phytotaxa 117 (2): 42 phytotaxa article phytotaxa

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Phytotaxa 117 (2): 42–50 (2013) www.mapress.com/phytotaxa/ Article

http://dx.doi.org/10.11646/phytotaxa.117.2.2

A new yellow-flowered ornithophilous Vriesea and an illustrated collection of the bromeliads from Pico Alto, Serra do Baturité, Ceará State, Northeastern Brazil

LEONARDO M. VERSIEUX1, EDUARDO C. TOMAZ1, MÁRCIA FORTUNATO2 & CHRISTIANO

VEROLA2

1Departamento de Botânica, Ecologia e Zoologia, Centro de Biociências, Universidade Federal do Rio Grande do Norte, 59078-900, Natal, Rio Grande do Norte, Brazil. E-mail: [email protected] and [email protected] de Biologia, Centro de Ciências, Universidade Federal do Ceará, 60455-760 Fortaleza, Ceará, Brazil. Email: [email protected] and [email protected]

Abstract

Vriesea baturitensis is described and illustrated as a new species. It is compared with V. friburgensis and V. rodigasiana, which we consider to be morphologically the most related species. The new taxon occurs in isolated Atlantic forest patches along the Baturité mountain range, in central-north Ceará state, Northeastern Brazil. It is characterized by the compact and regular rosette, a rounded leaf apex, stiff and erect peduncle, peduncle and primary bracts bright yellow, and the particular colors and sizes of the floral bracts and sepals. The humid habitat where the new species was found, known in Brazil as brejo de altitude, is surrounded by the Caatinga (Brazilian dry woodland) and due to its climatic conditions supports a rich flora of epiphytes. In Pico Alto we collected and photographed six different species of bromeliads, two of each being Guzmania and Vriesea, one of each being Aechmea and Racinaea. We conclude that the area of Pico Alto is an important remnant of humid forest and conservation measures to protect its epiphytes are urgently needed.

Key words: Baturité range, brejo de altitude, epiphytes, Poales, Tillandsioideae, Vrieseeae

Introduction

The Atlantic rainforest may be popularly known in Northeastern Brazil as brejos de altitude. Such a name is used when the humid forest is isolated within the semi-arid climate zone and is surrounded by an array of dry woodland Caatinga (Andrade-Lima 1982). According to Lins (1989), the brejos are “areas of exception” within the semi-arid climate and the establishment of this vegetation type occurred due to high annual rainfall, favored by high (> 1,000 m a.s.l.) elevations.

These forest patches are still understudied regarding their biodiversity, despite the critical conservation status of most of these fragments which have been exploited for timber or for pastures for cattle raising activities. An entire book (Siqueira-Filho & Leme 2006) has been devoted to the Bromeliaceae from the Atlantic Forest in Northeastern Brazil, focusing on areas within Pernambuco and Alagoas states. However, almost no information is available in the literature regarding the bromeliads from Ceará and Rio Grande do Norte states, considered by different authors as the northern limit of the Atlantic forest. Although still controversial, the inland brejos from Ceará could be considered floristically similar and therefore included in the official classifications of the Atlantic forest domain that typically follows the eastern Brazilian coast (Oliveira & Araújo 2007). Several works highlight the great biodiversity harbored by these small fragments (e.g., Oliveira & Araújo 2007). As a whole, the Brazilian Atlantic forest has been considered one of the most important diversification centers for bromeliads, hosting recently evolved lineages and several endemic genera (Smith 1955; Givnish et al. 2011), thus being a priority area in need to be carefully inventoried.

42 Accepted by Eric Gouda: 28 June 2013; published: 17 July 2013

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After conducting a floristic survey of the Bromeliaceae from the region of Pico Alto, Guaramiranga, which represents the most elevated area within the Baturité mountain range (1,100 m a.s.l.; IPECE 2010), we concluded that one taxon of Vriesea is a new species that is described in the present work. Additionally we provide pictures of all the bromeliads collected within this region, hoping to contribute to their conservation within this area. We have observed over the past few years a noticeable decline in the quality of the habitats in this area along with a concomitant reduction in the number of epiphytes caused by collection pressures.

Material and methods

The Guaramiranga municipality and its associated mountains, belonging to the Baturité mountain range, is located approximately 110 km southwester of Fortaleza, the capital of Ceará state. It occurs within a region classified as BSh having an arid, hot and dry climate (Köppen 1948). However the mountainous topography favors a humid tropical climate, classified as Af (Köppen 1948). Guaramiranga has two distinct seasons, a rainy season from December to May and a pronounced dry season between September and November. The average rainfall is about 1,750 mm. The average temperature is mild, around 25°C (IPECE 2010).

