Three new taxa in Cyrtopodium () from central and southeastern Brazil Author(s): João A. N. Batista and Luciano B. Bianchetti Source: Brittonia, 56(3):260-274. Published By: The New York Botanical Garden https://doi.org/10.1663/0007-196X(2004)056[0260:TNTICO]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1663/0007-196X%282004%29056%5B0260%3ATNTICO %5D2.0.CO%3B2

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Three new taxa in Cyrtopodium (Orchidaceae) from central and southeastern Brazil

JOAÄ O A. N. BATISTA AND LUCIANO B. BIANCHETTI

Batista, J. A. N. & L. B. Bianchetti (Embrapa Recursos GeneÂticos e Biotec- nologia, Parque EstacËaÄo BioloÂgica, P.O. Box 02372, BrasõÂlia, DF 70770-901, Bra- zil; emails: [email protected]; [email protected]). Three new taxa in Cyrtopodium (Orchidaceae) from central and southeastern Brazil. Brittonia 56: 260±274. 2004.ÐCyrtopodium brunneum, C. lamellaticallosum and C. poecilum var. roseum, from the cerrado and campo rupestre vegetation of central and southeastern Brazil are described and illustrated. Although the description of Cyrtopodium gonzalezii indicated that it is a distinct , the holotype is referrable to C. brandonianum. The undescribed are now de- scribed here as C. brunneum. Cyrtopodium gonzalezii is placed in the synonym of C. brandonianum. Key words: Orchidaceae, Cyrtopodium, cerrado vegetation, Brazil.

Cyrtopodium is a neotropical with characterization, we found that the holotype about 42 species ranging from southern of C. gonzalezii L. C. Menezes is unambig- Florida to northern Argentina. The center of uously C. brandonianum Barb.Rodr., al- diversity of the genus is in the Brazilian though the description does indicate that it cerrado, where at least 25 species occur. In is a new species, here described as C. brun- recent years, several new species have been neum. described in the genus (Batista & Bian- chetti, 2001; Bianchetti & Batista, 2000; Cyrtopodium brunneum J. A. N. Bat. & Menezes, 2000), most from central Brazil. Bianchetti, sp. nov. (Figs. 1 & 2) Based on our further exploration of the re- gion and examination of herbarium speci- TYPE: BRAZIL. Distrito Federal, BrasõÂ- mens of the genus, we describe two new lia, Plano Piloto, ®nal da Asa Norte, aÂrea species, C. brunneum and C. lamellatical- entre o Parque EcoloÂgico Norte e o Parque losum, and a new variety, C. poecilum var. Nacional de BrasõÂlia, local do futuro Setor roseum, below. Noroeste, 8 Sep 2001 (¯), J. A. N. Batista Traditionally, the of Cyrtopo- 1242 (HOLOTYPE: CEN; ISOTYPES: AMES, dium has been based almost completely on BHCB, ESA, HB, HUEFS, HUFU, K, reproductive characters, with the vegetative MBM, MO, NY, RB, SP, SPF, UB, UEC). parts, particularly the leaves, being neglect- Cyrtopodium gonzalezii L. C. Menezes, Bol. CAOB ed. This is because botanical studies of the 6(1): 9. 1995. pro parte, excluding type. genus have relied almost entirely on dried specimens collected mainly during the Cyrtopodio tristi Rchb. f. & Warm. similis sed in- ¯orescentia semper simplici, ¯oribus minoribus, sepal- ¯owering season, when most Cyrtopodium is oblongo-lanceolatis vel paulo lanceolato-ovatis, la- species do not have fully developed leaves. belli lobis lateralibus falcatis differt; etiam C. dusenii The examination of live plants throughout Schltr. similis sed foliis sub anthesi redactis, in¯ores- the developmental cycle has allowed char- centia semper simplici, ¯oribus majoribus, sepalis et acterization of the vegetative parts of most petalis atrobrunneis differt. species and, in many cases, the species can Terrestrial herb. Roots many, whitish, be identi®ed vegetatively. Based on this glabrous. Pseudobulbs completely buried,

Brittonia, 56(3), 2004, pp. 260±274. ISSUED: 2 August 2004 ᭧ 2004, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. 2004] BATISTA & BIANCHETTI: CYRTOPODIUM (ORCHIDACEAE) 261

FIG.1. Cyrtopodium brunneum. A. Habit at anthesis. B. Flower. C. Side view of ¯ower. D. Callus. E. Perianth. a±e. Variations in lip morphology, each one from a different individual of the same population. A±D drawn from Batista 1242, E and a±e drawn from Batista 208 by Simone C. Souza e Silva. 262BRITTONIA [VOL. 56

FIG.2. Cyrtopodium brunneum. A. Habitat at the time of ¯owering. Burnt cerrado at the type locality, about 2 weeks after ®re, with the vegetation beginning to regrow, at the beginning of September. B. In¯orescence. Note the poorly developed leaves at anthesis. C. Fully developed leaves, in December, about 2 months after ¯owering. The surrounding vegetation has been removed to expose the plants. D. Uprooted mature (Jan- uary), showing the fully developed leaves and underground pseudobulbs. E and F. Flowers, each from a different specimen. Scale bars ϭ 5 cm (B±D) and 5 mm (E, F). 