Fig. 1

Cyrtopodium brunneum A. Habit at anthesis. B. Frontal view of flower. C. Side view of flower. 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.

ABSTRACT. Cyrtopodium brunneum, C. lamellaticallosum and C. poecilum var. roseum, two new species and a new variety from the cerrado and campo rupestrian vegetation of central and southeastern Brazil are described and illustrated. Although the description of Cyrtopodium gonzalezii indicates that it is a distinct species, the holotype is referable to C. brandonianum. The undescribed plants are now described here as Cyrtopodium brunneum and C. gonzalezii is placed in the synonym of C. brandonianum.

Key words: Orchidaceae, Cyrtopodium, cerrado vegetation, Brazil.

Cyrtopodium is a Neotropical genus with about 42 species ranging from southern Florida to northern Argentina. The center of diversity of the genus is in the Brazilian cerrado, where at least 25 species occur. In recent years, several new species have been described in the genus (Bianchetti & Batista, 2000; Batista & Bianchetti, 2001; Menezes, 2000), most from central Brazil. Based on our further exploration of the region and examination of herbarium specimens of the genus, we describe bellow two new species C. brunneum and Cyrtopodium lamellaticallosum and and a new variety, C. poecilum var. roseum.
Traditionally, the taxonomy of Cyrtopodium has been based almost completely on reproductive characters, with the vegetative parts, particularly the leaves, being neglected. This is because botanical studies of the genus have relied almost entirely on dried specimens collected mainly during the flowering season, when most Cyrtopodium species do not have fully developed leaves. The examination of live plants throughout the developmental cycle has allowed characterization of the vegetative parts of most species and, in many cases, the species can be identified vegetatively. Based on this characterization, we found that the holotype of C. gonzalezii L. C. Menezes is unambiguously C. brandonianum Barb. Rodr., although the description does indicate that it is a new species. This new species is described here as C. brunneum.

Type: BRAZIL. Distrito Federal, Brasília, Plano Piloto, final da Asa Norte, area between the "Parque Ecológico Norte" and "Parque Nacional de Brasília", the futur "Setor Noroeste", 8 Sep 2001 (fl), J. A. N. Batista 1242 (HOLOTYPE: CEN; ISOTYPES: AMES, BHCB, HB, ESA, HUEFS, HUFU, K, MBM, MO, NY, RB, SP, SPF, UB, UEC).

Cyrtopodium gonzalezii L.C.Menezes in Boletim CAOB 6(1): 9. 1995. pro parte, excluding type.

Cyrtopodio tristi Rchb. f. & Warm. similis sed inflorescentia semper simplici, floribus minoribus, sepalis oblongo-lanceolatis vel paulo lanceolato-ovatis, labelli lobis lateralibus falcatis differt; etiam C. dusenii Schltr. similis sed foliis sub anthesi redactis, inflorescentia semper simplici, floribus majoribus, sepalis et petalis atrobrunneis differt.

