Two new of (, ) from Brazil

Leonardo M. Versieux1 and Maria das Graças Lapa Wanderley2

1 Instituto de Botânica, Seção de Curadoria do Herbário, Av. Miguel Stéfano 3687, São Paulo, SP, 04301-902, Brazil; e-mail: [email protected] 2 Instituto de Botânica, Seção de Curadoria do Herbário, Av. Miguel Stéfano 3687, São Paulo, SP, 04301-902, Brazil; e-mail: [email protected]

Abstract. Alcantarea turgida and A. tortuosa are described and illustrated as new species from southeastern Brazil. The relationships of A. turgida with A. extensa and with A. vinicolor and of A. tortuosa with A. farneyi and with A. benzingii are dis- cussed. Key words: Alcantarea, Brazil, Bromeliaceae, Minas Gerais, Rio de Janeiro, Tillandsioideae.

Resumo. Alcantarea turgida e A. tortuosa são descritas e ilustradas como novas es- pécies do Sudeste do Brasil. Os relacionamentos de A. turgida com A. extensa e com A. vinicolor e os de A. tortuosa com A. farneyi e com A. benzingii são discutidos.

In their monograph of Bromeliaceae sub- Alcantarea (sensu Grant, 1995a, 1995b, family Tillandsioideae, Smith and Downs 1995c; Till, 2000) is a morphologically well (1977) treated Alcantarea (E. Morren ex defined , which differs with in Mez) Harms as a subgenus of Vriesea Lindl. the long-linear ephemeral petals (ca. 10 to 15 (Vriesea subgen. Alcantarea E. Morren ex times longer than wide) that become flaccid Mez), following the classification used by or spiraled right after anthesis, and in the Mez (1894) in Flora Brasiliensis. Harms seeds that have both an apical and a basal (1929) was the first author to treat Alcantarea coma. Although easily separated from as an independent genus from Vriesea. Later, Vriesea by its morphological traits, recent Grant (1995a, 1995b) excluded one Central molecular studies by Barfuss et al. (2005) in- American and one West Indian species that dicate that Alcantarea is a monophyletic had been maintained within the subgenus in group, but its recognition as an independent previous works due to misinterpretations of genus is questionable because it turns floral characters and re-established Al- Vriesea into a paraphyletic group. cantarea as a genus. This geographically nar- Including the two new species proposed rower definition turned Alcantarea into a here and one recently published by Grant genus endemic to eastern Brazil, where it is (2003), Alcantarea comprises 20 species of confined to the states of Bahia, Minas Gerais, saxicolous , usually forming very large Espírito Santo, and Rio de Janeiro (Grant, rosettes and frequently reaching two to five 1995a, 1995b, 1995c); it was recently re- meters high while flowering. Alcantarea discovered in the state of São Paulo, which is species grow from near sea level up to the southernmost distributional limit of the 1900 m, mainly on gneiss-granitic inselbergs genus (Baker, 1889; Versieux & Wanderley, inside the Atlantic rainforest habitats of east- unpubl. data). ern Brazil, and disjunctly on quartzite out-