2004] BATISTA & BIANCHETTI: CYRTOPODIUM (ORCHIDACEAE) 263 ovoid, lea¯ess from the second year on- apex with a protruding fold, retuse when wards, externally white, (3±)4±6(±7) ϫ ¯attened, the margin smooth, occasionally (1±)1.5±2.5 cm. Leaves at ¯owering 3±6, slightly verrucose at the apex. Column little developed, 7±15 cm long, when fully erect, slightly arcuate, trigonous, 6±7(±8) developed 5±8, spreading, coriaceous, lan- mm long, yellow at the base, reddish white ceolate, the lowermost 1±2 leaves sheath- at the middle, and greenish purple towards like, the uppermost 4±7 leaves 13±33 ϫ 1± the apex; column foot (2.5±)3±4 mm long, 2.6 cm, articulate 1±2 cm from the surface yellow at the base, white at the middle with of the soil, the apex acuminate. In¯ores- small numerous red dots that merge to- cence lateral, erect, racemose, lax, 26±56 wards the apex. Anther ca. 2±2.5 ϫ 1.5 mm, cm, brownish to dark brown; peduncle 11± yellowish, apex green; pollinia 2, waxy, 26 cm, the 2 sheath-like bracts strongly ad- sulcate, ca. 1 ϫ 0.5±0.6 mm, yellow; stipe pressed, 1.7±3.5 cm long; rachis 12±26 cm triangular, hyaline, ca. 1 ϫ 1 mm at base. long; ¯oral bracts oblong-lanceolate, 1±3 ϫ Fruit de¯exed, green, oblong to elongate, 0.5±0.9 cm, greenish brown, acute to acu- sulcate, ca. 6±8.5 ϫ 1±1.5 cm when young, minate, the margins undulate, incurved; including the pedicel. ovary with pedicel 1.4±3.9 cm long, dark Etymology. Named after the dark brown brown. Flowers 9±15(±20) per in¯ores- color of the sepals and petals. cence, sweet-scented. Sepals oblong-lance- Additional specimens examined. BRAZIL. Distrito olate to slightly lanceolate-ovate, light to Federal: BrasõÂlia, Vila Maury, 6 Sep 1960 (¯), Andra- dark brownish or brownish green, lighter at de 412 & Emmerich 404 (HB, R); BrasõÂlia, Lago Nor- the base, frequently with dark-brown spots, te, 8 Oct 1990 (¯), Batista 122 (CEN), 1 Oct 1991 (¯), the apex apiculate, the margin undulate; Batista 208 (CEN), 7 Sep 1992 (¯), Batista 325 (CEN); GuaraÂ, Reserva EcoloÂgica do GuaraÂ, 13 Sep dorsal sepal (8±)11±13(±15) ϫ 4±5(±6) 1992 (¯), Batista 330 (CEN); BrasõÂlia, area between mm; lateral sepals slightly oblique, (8±)11± Parque EcoloÂgico Norte and Parque Nacional de Bra- 13(±15) ϫ 4±6(±7) mm. Petals concave, sõÂlia, 7 Oct 1994 (¯), Batista 421 (CEN), 11 Dec 1994 broadly elliptic to somewhat broadly lan- (veg), Batista 434 (CEN); BrasõÂlia, Lago Norte, 17 Feb ceolate, (8±)9±12(±13) ϫ 6±8 mm, the low- 1995 (veg), Batista 521 (CEN), QL 15, near Clube do Congresso, 17 Feb 1995 (veg), Batista 523 (CEN); NuÂ- er half greenish to light-brownish, occa- cleo Bandeirante, SantuaÂrio EcoloÂgico do Riacho Fun- sionally with few to many small brown do, 14 Sep 1995 (¯), Batista & Bianchetti 576 (CEN); dots, the upper half dark-brown, color tran- BrasõÂlia, area between Parque EcoloÂgico Norte and sition usually sharply delimited, the apex Parque Nacional de BrasõÂlia, 28 Sep 1998 (¯), Batista 792 (CEN, HB, MBM, SP), 30 Dec 2000 (veg), Batista obtuse to rounded, slightly apiculate, the 1112 (CEN); BrasõÂlia, area between Lago Norte and margins slightly undulate. Lip 3-lobed, EPIA, 6 Jan 2001 (veg), Batista 1126 (CEN); BrasõÂlia, (7±)9±11(±12) mm long, 10±12(±14) mm Lago Sul, condomõÂnio Prive Morada Sul II, above QI- wide between the apices of the side lobes 28, close to Paranoa dam, (¯. in cult.) 22 Sep 2001, when spread, the base shortly unguiculate, Batista et al. 1184 (CEN); Gama, BR-060, near the GDF check point and the access to Santo AntoÃnio do 1 mm long, yellowish; lateral lobes erect, Descoberto, 2 Oct 2001 (¯), Batista & Oliveira-Neto parallel, oblong-falcate, (3±)4±5(±6) ϫ 2± 1252 (CEN); BrasõÂlia, Plano Piloto, Asa Norte, 713± 3(±4) mm, reddish to reddish purple, the 714 N, area of University of BrasõÂlia, between Parque apex obtuse, the base unconstricted, the Olhos D'A gua and the CAESB sewage treatment plant, 7 Oct 2001 (¯), Batista 1254 (CEN); BrasõÂlia, margins entire, smooth; callus verrucose, Setor de MansoÄes do Lago Norte, 17 Sep 1990 (¯), slightly sulcate, extending from the base of Bianchetti & Batista 953 (CEN), trecho 2, 20 Oct 1991 the central lobe to the column foot, white, (¯), Bianchetti & Batista 1176 (CEN), near MI-7, 6 occasionally reddish white; isthmus sepa- Oct 1991 (¯), Bianchetti & Batista 1168 (CEN); rating the lateral lobes from the central lobe GuaraÂ, Reserva EcoloÂgica do GuaraÂ, 26 Aug 1990 (¯), Bianchetti & Batista 950 (CEN); BrasõÂlia, Plano Pil- usually prominent, sometimes elongated, 1± oto, 10 Nov 1961 (¯), Heringer 8756 (HB, UB); Bra- 2(±3) mm long; central lobe variable, ob- sõÂlia, EstacËaÄo Florestal CabecËa do Veado, 8 Oct 1965 cordate to obovate-oblong to somewhat re- (¯), Heringer 10616 (HB, UB); Santa Maria, near the niform, 5±7 ϫ (5±)6±8(±10) mm, yellow microwave towers, 28 Oct 1972 (¯), Heringer 12206 (HB, UB); ca. 20 km S of BrasõÂlia, on rd. to GoiaÃnia, with red to reddish-purple margins, occa- near Rio Melchior, 1125 m, 25 Sep 1965 (¯), Irwin et sionally with a few red dots at the base, the al. 8652 (UB); BrasõÂlia, con¯uence of Rio Torto with yellow becoming fainter towards aging, the Lake ParanoaÂ, 975 m, 9 Oct 1965 (¯), Irwin et al. 9086 264BRITTONIA [VOL. 56