Terrestrial herb. Roots many, whitish, glabrous. Pseudobulbs completely buried, ovoid, leafless from the second year onwards, externally white, (3-)4-6(-7) x (1-)1.5-2.5 cm. Leaves at flowering 3-6, little developed, 7-15 cm long, when fully developed 5-8, spreading, coriaceous, lanceolate, the lowermost 1-2 sheath-like, the uppermost 4-7 13-33 x 1-2.6 cm, articulate 1-2 cm from the surface of the soil, the apex acuminate. Inflorescence lateral, erect, racemose, lax, 26-56 cm, brownish to dark brown; peduncle 11-26 cm, the 2 sheathlike bracts strongly adpressed, 1.7-3.5 cm long; rachis 12-26 cm long; floral bracts oblong-lanceolate, 1-3 x 0.5-0.9 cm, greenish brown, acute to acuminate, the margins undulate, incurved; ovary with pedicel 1.4-3.9 cm long, dark brown. Flowers 9-15(-20) per inflorescence, sweet-scented. Sepals oblong-lanceolate to slightly lanceolate-ovate, light to dark brownish or brownish green, lighter at the base, frequently with dark-brown spots, the apex apiculate, the margin undulate; dorsal sepal (8-)11-13(-15) x 4-5(-6) mm; lateral sepals slightly oblique, (8-)11-13(-15) x 4-6(-7) mm. Petals concave, broadly elliptic to somewhat broadly lanceolate, (8-)9-12(-13) x 6-8 mm, the lower half greenish to light-brownish, occasionally with few to many small brown dots, the upper half dark-brown, color transition usually sharply delimited, the apex obtuse to rounded, slightly apiculate, the margins slightly undulate. Lip 3-lobed, (7-)9-11(-12) mm long, 10-12(-14) mm wide between the apices of the side lobes when spread, the base shortly unguiculate, 1 mm long, yellowish; lateral lobes erect, parallel, oblong-falcate, (3-)4-5(-6) x 2-3(-4) mm, reddish to reddish-purple, the apex obtuse, the base unconstricted, the margins entire, smooth; callus verrucose, slightly sulcate, extending from the base of the central lobe to the column foot, white, occasionally reddish-white; isthmus separating the lateral lobes from the central lobe usually prominent, sometimes elongated, 1-2(-3) mm long; central lobe variable, obcordate to obovate-oblong to somewhat obreniform, 5-7 x (5-)6-8(-10) mm, yellow with red to reddish-purple margins, occasionally with a few red dots at the base, the yellow becoming fainter towards aging, the apex with a protruding fold, retuse when flattened, the margin smooth, occasionally slightly verrucose at the apex. Column erect, slightly arcuate, trigonous, 6-7(-8) mm long, base yellow, middle reddish white and greenish purple towards the apex; column foot (2.5-)3-4 mm long, yellow at the base, white at the middle with small numerous red dots that merge towards the apex. Anther ca. 2-2.5 x 1.5 mm, yellowish, apex green; pollinia 2, waxy, sulcate, ca. 1 x 0.5-0.6 mm, yellow; stipe triangular, hyaline, ca. 1 x 1 mm at base. Fruit deflexed, green, oblong to elongate, sulcate, ca. 6-8.5 x 1-1.5 cm when young, including the pedicel.

Etymology. - Named after the dark brown color of the sepals and petals.