Brittonia, 59(1), 2007, pp. 57Ð64. ISSUED: 30 March 2007 © 2007, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. 58 brittonia [VOL. 59 crops in the Espinhaço mountain range, be- 0.8Ð1.2 m long, with internodes of 3.5Ð tween the states of Bahia and Minas Gerais. 6.5 cm long, green underneath the bracts, Despite its great ornamental appeal and its wine-colored maculate on sun exposed por- broad use as a garden , Alcantarea is tions, glabrous or nearly so; scape bracts: the poorly represented in herbaria, probably due proximal ones subfoliaceous and suberect, to the large size of many of the species, the distal ones spreading, recurved toward which makes collection of specimens diffi- apex, broadly lanceolate, with an ovate base cult. Species delimitation and identification and a caudate and a dark wine-colored mu- of incomplete specimens are also a chal- cronate apex, completely green maculate lenge. The conservation of natural popula- with red or totally red at the base and green tions and habitats of Alcantarea is of consid- maculate with wine-red toward apex, brown erable concern due to fires and the extraction lepidote, involute, distinctly nerved; inflores- of adult plants from the wild for the horticul- cence once branched, pyramidal to elliptic in ture trade. outline, 0.6Ð1.3 × 0.7Ð1 m, main axis with 4Ð The descriptions of two species of Al- 9 cm long internodes; flowering branches cantarea new to science presented here con- subdensely arranged, 14Ð32 in number, (3Ð) stitute partial results of a large scale system- 10–18-flowered, peduncle arcuate, the mid atic revision of the genus undertaken by the portion patent, and the terminal portion authors. slightly erect or the entire branch weakly ar- cuate; rachis geniculate, 18Ð27 cm long, Alcantarea turgida Versieux & Wand., sp. 4 mm diam., olive-green toward apex, nov. Type: Brazil. Minas Gerais: Con- glabrous, with 3.5 (proximal) to 1 cm (distal) ceição do Mato Dentro, alto da cachoeira long internodes separating the flowers; pri- do Tabuleiro, 19¡08'58.7"S, 43¡33'05.3"W, mary bracts broadly ovate, 6Ð9 × 6 cm at the 1300 m, 22 Dec 2005, Versieux et al. 260 base, with a 2Ð6 × 0.6 cm narrowly triangular (holotype: SP; isotypes: BHCB, MBM, and caudate apex, apex with a prominent vein RB, SEL, SPF, VIC). (Figs. 1, 2) and a recurved and rigid apiculus, shorter than the peduncles, green at the central por- A (L.B. Sm.) J.R. Grant et a A. tion and wine-colored along the margins and vinicolor (E. Pereira & Reitz) J.R. Grant affinis, sed plantis altioribus, inflorescentiis congestioribus, bracteis at the top or totally wine-colored, brown lepi- florum, sepalis, petalis et appendicibus petalorum ma- dote, involute, spreading, coriaceous, dis- joribus differt. (Carrière) Harms tinctly nerved; peduncles cylindrical to similis, sed vaginis et laminis foliorum angustioribus, slightly flattened, 9Ð22 cm long, 6Ð7 mm bracteis primariis quam pedunculis valde minoribus, al- abastris imbricatis, turgidis, nec complanatis differt. diam. at the base, glabrescent, bearing 1Ð3 sterile bracts; sterile bracts suborbicular, Plants saxicolous, short caulescent, flower- apex obtuse, discretely mucronate, 3Ð ing up to 2.5 m high; rosette (0.7Ð)1.3Ð1.7 m 3.7 × 3Ð3.2 cm, glabrous abaxially, brown diam., crateriform; leaves numerous, spread- lepidote adaxially, involute, erect, symmetric ing, arcuate; sheaths broadly ovate 19Ð or nearly so, coriaceous, distinctly nerved, 27 × 8Ð19 cm, pale brown adaxially, dark carinate toward apex; floral bracts convex, brown becoming pale brown with dark wine- suborbicular, apex obtuse, discretely mu- red color inclined lines at the top abaxially, cronate, 3.4Ð3.7 × 3.6Ð4(Ð4.4) cm, glabrous densely brown lepidote on both surfaces; and lustrous abaxially, brown lepidote adaxi- blades linear, long-attenuate, 61Ð75 × 7.5Ð ally, green at the central portion and wine- 11 cm, green, lustrous, inconspicuously dark colored along the margins, membranaceous wine-red along the margins, wine-colored and pale yellow along the margins and coria- spotted on both sides or only abaxially, lepi- ceous at the central portion on the dried ma- dote on both sides but glabrescent toward the terial, distinctly rugose-nervate, carinate, in- apex adaxially, coriaceous, distinctly nerved flated and densely imbricate at the branches with some prominent longitudinal veins; apical portions; pedicels obconic, 8Ð11 mm blade apex acute to cuspidate, totally green long, ca. 4 mm diam.; flowers odorless, not or dark wine-red colored, recurved, apiculus secund; sepals obovate, apex acute and twisted; scape erect, stout, 2Ð4 cm diam., minutely apiculate, 3.6Ð4(Ð4.4) × 1.4Ð1.8 cm, 2007] versieux & wanderley: alcantarea (bromeliaceae) 59

