A revision of Habenaria section Macroceratitae (Orchidaceae) in Brazil

Zenilton Gayoso

Habenaria sect.Macroceratitae from Brazil is revised, and seven species are recognized:H. bractescens, H. gourlieana, H. johannensis, H. longicauda, H. macronectar, H. nabucoi, and the newly describedH. paulistana. A main feature of these species is the presence of long, separated, involute stigmatic processes.Habenaria bractescens, H. gourlieana, H. johannensis, andH. macronectar are distributed mainly from central and southeastern Brazil to southern Brazil and southern South America, whereasH. longicauda andH. nabucoi are distributed mainly from west central, southeastern and northeastern Brazil to northern South America.Habenaria paulistana is restricted to the state of São Paulo.Habenaria bradei, H. juergensii, andH. sartoroides are lectotypified, andH. kleyi is neotypified. The identity ofH. fastor is discussed andH. nabucoi is recognized as the valid name for this species. Seven other species previously placed in sect.Macroceratitae are sufficiently distinct and are removed fr...

A revision of Habenaria section Macroceratitae (Orchidaceae) in Brazil JOAO A . N . BATISTA, L U C I A N O DE B E M B I A N C H E T T I , AND Z E N I L T O N DE J. G . M I R A N D A Batista, J. A.N., L. B. Bianchetti (Embrapa Recursos Gen6ticos e Biotecnologia, Parque Estaqao Biol6gica, Final Av. W5 Norte, C.P. 02372, Brasflia, DF, 70770-901, Brazil; e-mail: janb@cenargen.embrapa.br, bianchet@cenargen.embrapa.br) & Z. J. G. Miranda (Embrapa Cerrados, C.P. 08223, Planaltina, DF, 73301-970, Brazil; e-mail: zenilton@cpac.embrapa.br). A revision of Habenaria sect. Macroceratitae (Orchidaceae) in Brazil. Brittonia 58: 10--41. 2006.--Habenaria sect. Macroceratitae from Brazil is revised, and seven species are recognized: H. bractescens, H. gourlieana, H. johannensis, 1"1. longicauda, H. macronectar, H. nabucoi, and the newly described I-I. paulistana_ A main feature of these species is the presence of long~ separated, involute stigmatic processes. Habenaria bractescens, H. gourlieana, ILl. johannensis, and 1t. macronectar are distributed mainly from central and southeastern Brazil to southern Brazil and southern South America, whereas H. longicauda and H. nabucoi are distributed mainly from west central, southeastern and northeastern Brazil to northern South America. Habenaria paulistana is restricted to the state of S~o Paulo. Habenaria bradei, H. juergensii, and H. sartoroides are lectotypified, and H. kleyi is neotypified. The identity of H. fastor is discussed and H. nabucoi is recognized as the valid name for this species. Seven other species previously placed in sect. Macroceratitae are sufficiently distinct and are removed from the section. Key words: Orchidaceae, Habenaria, section Macroceratitae, Brazil. Batista, J. A. N., L.B. Bianchetti (Embrapa Recursos Gen6ticos e Biotecnologia, Parque Esta~ao Biol6gica, Final Av. W5 Norte, C.P. 02372, Brasflia, DF, 70770-901, Brasil; email: janb@cenargen.embrapa.br, bianchet@cenargen_embrapa.br) & Z. J. G. Miranda (Embrapa Cerrados, CP. 08223, Planaltina, DF, 73301-970, Brasil; email: zenilton@cpac.embrapa.br). A revision of Habenaria sect. Macroceratitae (Orchidaceae) in Brazil. Brittonia 58: 10-41. 2006.----E apresentada uma revisao de Habenaria seato Macroceratitae de ocorr~ncia no Brasil, sendo reconhecidas sete espfcies: H. bractescens, H. gourlieana, H. johannensis, 1t. longicauda, H. macronectar, H. nabucoi e uma nova esprcie descrita como I-L laaulistana. Uma das principals caracter~sticas destas esprcies s~o os estigmas separados, longos e corn as bordas involutas. Habenaria bractescens, H. gourlieana, H~ johannensis e H. macronectar distribuem-se principalmente do Centro-Oeste e Sudeste do Brasil ao Sul do Brasil e Sul da Amrrica do Sul, enquanto H. longicauda e H. nabucoi distribuem-se principalmente do Centro-Oeste, Sudeste e Nordeste do Brasil ao Norte da Amrrica do Sul. Habenaria paulistana 6 restrita ao estado de S~o Paulo. Habenaria bradei, H. juergensii, e H. sartoroides s~o lectotipificadas e H. kleyi 6 neotipificada. A identidade de H. fastor 6 discutida e H. nabucoi 6 reconhecido como o nome v~ilido para esta esp~cie. Outras sete esprcies previarnente inclufdas na se~ao Macroceratitae s~o suficientemente distintas e s~o excluidas da seq~o. Brittonia, 58(1), 2006, pp. 10-41. 9 2006, by The New YorkBotanicalGardenPress, Bronx, NY 10458-5126U.S.A. ISSUED: 31 March 2006 2006] BATISTA ET AL.: HABENARIA SECT. MACROCERATITAE (ORCHIDACEAE) Habenaria is the largest genus in subtribe Habenariinae, characterized by convex stigmas, which may be sessile or stalked, and the usually long rostellar arms and long caudicles. The subtribe is most diverse in Africa (Dressier, 1993), but several genera also occur in Asia. In the Americas, the group is represented only by Habenaria. Current counts estimate the number of Habenaria species at about 600 (Pridgeon et al., 2001), which makes it the largest genus of terrestrial Orchidaceae. The genus has a temperate and pantropical distribution; the main centers of diversity are in Brazil, southern and central Africa, and East Asia (Kurzweil & Weber, 1992). Several genera such as Bonatea, Centrostigma, Platycoryne, Roeperocharis, and Kryptostoma, have been segregated from Habenaria, but the generic separation is questionable (Kurzweil & Weber, 1992). The main distinctive features of Habenaria are the often bifid petals that are not fused to the other organs, the lip usually deeply divided and lacking a callus, the distinctly stalked stigmas, and the entire stigma lobes, which are usually free and not adnate to the petals or lip (Pridgeon et al., 2001). The only worldwide revisions of Habenaria were that of Kraenzlin (1892, 1901), in which 32 sections were recognized. Characterization of the sections was based primarily on the degree of dissection of the petals and lip and on column structure, particularly the length of the stigmas. Among the 165 to 170 Habenaria species known from Brazil (Hoehne, 1940; Pabst & Dungs, 1975) there are some very distinct groups of species, one of which is characterized by large flowers, long stigmatic processes, and several other characters. These species were placed by Kraenzlin (1892) and Cogniaux (1893) mainly into section Macroceratitae Kraenzl. As part of an ongoing revision of the Brazilian Habenaria, we now present our results on the systematics, nomenclature and biogeography of the Brazilian species in this section. Taxonomic history of Habenaria sect. Macroceratitae Habenaria sect. Macroceratitae was established by Kraenzlin (1892) in his revision of the genus. He placed 13 species in the sec- 11 tion, of which nine occur in Brazil (Table I). Cogniaux (1893), in his treatment of Habenaria in Flora Brasiliensis followed Kraenzlin's characterization of sect. Macroceratitae and included three more Brazilian species in the section, including Habenaria sartor, which had been placed by Kraenzlin in sect. Sartores Kraenzl. (Table I). Cogniaux (1893) eliminated sect. Sartores and transferred most of its members to sect. Nudae Cogn. Later, Kraenzlin (1901) accepted most of the modifications proposed by Cogniaux (1893), and incorporated H. sartor and one more species from Mexico, H. pringlei Robinson, into sect. Macroceratitae. Using a different approach, starting with the vegetative parts and then advancing towards the details of the flowers, Hoehne (1940) divided the species of Brazilian Habenaria into nine groups. The species in sect. Macroceratitae were placed mainly in the "Macroceratitis" group (Table I). The group comprised 12 species, of which six represented new combinations or taxa described after the publication of Flora Brasiliensis. Although the groups proposed by Hoehne were intended to be artificial subdivisions whose main purpose was for establishing keys to the species, some groups, such as the "Macroceratitis" group, contained species that seem to be phylogenetically related and were thus probably more natural than many of the sections established by Kraenzlin. In the last major survey of Brazilian orchids, Pabst and Dungs (1975) basically followed the divisions established by Hoehne, and used some new characters to distinguish several groups called alliances. Some of the species previously placed in sect. Macroceratitae and most of the species in the "Macroceratitis" group were placed in the H. gourlieana alliance (Table I), which consisted of eight species. The main differences in relation to Hoehne's (1940) Macroceratitis group were the reduction of some of the species to synonyms and the exclusion of H. macroceratitis, which does not occur in Brazil. Recently, Szlachetko (2004) proposed elevating some sections and groups of species of Habenaria to the genus level, and transferred the species concerned. In this context, H. vaupellii was transferred to the new genus Bertauxia, while H. bractescens, H. burkar- 12 BRITTONIA [VOL. 58 TABLEI m COMPARISON OF THE TREATMENTS OF SECT. Classification Characters Plant Flowers Sepals Petals Petal anterior segment Lip Stigmas Rostellum Distribution Species Macroceratitae AMONG DIFFERENT AUTHORS Kraenzlin (1892) Cogniaux (1893) Kraenzlin (1901) Hoehne (1940) Pabst & Dungs (1975) Sect. Sect. Sect. H. gourlieana Macroceratitae Macroceratitae Macroceratitae "Macroceratitis" group -glabrous reflexed, dorsal and laterals of equal length bipartite -- -glabrous reflexed,dorsal and laterals of equal length bipartite narrow and long -glabrous reflexed,dorsal and laterals of equal length bipartite -- bipartite -- tripartite long non-cucullate neotropical tripartite long non-cucullate -- tripartite long non-cucullate neotropical tripartite ---- H. H. H. H. H. H. H. H. H. H. H. H. H. H. araneiflora H. bractescens H. cryptophila H. gourlieana 1-1. helodes H. hydrophila H. Iongicauda H. macroceratitis H. melvillei H. rodeiensis H. sartor H. vaupelli H. araneiflora H. bicornis H. bractescens H. conopsea H. cryptophila H. distans H. gourlieana 1-1. helodes H. hydrophila H. longicauda H. macroceratitis H. melvillei H. pringlei H. sartor H. vaupelli H. aricaensis H. bractescens H. burkartiana H. fastor H. gourlieana H. helodes H. juergensii 1-1. kleyi 1-1. longicauda H. macroceratitis H. macronectar H. vaupellii anaplectron bicornis bractescens conopsea distans gourlieana helodes hydrophila longicauda linguicruris macroceratitis melvillei vaupelli alliance robust, leafy -large, white, showy - -- -long -H. aricaensis H. bractescens H. fastor H. gourlieana H. helodes H. Iongicauda H. macronectar H. vaupelli The em-dash (--) represents no data available. t i a n a , H. f a s t o r , H. g o u r l i e a n a , H. l o n g i c a u d a , H. m a c r o n e c t a r , and H. s a r t o r were transferred to the new genus K u s i b a b e l l a . Al- though some p r e l i m i n a r y data u s i n g m o l e c u lar markers indicated that the genus H a b e n a r i a is clearly polyphyletic ( B a t e m a n et al., 2003), in our opinion, for the m o m e n t , the proposed transferences are hasty and superficial as they have no support even at the morphological level. For instance, H. v a u p e l lii is separated from the bulk of species in sect. M a c r o c e r a t i t a e , which are very similar to each other, and is grouped with species with which it shares few similarities. The Brazilian species in sect. Macroceratitae The exact circumscription of sect. M a c r o c e r a t i t a e is still not clearly defined, as K r a e n - zlin did not select a type species for the section, and H. m a c r o c e r a t i t i s Willd. does not have the long stigmatic processes used by Kraenzlin as one of the major distinctive characters of the section. However, a m o n g the species placed by Kraenzlin (1892, 1901) and C o g n i a u x (1893) in sect. M a c r o c e r a t i t a e , there is a very distinct and certainly natural group characterized by large flowers with long spurs and long, involute stigmas. This is the group that best fits K r a e n z l i n ' s definition of sect. M a c r o c e r a t i t a e and best represents the species that are assigned to the section. As we show below, this group is represented in Brazil by seven species: H. b r a c t e s c e n s , H. g o u r l i e a n a , H. j