(HB); GuaraÂ, Reserva EcoloÂgica do GuaraÂ, 15Њ50ЈS, ear, erect, very narrow leaves. Compared 47Њ57ЈW, 1050 m, 20 Sep 1994 (¯), Oliveira 2 (UB). with the 25 Cyrtopodium species known GoiaÂs: Chapada dos Veadeiros, ca. 26 km N from Alto ParaõÂso, GO-118, 11 Oct 1999 (¯), Batista 945 (CEN); from the cerrado, the buried pseudobulb ca. 8 km from NiquelaÃndia, Tocantins Niquel Com- characteristic excludes several species with pany, 14Њ23Ј48ЉS, 48Њ25Ј59ЉW, 17 Sep 1996 (¯), Fon- exposed pseudobulbs, such as C. aliciae seca et al. 1154 (IBGE); Municipality of Mineiros, Linden & Rolfe, C. cardiochilum Lindl., C. BR-364, near CoÂrrego Alegre, 20 Sep 1974 (¯), Hatschbach & Kummrow 35014 (MBM); GoiaÃnia, 30 cipoense L. C. Menezes, C. cristatum Nov 1963 (¯), Heringer 9289 (HB); Municipality of Lindl., C. eugenii Rchb. f., C. hatschbachii LuziaÃnia, 16 Sep 1974 (¯), Heringer 13971 (HB, UB); Pabst, C. lissochiloides Hoehne & Schltr., 12 km NW of Veadeiros, on rd. to Cavalcante, 1200 C. palmifrons Rchb. f. & Warm., C. palu- m, 21 Oct 1965 (¯), Irwin et al. 9444 (HB, UB); Apa- dicolum Hoehne, C. parvi¯orum Lindl., C. recida de GoiaÃnia/HidrolaÃndia, 10 Aug 2002 (¯), Pas- tore 32 (CEN). Mato Grosso: 84 km from Alto Ara- saintlegerianum Rchb. f., C. virescens guaia, towards RondonoÂpolis, BR-364, Sep-Nov 1983 Rchb. f. & Warm., and C. vernum Rchb. f. (¯), Hutchison 8550 (UEC). Minas Gerais: UberlaÃn- & Warm. dia, EstacËaÄo EcoloÂgica do Panga, 24 Sep 1992 (¯), According to the protologue of C. gon- Araujo et al. 276 (HUFU); Morro Pelado, near Cam- panha, Aug 1896 (¯), BrandaÄo in CGGMG 1719 (R) zalezii (Menezes, 1995) and to Menezes (p.p., with C. cristatum); Municipality of Carmo do herself (L. C. Menezes, pers. comm.), the Rio Claro, Serra da Tormenta, 3 Nov 1990 (¯), Cam- holotype was collected on the peninsula do pos s.n. (CEN, HRCB); Serra do CipoÂ, 23 Aug 1958 Lago Norte, a district of BrasõÂlia where we (¯), Heringer 6438 (HB) (p.p., with C. parvi¯orum); have collected intensively during the past FelixlaÃndia, TreÃs Marias basin, 23 Aug 1958 (¯), Her- inger 6438 (UB), 2 Oct 1959 (¯), Heringer 6438 (UB), 12 years. Six species of Cyrtopodium are margins of Rio Paraopeba, 11 Oct 1959 (¯), Heringer known from this district: C. blanchetii 6438 (UB); Santana do Riacho, Km 107 on the Belo Rchb. f., C. brandonianum Barb.Rodr., C. Horizonte to ConceicËaÄo do Mato Dentro rd., 4 Oct brunneum, C. caiapoense L. C. Menezes, 1981 (¯), Pirani et al. in CFSC 7472 (SP, SPF). C. poecilum Rchb. f. & Warm., and C. vi- Pabst was probably the ®rst orchid tax- rescens Rchb. f. & Warm. This further ex- onomist to examine material of C. brun- cludes from the comparison those species neum, but misidenti®ed it ®rst as C. falci- that do not occur in the Distrito Federal, lobum Hoehne & Schltr. (now a synonym such as C. braemii L. C. Menezes and C. of C. parvi¯orum Lindl.) and later as C. dusenii Schltr., or those that occur in Bra- poecilum Rchb. f. & Warm. f. minor Hoeh- sõÂlia but not at this particular site, such as ne (now a synonym of C. fowliei L. C. Me- C. fowliei L. C. Menezes, C. latifolium nezes). He was apparently also the ®rst tax- Bianchetti & J. A. N. Bat., C. pallidum onomist to suspect that the specimens might Rchb. f. & Warm., C. triste Rchb. f. & represent a new species. In his personal ®les Warm., and C. linearifolium J. A. N. Bat. at the Herbarium Bradeanum, there is a card & Bianchetti. Additionally, the pseudobulbs with a sketch of C. brunneum indicated at of C. braemii, C. fowliei and C. latifolium the side as C. aff. parvi¯orum and at the are reddish-purple externally, while in the top as Cyrtopodium heringeri, but he never holotype of C. gonzalezii they are whitish. published a valid description of this species. The characterization of the vegetative More recently, Menezes described this spe- parts of the remaining species for compar- cies as Cyrtopodium gonzalezii L. C. Me- ison, including the holotype of Cyrtopo- nezes (Menezes, 1995), but unfortunately dium gonzalezii, is shown in Table I. Be- designated a holotype that is Cyrtopodium cause of its similarity to C. brunneum, C. brandonianum Barb.Rodr. This situation triste is also included in the comparison. leaves the new species without valid pub- All measurements were taken from living lication. specimens under ®eld conditions, except for The holotype of C. gonzalezii at the Uni- the measurements of pseudobulbs and the versity of BrasõÂlia herbarium is a single, holotype of C. gonzalezii, which were ob- sterile specimen. However, on the basis of tained from dried specimens. Only mature its vegetative characteristics this specimen and adult specimens and fully developed is C. brandonianum. The holotype has leaves were considered in the analysis. Leaf small, narrow, buried pseudobulbs and lin- length was measured from the surface of 2004] BATISTA & BIANCHETTI: CYRTOPODIUM (ORCHIDACEAE) 265 7.5±8 0.8±1.2 1.2±2.7 2±3.5(±6) RAZIL buried white (6±)8±10 (12±)25±45 15.3±18.5(±21.6) 10±22(±31.7) lanceolate erect present ,B LIA õ Â RAS B (4.5±)5±8(±8.5) 1.5±2.5(±3.5) buried white (3±)4±5(±6) (18.5±)35±65(±85) (1.3±)2±2.9(±3.4) (11.6±)18.5±25(±33.2) (8.8±)14±30(±47.2) lanceolate erect present (3±)4.5±6.5(±8) ORTE DISTRICT OF N AGO L C. poecilum C. blanchetii C. caiapoense (5±)6±8(±11.5) 1.5±2.8 buried reddish purple (12.7±)16.5±18.5(±30) (7.6±)11.3±20.8(±30.6) lanceolate spreading present (2.5±)3.5±6.5(±8) (3±)5±6 (14±)35±55(±81) (1.3±)2.5±3.5(±5.1) KNOWN FROM THE C. 3.5±4 1.2±1.3 4±4.5 5 0.5±0.9 gonzalezii (holotype) I buried white 55 55 linear erect present 22±42 ABLE T Cyrtopodium 3.5±5.5(±6.5) 0.8±1.5(±2) buried white (31.2±)37±42(±60) (15±)30±58(±76) linear