Fig. 1. Alcantarea turgida. A. Habit. B. Leaf (abaxial surface). C. Flowering branch. D. Floral bract. E. Sepal. F. Petal and antipetalous stamen. G. Gynoecium. H. Ovary and hypanthium (longitudinal section). I. Stigma. J. Ovule. (From the holotype.) 60 brittonia [VOL. 59

Fig. 2. Map showing the distribution of Alcantarea turgida (black squares) in Minas Gerais, and of A. tortuosa (black dot) in Rio de Janeiro.

exceeding the floral bracts for ca. 1 cm, free, with an elliptic apical appendage, restricted sparsely brown lepidote adaxially, green with to the upper ¹⁄₂-portion of the ovary; fruit an membranaceous hyaline margins, ecarinate; ellipsoid, rugose, dark castaneous, 5.3Ð petals long-linear with an acute apex, yellow 6.2 cm long capsule; seeds narrowly elliptic, in flower buds and immediately after the an- brown, ca. 6 × 1 mm; seed appendages: fim- thesis, fading quickly to creamy white, briate, the apical ones pale brown, ca. 15 mm spreading, weakly twisted, 9.3Ð11 × 1Ð long, the basal ones whitish, ca. 7 mm long. 1.2 cm, free, glabrous, inconspicuously white Distribution and ecology. Alcantarea spotted, margins entire with the distal ¹⁄₂-por- turgida grows as a heliophyte or semi- tion slightly undulate; petal appendages 2, sciophyte on rocky outcrops in the southern ligulate, 3.4Ð3.7 × ca. 0.3 cm, longitudinally portion of the Espinhaço range, Rio Doce adnate to the petal for 2.9Ð3.1 cm, the apex drainage basin, state of Minas Gerais, in free, apices overlapping and completely hid- southeastern Brazil. It was observed occur- ing the ovary from a top view; stamens free; ring in semi-deciduous montane forest and in filaments terete, 9Ð10 cm long, ca. 2 mm campo rupestre (i.e., grassland on rocky thick, white; anthers sagittate, 1.2Ð1.6 cm soils) vegetation of the Cipó range, usually long, ca. 1 mm thick, dorsifixed near the above 1000 m elevation. Its flowers are fre- base, rimose; style terete, white, 9.3Ð10 cm quently visited by hummingbirds, especially long ca. 2 mm diam.; stigma lobes white, ca. by Phaethornis pretrei. Bees are also com- 4 × 1 mm, spreading; ovary ca. 22 × 6 mm, mon visitors, perforating the petals to pillage narrowly ovate, hypanthium ca. 8 mm in pollen grains. Based on the IUCN criteria length; ovules numerous, ca. 1.6 mm long, (IUCN, 2001), A. turgida can be considered 2007] versieux & wanderley: alcantarea (bromeliaceae) 61

“Near Threatened” (NT). Regardless of its Minas Gerais—treated here as additional ma- occurrence within three different protected terial of A. turgida—as Vriesea geniculata areas, the common fires in adjacent grazing (Wawra) Wawra [basionym of Alcantarea areas as well as the collection pressure for geniculata (Wawra) J.R. Grant]. In contrast to the ornamental trade constitute real threats A. turgida, A. geniculata is an endemic species for this species in the near future. of Rio de Janeiro that occurs mainly at serra Phenology. Flowering individuals were dos Órgãos (Leme, 1997), and has smaller observed in June and from December to Feb- sepals, peduncles, and petal appendages; also, ruary. Fruiting in April. the primary bracts equal or surpass the sterile Etymology. The specific epithet is an allu- base of the branches, the inflorescence is more sion to the floral bracts that are densely im- densely arranged, and the foliage is com- bricate and turgid, and completely filled with pletely green while it is wine-colored spotted copious transparent viscose nectar that drips abaxially and may be totally wine-red-colored from the apex of the flowering branches. on the distal adaxial surface in A. turgida.