o h a n n e n s i s Barb. Rodr., H. longicauda, H. macronectar, H. nabucoi Ruschi, and the new species described here as H. p a u l i s t a n a J . A . N . Bat. & Bianchetti. A m o n g the other New World H a b e n a r i a , 2006] BATISTA ET AL.: HABENARIA SECT. MACROCERATITAE (ORCHIDACEAE) only one other species, H. p r i n g l e i , from Mexico and Central America, belongs with certainty to this same group. A species recently described from Brazil, n a m e l y H. p a b stii J. A . N . Bat. & Bianchetti (Batista et al., 2003), differs from the others species in this group in some i m p o r t a n t characters and has not been included here since its exact infrageneric position is unclear. Currently, this group of species in sect. M a c r o c e r a t i t a e can be recognized by the following characters: plants usually tall, robust, leafy, growing in h u m i d places or in water at the margins of small streams, ponds, and lakes; leaves linear to linear-lanceolate, usually conduplicate and keeled; flowers m e d i u m to large for the genus (the largest flowers a m o n g New World H a b e n a r i a ) ; spur long, pendent, longer than the pedicellate ovary, partially or c o m p l e t e l y enclosed by the bracts; dorsal and lateral sepals approximately the same size; petals bipartite, lip tripartite; lateral segments of the petals and lip 13 usually linear and elongated, and of the same size as or usually longer than the other segment; anther canals long; stigmas long, separated, or in contact only at the apex, base involute, that is, the margins rolled spirally inward on each side (Steam, 1983), the apex wide, not involute, the receptive surface turned to the sides or frontward. Another species in sect. M a c r o c e r a t i t a e , 1t. q u i n q u e s e t a (Michx.) A. Eaton (conspecific with H. m a c r o c e r a t i t i s ) , is k n o w n from southe m North America, Central America, the A n tilles, and northern South America, and has distinct stigmatic processes that are not particularly long, or involute, but are closely parallel, forming a u n i q u e receptive surface. The exact characterization of sect. M a c r o c e r a t i t a e and the infrageneric classification of 11. q u i n q u e s e t a are b e y o n d the scope of this work, but it is clear that further studies are necessary to define and delimit sect. M a c r o c e r a t i t a e and the infrageneric position of some species such as H. q u i n q u e s e t a and H. p a b s t i i . Key to Habenaria sect. Macroceratitae in Brazil 1. Spur shorter than 8.5 cm. 2. Flowers mostly white, lateral sepals not reflexed, lateral segments of petals and lip approximately of the same length and width as the posterior petal and midlobe of the lip ............................................... H. bractescens 2. Flowers mostly green, lateral sepals deflexed, lateral segments of petals and lip narrower and longer than the posterior petal and midlobe of the lip............................................................................... H. macronectar 1. Spur longer than 10 cm. 3. Midlobe of the rostellum prominent, mostly projected beyond the anthers; apex of rostellum broad, dentate to praemorse; viscidia very close, almost touching each other ................................................. H. johannensis 3. Midlobe of the rostellum partially or totally enclosed between the anthers; apex of rostellum narrow, acute to obtuse; viscidia usually spaced ca. 2-10 mm from each other (except H. paulistana). 4. Flowers mostly white; spur clavate, thickened towards the apical part (2-4 mm wide); midlobe of rostellum tall (8-9 mm), partially projected beyond the anthers....................................................... 11. nabucoi 4. Flowers mostly green; spur linear, not thickened towards the apical part (ca. 1 mm wide); midlobe of rostellum short (3.5-5 ram) completely enclosed between the anthers. 5. Lateral sepals not reflexed; anterior petal and lip lateral segments 1.8-2.6 times as long as the posterior petal and midlobe of the lip ............................................................................................14. gourlieana 5. Lateral sepals strongly reflexed or deflexed; anterior petal and lip lateral segments 1.2-1.8 times as long as the posterior petal and midlobe of the lip. 6. Lip segments and anterior petals reflexed; lateral sepals reflexed, forming a 90~ angle with the dorsal sepal; spur 13-15.5(-20) cm long; NE and N. Brazil.............................................. 14. longicauda 6. Lip segments and anterior petals not reflexed; lateral sepals deflexed, forming a 180~ angle with the dorsal sepal; spur 11.5-13 cm long; state of S~o Paulo ........................................ H. paulistana 1. HABENARIA BRACTESCENS Lindl., Gen. Sp. Orchid. P1. 308. 1835. K u s i b a b e l l a bractescens (Lindl.) Szlachetko, Richardiana 4(2): 59. 2004. TYPE: A R G E N TINA. Buenos Aires, J. T w e e d i e s . n . (HOLOTYPE; K, image seen; ISOTYPE; OXF). (Figs. 1 & 2) Habenaria sartor Lindl. forma minor Cogn., Bull. Herb. Boissier, set. 2, 3(10): 929. 1903. TYPE: PARAGUAY. in regione Cordillerae centralis, 1900, E. Hassler 6594 (HOLOTYPE:presumably at BR, n.y.; ISOTYPE:BM, image seen). Habenaria hauman-mercki Hickens, Anales Soc. Ci. Argent. 65: 304. 1908. SYNTYPES:ARGENTINA. Islas del Tigre, Feb 1902, C.M. Hickens s . n . (LECTOTYPE, designated by Toscano, 1994: SI); Bel- 14 BRITTONIA [ V O L . 58 B 5 ]lmm //'11 5mm lo ~ . . . 5mm E ........................................................................................................................................................................................................................... . 5r.rn lmm s.1 nl .................................................................................................. lmm FIG. 1. Dissected flower segments. A. Habenaria bractescens. B. H. gour/ieana. C. H. johannensis. D. H. longi cauda. E. H. rr~cronectar. F. H. nabucoi. 1. Perianth. 2. Ovary and pedicel, column and spur, side view. 3. Co]un'm, side view. 4. Column 3/4 side view. 5. Column apex, front view showing the connective and anther apex. 6. Rostellum, side or 3/4 side view. 7. Rostellum apex, front view. (A1, Sidney & Onishi 1560, UB; A2, Rojas 3961, SP; A3, Hatschbach & Guimardes 20530, MBM; A5-6, Camargo 18, MBM; B1, Batista 128, CEN; B2, Batista 344, CEN; B3-6, Batista 948, CEN; C1-2, Barros 2231, SP; C3.1, Pansarin 763, UEC; C3.2 and C5-7, Batista & Bianchetti 908, CEN; D1-6, Batista 1039, CEN; El, Scariot 344, CEN; E2, Vidal et aL 609, VIC; E3-6, Borba et al. s.n., UEC 118854; F1, Batista 1040, CEN; F2, Batista 253, CEN; F3-6, Batista & Oliveira 848, CEN. Perianth, pedicel and ovary drawn from dried material, other structures from alcohol-fixed material, except for H. bractescens where all drawings were made from dried specimens.) 2006] BATISTA ET AL.: HABENARIA A ~ f #8 SECT. M A C R O C E R A T I T A E ' 9 (ORCHIDACEAE) H 15 B "',~.~ ! C FIG. 2. Flowers, photographed from live specimens 9 A. Habenaria bractescens, inflorescence, from Foz do Iguagu, Paran~i. B. H. nabucoi, flower 3/4 side view, from Lagoa Bonita, Planaltina, Distrito Federal (Batista & Oliveira 848, CEN). C. H. gourlieana, flower side view, from Chapada dos Veadeiros, Goi~is (Batista 948, CEN). Scale bar = 1 cm. 16 BRITTONIA grano, cerca del rio, 1906, L. L. Hauman-Merck s.n. (drawings: SI, photocopy seen). Habenaria pontagrossensis Kraenzl., Kongl. Svenska Vetenskaps. Handl. 46(10): 6, taf. 2, Fig. 1. 1911. TYPE: BRAZIL. Paran,'i, Ponta Grossa, Rio Tibagy, 26 Jan 1909 (fl), P K. H. Dusen 7837 (HOLOTYPE: S, image seen; ISOTYPE: HBG, image seen). Habenaria kleyi Schltr., Repert. Spec. Nov. Regni Veg. Beih. 35: 20. 1925. TYPE: BRAZIL. Rio Grande do Sul, in der Umgebung von Porto Alegre, Jan 1919, U. Kley 01 (HOLOTYPE: B, destroyed; no isotype known). NEOTYPE, here designated: BRAZIL. Porto Alegre, U. Kley in Herb. J. Dutra 994 (SPF). Semi-aquatic herb. Stem short to medium, leafy, 20-56(-87) • 0.2~3.5(~3.7) cm including the inflorescence. Leaves (6-)8-10(-12), spreading, linear-lanceolate to lanceolate, elongated, largest at the center of the stem, 6.5-21 • 1-1.6(-2.4) cm, the upper leaves similar to the lower bracts. Inflorescence 616 cm long, usually short and few flowered; bracts 2-4.5 • 0.7-1.7 cm, shorter than the pedicel and ovary, decreasing in size towards apex of inflorescence, ovate-lanceolate to lanceolate, green. Flowers 2-6(-15), superposed, medium sized for the genus, but small for the section, whitish; pedicel and ovary 3.3-5.5 cm, green. Sepals 11-14 • 7-8 mm, mucronate, abaxial side greenish white, adaxial 'side white; dorsal sepal ovatelanceolate; lateral sepals 13-15 • 7-7.5 mm, not reflexed, obliquely ovate, apiculate. Petals bipartite, white, towards the apex creamy white or light green; posterior segment linear, slightly falcate, 13-15 • 1.2-1.5 mm; anterior segment 15-16 • 0.8-1.5 ram, linear. Lip tripartite, white, towards the apex creamy white or light green, pendent; the undivided basal part 3-4 • 1.5-2.5 mm; lateral segments linear, 13-14(-19) • 0.8-1 mm; central segment 12-15 • 1.5-1.8 mm, linearligulate. Spur 4-6.2(-7.5) cm long, 1.5-2 mm wide at base, 2-3.5 mm at apex, pendent, partially exposed, with a small erect triangular tooth ca. 1 mm tall, obtuse at the entrance of the orifice, thickened towards the apical part, base white, towards the apex greenish. Column 4-6.5 mm tall, white; anther 3.5-5 mm tall, the canals slightly erect, 3-5 mm long; connective emarginate; rostellum 5-7 mm long, the midlobe short, ca. 2 mm long, 3-4 mm tall, triangular, fleshy, completely enclosed between the anthers, acute, side lobes ca. 4-5 mm long, the base [VOL. 58 broad, ca. 1.5 mm wide, the apex 0.5-0.7 mm wide, truncate; auricles erect, triangular, fleshy, verrucose, flattened, with a broad base and acuminated towards the apex, ca. 2-2.5 mm tall, acute; stigmas 2, 8-9 mm long, the free projected part 5-6 mm long, the margins curved, involute, the apex not curved, ca. 1.5-2.5 mm wide. Pollinaria 2, yellow, the viscidia ca. 0.8 • 0.5 mm; caudicles elongate, ca. 5 mm long; pollinia 2 per pollinarium, laterally flattened, ca. 4 mm long. Distribution.---Argentina; Brazil (Bahia, Goi~is, Minas Gerais, Mato Grosso do Sul, Paran~i, Santa Catarina, Rio Grande do Sul); Paraguay; and Uruguay. Habenaria bractescens is concentrated mainly in southern South America (southern Brazil, Argentina, and Uruguay), from where there is abundant material, but can occasionally occur in central and northeastern Brazil (Fig. 3), where it has been rarely collected. Considering the known distribution of H. bractescens, the reports for northern South America, including Venezuela (Foldats, 1969) and Ecuador (Dodson, 1988) are doubtful. The species reported from Colombia (Hern~indez, 1958), based on the size of the spur (12-14 cm), clearly is not H. bractescens. The reports of H. bractescens from Central America are all referable to H. pringlei (Toscano de Brito, 1994). Habitat, Ecology and Phenology.--The species grows in water along the margins of rivers, lakes or marshes. In central Brazil, H. bractescens is an early-flowering species, with flowering at the very beginning of the rainy season, in October. In southern Brazil, Argentina, and Uruguay, flowering extends until February and March. Illustrations.--Kraenzlin (1911, plate 2, Fig. 1, as H. pontagrassensis), Hauman (1920, Fig. 1), Mansfeld (1930, plate 3, Fig. 12, as H. kleyi), Hoehne (1940, plate 12; plate 13, as H. kleyi), Cabrera (1942, Fig. 3), Cocucci (1954, Fig. 3), Pabst and Dungs (1975, Fig. 8), Artucio (1984, plate 167, Fig. 1), Toscano de Brito (1995, Fig. 46Jg-Jh). Additional specimens examined: BRAZIL. BAHIA: between 2.5 and 5 km S of Vila do Rio de Contas on side road to W. of the road to Livramento, leading to the Rio Brumado, 41~ 13~ 980 m, 28 Mar 1977 (fl), Harley et al. 20099 (CEPEC, HB, SPF); Mun. Jussiape, margin of Rio de Contas, Cachoeira do Fraga, 16 Feb 2006] B A T I S T A E T AL." H A B E N A R I A S E C T . M A C R O C E R A T I T A E ( O R C H I D A C E A E ) /"71 * 8oi " _\ ~ .. 'N.Ir~,,, ~.,r ......... ;o1_/ o,i i o ~ 0 ,oo lO0 ,oo 200 300 ,oo ,~ 17 ,~o,~ 4 0 0 5 0 0 61iO m l l l l i ,oi FIG. 3. Distribution of Habenaria bractescens (star), H. gourlieana (circle), and H. johannensis (square). Based on Flora Neotropica base map no 1. 