Revision of Orthophytum (Bromeliaceae): the Species with Sessile Inflorescences

Revision of Orthophytum (Bromeliaceae): the species with sessile inflorescences

RAFAEL BATISTA LOUZADA1,2 & MARIA DAS GRAÇAS LAPA WANDERLEY2 1Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, Cx. Postal 11461, 05422-970, São Paulo, SP, Brazil; email: [email protected] 2Instituto de Botânica, Secretaria do Meio Ambiente, Cx. Postal 3005, 01061-970, São Paulo, SP, Brazil; email: [email protected]

Abstract

Orthophytum is a bromeliad genus restricted to Brazil with records for the states of Paraíba, Pernambuco, Alagoas, Sergipe, Bahia, Espírito Santo and Minas Gerais. The genus is usually divided in two informal groups based on the presence or absence of a peduncle. This paper presents a taxonomic revision of the 12 species in the group with sessile inflorescences and descriptions of the genus and species, nomenclatural and ecological notes, as well as illustrations are provided. The new species Orthophytum ulei is described. The conservation status for each species is evaluated using the World Conservation Union (IUCN) criteria.

Key words: Brazil, Bromelioideae, Chapada Diamantina, endangered species, endemism, Espinhaço Range, inselbergs

Introduction

The Bromeliaceae is the largest Neotropical plant family of monocots. Currently, 58 genera and 3172 species are recognized (Luther 2008), the majority of these occurring in Brazil. Within Brazil, the family ranks second among the monocots with the highest number of endemic genera (16), only after Orchidaceae with 35 genera, (Smith & Downs 1974, 1977, 1979, Forzza 2005, Giulietti et al. 2005). The generic boundaries in Bromeliaceae are still not well-defined, particularly within the subfamily Bromelioideae, to which the genus Orthophytum Beer (1854: 347) has been assigned (Smith & Downs 1979, Wendt 1997, Canela et al. 2003, Faria et al. 2004, Schulte et al. 2005, Horres et al. 2007, Schulte et al. 2009). The genus Orthophytum comprises about 55 species that are all endemic to Brazil, distributed from the northeastern to the southeastern regions of the country. Two centres of diversity can be recognized, one in the Espinhaço range in the states of Bahia and Minas Gerais, and the other in the inselberg region in the states of Espírito Santo and Minas Gerais, which is part of the Atlantic rain forest area (Wanderley 1990, Leme 2004a, Louzada & Wanderley 2008, Martinelli et al. 2008). Orthophytum species grow on rocky outcrops in ‘campos rupestres’ (rocky fields), the 'caatinga', and on the inselbergs in the Atlantic rain forest area (Leme 2004a, Louzada & Wanderley 2008). When Beer (1854) described the genus, he did not assign any previously described species to it, nor published any new species. But he did cite one unnamed collection: Pohl 3436, today known as Orthophytum glabrum (Mez) Mez (1896: 116). Almost four decades later, Mez (1892) described the genus Prantleia, including P. glabra Mez in Martius (1892: 258, based on Pohl 3436 and P. leprosa Mez in Martius (1892: 259; based on Pohl 5229 and Glaziou 14035). When Mez (1896) realized that Pohl 3436 had already been applied to Orthophytum by Beer (1854), he synonymized his Pratleia under that genus (Mez 1896). Botanic expeditions to Bahia by Ernst Heinrich Georg Ule at the end of 19th the century, led to the description of two new genera: Sincoraea Ule (1908: 191) and Cryptanthopsis Ule (1908: 193), which were

Accepted by E. Gouda: 30 Sep. 2010; published: 6 Dec. 2010 1 subsequently proposed as synonyms of Orthophytum by Smith (1955), without providing any justification. In the same work Smith (1955) presented a key for 10 species of Orthophytum. Morphologically, Orthophytum is divided into two groups, one with pedunculate inflorescences (pedunculate inflorescence group), and the other without a peduncle (sessile inflorescence group). In the sessile inflorescence group, we recognize two separate complexes corresponding to two centres of diversity. The first species complex is composed of three species distributed in the inselbergs of Espirito Santo and is characterized by a long-caulescent habit. The second species complex displays an inconspicuous stem growing in sandstone outcrops in the Espinhaço range with one single species occurring in Minas Gerais and nine species in Bahia. Orthophytum stands out among other bromeliad genera by its high proportion of microendemics, with most species restricted to one or two localities. In the state of Minas Gerais, 68% of Orthophytum species are endemic, 47% are considered vulnerable, but none of these species can be found within the borders of any reserve (Versieux & Wendt 2006, 2007). The present work is a taxonomic revision of the sessile inflorescence group of Orthophytum. This is a first step to better understand the generic boundaries, propose conservation measures, and to provide new data for the revision of this Brazilian genus.

Material and Methods

Morphological analyses and geographical distributions were based on herbarium specimens from the following herbaria: ALCB, CEPEC, HB, MBM, MBML, RB, SP, SPF, HUEFS. To identify the material we studied original descriptions, photos and type-material. Five field trips to type localities and other areas in the states of Bahia, Minas Gerais and Espírito Santo were carried out to collect additional specimen. Plants collected in the field were incorporated into the living collection of the Instituto de Botânica in São Paulo and voucher specimen were deposited at the SP herbarium. Conservation assessments for species were based on the herbarium material and field observations, applying the IUCN red list category criteria (IUCN 2001).

Taxonomic Treatment

Orthophytum Beer (1854: 347)

Prantleia Mez in Martius (1881: 257). Type:—Prantleia glabra Mez in Martius (1892: 258) [currently accepted name = Orthophytum glabrum (Mez) Mez (1896: 117)] Sincoraea Ule (1908: 191). Type:—S. amoena Ule (1908: 191) [currently accepted name = Orthophytum amoenum (Ule) L.B.Sm. (1955: 33)] Cryptanthopsis Ule (1908: 193). Type:— C. saxicola Ule (1908: 193) [currently accepted name = Orthophytum saxicola (Ule) L.B.Sm. (1955: 34)]

Plants rupicolous or terrestrial, stoloniferous; stems inconspicous, covered by the leaf sheaths. Leaves rosulate, patent or arching sometimes falcate. Leaf sheaths imbricate, triangular to ovate, green, whitish-green or greenish-yellow, lepidote, glabrous or sparsely lepidote, margins spinose. Leaf blades coriaceous to subcoriaceous, linear-triangular to narrowly-triangular, plane or concave, lepidote on both surfaces or sparsely lepidote, margins serrate or spinose, spines congested to sparse. Peduncles present or absent, terminal, erect, glabrous or lanate, with foliaceous bracts longer than the internodes. Inflorescences sessile or pedunculated, compound or simple. Primary bracts foliaceous or subfoliaceous. Vestigial floral bract present or absent. Floral bracts green or red, margins spinose to serrate, pungent. Flowers sessile or short-pedicellate. Sepals free, erect or suberect, asymmetrical or rarely symmetrical, acute, acuminate, mucronate or mucronulate. Petals free, white to greenwish-white, sublinear-spatulate, margins entire, with two sacciform or cuppuliform appendages lacerate or digitate and two callosities lateral along the filaments. Epigenous tube present or

2 • Phytotaxa 13 © 2010 Magnolia Press LOUZADA & WANDERLEY absent. Stamens not surpassing the petals, filaments filiform, first whorl free and opposite the sepals, second whorl adnate to the petals; anthers linear. Styles tricostate. Stigmas simple-erect. Fruits with sepals persistent. Seeds ovate and striped.

