Tillandsia bonita (Bromeliaceae), a New Species from Mato Grosso do Sul, Brazil,with Notes on Leaf Anatomy
Leonardo M. Versieux,1,3 Nara Vasconcellos,2 and Gustavo Martinelli2
1Universidade Federal do Rio Grande do Norte, CB, Departamento de Botanica, Ecologia e Zoologia, Natal,Campus Universitario, Lagoa Nova, Natal, RN, Brazil, 59078-900
2Jardim Botanico do Rio de Janeiro, Pacheco Leao 915, Rio de Janeiro, RJ, Brazil, 22460-0303Author for correspondence ([email protected])
Communicating Editor: Austin Mast
Abstract—Tillandsia bonita is described as a new species and is illustrated. It is compared with Tillandsia didisticha, which we assume is themorphologically most similar species. The new species is the ninth taxon of this genus recorded for the flora of the state of Mato Grosso do Sul,central-western Brazil. Illustrations and a brief discussion on the leaf anatomy of the species are provided. Two additional species that havebeen overlooked by previous authors are included in the list of Bromeliaceae of Mato Grosso do Sul and an updated key for Tillandsia fromthat state is presented.
Keywords—Cerrado, new species, Parque Nacional da Serra da Bodoquena, plant anatomy, seasonal forest, Tillandsioideae.
The diversity of Bromeliaceae in Brazil is highest in areasof Atlantic rainforest, and a decrease in species richnessis observed toward the drier inland territory (Reitz 1983;Versieux and Wendt 2007; Martinelli et al. 2008; Versieux et al.2012). Nevertheless, drier areas may harbor endemic and xero-phytic species, the systematics, taxonomy, and conservation ofwhich have yet not received much attention (e.g. Miranda2001; Versieux and Wendt 2007; Versieux et al. 2010b).
The state of Mato Grosso do Sul (MS) in central-westernBrazil is covered by three different biomes: cerrado (savanna),the Pantanal wetland (including part of the chaco) and part ofthe Atlantic rainforest (IBGE 2004). The cerrado savanna has amosaic of physiognomies. It is severely threatened, and it isconsidered to be one of the areas of heaviest human impact inthe state of MS (Cavalcanti and Joly 2002). The region of theBodoquena Mountain Range in the western portion of MShas, in addition to the savannas, both tropical seasonal decid-uous and semi-deciduous forest along the rivers that areseverely threatened due to deforestation for logging or expan-sion of agricultural and cattle raising areas, making the foresta high priority for biodiversity conservation (Baptista-Mariaet al. 2009). The Bodoquena Range is nearly 300 km in length,varying from 20–50 km in width. It is mainly composed oflimestone and is surrounded by the Paraguay River depres-sion (Pantanal Plain) towards the north and west, the MirandaRiver basin to the east, and the Apa River basin to the south(Boggiani and Coimbra 1995).
Tillandsia L. encompasses nearly 610 species (excludingseveral natural hybrids), and it is the largest and most widelydistributed genus within Bromeliaceae, occurring through-out the American tropics and subtropics (Smith 1934; Luther2010). Tillandsia is an extremely polymorphic genus and isconsidered to be a young lineage within the monophyleticsubfamily Tillandsioideae (Barfuss et al. 2005). Tillandsiawas traditionally divided into seven subgenera: Allardtia,Anoplophytum, Diaphoranthema, Phytarrhiza, Pseudalcantarea,Pseudocatopsis, and Tillandsia (Smith and Downs 1977;Gilmartin 1983). More recently, systematic changes segregatedT. subgen. Pseudocatopsis as the genus RacinaeaM.A. Spencer &L.B. Sm. (Spencer and Smith 1993), and six species formerlyincluded in T. subgen. Allardtia were grouped into the genusViridantha Espejo (Espejo-Serna 2002). So, Tillandsia is cur-rently divided into six subgenera: Allardtia, Anoplophytum,Diaphoranthema, Phytarrhiza, Pseudoalcantarea, and Tillandsia
(Till 2000). However, recent molecular phylogenetic workindicates the para- or polyphyly of all these subgenera(Barfuss et al. 2005). Racinaea and Viridantha appear mono-phyletic, but separation of these from Tillandsia makes theremainder paraphyletic (Barfuss et al. 2005). In summary, thesubgeneric classification of Tillandsia is far from being satis-factory (Espejo-Serna 2002; Barfuss et al. 2005).After examining a blooming specimen of Tillandsia col-
lected during recent field expeditions to the Parque Nacionalda Serra da Bodoquena, close to the border of Brazil withBolivia, we concluded that it is a new species. This taxoncorresponds to the ninth species of Tillandsia recorded forthe state of Mato Grosso do Sul, including two other species(Tillandsia buchlohii Rauh and T. streptocarpa Baker), whichhave been cited in the literature (Smith and Downs 1977;Till 1996) but were missing in the Brazilian Bromeliaceaechecklist (Forzza et al. 2011). Here we provide an updatedkey for the species of Tillandsia from Mato Grosso do Sul,aiming to contribute to the future monograph of Bromeliaceaefor the Flora of the State of Mato Grosso do Sul.