1987 (fi), Harley et al. 24332 (SPF); Mun. Rio de Contas, ca. 1 km from Rio de Coritas on the road to Cachoeira do Fraga, 13~ 41~ 23 Mar 1996 (fl), Jardim et aL 768 (CEPEC). GoI~s: Rio Verdfio, 15 Oct 1968 (fi), Sidney & Onishi 1560 (UB). MATO GROSSO DO SUL: Sete Quedas, 17 Aug 1937 (fl), Cullen s.n. (HB 70204, PACA 89588, RB 35466, SP 314386). MINAS GEgAIS: Mun. Ituiutaba, canal de Sao Sim~o, 15 Oct 1950 (fl), Macedo 2631 (CEN, HB, RB, SP, SPF); Canal de S~o Sim~o, 9 Sept 1976, Raspalho s.n. (OUPR). PAgAN,g,:Ponta Grossa, Lagoa Dourada, 7 Dec 1978, Dombrowski 10109 & Scherer-Neto 616 (HB, MBM); Parque Nacional de Sete Quedas, Guaira, 9 Sept 1980 (fl), Fontetta 1164 (RB); Mun. Guaira, Parque Nacional Sete Quedas, 200 m, 16 Oct 1962, Hatschbach 9332 (HB, MBM); Mun. Ponta Grossa, Parque Vila Velha, Lagoa Dourada, 788 m, 21 Dec 1962, Hatschbach 9611 (HB, MBM); F6z do Igua~u, Parque Nacional Cataratas do lguagu, 200 m, 20 Feb 1963, Hatschbach 9900 (MBM); Mun. Guaira, Sete Quedas, 16 Sept 1981, Hatschbach 43974 (MBM, SPF), 200 m, 22 Apr 1968, Hatschbach 19094 & Guimaraes 182 (MBM, UPCB); 18 BRITTONIA Mun. Dois Vizinhos, Foz do Chopim, 6 Dec 1968, Hatschbach & Guimaraes 20530 (MBM); Sete Quedas, Guaira, 22 Jun 1967, Hatschbach & Haas 5495 (HB); Ponta Grossa, Lagoa Dourada, 2 Jan 1976, Krieger 14365 (CESJ, HB); Vila Velha, Lagoa Dourada, 28 Dec 1981, Krieger 15492 (CESJ, HB); Mun. Ponta Grossa, Lagoa Dourada, 26 Feb 1985, Kurnmrow et aL 2559 (MBM); BR 376, Lagoa Dourada, just after turning to Vila Velha in direction of Ponta Grossa, ca. 50~ 25~ 815 m, 28 Jan 1985, Lewis et al. 1390 (MBM, SPF); Foz do Iguaqu, banks of Floriano Falls, 10 Dec 1967, Pabst & Emmerich 9029 (HB); Mun. Ponta Grossa, Vila Velha, Lagoa Dourada, 19 Jan 1974, Moreira-Filho s.n. (UPCB 25088); Mun. Guafra, Salto das Sete Quedas, 23 Jul 1977, Kuniyoshi 4289 (MBM); Sete Quedas, Guaira, 9 Jul 1950, L. Camargo 18 (MBM). RIo GRANDE DO SUL: Arroio dos Patos, Fazenda Faxinal, 19 Jan 1976, Hagelund 12252 (HB), 29 Jan 1982 (fl), Hagelund 13787 (MBM); Porto Alegre, Juergens 1044 (SP); Vacaria, Encanados, 18 Dec 1997, Nauhs s.n. (PACA 85111); Morro do Sabid, near Porto Alegre, 28 Dec 1948, Rambo s.n. (HB 231, PACA 39245); Rio Guira, near $5o Francisco de Paula, 10 Feb 1941, Rambo s.n. (PACA 4746). PARAGUAY. DEPT. CANINDEYI~: Paraguaria euroaustra, in regione Yerbalium de Maracayd, 1898-1899, Hassler 5950 (BM); Guaird Falls, Sept 1921 (fl), Rojas 3961 (SP). URUGUAY. DEPT. CERRO LARGO: Jan 1926, Herter 18370 (HB, SP); Cerro Largo, 23 Feb 1938, Rosengartt B2577 (RB). DEPT, MINAS: Mar 1924, Herter 17166 (SP). DEPT. SORIANO:Berro 1472 (HB). ARGENTINA. PROV. BUENO AIRES: delta of the Parand, Feb 1931, Burkart 3812 (SP); delta of the Paranfi, 31 Jan 1932, Burkart 4510 (SP); delta of the Paranfi, 18 Jan 1931, Cabrera 1623 (SP); Bueno Aires, 6 Dec 1883 (fl), Spegazzini 970 (SP). PROV. CHACO: Colonia Benftez, Jan 1937, Schulz 582 (SPF). PROV. CORRIENTES: Dept. Ituzaing6, Ruta 12, 13 km W of the boundary with Misiones, 9 Jan 1976 (fl), Krapovickas & Crist6bal 28649 (MBM); Dept. San Miguel, Ruta 17, Curuzfi Laurel, 11 km NE of San Miguel, 19 Jan 1933 (fr), Schinini 27559 (MBM, SPF). PRov. MIsIOr~ms: Dept. Iguazd, Isla San Martfn, in front of San Martin Falls, 8 Aug 1991 (fl), Vanni et aL 2826 (MBM); Dept. Iguazti, Parque Nacional del Iguazd, cataratas, 14 Dec 1986, Xifreda & Maldonado 484 (MBM). can b e distinthe other species in sect. M a c r o c e r a t i t a e b y the ratio b e t w e e n the posterior and a n t e r i o r segments o f the petals, and b y the lateral lobes and m i d - l o b e o f the lip, which in each case are about the s a m e size and w i d t h (Fig. 1). The small flowers and short spurs o f H. b r a c t e s c e n s are s i m i l a r to H. m a c r o n e c t a r , but H. b r a c t e s c e n s can be d i s t i n g u i s h e d b y its white flowers (vs. u s u a l l y g r e e n - y e l l o w in H. m a c r o n e c t a r ) a n d b y nonreflexed lateral sepals (vs. p r o m i n e n t l y deflexed). A l s o , in H. m a c r o n e c t a r , the lateral Habenaria guished from bractescens [VOL. 58 segments o f the petals and those o f the lip are n a r r o w e r and l o n g e r than the petal p o s t e r i o r s e g m e n t and lip m i d - l o b e , respectively. O u r o b s e r v a t i o n s o f the flower c o l o r o f H. b r a c t e s c e n s are b a s e d on a c o l o r p h o t o g r a p h , k i n d l y p r o v i d e d b y W a l t e r Rossi, a n d on the i n f o r m a t i o n f o u n d on the h e r b a r i u m label o f the m a t e r i a l e x a m i n e d . The flowers are pred o m i n a n t l y w h i t e or s o m e w h a t green-white. In general habit and flower morphology, H a b e n a r i a b r a c t e s c e n s appears to b e similar to H. n a b u c o i in the same w a y as H. m a c r o n e c t a r is related to H. p a u l i s t a n a , i.e., it looks like a smaller form o f the latter. Additionally, H. b r a c t e s c e n s a n d H. n a b u c o i are f r o m waterlogged places, and have inflorescences with usually only a few predominantly white flowers and non-reflexed lateral sepals (Fig. 2). However, the two species differ in colunm structure: H. b r a c t e s c e n s does not have the tall rostellum as in H. n a b u c o i (Fig. 1). The e x a m i n a t i o n o f an i m a g e o f the isotype o f H. s a r t o r f. m i n o r C o g n . f r o m Paraguay, l o c a t e d at the Natural H i s t o r y M u seum H e r b a r i u m (BM), r e v e a l e d it is referable to H. b r a c t e s c e n s . The H a u m a n - M e r c k s y n t y p e o f H. h a u m a n - m e r c k i is j u s t a specim e n o f H. b r a c t e s c e n s with tripartite petals, a character that appears o c c a s i o n a l l y in H. b r a c t e s c e n s as well as in several other species in the genus, and is u n r e l i a b l e for the segregation o f a species (see also the notes for H. p a u l i s t a n a ) . A critical analysis o f the original d e s c r i p t i o n and illustration o f H. k l e y i and e x a m i n a t i o n o f i m a g e s o f the t y p e o f H. p o n t a g r o s s e n s i s c o n f i r m e d the p o s i t i o n o f other authors (Hoehne, 1940; Pabst & Dungs, 1975; Toscano de Brito, 1994) w h o i n c l u d e d one o r both species u n d e r the syno n y m o f H. b r a c t e s c e n s . A s w e have b e e n unable to locate an i s o t y p e o f H. k l e y i , w e select as n e o t y p e a s p e c i m e n from the s a m e locality and c o l l e c t o r (Porto A l e g r e , U. K l e y ) , located at the S P F h e r b a r i u m u n d e r the n u m b e r 994 o f the Jo~o Dutra collection. 2. HABENARIA GOURLIEANA Gill. ex Lindl., Gen. Sp. Orchid. P1. 309. 1835. K u s i b a b e l l a g o u r l i e a n a (Gill. ex Lindl.) Szlachetko, R i c h a r d i a n a 4(2): 59. 2004. TYPE" ARGENTINA. Mendoza, on m a r s h y ground, J. G i l l i e s s . n . (HOLOTYPE: K, i m a g e seen; ISOTYPE: OXF). ( H o l o t y p e 2006] B A T I S T A E T A L . : H A B E N A R I A SECT. MACROCERATITAE A vs. Ro t,An An :: Ac cn , B ~ .,, S t ~~ ' / So/ Vs ~ ~ Ro / Ca R.~ ~ I F ~ Sp Cn ~ , ' ....I.. .... / m E 19 ./../Cn Au St (ORCHIDACEAE) t~. / A n / " " / ~...~ / : An ov 9 st ....... Au Vs~. / Sp m f "-- ,/ Ra D An Vs Ro ~/ Cn Au ~"~ / / / Ra 2 _ ~ ~ ~ Au / Sp St St-'----- . .,~ " Ov / Sp ~ FIG. 4. Columns, photographed from live specimens. A. Habenaria nabucoi (Batista & Oliveira 848, CEN). B. H. gourlieana (Batista 948, CEN). C. H. johannensis (Batista & Bianchetti 908, CEN). D. H. longicauda (Batista 1039, CEN). E. H. paulistana (Batista & Pansarin 1089, CEN). Ac = anther canals; An = anthers; Au = auricles; Ca = caudicles; Cn = connective; Ov = ovary; Ra = rostellum arms; Ro = rostellum; Sp = spur; St = stigmas; Vs = viscidium. Scale bar = 2 mm. sheet at K m o u n t e d t o g e t h e r with a specim e n o f H. johanennsis.) (Figs. 1, 2, 4) Macrocentrurn mendocinum Phil., Annal. Univ. Chile 200. 1870. TYPE: unknown. Habenaria spegazziniana Kraenzl., Bot. Jahrb. Syst. 36(80): 10. 1905. TYPE: ARGENTINA. auf Berg- wiesen am Wege zwischen E1 Santuario und Carmen Calchaqui, C.L. Spegazzini 104753 (HOLOTYPE: HBG, image seen). Habenaria burkartiana Hoehne, Bot. Jahrb. Syst. 68: 127, Taf. 11. 1937. Kusibabella burkartiana (Hoehne) Szlachetko, Richardiana 4(2): 59. 2004. SYNTYPES: ARGENTINA. Concordia, provincia 20 BRITTONIA Entre-Rios, 22 Jan 1927 (fl), A. Burkart 946 (SYNTYPE: SP); Buenos Aires, Juancho, 13 Jan 1933 (fl), A. L. Cabrera 2710 (SYNTYPE:SP). Habenaria fastor Warm. ex Hoehne, Flora Brasilica 12(1): 71, Tab. 15. 1940. SYNTYPES: BRAZIL. Minas Gerais, Serra de S~o Calixto, Gr~o Mogol, 11 Nov 1938 (fl), F. Markgraf 3350, A. C. Brade & M. Barreto 12092 (SVNTVPES: BHMH, RB, SP); Mato Grosso, Rio Sap6, 10 Oct 1938 (fl), J.E. Rombouts s . n . (SYNTYPES:IAC 2756, SP 40966). Terrestrial herb. Stem medium and somewhat slender to tall and robust, leafy, (42-) 64-115 x 0.4-0.9 cm including the inflorescence. Leaves 8-12, spread, lanceolate, not elongated, lowermost sheath-like, largest around the center of stem, 9-18(-25) x 1.63.4 cm, upper leaves similar to the lower bracts. Inflorescence 9-25 cm long, usually few-flowered; bracts green, shorter than the pedicel and ovary, decreasing in size towards the apex of the inflorescence, lanceolate to oblong-lanceolate, 4-5.3 x 1.3-1.7 cm. Flowers (1-)3-12, usually few, spreading, superposed, large for the genus, greenish; pedicel and ovary green, slightly arched, 6-8 cm. Sepals light green to greenish white; dorsal sepal largely ovate-lanceolate, apiculate, (11-)15-19 x 9-13 mm; lateral sepals not reflexed, obliquely lanceolate to oblonglanceolate, apiculate, (14-)18-24 • (6-)8-11 mm. Petals light green or greenish white, the basal part whitish, bipartite; posterior segment linear to near-lanceolate, falcate, connivent with the dorsal sepal, 15-21 x 2-2.5 (-3) mm, anterior segment projected forward, somewhat parallel to the lip lateral segments, 1.8-2.6 times as long as the posterior segment, straighter, linear, 34-48 x 1 mm. Lip light green or greenish white, the basal part whitish, tripartite, projected forward, somewhat concave; undivided basal part whitish, 5-8 • 2-3 mm; lateral segments linear, (2-) 2.2-3 times as long as the middle segment, (25-)35-52 x 0.8-1(-1.5) mm; middle segment shorter and slightly wider, linearligulate, (11-)15-21 x 1.5(-2) ram. Spur whitish at the base and becoming greenish towards the apex, without a triangular tooth at the entrance of the orifice, pendent, arranged between the bracts, linear, 11-14.5 • 0.15-0.2 cm. Column 6-7 mm tall; anthers whitish, apparently yellow because of the pollinia, slightly arched, 4-5 mm long, canals white, slender, erect, ca. 7 mm long; [VOL. 58 connective green, retuse to apiculate; rostellum green, 8-9 mm long, middle lobe fleshy, triangular, slightly shorter than the anthers, about half the size of the column, 4-5 mm tall, completely arranged between the anthers, acute, side lobes erect, 6-7 mm long; auricles white, erect, fleshy, somewhat verrucose, triangular, ca. 2-3.5 mm tall, obtuse; stigmas 2, green, 12-13 mm long, the free receptive part 7-9 mm long, margins curved, somewhat involute, apex not curved, truncate, 2-3 mm wide. Pollinaria 2, yellow, viscidia 0.8-0.9 x 0.5-0.6 mm, spaced 2-3 mm apart; caudicles elongate, ca. 8 mm long; pollinia 2 per pollinarium, laterally flattened. Distribution.--Argentina; Brazil (Distrito Federal, Goi~is, Mato Grosso, Minas Gerais, Rio Grande do Sul, Tocantins); and Uruguay. Habenaria gourlieana is apparently a common species in Argentina and Uruguay, but it is rare in southern Brazil, from where we have seen only one collection from the state of Rio Grande do Sul (Fig. 3). The species was reported for Santa Catarina (Pabst & Dungs, 1975) and Parand (Hatschbach, 1962; Angely, 1965; Pabst & Dungs, 1975), but all the material we have seen from these states is misidentified. However, it should probably be expected for the other states in southern Brazil. The species has also been recorded from Bolivia (Schlechter, 1922; Pabst & Dungs, 1975) and Paraguay (Cogniaux, 1893, 1903, 1905), but we have been unable to confirm these reports. The only collection from Paraguay that we have seen identified as H. gourlieana by Cogniaux (Hassler 8703, BM) is referable to H. johannensis. Habitat, Ecology, and Phenology.