Key to the species of Orthophytum with sessile inflorescence

1 Plants with short stem, inconspicuous, 1–4.2 cm long; petals white, obtuse to subacute not cucullate...... 2 - Plants with long stem, ca 10 cm long; petals green with white margins, obtuse-cucullate...... 11 2 Compound inflorescences (rarely simple in O. ulei) ...... 3 - Simple inflorescences ...... 4 3 Floral bracts and sepals pink to red ...... 7 - Floral bracts and sepals green...... 10 4 Sepals without glandular trichomes; prickles of leaf blade (1.7–)2–3.5 mm long ...... O. hatschbachii - Sepals with glandular trichomes; prickles of leaf blade 0.3–1.5 mm long...... 5 5 Leaf blades densely lepidote; sepals narrowly triangular...... O. humile - Leaf blades sparsely lepidote; sepals triangular-lanceolate...... 6 6 Leaf blades 8–23 × 0.4–0.7 cm; petals c. 2.5 cm long ...... O. navioides - Leaf blades 2.5–6.7 × 0.3–0.4 cm; petals c. 1.9 cm long ...... O. mucugense 7 Leaf blades adaxially lepidote, not lucid ...... 8 - Leaf blades adaxially sparsely lepidote, lucid ...... 9 8 Leaf blades adaxially glabrous at base; petals 1.8–2.5 cm long, apices subacute...... O. burle-marxii - Leaf blades adaxially densely lepidote at base; petals 1.3–1.7 cm long, apices obtuse ...... O. ulei 9 Leaf blades coriaceous, 0.5-1 cm wide; prickles 1.5–3 mm long; floral bracts c. 1.8 × 1 cm; anthers 3.2–3.5 mm long ...... O. amoenum - Leaf blades subcoriaceous, 0.2–0.4 cm wide; prickles 0.3-0.4 mm long; floral bracts 0.9–1.2 × 0.6–0.7 cm; anthers 2.3–2.5 mm long ...... O. ophiuroides 10 Leaf blades margins serrate, adaxial surfaces with white-lanate ring at base; prickles 0.5–3.5 cm long; sepals and petals without glandular trichomes ...... O. albopictum - Leaf blades margins serrulate, adaxial surfaces without white-lanate ring at base; prickles 0.3–0.5 mm long; sepals and petals with glandular trichomes ...... O. heleniceae 11 Simple inflorescences ...... O. vagans - Compound inflorescences...... 12 12 Leaf blades densely lepidote on both surfaces; prickles 0.6-1 mm long ...... O. zanonii - Leaf blades adaxially glabrous toward the; prickles 2-3.5 mm long...... O. pseudovagans

Orthophytum albopictum Philcox (1985: 357). Figs. 1A–H, 7A.

Type:—BRAZIL. Bahia: near Cumbuca, about 3 km N of Mucugê on road N to Andaraí, Storr 122 (holotype CEPEC!, isotype K!).

Plants rupicolous or terrestrial, stoloniferous, stems short, 1.1–3 × 2.5–3.5 cm. Leaves patent or slightly arching. Leaf sheaths greenish yellow, broadly triangular, 0.6–3 × 1.1–3.2 cm, glabrous on both surfaces or lepidote at the abaxial apices, margins serrate, prickles 0.2–1 mm long. Leaf blades strongly coriaceous, plane, red, purple at, linear-triangular, 8–24 × 0.8–2 cm, attenuate, abaxial surfaces densely lepidote, adaxial surfaces glabrous and lanate at the base, margins serrate, prickles antrorse, 0.4–1.2(–2) mm long, apices mucronulate. Inflorescences compound, many-flowered. Primary bracts coriaceous, red, deltoid or lanceolate, 1.7–3.8 × 1.6–1.7 cm, lepidote at the apices or similar to leaf blades, margins serrate, mucronulate. Sterile bract present. Floral bracts subcoriaceous, green, asymmetrical, ecarinate, triangular, c. 1.5 × 1 cm, lepidote, margins serrate, apices mucronulate. Sepals green, asymmetrical, carinate, narrowly triangular, 1.2– 1.4 × 0.4–0.5 cm, lepidote, margins entire, apices mucronulate. Petals white, linear-spatulate, c. 1.8 × 0.4 cm, with simple trichomes at base, apices obtuse. Petal appendages sacciform, lacerate c. 6 mm from the base. Callosities conspicous. Stamens with antepetalous filaments ca. 3 mm long and adnate to the petals, the

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 3 antesepalous ones free, ca 9 mm long. Anthers yellow, ca 4 mm long, apices not apiculate. Ovaries trigonous, styles ca 1.3 mm long. Epigenous tube present, ca 2 mm long. Ovules numerous. Fruits and seeds not seen.

FIGURE 1. A–H. Orthophytum albopictum Philcox (Wanderley et al. 2538, SP). — A. Habit. — B. Primary bract. — C. Vestigial floral bract. — D. Flower. — E. Sepal and free stamen. — F. Petal with stamen adnate, petals appedages sacciform and petals callosities. — G. Detail of the trichomes below the petals appendages. — H. Style and stigma.

4 • Phytotaxa 13 © 2010 Magnolia Press LOUZADA & WANDERLEY Distribution and habitat:—Orthophytum albopictum is known from Andaraí and Mucugê counties in Bahia. It grows rupicolous or terrestrial, frequently exposed to the sun on rocky outcrops in grassland in campos rupestres between 700–1500 m. It was collected in the Parque Nacional da Chapada Diamantina and Parque Municipal de Mucugê. IUCN conservation status:—EN B1a. Based on collections and field observations throughout the range of this species, we noticed that it is restricted to an estimated area of less than 5000 km2. It is only known from two counties in Bahia, occurring within and outside of protected areas. Comments:—Orthophytum albopictum is similar to O. ulei, the newly described species in this paper(see below), but the floral bracts and sepals of O. albopictum are green (vs. red or pink), the leaf blades are lepidote only at the base (vs. completely lepidote) and with short prickles (0.4–1.2(–2) vs. 2–3.5 mm). This species is easily recognized by a lanate band in the centre of the rosette at anthesis, a result of a high density of leaf scales. While this band around the inflorescences is a good character to distinguish it from other species in the field, in dried material this character is sometimes obscure. Additional specimens examined:—BRAZIL. Bahia: Andaraí, 4 May 2002 (fl), Loureiro 625 (ALCB). Mucugê, 21 October 1988 (fl), Eggli 1249 (CEPEC); Estrada para Cascavel, 27 March 1980 (fl), Martinelli et al. 6689 (RB); Estrada Mucugê-Andaraí, 28 April 2001 (fl), Assis et al. s.n. (ALCB); Guiné, 29 June 2002 (fl), Conceição 1094 (SP); 29 June 2002 (fl), Conceição 1097 (SP); Parque Municipal de Mucugê, 7 January 2003 (fl), Wanderley & Conceição 2363 (SP); 7 January 2003 (fl), Wanderley & Conceição 2364 (SP); 17 January 2006 (fl), Wanderley & Conceição 2366 (SP); 17 January 2006 (fl), Wanderley et al. 2538 (SP); Rio Cumbuca, 4 February 1974 (fl), Harley 15982 (CEPEC, RB); Rio Piaba, 9 November 1988 (fl), Kral & Wanderley 75623 (SP); 7 January 2003 (fl), Wanderley & Conceição 2362 (SP); Serra do Capa Bode, 28 October 1978 (fl), Martinelli 5525 (RB); 30 October 1978 (fl), Martinelli 5519 (RB); 6 October 1980 (fl), Furlan et al. CFCR 433 (SPF).

Orthophytum amoenum (Ule) Smith (1955: 33), Figs. 2A–J, 7B.