The Guaramiranga mountain range has fragments of humid forests that have survived human interference and are surrounded by open and drier areas along the lower valleys (Fig. 1 A). This forest is severely fragmented and the areas where the newly discovered species occur are not covered by the usual high forest, instead they contain sparse relatively short trees rich in epiphytes along with shrubs and rocky outcrops affected by strong winds.

Plant material has been collected and studied in the field since 2010 by the authors along trails from 800 m elev. up to the highest peak, Pico Alto, 1,100 m elev. (4°16‘37,01”S, 38°58’10,98”W) as part of an ongoing project to document the bromeliad flora from Ceará state (Versieux, in prep.). Ecological observations were carried out within a larger project designed to observe the floral biology of all the bromeliads occurring in the Pico Alto region (Fortunato 2013). Specimens were deposited in the herbaria of the Universidade Federal do Ceará (EAC) and Universidade Federal do Rio Grande do Norte (UFRN) and their previous herbarium collections were also investigated. Freshly collected flowers were preserved in ethanol 70% and studied and illustrated using the stereomicroscope.

Results and Discussion

During the sampling of bromeliads in the area we observed and collected the following six sympatric bromeliad species in the region of Pico Alto including the new Vriesea currently being described: Aechmea aquilega(Salisbury 1806: 40) Grisebach (1864a: 592; Fig. 1 B–E), Guzmania sanguinea (André 1879: 367) André ex Mez (1896: 901; Fig. 1 F), G. monostachia (Linnaeus 1753: 287) Rusby ex Mez (1896: 905; Fig. 1 G), Vriesea baturitensis sp. nov. (Figs. 2 A–D, 3, 4), V. aff. wawranea Antoine (1884: t. 1, 2; Fig. 1 H), and Racinaea spiculosa (Grisebach 1864b: 17) Spencer & Smith (1993: 157; Fig. 1 I–J). Among these species the epiphytism is the prevalent life form, although Aechmea aquilega could also be seen growing on bare rocks (Fig. 1 B).

Complexes of species are commonly seen in Vriesea (e.g., Versieux 2011), and misinterpretations of floral characters is a common problem when only dried material is examined. After studying the specimens identified as Vriesea rodigasiana (Morren 1882: 171) from the Pico Alto region deposited in herbarium EAC and comparing them with the living material obtained during our field work at the same area and with the typical V. rodigasiana material frequently collected in southern and southeastern Brazil (Fig. 2 E), we concluded that the plants from Ceará state do not match the circumscription of V. rodigasiana. It is likely that

Phytotaxa 117 (2) © 2013 Magnolia Press • 43NEW VRIESEA FROM CEARÁ

FIGURE 1. Bromeliaceae of Pico Alto, Guaramiranga, Ceará. A. Overview of the green forest at the top of the Baturité range, surrounded by drier valleys. B–E. Aechmea aquilega; B. Rupicolous individuals; C. Epiphytic individuals; D. Detail of one inflorescence exhibiting spaced branches; E. Close up of an inflorescence branch and flower. F. Guzmania sanguinea with floral buds. G. Guzmania monostachia. H. Vriesea aff. wawranea. I-J. Racinaea spiculosa; I. Habit; J. Close up of the flower. (Photos: A–E, H:

L.M. Versieux; F–J: M. Fortunato).

VERSIEUX ET AL.44 • Phytotaxa 117 (2) © 2013 Magnolia Press

all the specimens cited by Smith & Downs (1977) as V. rodigasiana for Ceará state correspond to the new species described here. A combination of characteristics that distinguish V. rodigasiana from V. baturitensisenabled us to describe it here as a new taxon: the rosette shape (not so regular vs. more rosulate, well organized and nearly radially symmetric in V. baturitensis); inflorescence shape (mostly pyramidal, branches slightly contorted, with the distal branches becoming shorter vs. inflorescence ellipsoidal or ovoidal, branches straight, divergent), primary bract color (red vs. yellow or reddish yellow), peduncle shape and position (usually contorted and suberect vs. straight and stiff, erect), and shape of the rachis of the branches (geniculate vs. flexuous to nearly straight). Finally, V. baturitensis has longer and wider floral bracts and sepals, among other distinct dimensions presented in Table 1. The new species can also be compared with and distinguished from V. friburgensis (Mez 1894: 537) but has shorter flowers and yellow primary bracts (Table 1), as well as bright yellowish green and thicker leaves (vs. dark green to reddish and thinner). Both species can also be separated by the length of the upper peduncle bracts, which are much exceeding the internodes in V. friburgensis and are usually shorter in the new species, sometimes exceeding only in few millimeters the internode.