--In central Brazil, Habenaria gourlieana is an early flowering species, with flowering at the very beginning of the rainy season, from October to November. In Uruguay and Argentina, flowering is somewhat latter, occurring from December to February. The flowers are green or light green, but never pure white as in H. nabucoi. Also, in contrast to H. nabucoi, we have found the species growing in humid places, such as in marshes with shrubs beside gallery forests, but not in water. Illustrations.--Cogniaux (1893, Fig. 4; reproduced in Hoehne 1940, plate 20), Hoehne (1919, plate 165 and plate 167, Fig. 2, as H. fastor), Hoehne (1937, plate 11, as H. 2006] BATISTA ET AL." HABENARIA SECT. MACROCERATITAE (ORCHIDACEAE) 21 the size of the plants and floral parts we have seen from Argentina is somewhat smaller than for material f r o m central Brazil, and the smaller values in the description are in m o s t cases referable to plants from Argentina. In the illustration provided by Singer and Cocucci (1997), which is based on material f r o m Argentina, the column is somewhat different from the material from central Brazil that we have examined. The rostellum mid-lobe is more prominent and is partially projected beyond the anthers. However, these Additional specimens examined: BRAZIL. DIS- differences are small and are insufficient to TRITO FEDERAL:Guar~i, Reserva Eco16gica do Guar~i, suggest any degree o f taxonomic recognition, 20 Oct 1990 (fl), Batista 128 (CEN), 11 Nov 1991 (fl), even at the subspecific level. Batista 215 (CEN), 18 Oct 1992 (fl), Batista 344 (CEN); A n analysis o f the syntypes of H. burkarBrasflia, area between Lago Sul QI-17 and Setor de Mansres Park Way quadra 25, near ribeir~o do Gama, 14 tiana didn't reveal any specific difference Dec 2002 (fr), Batista & Pellizzaro 1320 (CEN); f r o m H. g o u r l i e a n a , thus confirming the preTaguatinga, Proflora marsh, 24 Nov 1988 (fl), Filgueiras vious inclusion o f H. b u r k a r t i a n a as a syn2349 (IBGE); Guard, Industrial sector, Heringer 8158 (IAN), 10 Jan 1963, Heringer 10025 (UB); Brasflia, o n y m of H. g o u r l i e a n a (Pabst & Dungs, Piano Piloto, Feb 1963, Heringer 10041 (HB). GoI~s: 1975). H a b e n a r i a s p e g a z z i n i a n a Kraenzl. Chapada dos Veadeiros, 23 km from GO-118 on the road was described as having tripartite petals, to S~o Jorge and Colinas do Sul, 11 Oct 1999 (fl), however the examination o f an image o f the Batista 948 (CEN); Formosa, 19 Oct 1965 (fi), A.P. holotype and a critical analysis o f the speDuarte & E. Pereira 10287 (HB); same locality, 19 Oct 1965 (fl), A.P. Duarte 9376 (RB); Mun. Alto Paraiso, cies original description didn't reveal any adfazenda Mat0 Fundo, 1200 m, 16 Oct 1990 (fl), ditional significant difference from H. Hatschbach et al. 54669 (HEPH, MBM). MINAS g o u r l i e a n a . Since this character is variable GERAIS: Caet6, Nov 1915 (fl), Hoehne 6420 & 6421 and represents a genetic abnormality (see the (R); Betim, 30 Jul 1955 (fl), Roth 16479 (CESJ, HB); Mun. Formiga, western Minas Gerais, Oct 1956 (fl), notes for H. p a u l i s t a n a ) , H. s p e g a z z i n i a n a is Welter 199 (HB). RIO GRANDE DO SUL: Mun. Caqa- treated here as a s y n o n y m of H. g o u r l i e a n a , pava do Sul, BR 153, Jan 1994 (fl), Falkenberg et aL as suggested by Pabst and Dungs (1975). We 6390 (BHCB). TOCANTINS:Alianqa, 7 km from the city, have not seen the type material o f M a c r o 12~176 14 Nov 1997 (fr), Proen~a & Lake c e n t r u m m e n d o c i n u m Phil. and the present 1874 (UB). URUGUAY. DEPT. FLORIDA:Arroyo Mansevillagra, inclusion o f the name in the s y n o n y m y Picada Castro, Estancia "Rinc6n de Santa Elena", Jan o f H. g o u r l i e a n a is based on previous 1947 (fl), Rosengurtt 5854 (HB). DEPT. MINAS: cerda authors (Cogniaux, 1893; Kraenzlin, 1901; de Currales, Feb 1921 (fl), Schroeder 15752 (HB). Schlechter, 1922; Hoehne, 1940; Pabst & DEPT. MONTEVIDEO" Montevideo, Feb 1875, Arechavaleta 2625 (BR). DEvr. Rio NEGRO: Rio Dungs, 1975). The creation o f the genus K u s i b a b e l l a Negro, l0 Feb 1914 (fl), Berro 7298 & 7299 (HB). DEPT. SALTO: Salto, 2 Feb 1937, Rosengurtt B1071 (Szlachetko, 2004) is in our opinion prema(RB). DEPT. SANJos~: Barra Santa Lucfa, 28 Jan 1919 ture. Some studies have shown that (fl), Osten 14723 (HB). DEPT. SORIANO: Soriano, H a b e n a r i a is highly polyphyletic (Bateman Rosengurtt et al. PE-4763 (HB). ARGENTINA. PRov. CORRIENTES: Dept. Empe- et al., 2003), but these were based primarily drado, Estancia La Yela, 21 Dec 1972 (fl), Pedersen in African and Asiatic species, where sub10271 (MBM). tribe Habenariinae is particularly diverse. However, this is still to be shown for tropical America, were H a b e n a r i a is the only repreHabenaria gourlieana can be distinguished from other species in the section by sentative of subtribe Habenariinae. Furthermore, considering that most Neotropical the green flowers, non-reflexed lateral sepals, and long, filiform lateral segments o f the H a b e n a r i a species are still poorly characterized, even at the morphological level, and petals and lip, which are 1.8-2.6 and 2 - 3 times as long as the posterior petals and lip that, consequently, infrageneric relationships within Neotropical H a b e n a r i a are poorly mid-lobe, respectively (Figs. 1, 2). Overall, burkartiana; reproduced in Hoehne 1940, plate 22), Cabrera (1942, Fig. 4, reproduced in Correa 1968, Fig. 121 and Cabrera & Zardini 1978, Fig. 52), Cocucci (1954, Fig. 4), Pabst and Dungs (1975, Fig. 10, based on R o s e n g u r t t et al. P E - 4 7 6 3 , HB; Fig. 13, as H. m a c r o n e c t a r , based on M a r k g r a f et al. 3 3 5 0 , RB), Artucio (1984, plate 167, Fig. 1), Singer and Cocucci (1997, Fig. 1A, B, column, Fig. 2 A - D , color plates), Galetto et al. (1997, Fig. 1A-D). 22 BRITTONIA known, any attempt to subdivide the genus in smaller genera should be preceded by a better characterization of the species and a reassessment of current sectional treatment, which is clearly equivocal. [VOL. 58 lel to the lateral segments of the lip, 1.5-2 times as long as the posterior segment, linear, 24-38 x 0.8-1(-1.5) mm. Lip whitish or greenish white, tripartite, projected forward, somewhat concave; undivided basal part 6-8 x 1-2 mm; lateral segments 1.6-2.1 times as long as the middle segment, linear, 33-47 x 3. HABENARIA JOHANNENSIS Barb. Rodr., Rev. Eng. 3: 74, Est. 2, Fig. B. [May] 1881. 0.8-1 mm; middle segment shorter and TYPE: BRAZIL. Minas Gerais, near S. slightly wider, linear-ligulate, 15-25 x 1.3-2 Joao d' E1-Rey, J. Barbosa Rodrigues s.n. mm. Spur whitish at the base and becoming (No original material is known. Typified greenish towards the apex, without a trianguby Cribb & Toscano de Brito, 1996: 1: 30. lar tooth at the base, pendent, linear, slightly Original illustration by Barbosa Rodrigues, thickened towards the apical part, enclosed reproduced in Sprunger, 1996: 1: 59B.) between the bracts, 10.8-13.5 cm, 1-1.5 mm (Figs. 1, 4, 5) in the middle, 2-3 mm in the apex. Column ca. 7-8 mm tall; anthers whitish, apparently Habenaria vaupellii Rchb.f & Warm., Otia Bot. yellow because of the pollinia, slightly Hamb. 2: 79. [August] 1881. Bertauxia vaupellii (Rchb.f & Warm.) Szlachetko, Richardiana 4(2): arched, ca. 4.5-6 mm tall, canals slender, 58. 2004. TYPE: BRAZIL. Minas Gerais, Lagoa partially erect, 10-13 mm long; connective Santa, E. Warming 131 (HOLOTYPE"C, photocopy green, apiculate; rostellum middle lobe seen). green, fleshy, prominent, elongated, forming Habenaria bradei Schltr., Anexos Mem. Inst. Butantan, Sec. Bot. 1(4): 13, Tab. 1, Fig. 4. 1922. TYPE: a tunnel-like structure projected beyond the BRAZIL. Sao Paulo, in uliginosis apertis Morca anthers, laterally compressed, 6-7 mm tall, prope urb. S~o Paulo, 9 Mar 1913 (fl), A. C. Brade base extended, slightly bent to form a chan6200 (HOLOTYPE: B, destroyed; LECTOTVPE: here nel that extends through the lateral lobes and designated, SP; ISOLECTOTYPE:HB). which supports the anther canals, apex diTerrestrial herb. Stem usually tall, robust, lated, tridentate or serrated, sometimes emarleafy, occasionally short and slender, (48-) ginate, ca. 1.8-2.5 mm wide, lateral lobes 79-148 • (0.4-)0.6-1.3 cm including the in- elongated, parallel, almost or in contact at the florescence. Leaves 7-12, spread, elongated, apex, 7-8 mm long, apex truncate; auricles linear, lowermost sheath-like, largest around white, ca. 8 mm long, the apical free part the center of stem, 10-30- (45) • 1-3.4 cm, erect, small, verrucose, obtuse, ca. 3 mm upper leaves similar to the lower bracts. In- long; stigmas green, ca. 13-14 mm long, the florescence 9-28 cm, few to many flowered; free part 6-7 mm long, margins curved, bracts green, shorter than the pedicel and somewhat involute, apex not curved, trunovary, decreasing in size towards apex of in- cate, ca. 3 mm wide. Pollinaria 2, yellowish; florescence, lanceolate, 3.5-6.5 • 1.3-2 cm. viscidia large, close to each other, ca. 1.5-1.7 Flowers (3-)7-14(-20), spreading, super- • 0.7-1 mm; caudicles elongate, 13-14 mm posed, large for the genus, greenish white; long; pollinia 2 per pollinarium, laterally flatpedicel and ovary green, slightly arched, 4.5- tened, 6 • 2 mm. Etymology.--Probably based on the local5.8 cm. Sepals green; dorsal sepal, ovatelanceolate to largely lanceolate, apiculate, ity were the type was collected, S~o Jo~o del 14-22 x '10-12 mm; lateral sepals non- Rei, in Minas Gerais, Brazil, one of the reflexed, the base superposed to the petal places from where Barbosa Rodrigues deposterior segment, obliquely oblong- scribed several new species. Distribution.--Bolivia; Brazil (Bahia, Dislanceolate, apiculate, 20-25 • 6-9 mm. Petals whitish or greenish white, bipartite; trito Federal, Espirito Santo, Goi~is, Mato posterior segment broad, connivent with the Grosso do Sul, Minas Gerais, Paran~i, Rio de dorsal sepal, 3.3-5 times as wide as the ante- Janeiro, Rio Grande do Sul, Sao Paulo, Santa rior segment, oblong-lanceolate, slightly fal- Catarina); and Paraguay. Habenaria johancate, 16-19 x 4-7 mm; anterior segment in- nensis is apparently most common in southserted 3--4 mm higher in the posterior ern and southeastern Brazil, particularly in segment, projected forward, somewhat paral- the states of S~o Paulo, Paran~i and Minas 2006] B A T I S T A E T A L . : H A B E N A R I A SECT. M A C R O C E R A T I T A E ( O R C H I D A C E A E ) 23 tt 9 B i~ ~ 9 ] C FIG. 5. Flowers, photographed from live specimens. A. Habenariajohannensis, flower front view, from Serra do Cip6, Minas Gerais (Batista & Bianchetti 908, CEN). B. H. longicauda, flower 3/4 side view, from Barreiras, Bahia (Batista 1039, CEN). C. H. paulistana, flower front view, from Serra do Japi, Jundiaf, Sao Paulo (Batista & Pansarin 1089, CEN). Scale bar = 1 cm. 24 BRITTONIA Gerais, from where there is abundant material (Fig. 3). However, the species can occasionally reach other parts of southern and central Brazil from where it has been collected more sporadically. Habenaria johannensis should also be expected for the state of Mato Grosso. According to Toscano de Brito (1995), the species also occurs in Bolivia. Habitat, Ecology, and Phenology.-Throughout the species distribution range, flowering is concentrated from December to February. In central Brazil, this period corresponds to the peak of the rainy season. As with other species in sect. Macroceratitae, Habenaria johannensis is typical of wet places, and grows in wet meadows, as well as in water along the margins of small rivers and lakes. Occasionally, the species can also colonize humid roadside banks and other man-made habitats. Illustrations.--Barbosa Rodrigues (1881, Fig. 2B), Warming (1884, plate 8, Fig. 5, as H. vaupellii), Schlechter and Hoehne (1922, plate 1, Fig. 4, as H. bradei), Hoehne (1940, plate 18), Brade (1951, plate 1, Fig. 2), Pabst and Dungs (1975, Fig. 14, column only), Toscano de Brito (1995, Fig. 46Js-Jt), illustration by Barbosa Rodrigues reproduced in Sprunger (1996, vol. I, Fig. 6B). Selected specimens examined: BRAZIL. BAHIA: 9 km S.W. of Mucuge, on the road from Cascavel, 950 m, 41~ 13~ 6 Feb 1974 (fl),Harley et al. 16086 (CEPEC, HB); 25 km WNW of Vila do Rio de Contas, 17 Feb 1977 (fl), Harley et al. 19597 (HB, SPF). DISTRITO FEDERAL: Zoobot~nico, 17 Jan 1967 (fl), A.P. Duarte 10173 (HB, RB); Brasflia, Rio Torto, near Sobradinho, 7 Jan 1966 (fl), Irwin et al. 11431 (HB, IAN, NY, UB). GoI.