Basionym: Sincoraea amoena Ule (1908: 191). Type:—BRAZIL. Bahia: Serra do Sincorá, 1400 m, E. Ule 7106, (lectotype B!).

Plants rupicolous, stoloniferous, stems short, 2–3.7 × 2–2.5 cm. Leaves arching. Leaf sheaths greenish-white, broadly ovate, 0.6–1.5 × 0.7–2.5 cm, lepidote on abaxial surfaces and sparsely lepidote on adaxial surfaces, margins serrate, prickles 0.5–1 mm long. Leaf blades coriaceous, concave, wine red or pink to red at base, linear-triangular or narrowly triangular , 4.7–17.3 × 0.5–1 cm, attenuate, densely lepidote abaxially, sparsely lepidote adaxially, margins serrate, prickles straight or antrorse, ca 1.5–3 mm long, apices mucronate. Inflorescences sessile, compound, many-flowered. Primary bracts coriaceous, red or pink, triangular to narrowly triangular, 3.3–6.5 × 1.4–1.5 cm, sparsely lepidote, margins serrate, apices mucronulate. Vestigial floral bract present. Floral bracts subcoriaceous, red or pink, asymmetrical, carinate, triangular, ca 1.8 × 1 cm, glabrous, margins serrate, apices mucronate. Sepals red or pink, asymmetrical, carinate, narrowly elliptic or lanceolate, 1.1–1.2 × 0.3 cm, glabrous, margins entire, apices acuminate. Petals white, linear-spatulate, 1.5– 1.6 × 0.4–0.5 cm, glabrous, apices obtuse. Petal appendages sacciform, lacerate, 3–4 mm from the base. Callosities inconspicous. Stamens with antepetalous filaments 2.5–3 mm long and adnate to the petals, the antesepalous ones free, ca 8 mm long. Anthers yellow, 3.2–3.5 mm long, apices apiculate. Ovaries trigonous, styles 12 mm long. Epigenous tubes present, 2 mm long. Ovules ca 20 per placenta. Fruits and seeds not seen. Distribution and habitat:—Orthophytum amoenum (Ule) L.B.Sm. is known from Lençóis and Palmeiras counties in Bahia. It grows rupicolous, frequently exposed to the sun in rocky outcrops in campos rupestres. IUCN conservation status:—CR B1ab (ii, iii). Orthophytum amoenum is well represented in herbaria, but all specimens refer only to a single locality, where it occurs on two adjacent hills: Morro do Pai Inácio and

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 5 Morro da Mãe Inácia. In recent field trips to these areas, a decline in the quality of the habitat was noted. The area is frequently visited by tourists, causing disturbance to the population of this species.

FIGURE 2. A–J. Orthophytum amoenum (Ule) L.B.Sm. (Wanderley et al. 2528, SP). — A. Habit. — B. Flower. — C. Floral bract. — D. Sepal. — E. Petal with stamen, showing petals appendages and petals callosities. — F. Detail of the petal lobe. — G. Petals appendages sacciform with lacerate margins. — H. Style and stigma. — I. Detail of the tricostate style. — J. Detail of the stigma.

6 • Phytotaxa 13 © 2010 Magnolia Press LOUZADA & WANDERLEY Comments:—Orthophytum amoenum is morphologically similar to O. navioides (L.B.Sm.) Smith (1955: 34), however the leaf blades of O. amoenum are wider and shorter (4.7–17.3 × 0.5–1 cm vs. 8–23 × 0.4–0.7 cm) and strongly coriaceous, presenting long and sparse prickles (1.5–3 vs. 1–1.5 mm long). Floral features of these species are easily distinguished, with obtuse petals apices in O. amoenum and subacute in O. navioides, and larger flowers in O. navioides. Orthophytum navioides occurs in shady places on rocks near streams and waterfalls forming dense mats, while O. amoenum grows in sunny habitats, in grassland on rocky fields, with fewer and more widely spaced individuals. Additional specimens examined:—BRAZIL. Bahia: Lençóis, BR 242, 9 November 1988 (fl), Wanderley et al. 1598 (SP); Chapadinha, 23 February 2003 (fl), Rapini & Fiaschi s.n. (SP 364464, HUEFS 70197); Palmeiras, Morro da Mãe Inácia, 14 January 2006 (fl), Wanderley et al. 2521, 2522, 2528 (SP); Morro do Pai Inácio, 29 February 1980 (fl), Mori 13293 (CEPEC); 16 November 1983 (fl), Martinelli et al. 9793 (RB); 19 February 1983 (fl), Noblick & Pinto 2822 (CEPEC, RB, SP); 21 July 1981 (fl), Pirani et al. CFCR 1620 (RB); June 1986 (fl), Santos s.n. (HUEFS 10067); 8 November 1997 (fl), Guedes et al. 5514 (ALCB); 26 October 2004 (fl), Carvalho et al. PCD 1023 (ALCB, SP); 15 April 2005 (fl), Cardoso et al. 428 (HUEFS).

Orthophytum burle–marxii Smith & Read (1979: 164), figs. 3A–G, 7C.

Type:—Brazil. Bahia: J. Bogner 1311 (holotype US!). Synonyms: Orthophytum burle-marxii L.B.Sm. & Read var. seabrae Rauh (1985: 79). Type: Brazil. Bahia: Seabra, 6 Oct 1981, Rauh 56 497a (holotype HEID!) syn. nov.

Plants rupicolous, stoloniferous, stems short, ca 2.5 × 4 cm. Leaves patent or slightly arching. Leaf sheaths white-greenish, broadly triangular or ovate-triangular, 1–3 × 1–4 cm, glabrous, rarely sparsely lepidote, margins serrate, prickles 0.5–1 mm long. Leaf blades strongly coriaceous, concave, red or green, cinereous, linear-triangular, 12.8–53 × 1.5–2.5 cm, attenuate, abaxial surfaces densely lepidote, adaxial surfaces lepidote with the base glabrous, margins serrate, prickles antrorse 1–2 mm long, apices mucronate. Inflorescences sessile, compound, many–flowered. Primary bracts coriaceous, red, triangular, or triangular-lanceolate, 2.2– 5.5 × 1–6 cm, sparsely lepidote, margins serrate, apices mucronulate. Vestigial floral bract present. Floral bracts subcoriaceous, red, symmetrical, ecarinate, triangular, 1.5–4.8 × 0.8–3.7 cm, glabrous, margins serrate, mucronulate. Sepals red, asymmetrical, carinate, narrowly triangular, 2 × 0.4–0.6 cm, sparsely lepidote, margins entire, acuminate, or mucronulate. Petals white, linear-spatulate, 1.8–2.5 × 0.6 cm, glabrous, subacute. Petal appendages sacciform, lacerate, ca 7 mm from base. Callosities conspicous. Stamens with antepetalous filaments ca 0.5 mm long and adnate to the petals, the antesepalous ones free, ca 14 mm long. Anthers yellow, ca 4.5 mm long, not apiculate. Ovaries trigonous. Epigenous tubes present, ca 2 mm long. Ovules numerous. Fruits and seeds not seen. Distribution and habitat:—Orthophytum burle-marxii is known from Lençóis and Morro do Chapéu counties in Bahia. It grows on rocky outcrops in grassland on rocky fields frequently exposed to the sun, from 900–1.400 m elevation. IUCN conservation status:—VU B1ab (iii). The estimated area for Orthophytum burle–marxii is less than 20,000 km2. Recently, a degradation of the habitat of this species was observed, mainly due to tourism in the region. Comments:—Orthophytum burle–marxii and O. heleniceae Leme (2004b: 67) are morphologically similar, but in O. burle–marxii the leaf blade is stiff coriaceous, with sepals, and floral bracts red, forming a red and glabrous band around of inflorescence, while in O. heleniceae the band is green. Living plants of the species can be easily distinguished by these characteristics, but in herbarium material these differences are more difficult to recognize.

