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Leaf blades of Vriesea species in cross section (A, E, K-L) and surface view (scanning electron microscopy, SEM, B-D; F-J). A. V. zildae, showing stoma only on abaxial surface (arrows). B. V. sucrei, stomata on abaxial surface. C V. jonesiana, stomata on abaxial surface. D. V. ensiformis, stomata and trichomes organised in linear series on abaxial surface. E. V. eltoniana, trichomes on both adaxial and abaxial surfaces. F. V. eltoniana, trichome on abaxial surface showing shield with four cells. G, H. V. pabstii, adaxial (G) and abaxial (H) surfaces densely covered by trichomes. I, J. V. longicaulis, trichomes remnants (arrows) on adaxial surface (I) and intact trichomes on abaxial surface (J). K, L. Peltate trichome in longitudinal section on abaxial surface showing stalk with five cells (asterisks) in V. botafogensis (K) and with six cells (asterisks) in V. psittacina (L). [Scale bars = 5 μm (C), 10 μm (B), 20 μm (F, K, L), 100 μm (A, D, E, I, J), 250 μm (G, H).]
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The Bromeliaceae are a largely Neotropical family originating in open, dry environments. Vriesea Lindl., the third largest genus of the family, is traditionally divided between two sections. About 90% of the species of the genus occur in Brazil, where the centre of diversity is the Atlantic Rainforest. Leaf morphoanatomical studies conducted on bro...
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... of all the Vriesea species analysed are hypostomatic (Fig. 1A). In surface view, the stomata have narrowly elliptical (V. eltoniana, V. flexuosa, V. guttata, V. paratiensis, V. philippocoburgii, V. procera, V. sincorana, V. sucrei (Fig. 1B) and V. vagans) to broadly elliptical outlines (V. billbergioides, V. botafogensis, V. cacuminis, V. ensiformis, V. flava, V. gradata, V. hydrophora,V. ...
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... of all the Vriesea species analysed are hypostomatic (Fig. 1A). In surface view, the stomata have narrowly elliptical (V. eltoniana, V. flexuosa, V. guttata, V. paratiensis, V. philippocoburgii, V. procera, V. sincorana, V. sucrei (Fig. 1B) and V. vagans) to broadly elliptical outlines (V. billbergioides, V. botafogensis, V. cacuminis, V. ensiformis, V. flava, V. gradata, V. hydrophora,V. incurvata, V. jonesiana (Fig. 1C), V. longicaulis, V. pabstii, V. pseudatra, V. psittacina. V. saundersii and V. zildae). The stomata, along with the trichomes, are organised in rows ...
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... surface view, the stomata have narrowly elliptical (V. eltoniana, V. flexuosa, V. guttata, V. paratiensis, V. philippocoburgii, V. procera, V. sincorana, V. sucrei (Fig. 1B) and V. vagans) to broadly elliptical outlines (V. billbergioides, V. botafogensis, V. cacuminis, V. ensiformis, V. flava, V. gradata, V. hydrophora,V. incurvata, V. jonesiana (Fig. 1C), V. longicaulis, V. pabstii, V. pseudatra, V. psittacina. V. saundersii and V. zildae). The stomata, along with the trichomes, are organised in rows (Fig. 1D), the rows alternating with epidermal bands having neither stomata nor trichomes. This characteristic was not observed in V. pabstii, in which the abaxial leaf surface is densely ...
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... (Fig. 1B) and V. vagans) to broadly elliptical outlines (V. billbergioides, V. botafogensis, V. cacuminis, V. ensiformis, V. flava, V. gradata, V. hydrophora,V. incurvata, V. jonesiana (Fig. 1C), V. longicaulis, V. pabstii, V. pseudatra, V. psittacina. V. saundersii and V. zildae). The stomata, along with the trichomes, are organised in rows (Fig. 1D), the rows alternating with epidermal bands having neither stomata nor trichomes. This characteristic was not observed in V. pabstii, in which the abaxial leaf surface is densely covered in ...
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... trichomes occur on both adaxial and abaxial leaf surfaces (Fig. 1E). In surface view, these comprise a circular shield of four triangular cells (Fig. 1F), surrounded by two concentric series of thin-walled, quadrangular cells: the first series with eight cells and the second with sixteen cells (Fig. 1F). The two series are surrounded by a variable number of radially elongated cells in symmetrical ...