Materials and Methods
The morphological description of the new species is based on a livingspecimen cultivated at the Rio de Janeiro Botanical Garden and on itsfreshly collected flowers, as well as on the herbarium specimens preparedwhen the plant was collected in the field (RB; Martinelli 16923, 16923a,16923b). Conservation assessment for the new species was based on fieldobservation, applying the IUCN red list category criteria (IUCN 2001).Transversal and paradermal sections were taken using a razor blade,bleached by the use of sodium hypochlorite, stained with 0.05% astrablue and safranin (Bukatsch 1972) and mounted in water for temporaryobservation and to obtain photomicrographs. To observe trichomes,the abaxial epidermis was gently scraped with the razor blade and thematerial was then stained as described above. A stigma and an ovuleconserved in 70% ethanol were directly mounted on temporary slidesand observed under the microscope.
Results
Tillandsia bonita Versieux & Martinelli, sp. nov.—TYPE:BRAZIL. Mato Grosso do Sul: Mun. Bodoquena, ParqueNacional da Serra da Boquena, estrada pela FazendaRemanso. Afloramento de rocha e matas ciliares do rioSalobra, 06 Dec. 2010, Martinelli 16923 (holotype: RB450408!; isotype: UFRN!).
75
Diagnosis—The new species is placed in the genus Tillandsiadue to the presence of petals lacking appendages and aninflorescence of a distichous flowered, double spike andin T. subgen. Anoplophytum due to the presence of symmet-ric sepals, included stamens, slender style, and stamenspresenting plicate filaments (Smith and Downs 1977). The fol-lowing combination of characters distinguishes Tillandsiabonita from T. didisticha Baker (sensu Tardivo 2002): the size ofthe flower (2.6 vs. 1.8–2.0 cm respectively), length of the sepals(1.5 vs. 1.1 cm), length of the petals (2.4 vs. 1.5–2.0 cm), lengthof the style (13 vs. 9 mm), floral bracts color (green vs. red),floral bract size (1.9–2.2 + 0.7–0.9 cm vs. 1.1–1.5 + 0.5 cm),and floral bract indumentum (glabrous to inconspicuouslylepidote vs. entirely lepidote or lepidote only toward the apex)(Table 1).Description—Rupicolous herb, ca. 30 cm tall when flow-
ering, short caulescent (Fig. 1). Rosette ca. 22 + 22 cm, nar-row infundibuliform, and slightly inflate at the base (Fig. 2).Leaves ca. 16, suberect; leaf sheath ca. 4 + 4 cm, broadlyovate to nearly orbicular, pale green passing to wine-red onboth surfaces in vivo, densely hyaline-lepidote on both facesin sicco, coriaceous; leaf blade 13–17 + 1.5–2.4 cm, narrowlytriangular, wine-red, concolorous on both surfaces, denselylepidote on both surfaces, trichomes densely arranged inrows abaxially, canaliculate, coriaceous, suberect, arched, dis-tinctly nerved with 1–2 prominent nerves; apex acute, atten-uate, wine-red, straight. Peduncle ca. 15 + 0.4–0.6 cm, erect,straight to slightly curved, terete, green, sparsely lepidoteclose to the nodes; internodes 8, 1.5–2.0 cm long; peduncle-bracts 2–5 + 1.7–2 cm, ovate, apex acuminate, the proximalones wine-red, the distal ones wine-red but green along themargins, erect, finely nerved, the proximal ones coriaceous,the middle and distal ones chartaceous and concealing theinternode, densely white-lepidote toward the apex; marginsgreen, thinner, white-lepidote. Inflorescence ca. 12 + 10 cm(not including the peduncle), a once-compound double-spike,obovoidal, erect, rachis