2016) and Leme et al. (2017) considered the stigma types as highly specific and valuable diagnostic character. However a precise interpretation of stigma morphology requires optimal conservation, and caution due to structural changes during the development. ...
Thesis
Full-text available
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. 5 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 6 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.
... Agt1 encodes a glyoxylate aminotransferase that is involved in the photorespiration pathway in Arabidopsis [15][16][17] and the locus was suggested as a marker for phylogenetic studies at low taxonomic levels [18]. Since then Agt1 has been successfully used in a number of phylogenetic studies covering a wide range of angiosperm plant groups [19][20][21][22][23][24][25]. In two recent studies that aimed at a revision of the "Cryptanthoid complex" [23] and the genus Ananas, including its closest relatives [25], Agt1 proved to be of great value for the reconstruction of Bromeliaceae genus/subgenus level phylogenies and thus prompted a further investigation of its potential suitability for barcoding. ...
... Since then Agt1 has been successfully used in a number of phylogenetic studies covering a wide range of angiosperm plant groups [19][20][21][22][23][24][25]. In two recent studies that aimed at a revision of the "Cryptanthoid complex" [23] and the genus Ananas, including its closest relatives [25], Agt1 proved to be of great value for the reconstruction of Bromeliaceae genus/subgenus level phylogenies and thus prompted a further investigation of its potential suitability for barcoding. ...
... The Agt1 marker region used for this study was initially amplified as described in previous studies [18,23]. As double bands occurred at high frequency, we designed bromeliad specific primers with higher annealing temperatures, to attain increased specificity and to impede the occurrence of unspecific bands. ...
Article
Full-text available
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.
... The majority of the AE55 species has been described in the past 20 years or so (Leme & al. 2017). There are two centres of diversity, one in the Serra do Espinhaço Range in Bahia and Minas Gerais, the other in the inselberg regions of Minas Gerais and Espírito Santo. ...
... Traditionally, Orthophytum was divided into 2 main groups, based on whether the inflorescence is sessile or long-pedunculate, and Leme (2004) provided a further break-down into 6 "subcomplexes". These groupings were only partly recovered in the molecular phylogenies of Louzada & al. (2014) and Leme & al. (2017). Based on their results, Leme & al. (2017) divide Orthophytum into 5 subgenera, but only Subgen. ...
... These groupings were only partly recovered in the molecular phylogenies of Louzada & al. (2014) and Leme & al. (2017). Based on their results, Leme & al. (2017) divide Orthophytum into 5 subgenera, but only Subgen. Orthophytum (in terminal sister-position to Subgen. ...
Chapter
Orthophytum Beer (Flora 37: 347, 1854). Type: Prantleia glabra Mez [typification according to L. B. Smith & Downs, Fl. Neotrop. 14(3): 1696, 1979]. — Bromelioideae — Lit: Smith & Downs (1979: 1696-1710, Fl. Neotropica); Leme (2008: key to the mello-barretoi-complex); Louzada & al. (2010: cytology); Louzada & Wanderley (2010: partial monograph of the vagans-clade); Louzada (2012: unpublished monograph, with key); Louzada & al. (2014: molecular phylogeny); Leme & al. (2017: classification). Distr: E Brazil; Atlantic Forest, Cerrado and Caatinga vegetations, usually saxicolous. Etym: Gr. ‘orthos’, erect, straight; and Gr. ‘phyton’, plant; for the erect stems with well-spaced leaves of the type species.
... were not recovered as a monophyletic group, although the segregation of the species of the genus in different clusters did not receive statistical support. In a more comprehensive phylogenetic analysis based on DNA data for Cryptanthus, Cruz et al. (2017) clarified the infrageneric relationships, suggesting that subgenus Hoplocryptanthus is paraphyletic, and Leme et al. (2017), based on morphological features and molecular data, proposed the recognition of subgenus Cryptanthus as Cryptanthus s.s. and divided subgenus Hoplocryptanthus into three new genera, Hoplocryptanthus (Mez) Leme et al. (2017) referred to these genera as the cryptanthoid complex. ...
... were not recovered as a monophyletic group, although the segregation of the species of the genus in different clusters did not receive statistical support. In a more comprehensive phylogenetic analysis based on DNA data for Cryptanthus, Cruz et al. (2017) clarified the infrageneric relationships, suggesting that subgenus Hoplocryptanthus is paraphyletic, and Leme et al. (2017), based on morphological features and molecular data, proposed the recognition of subgenus Cryptanthus as Cryptanthus s.s. and divided subgenus Hoplocryptanthus into three new genera, Hoplocryptanthus (Mez) Leme et al. (2017) referred to these genera as the cryptanthoid complex. ...
... The phylogenetic tree used in the reconstruction of ancestral character states was based on the AFLP data obtained by Cruz et al. (2017). For the tree reconstruction, we considered Hoplocryptanthus as the earliest-diverging group, according to Leme et al. (2017). The maximum likelihood tree with all previously analysed accessions was adjusted by trimming the species without any information on genome size in Mesquite v.3.5 (Maddison & Maddison, 2018). ...
Article
We describe the chromosome numbers and genome sizes of species of the cryptanthoid complex of Bromeliaceae in a phylogenetic framework and their relationship with habitat preferences. The 2C DNA contents varied 2.13-fold among species, ranging from 0.76 to 1.66 pg. A significant difference in DNA content was found among Cryptanthus, Hoplocryptanthus and Rokautskyia. Moreover, species from campos rupestres and the Atlantic Forest had lower and higher genome size values, respectively. The smaller genome sizes of Hoplocryptanthus spp. from campos rupestres may be related with the large genome constraint. The species show a highly conserved ploidy (with 2n = 32 and 34), although the genome sizes varied considerably. The observed variation in chromosome numbers seems to be influenced by dysploidy, but additional investigations are needed. Our study demonstrates that the genome size variation in the cryptanthoid complex species is not strictly related to the phylogenetic relationships and has probably been influenced by different evolutionary processes.
... Currently, there are new approaches aiming to integrate molecular data with geographical distribution data and previously neglected or misunderstood morphological features (e.g. presence of chalazal ovule appendage, stigma type or calyx persistence) (Cruz et al., 2017;Leme et al., 2017;Aguirre-Santoro et al., 2016). It becomes clear that, despite the possible e ects of convergence, reassessing the available morphological characters and recognizing up to now neglected characters of potential phylogenetical value will be necessary for a full understanding of the systematics and evolution of Bromeliaceae. ...
... c b a does not have pollen in tetrads. is expands the morphological limits of the genus, but on the other hand, puts in doubt the reliability of the characters used for its de nition. Androlepis fragrans is sympatric with other species of the clade, which suggest a possible relationship, according to previously observed geographical tendencies on the subfamily (Sass and Specht, 2010;Aguirre-Santoro et al., 2015;Leme et al., 2017). However, its phylogenetic position inside the clade, including its placement within Androlepis, needs to be evaluated using molecular and morphological data. ...
... In the opposite case, in widely distributed species, such as Aechmea mexicana, the study of the biological entity throughout its distribution range requires a great sampling e ort (not only to consult herbarium specimens, but also to obtain DNA samples). e most recent Bromelioideae taxonomic updates have highlighted the utility of some oral characters in the delimitation of genera, such as stigma type, appendage architecture, ower compression and chalazal appendages (Aguirre-Santoro et al., 2016;Aguirre-Santoro, 2017, 2018Leme et al., 2017). According to our current observations and the revision of literature (Brown & Gilmartin, 1984, 1989Leme et al., 2017), stigma type in species such as A. mexicana, A. lueddemanniana and Androlepis skinneri are conduplicate-spiral, whereas both Ursulaea and Billbergia viridi ora exhibit a conduplicate-patent stigma type. ...
... Cryptanthus is endemic to Brazil and comprises 55 species distributed through the Atlantic Forest, Caatinga and Cerrado, with distribution concentrated in the Southeast and Northeast regions (Leme & Siqueira-Filho 2006, Leme et al. 2017, Flora do Brasil 2020. Only one species occurs in the state of RN, restricted to fragments of Atlantic Forest. ...
... This genus has notably two centers of diversity: the rocky outcrops of the Espinhaço Range (BA and MG) and inselbergs in the Atlantic Forest of MG and ES (Wanderley 1990, Louzada & Wanderley 2010, Gouda & Butcher 2018, Flora do Brasil 2020. Based on morphologic and molecular phylogenic studies, current classification supports the subdivision of Orthophytum in five subgenera (Leme et al. 2017), as well as the segregation of several species to the re-established genus Sincoraea (Louzada & Wanderley 2016). In RN, the genus is represented by only one species. ...
... Comments:-Orthophytum disjunctum was part of the traditional "pedunculate inflorescence group" (Louzada & Wanderley 2010, Louzada et al. 2014 and is now placed under Orthophytum subg. Orthophytum (Leme et al. 2017). The leaves with densely white-lepidote faces, triangular and conspicuously serrate blades can distinguish this species, as well as the foliaceous peduncle bracts and primary bracts and the pedunculate compound inflorescence. ...