~S: about 28 km from Cavalcante, at the top of the Chapada, towards $5o Domingos region, 1 Jan 1999 (bud), Batista 836 (CEN); Mun. Pinheiros, Emas National Park, near Rio Formoso, 4 Dec 1994, Cesar et al. 226 (UFG). MATO GROSSO DO SUL: Mun. Rio Brilhante, fazenda Bela Vista, 24 Jan 1971, Hatschbach 26101 (HB, MBM, UPCB); Mun. Iguatemi, MS-295, Rio Panduf, 7 Feb 1993, Hatschbach et al. 58616 (MBM). MINAS GERAIS: Serra do Cipr, about 8.6 km after the crossroad to Morro do Pilar, km 137, towards Concei~5o do Mato Dentro, 13 Mar 1999, Batista & Bianchetti 908 (CEN); Monte Silo, 22 Jan 1981 (SP 200235); Serra de Caldas, 10 Oct 1987, Brandao 13196 (PAMG); 4 km SW of Extrema, BR-381, 25 Feb 1976, Davidse & Ramamoorthy 10545 (HB, SP); Mun. Extrema, border between Minas Gerais and S~o Panlo, 28 Jan 1996, Forzza 151 (SPF); between Monte Silo and Ouro Fino, 22 Jan 1973, Gottsberger s.n. (HB 63906); Mun. Tr~s Cora~res, Sao Tom6 das Letras, 4 Feb 1973, [VOL. 58 Hatschbach & Ahumada 31255 (HB, MBM); Serra do Congo Socco, 11 Jan 1921, Hoehne s.n. (SP 4891); Miguel Burnier, 30 Jan 1921, Hoehne s.n. (SP 30036); 40 km E. of Belo Horizonte, BR-31, 17 Jan 1971, Irwin et al. 30637 (NY, UB); Bastos, Rio das Velhas, "Schwacke expedition," Mar 1893, Magalhaes s.n. (OUPR); Mun. Formoso, Grande Sert~o Veredas National Park, 15~176 720 m, 17 Feb 1999, Mendonga et al. 3838 (IBGE); Conceiq~o do Mato Dentro, Parque Natural Municipal do Ribeir~o do Campo, 10 Feb 2003, Mota & Viana 1709 (BHCB); Belo Horizonte, ICB campus, 29 Jan 1980, Oliveira s.n. (BHCB 1264); Serra do Caraqa, 20 Mar 1957, Pabst 3216 (HB); Caetr, 20 Mar 1957, Pereira 2495 & Pabst 3331 (HB, RB); Ouro Preto, 1050 m, 5 Mar 1895 (fl), Schwacke 11472 (BR); S~o Roque de Minas, 17 km from the city, towards Piumhi, ribeir~o Cachoeira, 780 m, 20~ 46~ 14 Jan 1994, Souza et al. 5063 (SPF); Morro do Ferro, 7 km from the city on the road to Santiago, 20~ 44~ 878 m, 15 Jan 1994, Souza et al. 5100 (SPF); Pogos de Caldas, ALCOA-PO(~OS, 15 Mar 2000, Tameirao Neto 2991 (BHCB); Viqosa, E.S.A., 11 Feb 1965, Vidal 293 (VIC); Widgren 754 (K) (mixed with the holotype of H. gourlieana); Serra da Moeda, 26 Feb 1976, Windisch & Ghillany 444 (HB); Barbacena (R 28915). PARAN.~: Mun. Jundia/do Sul, fazenda Monte Verde, 28 Jan 1997, Carneiro 290 (MBM); S~o Jerrnimo da Serra, on the Sao Jerrnimo da Serra-Sapopema road, 11 Mar 1989, Cruz s.n. (FUEL 6759); desvio Ribas, 17 Feb 1911, Dusen 11575 (MBM); Mun. Tibagi, Santo Antonio, 31 Jan 1959, Hatschbach 5472 (HB, MBM); Mun. S~o Mateus do Sul, Rio Potinga, 9 Feb 1976, Hatschbach et al. 13808 (HB, MBM, SP); Mun. Ponta Grossa, Porto Sao Luis, Rio Tibagi, 25 Feb 1967, Hatschbach 16065 (MBM); Mun. Curitiba, Xisto highway, Rio Barigui, 21 Dec 1967, Hatschbach 18231 (MBM); Mun. Piraquara, Guarituba, 26 Dec 1977, Hatschbach 40715 (MBM, UEC); Mun. Jaguariafva, PR-11, Rio Jaguariafva, 27 Jan 1989, Hatschbach & Manosso 52678 (MBM); Mun. Senges, Rio Pelame, 17 Feb 1982, Kummrow & Stutts 1776 (MBM). RIO DE JANEIRO" Itatiaia, Benfica, 3 Feb 1921, Porto 1027 (RB, SP). RIO GRANDE DO SUL: Pelotas, 27 Jan 1950, Gilberto 38 (HB); Torres, near Trfis Forquilhas, 30 Jan 1952, Pabst 1340 (HB); Tupanciretan, Ivai, 28 Jan 1942, Rambo s.n. (PACA 9693); fazenda Ronda, near Vacaria, 10 Jan 1947, Rambo s.n. (PACA 34840). SANTA CATARINA" Florian6polis, 20 Feb 1974, Bresolin 1119 (HB); Sombrio, 28 Dec 1945, Reitz 1359 (HB, RB). $7,o PAULO: Pardinho, Rio Santo Ign~icio, 5 Mar 1974, Amaral J~. 1761 (BOTU); Paraibuna, Tamoios highway, SP-99, km 54, 4 Feb 1996, Borba 174 (CEN, UEC); 9 km S of Atibaia, 25 Feb 1976, Davidse & D'Arcy 10530 (SP); Vargem Grande do Sul, on the road to Casa Grande, 14 Nov 1997, Dutilh & Marcondes-Ferreira 14 (UEC 94941); Sao Paulo, Ipiranga station, Mar 1899, Edwall in Comm. Geogr. Geol. Sao Paulo 4057 (SP); Moji das Cruzes, 8 Mar 190?, Edwall & Porto s.n. (SP 28987); Angatuba, forestry service farm, 20 Feb 1966, Emmerich & Dressier 2803 (HB, R); 25 km NW from Moji-Gua~ti, Moji-Gua~6 experimental station, fazenda Campininha, 18 Jan 1977, Gibbs & Leitao-Filho 4294 (MBM, UEC); near Jaragu~t, 7 Aug 1946, W. Hoehne s.n. (SPF 13331); Monte Alegre, Amparo, 25 Mar 1943, Kuhlmann 331 (SP); Serra do Que- 2006] BATISTA ET AL." HABENARIA SECT. MACROCERATITAE (ORCHIDACEAE) bra Cangalhas, 15 Mar 1939, Kuhlmann & Gehrt s.n. (SP 40026, SPF 65056); S~o Jos6 dos Campos, 27 Feb 1962, Mimura 303 (SP); Campinas, Apr 1895, Novaes 541 (SP 28981); km 139 on the Sorocaba-Itapetinga road, 20 Jan 1968, Pabst 9081 (HB); Jarinu, km 68 of the highway from Jarinu to Campo Limpo Paulista, 15 Jan 2003, Pansarin & Mickeliunas 962 (CEN, UEC); Jundiai, Serra do Japi, 13 Feb 2003, Pansarin & Mickeliunas 990 (UEC); Sumarr, Feb 1997, Singer & Dittrich s.n. (MBM 226987); Mun. ,~guas de Prata, S~o Roque da Fatura district, on the road to Vargem Grande do Sul, 21~ 46~ 1170 m, 11 Jan 1994, Souza et al. 4998 (SPF); Queluz, Feb 1894, Ule 40 (R); Sales6polis, Serra do Mar, 12 Jan 1975, Windisch & Ghillany 259 (HB); Campos da Bocaina, 2000 m, 21 Apr 1975, Windisch & Ghillany 279 (HB). PARAGUAY. DEPT. CORDILLERA:Valenzuela, Mar 1942, Rojas 9585 (SP). DEPT. GUAIR~: Cordillera de Villa Rica, Jan 1905 (fl), Hassler 8703 (BM). DEPT. I~IEEMIaUCO:Tebicuary, Nov 1941, Rojas 9334 (SE SPF). H a b e n a r i a j o h a n n e n s i s can be distinguished easily from other species in the section by the distinct structure of the rostellum, with its prominent mid-lobe projected almost completely beyond the anthers (Figs. 1, 4). Additionally, the posterior petal segments are wider (4-7 mm) than in most other species in the section (usually 1--4 mm), and are overlapped by the non-reflexed lateral sepals (Fig. 5). The sepals are green and the petals and lip green-white or green-yellow. The examination of the isotype of H. bradei and of a photocopy of the holotype of H. vaupellii has confirmed that they are conspecific with H. j o h a n n e n s i s . H a b e n a r i a vaupellii and H. j o h a n n e n s i s were both described in 1881, but Cogniaux (1893) considered H. j o h a n n e n s i s a synonym of H. vaupellii and was followed by all subsequent authors for 103 years. However, according to Sprunger (1996) H. j o h a n n e n s i s should have priority, since it was published in May and H. vaupellii in August. Curiously, around the same period G a l e a n d r a lagoensis Rchb.f. & Warm, described in the same work as H. vaupellii, was considered Galeandra montana a synonym of Barb.Rodr., which was described in the same work as H. j o h a n n e n s i s . The transfer of H. j o h a n n e n s i s (as H. vaupellii) to the new genus Bertauxia (Szlachetko, 2004) is unwarranted. The two other species transferred to Bertauxia, H. rodeiensis Barb. Rodr. and H. nasuta Rchb.f. & Warm., have little similarity to each other 25 and with H. j o h a n n e n s i s and certainly do not belong to the same section within the genus. Additionally, one of the main diagnostic characters of the new genus, the well developed mid-lobe of the rostellum, is extremely variable in H a b e n a r i a and appears in several other species such as H. anisitsii Kraenzl., H. magniscutata Catling, H. pabstii J.A.N. Bat. & Bianchetti, and H. pratensis (Salzm. ex Lindl.) Rchb.f. These species differ widely between each other, and certainly belong to different sections of the genus. 4. HABENARIA LONGICAUDA Hook., Bot. Mag. 4, t. 2957. 1829. Kusibabella longicauda (Hook.) Szlachetko, Richardiana 4(2): 61. 2004. TYPE: GUYANA. Demerara, C. S. Parker s . n . (HOLOTYPE:K, image seen; ISOTYPE: K, image seen). (Figs. 1, 4, & 5). Semi-aquatic herb. S t e m tall, somewhat slender to robust, leafy, 70-134 • 0.4-1.3 cm including the inflorescence. L e a v e s 8-11, spreading, elongate, linear to linearlanceolate, lowermost sheath-like, largest around the center of stem, 11-22(-30) • 1.42.6 cm, upper leaves similar to the lower bracts. Inflorescence (8-)14-19(-30) cm long, few to many-flowered; bracts green, shorter than the pedicel and ovary, decreasing in size towards the apex of the inflorescence, lanceolate, 2.5-7 • 0.9-1.9 cm. F l o w e r s (1-) 4-15(-22), spreading, superposed, medium to large sized for the genus, greenish; pedicellate ovary green, slightly arched, 6.5-10 cm. Sepals green with the base light green to whitish; dorsal sepal oval-lanceolate, obscurely apiculated, 13-17 • 9-11 mm; lateral sepals obliquely oblong-lanceolate, strongly reflexed, forming a 90 ~ angle with the dorsal sepal, apiculate, 16-20 x 7.5-9 mm. Petals light green becoming greenish yellow with aging, the base whitish, bipartite; posterior segment linear, slightly falcate, apex connivent with the dorsal sepal, (12-)14-16 x 1.5(-2) mm; anterior segment reflexed, somewhat parallel to the lateral sepals, 1.21.5 times as long as the posterior segment, straighter, linear, (15-)20-28 • 0.8-1.5 mm. Lip light green becoming greenish yellow with aging, the base whitish, tripartite, strongly reflexed, convex; undivided basal region 4 - 8 • 3-4 mm; lateral segments, linear, 26 BRITTONIA 1.2-1.8 times as long as the middle segment, 18-28 • 1 mm; central segment a little shorter and wider, linear-ligulate, 14-20 x 12 mm. Spur whitish at the base, becoming green towards the apex, pendent, without a triangular tooth at the entrance of the orifice, usually partially arranged between the bracts, straightened towards the apical part, 1315.5(-20) cm, 2-3 m m in the base, 1 m m in the apical part. Column 6.5-9 m m tall; anthers whitish, apparently yellow because of the pollinia, slightly arched, 4.5-7 m m tall, canals white, base slightly thickened, ca. 1.5 m m wide, slender towards the apex, erect, 5 7 m m long; connective green, retuse to emarginate; rostellum light green, 9 m m long, mid lobe fleshy, triangular, small, 3.5-4 m m tall, completely enclosed between the anthers, apex subacute, side lobes erect, 6 - 7 m m long; auricles whitish, erect, fleshy, somewhat verrucose, triangular to hom-shaped, ca. 3-3.5 m m tall, obtuse; stigmas green, 6-7 m m long, margins curved, somewhat involute, apex not curved, truncate, 2.5-3 m m wide. Pollinaria 2, yellow, viscidia 0.8-0.9 • 0.5 mm, spaced 5.5-7.5 m m apart; caudicles elongate, ca. 8 m m long; pollinia 2 per pollinarium, laterally flattened. Etymology.--The name is a reference to the long spur of this species. Distribution.--Brazil (Bahia, Maranhao, Parfi, Tocantins); French Guiana; Guyana; Surinam; and probably also Venezuela. Habenaria longicauda is apparently common in northern South America, particularly in the Guianas, and reaches northern and northeastern Brazil only sporadically (Fig. 6). The illustration in Foldats (1969, Fig. 29) and identified as H. sartor undoubtedly represents H. longicauda and, if based on Venezuelan material, indicates that the species also occurs in Venezuela. Habenaria longicauda was collected in "northern central Brazil only recently, and most certainly should also be expected for the state of Mato Grosso. The species has been reported from the state of Amazonas (Renz 1992; Pabst & Dungs 1975), but the specimens cited (Prance et al. 8057, 13665), which are the only specimens of sect. Macroceratitae that we have seen from this state, are, in our opinion, referable to H. nabucoi. Habitat, Ecology, and Phenology.---Flow- [VOL. 58 ering in northern central Brazil occurs at the peak of the rainy season, from late December to early February. The species is typical of wet places, growing in water at the margins of lakes and streams. Illustrations.