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 7 FIGURE 3. A–G. Orthophytum burle-marxii L.B.Sm. & Read (Louzada & Moreira 45, SP). — A. Habit. — B. Primary bract. — C. Floral bract. — D. Sepal. — E. Sepal with stamen. — F. Petal showing an adnate stamen, petals appendages and callosities. — G. Style and stigma.

Additional specimens examined:—BRAZIL. Bahia: Lençóis, Morro da Chapadinha, Chapadinha, 22 November 1994 (fl), França et al. 1257 (SP); Serra Larga, 19 December 1984 (fl), Lewis et al. CFCR 7268

8 • Phytotaxa 13 © 2010 Magnolia Press LOUZADA & WANDERLEY (RB); Rio Mucugesinho, 21 December 1981 (fl), Carvalho et al. 1065 (RB); 10 December 2003 (fl), Wanderley & Conceição 2371 (SP); 8 January 2003 (fl), Wanderley & Conceição 2379 (SP); 14 January 2006 (fl), Wanderley et al. 2530 (SP); Morro da Chapadinha, 22 November 1994 (fl), Melo et al. 1257 (CEPEC); Ribeirão do Meio, 08 January 2003 (fl), Wanderley & Conceição 2368 (SP); 17 December 2006 (fl), Louzada & Moreira 45 (SP); Mirante, 13 January 2006 (fl), Louzada & Moreira 11 (SP); Morro do Chapéu, Cachoeira do Ferro Doido, 27 October 1978 (fl), Martinelli 5335 (RB).

Orthophytum hatschbachii Leme (1995: 120), figs. 4A–G, 7D.

Type:—BRAZIL. Bahia: Rio de Contas, Mato Grosso 1300 m, 7 April 1992, G. Hatschbach & Barbosa 56827 (holotype MBM!).

Plants rupicolous, stoloniferous, stems short, 3–4.2 × 3 cm. Leaves patent. Leaf sheaths greenish–white, broadly triangular or broadly ovate, 0.5–3 × 0.8–4 cm, glabrous, margins serrate, prickles 0.1–0.8 mm long. Leaf blades stiff coriaceous, slightly concave, red, base greenish white, linear-triangular, 6.5–33 × 0.7–1.8 cm, attenuate, abaxial surfaces sparsely lepidote, adaxial ones glabrous or lepidote, margins strongly serrate, prickles straight or antrorse, (1.7–)2–3.5 mm long, apices mucronate. Inflorescences sessile, simple, many- flowered. Vestigial floral bract absent. Floral bracts coriaceous, green, symmetrical or asymmetrical, carinate, triangular, 2–2.5 × 0.9–2.3 cm, glabrous, margins serrate, apices acuminate. Sepals green, asymmetrical, carinate, narrowly ovate, 1.2 × 0.3 cm, glabrous, margins entire, apices mucronulate. Petals white, linear- spatulate, 1.7 × 0.4 cm, glabrous, obtuse. Petal appendages sacciform, lacerate, 4 mm from base. Callosities conspicous. Stamens with antepetalous filaments ca 2.5 mm long and adnate to the petals, , the antesepalous ones free 10 mm long. Anthers yellow, 3 mm long, apices apiculate. Ovaries trigonous, styles 10 mm long. Epigenous tubes present, 2.5 mm long. Ovules numerous. Fruits and seeds not seen. Distribution and habitat:—Orthophytum hatschbachii is known from Abaíra and Rio de Contas counties in the state of Bahia. It grows on rocky outcrops either exposed to the sun or in the shade near streams. IUCN conservation status:—EN B1ab (iii). The expansion of agricultural areas substantially reduced the available habitat of Orthophytum hatschbachii, and it is currently known from an estimated area of less than 5000 km2. Comments:—Orthophytum hatschbachii is similar to O. burle-marxii and O. heleniceae, differing from O. burle-marxii by simple inflorescences (vs. compound), petal appendages to 4 mm from the petals base (vs. 7 mm), leaf blades green at base (vs. red) and bracts and sepals green (vs. red). Orthophytum heleniceae may be confused with O. hatschbachii because of the colour of the leaves and bracts, but they differ in the margins of leaf blades, which are serrate in O. hatschbachii and serrulate on O. heleniceae. In addition, the length of prickles, the branching of inflorescences and the presence or absence of glandular trichomes are additional characteristics that separate these species. In O. hatschbachii the prickles are larger (2–3.5 vs. 0.5 mm long), the inflorescences are simple (vs. compound) and floral bracts, petals and sepals do not bear glandular trichomes. Additional specimens examined:—BRAZIL. Bahia: Abaíra, Catolés de Cima, 25 December 1992 (fl), Harley et al. 50366 (SP); Cabaceira, Riacho Fundo, 25 October 1993 (fl), Ganev 2314 (HUEFS); Rio de Contas, Estrada Real, 3 March 2002 (fl), Souza & Konno 503 (SP); Brumadinho, 21 December 2006, (fl), Louzada & Moreira 59 (SP); Fazenda Brumadinho, 5 November 1988 (fl), Wanderley & Kral 1532 (SP); Mato Grosso, 25 December 2004 (fl), Forzza & Silva 3825 (RB); Pico das Almas, 14 December 1984 (fl), Arraes et al. CFCR 6917 (RB); 30 October 1988 (fl), Harley et al. 25779 (SP, CEPEC); 20 November 1988 (fl), Harley et al. 26225 (SP, CEPEC); 22 January 1974 (fl), Harley 15413 (CEPEC).

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 9 FIGURE 4. A–G. Orthophytum hatschbachii Leme (Louzada & Moreira 59, SP). — A. Habit. B. Floral bract. — C. Flower with floral bract. — D. Section showing epigenous tube. — E. Sepal and stamen. — F. Petal showing an adnate stamen, petals appendages and petals collosities. — G. Style and Stigma.

Type:—BRAZIL. Bahia: field collection in Andaraí, Cachoeira da Garapa, 1300 m, O. Ribeiro & H. Ribeiro s.n. (holotype HB!).

Plants rupicolous, stoloniferous, stems short, ca 3 × 2 cm. Leaves patent or arching, sometimes falcate. Leaf sheaths greenish–white, broadly ovate, 1.5–1.7 × 2–3.4 cm, glabrous, margins serrulate, prickles 0.3–0.5 mm long. Leaf blades subcoriaceous, concave, red at anthesis, greenish–yellow at base, linear–triangular, 43.5–69 × 1.5 cm, attenuate, glabrous, sparsely lepidote, margins serrulate, prickles antrorse, ca 0.5 mm long, apices mucronate. Inflorescences sessile, compound, many–flowered. Primary bracts subcoriaceous, green, triangular to linear–triangular, 3.3–29 × 2–4.6 cm, glabrous, margins serrate, apices mucronate. Vestigial floral bract present. Floral bracts subcoriaceous, green, symmetrical, ecarinate, triangular, 1.6–2.8 × 1–2 cm, with glandular trichomes, margins serrate, apices mucronate. Sepals green, asymmetrical, carinate, narrowly triangular, ca 1.5 × 0.3 cm, with glandular trichomes, margins entire, acuminate. Petals white, linear-spatulate, ca 2 × 0.4 cm, with simple trichomes, obtuse. Petal appendages sacciform, lacerate, ca 6.5 mm from base. Callosities conspicous. Stamens with antepetalous filaments ca 2 mm long and adnate to the petals, the antesepalous ones free, ca 11 mm long. Anthers yellow, ca 3.5 mm long, apices not apiculate. Ovaries trigonous, styles 1.3 mm long. Epigenous tubes present, ca 2 mm long. Ovules ca 20 per placenta. Fruits and seeds not seen. Distribution and habitat:—Orthophytum heleniceae is only known from Andaraí county, where it grows on humid rocks near streams and waterfalls. IUCN conservation status:—CR B2ab (iii). Orthophytum heleniceae is only known from two collections. Based on these collections, it is estimated that the species occurs in an area of ca 10 km2, where the habitat quality is declining. Comments:—Orthophytum heleniceae is characterized by rosettes of leaves that are red at anthesis, with primary bracts and the bases of the leaves green, forming a coloured band around the inflorescence. The floral bracts and sepals are green and the leaf margins are serrated with short prickles. Our study at the type locality enabled us to substantially extend our understanding of this taxon, previously know only from one collection. Additionally, we observed that the flowers are visited by hummingbirds. Additional specimens examined:—BRASIL. Bahia: Andaraí, Cachoeira da Garapa, 18 January 2006 (fl), Wanderley et al. 2544 (SP).