Article
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The Bromeliaceae Flora for the state of Rio Grande do Norte, Northeastern Brazil, is presented, based on extensive fieldwork, morphological analyses using herbarium and freshly collected material, and specialized literature. Twenty-six species of bromeliads were recorded in Rio Grande do Norte, distributed in ten genera and in three subfamilies. Bromelioideae was the richest subfamily (eight genera/14 species), followed by Tillandsioideae (one genus/12 species), and Pitcairnioideae (one genus/one species). Aechmea mertensii, Hohenbergia horrida and Tillandsia tenuifolia are new records for Rio Grande do Norte. Eight species (31%) are restricted to the Eastern portion of the state, in the Atlantic Forest. Caatinga dry woodlands harbor 18 species, with remarkable presence of Bromelia laciniosa, Encholirium spectabile, Tillandsia recurvata and T. streptocarpa, the four most widely distributed taxa. We discuss problems related to unclear taxonomic circumscriptions of species or diverging information between authors, more expressively in Hohenbergia, but also in Aechmea, Cryptanthus and Tillandsia. The data presented here might contribute to better understand the morphological variation of these taxa and suggest additional research on their taxonomy. Morphological descriptions, general comments, a map, photo plates and an identification key for all taxa are provided.
... The majority of the AE55 species has been described in the past 20 years or so (Leme & al. 2017). There are two centres of diversity, one in the Serra do Espinhaço Range in Bahia and Minas Gerais, the other in the inselberg regions of Minas Gerais and Espírito Santo. ...
... Traditionally, Orthophytum was divided into 2 main groups, based on whether the inflorescence is sessile or long-pedunculate, and Leme (2004) provided a further break-down into 6 "subcomplexes". These groupings were only partly recovered in the molecular phylogenies of Louzada & al. (2014) and Leme & al. (2017). Based on their results, Leme & al. (2017) divide Orthophytum into 5 subgenera, but only Subgen. ...
... These groupings were only partly recovered in the molecular phylogenies of Louzada & al. (2014) and Leme & al. (2017). Based on their results, Leme & al. (2017) divide Orthophytum into 5 subgenera, but only Subgen. Orthophytum (in terminal sister-position to Subgen. ...
... Cryptanthus is monophyletic and includes 59 species, restricted to the north-eastern and southeastern regions of Brazil (Leme et al., 2017;Leme et al., 2020). Most of its species occur in the Brazilian Atlantic Forest, and a few occur in the caatinga (Leme et al., 2017). ...
... Cryptanthus is monophyletic and includes 59 species, restricted to the north-eastern and southeastern regions of Brazil (Leme et al., 2017;Leme et al., 2020). Most of its species occur in the Brazilian Atlantic Forest, and a few occur in the caatinga (Leme et al., 2017). Cryptanthus burle-marxii Leme and C. zonatus (Vis.) ...
Article
Morphological variations of individuals and populations of plants have hampered taxonomists from understanding whether such variations are intra- or interspecific. In this research, we study Cryptanthus burle-marxii and C. zonatus, the morphological variations of which overlap, making it difficult to identify them. Both taxa are restricted to the north of the Brazilian Atlantic Forest and are included in the C. zonatus species complex. We applied different methods such as morphological comparison, population genetics using ten nuclear microsatellite markers and ecological niche modelling to study population genetic structure and species delimitation of this species complex. Our analysis revealed that the two genetic clusters were formed possibly because of the presence of a historical geographical barrier in a region called ‘depressão do Abiaí’. The two genetic clusters were concordant with the northern and southern distribution of the complex but incongruent with a morphological variation or current taxonomic delimitation. Thus, we synonymize C. burle-marxii with C. zonatus and re-evaluate its conservation status as endangered.
... However, classification systems as a whole and taxonomy as routinely practised in herbaria have largely been based on floral characters (Stace, 1989;Cutler, Botha & Stevenson, 2008;Mendes et al., 2012;Albert et al., 2014). Specifically in bromeliads, the lack of in-depth knowledge of floral morphology has been pointed out by different researchers as a limitation to the correct understanding not only of morphology itself, but also of systematics and ecology, since much of the research that has been done has been based on flowers of herbarium material, with poor preservation of intracalyx structures (Brown & Gilmartin, 1984;Kuhn et al., 2016;Leme et al., 2017). However, floral morphology and particularly detailed analyses of stamens, stigmas, flower shapes and petal appendages has been shown to be fundamental in the delimitation of taxa, such as new genera (e.g. ...
... However, floral morphology and particularly detailed analyses of stamens, stigmas, flower shapes and petal appendages has been shown to be fundamental in the delimitation of taxa, such as new genera (e.g. Louzada & Versieux, 2010;Barfuss et al., 2016;Carvalho et al., 2016;Leme et al., 2017). ...
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.
... The description is based on living material, herbarium specimens, flowers preserved in liquid, field data and photographs. Descriptive terminology is based on Smith & Downs (1979), with amendments from Scharf & Gouda (2008) for inflorescence structures, Leme et al. (2017) for the description of petal appendages, and Halbritter et al. (2018) for pollen morphology. For the observation of micromorphological structures, anthers, pollen, and stigmas, samples were coated in high vacuum with palladium-gold with a DENTON VACUUM DESK II sputter system, and subsequently observed in a JOEL Scanning Electron Microscope JSM-6360LV. ...
... The morphological characters traditionally used in generic delimitations, such as inflorescence branching, presence/absence of petal appendages, and sepal symmetry, have apparently evolved independently several times within the subfamily (Schulte & Zizka 2008, Sass & Specht 2010, Aguirre-Santoro et al. 2016 and are, therefore, problematic. Recent systematic studies within the subfamily have disclosed the taxonomic relevance of some ignored characters, such as petal appendage type, size and number of seeds per fruit, among others (Schulte & Zizka 2008, Aguirre-Santoro et al. 2016, Leme et al. 2017. For this reason, the generic assignment of the new taxon is based upon characters that have been rarely used in Bromelioideae: dioecy and the distal filament ornamentation close to the connection with the anther. ...
Article
Androlepis najarroi (Bromelioideae, Bromeliaceae), a third dioecious species identified in the subfamily, is proposed as new from the Chiapas Highlands and Veracruzan biogeographical provinces, southern Megamexico. The novelty is superficially similar to Aechmea lueddemanniana but features characteristics of Androlepis, such as unisexual flowers and apically divided filaments. Based upon available information on the species and current generic concepts, assigning this species to Androlepis might only be provisional. The present contribution includes distribution maps, iconography, an identification key to the species of Androlepis, and a preliminary evaluation of the conservation status of this new species.
... Descriptive terminology follows Smith & Downs (1979), with adaptations suggested by Scharf & Gouda (2008). Definitions for the genera of the "Cryptanthoid" complex follow Leme et al. (2017) Specimens were herborized according to Fidalgo & Bononi (1989), and deposited in herbaria HVASF and HUEFS. Live specimens were introduced for cultivation at the Coleção de Plantas Vivas do Vale do São Francisco (Vivasf) in Petrolina, state of Pernambuco, and in Refúgio dos Gravatás, in Teresópolis, state of Rio de Janeiro, following guidelines recommended by the Convention on Biological Diversity (1992). ...
Article
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A new species of Cryptanthus, known exclusively from Chapada Diamantina, in the municipalities of Miguel Calmon and Jacobina, state of Bahia, Northeast Brazil, is described and illustrated. Cryptanthus euglossinii is characterized by having leaves that are reddish in the marginal region, white-scaly near the base and glabrous adaxially towards the apex, with the abaxial surface covered by trichomes that obscure the color of the leaf. Comparisons are made with two similar species, C. reisii and C. bibarrensis, that are considered closely related. Euglossinii bees were observed visiting the flowers of C. euglossinii and collecting floral essences on petals, a relationship that is the basis of the epithet of the new species. Cryptanthus euglossinii is considered Endangered (EN) based on an extent of occurrence of 860 km2, an area of occupancy of 1,000 km2, and criteria established by the IUCN.
... Pleroma miconiifolium is rupicolous, growing on shallow pockets of soil over exposed rock at elevations around 1000 m. At the type locality, the species is syntopic with another endemic and recently described species, the bromeliad Orthophytum santaritense Leme, S.Heller & Zizka (Bromeliaceae; Leme et al., 2017). ...
Article
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Eastern Minas Gerais has been historically neglected regarding biodiversity sampling effort. However, recent botanical explorations have revealed several new taxa for its flora, especially from disjunct fragments of campos rupestres vegetation, which form a mosaic with granitic inselbergs in the region. In this article, we add four new species of Pleroma (Melastomataceae) to the growing list of new taxa: P. brevicomosum, P. caetanoi, P. miconiifolium and P. petrophylax. We provide descriptions, detailed photographs, taxonomic comparisons, a distribution map and conservation status assessments for these new taxa. All four species are assessed under a category of threat, highlighting the need for conservation actions in these biodiverse albeit still poorly known areas.