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... trichomes occur on both adaxial and abaxial leaf surfaces (Fig. 1E). In surface view, these comprise a circular shield of four triangular cells (Fig. 1F), surrounded by two concentric series of thin-walled, quadrangular cells: the first series with eight cells and the second with sixteen cells (Fig. 1F). The two series are surrounded by a variable number of radially elongated cells in symmetrical arrangement. Among the species analysed, only V. botafogensis, V. flava and V. pabstii show ...
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... trichomes occur on both adaxial and abaxial leaf surfaces (Fig. 1E). In surface view, these comprise a circular shield of four triangular cells (Fig. 1F), surrounded by two concentric series of thin-walled, quadrangular cells: the first series with eight cells and the second with sixteen cells (Fig. 1F). The two series are surrounded by a variable number of radially elongated cells in symmetrical arrangement. Among the species analysed, only V. botafogensis, V. flava and V. pabstii show intact trichomes on both adaxial and abaxial leaf surfaces (Fig. 1G, H). In the other species, intact trichomes occur only on the abaxial surfaces ...
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... quadrangular cells: the first series with eight cells and the second with sixteen cells (Fig. 1F). The two series are surrounded by a variable number of radially elongated cells in symmetrical arrangement. Among the species analysed, only V. botafogensis, V. flava and V. pabstii show intact trichomes on both adaxial and abaxial leaf surfaces (Fig. 1G, H). In the other species, intact trichomes occur only on the abaxial surfaces (Fig. 1I, J). In longitudinal section, the trichome stalk has five cells (V. billbergioides, V. botafogensis (Fig. 1K), V. cacuminis, V . e n s i f o r m i s , V . f l a v a , V . g u t t a t a , V. hydrophora, V. philippocoburgii, V. saundersii a n d V . s i n ...
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... (Fig. 1F). The two series are surrounded by a variable number of radially elongated cells in symmetrical arrangement. Among the species analysed, only V. botafogensis, V. flava and V. pabstii show intact trichomes on both adaxial and abaxial leaf surfaces (Fig. 1G, H). In the other species, intact trichomes occur only on the abaxial surfaces (Fig. 1I, J). In longitudinal section, the trichome stalk has five cells (V. billbergioides, V. botafogensis (Fig. 1K), V. cacuminis, V . e n s i f o r m i s , V . f l a v a , V . g u t t a t a , V. hydrophora, V. philippocoburgii, V. saundersii a n d V . s i n c o r a n a ) o r s i x c e l l s ( V. eltoniana, V.flexuosa V. gradata, V. incurvata, ...
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... arrangement. Among the species analysed, only V. botafogensis, V. flava and V. pabstii show intact trichomes on both adaxial and abaxial leaf surfaces (Fig. 1G, H). In the other species, intact trichomes occur only on the abaxial surfaces (Fig. 1I, J). In longitudinal section, the trichome stalk has five cells (V. billbergioides, V. botafogensis (Fig. 1K), V. cacuminis, V . e n s i f o r m i s , V . f l a v a , V . g u t t a t a , V. hydrophora, V. philippocoburgii, V. saundersii a n d V . s i n c o r a n a ) o r s i x c e l l s ( V. eltoniana, V.flexuosa V. gradata, V. incurvata, V. jonesiana, V. longicaulis, V. pabstii, V. paratiensis, V. procera, V. pseudatra, V. psittacina (Fig. ...
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... (Fig. 1K), V. cacuminis, V . e n s i f o r m i s , V . f l a v a , V . g u t t a t a , V. hydrophora, V. philippocoburgii, V. saundersii a n d V . s i n c o r a n a ) o r s i x c e l l s ( V. eltoniana, V.flexuosa V. gradata, V. incurvata, V. jonesiana, V. longicaulis, V. pabstii, V. paratiensis, V. procera, V. pseudatra, V. psittacina (Fig. 1L), V. sucrei, V. vagans and V. ...
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... cross-section, the leaf blade presents a onelayered epidermis on both surfaces (Fig. 2E). Epidermal cells have thickened anticlinal and inner periclinal walls, both of which present pectin (Appendix 1, Fig. 2E). The stomata are positioned at the same level as the epidermal cells (Fig. 2F). ...