Additional specimens examined. BRAZIL, Minas Alcantarea tortuosa Versieux & Wand., sp. Gerais: São Gonçalo do Rio Abaixo, Estação Ecológ- nov. Type: Brazil. Rio de Janeiro: Santa ica de Peti - Cemig, Mirante do Cruzeiro, 23 Apr 2002, Lombardi et al. 4748 (BHCB, RFA); Itabira do Campo, Maria Madalena, afloramento rochoso Jun 1902, Mello-Mattos s.n. (R 46318); Santana do Ria- próximo ao pico da pedra do Desengano, cho, Parque Nacional da Serra do Cipó, 19¡13'23.4"S, 21¡53'59"S, 41¡54'40.2"W, 1900 m, 25 43¡30'06.6"W, 1200 m, 11 Dec 2005, Wanderley et al. Nov 2005, Versieux & Calvente 240 (holo- 2488 (SP). type: SP). (Figs. 2, 3) Alcantarea turgida is vegetatively similar to Alcantarea farneyi (Martinelli & And. Costa) J.R. A. extensa (L. B. Sm.) J. R. Grant and to A. Grant affinis, sed habita majore, foliis rosulatis, in- fundibuliformibus, latioribus, inflorescentiis longioribus, vinicolor (E. Pereira) J. R. Grant. In compari- paniculatis et bracteis florum carinatis differt. son to the latter two species, A. turgida has a different type of inflorescence, with the main Plants saxicolous, short caulescent, flower- axis almost straight (vs. slightly geniculate), ing 0.8Ð1.4 m high, propagating by basal the flowering branches more numerous, shoots; leaves ca. 15 in number, suberect, shorter (also more densely arranged), and densely arranged in a funnelform rosette; patent and ascending toward the apex or sheaths broadly ovate, 12Ð17 × 9Ð10.5 cm, weakly arcuate (vs. lax inflorescences with pale brown becoming soft green toward apex long and spreading flowering branches). Al- abaxially, whitish becoming greenish adaxially, cantarea turgida also has longer sepals, petals, densely brown lepidote on both sides; blades and petal appendages. Also, in the dried speci- sublinear, apex cuspidate, 24Ð42 × 4.5Ð6 cm, mens of A. turgida, the floral bracts are not as lustrous, green on both surfaces, with a narrow dark as in A. extensa. Alcantarea turgida also (i.e., ca. 0.8 mm wide) dark wine-colored line resembles A. imperialis (Carrière) Harms, but along the margins, glabrous, involute, suberect it can be separated from the latter by the with recurved apices, coriaceous, distinctly length of the primary bracts, which are shorter nerved, weakly plicate in cross section; scape than the peduncles, by the inflated floral bracts 60Ð75 cm long, exceeding the rosette, with and floral buds at the apex of the branches (vs. 4.5Ð7 cm long internodes, 1.3Ð1.7 cm diam., totally flattened floral buds), by the lustrous pale green underneath the bracts but dark foliage (vs. glaucous), by the flowers not se- wine-colored at exposed portions, glabrescent, cund (vs. secund), and by the form of the distinctly nerved, slightly curved; scape bracts: flowering branches, which are patent and as- the proximal ones subfoliaceous, the distal cending toward the apex or weakly arcuate ones ovate, apex long-attenuate and slightly re- (vs. pendulous, and with a sigmoid shape). It curved, 8Ð14 × 3Ð3.6 cm, densely white lepi- is also important to distinguish A. turgida dote and totally green becoming purple toward from A. geniculata (Wawra) J.R. Grant, since apex adaxially, sparsely lepidote and deep pur- Smith and Downs (1977) cited the collection ple becoming greenish with purplish-red dots Mello-Mattos s.n. (R) from Itabira do Campo, toward apex abaxially, margins with a very 62 brittonia [VOL. 59