... 3.3.17. Poales: Bromeliaceae, Cyperaceae, Eriocaulaceae, Juncaceae, Poaceae, Rapateaceae Bromeliaceae are well known for forming plantlets in their inflorescences (Leme et al., 2017). There are several species of Cyperaceae and Juncaceae that are pseudoviviparous (Leck and Leck, 2005). ...
Article
Aerial vegetative diaspores — often simply called “bulbils” in the botanical and horticultural literature — are diverse in morphology, origin, position, and modes of dispersal. This review examines their occurrence in the Angiosperms, their morphology and site of origin, and possible ecological and evolutionary advantages. Moreover, a standard terminology is proposed based on three criteria: dormancy, polarity, and whether the storage tissue is leaf or stem. We review the taxonomic and geographic distribution of bulbils, cormlets, tubercles, and gemmae, their modes of dispersal, and whether dispersal mode differs from that of the seeds. We detect geographic biases in the distribution of plantlets and tubercles (mostly tropical in distribution) versus bulbils, cormlets, and gemmae (mostly temperate). We note the physiological differences between seeds and aerial vegetative diaspores, which may account for differences in modes of dispersal, which for aerial vegetative diaspores includes anemochory, epizoochory, endozoochory, barochory, and hydrochory. Additional research is suggested so that gaps in our understanding of this common form of asexual reproduction can be filled.
... 76 Cryptanthus burle-marxii and other congenerics are andromonoecious, a derived character among Bromeliaceae that is exclusive to this genus. 10,77 The six stamens of both staminate and hermaphrodite flowers extend from the center of the floral axis, raising the anthers symmetrically toward all directions above the nectariferous chamber. This builds a clear actinomorphic symmetry in male flowers. ...
Article
Perfume flowers (sensu Vogel) produce intense scents that function both as attractants and as the sole rewards for pollinators. The scent is collected exclusively by male euglossine bees and used during pre-mating behavior. Perfume flowers have evolved independently in 15 angiosperm families, with over 1,000 reported species across the Neotropical region. Members of Cryptanthus (Bromeliaceae) represent a puzzling exception among perfume flowers, as flowers produce nectar and do not emit a noticeable scent yet still attract euglossine males. Here, we studied the pollination ecology of Cryptanthus burle-marxii and decode the chemical communication between its flowers and euglossine males. Field observations revealed euglossine males and hummingbirds as potential pollinators. The bees always contacted anthers/stigma of C. burle-marxii while scraping the petals to obtain chemicals, whereas nectar-seeking hummingbirds normally only contacted the anthers. Based on gas chromatography-mass spectrometry/nuclear magnetic resonance analyses of flower scent samples and bioassays, we identified the diterpene copalol as the only floral scent compound triggering scent-gathering behavior in euglossine males. Unlike euglossine-bee-mediated pollination, hummingbird pollination is ancestral in the Cryptanthus clade, suggesting a case of an ongoing pollinator shift mediated by the evolution of perfume as a reward. Copalol was previously unknown as a floral scent constituent and represents the heaviest and least-volatile compound known to attract euglossine males. Our study provides the first experimental evidence that semivolatile floral compounds can mediate euglossine bee interactions. Male euglossine pollination in other plant species lacking noticeable floral scents suggests that semivolatile- mediated pollinator attraction is more widespread than currently appreciated.
... As a result of the low rates of substitution in plastid regions, the tree topologies are often poorly resolved not helping in the circumscription of some genera in Bromeliaceae (Smith & Donoghue, 2008;Maia et al., 2012;Palma-Silva et al., 2016); non-natural genera are frequently reported in Bromeliaceae (Barfuss et al., 2005;Evans et al., 2015;Schütz et al., 2016;Maciel et al., 2018) and only 30% of the currently recognized genera are monophyletic (Escobedo-Sarti et al., 2013). To overcome these problems, recent studies have increased the amount of analysed data either by increasing the number of molecular markers and their phylogenetic informativeness (e.g. more quickly evolving nuclear regions or even AFLP) or increasing the numbers of terminals (Krapp et al., 2014;Heller et al., 2015;Barfuss et al., 2016;Pinangé et al., 2016;Schütz et al., 2016;Goetze et al., 2017;Gomes-da-Silva & Souza-Chies, 2017;Leme et al., 2017;Maciel et al., 2018;Matuszak-Renger et al., 2018). Nevertheless, large and complex to delimit genera such as Tillandsia L. (741 species), Pitcairnia L'Heritier (408 species), Vriesea Lindl. ...
Article
Vriesea is the second largest genus in Tillandsioideae, the most diverse subfamily of Bromeliaceae. Although recent studies focusing on Tillandsioideae have improved the systematics of Vriesea, no consensus has been reached regarding the circumscription of the genus. Here, we present a phylogenetic analysis of core Tillandsioideae using the nuclear gene phyC and plastid data obtained from genome skimming. We investigate evolutionary relationships at the intergeneric level in Vrieseeae and at the intrageneric level in Vriesea s.s. We sampled a comprehensive dataset, including 11 genera of Tillandsioideae and nearly 50% of all known Vriesea spp. Using a genome skimming approach, we obtained a 78 483-bp plastome alignment containing 35 complete and 55 partial protein-coding genes. Phylogenetic trees were reconstructed using maximum-likelihood based on three datasets: (1) the 78 483 bp plastome alignment; (2) the nuclear gene phyC and (3) a concatenated alignment of 18 subselected plastid genes + phyC. Additionally, a Bayesian inference was performed on the second and third datasets. These analyses revealed that Vriesea s.s. forms a well-supported clade encompassing most of the species of the genus. However, our results also identified several remaining issues in the systematics of Vriesea, including a few species nested in Tillandsia and Stigmatodon. Finally, we recognize some putative groups within Vriesea s.s., which we discuss in the light of their morphological and ecological characteristics.
... Nonetheless, the subfamily can be divided into two major groups: several informal early diverging tank-less lineages and mostly tank-forming core bromelioids (Schulte et al., 2009;Silvestro et al., 2014). The tank-less lineages are terrestrial or lithophytic usually characterized by succulent leaves with considerable water storage tissue and CAM photosynthesis, except the earliest diverging Greigia, Ochagavia, Fascicularia, as well as several other species from the Cryptanthoid complex and the genus Fernseea, which are C3 (Horres and Zizka, 1995;Silvestro et al., 2014;Crayn et al., 2015;Leme et al., 2017). In the tank-forming lineages of the core Bromelioideae, one large and/ or several small tanks are formed by the tightly clasping leaf sheaths, which can thus impound up to several liters of water in larger plants (e.g. ...
Article
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The subfamily Bromelioideae is one of the most diverse groups among the neotropical Bromeliaceae. Previously, key innovations have been identified which account for the extraordinary radiation and species richness of this subfamily, especially in the so-called core Bromelioideae. However, in order to extend our understanding of the evolutionary mechanisms, the genomic mechanisms (e.g. polyploidy, dysploidy) that potentially underlie this accelerated speciation also need to be tested. Here, using PI and DAPI staining and flow cytometry we estimated genome size and GC content of 231 plants covering 30 genera and 165 species and combined it with published data. The evolutionary and ecological significance of all three genomic characters was tested within a previously generated dated phylogenetic framework using ancestral state reconstructions, comparative phylogenetic methods, and multiple regressions with climatic variables. The absolute genome size (2C) of Bromelioideae varied between 0.59 and 4.11 pg, and the GC content ranged between 36.73 and 41.43%. The monoploid genome sizes (Cx) differed significantly between core and early diverging lineages. The occurrence of dysploidy and polyploidy was, with few exceptions, limited to the phylogenetically isolated early diverging tank-less lineages. For Cx and GC content Ornstein-Uhlenbeck models outperformed the Brownian motion models suggesting adaptive potential linked to the temperature conditions. 2C-values revealed different rates of evolution in core and early diverging lineages also related to climatic conditions. Our results suggest that polyploidy is not associated with higher net diversification and fast radiation in core bromelioids. On the other hand, although coupled with higher extinction rates, dysploidy, polyploidy, and resulting genomic reorganizations might have played a role in the survival of the early diverging bromelioids in hot and arid environments.
... On the Minas Gerais side of the Espinhaço Mountains, Versieux et al. (2008) listed 141 species in 23 genera, totaling 45.9% of the species recorded for the state. This list is increasing progressively through the different floristic inventories of new areas (e.g., Encholirium viride Braun & Esteves 2017; Orthophytum santaritense Leme et al. 2017;Dyckia nobilis Buneker et al. 2016;and Dyckia sulcata Guarçoni et al. 2014). According to Versieux et al. (2008), 62% of the endemic species of Bromeliaceae of Minas Gerais are restricted to the Espinhaço range, the most important area of endemism for the family in the state (Versieux & Wendt 2007). ...