Orthophytum humile Smith (1968: 75), figs. 5A–I, 7E.

Type:—BRAZIL. Minas Gerais: Grão Mogol, 18 August 1960, B. Maguire, G.M. Magalhães & C.K. Maguire 49288 (holotype US!, isotype NY!).

Plants rupicolous, stoloniferous, stems short, 0.8–2 × 1–2 cm. Leaves slightly arching. Leaf sheaths green, deltoid, 0.4–1.2 × 0.4–1 cm, glabrous, margins serrate, prickles 0.1–0.3 mm long. Leaf blades subcoriaceous, plane, red, wine red to pink, and white and green toward the base, wine red, middle pink and green at the base, linear–triangular, 2.5–21.5 × 0.2–0.5 cm, attenuate, abaxial surface densely lepidote, adaxial surface lepidote, margins serrate, prickles antrorse 0.7–1.4 mm long, apices mucronate. Inflorescences sessile, simple, many- flowered. Vestigial floral bract absent. Floral bracts subcoriaceous, green, asymmetrical, ecarinate, triangular, 1.3–2.1 × 0.4–1 cm, lepidote, sometimes with glandular trichomes too, margins serrate, mucronulate. Sepals green, asymmetrical, carinate, narrowly triangular, ca 1.5 × 0.3 cm, with glandular trichomes, margins entire, mucronulate. Petals white, linear-spatulate, ca 1.8 × 0.6 cm, with simple trichomes, obtuse. Petal appendages sacciform, deeply fimbriate, ca 4 mm from base. Callosities conspicuous. Stamens with the antepetalous filaments ca 0.4 mm long and adnate to the petals, the antesepalous ones free, ca 11 mm long. Anthers yellow,

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 11 ca 3.5 mm long, apices not apiculate. Ovaries trigonous, styles ca 11 mm long. Epigenous tubes present, ca 1 mm long. Ovules numerous. Fruits and seeds not seen.

FIGURE 5. A–I. Orthophytum heleniceae Leme. (Wanderley et al. 2544, SP). — A. Habit. — B. Primary bract. — C. Floral bract. — D. Flower. — E. Section showing Ovaries with axial placentation and epigenous tube. — F. Sepal and free stamen. — G. Petal showing an adnate stamen, petals appendages and petals callosities. — H. Detail of the glandular trichomes in petal. — I. Style and stigma.

FIGURE 6. A–I. Orthophytum humile L.B.Sm. (Louzada & Moreira 41, SP). — A. Habit. B. Floral Bract. — C. Flower. — D. Section showing Ovaries with axial placentation and epigenous tube. — E. Sepal and free stamen. — F. Sepal abaxial surface. — G. Petal showing an adnate stamen, petals appendages and petals callosities. — H. Detail of the sepal showing the glandular trichomes. — I. Detail of the style and stigma.

Distribution and habitat:—Orthophytum humile is known from Cristália and Grão Mogol counties in the state of Minas Gerais. It grows on humid or dry rocks near streams at 850–1000 m elevation, often associated with termite mounts. IUCN conservation status:—EN B1ab (iii, iv). Orthophytum humile occurs in an estimated area less than 5000 km2. A decline in the number of populations was observed, as well as a decrease in the quality of the habitat.

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 13 FIGURE 7. — A. Orthophytum albopictum. — B. Orthophytum amoenum. — C. Orthophytum burle–marxii. — D. Orthophytum hatschbachii. — E. Orthophytum heleniceae. — F. Orthophytum humile. (Photos: Abel A. Conceição [A– B] and Suzana E. Martins [E]).

14 • Phytotaxa 13 © 2010 Magnolia Press LOUZADA & WANDERLEY Comments:—Orthophytum humile is similar to Orthophytum mucugense Wanderley & Conceição (2006: 4), differing mainly by the leaf blades that are densely lepidote (vs. sparsely lepidote), and prickles that are 0.7–1.4 cm long (vs. 0.3–0.4 mm long). Orthophytum humile is also similar to O. ophiuroides Louzada & Wanderley (2008: 406), but can be separated mainly by its simple inflorescence (vs. branched inflorescence) and green coloured sepals and bracts (vs. red). Additional specimens examined:—BRAZIL. Minas Gerais: Cristália, Serra do Batieiro, 14 September 1991 (fl), Carvalho 575 (SP); Grão Mogol, 25 February 1986 (fl), Semir et al. CFCR 9659 (RB, SPF); Bacia do Ribeirão da Morte, 4 November 1987 (fl), Cordeiro et al. CFCR 11484 (SPF); Córrego das Mortes, 13 October 2006 (fl), Louzada & Moreira 41 (SP).

Orthophytum mucugense Wanderley & Conceição (2006: 4), figs. 8A–N, 13A.

Type:—BRAZIL. Bahia: Mucugê, Parque Municipal Sempre-viva, cachoeira do Rio Tiburtino, 12°59’83”S, 42°20’91”W, 17 January 2006, M.G.L. Wanderley, A. Conceição, R. Louzada, S. Martins 2540 (holotype SP!, isotype HUEFS!).

Plants rupicolous, stoloniferous, stems short, ca 1 × 1.3 cm. Leaves patent or arching. Leaf sheaths reddish- white, ovate, 0.5–1 × 0.4–1 cm, glabrous, margins serrate, prickles ca 0.4 mm long. Leaf blades subcoriaceous, plane or slightly concave, green, partially to completely red at anthesis, linear-triangular, 2.5– 6.7 × 0.3–0.4 cm, attenuate, sparsely lepidote on both surfaces and with glandular trichomes on the base of the abaxial surfaces, margins serrate, prickles antrorse, ca 0.3–0.4 mm long, apices mucronulate. Inflorescences sessile, simple, many-flowered. Vestigial floral bract absent. Floral bracts subcoriaceous, green, symmetrical, carinate, ovate-lanceolate, 1–1.2 × 0.4–0.7 cm, with glandular trichomes, margins serrate, apices mucronulate. Sepals green, asymmetrical, carinate, narrowly triangular, ca 0.8 × 0.3 cm, with glandular trichomes, margins entire, apices acuminate. Petals white, linear–spatulate, ca 1.9 × 0.4 cm, glabrous, obtuse. Petal appendages sacciform, lacerate, ca 5 mm from base. Callosities inconspicuous. Stamens with antepetalous filaments ca 3 mm long and adnate to the petals, the antesepalous ones free, ca 7 mm long. Anthers yellow, ca 1.8 mm long, apices apiculate. Ovaries trigonous, styles ca 8 mm long. Epigenous tubes present, ca 0.5 mm long. Ovules numerous. Fruits subtrigonous. Seeds obtuse. Distribution and habitat:—Orthophytum mucugense is only known from Mucugê county in Bahia. It grows on moist and shaded rocks near streams and waterfalls. IUCN conservation status:—VU B2ab (ii, iii). Despite occurring within a protected area, Orthophytum mucugense is threatened due to the small population size and its limited distribution. Comments:—Orthophytum mucugense is morphologically similar to O. humile, but the leaf shapes, colour and trichomes can be used to separate these two species. Orthophytum mucugense has leaves that are entirely red at anthesis, while O. humile has leaves that are wine red to pink at the apices, but white and green toward the base. The leaf blades of O. humile are densely lepidote on both surfaces, differing from O. mucugense with glabrous to sparsely lepidote leaf blades. Additional specimens examined:—BRAZIL. Bahia: Mucugê, Serra do Capa Bode, 28 October 1978, (fl), Martinelli 5521 (RB); Parque Municipal Mucugê, cachoeira do Rio Tiburtino, 7 January 2003 (fr), Wanderley et al. 2367. (SP); 17 December 2006 (fr), Louzada & Moreira 58 (SP).