Citations
... Floral bracts of Tillandsioideae species seem to share several features with the overall structure of bromeliads leaves, mostly notably xeric traits such as sclerotic epidermis and the presence of hypodermis, which is often differentiated in a water-storage tissue (Tomlinson 1969;Santos-Silva et al. 2013;Faria et al. 2021). In leaves, sclerification of the adaxial epidermal surface is common (Tomlinson 1969;Santos-Silva et al. 2013;Faria et al. 2021), a feature mostly absent in the studied floral bracts. ...
... Floral bracts of Tillandsioideae species seem to share several features with the overall structure of bromeliads leaves, mostly notably xeric traits such as sclerotic epidermis and the presence of hypodermis, which is often differentiated in a water-storage tissue (Tomlinson 1969;Santos-Silva et al. 2013;Faria et al. 2021). In leaves, sclerification of the adaxial epidermal surface is common (Tomlinson 1969;Santos-Silva et al. 2013;Faria et al. 2021), a feature mostly absent in the studied floral bracts. The presence of this trait in Go. chrysostachys comprise another distinction of this species with G. ospinae, which might corroborate the position of G. chrysostachys as more closely related to the Cipuropsis-Mezobromelia Complex (Machado et al. 2020). ...
... Our results suggest that the presence of a sclerotic hypodermis might be more associated with a structural function (mechanical support or protection against herbivores) than a xeric trait (see also Santos-Silva et al. 2013;Faria et al. 2021). Many species that inhabit xeric environments or are usually considered highly xeromorphic did not show a mechanical hypodermis (e.g. A. compacta, A. farneyi, T. loliacea and T. stricta). ...
Bromeliaceae display many water-use strategies, from leaf impounding tanks to CAM photosynthesis and absorbing trichomes. Recent studies show that trichomes in inflorescences of bromeliads can exude viscous secretions, protecting against various stresses, including excessive water loss. In light of this, and considering the knowledge gap regarding inflorescence trichomes in bromeliads, we aimed to investigate the presence, source, and chemical nature of inflorescence secretions in species of the Tillandsioideae (Bromeliaceae) and to describe the anatomy of their floral bracts focusing on trichome structure and position.We conducted a prospection of secretory activity and anatomy in floral bracts in 52 species of Tillandsioideae and one early-divergent Bromeliaceae species. We used histochemical tests to investigate the presence and nature of secretion combined with standard light microscopy methods. Secretion appears in all studied species of tribe Vrieseeae, in Guzmania species, Wallisia cyanea, Tillandsia streptopylla (Tillandsieae), and Catopsis morreniana (Catopsideae). It is absent in Vriesea guttata (Vrieseeae), Racinaea crispa, and various Tillandsia species (Tillandsieae). Secretion is produced by peltate trichomes on the adaxial surface of young bracts and comprises hydrophilic and lipophilic substances. Bract anatomy revealed an internal mucilage-secreting tissue with wide distribution within subtribe Vrieseinae. Our results point to a broad occurrence of secretion associated with bracteal scales in inflorescences of Tillandsioideae. Secretory function is strongly related to trichomes of the adaxial surface, whereas the indumentum of the abaxial side is lacking or likely associated with water absorption; the latter case is especially related to small, xeric plants. Exudates might engage in colleter-like roles, protecting against desiccation, high-radiation, and herbivores. Directions for future research are presented.
... According to Giuliani and Bini (2008), starch grains were found in the trichomes of leaves and floral verticils in the subfamily Lamioideae (Lamiaceae) in association with the initial phase of the secretion process. In Bromeliaceae, starch grains can be found in chlorenchyma cells for some species of Vriesea and may be related to energy storage, osmoregulation and tolerance to water stress (Faria et al., 2021). However, its presence in the stigmatic region in Vriesea, as analyzed here, is novel. ...
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.
... In this line of thought, based on several reports of environmental effects on the composition of epicuticular waxes, (Hull et al., 1979;Shepherd et al., 1995;Medina et al., 2004;Domínguez et al., 2011;Ortúñez and Cano-Ruiz, 2013;Rajčević et al., 2014;Menzel et al., 2017;Xue et al., 2017;Sharma et al., 2018) as well as on the close link established between the characteristics of epicuticular waxes of many taxa and taxonomic, ecological or evolutive issues, (Li and Christophel, Frontiers in Plant Science | www.frontiersin.org 2000; Versieux et al., 2012;Li et al., 2016;Mitić et al., 2016;Harris et al., 2017;Klimko et al., 2018;Lindelof et al., 2020;Weber and Schwark, 2020;Faria et al., 2021) we propose in this work an unprecedented strategy to contribute to the study of vicariant phenomena in plant science. This new strategy involves a comprehensive physical-chemical analysis of the surface of plant leaves that strengthens the more usual phylogenetic, biogeographic and ecological approaches. ...