Article
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An illustrated checklist of the species of Bromeliaceae from the Serra do Cabral, Minas Gerais state, southeastern Brazil is presented, including 20 species in 11 genera from three subfamilies. Subfamilies Tillandsioideae and Bromelioideae each contain eight species, which mainly inhabit rocky fields. Tillandsia is the most diverse genus, with six species. Subfamily Pitcairnioideae contains four species, including Dyckia spinulosa, as well as Dyckia oscari, a new species described and illustrated herein.
... As a result of the low rates of substitution in plastid regions, the tree topologies are often poorly resolved not helping in the circumscription of some genera in Bromeliaceae (Smith & Donoghue, 2008;Maia et al., 2012;Palma-Silva et al., 2016); non-natural genera are frequently reported in Bromeliaceae (Barfuss et al., 2005;Evans et al., 2015;Schütz et al., 2016;Maciel et al., 2018) and only 30% of the currently recognized genera are monophyletic (Escobedo-Sarti et al., 2013). To overcome these problems, recent studies have increased the amount of analysed data either by increasing the number of molecular markers and their phylogenetic informativeness (e.g. more quickly evolving nuclear regions or even AFLP) or increasing the numbers of terminals (Krapp et al., 2014;Heller et al., 2015;Barfuss et al., 2016;Pinangé et al., 2016;Schütz et al., 2016;Goetze et al., 2017;Gomes-da-Silva & Souza-Chies, 2017;Leme et al., 2017;Maciel et al., 2018;Matuszak-Renger et al., 2018). Nevertheless, large and complex to delimit genera such as Tillandsia L. (741 species), Pitcairnia L'Heritier (408 species), Vriesea Lindl. ...
Article
Vriesea is the second largest genus in Tillandsioideae, the most diverse subfamily of Bromeliaceae. Although recent studies focusing on Tillandsioideae have improved the systematics of Vriesea, no consensus has been reached regarding the circumscription of the genus. Here, we present a phylogenetic analysis of core Tillandsioideae using the nuclear gene phyC and plastid data obtained from genome skimming. We investigate evolutionary relationships at the intergeneric level in Vrieseeae and at the intrageneric level in Vriesea s.s. We sampled a comprehensive dataset, including 11 genera of Tillandsioideae and nearly 50% of all known Vriesea spp. Using a genome skimming approach, we obtained a 78 483-bp plastome alignment containing 35 complete and 55 partial protein-coding genes. Phylogenetic trees were reconstructed using maximum-likelihood based on three datasets: (1) the 78 483 bp plastome alignment; (2) the nuclear gene phyC and (3) a concatenated alignment of 18 subselected plastid genes + phyC. Additionally, a Bayesian inference was performed on the second and third datasets. These analyses revealed that Vriesea s.s. forms a well-supported clade encompassing most of the species of the genus. However, our results also identified several remaining issues in the systematics of Vriesea, including a few species nested in Tillandsia and Stigmatodon. Finally, we recognize some putative groups within Vriesea s.s., which we discuss in the light of their morphological and ecological characteristics.
... Using a much larger dataset than in all previous studies, we aim to test the phylogenetic placement of O. elegans and to assess the relationships among other taxa in the FasciculariaOchagavia group. Additionally, we include new data on pollen morphology, which has been demonstrated to be particularly useful in taxonomy of Bromelioideae (Halbritter 1992;Leme et al., 2017), and discuss evolutionary, systematic and nomenclatural consequences of our results. ...
Article
Ochagavia (four species) and Fascicularia (one species) form a well-supported clade of the early-diverging Bromelioideae. The two genera are morphologically similar, but they can be easily discerned on the basis of generative characters. Besides the species distributed on the Chilean mainland, the group includes O. elegans, endemic to the Robinson Crusoe Island of the Juan Fernández Islands. In previous molecular phylogenetic studies, O. elegans formed a sister clade to the remainder of Fascicularia and Ochagavia. A phylogenomic approach, including nearly complete and, in five cases, full plastomes (c. 160 kbp) and the nuclear rDNA cistron (c. 6 kbp), and scanning electron microscope (SEM) images of pollen were used to analyse relationships in the Fascicularia-Ochagavia group. Plastome and nuclear trees were largely congruent and supported previous phylogenetic analyses of O. elegans being sister to the remainder of the group. A divergent phylogenetic position was suggested for O. carnea using different organellar trees. SEM analysis of pollen supported the division of Fascicularia and Ochagavia. Evolutionary and taxonomic implications of our results are discussed.
... focused on individual taxa (Canela et al., 2003;Leme, Heller, Zizka, & Halbritter, 2017;Peters, 2009;Zizka, Horres, Nelson, & Weising, 1999;Zizka, Trumpler, & Zöllner, 2002) or geographic regions (Cáceres, 2012;Judith et al., 2013;Zizka et al., 2009; www.flora dobra sil.jbrj.gov.br). Our distribution maps are available in Appendix S2, and we supply all species ranges under a CC-BY license via the bromeliad r package, which also includes functions for publication-level species richness maps for individual genera, traits or conservation categories (Appendix S4). ...
Article
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Aim To provide distribution information and preliminary conservation assessments for all species of the pineapple family (Bromeliaceae), one of the most diverse and ecologically important plant groups of the American tropics—a global biodiversity hotspot. Furthermore, we aim to analyse patterns of diversity, endemism and the conservation status of the Bromeliaceae on the continental level in the light of their evolutionary history. Location The Americas. Methods We compiled a dataset of occurrence records for 3,272 bromeliad species (93.4% of the family) and modelled their geographic distribution using either climate‐based species distribution models, convex hulls or geographic buffers dependent on the number of occurrences available. We then combined this data with information on taxonomy and used the ConR software for a preliminary assessment of the conservation status of all species following Criterion B of the International Union for the Conservation of Nature (IUCN). Results Our results stress the Atlantic Forest in eastern Brazil, the Andean slopes, Central America and the Guiana Highlands as centres of bromeliad diversity and endemism. Phylogenetically ancient subfamilies of bromeliads are centred in the Guiana highlands whereas the large radiations of the group spread across different habitats and large geographic area. A total of 81% of the evaluated bromeliad species are Possibly Threatened with extinction. We provide range polygons for 3,272 species, as well as newly georeferenced point localities for 911 species in the novel “bromeliad” r package, together with functions to generate diversity maps for individual taxonomic or functional groups. Main conclusions Diversity centres of the Bromeliaceae agreed with macroecological patterns of other plant and animal groups, but show some particular patterns related to the evolutionary origin of the family, especially ancient dispersal corridors. A staggering 2/3rds of Bromeliaceae species might be threatened with extinction, especially so in tropical rain forests, raising concerns about the conservation of the family and bromeliad‐dependent animal species.
... focused on individual taxa (Canela et al., 2003;Leme, Heller, Zizka, & Halbritter, 2017;Peters, 2009;Zizka, Horres, Nelson, & Weising, 1999;Zizka, Trumpler, & Zöllner, 2002) or geographic regions (Cáceres, 2012;Judith et al., 2013;Zizka et al., 2009; www.flora dobra sil.jbrj.gov.br). Our distribution maps are available in Appendix S2, and we supply all species ranges under a CC-BY license via the bromeliad r package, which also includes functions for publication-level species richness maps for individual genera, traits or conservation categories (Appendix S4). ...
Conference Paper
Our ongoing project involves a detailed morphological, biogeographical and molecular systematic analysis of Deuterocohnia (Bromeliaceae). The genus is mainly distributed in the Andes of southern Bolivia and northern Argentina. Typical habitats of these plants are dry, rocky slopes or rock faces. Succulent leaves and CAM photosynthesis are some of the adaptations related to their xerothermic habitat. Our recently conducted revision of the genus resulted in 17 species, four subspecies and four varieties. Comparative sequence analysis of three intergenic chloroplast DNA-regions (trnS-ycf3, rps16-trnK, rpl32-trnL) and two nuclear single copy marker (PRK exon2-5 and PHYC) show contradicting tree topologies. According to the chloroplast data, Deuterocohnia proved to be deeply paraphyletic, with five species forming a sister group to the closely related genus Dyckia, and all other species being sister to the Dyckia/Deuterocohnia clade. In contrast to the chloroplast DNA data, preliminary phylogenetic analyses based on low-copy nuclear markers provide good support for the monophyly of Deuterocohnia. The pattern of the cpDNA-based phylogeny and haplotype network supports biogeographical pattern rather than morphology. In addition to the sequence analyses, first steps towards microsatellite analyses of Deuterocohnia are presented.