--Hooker (1829, t. 2957), Foldats (1969, Fig. 29, as H. sartor), Snuverink and Westra (1983, Fig. 9 and Fig. 10, as H. longicauda subsp, ecalcarata), Werkhoven (1986, p. 141, color plate). Additional specimens examined: BRAZIL. BAHIA: Barreiras, Acaba Vidas waterfall, 9 Feb 1999 (fr), Batista & Lemos 867 (CEN); same locality, 24 Jan 2000 (fl), Batista 1039 (BHCB, CEN); Barreiras, Acaba Vidas waterfall, 11~ 45~ 22 Jan 1998 (fl), Gongalves 177 (CEN, UB). MARANrlTLO: 50 km from Balsas towards Piauf, 17 Jan 1998 (fl), Oliveira & Gon~alves 282 (CEN, UB). PAR~,: Almeirim, Rio Jutahy, 16 Apr 1923 (fl), Ducke s.n. (RB 18720, SP 31494); Bel6m, Catfi, 7 Nov 1945, Pires & Black 547 (IAN); Rio Guarani, Utinga, igap6, below Bussuquara, 1 May 1947 (fl), Pires & Black 1538 (IAC, IAN, RB); Mun. Meuga~o, Ferreira Pena scientific station, Caxiuan~, 13 Oct 1995 (fl), Silva 476 (MG). TOCANTINS: Dianopolis, Nova Igua~u farm, 11~176 610 m, 25 Set 2003 (fl), Scarlot et aL 718 (CEN). In specimens with fully opened flowers, H. longicauda can be distinguished from other species in the section by the reflexed floral segments, including the anterior petal segments, the lip segments, and the lateral sepals which form a 90 ~ angle with the dorsal sepal (Fig. 5). In addition, the spur can be longer than in other species of the section, reaching up to 25 cm according to Snuverink and Westra (1983), and is apically acuminate. The flowers are green or green-white, becoming green-yellow with age. In central and northern Brazil, the distribution of Habenaria longicauda overlaps that of H. nabucoi, and in western Bahia we have found populations of both species flowering at the same time and separated from each other by no more than 10 km. It would be interesting to investigate the mechanisms of reproductive isolation of both species and to determine whether there is genetic exchange between them. Habenaria longicauda is the least collected and apparently the rarest species of sect. Macroceratitae in Brazil. However, this could simply be an artifact of sampling since its geographical distribution covers a rather large area that has been little collected. 2006] B A T I S T A E T A L . : H A B E N A R I A SECT. M A C R O C E R A T I T A E ( O R C H I D A C E A E ) 27 O t 1 '.~. ...),,; A"; . ..~, ....... k '~'~'-.~ ..r ,~ '. i .... i9, . , d . . L , f i .............. ....... ......... 3O / l ? .oo .o,o ,oo ~ . . . . 0 100 200 300 400 500 BOOmtln Pr~ GO / ,ooo,,., by t-tendr~k~ 4o / FIG. 6. Distribution of Habenaria longicauda (circle), H. macronectar (star), H. nabucoi (triangle), and H. paulistana (square). Based on Flora Neotropica base map no 1. 5. HABENARIA MACRONECTAR (Veil.) Hoehne, Engl. Jahrb. 68: 128. 1937. Orchis Vell., F1. Flum. 9, Tab. 45. 1831; Arch. Mus. Nac. Rio de Janeiro 5: macronectar 368. 1881. Kusibabella macronectar (Veil.) S z l a c h e t k o , R i c h a r d i a n a 4(2): 61. 2004. HOLOTYPE2 u n k n o w n , p r e s u m a b l y lost. LECTOTYPE h e r e d e s i g n a t e d , Veil. F1. F l u m . 9, t. 45. 1831. (Fig. 1) Habenaria sartor Lindl., London J. Bot. 2: 662. 1843. Kusibabella sartor (Lindl.) Szlachetko, Richardiana 4(2): 61. 2004. TYPE: BRAZIL. Rio de Janeiro, Organ Mountains, 1837 (fl), G. Gardner 676 (HOLOTYPE:K, image seen; ZSOT?PES:AMES, BM, BR, NY, US). Habenaria helodes Rchb.f., Linnaea. 22: 813. 1849. TYPE: BRAZIL. Rio de Janeiro, Neu-Freiburg, H.K. Beyrich s.n. (HOLOTYPE:W, image seen; ZSOTYPE: LE, n.v.). Habenaria dolichosceras Barb. Rodr., Gen. Sp. Or- 28 BRITTONIA chid. 1: 153. 1877. TYPE: BRAZIL. Minas Gerais, dans les champs humides de Caldas, et dans la Serra da Mantiquyra, J. Barbosa Rodrigues s.n. (No original material is known. Typified by Cribb & Toscano de Brito, 1996: 1: 30. Original illustration by Barbosa Rodrigues, reproduced in Sprunger, 1996: 1: 59A) Habenaria sartoroides Schltr., Repert. Spec. Nov. Regni Veg. 16: 248. 1919. TYPE: BRAZIL. Paran~i, Serra do Mar, Banhado, 3 Mar 1914 (fl), P K . H . Dusen 14571 (HOLOTYPE: B, destroyed; LECTOTYPE: here designated, AMES). Habenaria juergensii Schltr., Repert. Spec. Nov. Regni Veg. Beih. 35: 20. 1925. SYNTVPES: BRAZIL. Rio Grande do Sul, Municfpio Rio Pardo, in etwas sumpfigem Boden der Campos bei Jo~o Rodrigues, 50 m, Feb 1921, C. Juergens 23 (B, destroyed; no duplicate located); San Leopoldo, Fazenda dos Prazeres, 25 Jan 1904 (fl), J. Dutra 673 (SVNTYPE:B, destroyed; LECTOTYPE:here designated, SP). Terrestrial herb. Stem variable, usually short to medium height, but also tall and robust, leafy, 30-70(-112) x 0.3-0.7 cm, including the inflorescence. Leaves ( 6 - ) 8 - 1 0 ( 12), spread, linear-lanceolate to lanceolate, largest around the center of the stem, 6.5-25 x 1.2-3(--4.2) cm, upper leaves similar to the lower bracts. Inflorescence (6.5-)12-30 cm long, few to many flowered; bracts slightly longer to shorter than the pedicel and ovary, decreasing in size towards the apex of the inflorescence, ovate-lanceolate to lanceolate 3 4(-8.5) x 1-2.4 cm. Flowers (3-)6-18(-30), superposed, large to medium for the genus, greenish white to greenish yellow; pedicellate ovary green, 3.5-6.3 cm. Sepals green; dorsal sepal largely lanceolate, apiculate, 1318 • 8-11 mm; lateral sepals deflexed, forming a 180 ~ angle with the dorsal sepal, obliquely ovate-lanceolate, apiculate, 15-21 x 6-7(-9) ram. Petals light green, greenish yellow or greenish white, bipartite; posterior segment linear-lanceolate, slightly falcate, 14-18 x 1-2 ram, the apical part conivent with the dorsal sepal; anterior segment projected forward, somewhat parallel to the lip lateral segments, 1.2-2 times as long as the posterior segment, straighter, linear, 20-28 x 0.5-1 mm. Lip light green, greenish yellow or greenish white, tripartite, projected forward; undivided basal part 3-6 x 1-2 mm; lateral segments somewhat concave, 1.2-1.7 times as long as the central segment, linear, 18-35 x 1 ram; central segment shorter and slightly wider, linear-ligulate, 11-22 • 1.5- [VOL. 58 1.8 mm. Spur whitish at the base, greenish towards the apical part, with a small erect triangular tooth at the entrance of the orifice, pendent, the apical part hidden between the bracts, linear, the basal part with the same width or slightly wider than the apical part, 4.8-8.5 x 0.1-0.2 cm. Column 5-6 mm tall; anthers spaced, ca. 4 mm tall, canals, erect, 4-5 mm long; connective broad, retuse; rostellum fleshy, ca. 7 mm long, mid lobe triangular, short, 2-2.5 mm tall, completely enclosed between the anthers, acute, side lobes 4.5-5 mm long; auricles, erect, fleshy, verrucose, triangular, flattened, acuminated, 2 mm tall, acute; stigmas 9-10 mm long, the free receptive part 6-7 mm long, margins curved, somewhat involute, apex not curved, 1.5-2 mm wide. Pollinaria 2, yellow, viscidia 0.8 • 0.6 mm, spaced 2-3 mm apart; caudicles elongate, ca. 4 mm long; pollinia 2 per pollinarium, laterally flattened. Etymology.--Named after the long spur. Distribution.--Brazil (Espirito Santo, Minas Gerais, Paran~i, Rio de Janeiro, Rio Grande do Sul, S~o Paulo, Santa Catarina); and Uruguay. Habenaria macronectar has been reported from Paraguay (Cogniaux, 1903, 1905; Hoehne, 1940; Brade, 1951) and Argentina (Pabst & Dungs, 1975), and although the species may occur in these countries, further investigation is necessary to verify these reports. The material identified as H. macronectar from these countries that we have seen in Brazilian herbaria was too poorly conserved to allow a definite conclusion. The species has also been reported from Suriname by Reichenbach f. (1859), Kraenzlin (1892) and Cogniaux (1893), but the material identified as H. sartor (Splitgerber 754) is, according to Snuverink and Westra (1983), referable to H. longicauda. Habitat, Ecology, and Phenology.--The species is typical of wet places, growing in humid soil, as well as in water at the margins of streams, small lakes, and ponds. Occasionally, the species can also occupy humid roadsides and other artificial habitats. Flowering throughout the distribution range occurs mainly from January to March. Throughout most of its distribution in Brazil, the species is typical of high altitude mountain meadows, growing at altitudes over 800 m. In the state of Rio Grande do Sul, in southern Brazil, 2006] B A T I S T A E T AL." H A B E N A R I A SECT. MACROCERATITAE b e l o w p a r a l l e l 3 0 ~, t h e s p e c i e s is a l s o f o u n d a t l o w e r a l t i t u d e s g r o w i n g at a l m o s t s e a l e v e l . lllustrations.--Vellozo ( 1 8 3 1 , Fig. 4 5 ) , C o g n i a u x ( 1 8 9 3 , p l a t e 5, Fig. 1, as H. s a r t o r , r e p r o d u c e d f r o m t h e B a r b o s a R o d r i g u e s ill u s t r a t i o n o f H. d o l i c h o s c e r a s ) , M a n s f e l d ( 1 9 3 0 , p l a t e 6, Fig. 2 2 , as H . s a r t o r o i d e s ) , H o e h n e ( 1 9 4 0 , p l a t e 14, as H. h e l o d e s ; p l a t e 16, r e p r o d u c e d f r o m t h e B a r b o s a R o d r i g u e s i l l u s t r a t i o n o f H . d o l i c h o s c e r a s ; p l a t e 19, as H . j u e r g e n s i i ) , B r a d e ( 1 9 5 1 , Fig. 1), P a b s t a n d D u n g s ( 1 9 7 5 , p a g e 177, Fig. 9; p a g e 2 4 2 , Fig. 9, as H . ( a s t o r , b a s e d o n A m a r a n t e 0 8 , R B ) , A l v e s ( 1 9 9 1 , p a g e 111, as H . ( a s t o r , anomalous material, based on Kolbek & Alves 1000, RB), Miller and Warren (1994, p a g e 33, lip o n l y a n d p l a t e 1, c o l o r p l a t e , as H. (a s t o r ) , illustration o f H. dolichosceras b y Barbosa Rodrigues reproduced in Sprunger ( 1 9 9 6 , vol. I, Fig. 6 A , as H . s a r t o r ) . Selected specimens examined: BRAZIL. ESPIRITO SANTO: Pedra Azul, 12 Feb 1970, Kautsky 228 (HB); Castelo, Foruo Grande, 26 Jan 1973. without collector (MBML 474). MINAS GERAIS: Ouro Preto, Parque Estadual do Itacolomi, 15 Feb 2003, Batista et al. 1378 (CEN): Caldas, Pedra Branca, 19 Jan 2001, Borba et al. s.n. (UEC 118854); Carandai, km 418 from E.EC. do BrasiL 15 Feb i962, Duarte 6330 (HB, RB); Morro do Ferro, near Polos de Caldas, 31 Mar 1968, Emmerich 3175 (HB); Serra do S~o Jos6, 20 Sept 1989, Kolbek & Alves I000 (RB); Pacau, Serra da Mantiqueira, Feb 1913, Lutz 415 (R); Alagoa, ca. 4 km from the Quaresmeira site, 3 Mar 2003, Pansarin & Mickeliunas 1006 (CEN); Santos Dumont, 15 Mar 1957, Pereira 2340 & Pabst 3174 (HB, RB); Caldas, 1 Mar 1877, Regnell 111-1186 (BR, SP); Ant6nio Carlos, 10 Feb 1950, Roth 16475 (CESJ); Serra da Mina, 12 km from Bom Jardim de Minas, 4 Mar 1964, Saleh I08 (HB); Mun. Barroso, BR265, Lavras-Barbacena, km 130, 21~ 44~ 1020 m, 5 Feb 1989, 8cariot et al. 344 (CEN, IAC, SPF); Estouro, Morro do Brigadeiro, 13 Feb 1985, Vidal et aL 609 (VIC). PARAN~{; Mun. Colombo, Santa M6nica Field Club, 8 Feb 1984, Bidd et aL 283 (UPCB); Serra da Esperanqa, 17 Feb 1949, Brade 19677 (HB, RB); Ipiranga, 9 Feb 1904, Dusen 3514 (R, SP); Pinhaes, 23 Jan 1904, Dusen s.n. (R 3503); Banhado, 1914, Dusen s.n. (MBM 253377); Mun. Curitiba, Pinhaes, 31 Jan 1947, Hatschbach 61I (MBM, RB); Piraquara, 22 Jan 1953, Hatschbach 3006 (MBM); S~o Jos6 dos Pinhaes, Matulfio, 2 Mar 1957, Hatschbach 3813 (MBM); Mun. Tijucas do Sul, Saltinho, 15 Nov 1957, Hatschbach 3818 (MBM); Mun. Morretes, Pilao de Pedra, 27 Feb 1965, Hatschbach 12387 (MBM); Mun. General Carneiro, Rio Iratin, Hatschbach 13679 (MBM); Mun. Guarapuava, Guarfi, 18 Jan 1968, Hatschbach 18320 (HB, MBM, UPCB); Mun. Imicio Martins, Monte Alto, 21 Jan 1998, Hatschbach et al. 67492 (MBM); 47 km from Horizonte. road to Palmas, BR-280, 1250 msm, 28 Jan 1985, Krapovickas & CristObal 39723 (MBM); Mnn. Porto (ORCHIDACEAE) 29 Amazonas, fazenda Sat Roque, 3 Feb 1976, Kummrow 1068 (MBM, UEC); Mun. Araucaria, Tindiquera, 11 Feb 1962, Leinig 279 (HB); Bituruna. 