Additional specimens examined. BRAZIL. Distrito Federal: Brasília, Vila Maury, 6 Sep 1960 (fl), Andrade 412 & Emmerich 404 (HB, R); Brasília, Lago Norte, 8 Oct 1990 (fl), Batista 122 (CEN), 1 Oct 1991 (fl), Batista 208 (CEN), 7 Sep 1992 (fl), Batista 325 (CEN); Guará, Reserva Ecológica do Guará, 13 Sep 1992 (fl), Batista 330 (CEN); Brasília, area between Parque Ecológico Norte and Parque Nacional de Brasília, 7 Oct 1994 (fl), Batista 421 (CEN), 11 Dec 1994 (veg), Batista 434 (CEN); Brasília, Lago Norte, 17 Feb. 1995 (veg), Batista 521 (CEN), QL 15, near Clube do Congresso, 17 Feb. 1995 (veg), Batista 523 (CEN); Núcleo Bandeirante, Santuário Ecológico do Riacho Fundo, 14 Sep 1995 (fl), Batista & Bianchetti 576 (CEN); Brasília, area between Parque Ecológico Norte and Parque Nacional de Brasília, 28 Sep 1998 (fl), Batista 792 (CEN, HB, MBM, SP), 30 Dec 2000 (veg), Batista 1112 (CEN); Brasília, area between Lago Norte and EPIA, 6 Jan. 2001 (veg), Batista 1126 (CEN); Brasília, Lago Sul, condomínio Prive Morada Sul II, above QI-28, close to Paranoá dam, (fl. in cult.) 22 Sep 2001, Batista et al. 1184 (CEN); Gama, BR-060, near the GDF check point and the access to Santo Antônio do Descoberto, 2 Oct 2001 (fl), Batista & Oliveira-Neto 1252 (CEN); Brasília, Plano Piloto, Asa Norte, 713-714 N, area of University of Brasília, between Parque Olhos D´Água and the CAESB sewage treatment plant, 7 Oct 2001 (fl), Batista 1254 (CEN); Brasília, Setor de Mansões do Lago Norte, 17 Sep 1990 (fl), Bianchetti & Batista 953 (CEN), trecho 2, 20 Oct 1991 (fl), Bianchetti & Batista 1176 (CEN), near MI-7, 6 Oct 1991 (fl), Bianchetti & Batista 1168 (CEN); Guará, Reserva Ecológica do Guará, 26 Aug 1990 (fl), Bianchetti & Batista 950 (CEN); Brasília, Plano Piloto, 10 Nov 1961 (fl), Heringer 8756 (HB, UB); Brasília, Estação Florestal Cabeça do Veado, 8 Oct 1965 (fl), Heringer 10616 (HB, UB); Santa Maria, near the microwave towers, 28 Oct 1972 (fl), Heringer 12206 (HB, UB); ca. 20 km S of Brasília, on rd. to Goiânia, near Rio Melchior, 1125 m, 25 Sep 1965 (fl), Irwin et al. 8652 (UB); Brasília, confluence of Rio Torto with Lake Paranoá, 975 m, 9 Oct 1965 (fl), Irwin et al. 9086 (HB); Guará, Reserva Ecológica do Guará, 15o50’S, 47o57’W, 1050 m, 20 Sep 1994 (fl), Oliveira 2 (UB). Goiás: Chapada dos Veadeiros, ca. 26 km N from Alto Paraíso, GO-118, 11 Oct 1999 (fl), Batista 945 (CEN); ca. 8 km from Niquelândia, Tocantins Niquel Company, 14o23’48”S, 48o25’59”W, 17 Sep 1996 (fl), Fonseca et al. 1154 (IBGE); Municipality of Mineiros, BR-364, near Córrego Alegre, 20 Sep 1974 (fl), Hatschbach & Kummrow 35014 (MBM); Goiânia, 30 Nov 1963 (fl), Heringer 9289 (HB); Municipality of Luziânia, 16 Sep 1974 (fl), Heringer 13971 (HB, UB); 12 km NW of Veadeiros, on rd. to Cavalcante, 1200 m, 21 Oct 1965 (fl), Irwin et al. 9444 (HB, UB); Aparecida de Goiânia/Hidrolândia, 10 Aug 2002 (fl), Pastore 32 (CEN). Mato Grosso: 84 km from Alto Araguaia, towards Rondonópolis, BR-364, Sep-Nov 1983 (fl), Hutchison 8550 (UEC). Minas Gerais: Uberlândia, Estação Ecológica do Panga, 24 Sep 1992 (fl), Araujo et al. 276 (HUFU); Morro Pelado, near Campanha, Aug 1896 (fl), Brandão in CGGMG 1719 (R) (p.p., with C. cristatum); Municipality of Carmo do Rio Claro, Serra da Tormenta, 3 Nov 1990 (fl), Campos s.n. (CEN, HRCB); Serra do Cipó, 23 Aug 1958 (fl), Heringer 6438 (HB) (p.p., with C. parviflorum); Felixlândia, Três Marias basin, 23 Aug 1958 (fl), Heringer 6438 (UB), 2 Oct 1959 (fl), Heringer 6438 (UB), margins of Rio Paraopeba, 11 Oct 1959 (fl), Heringer 6438 (UB); Santana do Riacho, km 107 on the Belo Horizonte to Conceição do Mato Dentro rd., 4 Oct 1981 (fl), Pirani et al. in CFSC 7472 (SP, SPF).