Article
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A phylogenomic analysis of the so far phylogenetically unresolved subfamily Bromelioideae (Bromeliaceae) was performed to infer species relationships as the basis for future taxonomic treatment, stabilization of generic concept, and further analyses of evolution and biogeography of the subfamily. A target-enrichment approach was chosen, using the Angiosperms353 v.4 kit RNA-baits and including 86 Bromelioideae species representing previously identified major evolutionary lineages. Phylogenetic analyses were based on 125 target nuclear loci, assembled off-target plastome as well as mitogenome reads. A Bromelioideae phylogeny with a mostly well-resolved backbone is provided based on nuclear (194 kbp), plastome (109 kbp), and mitogenome data (34 kbp). For the nuclear markers, a coalescent-based analysis of single-locus gene trees was performed as well as a supermatrix analysis of concatenated gene alignments. Nuclear and plastome datasets provide well-resolved trees, which showed only minor topological in-congruences. The mitogenome tree is not sufficiently resolved. A total of 26 well-supported clades were identified. The genera Aechmea, Canistrum, Hohenbergia, Neoregelia, and Quesnelia were revealed polyphyletic. In core Bromelioideae, Acanthostachys is sister to the remainder. Among the 26 recognized clades, 12 correspond with currently employed taxonomic concepts. Hence, the presented phylogenetic framework will serve as an important basis for future taxonomic revisions as well as to better understand the evolutionary drivers and processes in this exciting subfamily.
Article
Bromeliaceae is one of the most diverse families in terms of morphology, especially its floral structure and stigmatic variability. Nineteen different stigmatic types are observed in the family, the margins of which differ in shape and presence or absence of epidermal appendages. Although many studies have classified stigma types, those that report on stigmatic margins are rare. Therefore, in this work, we aimed to study in detail the morphology of stigmatic margins in Bromeliaceae, including micromorphology and histochemistry, characterizing the secretions produced. To accomplish this, we used both optical and electron microscopy. Stigmatic margins of the twenty-two species studied were described as crenate, laciniate, lobate, undulate, and entire. We also observed unicellular, bicellular and multicellular secretory trichomes, branched or unbranched, with two to five arms. Stigmatic secretions, such as mucilage, total lipids, starch, polysaccharide, alkaloid and essential oil, were found. By addressing the diversity of stigmatic margins, we have added to the systematics of Bromeliaceae, including the morphology and histochemistry of epidermal appendages.
Article
The morphology of stigma has taxonomic values. To further explore the taxonomy of family Asteraceae, the morphological characteristics of stigma of 28 genera, 32 species, and two varieties in the family were observed using scanning electron microscopy. The results indicated that the stigma morphology of these Asteraceae plants could be divided into 10 types, of which eight are reported for the first time. The morphological characteristics of stigma support the close relationship between genera Aster and Erigeron and among genera Sonchus, Taraxacum, and Youngia. Our results enriched the stigma type diversity data and provided a morphological basis for the study of the phylogenetic evolution of Asteraceae. Scanning electron microscopy was used to provide high‐quality figures for observing stigma morphology. Ten types of Asteraceae stigmas enriched the stigma type diversity data.
Article
Plant evolution may be triggered by significant chromosome changes. In some plant groups, karyoevolution played an important role, influencing speciation processes. Hohenbergia comprises 48 species distributed through eastern Brazil. Previous cytological information includes few species and only chromosome counts, lacking information about genome size and more accurate karyomorphological investigation. Here, we compare cytomolecular features and genome sizes of 12 Hohenbergia spp. Besides, new measurements of genome sizes of 32 species are reported. All studied species presented 2n = 50, a number prevalent in Bromelioideae. The genome sizes (2C) varied from 0.74 to 1.52 pg. Despite the apparent homogeneity in chromosome number and genome size in Hohenbergia, significant polymorphism was observed in regard to the distribution of CMA+/DAPI0 bands and sites of 35S and 5S rDNA in metaphase chromosomes. Seven out of 12 analysed species presented heteromorphic pairs regarding 35S rDNA and/or 5S rDNA. Hohenbergia thus shows karyotypic diversity despite the conservation in chromosome number.
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We describe the chromosome numbers and genome sizes of species of the cryptanthoid complex of Bromeliaceae in a phylogenetic framework and their relationship with habitat preferences. The 2C DNA contents varied 2.13-fold among species, ranging from 0.76 to 1.66 pg. A significant difference in DNA content was found among Cryptanthus, Hoplocryptanthus and Rokautskyia. Moreover, species from campos rupestres and the Atlantic Forest had lower and higher genome size values, respectively. The smaller genome sizes of Hoplocryptanthus spp. from campos rupestres may be related with the large genome constraint. The species show a highly conserved ploidy (with 2n = 32 and 34), although the genome sizes varied considerably. The observed variation in chromosome numbers seems to be influenced by dysploidy, but additional investigations are needed. Our study demonstrates that the genome size variation in the cryptanthoid complex species is not strictly related to the phylogenetic relationships and has probably been influenced by different evolutionary processes.
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Lezak B, Varacallo M. Anatomy, Bony Pelvis and Lower Limb, Foot Veins. [Updated 2019 Jun 6]. In: StatPearls [Internet]. Treasure Island (FL): StatPearls Publishing; 2019 Jan-. Available from: https://www.ncbi.nlm.nih.gov/books/NBK542295/
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This open access book offers a fully illustrated compendium of glossary terms and basic principles in the field of palynology, making it an indispensable tool for all palynologists. It is a revised and extended edition of “Pollen Terminology. An illustrated handbook,” published in 2009. This second edition, titled “Illustrated Pollen Terminology” shares additional insights into new and stunning aspects of palynology. In this context, the general chapters have been critically revised, expanded and restructured. The chapter “Misinterpretations in Palynology” has been extended with new research data and additional ambiguous terms, e.g., polyads vs. massulae; the chapter “Methods in Palynology” has been extensively enhanced with illustrated protocols showing the majority of the methods and techniques used when studying recent and fossil pollen with LM, SEM and TEM. Moreover, additional information about the description and publication of pollen data is provided in the chapter “How to Describe and Illustrate Pollen Grains.” Various other parts of the general chapters have now been updated and/or extended with more comprehensive textual passages and new illustrations. The chapter “Illustrated Pollen Terms” now features new and more appropriate examples of each term, including additional LM micrographs. Where necessary, the entries for selected pollen terms have been refined by rewording or adding definitions, illustrations, and new micrographs. Lastly, new terms are included, such as “suprasculpture” and the prefix “nano-“ for ornamentation features. The chapter “Illustrated Pollen Terms” is the main part of this book and comprises more than 300 widely used terms illustrated with over 1,000 high-quality images. It provides a detailed survey of the manifold ornamentation and structures of pollen, and offers essential insights into their stunning beauty. Springer link: http://www.springer.com/de/book/9783319713649
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From 1990 to 2006, 2,875 new angiosperm species were described in Brazil, including 280 newBromeliaceae species. This publication rate is considered to be a useful indicator of floristic richness andalso reveals the huge gaps in our knowledge of species that make up Brazilian biomes and the importanceof taxonomy as a basic tool to assess biodiversity and conservation. The goal of modern taxonomists is ina race against time ordained by an unprecedented rate of global biodiversity loss, and therefore collaborationis vital to successfully close these gaps. This paper is the result of a broad cooperative research effortundertaken specifically to provide basic data on the identity of new components of Brazilian biologicaldiversity. The authors describe and illustrate 22 new Bromeliaceae species from three subfamilies: Bromelioideae - Aechmea guaratingensis, A. paratiensis, A. rubroaristata, Cryptanthus capitellatus, C. venecianus, C. viridovinosus, Hohenbergia aechmeoides, H. arcuata, H. barbarespina, H. reconcavensis, Nidularium alegrense, Orthophytum teofilo-otonense, O. cearence; Pitcairnioideae - Dyckia espiritosantensis, D. nana, Pitcairnia capixaba; Tillandsioideae - Tillandsia castelensis, Vriesea euclidiana, V. fontanae, V. multifoliata, V. sanctateresensis and V. teresopolitana.