Classically, vicariant phenomena have been essentially identified on the basis of biogeographical and ecological data. Here, we report unequivocal evidences that demonstrate that a physical–chemical characterization of the epicuticular waxes of the surface of plant leaves represents a very powerful strategy to get rich insight into vicariant events. We found vicariant similarity between Cercis siliquastrum L. (family Fabaceae, subfamily Cercidoideae) and Ceratonia siliqua L. (family Fabaceae, subfamily Caesalpinoideae). Both taxa converge in the Mediterranean basin (C. siliquastrum on the north and C. siliqua across the south), in similar habitats (sclerophyll communities of maquis) and climatic profiles. These species are the current representation of their subfamilies in the Mediterranean basin, where they overlap. Because of this biogeographic and ecological similarity, the environmental pattern of both taxa was found to be very significant. The physical–chemical analysis performed on the epicuticular waxes of C. siliquastrum and C. siliqua leaves provided relevant data that confirm the functional proximity between them. A striking resemblance was found in the epicuticular waxes of the abaxial surfaces of C. siliquastrum and C. siliqua leaves in terms of the dominant chemical compounds (1-triacontanol (C30) and 1-octacosanol (C28), respectively), morphology (intricate network of randomly organized nanometer-thick and micrometer-long plates), wettability (superhydrophobic character, with water contact angle values of 167.5 ± 0.5° and 162 ± 3°, respectively), and optical properties (in both species the light reflectance/absorptance of the abaxial surface is significantly higher/lower than that of the adaxial surface, but the overall trend in reflectance is qualitatively similar). These results enable us to include for the first time C. siliqua in the vicariant process exhibited by C. canadensis L., C. griffithii L., and C. siliquastrum.
... The features of leaf anatomy, such as the hypostomatic leaves with peltate trichomes, epidermal cells with thickened cell walls, covered with cuticle and epicuticular wax, mechanical hypodermis, water-storage parenchyma, and air lacunae associated with diaphragms constituted by arm-cells (Tomlinson 1969) support the hypothesis of the origin of Bromeliaceae in dry and open environments (Bouchenak-Khelladi et al. 2014). Within Vrieseinae, these features have been reported for Alcantarea (Versieux et al. 2010) and Vriesea (Arruda and Costa 2003;Proença and Sajo 2007;Faria et al. 2021). The leaf anatomy of Stigmatodon and Waltillia has not been studied, but its occurrence in open environments, together with the presence of coriaceous leaves and the high density of trichomes on the blade, suggest that the features of the leaf anatomy are typically xeromorphic (Tomlinson 1969;Givnish et al. 2011). ...
... The vouchers used for anatomical studies are indicated in Appendix S1. We used the specific terminology for both macro-and micromorphological descriptions of the family (Tomlinson 1969;Smith and Downs 1977;Arruda and Costa 2003;Scharf and Gouda 2008;Versieux et al. 2010;Gomes-da-Silva et al. 2012;Barfuss et al. 2016;Leme et al. 2017a;Faria et al. 2021) (Table S1). ...
... Anatomical leaf traits were synapomorphic for the xeric clade of Pitcairnioideae and they were important to support a more coherent circumscription of this group (Santos- Gomes-da-Silva et al. 2019). Here we investigated the evolution of two anatomical characters (character iii and iv) that were associated to xeric environments by other bromeliad genera that occur in dry habitats as does Stigmatodon (Tomlinson 1969;Benzing 2000;Versieux et al. 2010;Santos-Silva et al. 2013;Silva et al. 2020;Faria et al. 2021). ...