Pabst was probably the first orchid taxonomist to examine material of C. brunneum, but misidentified it first as C. falcilobum Hoehne & Schltr. (now a synonym of C. parviflorum Lindl.) and latter as C. poecilum Rchb. f.& Warm. f. minor Hoehne (now a synonym of C. fowliei L.C.Menezes). He was apparently also the first taxonomist to suspect that the specimens might represent a new species. In his personal files at the Herbarium Bradeanum, there is a card with a sketch of C. brunneum indicated at the side as C. aff. parviflorum and at the top as Cyrtopodium heringeri; but he never published a valid description of this species. More recently, Menezes described this species as Cyrtopodium gonzalezii L.C.Menezes (Menezes, 1995), but unfortunately designated a holotype that is Cyrtopodium brandonianum Barb. Rodr. This situation leaves the new species without valid publication.
The holotype of C. gonzalezii, at the University of Brasília herbarium is a single, sterile specimen. However, on the basis of its vegetative characteristics this specimen is C. brandonianum. The holotype has small, narrow, buried pseudobulbs and linear, erect, very narrow leaves. Compared with the 25 Cyrtopodium species known from the cerrado, the buried pseudobulb characteristic excludes several species with exposed pseudobulbs, such C. aliciae Linden & Rolfe, C. cardiochilum Lindl., C. cipoense L.C.Menezes, C. cristatum Lindl., Cyrtopodium eugenii Rchb. f., C. hatschbachii Pabst, C. lissochiloides Hoehne & Schltr., C. palmifrons Rchb. f.& Warm., C. paludicolum Hoehne, C. parviflorum Lindl., C. saintlegerianum Rchb. f., C. virescens Rchb. f.& Warm., and C. vernum Rchb. f.& Warm.
According to the protologue of C. gonzalezii (Menezes, 1995) and to Menezes herself (L. C. Menezes, pers. comm.), the holotype was collected on the peninsula do Lago Norte, a district of Brasília where we have collected intensively during the past 12 years. Six species of Cyrtopodium are known from this district: Cyrtopodium blanchetii Rchb. f., C. brandonianum Barb. Rodr., C. brunneum, C. caiapoense L.C.Menezes, Cyrtopodium poecilum Rchb. f.& Warm. and C. virescens Rchb. f.& Warm. This further excludes from the comparison those species that do not occur in the Distrito Federal, such as C. braemii L.C. Menezes and Cyrtopodium dusenii Schltr., or those that occur in Brasília but not at this particular site, such as C. fowliei L.C.Menezes, C. latifolium Bianchetti & J.A.N.Bat., C. pallidum Rchb. f.& Warm., C. triste Rchb. f.& Warm. and C. linearifolium J.A.N.Bat. & Bianchetti. Additionally, the pseudobulbs of C. braemii, C. fowliei and C. latifolium are reddish-purple externally, while in the holotype of C. gonzalezii they are whitish.
The characterization of the vegetative parts of the remaining species for comparison, including the holotype of Cyrtopodium gonzalezii, is shown in Table I. Because of its similarity to C. brunneum, C. triste is also included in the comparison.

All measurements were taken from living specimens under field conditions, except for the measures of pseudobulbs and the holotype of C. gonzalezii, which were obtained from dried specimens. Only mature and adult specimens and fully developed leaves were considered in the analysis. Leaf length was measured from the surface of the soil, which corresponds approximately to the distance from the apex of the pseudobulb to the apex of the leaf.

Fig. 7. Cyrtopodium poecilum. Fully developed (mature) leaves. Plant in campo sujo vegetation.

Cyrtopodium poecilum (Fig. 7) and C. blanchetii (Fig. 8) have much broader leaves than Cyrtopodium gonzalezii.

Fig. 8. Cyrtopodium blanchetii. Plant with mature leaves and fruits.

Fig. 9. Cyrtopodium poecilum. Pseudobulbs.

Fig. 10. Cyrtopodium blanchetii. Pseudobulbs.