TABLE 1. Morphological comparison of Vriesea baturitensis with related taxa. All measures are in cm and were obtained from the examined material for each species (Appendix 1).

According to Rocca and Sazima (2013) V. rodigasiana is pollinated mainly by the hummingbirds Thalurania glaucopis and Ramphodon naevius, the later considered to be the most efficient pollinator based on percentage of pollen germination. Vriesea baturitensis is visited by Chlorostilbon notatus, Glaucis hirsutusand mainly by Phaethornis pretrei (Fortunato 2013).

The second species of Vriesea found in Pico Alto (Fig. 1 H) is treated here as another unpublished species. It has earlier been identified either as V. bituminosa (Wawra 1862: 347; Smith & Downs 1977;) or as V. oleosa(Leme 1999: 160; Siqueira Filho & Leme 2006: 373), but according to a recent revision of this complex, the plants from Ceará state would be considered a new species related to V. wawranea (Moura 2011: 175). Both species of Guzmania recorded here are only known in Brazil by their occurrence in Ceará state, highlighting, together with the two new species of Vriesea the importance of this area for Bromeliaceae conservation.

Taxonomic treatment

Vriesea baturitensis Versieux & Tomaz, spec. nov. (Figs. 2–4).Vriesea baturitensis is related to V. rodigasiana but differs by the yellow primary bracts and by the wider and longer

floral bracts and sepals, by the rounded leaf apex (in vivo), by the slightly spreading petals and spreading anthers (vs. erect corolla and strict anthers) and by the shape of the inflorescence. It is also similar to V. friburgensis, but can be separated by the shorter flowers and yellow primary bracts.

Type:—BRAZIL. Ceará state: Guaramiranga, Estrada de acesso às torres de TV, 42.261° S, 38.933° W, 8 May 2012, L. M. Versieux 525, M. E. M. Fortunato & I. Costa (holotype [2 sheets] UFRN!, isotype EAC!).

Epiphyte, heliophyte or semi-sciadophyte. Leaves 15–19 cm long, yellowish green, nearly symmetrically arranged in a dense funnel-form or crateriform rosette. Leaf sheath (4–)5.6–7.1 × 4–5.8 cm, elliptic, densely brown lepidote in the center to pale brown along the margins on both surfaces. Leaf blade 10–17 × 2.6–3 cm, ligulate with apex rounded in vivo, apiculate, lustrous, coriaceous, subdensely lepidote abaxially and sparsely lepidote adaxially. Peduncle 14–31 × ca. 0.4 cm, slender, stiff, sparsely lepidote, internodes 3–4 cm long. Peduncle bracts 3.8–4.5 × 1.5–1.8 cm,

V. baturitensis V. friburgensis V. rodigasiana

Primary bract color yellow red red

Floral bracts 2.1–2.3 × 1.6–2.1 2.8-3.2 1.0–2.0 × 1–1.4

Flower length 3.9–4 3.5-4 2.8–3.2

Sepals 2.6–3.2 × 0.8–1.1 2.7 × 0.5 1.9–2.2 × 0.5–0.7


FIGURE 2. Vriesea baturitensis. A. Habit of the plant used to prepare the holotype; B. Detail of young inflorescence with divergent branches; C. Flower (frontal view); D. Flower (side view); E. Vriesea rodigasiana, blooming individuals photographed in Santa Catarina State, southern Brazil (Photos: A: L.M. Versieux; B–D: M. Fortunato; E: L.A. Funez).


FIGURE 3. Vriesea baturitensis. A. Leaf; B. Peduncle bract (middle one); C. Primary bract; D. Floral bract; E. Sepal (abaxial view); F. Sepal (adaxial view); G. Stigma; H. Petal with antisepalous and antipetalous stamen; I. Details of the petal appendages. (From the holotype).