straight, internodes 0.7–1.4 + 0.6 cm;primary bracts longer than the stipes, green, densely white-lepidote toward the apex, erect, tightly clasping the base of thespike, finely nerved, chartaceous; inflorescence branches ca. 8,5–13-flowered; stipes ca. 15–20 + 5 mm, weakly complanate,distinctly nerved, green, glabrous; sterile bracts 1–3, similarto the floral bract; rachilla 3.0–5.5 cm, flexuous, green, glabrous,not visible in vivo but with the nodes apparent in sicco; inter-nodes ca. 0.8 + 0.2 cm; floral bract 1.9–2.2 + 0.7–0.9 cm,elliptic, apex acute, green, sparsely hyaline-lepidote abaxi-ally, inconspicuously lepidote to glabrous adaxially, chartaceous,ecarinate, margins membranaceous, hyaline in sicco. Flowersca. 2.6 cm long, odorless, diurnal, distichously arranged,
erect, shortly pedicellate; pedicels ca. 2 + 1 mm; sepalsca. 1.5 + 0.4 cm, shortly connate (ca. 2 mm) at the baseadaxially, elliptic, apex rounded, symmetric or nearly so, notexceeding the floral bracts, green to greenish-hyaline along themargins, glabrous, ecarinate, margins membranaceous; petalsca. 2.4 + 0.25 cm, ligulate, apexobtuse, slightly recurvedat anthe-sis, white; stamens included; filaments 1.3–1.5 cm + 1 mm,complanate, shortly plicate below the connective,white; anthersca. 13 + 1 mm, linear in vivo to sagittate in sicco, yellow,dorsifixed near the base; style ca. 13 mm long, graduallydiminishing in diameter toward the apex, terete, white;stigma trifid, lobes spreading (Fig. 3H), ca. 4 + 1 mm, white,simple-erect (type I sensu Brown and Gilmartin 1984, 1989) withtubular papillae (Fig. 3I); ovary ca. 4 + 2 mm, ovoidal, trisulcate,brownish-green in sicco; ovules (Fig. 3J) ca. 0.05 cm long, ovate,numerous. Figures 1–2, 3H–J.
Phenology—Flowering in cultivation fromAugust throughOctober.
Distribution—So far the species is only known fromParque Nacional da Serra da Bodoquena, Bodoquena, in thewestern portion of the state of Mato Grosso do Sul, central-western Brazil.
Ecology—It grows forming dense heliophilous populationsover limestone outcrops along the semi-evergreen riverinetropical forest.
Conservation—IUCN (2001): provisionally treated as LeastConcern (LC), because it is protected within a Braziliannational park.
Etymology—The epithet chosen is a reference to the munic-ipality of Bonito, which is a famous place and an ecologicalsanctuary along the Serra da Bodoquena. This region has sev-eral rivers with clear and bicarbonated water and harbors anoteworthy biodiversity (e.g. Bueno et al. 2007; Uetanabaroet al. 2007; Dalzochio et al. 2011) including ornithophilousspecies of Bromeliaceae (e.g. Araujo and Sazima 2003; Fariaand Araujo 2010). Additionally, the word “bonita” is thePortuguese adjective equivalent to good-looking, which isindeed an attribute of the contrasting wine-red foliage, greenbracts, and white flowers of this plant.
Paratypes—BRAZIL. Mato Grosso do Sul. Bodoquena.Parque Nacional da Serra da Bodoquena, estrada pelaFazenda Remanso, afloramento de rocha e matas ciliares dorio Salobra, 04 Dec 2010, G. Martinelli 16923a (RB 545183).Parque Nacional da Serra da Bodoquena, estrada pelaFazenda Remanso, afloramento de rocha e matas ciliares dorio Salobra, living plant for cultivation, bloomed at the Rio deJaneiro Botanic Garden inside the ‘Burle Marx’ Greenhouseunder the number RBvb 1226, 02 Oct 2011, G. Martinelli16923b (RB 547839, UFRN).