erect present (2.5±)4±6(±10.5) (3±)4±5(±7) (24±)41±60(±81) (0.7±)1±1.5(±2.1) (3±)4±6 1±2.3 buried white 12.9±15.5(±21.9) (7.2±)10±21(±29.3) linear-lanceolate spreading present (0.5±)1±2(±2.5) (3±)4±5(±6) (5±)12±21(±31) (0.8±)1.1±1.6(±2.0) C. brunneum C. triste C. brandonianum (4±)5±6.5(±7.5) (1.4±)1.7±2.3(±2.8) buried white (8.1±)9.5±15.5(±17.2) (6.7±)8.5±16.7(±22) lanceolate spreading present (0.5±)1±2(±3) (4±)5±7(±8) (6±)16±25(±33) (0.6±)1.1±2.1(±2.6) EGETATIVE CHARACTERISTICS IN SEVEN TERRESTRIAL SPECIES OF V 1 2 Mean ratio per plant. Ratio per leaf. 1 2 Length (cm) Width (cm) Position Color Ratio L/W Shape Position Articulation Length (cm) Ratio L/W Number Length (cm) Width (cm) Pseudobulbs Leaves 266BRITTONIA [VOL. 56 the soil, which corresponds approximately Cyrtopodium species in central Brazil, to the distance from the apex of the pseu- ¯ower at the end of the dry season and the dobulb to the apex of the leaf. Cyrtopodium beginning of the rainy season (September poecilum and C. blanchetii have much to November), while C. brandonianum broader leaves than C. gonzalezii. The ¯owers mainly during the peak of the rainy pseudobulbs of C. poecilum are reddish- season (December and January). However, purple externally, and therefore distinct in common with most other species, the from all the other species in this compari- vegetative growth of C. brandonianum son; the pseudobulbs of C. blanchetii are starts at the beginning of the rainy season, larger than those of C. gonzalezii. Cyrto- usually during October, so that in Novem- podium caiapoense has more and wider ber, plants have already initiated vegetative leaves and usually has longer pseudobulbs growth but have incompletely developed than C. brunneum. On the other hand, C. leaves. brunneum has shorter and broader leaves. Of the remaining terrestrial species of The mean leaf length-to-width ratio was Cyrtopodium with small, buried pseudo- 9.5±15.5 for C. brunneum, and 55 for C. bulbs, only two have long, linear, straight gonzalezii. Additionally, the leaves of C. leaves like the holotype of C. gonzalezii. brunneum are typically spreading, while in The ®rst, C. linearifolium, occurs in dark, C. gonzalezii they are erect. The pseudo- sandy-clay soil associated with campo ru- bulbs of C. brunneum are usually slightly pestre vegetation found at higher altitudes, longer and broader than those of C. gon- has nonarticulated leaves, and, in the Dis- zalezii. Another difference involves the dis- trito Federal, is known only from the higher tance of the leaf articulation relative to the altitude meadows of the Chapada da Con- apex of the pseudobulb, which was shorter tagem. The second, C. pallidum, occurs at in C. brunneum than in C. gonzalezii.A several sites throughout the Distrito Federal plot of leaf length versus width for C. brun- but typically grows in dark, sandy-clay soil neum, the holotype of C. gonzalezii, and C. found in wetter areas (close to gallery for- brandonianum is shown in Figure 3. These ests) usually associated with regularly results show that the vegetative character- spaced mounds of earth known as murun- istics of the holotype of C. gonzalezii are dus, and does not occur on the dark red distinct from those of C. brunneum, C. poe- latosol of typical cerrado of the Lago Norte cilum, C. blanchetii, and C. caiapoense, but district. In summary, we conclude that the fall within the range of variation of C. bran- type specimen of C. gonzalezii is an indi- donianum. vidual of C. brandonianum with leaves that The slightly smaller values for C. gon- are not fully developed. zalezii, particularly for leaf width, can be A critical analysis of the description of explained by the fact that the measurements C. gonzalezii (Menezes, 1995) reveals a were taken from a dried specimen, in which mixture of characteristics, with the vegeta- the leaves may have shrunk during drying. tive parts corresponding to C. brandonian- The size and number of pseudobulbs indi- um and the reproductive parts to C. brun- cate that the type specimen of C. gonzalezii neum. Menezes (1995) intended to describe is most certainly an adult plant, but with a new species, and, indeed, the pictures of immature leaves. To further examine this ¯owers in the original description are from possibility, the length and width of imma- C. brunneum. However, according to the In- ture leaves of C. brunneum and C. bran- ternational Code of Botanical Nomencla- donianum were plotted with the data for the ture (Greuter et al., 2000), the nomencla- holotype of C. gonzalezii (Fig. 3). The leaf tural type is that element to which the name measurements for the holotype of C. gon- of a taxon is permanently attached, whether zalezii fell within the range of measure- as a correct name or as a synonym. The ments for C. brandonianum. In agreement holotype of C. gonzalezii is C. brandonian- with these results, the collection date for the um, thus C. gonzalezii is a synonym of C. holotype of C. gonzalezii as indicated on brandonianum. the holotype label is early November. Most Cyrtopodium brunneum inhabits the 2004] BATISTA & BIANCHETTI: CYRTOPODIUM (ORCHIDACEAE) 267