The genus Stigmatodon occurs in vertical and bare granite slopes, typical of the inselbergs of the Brazilian Atlantic Forest. Here, we present the first broad phylogenetic analysis focused on Stigmatodon , sampling a total of 83 terminals, including 16 of the 20 species of the genus and the morphologically similar species of Vriesea . We conducted a phylogenetic analysis using two plastid markers ( matK and rps16-trnK ) and the nuclear gene PHYC to infer phylogenetic relationships and reconstruct ancestral states for ecological and morphological characters. Our results suggest the monophyly of Stigmatodon as originally circumscribed is only possible with the inclusion of morphologically and ecologically similar Vriesea species. In addition, the morphological and anatomical traits led us to propose a new circumscription for the genus, combining eight species of Vriesea to Stigmatodon as S. andaraiensis , S. freicanecanus , S. lancifolius , S. limae , S. oliganthus , S. pseudoliganthus , S. vellozicolus , and S. zonatus . The stomata positioned above the ordinary epidermal cells, the adaxial water-storage parenchyma with axially elongated cells, the stamens positioned in two groups of three on each side of the corolla, and the tubo-laciniate stigma are exclusive to Stigmatodon in its new circumscription. These new morphological and phylogenetic results constitute a relevant contribution to the taxonomy and evolution of Bromeliaceae, one of the most diverse and ecologically important families of flowering plants of the Neotropics.
... In this work, the leaf and root anatomy of the plants were in accordance with previous reports of A. imperialis as well as other species of the Bromeliaceae family (Zorger et al. 2019, Martins et al. 2020a, Silva et al. 2020b, Faria et al. 2021. Even though Se modulated the root architecture of A. imperialis plants, this metalloid did not interfere with the roots' anatomical traits. ...
Elements not usually included in culture medium formulations, such as selenium (Se), may have beneficial effects on micropropagated plants. We evaluated the effects of Se on the physiological and anatomical responses of Alcantarea imperialis during in vitro culture. Plants were cultured in a medium containing a gradient of Se concentrations (0, 4, 8, 16, or 32 µM Se). After 56 d, the growth traits, chlorophyll a fluorescence, and root and leaf anatomy were analyzed. The fresh mass declined at the highest Se concentration. Higher Se concentrations induced bigger stomata, while the stomatal density decreased. Plants cultured with Se had improved PSII and PSI electron transport. This led to higher values of the total performance index. Thus, Se-induced plants showed a higher electron transport dynamics and energy conservation from water to PSI and developed anatomical traits that can favor tolerance to water deficit.
... Most Tillandsia species are epiphytes and adapted to xeric environments and have in their leaves epidermic trichomes capable of absorbing water and nutrients (Faria et al., 2021;Ha et al., 2021;Papini, Tani, Falco, & Brighigna, 2010;Scatena & Segecin, 2005). It is also possible to find species with rupicolous or saxicolous habits (Stefano et al., 2008). ...
... Together with the roots, they are used for fixation, which justifies the more elaborate character of the trichomes, enhancing the shield-fluid contact area. Thus, it favors water and nutrients' absorption as the Tillandsia species use the trichomes instead of the roots for this function (Faria et al., 2021;Ha et al., 2021;Scatena & Segecin, 2005). ...
Tillandsia L. is the largest genus of the family Bromeliaceae, containing 755 species and seven subgenera. Morphoanatomical studies of leaves provide useful characteristics to phylogenetic, taxonomic, and ecological analyses. This study aims to characterize and compare the leaves of 24 species of the four subgenera of Tillandsia that occur in Bahia and also perform adaptative inferences to environmental responses. The results of the species' morphoanatomical studies were compared through dissim-ilarity analysis. The species have rosulate leaves with varying lengths and widths. The peltate trichomes present variation in the indument density and the length of their wing and central disk. The stomata are longitudinally distributed in one or both sides of the limb. The mesophyll is dorsiventral and presents aquiferous and chlorophyllic parenchymas. The vascular bundles are collateral and partially covered by fibers, except for Tillandsia linearis. Based on the dissimilarity analysis, it was possible to identify the formation of five groups. Group G1 was composed of T. linearis, which diverged from the other species of the subgenus. Group G2 was formed by the remaining species of the subgenus Phytarrhiza. G3 and G4 presented the species of the subgenus Diaphoranthema and Tillandsia, respectively. Group G5 gathered 11 species of the subgenus Anoplophytum and presented higher variability than the other subgenera. Based on the results, the morphoanatomical characteristics can be used to characterize and group Tillandsia species, besides confirming the morphological variability of these species to the epiphyte habit in different environments, especially xeric ones.