Fig. 11. Cyrtopodium caiapoense. Vegetative parts

The pseudobulbs of C. poecilum are reddish purple externally (Fig. 9), and therefore distinct from all the other species in this comparison; the pseudobulbs of C. blanchetii (Fig. 10) are larger than those of C. gonzalezii.
Cyrtopodium caiapoense (Fig. 11) has more and wider leaves and usually has longer pseudobulbs than Cyrtopodium gonzalezii, On the other hand, C. brunneum and C. triste has shorter and broader leaves. In summary, the vegetative parts of neither of the five previous species is compatible with the holotype of Cyrtopodium gonzalezii (Table 1).
Specifically comparing C. brunneum and C. gonzalezii, the mean leaf length-to-width ratio was 9.5-15.5 for Cyrtopodium brunneum, and 55 for C. gonzalezii (Table 1). Additionally, the leaves of C. brunneum are typically spreading (Figs. 4-5), while in C. gonzalezii, they are erect.

Fig. 5a. Cyrtopodium brunneum. Uprooted mature plant (January), showing the fully developed leaves and underground pseudobulbs. Lateral view.

Fig. 4. C. brunneum. Fully developed leaves, in December, about 2 months after flowering. The surrounding vegetation has been removed to expose the plants.

Fig. 5b Cyrtopodium brunneum. Uprooted mature plant (January), showing the fully developed leaves and underground pseudobulbs. Front frontal.

The pseudobulbs of C. brunneum are usually slightly longer and broader than those of C. gonzalezii. Another difference involves the distance of the leaf articulation relative to the apex of the pseudobulb, which was shorter in C. brunneum than in C. gonzalezii.

Fig. 12. Cyrtopodium brandonianum. Plant with fully developed leaves.

Of the species presented in table 1 the only that has characteristics compatible with the holotype of Cyrtopodium gonzalezii is Cyrtopodium brandonianum (Figs. 12-13).

Fig. 13. Cyrtopodium brandonianum. Pseudobulbs.

Fig. 14. 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.

Fig. 2 - Cyrtopodium brunneum. Habitat at the time of flowering. Burnt cerrado at the type locality, about 2 weeks after fire, with the vegetation beginning to regrow, at the beginning of September.

Less often, C. brunneum is found on rocky, shallow, sandy clay soil, where it grows with C. vernum Rchb. f. & Warm and C. cristatum Lindl. Cyrtopodium brunneum also occurs in the typical cerrado, under partial shade. In all of these habitats, the pseudobulbs of C. brunneum are completely buried in the ground, sometimes with only the upper part exposed.
Flowering in Cyrtopodium brunneum extends from the end of the dry season to the beginning of the rainy season, from late August to November, but it flowers mainly during September and October. This flowering coincides with that of C. poecilum, C. blanchetii, C. caiapoense, C. cristatum and C. vernum; C. triste usually flowers a little later (late October to early December), as does C. brandonianum (mainly from December to January).02*

Fig. 3. Cyrtopodium brunneum. Inflorescence. Note the poorly developed leaves at anthesis.

As with other terrestrial Cyrtopodium, flowering is enhanced by fire, and among the species with small, buried pseudobulbs C. brunneum is usually the first to flower after a fire. The inflorescence appears a few days after a fire and the plants are in full bloom 2-3 weeks later (Fig. 3). Flowering of the species in unburned places is very rare or infrequent; we have never seen plants flowering at unburned sites. Cultivated plants can flower without fire, very infrequently, and only when exposed to hydric stress and full sun. The flowers have a slightly sweet scent. Removal of the pollinarium occurs somewhat frequently, indicating that deposition on the stigma may be a more limiting factor than lack of pollinator visits. Fruit set is low in nature. The species most closely related to Cyrtopodium brunneum is C. triste. In their overall vegetative aspects, C. brunneum and C. triste are very similar and difficult to separate.