Leaf Anatomical Characterization—The epidermis infront view presents rectangular cells with sinuous and thickwalls (Fig. 3F). The abaxial and adaxial epidermis are coveredby nearly circular trichomes, which are radially symmetric ornearly so (Fig. 3C, F, G). Trichome cells are organized concen-trically, following the 4 + 8 + 16 + 64 pattern (Fig. 3G). Thetrichomes emerge from a lower level than the other epidermalcells, and the stalk is uniseriate and composed of few cells.The mesophyll is composed of a multicellular water storageparenchyma that usually presents elongated or rounded, thin-walled cells and occupies nearly 1/3 of the upper mesophyll(Fig. 3A). The remaining 2/3 of the mesophyll are filledwith rounded chlorenchymatous cells and a water storageparenchyma. The first is positioned immediately below
Table 1. Morphological comparison between Tillandsia bonita andT. didisticha sensu lato (according to Tardivo, 2002).
Fig. 1. Tillandsia bonita. A. Habit of the cultivated plant. B. Detail of the inflorescence at full-bloom. C. Floral bract. D. Sepal. E. Flower. F. Detail of thepetal and stamens. G. Stamen. H. Pistil. I. Ovary. J. Detail of the trifid stigma. From Martinelli 16923b. Illustration by Paulo Ormindo.
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Fig. 2. Tillandsia bonita. A. Habit of the cultivated plant. B. Detail of the inflorescence in the beginning of blooming. C. Final stage of blooming.D. Photomicrograph of abaxial surface of the leaf blade showing the densely arranged appressed trichomes. E. Detail of the peduncle, showing thedenser indumentum close to the node. F. Detail of the flowering bracts margins. G. Detail of the leaf apex. H. Detail of the petal apex. I. Detail of theliquid preserved ovary (all photos from Martinelli 16923b).
78 SYSTEMATIC BOTANY [Volume 38
the upper water storage parenchyma and between the vas-cular bundles. The water storage parenchyma is also depos-ited below each vascular bundle (Fig. 3B) and idioblastcontaining raphides can be seen within this tissue (Fig. 3D).
Collateral vascular bundles are present in a single series andhave similar diameter (Fig. 3A). Stomata complexes aretetracytic and restricted to the abaxial face. The subsidiarycells are parallel to the guard cell (Fig. 3E).
Artificial key to the species of TILLANDSIA from MATO GROSSO DO SUL, BRAZIL
Fig. 3. Leaf anatomy and floral micrographs of Tillandsia bonita. A. Cross-section of the mid portion of the leaf blade (x40). B. Detail of the abaxialmesophyll and vascular bundles (x100). C. Detail of the abaxial epidermis covered by trichomes (x400). D. Idioblast containing raphides (x400). E. Stomain the abaxial surface (x400). F. Detail of the sinuous walls of the abaxial epidermis cells and the central portion of the trichomes (x400). G. Detail of anisolated trichome (x100). H. Stigma, liquid preserved (x100). I. Detail of the tubular projections of the stigma papillae (x400). J. Ovule, liquid preserved(x100) (all photos from Martinelli 16923b).