Fig. 3. Alcantarea tortuosa. A. Habit. B. Leaf. C. Flowering branch. D. Floral bract. E. Sepal. F. Petal and antipetalous stamen. G. Gynoecium. H. Ovary and hypanthium (longitudinal section). I. Stigma. J. Ovule. (From the holotype.) 2007] versieux & wanderley: alcantarea (bromeliaceae) 63 narrow purple line all along, involute, suberect, neyi (Martinelli & And. Costa) J. R. Grant densely imbricate, the basal ¹⁄₃ firmly conceal- and Vellozia variegata Goeth. & Henrard ing the scape, distinctly nerved with some near the peak of the main inselberg of the prominent veins, weakly plicate in cross sec- Desengano State Park, Rio de Janeiro state, tion; inflorescence once branched, elliptic in Brazil. It has only been registered from this outline, 46Ð54 × 23Ð26 cm, main axis with 3Ð park, where it grows inside a unique habitat 4 cm long internodes; flowering branches laxly known as campo de altitude (i.e., high alti- to subdensely arranged, 4Ð12 in number, 6Ð14- tude grassland), which is associated with the flowered, pendulous to arcuate, some of them Atlantic rain forest domain and is very rich in distinctly tortuous or somewhat twisted; rachis endemic species (Martinelli, 1989). Based on geniculate, almost quadrangular in cross sec- the IUCN criteria (IUCN, 2001), A. tortuosa tion, 8Ð20 cm long, 4 mm diam., green, can be included in the conservation status of glabrous, with 2.7 (proximal) to 1.2 cm (distal) Endangered EN B1ab(iii) due to its restricted long internodes separating the flowers; primary area of occurrence, small population size, bracts broadly ovate, 5Ð6 × 3.2Ð3.6 cm, apicu- and projected decline in the quality of its late, slightly inflated, dark wine-colored at the habitat, since it grows right along the trail insertion line, along the margins and toward leading to the Desengano peak. the apex, and greenish in the central portion, Phenology. In floral bud during November, distinctly nerved, lepidote; peduncles flattened, blooming from December to January. elliptic in cross section, 4Ð9 cm long, 7 mm Etymology. It is named after the shape of diam. at the base, green, glabrous, bearing 1Ð2 the scape and of the inflorescence, which is sterile bracts; sterile bracts broadly ovate, tortuous. shortly attenuate, 3Ð4.2 × 2.5Ð3.5 cm, purplish green, glabrous or nearly so, symmetric, cari- Additional specimen examined. BRAZIL. Rio de nate; floral bracts broadly ovate, apex obtuse, Janeiro: Santa Maria Madalena, afloramento rochoso × próximo ao pico da pedra do Desengano, 21¡53'59"S, 2.8Ð3.6 2.5Ð2.8 cm, glabrous or nearly so, 41¡54'40.2"W, 1900 m. 25 Nov 2005, Versieux & Cal- green with few purple spots along the margins, vente 241 (SP). symmetric, membranaceous, carinate toward the apex; pedicels obconic, 4Ð6 mm long, 4Ð Alcantarea tortuosa is probably related to 5 mm diam., green; flowers odorless, not se- A. farneyi, but its unique set of characters cund; sepals elliptic, apex acute, 3.5Ð sets it apart from all other members of the 3.8 × 1.2Ð1.8 cm, equaling the floral bracts to genus. Future molecular work should indi- exceeding them for ca. 2 mm, free, symmetric, cate better its taxonomic relationships. Al- glabrescent, green, ecarinate; petals long-linear though both species are sympatric, and the with an acute apex, golden-yellow, inconspicu- blooming periods almost overlap, A. farneyi ously whitish at the base, recoiling and spiral- exhibits a graminoid habit, forming a sub- ing, 5.7Ð6 × 0.6Ð0.7 cm, free, symmetric, bulbous rosette, with very narrow (i.e., ca. glabrous; petal appendages 2, ligulate, 14Ð 1.5 cm wide) and involute leaves, ecarinate 20 × 1Ð2 mm, longitudinally adnate to the floral bracts, and a simple, pendulous, and petals for 1.1Ð1.6 cm; stamens free; filaments shorter (ca. 11 cm) inflorescence. On the terete, slightly flattened, 5.2Ð5.5 cm long, ca. other hand, A. tortuosa has a funnelform 1 mm thick, white; anthers sagittate, 1.2Ð rosette that accumulates water, much wider 1.5 cm long, ca. 1 mm thick, dorsifixed near the leaf blades, floral bracts carinate toward the base, rimose; style terete, white, 7.5Ð7.7 cm apex, and a branched and longer inflores- long, ca. 1 mm diam.; stigma lobes white, ca. cence. The similarities between these two 3 × 1 mm, erect; ovary ca. 11 × 3Ð4 mm, ovate, species reside in the presence of very narrow hypanthium ca. 5 mm in length; ovules numer- dark red margins along the leaf blades, col- ous, ca. 1.4 mm long, with an elliptic apical ap- oration and imbrication of bracts, and in pendage, restricted to the upper ¹⁄₂-portion of some floral characters such as the size of the the ovary; fruits and seeds unknown. sepals, the petals, and the petal appendages. Distribution and ecology. Alcantarea tor- Alcantarea tortuosa can also be distin- tuosa grows as a heliophyte on bare rocks guished from A. benzingii Leme, a species above 1900 m together with Alcantarea far- that occurs in similar habitat in Espírito 64 brittonia [VOL. 59