FIG. 3. Plot of leaf length vs. width for mature, fully developed leaves and young, incompletely developed leaves of C. brunneum, C. brandonianum and the holotype of C. gonzalezii. Measures of the holotype of C. gonzalezii are the same and repeated for both mature and young leaves. campo limpo and campo sujo vegetation, on This soil is moist for brief periods during deep, reddish, clay soil (dark red latosol), the rainy season, but never retains water for exposed to full sun, being protected only by long periods and dries completely during the surrounding herbaceous vegetation. the dry season. Cyrtopodium poecilum 268BRITTONIA [VOL. 56

Rchb. f. & Warm., C. brandonianum and tend to fall earlier than those of C. Barb.Rodr., C. triste Rchb. f. & Warm., C. triste.InCyrtopodium triste the leaves are blanchetii Rchb. f., and C. caiapoense L. slightly more rigid and coriaceous, and usu- C. Menezes all occur in the same habitat ally crack when bent; the base ¯atter than and are frequently found with C. brunneum. in C. brunneum; and change little in color Less often, C. brunneum is found on rocky, and turn from green to brownish during se- shallow, sandy clay soil, where it grows nescence. However, many of these charac- with C. vernum Rchb. f. & Warm and C. teristics are discrete and depend upon the cristatum Lindl. Cyrtopodium brunneum age and development of the plants. As a also occurs in the typical cerrado, under result, separating the two species based partial shade. In all of these habitats, the only on their vegetative parts is dif®cult. pseudobulbs of C. brunneum are complete- Development of the leaves of C. brun- ly buried in the ground, sometimes with neum is dependent upon habitat conditions. only the upper part exposed. Plants growing under shade in typical cer- Flowering in Cyrtopodium brunneum ex- rado on deep clay soil usually show maxi- tends from the end of the dry season to the mum leaf development in length and width. beginning of the rainy season, from late Plants growing on rocky slopes, on sandy, August to November, but it ¯owers mainly shallow, poor soil, in almost full sun, are during September and October. This ¯ow- much smaller, and resemble C. triste in leaf ering coincides with that of C. poecilum, C. length, but not in width (small specimens blanchetii, C. caiapoense, C. cristatum, and of C. triste have very narrow leaves). The C. vernum; C. triste usually ¯owers a little development of the leaves at anthesis is in- later (late October to early December), as cipient and they only become fully devel- does C. brandonianum (mainly from De- oped 1±2 months after ¯owering. As with cember to January). As with other terrestri- most other Cyrtopodium species of central al Cyrtopodium, ¯owering is enhanced by Brazil, growth of C. brunneum is during the ®re, and among the species with small, bur- rainy season and dormancy is during the ied pseudobulbs C. brunneum is usually the dry season, when the leaves are lost. Plants ®rst to ¯ower after a ®re. The in¯orescence very frequently appear in groups, with sev- appears a few days after a ®re and the eral specimens (from 3 to more than 15) plants are in full bloom 2±3 weeks later. growing close to each other. Cyrtopodium Flowering of the species in unburned places brunneum and C. triste are also very similar is very rare or infrequent; we have never in terms of ¯ower color. Both species have seen plants ¯owering at unburned sites. dark brownish sepals and petals and a yel- Cultivated plants can ¯ower without ®re, lowish red lip. However, C. triste frequently very infrequently, and only when exposed has a branched in¯orescence with up to to hydric stress and full sun. The ¯owers three lateral branches, larger ¯owers, and a have a slightly sweet scent. Removal of the lip with broadly obovate lateral lobes. pollinarium occurs somewhat frequently, Cyrtopodium brunneum is also somewhat indicating that deposition on the stigma similar in overall aspect to C. dusenii Schltr. may be a more limiting factor than lack of However, the leaves of C. dusenii are al- pollinator visits. Fruit set is low in nature. ready well developed at anthesis, there is The species most closely related to Cyr- usually a branched in¯orescence with topodium brunneum is C. triste. In their (0±)1±2(±3) lateral branches, and the small- overall vegetative aspects, C. brunneum and er ¯owers are completely yellow with C. triste are very similar and dif®cult to brown spots. Cyrtopodium brunneum also separate. The main vegetative differences has been confused with C. parvi¯orum, between the two species are presented in which has a similar ¯ower color and falcate Table I. In addition, the leaves of C. brun- lateral lobes of the lip. However, C. parvi- neum are slightly more slender, ¯exible, ¯orum has larger and exposed pseudobulbs less coriaceous and rarely crack when bent; (7±20 cm long), nonarticulate leaves, a lon- have the base laterally compressed; usually ger and frequently branched in¯orescence turn light green and yellow before drying, (0.6±1.2 m long and with (0±)2±4 lateral 2004] BATISTA & BIANCHETTI: CYRTOPODIUM (ORCHIDACEAE) 269 branches), usually larger ¯owers, and lon- C. Mota & P. L. Viana 1262 (HOLOTYPE: ger lateral lobes of the lip (8±10 mm long). CEN; ISOTYPES: AMES, K, RB, SP). Cyrtopodium