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This new species is placed in T. subgen. Anoplophytum dueto its transversely plicate filaments (Smith 1934). Althoughoriginally published as a genus by Beer (1854), Anoplophytumis now treated as a subgenus of Tillandsia (Smith and Downs,1977; Till, 2000). It is characterized by the presence of trans-versely plicate filaments (Evans and Brown 1989), spiralphyllotaxis, odorless flowers, symmetric sepals, and usuallyblue, rose, or white flowers with a tubular corolla (Till 2000).Simple erect stigmas are also frequent within this subgenus(Brown and Gilmartin 1989) and are present in T. bonita.Species of T. subgen. Anoplophytum occur in mesic or xeric
environments (Till 2000). According to Smith and Downs(1977), as reviewed by Till (2000), this subgenus groups twoassemblages of species: (1) strongly xeromorphic speciesmostly bearing dense distichously white-flowered spikesand concentrated in the Andes and (2) a Brazilian group ofxeromorphic and mesomorphic species bearing compounddistichous or polystichous spikes mostly with dark rose orblue petals. The first group mentioned above was transferredto T. subgen. Allardtia by Till (2000), and the separation ofthese two subgenera is still considered problematic anddifficult (Tardivo 2002). It now appears that neither ofthese subgenera is monophyletic (Barfuss et al. 2005), and sowe take what we consider to be the more conservativeapproach of recognizing a broadly circumscribed T. subgen.Anoplophytum (sensu Smith and Downs, 1977; Tardivo 2002),while awaiting the publication of ongoing molecular phylo-genetic investigations in Tillandsia (Barfuss et al. 2011).However, if we were to follow the segregation proposed byTill (2000), then we would place T. bonita within T. subgen.Allardtia with T. didisticha.In Tardivo’s (2002) revision of the T. subgen. Anoplophytum,
a broad circumscription was adopted for T. didisticha, pro-posing three other species as synonyms: Tillandsia barrosoaeW. Till, T. pucaraensis Ehlers and T. vernicosa Baker. Tardivopoints out that T. didisticha has a broad area of occurrenceencompassing Bolivia, Argentina, Paraguay, and Brazil, inthe states of Distrito Federal, Goias, Mato Grosso, MatoGrosso do Sul, and Parana. According to this author thebroad occurrence of T. didisticha has an influence on thegreat variability seen in leaf position and rosette shapes,size and number of inflorescence branches, and floral bractssize and indumentum. In our diagnosis we adopted for com-parison purposes all the measurements presented by Tardivo(2002), who examined more than 50 specimens (not includingduplicates). Basically, the main differences seen between thetwo species are related to flower and flower parts size, aswell as the color of the floral bracts. Additionally, T. bonita
has nearly glabrous floral bracts, in contrast with mostmorphotypes seen of T. didisticha.
The leaf anatomy of T. bonita follows patterns seen for theentire genus (e.g. Braga 1977; Loeschen et al. 1993; Mosti et al.2005; Scatena and Segecin 2005). Many of the characteris-tics observed here, such as epidermis with thick and sinuouswalls, extensive water storage parenchyma, and the densityof trichomes may be seen as adaptations to endure alithophytic lifestyle, as seen in other Bromeliaceae (Freschiet al. 2010; Versieux et al. 2010a). This species is the semi-mesic type of Gilmartin (1983), presenting a poorly devel-oped or no tank and narrow leaf blades with inconspicuous,appressed scales.
Another distinctive characteristic of the new taxon worthnoting is the complete dominance of the wine-red leaves, acharacter that was observed in all individuals in differentpopulations and that has not changed under cultivation.Anthocyanin accumulation in leaves may be related to abi-otic effects such as nutrient deficiency, excess light, UV-Bradiation, water stress (osmoregulation), and herbivory(Close and Beadle 2003; Manetas 2006), which may beavoided due to aposematic coloration in red leaves (Cooneyet al. 2012). In the case of T. bonita, 10 individuals were takento Rio de Janeiro Botanic Garden greenhouses and afternearly one year of cultivation no change in color of the leaveswas noticed. It would be interesting to check in the futurewhether the seedlings of this species are already wine-redcolored. This color morph pattern has been noticed for otherspecies of Tillandsia (e.g. T. tenuifolia L.) and is much appreci-ated by horticulturists (Isley 2009).
Acknowledgments. The authors thank Derek Butcher, Len Colgan,two anonymous reviewers, and associate editor Austin Mast for theirconstructive comments on the manuscript. We are grateful to PauloOrmindo for the beautiful illustration. Instituto Chico Mendes (formerlyIBAMA) provided collection permits. The bromeliad living collection ofRio de Janeiro Botanic Garden is supported by AMIL (AssistenciaMedica Internacional SA). Funding for G. Martinelli to conduct field-work was provided by the Brazilian National Centre for Flora Conserva-tion (CNCFlora) and by the Global Environmental Facility (GEF/PROBIOII).
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