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 15 FIGURE 8. A–P. Orthophytum mucugense Wand. & A.A.Conc. (Wanderley et al. 2540, SP). — A. Habit. — B. Lateral view of external flower. — C. Ventral view of external flower. — D. Floral bracts. — E. Section of external flower. — F. Flower and floral bract. — G. Floral bract. — H. Sepal. — I. Petal. — J. Petal showing the petals appendages. — K. Central flower. — L. Apical bud. — M. Section of the Ovaries. — N. Seeds.

Basionym: Cryptanthopsis navioides Smith (1940: 31). Type:—BRAZIL. Bahia: Jacobina, on perpendicular rocks above stream in isolated ravine, 500 m, 16 June 1939, M. Foster & R. Foster 90 (holotype GH not seen, isotype R!, SP!).

FIGURE 9. A–F. Orthophytum navioides (L.B.Sm.) L.B.Sm. — A. Habit. — B. Section from the base of flower showing the epigenous tube. — C. Sepal and free stamen. — D. Sepal abaxial surface. — E. Petal showing an adnate stamen, petals appendages and the petals collosities. — F. Style and stigma (Jardim et al. 729).

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 17 Plants rupicolous, stoloniferous, stems short, ca 2 × 1.5 cm. Leaves patent or arching. Leaf sheaths white, triangular, ca 1 × 1.5 cm, glabrous, margins serrate, prickles ca 0.5 mm long. Leaf blades coriaceous, plane or slightly concave, green, internal one reddish, linear-triangular, 8–23 × 0.4–0.7 cm, attenuate, sparsely lepidote on both surfaces, margins serrate, prickles antrorse or retrorse, 1.1–5 mm long, apices long-attenuate. Inflorescences sessile, simple, many-flowered. Vestigial floral bract absent. Floral bracts subcoriaceous, green, symmetrical, carinate, triangular, ca 2 × 1 cm, lepidote, margins serrate, apices mucronulate. Sepals green, symmetrical, carinate, narrowly triangular, 2–3 × 0.5 cm, with glandular trichomes, margins entire, apices acuminate. Petals white, linear-spatulate, 2.5–3 × 0.4 cm, glabrous, apices subacute. Petal appendages sacciform, lacerate, ca 3 mm from the petal base. Stamens with antepetalous filaments ca 9 mm long and adnate to the petals, the antesepalous ones free, ca 20 mm long. Anthers yellow, 2.3–3 mm long, apices not apiculate. Ovaries trigonous, styles ca 20 mm long. Epigenous tubes present, ca 1.5 mm long. Ovules numerous. Fruits and seeds not seen. Distribution and habitat:—Orthophytum navioides is only known from Jacobina county in the state of Bahia, where it grows there on shaded rocks in dry forests near streams. IUCN conservation status:—CR B2ab (iii). Orthophytum navioides is only known from four herbarium specimens, including the type material. All collections were made at one locality with an estimated area of ca 10 km2. A decline in the quality of the habitat was observed, which justifies the conservation status proposed here. Comments:—Orthophytum navioides is similar to O. mucugense in habit, colour and texture of the leaves, bracts and sepals, but these species differ in the length of the leaf blades (8–23 vs. 2.5–6.7 cm in O. mucugense) and size of the flowers (longer in O. navioides). Additional specimens examined:—BRAZIL. Bahia: Jacobina, 4 August 2003 (fl), Ribeiro & Ribeiro s.n. (HB); Cachoeira Véu da Noiva, 27 October 1995 (fl), Jardim et al. 729 (CEPEC); 24 November 2001 (fl), Loureiro et al. 477 (CEPEC).

Orthophytum ophiuroides Louzada & Wanderley (2008: 406), figs. 10A–G, 13D.

Type:—BRAZIL. Bahia: Lençóis, Rio Mandassaia, 15 January 2006, M.G.L. Wanderley, A.A. Conceição, S.E. Martins & R.B. Louzada. 2532 (holotype SP!, isotype HUEFS!).

Plants rupicolous, stoloniferous, stems short, 1.2–3 × 1.5–2 cm. Leaves arching. Leaf sheaths white, broadly triangular or ovate-triangular, 0.4–0.8 × 0.7–0.8 cm long, glabrous, margins serrate, prickles 0.5 mm long. Leaf blades subcoriaceous, plane or slightly concave, green or partially red at anthesis, linear-triangular, 0.6– 26 × 0.2–0.4 cm, attenuate, glabrous or sparsely lepidote, margins serrate, prickles antrorse 0.3–0.4 mm long, apices long-attenuate. Inflorescences sessile, compound, many–flowered. Primary bracts subcoriaceous, red or pink, triangular-lanceolate, 2–8.5 cm long, lepidote, margins serrate, apices acuminate. Vestigial floral bract present. Floral bracts subcoriaceous, red, asymmetrical, carinate or ecarinate, triangular, 0.9–1.2 × 0.6– 0.7 cm, lepidote, margins serrate or entire, apices mucronulate. Sepals red, asymmetrical, carinate, narrowly triangular-ovate, ca 1 × 0.3 cm, glabrous, margins entire, apices acuminate. Petals white, linear-spatulate, ca 1.5 × 0.4 cm, glabrous, obtuse, apices apiculate. Petal appendages sacciform, lacerate, ca 6 mm from base. Callosities conspicous. Stamens with antepetalous filaments ca 3.5 mm long and adnate to the petals, the antesepalous ones free, ca 9 mm long. Anthers yellow, 2.3–2.5 mm long, apices apiculate. Ovaries trigonous, styles ca 10 mm long. Epigenous tubes absent. Ovules numerous. Fruits and seeds not seen. Distribution and habitat:—Orthophytum ophiuroides is only known from Lençóis county in Bahia. It occurs on rocky outcrops in shaded habitats near streams. IUCN conservation status:—EN B2ab (ii, iii). Orthophytum ophiuroides is known from a few herbaria collections and it is a species with a limited distribution; an area less than 500 km2 was estimated. It is considered threatened with a risk of becoming extinct in the wild, because it occurs in less than five localities and its suitable habitat is declining.

18 • Phytotaxa 13 © 2010 Magnolia Press LOUZADA & WANDERLEY FIGURE 10. A–G. Orthophytum ophiuroides Louzada & Wand. (Wanderley et al. 2532, SP). — A. Habit. — B. Branch of inflorescence and primary bract. — C. Flower and floral bract. —D. Floral bract. — E. Sepal. — F. Petal and stamen showing the lacerate petal appendages. — G. stigma.