parvi¯orum occurs in wet places and, when found in drier places, is Cyrtopodio cristato Lindl. similis sed bracteis ¯or- alibus longioribus et de¯exis, petalis et sepalis brun- usually from sandy soils in campo rupestre neo-purpureis, labelli lilacino-purpureo cum callo la- vegetation; it is never found in red clay lat- mellato marginibus irregulariter dentatis praedito dif- osol in typical cerrado vegetation. Cyrto- fert. podium brunneum has a relatively constant ¯ower color, especially when compared to Terrestrial herb. Roots many, whitish, other species such as C. braemii L. C. Me- glabrous. Pseudobulbs exposed or occa- nezes and C. pallidum Rchb. f. & Warm., sionally partially buried, ovoid, acuminate, which are highly variable in color. How- lea¯ess from the second year onwards, ex- ever, C. brunneum is extremely variable in ternally greenish, 5±11 ϫ 1.4±2.4 cm. terms of lip morphology (Fig. 1E: a±e), par- Leaves at ¯owering 7±8, incompletely de- ticularly the mid-lobe, and is probably one veloped but already apparent, 11±23 ϫ 0.6± of the most variable species in the genus 1.4 cm, when fully developed 9±10, erect, for this character. coriaceous, linear-lanceolate, the lowermost Cyrtopodium brunneum is not an uncom- 1±2 leaves sheath-like, the uppermost 8±9 mon species, but the buried pseudobulbs leaves 16±35 ϫ 1±1.8 cm, articulate 4.8± and small size of the plants, which grow 6.2 cm from the leaf base, the apex acu- almost completely obscured by grasses, can minate. In¯orescence lateral, erect, usually make ®nding plants in the ®eld a dif®cult simple, occasionally with a lateral branch task. Plants are most easily located when up to 13 cm long, lax, 19±35 cm, greenish they are in ¯ower in vegetation recently to brownish green; peduncle 11±20 cm, the burned. The species is found in the states 1±2 sheath-like bracts loosely adpressed, of GoiaÂs, Mato Grosso, Minas Gerais, and 1.5±3 cm long, frequently also with 1±2 the Distrito Federal, in the central-western, pair of opposite bracts with abortive ¯ow- central and southeastern Brazilian cerrado. ers; rachis 9±18 cm long; ¯oral bracts de- The species may also be expected to occur ¯exed, parallel and partially covering the in the cerrado region of Tocantins and west- rachis, broad, ovate to ovate-lanceolate, oc- ern Bahia. casionally opposite, 2±2.7 ϫ 1.4±2 cm, brownish green, the apex acute, discreetly CYRTOPODIUM BRANDONIANUM Barb.Rodr., apiculate, the margins undulate; ovary with Gen. Sp. Orchid. 1: 132. 1877. TYPE: pedicel 1.9±4.7 cm long, brownish green. BRAZIL. Minas Geraes, Capivary, Bar- Flowers 10±15 per in¯orescence, showy, bosa Rodrigues s.n. (No original material discreetly sweet-scented. Sepals re¯exed in is known. Typi®ed by Cribb & Toscano fully opened ¯owers, broadly lanceolate to de Brito, 1996: 1: 30. Original illustra- ovate or elliptical, 15±20 ϫ 9±14 mm, dark tion by Barbosa Rodrigues, reproduced brownish-purple, the apex discreetly apic- in Sprunger, 1996: 1: 332). ulate, the margin slightly undulate; lateral sepals slightly oblique. Petals usually re- Cyrtopodium gonzalezii L. C. Menezes, Bol. CAOB ¯exed in fully opened ¯owers, slightly con- 6(1): 9. 1995. TYPE: BRAZIL. Distrito Federal, penõÂnsula do Lago Norte, 9 Nov 1994, L. C. Me- cave, broadly obovate-oblong, 14±16 ϫ nezes UB-54 (HOLOTYPE: UB). 11±14 mm, brownish purple, base lighter, the apex rounded, slightly apiculate, the Cyrtopodium lamellaticallosum J. A. N. base attenuate, the margins entire, smooth. Bat. & Bianchetti, sp. nov. (Figs. 4 & 5) Lip 3-lobed, 10±12 mm long, 13±16 mm wide between the apices of the side lobes TYPE: BRAZIL. Minas Gerais, MunicõÂpio when spread, the base unguiculate, ca. 2 de Moeda, Serra da Moeda, nos morros a mm long, yellow; lateral lobes erect, not margem da estrada vicinal que liga a BR- parallel, turned ca. 45Њ forward, reniform, 040 a Moeda, ca. 1400 m, 24 Oct 2001, J. (4±)5.5 ϫ 6±7(±8) mm, the base and center A. N. Batista, L. B. Bianchetti, A. Salino, R. lilac, towards the borders pinkish, the base 270BRITTONIA [VOL. 56

FIG.4. Cyrtopodium lamellaticallosum. A. In¯orescence. B. Flower, side view. C. Flower, frontal view. D. Callus. E. Perianth. a±b. Variations in lateral sepals and lip morphology, from different individuals of the same population. F. Habit with almost fully developed leaves about 1±2 months after ¯owering. The apex of a one year old pseudobulb showing the remaining sheet of the leaves and the articulation pattern is enlarged. Drawn from Batista et al. 1262 by Simone C. Souza e Silva. 2004] BATISTA & BIANCHETTI: CYRTOPODIUM (ORCHIDACEAE) 271