nearly ovate, apex obtuse to apiculate, sparsely brown lepidote toward the apex, erect with the apex recurved, enfolding the internodes, the proximal ones slightly longer than the internodes and the distal ones shorter. Inflorescence (excluding the peduncle) 20–30 cm long, compound, once-branched of 3–9 branches, erect, internodes of the axis 1.7–4 cm long; branches 9–14 cm long, 5–7-flowered, divergent. Primary bracts 2–4 × 1.9–2.1 cm, ovate, apex acute to shortly acuminate, subdensely lepidote adaxially and sparsely lepidote to nearly glabrous at the base abaxially, yellow, exceeding the stipes of the branches. Stipes 0.9–1.6 × 0.2–0.3 cm. Floral bracts 2.1–2.3 × 1.6–2.1 cm, ovate, apex obtuse, carinate, sparsely lepidote adaxially, the abaxial surface with a few scales concentrated centrally on the keel mainly near the apex. Flowers 3.9–4 cm long, distichous, sessile. Sepals 2.6–3.2 × 0.8–1.1 cm, symmetric, free, yellow, subdensely lepidote adaxially, glabrous abaxially, ecarinate, but with a thicker central area. Corolla nearly tubular with the petal apex recurved. Petals 3–3.5 × 0.4–0.5 cm, ligulate, apex retuse, yellow. Petal appendages 0.8–0.9 × 0.1


cm, narrowly triangular, asymmetric, the distal part free for 3–4.5 mm. Stamens exserted, distinctly spreading. Filaments 3–4 cm long, yellowish. Anthers 4–5 × ca. 0.5 mm, linear, dorsifixed at the base, blackish, the pollen grains yellow. Style 3.4–3.6 cm long. Stigma 0.2 cm diam, with convolute blades, yellow. Ovary ca. 4 mm long, nearly conical. Capsule 3–3.5 cm long, ovoidal, pale brown.

Distribution and conservation:—So far restricted to the Baturité mountain range, Ceará. The species appears to grow only at the higher elevations (> 800 m.a.s.l.) where mist formation is frequent. More field studies are necessary in order to clearly establish the conservation status of the species, but we observed a decline in the habitat quality and that adult individuals are being collected from the wild by collectors.

Ecology:—Vriesea baturitensis is a heliophilous or semi-sciadophilous epiphyte. Anthesis is diurnal, with the flowers opening at 06:00 AM and withering at 06:00 PM the same day. The flowering is continuous, with a peak in the dry season (December to May). Fruiting is also continuous and the fruit ripening period takes about four months. The morphology of the flowers indicates ornithophily and they are visited by three different species of hummingbirds. A continuous supply of floral resources by this species favors the maintenance of the pollinator guild.

Etymology:—The species is named after the Baturité mountain range.Paratypes:—BRAZIL: Ceará, Mun. Pacoti, Serra do Baturité, 17 June 1989, Figueiredo s.n. (EAC

20333!); Serra do Baturité, Morro Alto, 22 November 1974, Fernandes s.n. (EAC 2520!); Mun. Guramira: Baturité, Pico Alto, 28 December 1997, Otoch s.n. (EAC 26156!); Serra de Baturité, Pico Alto, 8 February 1990, Fernandes & Fontella s.n. (EAC 16286!); Estrada do Pico Alto, 24 April 2000, Lima-Verde s.n. (EAC 31513!).

FIGURE 4. Holotype of Vriesea baturitensis deposited in the UFRN herbarium.


Acknowledgments

We are very grateful to Walter Till, Andrea Costa and to associate editor Eric Gouda for their much valuable comments on the manuscript and constructive reviews. We thank R. Smythe and A. Calvente for proofreading and biologist Luís Adriano Funez who provided the picture of Vriesea rodigasiana from Santa Catarina used to illustrate this taxon in Figure 1. Line drawings were prepared by Rudson Cruz. I. Costa helped during the field work. We thank curators of EAC, HB and UFRN for multiple assistances. The second author had a REUNI/PIBIC/UFRN fellowship (project 6738-2011) and the third author had a CAPES M.Sc. fellowship.

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http://dx.doi.org/10.1007/BF01962979


http://www.ipece.ce.gov.br/publicacoes/perfil_basico/pbm-2010/Guaramiranga.pdf/view

APPENDIX 1: Examined material for obtaining the values for Table 1.

Vriesea friburgensis: BRAZIL. Minas Gerais: Lima Duarte, Leme 1473. São Paulo: Botucatu, Amaral Jr. 1387. Paraná: Serra da Graciosa, Brito 946, 949; Tibagi, Silva et al. 1815 (all at HB). Santa Catarina: Campo Alegre, Silva s.n. (HB 71747).

Vriesea rodigasiana: BRAZIL. Bahia: Una, Leme 3032. Espírito Santo: Santa Teresa, Seidel 745. Rio de Janeiro: Baixada Fluminense, Pereira 10598. Paraná: Serra da Graciosa, Brito 941; Praia do Leste, Leinig 503 (all at HB).


phytotaxa 117 (2): 42 phytotaxa article phytotaxa

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