Comments:—Orthophytum ophiuroides is similar to O. navioides, but differs from that species by having shorter petals (ca. 1.5 vs. 2.5–3 cm) and sepals (ca. 1 vs. 2–3 cm), without glandular trichomes. In addition, the bracts and sepals are red (not green).

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 19 Erroneous identifications of Orthophytum ophiuroides in herbarium collections are quite common. Vouchers are sometimes identified as O. humile, but O. ophiuroides has wider and glabrous leaf blades (vs. narrow and densely lepidote). Additional specimens examined:—BRASIL. Bahia: Lençóis, Poço Harley, 13 January 2006 (fl), Louzada & Moreira 7 (SP); 15 January 2006 (fl), Wanderley et al. 2531 (SP); 8 April 2008 (fl), Louzada & Wanderley 88 (SP).

Orthophytum pseudovagans Leme & Kollmann (2007: 155), fig. 13C.

Type:—BRAZIL. Espírito Santo: Águia Branca, propriedade de Ciro Ferreira, 18º 58’40.5”S, 40º 39’56.1”W, 250 m, 27 April 2006, V. Demuner, L.F.S. Magnago, T. Cruz & E. Bausen 2270 (holotype MBML!, isotype HB not seen).

Plants rupicolous or terrestrial, stoloniferous, stems elongate, 12–15 × 1.8 cm. Leaves slightly arching. Leaf sheaths green, ovate, 3.6 × 2–2.5 cm, lepidote at base, margins serrate, prickles ca 3 mm long. Leaf blades coriaceous, canaliculate, green, internal one reddish, narrowly triangular, 30–55 × 1.5–1.7 cm, attenuate, lepidote, glabrous and bright at base, margins serrate, prickles straight or retrorse, 2–3.5 mm long, apices acute. Inflorescences sessile, compound, many–flowered. Primary bracts subcoriaceous, green or red, very narrowly triangular, 3.2–12.7 × 1.8 cm, margins serrate, apices acuminate. Floral bracts subcoriaceous, green, asymmetrical, carinate, narrow triangular, ca 1.6 × 0.8 cm, lanate, margins serrate, apices acuminate. Sepals green, asymmetrical, carinate, triangular-lanceolate, 1.6–1.8 × 0.4 cm, densely lepidote, margins entire, apices acuminate. Petals green with white margins, spatulate, 1.7–1.8 × 0.5–0.6 cm, glabrous, obtuse, apices cucullate. Petal appendages cupuliform, lacerate, ca 3 mm from base. Callosities conspicuous, ca 2.5 mm long, apices not apiculate. Ovaries trigonous. Epigenous tubes absent. Ovules numerous. Fruits and seeds not seen. Distribution and habitat:—Orthophytum pseudovagans is only known from Águia Branca county in the state of Espírito Santo, where it grows on rocky outcrops in the Atlantic rain forests at ca 250 m elevation. IUCN conservation status:—DD. Orthophytum pseudovagans still lacks sufficient distribution data, because it was discovered only recently and is currently known only from the type material. Comments:—According to Leme & Kollmann (2007), O. pseudovagans forms the 'vagans subcomplex' together with O. vagans Foster (1960: 59) and O. zanonii Leme (2004b: 72). Long stems and obtuse–cucullate petal apices characterize this group of species. Orthophytum pseudovagans differs from O. vagans mainly by its compound (vs. simple) inflorescences. It differs from O. zanonii by longer leaf blades (30–55 cm vs. 12–14.6), which are glabrous, bright green– spotted and red at the base (vs. lepidote and concolorous green).

Orthophytum ulei Louzada & Wand. sp. nov., figs. 11A–F, 13E

Ortophytophytum albopicto affine sed bracteis primariis, bracteis floralibus et sepalis roseis (vs viridibus), bracteis floralibus carenatis (vs non carenatis) et petalis glabris basim versus (vs petalis trichomatibus praeditis basim versus) differt. Type:—BRAZIL. Bahia: Mucugê road to Andaraí, 12o56’41.2”S, 41o17’28.5”W, ca 867 m. 9 April 2008 (fl), R.B. Louzada & M.G.L. Wanderley 91 (holotype SP, isotype HUEFS).

Plants rupicolous, stoloniferous, stems short ca 2.6 cm long. Leaves patent or arching. Leaf sheaths greenish- white, triangular or ovate, 1–1.4 × 1.3–2.2 cm, glabrous, margins serrate, prickles ca 0.5 mm long. Leaf blades coriaceous, slightly concave, pink at base, wine for the, very narrowly triangular, 5–15.5 × 1.2–1.6 cm, attenuate, abaxially densely lepidote, adaxially lepidote, scales densely arranged at base, margins serrate, prickles antrorse 2–3.5 mm long, apices mucronate. Inflorescences sessile, simple or compound, many-

20 • Phytotaxa 13 © 2010 Magnolia Press LOUZADA & WANDERLEY FIGURE 11. A–F. Orthophytum ulei Louzada & Wand. (from the holotype). — A. Habit. — B. Outer floral bract. — C. Inner floral bract. — D. Flower showing the petals and in the detail the apices of the posterior connate sepals. — E. Sepal and free stamen. — F. Petal, showing petal appendages, callosities and stamen.

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 21 flowered. Primary bracts subcoriaceous, pink, triangular or lanceolate, 2–5 × 0.8 cm, lepidote, margins serrate, apices mucronate. Vestigial floral bract present. Floral bracts subcoriaceous, pink, asymmetrical, triangular, 1.2–1.7 × 0.7–1 cm, lepidote, margins serrate, apices mucronate. Sepals pink, asymmetrical, carinate, narrowly triangular, 1.3–1.6 × 0.3–0.6 cm, lepidote, margins entire, apices mucronate, anterior one free, posterior connate. Petals white, linear-spatulate, 1.3–1.7 × 0.4 cm, glabrous, apices obtuse. Petal appendages sacciform, lacerate, ca 5 mm from the base. Callosities conspicous. Stamens with antepetalous filaments ca 5 mm long and adnate to the petals, the antesepalous one free ca 10 mm long. Anthers, yellow, ca 3 mm long, apices apiculate, apiculus absent in dry material. Ovaries trigonous. Epigenous tubes present, ca 2 mm long. Ovules numerous. Fruits and seeds not seen. Distribution and habitat:—Orthophytum ulei is only known from Mucugê county in Bahia. It grows rupicolous in xeric environments, frequently exposed to the sun on rocky outcrops and in grassland on rocky fields at ca 870 m elevation. IUCN conservation status:—EN B1ab (ii, iii). Based on collections and field observations in the areas of occurrence, we noticed that O. ulei is restricted to an area estimated to be less than 5000 km2. It is only known from one county in Bahia, occurring outside of protected areas. Comments:—Orthophytum ulei is similar to O. amoenum and O. albopictum. It differs from O. amoenum mainly in having larger leaf blades, (1.2–1.6 vs. 0.5–1 wide), that are adaxially lepidote (vs. sparsely lepidote), lepidote floral bracts and sepals (vs. glabrous) and narrowly triangular (vs. elliptic) sepals. It differs from O. albopictum mainly in its primary bracts, floral bracts and sepals: sepals are pink (vs. green), floral bracts carinate (vs. ecarinate) and petals are glabrous (vs. trichomes below the petal appendages). Orthophytum albopictum has a white-lanate well-delimited band around the inflorescence, formed by dense scales (peltate trichomes) at the base of leaves and primary bracts, the prickles are not longer than 2 mm, while in O. ulei the prickles are 2–3.5 mm long. Etymology:—The name of this new species is attributed to Ernst Heinrich Georg Ule (1854–1915), German botanist and explorer, who undertook botanical expeditions to southern and central Brazil at the end of the 19th the century. In Chapada Diamantina, previously known as Serra do Sincorá, Ule discovered a new species of Bromeliaceae and described it in a new genus, Sincoraea (Ule 1908). Later, Smith (1955) synonymized this genus under Orthophytum (Smith 1955). Additional specimens examined:—BRASIL. Bahia: Mucugê, 22 km from Andaraí in road to Mucugê, 16 February 1977 (fl), Harley 18746 (RB); Serra do Capa Bode, 30 October 1972 (fl), Martinelli 5545 (RB); 29 October 1978 (fl), Martinelli 5546 (RB).