9 Feb 1948, MelloFilho 758 (R); Maringfi, without date and collector (CEN, HB 75755). RIo D~ JANEtRO: Nova Friburgo, Morro da Cruz, Dec 1935, Amarante 08 (RB); Friburgo, Jan 1928, Freire 293 (R, SP); Alto Macah~, 1878 (fl), Glaziou 6916 (BR); Organ Mountains, 1839 (ill Miers 2248 (BM); Teres6polis, Jan 1888 (fl), Moura 68 (BR); Mun. Petr6polis, km 20 on the Itaipava-Teres6polis road, 24 Feb 1963, Pabst 7305 (AMES, HB); Serra dos Orgaos, 13 Apr 1944, Pereira s.n. (HB 5572); Itatiaia, trail to inveruada Mau~i, 5 Feb 1925, Porto 1522 (RB, SP). Rio GRANDE DO SUL: Vila Oliva, near Caxias, Feb 1945, Buck s.n. (PACA 28069); Lageadinho, near Born Jesus, 20 Jan 1958, Camargo 3110 (PACA); Caracol, near Canela, 30 Jan 1936, Emrich s.n. (PACA 26858); Canoas, 17 Jan 1949, Miguel s.n. (SE SPF 14303); Morro da Policia, near Porto Alegre, 6 Jan 1933, Orth s.n. (PACA 568); near TrEs Forquilhas, Osorio Torres, 30 Jan 1952, Pabst 1339 (HB); between Canela and Born Jesus, 37 km from Canela, 900 msm, 10 Feb 1952, Pabst 1355 (HB, MBM, RB); Passo Fundo, 28 Jan 1964, Pereira 8687 & Pabst 7962 (HB, RB); Pestana, near Iju/, 7 Feb 1956, Pivetta 1179 (PACA); near S~o Francisco de Paula, 8 Feb 1941, Rambo s.n. (PACA 4434); near Quaraf, fazenda Jarau, Jan 1945, Rambo s.n. (PACA 26230); Nonoai ad ft. Uruguay, Mar 1945, Rambo s.n. (PACA 28514); Cambarfi, near S~o Franscisco de Paula, Feb 1948, Rambo s.n. (PACA 36566); Cachoeirinha, near Gravataf, 7 Jan 1949, Rambo s.n. (PACA 39560); Jaquirana, near S~o Francisco de Paula, 20 Feb 1952. Rambo s.n. (HB 1729, PACA 52107); Pelotas, 27 May 1959, Sacco 1288 (HB); S~o Leopoldo, 1907, Theissen 629 (PACA 7470). SANTA CATAmNA: Lages, 1935, Bruxel s.n. (PACA 6827, PACA 6834); Sat Joaquim, 20 Jan 1958, Mattos 5008 (HB); near Ponte Alta, on the road to Rio do Sul, 29 Jan 1964, Pereira 8727 & Pabst 8002 (HB); Santa Terezinha do Boqueirao, Lages, 1 Feb 1963, Reitz 6462 (HB); Serra do Oratorio, Born Jardim, S~o Joaquim, 19 Feb 1959, Reitz & Klein 8437 (HB); Porto Uni~o, 25 Feb 1962, Reitz & Klein 12408 (HB); Mun. Campos Novos, BR-470, fazenda Carazinho, 11 Feb 1996, Ribas et aL 1251 (MBM); Mun. Campo Alegre, Serra Quiriri, margins of the Rio Negro, 11 Feb 2001, Ribas & Silva 3278 (MBM); Mun. Caqador, 7 Feb 1957. Smith & Klein 10989 (RB). S~o PAULO: Serra da Bocaina, fazenda Bonito, 15-30 Jan 1913, Lutz 366 (R); Campos do Jordao, Mantiqueira, 10--12 Feb 1921, Lutz & Lutz 1615 (R); between Santo Andr6 and Nova Utinga, 18 Jan 1952, Pabst 1314 (HB); Atibaia, Pedra Grande, 18 Feb 2004, Pansarin & Mickeliunas 1128 (CEN, UEC); Cunha, forest reserve, 13 Feb 1981, Wanderley 284 (SP, UB). URUGUAY. DEPT. R f o NEGRO: Feb 1914, Berro 7299 (HB 1200). DEPT. SAN JOSE: San Jos6, 23 Feb 1907, Berro 3793 (HB). H a b e n a r i a m a c r o n e c t a r a n d H. bractescens can be separated from the other Brazili a n s p e c i e s i n sect. M a c r o c e r a t i t a e b y t h e s h o r t spur, u s u a l l y ca. 5 - 6 ( - 8 . 5 ) c m l o n g ( F i g . 1). A p a r t f r o m t h i s c h a r a c t e r i s t i c , b o t h species show few similarities and can be sep- 30 BRITTONIA arated by the color of the flowers (white in H. bractescens, vs. green in H. macronectar), the position of the lateral sepals (non-reflexed vs. strongly deflexed), and the lateral segments of the petals and lip (about the same size and width as the posterior petal and lip lateral segments vs. longer and narrower). In Brazilian herbaria, Habenaria macronectar has been most frequently confused with H. gourlieana and H. nabucoi (identified as H. fastor), but can be distinguished from both species by the shorter spur and the deflexed lateral sepals. Curiously, some specimens of H. m a c r o n e c t a r from the states of Espirito Santo ( M B M L 474) and Minas Gerais (Kolbek & A l v e s 1000, RB) have bi- or tripartite apex of the lateral segments of the lip. The flowers of H. macronectar are green or green-white, becoming green-yellow with age. We follow here Hoehne's (1940) position to consider Vellozo's epithet as that valid for the species since, according to article 44.1 of the International Code of Botanical Nomenclature (Greuter et al., 2000), an illustration with analysis published before 1908 is equivalent to a diagnosis and, according to Carauta (1973), the effective date of publication of Vellozo's plate was 1831, four years earlier than Lindley's description of H a b e n a r i a sartor. Vellozo's plate is somewhat schematic, but the deflexed lateral sepals, the "short" spur and the origin of the plant, from the state of Rio de Janeiro, where H. macronectar is the only "short-spurred" species of the sections known to occur in the state, leaves no doubt about the identity of the species. Examination of an image of the type of H. helodes Rchb.f., also from the state of Rio de Janeiro, showed that it is not specifically different from H. macronectar. A critical analysis of the original description and illustration of H. dolichosceras Barb. Rodr., and examination of a isotype of H. sartoroides Schltr. and a syntype of H. j u e r g e n s i i Schltr. did not reveal any specific difference when compared to H. macronectar. For instance, in all these species the sepals are deflexed, the petal anterior segment are slender and longer than the petal posterior segment and the spur is 5.3-6 cm long. Accordingly, H. dolichosceras was treated as a synonym of H. sartor [VOL. 58 (Cogniaux, 1893), and H. dolichosceras and H. sartoroides as synonyms of H. macronectar (Hoehne, 1940; Brade, 1951). Pabst and Dungs (1975) considered H. j u e r g e n s i i a synonym of H. gourlieana, but the spur about 5.5 cm long and the petal anterior segment 1.9 times as long as the petal posterior segment leave no doubt that H. j u e r g e n s i i is referable to H. macronectar. 6. HABENARIA NABUCOI Ruschi, Bol. Mus. Biol. Prof. Mello-Leitao, ser. Bot. 78: 1. 1973. TYPE: BRAZIL. Espirito Santo, in campis humidis ad Serra, 20 m , Jan 1973 (fl), E. Colnago s.n. (HOLOTYPE: MBML, photo seen). (Figs. 1, 2, 4) gourlieana Gill. ex Lindl. f. magna Hoehne, Comm. Linhas Teleg. Estrateg. Matto Grosso Annexo 5, Bot. 9: 25, Tab. 168, Fig. 3. 1919. SYNTYPES: BRAZIL. Mato Grosso~ margins of a Rio Arinos waterfall, Nov 1914 (fl), J.G. Kuhlmann 124 & 125 (SYNTYPES: R). Habenaria fastor Warm. ex Hoehne, Flora Brasilica 12(1): 71, Tab. 15. 1940. Kusibabella fastor (Warm. ex Hoehne) Szlachetko, Richardiana 4(2): 59. 2004. SYNTYPES: BRAZIL. $5o Paulo, Paranapanema valley, ,~gua Boa lake, 3 Dec 1899 (fl), A. Habenaria Loefgren in Comm. Geogr. Geol. S~o Paulo s.n. (SYNTYPE: SP 28977); Minas Gerais, Mun. Santa Luzia, Lagoa Santa, 5 Feb 1935 (fl), M. Barreto 4869 (SYNTYPES:BHMH, R, RB, SP). Semi-aquatic herb. Stern tall, robust, leafy, 64-128 x 0.5-1.3 cm including the inflorescence. L e a v e s 8-12, spread, narrow, linear, elongated, lowermost sheath-like, largest around the center of stem, 9-23(-45) x 1.21.8(-2.6) cm, upper leaves similar to the lower bracts. Inflorescence 9 - 1 9 cm long, short, usually few flowered; bracts lanceolate to ovate-lanceolate, 4.3-6.5 x 1.1-2.1 cm, shorter than the pedicel and ovary, decreasing in size towards the apex of the inflorescence, green. F l o w e r s 2-8(-1 l) spreading, superimposed, large for the genus, whitish; pedicellate ovary 5-8.5 cm long. Sepals mucronate, externally greenish white with the margins white and the veins marked in green, internally white or creamy white with the veins marked in light green; dorsal sepal ovate to ovate-lanceolate, 20-23 x 10-15 mm; lateral sepals, obliquely oblong-lanceolate, 21-30 x 9-13 mm, non-reflexed. Petals bipartite, completely white; posterior segment linearlanceolate, falcate, 19-25 x ( 2 - ) 3 - 4 mm, 2006] BATISTA ET AL.: HABENARIA SECT. MACROCERATITAE (ORCHIDACEAE) 31 apex connivent with the dorsal sepal; anterior segment linear, 30-43 x 1-1.8 mm, 1.4-1.9 times as long as the posterior segment, nonreflexed, projected forwards or downwards, somewhat parallel to the lateral segments of the lip. Lip tripartite, projected forwards or downwards, white; undivided base 5-8 • 2-3 ram; lateral segments linear, 25-38(-45) x 1.5(-2) mm, 1.3-1.7 times as long as the medium segment; medium segment shorter and slightly wider, linear-ligulate, ( 1 9 - ) 2 4 32 • (1.5-)2,5(-3) ram. Spur pendent, with a small erect triangular tooth at the base, tooth ca. 0.8-1 mm tall, acute, the apex sometimes free from the bracts, base narrow, thickened towards the apex, 11-15.5 cm long, base 1.52 mm wide, near the apex (2-)3(-4) mm wide, base white and becoming greenish towards the apex. Column 8-11 mm tall; anthers slightly arched, 7 - 9 • 3 4 ram, whitish, apparently yellow because of the pollinia, canals slender, erect, 7 - 1 0 nun long; connective light green, apex variable but always prominent, obtuse to slightly apiculate; rostellum ca. 11 mm long, creamy white, midlobe large, prominent, fleshy, triangular, 8-9 m m tall, base 5-6 m m wide, about the same size as the anthers, partially projected beyond the anthers, acuminate, acute; rostellum arms slender, erect, ca. 7 m m long; auricles erect, triangular, 4-5 m m tall, fleshy, somewhat verrucose, acuminate, acute, white; stigmas ca. 12 mm long including the base, the free part 7-9 mm long, involute, apex not curved, truncate, 3-4 mm wide, creamy white. Pollinaria 2, yellow, viscidia ca. 1 • 0.8 mm, spaced 6.5-10 mm apart; caudicles elongate, ca. 10 mm long; pollinia 2 per pollinarium, laterally flattened. E t y m o l o g y . i N a m e d after Jo~o Mauricio Nabuco, director of the Brazilian Institute for Forest Development (IBDF), and responsible for the federal environment policy at the time the species was described. D i s t r i b u t i o n . I B r a z i l (Amazonas, Bahia, Distrito Federal, Espirito Santo, Mato Grosso, Minas Gerais, Piaui and S~o Paulo); probably also in Venezuela. The extent of the geographical distribution and the relationship of H. nabucoi with other extra-Brazilian long-spurred plants reported from Paraguay (Cogniaux, 1903, 1905), Peru (Schweinfurth, 1958; as H. sartor), Colombia (Hernfindez, 1958; as H. bractescens) and Bolivia (Schlechter, 1922) are still unclear. The illustration presented by Foldats (1969, Fig. 30) and identified as H. sartor is undoubtedly referable to H. nabucoi. If this illustration is based on a Venezuelan plant, this further indicates that the species also occurs in Venezuela. We have not seen any material of H. nabucoi from Pernambuco, but the collection Gardner s.n. mentioned by Kraenzlin (1892) and identified as H. sartor is most probably referable to H. nabucoi since H. macronectar does not reach northeastern Brazil. Habitat, Ecology, and P h e n o l o g y . i I n central Brazil, H. nabucoi flowers during the peak of the rainy season, mainly from December to February. This is somewhat later than for some other species in sect. Macroceratitae, such as H. gourlieana and H. bractescens, which flower early in the rainy season. Habenaria nabucoi is typical of very wet places and, in agreement with Warming's notes (Warming, 1884), we have found the species growing partially immersed in water at the margins of lakes and small streams. Plants of H. nabucoi usually occur in groups, and can develop a lateral root-like appendage, with a condensed shoot at the apex, which grows underwater and can reach at least 1-2 m long, suggesting that vegetative propagation is a common feature of the species. I l l u s t r a t i o n s . i W a r m i n g (1884, plate 8, Fig. 4, as H. fastor), Hoehne (1919, plate 168, Fig. 3, as H. gourlieana vat. magna), Foldats (1969, Fig. 30, as H. sartor), Ruschi (1973, pages 2-3), Pabst and Dungs (1975, Fig. 12, as H. longicauda, based on Prance 8057, HB), Ruschi (1986, page 55, as H. longicaudata, color plate). Additional specimens examined: BRAZIL. AMAZONAS:Mun. Labrea, 3 km N of Labrea, Lago Preto, Rio Punts, 29 Oct 1968 (fl), Prance et al. 8057 (HB, INPA, NY); Rio Punts, S of Lago Preto, 2 km N. of Labrea, 25 Jun 1971 fir), Prance et al. 13665 (HB, INPA). BAHIA: Barreiras, Rio Branco, about 90 km NW of Barreiras, 29.5 km after the crossroad with BR-020, on the road towards Cotia, 9 Feb 1999 (fl), Batista & Lemos 869 (CEN, SP, UEC), 24 Jan 2000 (fl), Batista 1040 (CEN); Mun. Marac~is, about 0.5-1.0 km N of Marac~is, towards Planaltino, 2 Jul 1993 (ill Queiroz & Fraga 3303 (HUEFS); km 5-10 on the Nilo Peqanha-Ituber~i road, 19 Feb 1975 (fl), Santos 2859 (CEPEC. HB); Mun. 32 BRITTONIA Itacarr, km 19 on the Ubaitaba-Itacar6 road, 14 Feb 1978 (fl), Santos 3167 & Mattos Silva 120 (CEPEC); Ubafra, outskirts of Trrs Braqos, Jan 1991 (fl), Sobral et al. 6700 (MBM); Maricas-Ituruqu, 29 Jan 1977 (fl), Windisch & Ghillany 645 (HB). DISTRITO FEDERAL: Planaltina~ Mestre D'Armas lake, 19 Apr 1979 (fr), Ashton s.n. (CH 829, HEPH 3325); Brasflia, Crrrego do Acampamento, 19 Jan 1992 (fl), Batista 253 (CEN); Planaltina, Reserva Ecol6gica de Aguas Emendadas, 6 Jan 1999 (fl), Batista & Oliveira 848 (CEN); Lagoa Bonita, 23 Nov 1980 (ill Bianchetti 466 (HEPH); Lagoa Bonita, 30 km NE of Brasflia. 