Thesis
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The monocot family Bromeliaceae comprises approximately 3,140 species distributed in tropical and subtropical regions of the New World from the southern United States to southern Argentina. The family is subdivided into eight subfamilies; the most species-rich are Tillandsioideae and Bromelioideae. Taxonomic concepts within Bromeliaceae are highly problematic, since discriminating morphological characters have been shown to be homoplastic or plesiomorphic. The present study aims to provide a robust phylogenetic framework for Bromeliaceae, especially for the most diverse and complicated subfamilies Bromelioideae and Tillandsioideae. Resulting phylogenies provide a basis to estimate the usefulness of morphological characters and to propose or strengthen hypotheses concerning evolutionary traits, biogeography, age and origins of bromeliads. The main questions raised are: (1) Do additional sequence data from the plastid genome and a wider sampling within Bromeliaceae provide a better resolved, robust phylogenetic framework? What are the reasons for the low DNA sequence divergence observed up to now? (2) Can nuclear DNA sequences be successfully implemented for phylogenetic reconstruction? What are the challenges to optimize nuclear markers and do they perform better than plastid loci? (3) Can the resulting phylogeny based on plastid and nuclear DNA sequences together with the re-evaluated morphological characters provide a reasonable, stable classification? To provide a more robust phylogenetic hypothesis for the classification of Bromeliaceae, eight rapidly evolving plastid DNA markers (atpB-rbcL, matK, ndhF, psbA-trnH, rpl32-trnL, rps16, trnL intron, and trnL-trnF) and 90 bromeliad species were included in the current study and analysed using maximum-parsimony, maximum-likelihood, and Bayesian approaches. Results support the formerly proposed eight-subfamily classification based on the single plastid gene ndhF. Support values for five of the subfamilies increase, but that for Lindmanioideae, Puyoideae, and Bromelioideae decrease as a result of expanded taxon sampling, including several more divergent species. The initially proposed monophyletic origin of Puyoideae cannot be unambiguously confirmed. Calibration of the resulting phylogeny against time and biogeographic analysis reveals that Bromeliaceae originated in the Guayana Shield about 100 million years ago (Ma) and spread radially into adjacent areas ca. 16—13 Ma. Extant lineages arose between 20 and 5 Ma. Andean uplift facilitated diversification of core Tillandsioideae about 14.2 Ma and Bromelioideae 10.1 Ma, the latter having their greatest diversity in the Brazilian Shield due to dispersal from the Andes. The most species-rich genera did not appear before 8.7 My with a high diversification between 5 and 4 Ma, which is most likely the reason for the comparatively low sequence divergence. To test the usefulness of nuclear regions for phylogenetic reconstruction in Bromelioideae, DNA sequences of part of the low-copy nuclear gene phosphoribulokinase (PRK) and five plastid loci (matK, 3'trnK intron, trnL intron, trnL-trnF, and atpB-rbcL) were investigated. Phylogenetic trees obtained from analyses of the PRK sequences do not contradict trees obtained from plastid markers. The PRK matrix shows a significantly higher number of potentially PICs (phylogenetically informative characters) than the plastid dataset (16.9% vs. 3.1%), which improves resolution and support in the resulting trees. Although PRK is not able to resolve relationships completely, the combined analysis with plastid markers yields good support for several uncertain relationships observed previously. The early diverging lineages can be identified (“basal bromelioids”) and the remainder of the subfamily clustered into a highly supported clade (“eu-bromelioids”). Results indicate that taxonomic circumscriptions within “core bromeliads” are still insufficient, and relationships complex and difficult to solve. Several genera appear polyphyletic, and Aechmea as well as Quesnelia remain the most complicated genera of the subfamily. Most-parsimonious character state reconstructions for two evolutionary traits (tank habit, sepal symmetry) indicate that both characters have undergone few transitions within the subfamily and thus are not as homoplasious as previously assumed. The comparative study of nuclear DNA sequences within Tillandsioideae shows that some nuclear markers are able to provide more information and a higher degree of resolution in phylogenetic trees than plastid markers. However, their utility does not depend only on sequence variability, but also on methodological challenges in using traditional Sanger-sequencing. The internal transcribed spacer of nuclear ribosomal DNA (ITS nrDNA) is not recommended as a suitable marker for phylogenetic investigations of Bromeliaceae due to the presence of strong secondary structures which create problems in amplification and sequencing as well as its low number of PICs for resolving deeper nodes. Amplified fragments of the genes malate synthase (MS) and RNA polymerase II, beta subunit (RPB2) are not helpful due to their small size and limited number of PICs. Glucose-6-phosphate isomerase (PGIC), nitrate reductase (NIA), and xanthine dehydrogenase (XDH) need to be further investigated. Phosphoribulokinase (PRK) and phytochrome C (PHYC) are useful nuclear markers and able to provide considerable resolution in phylogenetic trees, but some relationships are poorly supported. The combined analysis of nuclear DNA sequence data (PRK, PHYC) with the already existing plastid DNA sequence data (atpB-rbcL, matK, rbcL, partial rbcL-accD, rps16 intron, partial trnK intron, trnL intron and trnL-trnF) shows a significant increase of resolution within phylogenetic trees of Tillandsioideae. Nine accepted genera can be re-circumscribed and three new genera are described taxonomically based on morphology. For morphologically distinct species groups within Racinaea and Tillandsia, two new subgenera are erected. Viridantha has been downgraded to subgeneric rank. Poor sampling within the Cipuropsis-Mezobromelia clade and missing support for clades within Tillandsia prevent the recognition of further taxonomic groups.
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Abstract— Genetic concepts within the Bromelioideae are highly problematic, in particular within the tank-forming “core bromelioid” clade. Previous molecular studies showed that the largest genus, Aechmea, and allied genera are polyphyletic and require revision. Here we focus on one group within the Aechmea alliance, the Portea/Gravisia group. To assess whether species of this group form a distinct lineage within the core bromelioid clade, and to clarify generic limits and interspecific relationships within the group, we generated and analyzed amplified fragment length polymorphisms (AFLP). In total, 69 species were sampled, including 26 species previously assigned to the Portea/Gravisia group. Neighbor joining, maximum likelihood and Bayesian analyses of the AFLP data consistently retrieved the Portea/Gravisia group as a monophyletic clade with several subclades, comprising species from the genera Aechmea, Canistrum and Portea. The phylogenetic distribution of polyporate pollen of the resulting trees indicates this character state arose once within the core bromelioids and thus can be regarded as a synapomorphy for the Portea/Gravisia clade. Further, our study shows that within the Portea/Gravisia group, subclades are characterized by petal color and geographic distribution. Thus, the present study is a further advance in the challenging task of elucidating phylogenetic relationships within the core bromelioids as the basis for a revised taxonomy of this ecologically important group.
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A phylogenetic study testing the monophyly of the geographically disjunct genus Ronnbergia (Bromeliaceae, Bromelioideae) is presented. The phylogenetic analyses were based on taxon sampling that included all but one species of Ronnbergia, and representative lineages across the subfamily Bromelioideae. Three chloroplast DNA sequence markers (matK, psbA-trnH, and trnL-trnF) and morphological data were used for the phylogenetic reconstruction. Both the molecular and morphological datasets supported the polyphyly of Ronnbergia, either independently or in combination. These findings suggest that the geographic disjunction of this genus is most likely a product of taxonomic misinterpretation. The results also indicate that the species currently circumscribed in Ronnbergia are closely related to species in the genus Aechmea with similar geographic ranges. The datasets do not have enough resolution power to reconstruct a deep phylogenetic history that involves all the species of Ronnbergia. Nevertheless, this study provides clues for future approaches that should focus on a larger species sampling and the use of multi-locus DNA sequence data.
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Adaptation to pollinators is a key factor of diversification in angiosperms. The Caribbean sister genera Rhytidophyllum and Gesneria present an important diversification of floral characters. Most of their species can be divided in two major pollination syndromes. Large-open flowers with pale colours and great amount of nectar represent the generalist syndrome, while the hummingbird-specialist syndrome corresponds to red tubular flowers with a less important nectar volume. Repeated convergent evolution toward the generalist syndrome in this group suggests that such transitions rely on few genes of moderate to large effect. To test this hypothesis, we built a linkage map and performed a QTL detection for divergent pollination syndrome traits by crossing one specimen of the generalist species Rhytidophyllum auriculatum with one specimen of the hummingbird pollinated R. rupincola. Using geometric morphometrics and univariate traits measurements, we found that floral shape among the second-generation hybrids is correlated with morphological variation observed between generalist and hummingbird-specialist species at the genus level. The QTL analysis showed that colour and nectar volume variation between syndromes involve each one major QTL while floral shape has a more complex genetic basis and rely on few genes of moderate effect. Finally, we did not detect any genetic linkage between the QTLs underlying those traits. This genetic independence of traits could have facilitated evolution toward optimal syndromes.
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A checklist of the 14 genera and 34 species of Bromeliaceae from the Parque Estadual do Rio Preto in São Gonçalo do Rio Preto municipality, Minas Gerais state, southeastern Brazil, is presented. The Tillandsioideae was the most diverse subfamily and was found to be concentrated in rocky field areas. Bromelioideae is also a species rich subfamily, but its taxa have shown a preference to forested areas and savannas at lower altitudes. Pitcairnioideae is highlighted by its level of endemism, but has only four species. Cryptanthus micrus, a new species found in this area is described and illustrated. Our cluster analysis indicated that the Rio Preto State Park has a Bromeliaceae flora more similar to that from Pico do Itambé and Grão Mogol State Parks. Taxa like Dyckia glandulosa, Orthophytum itambense and Vriesea medusa, which were previously considered to be endemic to Pico do Itambé, now have their area of occurrence extended to Rio Preto. These new occurrences highlight the importance to create a corridor joining these neighboring reserves to connect populations of narrowly ranged or rare species. In this work we present pictures of 19 species in their habitats within the park, and we hope that these illustrations will help in the identification and conservation of these taxa.