Orthophytum vagans Foster (1960: 59).

Type:—BRAZIL. Espírito Santo: R.G. Wilson 578 (holotype US!).

Plants rupicolous, stems elongate, ca 19 cm long. Leaves slightly arching. Leaf sheaths ovate, 1.5–2 × 1.7–2.5 cm, margins serrate, prickles 0.5–3 mm long. Leaf blades subcoriaceous, canaliculate, green, internal one reddish white, narrowly triangular, attenuate, sparsely lepidote, margins serrate, prickles antrorse, apices mucronate. Inflorescences sessile, simple, few-flowered. Floral bracts subcoriaceous, narrowly triangular, margins serrate, apices acuminate. Sepals white, narrowly triangular, lanate, margins entire, apices acuminate. Petals green with white margins, linear-spatulate, apices obtuse-cucullate. Petals appendages cupuliform, lacerate. Stamens with antepetalous filaments ca 10 mm long and adnate to the petals,the antesepalous ones free 18 mm long. Ovaries trigonous. Fruits and seeds not seen. Distribuition and habitat:—unknown IUCN conservation status:—DD. Orthophytum vagans was described from cultivated material found in a garden in Rio de Janeiro. It has not yet been found in the wild.

22 • Phytotaxa 13 © 2010 Magnolia Press LOUZADA & WANDERLEY Comments:—Orthophytum vagans is similar to O. zanonii as both are having an elongated stem and obtuse-cucullate petals. It differs from O. zanonii mainly by having a simple inflorescence (vs. compound). Herbarium collections of Orthophytum vagans other than the type material are not known. Live material was brought to the United States by R.G. Wilson, and initially, the author of this species thought it was Cryptanthus glazioui Mez (1891–94: 202), due to the elongated stems. However, after observing the fertile plants in cultivation, Foster (1960) concluded that this plant was a new species of Orthophytum, easily recognized by the elongated stems. The presence of elongated stems forming a clump, inspired the name vagans, meaning 'wandering'.

Orthophytum zanonii Leme (2004b: 72), figs. 12A–J, 13F.

Type:—BRAZIL. Espírito Santo: Pancas, Laginha de Pancas, Pedra do Vidal, property of Vidal Krause, ca. 300 m, 16 August 2003, E.M.C. Leme, M. Zanoni & E. Malgnago 5930 (holotype HB!).

Plants rupicolous, stems elongate, ca 10 × 2 cm. Leaves arching. Leaf sheaths greenish-white, ovate, 2.3–3 × 2.1–3 cm, sparsely lepidote, margins serrate, prickles 0.2–0.3 mm long. Leaf blades coriaceous, slightly concave, red or wine when young, linear–triangular, 12–14.6 × 1.1–1.5 cm, attenuate, densely lepidote on both surfaces, margins serrate, prickles antrorse 0.6–1 mm long, apices mucronate. Inflorescences sessile, compound, many-flowered. Primary bracts coriaceous, green, triangular to very narrowly triangular, 3.2–12.7 × 1.8 cm, densely lepidote, margins serrate, apices mucronate. Vestigial floral bract present. Floral bracts coriaceous, green, asymmetrical, carinate, triangular, 2.1–2.5 × 1.2 cm, lepidote, margins serrate, apices mucronate. Sepals green, symmetrical, ecarinate, triangular-ovate, ca 1.6 × 0.5 cm, densely lepidote, margins entire, apices mucronulate. Petals green with white margins, linear-spatulate, ca 1.9 × 0.4 cm, glabrous, obtuse, apices cucullate. Petal appendages cupuliform, lacerate, ca 6 mm from base. Callosities conspicous. Stamens with antepetalous filaments ca 6 mm long and adnate to the petals, the antesepalous ones free, ca 15 mm long. Anthers green, ca 2.2 mm long, apices apiculate. Ovaries trigonous, styles ca 1.7 mm long. Epigenous tube absent. Ovules numerous. Fruits and seeds not seen. Distribuition and habitat:—Orthophytum zanonii is only known from Pancas county, district of Laginha in the state of Espírito Santo. It grows on inselbergs in Atlantic rain forest between 430–550 m elevation. IUCN conservation status:—EN B2ab (ii, iii). Orthophytum zanonii is known from a single locality with an estimated range of less than 500 km2. A decline of the extent of its habitat may bring this species in danger of extinction. Comments:—Orthophytum zanonii is similar to O. vagans and O. pseudovagans, all having an elongated stem. Orthophytum zanonii differs from O. vagans in its densely lepidote (vs. sparsely lepidote) leaves and coriaceous (vs. subcoriaceous) and concolorous leaf blades that become red (vs. green) at anthesis. In O. zanonii the leaf margins are serrated with prickles that are half as long as those of the other two species (Leme 2004b). The species presents a peculiar vegetative reproduction, forming extensive populations with lateral shoots emerging from the leaf axils and bract axils of the inflorescence. The leaves may be densely lepidote with gray hairs when exposed to intense light conditions, but become green and bright in some localities of the inselbergs that are more shaded. Additional specimens examined:—BRAZIL. Espírito Santo: Pancas, Laginha de Pancas, Pedra do Vidal, 16 August 2003 (fl), Leme et al. 5941 (HB); 5 August 2006 (fl), Louzada et al. 18 (SP).

Acknowledgements

We acknowledge the support from several Brazilian agencies: Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq). We

REVISION OF ORTHOPHYTUM (BROMELIACEAE) Phytotaxa 13 © 2010 Magnolia Press • 23 thank Dr Tarciso Filgueiras for language editing, for latin diagnosis and suggestions to the manuscript, and Jan Schnitzler, Maria Cláudia Medeiros and Pedro Schwartsburd for language editing; Klei Souza for the drawings; Suzana E. Martins, Abel A. Conceição, Oscar Ribeiro, Cyrio Santana and Luis F. Magnago for the photographs. We also thank curators of the herbaria (ALCB, CEPEC, HB, MBM, MBML, RB, SP, SPF, HUEFS) for access to specimens.

FIGURE 12. A–J. Orthophytum zanonii Leme. (Louzada et al. 18, SP). — A. Habit. — B. Floral bract. — C. Branch of inflorescence with two flowers. — D. Flower. — E. Sepal. — F. Sepal showing and free stamen. — G. Petal, showing the petal appendages and callosities. — H. Detail of the petal appendages. — I. Style and stigma. — J. stigma.

24 • Phytotaxa 13 © 2010 Magnolia Press LOUZADA & WANDERLEY FIGURE 13. — A. Orthophytum mucugense. — B. Orthophytum navioides. — C. Orthophytum pseudovagans. — D. Orthophytum ophiuroides. — E. Orthophytum ulei. — F. Orthophytum zanonii. (Photos: Cyrio Santana [A], Oscar Ribeiro [B], Luiz Magnago [C]).

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