FIG.5. Cyrtopodium lamellaticallosum. A. In¯orescence. Note the large, de¯exed, ¯oral bracts partially covering the rachis and the re¯exed ¯ower segments. B. Pseudobulbs and young, incompletely developed leaves at the time of ¯owering (October). C. Flower, front view. D. Callus, lateral view. Note the rounded lamellae with irregularly dentate margins. Scale bars ϭ 5 cm (A, B), 5 mm (C), 3 mm (D). 272BRITTONIA [VOL. 56 constricted, the margins usually entire to months later. Non¯owering specimens usu- occasionally slightly serrate; callus promi- ally have larger leaves which reach full de- nent, formed by (4±)6(±8) parallel lamellae, velopment earlier. Plants grow during the longitudinally extending from the base of rainy season and are dormant during the dry the central lobe to the unguicule, the central season, when they loose their leaves. Cyr- 2(±4) lamellae rounded, thicker, yellow topodium lamellaticallosum is sympatric with the margins entire or slightly dentate, with C. poecilum Rchb. f. & Warm., C. eu- the outermost 2(±4) larger, slender, slightly genii Rchb. f., and C. pallidum Rchb. f. & spread, discreetly to profusely irregularly Warm. Flowering of C. lamellaticallosum dentate, dark lilac; isthmus separating the and C. poecilum is concurrent, while C. eu- lateral lobes from the central lobe discreet, genii ¯owers earlier, from June to Septem- ca. 1 mm long; central lobe, reniform to ber, and C. pallidum ¯owers later, from late lunate and then with the lateral edges au- November to early January. riculate, 4±5 ϫ 9±11 mm, the base and cen- Flower color is variable and changes tral part dark lilac, apex slightly orange, lat- with age. The lilac color turns to faint or- eral edges yellowish, the lilac becoming ange and the lip ends up with a yellowish faint orange with aging, the apex rounded orange color in older ¯owers. Some color or very discreetly retuse when ¯attened, the variants with a completely lilac lip have margin re¯exed, slightly verrucose to com- also been observed. As in most other spe- pletely smooth. Column erect, slightly ar- cies in the genus, the column foot has a cuate, trigonous, 9±10 mm long, base yel- conspicuous dark spot at the base near the lowish white, middle lilac, towards the apex claw. The function of this spot is unknown; purple greenish; column foot 2±3(±4) mm it might facilitate pollination. long, at the very end immediately before Cyrtopodium lamellaticallosum is a very the unguicule with a conspicuous dark distinct species; it is not clear which species brown stripe. Anther 3 ϫ 2.5 mm, yellow- is most closely related to it. In overall as- ish, apex green; pollinia 2, waxy, sulcate, pect, with the small, exposed pseudobulbs ca. 1.3±1.5 ϫ 0.8±1 mm, yellow, attached and the usually simple in¯orescence, C. la- to a triangular stipe. Fruit not known. mellaticallosum (particularly dried speci- Etymology. The epithet refers to the la- mens), somewhat resembles C. cristatum mellate callus with irregular dentate mar- Lindl. However, C. lamellaticallosum can gins, a very distinctive and unique charac- easily be distinguished by the usually teristic in the genus. slightly larger ¯owers, the larger and de- ¯exed ¯oral bracts, the brownish purple se- Additional specimens examined. BRAZIL. Minas Gerais: Ouro Preto, TreÃs Moinhos, 10 Oct 1977 (¯), pals and petals, and the lamellate callus Badini s.n. (OUPR); Mun. Moeda, Serra da Moeda, with irregularly dentate margins. Lamellate Nov 1988 (¯), Martens s.n. (BHCB), near the rd. con- calli are uncommon in the genus and only necting Moeda to BR-040, 18 Oct 1997 (¯), Salino C. blanchetii Rchb. f. and C. hatschbachii 3610b (BHCB) (p.p., with C. poecilum var. roseum). Pabst, share this characteristic. In C. blan- Cyrtopodium lamellaticallosum was chetii, the overall aspect of the callus, par- found in the campo rupestre, on dark, ticularly of the central lamella, is similar to sandy-clay soil. This soil can retain water that of C. lamellaticallosum;inC. blanche- for a few days or weeks during the rainy tii there are always four lamellae, but are season, November to March, but dries out frequently less prominent, more elongated, completely during the dry season, May to and less rounded, and have margins that are September. Like most other Cyrtopodium entire or slightly irregular. In C. hatschba- species of central and southeastern Brazil, chii, there are just two lamellae, which are C. lamellaticallosum ¯owers during Octo- larger, farther apart, and rounded, with ber and November, the beginning of the smooth, sometimes slightly undulate mar- rainy season. Similar to most other Cyrto- gins. Beyond these differences in the callus, podium species, a new vegetative shoot is Cyrtopodium blanchetti is usually from typ- formed at the time of ¯owering, but leaves ical cerrado and has completely buried only become fully developed about two pseudobulbs; petals and sepals spotted 2004] BATISTA & BIANCHETTI: CYRTOPODIUM (ORCHIDACEAE) 273 brown over a greenish background; lateral Distrito Federal have a discreet but unpleas- lobes of the lip spathulate, brownish; and ant smell somewhat resembling rancid but- the midlobe obovate-subrhombiform, yel- ter, C. poecilum var. roseum has a slightly low. Cyrtopodium hatschbachii is from per- sweet scent. This difference in ¯ower scent manently wet areas; has larger pseudobulbs, may attract different pollinators. The polli- 8±21 ϫ 2.5±5 cm; the sepals and petals nators and pollination mechanisms of C. completely pink; and the lateral lobes of the poecilum are unknown. lip elliptical and subfalcate. The holotype of C. poecilum var. poecil- Cyrtopodium lamellaticallosum is a rare um is from Lagoa Santa, central Minas Ger- species. It is known from just two localities ais, which is closer to the Serra da Moeda at the southernmost Serra do EspinhacËo in