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Communication of any sort is complex and communication between plants and animals is particularly so. Plant-pollinator mutualisms are amongst the most celebrated partnerships that have received a great deal of attention for many centuries. At the outset, most pollination studies focused on phenotypic matches and invoked co-evolution to explain plant- pollinator interactions, which gave rise to the concept of pollination syndromes. A few centuries later, there has been a substantial shift in the way we view these mutualistic interactions. In a significant departure from a co-evolutionary framework, numerous studies subsequently showed that there is usually only a loose, non-exclusive matching between flowers and their pollinators. Concurrently, the global prevalence of generalized pollination systems was demonstrated repeatedly. However, our understanding of the evolutionary processes that underlie these mutualisms is still limited. Here, we provide a concise review of the state of our knowledge on the evolution of floral traits and pollinator sensory perception and how these together shape the structure and organization of pollination networks.
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Abstract— Of the eight subfamilies currently recognized in Bromeliaceae, Bromelioideae is perhaps the most poorly understood. Generic circumscriptions are unclear, and an exceptionally diverse morphology coupled with an unusually low rate of sequence divergence within Bromeliaceae has made it difficult to resolve phylogenetic relationships within the subfamily. Although recent molecular studies have begun elucidating relationships among species in Bromeliaceae, most have not sampled deeply and/or broadly across Bromelioideae. The purpose of this study was to conduct a phylogenetic analysis within subfamily Bromelioideae using three chloroplast DNA regions (matK, psbA-trnH, and trnL-trnF), with the inclusion of multiple species from a broad sampling of bromelioid genera. Ochagavia, Deinacanthon, Fascicularia, Bromelia and Fernseea diverged relatively early in the history of the subfamily, with the remaining taxa being placed in a large and poorly resolved eubromelioid clade. Bromelia and Cryptanthus were found to be monophyletic, while 13 other genera were polyphyletic. Aechmea, the most morphologically diverse genus within the subfamily, was highly polyphyletic, with species distributed among 12 different lineages, with little support for subgeneric circumscriptions.
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This is the first in a series of papers that will review generic limits in the Tillandsioideae. The intent is to circumscribe the boundaries of individual genera in order to develop a natural and phylogenetic system of generic-level classification within the subfamily. Here, Alcantarea (Morren ex Mez) Harms is resurrected to the generic rank from its previous subgeneric position under Vriesea. It is distinguished from Vriesea by its spectacular, linear-long, fusiform, ephemeral, distinctly flaccidescent, spiralescent petals, and seeds with both basal and apical comas. The history, nomenclature, taxonomy, and generic relationships of the genus are discussed. Alcantarea is restricted to the Brazilian states of Rio de Janeiro, Espírito Santo, Minas Gerais, and Bahia, the are of the enter of diversity of Vriesea. A new genus of Bromeliaceae is described for sixty-six species and one subspecies previously attributed to Vriesea Lindley subgenus Vriesea section Xiphon (E. Morren) E. Morren ex Mez. In honor of Prof. Dr. Werner Rauh, it is named Werauhia J.R.Grant, gen. nov. It is characterized by plants with mostly nocturnal anthesis, a general lack of brilliant coloration, mostly fleshy, frequently secund bracts and flowers, bilaterally symmetric, often zygomorphic corolla, androeciae and gynoeciae, the latter two well included within the corolla, dactylic petal appendages with one to five fingers of varying length, stigmata with the cupulate type morphology, lacking papillae, and stout, dark-colored capsules. Within Werauhia, two sections are described: Werauhia for thirty taxa representing elements of Utley's "Thecophylloid allies", and Jutleya J.Grant sect. nov. in honor of Dr. John F. Utley, for thirty-six species and one subspecies representing Utley's "Thecophylloid vrieseas". The genus ranges from southern Mexico through Mesoamerica to the West Indies, Peru, and northeastern Brazil. Its center of diversity and highest concentration of species is in the mountain range that includes the Cordillera de Guanacaste, Cordillera de Tilaran, Cordillera Central and Cordillera de Salamanca in Costa Rica and western Panama.
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Significance By using structural characteristics, such as long tubular flowers, plants are known to achieve selective visitation by certain pollinator species. These morphological traits typically arise over evolutionary timescales. We show for the first time, to our knowledge, that at least one plant has also evolved the capacity to recognize pollinator species immediately after visitation, thereby increasing the likelihood that a flower visitor has delivered high-quality pollen. This novel responsiveness by the plant leads to functional specialization in an apparently generalized tropical plant–pollinator network. Such specialized linkages likely facilitate coevolution but also, render pollination mutualisms more vulnerable to environmental change.
Article
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Aechmea (ca. 220 species) is the largest and most diverse genus in Bromelioideae (Bromeliaceae), and several dissimilar generic concepts and infrageneric classifications have been proposed, frequently involving other closely related Bromelioideae. A morphology-based phylogenetic analysis using parsimony was conducted with 86 taxa, including 52 Aechmea (7 of the 8 recognized subgenera represented) and 34 exemplars from 9 closely related genera as the ingroup. Two species of Cryptanthus were included as the outgroup. The main objectives were to assess the validity of the major infrageneric classification systems proposed for Aechmea and to elucidate the phylogenetic position of Aechmea and putatively related genera in subfamily Bromelioideae. The topology of the consensus tree suggests that Acanthostachys, Billbergia, Portea, and non-Brazilian Ronnbergia may be monophyletic. Hohenbergia, Streptocalyx, and Quesnelia are paraphyletic or polyphyletic, as are most subgenera of Aechmea, except for subgenera Chevaliera and Macrochordion, which appear monophyletic. Characters traditionally emphasized in classifications of Bromelioideae displayed high levels of homoplasy, and this may be a reason for the artificiality of the taxonomic systems proposed for these taxa. Due to weak internal support, we refrain from recognizing any new taxonomic rearrangements. These results do provide new insights into the relationships within a number of Bromelioideae genera and suggest directions for future studies.
Article
A taxonomic revision of Bromeliaceae subfam. Tillandsioideae is presented based on a multi-locus DNA sequence phylogeny (viz., plastid DNA loci rpoB-trnC-petN, trnK-matK-trnK, and ycf1, and the nuclear DNA gene PHYC) and new or re-evaluated morphology (e.g., leaf, inflorescence, sepal, petal, ovary, stigma, stamen, pollen, ovule, and seed morphology). This enables the circumscription of monophyletic units using synapomorphic combinations of diagnostic morphological characters. Stigma morphology has proven to be indicative for super-specific taxa in Tillandsioideae. One new stigma type and several subtypes of previously described stigmas were found. The four tribes proposed earlier are mostly confirmed, but Catopsideae replaces the formerly used name Pogospermeae for the monotypic tribe of Catopsis. In addition, the two new subtribes Cipuropsidinae and Vrieseinae are proposed within tribe Vrieseeae. Several new genera are established to render taxonomic units monophyletic and morphologically well circumscribed. They represent segregates of either Mezobromelia (Gregbrownia: 4 spp.), Tillandsia (viz., Barfussia: 3 spp., Josemania: 5 spp., Lemeltonia: 7 spp., Pseudalcantarea: 3 spp., and Wallisia: 4 spp. and 1 hybrid), or Vriesea (viz., Goudaea: 2 spp., Jagrantia: 1 sp., Lutheria: 4 spp., Stigmatodon: 18 spp., and Zizkaea: 1 spp.). The new subgenera Tillandsia subg. Pseudovriesea and T. subg. Viridantha are established, and T. subg. Aerobia is resurrected. An identification key to all accepted genera of Bromeliaceae subfam. Tillandsioideae is provided. Furthermore, to clarify nomenclatural uncertainties, typifications are proposed for Catopsis subg. Tridynandra, Thecophyllum [unranked] Biflorae, Tillandsia subg. Aerobia, T. sect. Caricifoliae, T. sect. Conostachys, T. sect. Cyathophora, T. sect. Eriophyllum, T. sect. Macrocyathus, T. sect. Platystachys Baker auct. non al., Tillandsia sect. Strepsia, Vriesea subg. Conostachys Mez auct. non al., T. lindenii K. Koch auct. non al., and T. macropetala.
Article
The tank-epiphytic clade of berry-fruited bromeliads, also known as the Core Bromelioideae, represents a remarkable event of adaptive radiation within the Bromeliaceae; however, the details of this radiation have been difficult to study because this lineage is plagued with generic delimitation problems. In this study, we used a phylogenetic approach to investigate a well supported, albeit poorly understood, lineage nested within the Core Bromelioideae, here called the "Ronnbergia Alliance." In order to assess the monophyly and phylogenetic relationships of this group, we used three plastid and three nuclear DNA sequence markers combined with a broad sampling across three taxonomic groups and allied species of Aechmea expected to comprise the Ronnbergia Alliance. We combined the datasets to produce a well-supported and resolved phylogenetic hypothesis. Our main results indicated that the Ronnbergia Alliance was a well-supported monophyletic group, sister to the remaining Core Bromelioideae, and it was composed by species of the polyphyletic genera Aechmea, Hohenbergia and Ronnbergia. We identified two major internal lineages with high geographic structure within the Ronnbergia Alliance. The first of these lineages, called the Pacific Clade, contained species of Aechmea and Ronnbergia that occur exclusively from southern Central America to northwestern South America. The second clade, called the Atlantic Clade, contained species of Aechmea, Hohenbergia and Ronnbergia mostly limited to the Atlantic Forest and the Caribbean. We also explored the diagnostic and evolutionary importance of 13 selected characters using ancestral character reconstructions on the phylogenetic hypothesis. We found that the combination of tubular corollas apically spreading and unappendaged ovules had diagnostic value for the Ronnbergia Alliance, whereas flower size, length of the corolla tube, and petal pigmentation and apex were important characters to differentiate the Pacific and Atlantic clades. This study opens new perspectives for future taxonomic reorganizations and provides a framework for evolutionary and biogeographic studies.