The main vegetative differences between the two species are presented in table 1. In addition, the leaves of Cyrtopodium brunneum are slightly more slender, flexible, less coriaceous and rarely crack when bent; have the base laterally compressed; usually turn light green and yellow before drying, and tend to fall earlier than those of C. triste. In Cyrtopodium triste the leaves are slightly more rigid and coriaceous, and usually crack when bent; have the base flatter than in C. brunneum; and change little in color and turn from green to brownish during senescence. However, many of these characteristics are discrete and depend upon the age and development of the plants. As a result, separating the two species based only on their vegetative parts is difficult.

Development of the leaves of C. brunneum is dependent upon habitat conditions. Plants growing under shade in typical cerrado on deep clay soil usually show maximum leaf development in length and width. Plants growing on rocky slopes, on sandy, shallow, poor soil, in almost full sun, are much smaller, and resemble C. triste in leaf length, but not in width (small specimens of C. triste have very narrow leaves). The development of the leaves at anthesis is incipient, and they only become fully developed 1-2 months after flowering. As with most other Cyrtopodium species of central Brazil, growth of Cyrtopodium brunneum is during the rainy season and dormancy is during the dry season, when the leaves are lost. Plants very frequently appear in groups, with several specimens (from 3 to more than 15) growing close to each other (Fig. 15). Cyrtopodium brunneum and C. triste are also very similar in terms of flower color. Both species have dark brownish sepals and petals and a yellowish red lip.

Fig. 15. Cyrtopodium brunneum. Group of plants. A common feature of the species.

Fig. 16 - C. triste. flower

However, C. triste frequently has a branched inflorescence with up to three lateral branches, larger flowers, and a lip with broadly obovate lateral lobes (Fig. 16). Cyrtopodium brunneum is also somewhat similar in overall aspect to C. dusenii Schltr. However, the leaves of C. dusenii are already well developed at anthesis, there is usually a branched inflorescence with (0-)1-2(-3) lateral branches, and the smaller flowers are completely yellow with brown spots. Cyrtopodium brunneum also has been confused with C. parviflorum, which has a similar flower color and falcate lateral lobes of the lip (Fig. 17 (Fig. 17).

Fig. 17 - C. parviflorum. Flower

However, C. parviflorum has larger and exposed pseudobulbs (7-20 cm long), nonarticulate leaves, a longer and frequently branched inflorescence (0.6-1.2 m long and with (0-)2-4 lateral branches), usually larger flowers, and longer lateral lobes of the lip (8-10 mm long). Cyrtopodium parviflorum occurs in wet places and, when found in drier places, is usually from sandy soils in campo rupestrian vegetation; it is never found in red clay latosol in typical cerrado vegetation.

Cyrtopodium brunneum has a relatively constant flower color (Figs. 6a-b), especially when compared to other species such as C. braemii L. C. Menezes and C. pallidum Rchb. f. & Warm., which are highly variable in color. However, Group of plants. A common feature of the species. brunneum is extremely variable in terms of lip morphology (Fig. 1E: a-e), particularly the mid-lobe, and is probably one of the most variable species in the genus for this character. Cyrtopodium brunneum is not an uncommon species, but the buried pseudobulbs and small size of the plants, which grow almost completely obscured by grasses, can make finding plants in the field a difficult task.

Fig. 6a - C. brunneum. Flower

Fig. 6b - -C. brunneum. Flower

Plants are most easily located when they are in flower in vegetation recently burned. The species is found in the states of Goiás, Mato Grosso, Minas Gerais, and the Distrito Federal, in the central-western, central and southeastern Brazilian cerrado. The species may also be expected to occur in the cerrado region of Tocantins and western Bahia.

Cyrtopodium gonzalezii L.C.Menezes in Boletim CAOB 6(1): 9. 1995. Typus: BRASIL. Distrito Federal, península do Lago Norte, 9 Nov. 1994, L. C. Menezes UB-54 (HOLOTYPUS: UB).

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