Santo state, and that was described as being Literature Cited closely related to A. farneyi. Alcantarea tor- Baker, J. G. 1889. Handbook of the Bromeliaceae. tuosa is recognized by the size and shape of George Bell & Sons, London. its scape, which is much thicker and tortuous Barfuss, M. H. J., R. Samuel, W. Till & T. F. Stuessy. (vs. slender and suberect), by the distinct 2005. Phylogenetic relationships in subfamily coloration of its deep reddish-purple bracts Tillandsioideae (Bromeliaceae) based on DNA se- quence data from seven plastid regions. American (vs. reddish-yellow), by the size and type of Journal of 92: 337Ð351. the inflorescence, which is branched (vs. Grant, J. R. 1995a. The resurrection of Alcantarea and shorter and simple inflorescence), by the size , a new genus. Tropische und Subtropische and type of the rosette, and by its wider Pflanzenwelt 91: 7Ð15. blades. ———. 1995b. Addendum to “The ressurection of Al- cantarea and Werauhia, a new genus” (Bromeli- aceae: Tillandsioideae). Phytologia 78: 119Ð123. ———. 1995c. Sinopse do gênero Alcantarea. Acknowledgments Bromélia 2: 24Ð26. ———. 2003. Alcantarea heloisae (Bromeliaceae), a We thank IBAMA and the Instituto Esta- new species from Rio de Janeiro State. Vidalia 1: 31Ð dual de Florestas do Rio de Janeiro for issu- 33. Harms, H. 1929. Bromeliaceae novae III. Notizblatt des ing the necessary plant collection permits and Botanischen Gartens und Museums zu Berlin- also the curators of the following herbaria for Dahlem 10: 801Ð802. providing loans of specimens or access to IUCN. 2001. IUCN Red list categories and criteria: Ver- collections: BHCB, HB, R, RB, RFA. Dr. sion 3.1. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, United Kingdom. Enrique Forero, Dr. Jason Grant, and one Leme, E. M. C. 1997. Contribuição ao estudo do gênero anonymous reviewer provided helpful com- Alcantarea II. Bromélia 4: 29Ð40. ments on the manuscript. We also thank Dr. Martinelli, G. 1989. Campos de altitude. Editora Index, Tarciso Filgueiras for helping with the Latin Rio de Janeiro, Brasil. diagnoses; Klei Sousa for the illustrations; Mez, C. 1894. Bromeliaceae. In: C. F. P. Von Martius, A. G. Eichler, & I. Urban (eds.), Flora Brasiliensis 3: Alice Calvente, Celso Paiva, and Pedro Viana 173Ð643. Leipzig, F. Fleischer, Germany. for assistance in the field; Dr. Renato de Smith, L. B. & R. J. Downs. 1977. Tillandsioideae Mello-Silva and Leonardo Lopes for the (Bromeliaceae). Flora Neotropica Monograph 14: identifications of the Vellozia and humming- 663Ð1492, New York, Hafner Press, USA. Till, W. 2000. Tillandsioideae. Pp. 555Ð571. In: D. H bird species, respectively. Financial support Benzing (ed.), Bromeliaceae: profile of an adaptive was provided by FAPESP to L. M. Versieux radiation. Cambridge University Press, Cambridge, and by CNPq to M. G. L. Wanderley. United Kingdom.