Article
We here propose Aechmea subintegerrima as a new combination for Ronnbergia brasiliensis, a taxon belonging to the Aechmea lingulata complex. Additionally, we describe and illustrate 14 new Bromeliaceae species from the Brazilian states of Bahia, Espirito Santo, Mato Grosso, Minas Gerais and Rio de Janeiro: Aechmea aiuruocensis, A. altocaririensis, A. limai, A. recurvipetala, Dyckia kranziana, Hohenbergia loredanoana, Neoregelia dactyloflammans, N. retrorsa, Orthophytum rafaelii, Vriesea minutiflora, V. nubicola, V. pulchra, V. santaleopoldinensis, and V. serranegrensis. The morphological affinities of the new taxa are also discussed.
Article
O fundador da Escola de Minas de Ouro Preto, o mineralogista Francês Claude Henrique Gorceix, definiu certavez o estado de Minas Gerais como aquele com o peito de aço e o coração de ouro. Essa comparação vale emespecial para o Quadrilátero Ferrífero (QF), uma região clássica da geologia e da mineração brasileira, que seestende entre as cidades de Belo Horizonte (NW), Itabira (NE), Ouro Preto (SE) e Congonhas (SW). Ocorremaqui jazidas de ferro (Fe), manganês (Mn), ouro (Au), bauxita e pedras preciosas, como topázio e esmeralda. A área foi descoberta pelos bandeirantes no final do século XVII quando buscavam pela esmeralda, raridadesobre a qual circulavam na época colonial os boatos mais insanos. Entretanto, eles encontraram o ouro, e esteera preto, motivo pelo qual a localidade do descobrimento passou a ser chamada de Ouro Preto. Os primeirosachados do metal nobre em torno de 1693 levaram a uma verdadeira febre aurífera. Houve naquele tempouma migração enorme em direção às montanhas ao redor desse lugar, denominado inicialmente Villa Rica.E essa migração trouxe todos os seus aspectos positivos e negativos. Assim, antigas crônicas mencionam que no norte do país monastérios inteiros eram despovoados, porque também os monges foram atraídospelo novo Eldorado. A procura dos aventureiros pelo metal nobre foi tão grande que a superpopulação daárea causou em 1701 uma enorme emergência de fome, que suprimiu grande parte da população. Muitosmorreram com os bolsos cheios de ouro, mas não havia nada comestível que pudesse ser adquiridos com seus tesouros. Uma vez que as ocorrências mais produtivas nos aluviões e sedimentos do rio do Carmo foramexploradas rapidamente, a febre aurífera chegou a um fim já após aproximadamente quarenta anos. Somentemuito mais tarde surgiu na região a exploração subterrânea organizada do ouro. Como conseqüência a regiãoem torno de Ouro Preto perdeu muito de sua importância econômica, ainda assim a cidade permaneceu por muito tempo o centro administrativo de Minas e posteriormente foi promovida à capital do Estado. Com oreconhecimento geológico e a exploração das enormes ocorrências do minério de ferro na área do QF após asegunda guerra mundial, Minas Gerais viveu um renascimento econômico e transformou-se num dos estados mais ricos do Brasil. Ouro Preto com seu centro histórico bem conservado, suas igrejas barrocas ricas em ouro e obras de arte, seus museus, entre eles o bem conhecido museu mineralógico da Escola de Minas, e outros monumentos e aspectos interessantes se transformou em uma jóia turística nacional.
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The genus Fascicularia Mez is revised as part of a study of the Bromeliaceae for the Flora de Chile . Morphological and anatomical investigation of herbarium and living material from cultivation as well as DNA-studies (RAPDs) in cultivated material has led us to conclude that Fascicularia bicolor (Ruiz & Pav.) Mez, the only one species in the genus, comprises two subspecies which are distinguished by their leaf anatomy and morphology.
Article
The author describes and illustrates two new outlier species in Bromeliaceae, Orthophytum roseolilacinum and O. vasconcelosianum, from Minas Gerais state, Brazil, which are members of the “Cryptanthoid complex”. The morphological affinities and discordant features in relation to the conceptual boundaries of the genus are also discussed.
Article
A multiple sequence alignment program, MAFFT, has been developed. The CPU time is drastically reduced as compared with existing methods. MAFFT includes two novel techniques. (i) Homo logous regions are rapidly identified by the fast Fourier transform (FFT), in which an amino acid sequence is converted to a sequence composed of volume and polarity values of each amino acid residue. (ii) We propose a simplified scoring system that performs well for reducing CPU time and increasing the accuracy of alignments even for sequences having large insertions or extensions as well as distantly related sequences of similar length. Two different heuristics, the progressive method (FFT‐NS‐2) and the iterative refinement method (FFT‐NS‐i), are implemented in MAFFT. The performances of FFT‐NS‐2 and FFT‐NS‐i were compared with other methods by computer simulations and benchmark tests; the CPU time of FFT‐NS‐2 is drastically reduced as compared with CLUSTALW with comparable accuracy. FFT‐NS‐i is over 100 times faster than T‐COFFEE, when the number of input sequences exceeds 60, without sacrificing the accuracy.
Article
We describe and illustrate 14 miscellaneous new Bromeliaceae species from the Brazilian states of Bahia, Espirito Santo, Minas Gerais, Rio de Janeiro and Sao Paulo: Aechmea ituberaensis, A. pendulispica, Alcantarea acuminatifolia, A. occulta, Cryptanthus aracruzensis, C. ilhanus, C. rigidifolius, C. tabuleiricola, C. viridipetalus, Neoregelia desenganensis, N. insulana, N. watersiana, Vriesea saltensis, and V. sanctaparecidae. The morphological affinities of the new taxa are also discussed.
Book
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
Article
A cladistic analysis of morphological data for the genus Canistropsis is presented. With the exception of two species, C. selloana and C. correia-araujoi, results support Canistropsis as monophyletic. Canistropsis correia-araujoi is a suspected, natural bigeneric hybrid. Canistropsis selloana is transferred back into the monotypic genus Andrea. A taxonomic treatment, including a greatly emended description, and discussion of the distribution, habitat, and conservation of Andrea selloana is presented.
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Over 400 taxa in the Bromeliaceae were surveyed for stigma morphology. Five morphological categories (conduplicate-spiral, simple-erect, cupulate, convolute-blade, and coralliform) account for all known variation in bromeliad stigma morphology. The coralliform type is described and illustrated for the first time. The subfamilies Bromelioideae and Pitcairnioideae appear to be nearly homogeneous for the conduplicate-spiral stigma type. All five of the stigma types are found in the third subfamily, Tillandsioideae, where stigma variability promises to be the most useful systematically. There appear to be correlations between the simple-erect stigma type and 1) derived sexual systems (i.e., dioecism, andromonoecism, cleistogamy) within the family, and 2) xericepiphytism in the genus Tillandsia. In conjunction with other floral features, stigma morphology may be useful in refining generic and infrageneric circumscriptions.
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Cryptanthus Otto & A. Dietrich, one of the most distinctive genera in the Bromeliaceae, is endemic to Brazil. The flowering phenology and floral biology of Cryptanthus dianae Lerne were studied in natural sites in Tapacurá Ecological Station and Natural heritage private reserve (RPPN) Frei Caneca, both located in Pernambuco. Cryptanthus dianae flowers once a year, with one to four flowers opening per day. The species is andromonoecious with a predominance of male flowers during the beginning of the blooming period. Flowers emerge in the central apical area of the inflorescence and more rarely in the peripheral basal area of the rosette where the hermaphrodite flowers open. Visits by hummingbirds (Phae thornis ruber), flies, diurnal moths (Saliana sp.), and bees (Euglossa cordata, Eulaema nigrita, Trigona sp.) were observed. According to the behavior and frequency of visits, the Euglossini males appear to be the main pollinators. During floral visits, Euglossa cordata and Eulaema nigrita scrape the petals, filaments and style with their forelegs, thus suggesting that odor is an additional floral resource in addition to the pollen and nectar. This is the first report of the presence of floral odor in Bromeliaceae as the primary attractive.
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The history of discovery of the single Bromeliaceae species outside the New World, Pitcairnia feliciana, is recounted by its first collector, Henri Jacques-Félix, for whom it was named. Notes on distribution and a description of the habitat where P. feliciana is found are included.