than to the Distrito Federal, where we have central Minas Gerais state. This species collected more intensively and where we might be expected to occur at other sites or have found most of the material we have on other mountains in the EspinhacËo range. examined in situ. However, examination of the original description (Reichenbach, Cyrtopodium poecilum Rchb. f. & Warm. 1881) and original illustration (Warming, var. roseum J. A. N. Bat. & Bianchetti, 1884) of C. poecilum var. poecilum re- var. nov. vealed that the color pattern of the type ma- terial matches the specimens from the Dis- TYPE. BRAZIL. Minas Gerais, MunicõÂpio trito Federal. The specimens from the Serra de Moeda, Serra da Moeda, nos morros a da Moeda with a distinct color pattern are margem da estrada vicinal que liga a BR- considered here as a color variant. In the 040 a Moeda, ca. 1400 m, 24 Oct 2001, J. Distrito Federal, C. poecilum occurs pref- A. N. Batista, L. B. Bianchetti, A. Salino, R. erentially on deep red clay latosol in typical C. Mota & P. L. Viana 1261 (HOLOTYPE: cerrado or campo sujo and campo limpo CEN; ISOTYPE: BHCB). vegetation. In the Serra da Moeda, the va- A varietate typica ¯oribus fragrantibus et labelli lob- riety was found growing on dark, sandy is lateralibus roseis et marginibus lobi centralis roseis clay soil in campo rupestre vegetation, differe. where it grew close to C. lamellaticallosum Etymology. The epithet refers to the and C. pallidum. Both varieties of C. poe- pink margins of the central lobe and pink cilum ¯ower mainly in October and ¯ow- lateral lobes of the lip. ering is enhanced by ®re. A visit to the Ser- ra da Moeda in one year revealed no spec- Additional specimen examined. BRAZIL. Minas imens in ¯ower, whereas in the next year, Gerais: Mun. of Moeda, Serra da Moeda, near the rd. after the area had burned, a great number connecting Moeda to BR-040, 18 Oct 1997 (¯), Salino of ¯owering specimens were observed. 3610b (BHCB) (p.p., with C. lamellaticallosum). In general, color variants are better treat- Cyrtopodium poecilum var. roseum is ed as forms rather than as varieties. How- morphologically identical to C. poecilum ever, for C. poecilum var. roseum, the color var. poecilum in vegetative and most ¯oral variation seems to be clearly ®xed in the characters. Both have mature pseudobulbs population, as most of the individuals had that are small, externally reddish purple, this character. Within this color variant, and completely buried; large and broad some differences in shade have been ob- leaves; a simple to 3-branched in¯ores- served, e.g., a more or less intense shade of cence; and sepals and petals profusely spot- pink, particularly in relation to the margins ted brownish purple over a greenish back- of the central lobe, or a fading of the pink ground. The principal difference is the color in older ¯owers, rendering the lip somewhat of the lateral lobes and margins of the cen- dull and discolored. tral lobe of the labellum, which are pink in the new variety and brownish, reddish Acknowledgments brown or reddish purple in C. poecilum var. poecilum. All specimens of C. poecilum The authors thank Alexandre Salino for var. poecilum that we have examined in the helping locate C. lamellaticallosum in the 274BRITTONIA [VOL. 56

®eld and Julio Lombardi for the loan of ma- Iconographie des orchideÂesduBreÂsil/JoaÄo Barbosa terial from BHCB. We also thank the cu- Rodriguez. Vol 1. Illustrations. Friedrich Reinhardt Verlag, Basle, Switzerland rators of HB, HRCB, HUFU, IBGE, MBM, Greuter, W., J. McNeill, F. R. Barrie, H. M. Burdet, OUPR, R, SP, SPF, UB and UEC for loans V. Demoulin, T. S. Filgueiras, D. H. Nicolson, P. or access to their collections; Simone C. C. Silva, J. E. Skog, P. Trehane, N. J. Turland Souza e Silva for preparing the illustrations; & D. L. Hawksworth, editors. 2000. International Code of Botanical Nomenclature (Saint Louis Tarciso Filgueiras for the Latin diagnoses; Code). Regnum Vegetabile. 138. Koeltz Scienti®c and Stephen Hyslop for reviewing the En- Books, KoÈnigstein. glish. Gustavo Romero and FaÂbio de Barros Menezes, L. C. 1995. Novas orquõÂdeas brasileiras read earlier versions of the manuscript and (New Brazilian orchids). Bol. CAOB 6: 8±13. provided several helpful suggestions. ÐÐÐ. 2000. Genus Cyrtopodium: espeÂcies brasileir- as. Ed. IBAMA, BrasõÂlia. Reichenbach, H. G. 1881. VIIÐNovitiae Orchidaceae Literature Cited Warmingianae. Cyrtopodium. Otia Botanica Ham- burgensia 2(1): 58±60. Batista, J. A. N. & L. B. Bianchetti. 2001. Cyrto- Sprunger, S., editor. 1996. Iconographie des orchideÂes podium linearifolium (Orchidaceae): a new species du BreÂsil/JoaÄo Barbosa Rodriguez. Vol 1. Illustra- from central Brazil. Lindleyana 16: 226±230. tions. Friedrich Reinhardt Verlag, Basle, Switzer- Bianchetti, L. B. & J. A. N. Batista. 2000. Cyrto- land. podium latifolium (Orchidaceae): a new species Warming, E. 1884. Symbolae ad ¯oram Brasiliae cen- from central Brazil. Lindleyana 15: 222±226. tralis cognoscendam, Part 30. Orchideae 2. Viden- Cribb, P. & A. Toscano de Brito. 1996. Introduction skabelige Meddelelser fra den naturhistoriske For- and history. Pages 23±32. In: S. Sprunger, editors. ening i kjoÈbenhavn 5±8: 86±99.