21 Jan 1983 (fl), Rarnos 179 (CEN, HEPH, UEC). ESPIRITO SANTO: Mun. Serra, near Rel6gio, 100 m, 30 Jan 1973 (fl), Colnago & Kautsky 395 (HB); Linhares, CVRD nature reserve, Jueirana Vermelha road, Rio Barra Seca, 5 Jan 1995 (fl), Folli 2457 (CVRD); Mun. Linhares, Urussuquara, Suruaca valley, 24 Feb 2000 (fl), Fraga 580 (CEN, MBML); Serra, 30 km N of Vit6ria, 24 Feb 1973 (fl), Lagasa 162 (MBML). MATO CROSSO: Silva 399 (CH). MINAS GERAIS: 40 km from Paraopeba, Lagoa Preta, margins of the Rio Paraopeba, Apr 1958 (fr), Heringer 6383 (HB, HUFU, UB); Paraopeba, 1959 fir), Heringer 10039 (HB); Mun. Marlieria, parque florestal do Rio Doce, margins ofD. Helvecio lake, 26 Jun 1983 (fl), Ikusima & Gentil s.n. (SP 192356); Mun. Formoso, Parque Nacional Grande Sert~o Veredas, vereda at Santa Rita stream, 710 m, 15~ 45~ 15 Feb 1999 (fl), Sevilha et al. 1802 (IBGE). PIAUf: Br-135, near "Redenq~o do Gurgu6ia," 09~ 44~ 22 Jan 1998 (fl), Gonqalves 175 (CEN). SXo PAULO: MojiGuaqu, Reserva do Instituto Florestal, fazenda Campininha, Mar 2000 (fl), Amaral et al. 2000/38 (UEC). This species first appeared in the literature as Habenaria fastor Lindl. in Warming's (1884) account on the flora of Lagoa Santa, Minas Gerais, Brazil. As mentioned by Hoehne (1919, 1940), the name H. fastor most certainly arose from a typographical error, since Lindley never described H. fastor, but H. sartor. The identification as H. fastor in Warming's work (1884) was probably made by Reichenbach f., since the identification of Warming's orchids was made by that author (Reichenbach f., 1881). At that time, despite extensive differences in the size of some of the flower parts, particularly the spur, Reichenbach f. (1859), followed by Kraenzlin (1892, 1901) and Cogniaux (1893), included specimens of H. longicauda and H. nabucoi under H. sartor. Hoehne (1919) was apparently the first author to notice that the long spurred (11-15.5 cm) plants of the Warming collection from Lagoa Santa were distinct from the shorter spurred H. sartor (4.8-8.5 cm). Since Warming (1884) clearly did not intend to describe [VOL. 58 H. fastor as a new species, Hoehne (1940) tried to validate the species by describing it as H. fastor Warm. ex Hoehne. However, a Latin diagnosis was not presented and hence, according to article 36.1 of the International Code of Botanical Nomenclature (Greuter et al., 2000), the name is not validly published. From the description and listing of the specimens examined, Hoehne (1919, 1940) included under H. fastor, in addition to the species represented by the Warming collection from Lagoa Santa, two other species with long spurs, H. gourlieana and the taxon now described as H. paulistana. Consequently, the protologue of H. fastor is a mixture of three distinct species: H. nabucoi, H. gourlieana, and H. paulistana. The species referable to H. fastor sensu Warm. was only validly described as a distinct species by Ruschi in 1973 as H. nabucoi. Unfortunately, Ruschi did not perceive the extent of the question and only compared his new species with H. macronectar, despite extensive similarities with the previously described H. fastor Warm. ex Hoehne. Habenaria nabucoi can be easily distinguished from H. macronectar by the more robust habit, the larger and pure white or creamy white flowers (vs. smaller and predominantly green in H. macronectar), the non-reflexed lateral sepals (vs. deflexed), the spur 11-15 cm long (vs. 5-8.5 cm), and geographical distribution ranging from southeastern Brazil, to west central, northeastern and northern Brazil, apparently reaching Venezuela (vs. restricted to southern and southeastern Brazil, also reaching Uruguay). Habenaria nabucoi can be distinguished from other similar long-spurred species in sect. Macroceratitae, such as H. longicauda, H. paulistana, and H. gourlieana, by the white flowers, non-reflexed lateral sepals (except for H. gourlieana), and the spur that is broadened at the apex (Figs. 1, 2). Also, the column structure of H. nabucoi is distinguished by the longer and slender anther canals and rostellum arms and the taller rostellum mid-lobe, which is partially projected beyond the anthers (Figs. 1, 2, 4). In some specimens of H. nabucoi the apex of the rostellum mid-lobe exceeds the apex of the anthers. 2006] BATISTA ET AL.: HABENARIA SECT. MACROCERATITAE (ORCHIDACEAE) 33 ment linear to linear-lanceolate, falcate, 1822 • (1.5-)2(-3) mm, base free and towards the apex connivent with the dorsal sepal; anterior segment linear, 25-35 • 1-1.5(-2) mm, projected forward, 1.3-1.7 times as long as the posterior segment. Lip tripartite, projected forward, whitish at the base and light green to greenish yellow in the segments; undivided base 6 - 9 x 1.5-2 mm; lateral segments linear, 25-36 • 1-1.5(-2) mm, somewhat parallel to the anterior petals, 1.3-1.6 times as long as the middle 7. H a b e n a r i a paulistana J. A . N . Batista & segment; medium segment shorter and Bianchetti, sp. nov. TYPE: BRAZIL. S~o slightly wider, linear-ligulate, 20-26 x 1Paulo, Municipality of Cunha, margin of 2(-2.5) ram. Spur pendent, enclosed by the Rio Parafba, along the road towards Rio bracts, with a small erect triangular tooth ca. Bonito trail, 23~ 45~ 14 1 m m tall at the base, tooth obtuse, wider at Dec 1996 (fl), J. P Souza, E C. Souza, R. R. the base and straighter towards the apex, 11.5-13 cm, 3-4.5 m m in the base, ca. 2 Rodrigues, J. Y Tamashiro & N. M. Ivanauskas 875 (HOLOTYPE: CEN; ISO- m m in the apex, base whitish and becoming green towards the apex. Column 7-8 mm TYPES: ESA, SE SPE SPSE UEC). (Figs. 4, 5, 7) tall; anthers 4-5 x 3 - 4 mm, whitish, apparently yellow because of the pollinia; antherHabenariae nabucoi Ruschi affinis sed inflorescentia frequenter multiflora, floribus viridibus, sepalis later- canals erect, 4 - 6 • 1 mm, thickened, white; aliter deflexis, calcari apicem versus recto, columnae connective broad, truncate to retuse, green; structura antheris brevioribus, lobo mediano rostelli bre- rostellum ca. 9 mm long; middle lobe 3.5-5 viori et omnino per antheras incluso et rostelli brachiis mm tall, fleshy, triangular, acute, small, crassis differt. Etiam H. macronectari (Vell.) Hoehne about half the size of the column, comaffinis sed floribus majoribus et calcari longiore differt. pletely enclosed by the anthers, green; rosTerrestrial herb. Stem tall, robust, leafy, tellum arms erect, ca. 6 mm long, white84-146 x 0.7-0.9(-1.4) cm including the in- green; auricles erect, triangular, 2.5-3 m m florescence. Leaves ca. 12, spreading, tall, fleshy, verrucose, white; stigmas 2, 1112 m m long including the base, the free part linear-lanceolate to lanceolate, lowermost sheath-like, largest around the center of 6 - 8 m m long, margins curved, somewhat instem, 9-27 • 1.3-3.8 cm, upper leaves simi- volute, whitish, apex not curved, truncate, 2-3.5 mm wide. Pollinaria 2, yellow, vislar to the lower bracts. Inflorescence 19-28 cm long, usually elongated and many flow- cidia ca. 1.3 x 0.8-1 mm, spaced 1-1.5 mm ered; bracts oblong-lanceolate to lanceolate, apart; caudicles slender, elongate, 5 - 6 mm 3.3-5.5 • 1-1.8 cm, shorter than the pedicel long; pollinia 2 per pollinarium, laterally and ovary, decreasing in size towards the flattened, ca. 4 m m long. apex of the inflorescence, green. Flowers E t y m o l o g y . I T h e new species name is a (5-)10-17(-27), spreading, superimposed, reference to the state of S~o Paulo, to which large for the genus, greenish to greenish yel- it is restricted. low; pedicellate ovary 5.2-7.5 cm, slightly Distribution.--Brazil (S~o Paulo). So far, arched, green. Sepals mucronate, green; the species is known only from the state of dorsal sepal ovate to ovate-lanceolate, 17- $5o Paulo (Fig. 6). Although it is very pos23 • 9-13 mm; lateral sepals deflexed, sible that the species may be found in the forming a 180 ~ angle with the dorsal sepal, neighboring states of Minas Gerais and obliquely ovate-lanceolate to largely lanceo- Paran4, we have so far not seen any specimen late, 22-27 x 8-11 mm. Petals bipartite, of H. paulistana in the many specimens of whitish at the base and light green to green- Habenaria that we have examined from these ish yellow in the segments; posterior seg- states. Examination of the syntypes of H. gourlieana f. magna Hoehne, located at the National Museum herbarium (R), revealed that this taxon is referable to H. nabucoi. Although this taxon is certainly not a variety of H. gourlieana, and was described earlier than H. nabucoi, according to article 11.2 of the International Code of Botanical Nomenclature (Greuter et al., 2000), in no case does a name have priority outside the rank at which it is published. 34 BRITTONIA [ V O L . 58 2cm 5mm 2cm[ 5mm i'"I/ t FIC. 7. Habenaria paulistana. A. Habit. B. Flower, side view. B1. Flower, front view. C. Ovary and pedicel, column and spur, side view. D. Perianth. E. Column, side view. El. Column, 3/4 side view. F. Column apex, front view showing the connective and anther apex. G. Rostellum, side view. (A, J. P Souza et al. 875, ESA, isotype; B-G, Batista & Pansarin 1089, CEN). Habit, perianth, ovary and pedicel drawn from dried material, other structures drawn from alcohol-fixed material. 2006] B A T I S T A E T AL." H A B E N A R I A S E C T . M A C R O C E R A T I T A E ( O R C H I D A C E A E ) Habitat, Ecology, and Phenology.-Habenaria paulistana is found in moist places but does not occur in water. If this finding is confirmed by further field studies, then it would represent another difference between H. paulistana and H. nabucoi and H. longicauda, since the latter two species always grow in water. The flowers of H. paulistana are green to light green, becoming green-yellow with age. Flowering occurs mainly during November and December. Illustrations.--Schlechter and Hoehne (1921, plate 2, Fig. 3, as H. sartoroides), Hoehne (1930: 79, as H. sartoroides, photo). Additional specimens examined: BRAZIL. S7~o PAULO: Reserva Biol6gica do Parque Estadual das Fontes do Ipiranga. 25 Nov 1981 (fl), Barros 657 (CEN, SP); JundiaL Serra do Japi, ecologic base camp, 7 Nov 2000 (fl), Batista & Pansarin 1089 & 1089A (CEN); Biritiba Mirim, Estaqao Biol6gica de Boracria, 23~ ', 23~ 45~ ', 45~ 890-950 m, 9 Dec 1983 (fl), Custrdio Filho 2040 (SP, UB), 29 May 1986, Custrdio Filho 2612 (SPSF), 19 Apr 1986, Custddio Filho 2561 (MBM, SPSF, UB); Sales6polis, Casa Grande, Esta~go Biol6gica de Boracria, 23~ 45~ 890-950 m, 17 Dec 1986 (fl), Custddio Filho 2796 (UEC); JacarehyMogy das Cruzes road, 25 Nov 1938 (fl), Germeck & Paolieri s.n. (IAC 4368, IAC 4461, SP 266611); Mun. Sales6polis, Estaq~o Biol6gica de Boracria, 21 Nov 1957 (fl), Handro 745 (HB, SP, SPF); Silo Paulo, capital, Av. Indianapolis, 16 Nov 1948 (fl), IV..Hoehne s.n. (SPF 13337); Jardim Morumbi, 6 Dec 1948, Joly 699 (SPF); Sales6polis, Boracria, margins of Rio Claro, 30 Nov 1951 (fi), Kuhlmann 2782 (SP); Franca, 22 Nov 1909 (fi), Loefgren 483 (RB); Atibaia, Pedra Grande, 20 Nov 1997 (fi), Pansarin et al. 97/86 (UEC); Jundiaf, Serra do Japi, near the DAE dam, 1 Nov 1999 (fl), Pansarin 559 (UEC), 9 Oct 2000 (fl), Pansarin 726 (UEC); S~o Paulo, botanical garden and state park, 12 Aug 1968 (fl), Sendulsk~y 989 (SP). In the general morphology and size of the plants and flowers, Habenaria paulistana is similar to H. nabucoi, the species with which it has most frequently been mistaken (identified as H. fastor). However, other than these characteristics, both species show few similarities. They can be distinguished by the inflorescence, which is usually long and many flowered in H. paulistana (vs short and few flowered in H. nabucoi), green flowers (vs white), deflexed lateral sepals (vs nonreflexed), the spur narrowed towards the apex (vs thickened), the anthers and the rostellum smaller and shorter, and the rostellum arms thickened in H. paulistana (Figs. 1, 2, 4, 5, 35 7). Also, the lateral segments of the petals and lip are usually narrower and slender in H. paulistana. Although Habenaria paulistana is similar to H. nabucoi in the size of the flowers and development of the spur, the pattern of the inflorescence, color of the flowers, arrangement of the lateral sepals and column structure place it close to H. macronectar, from which it differs mainly by the larger size of the plants and flowers (Figs. 1, 7). It is possible that more detailed studies may show that H. paulistana is a subspecies or variety of H. macronectar. However, because of the difficulties in identifying and defining infraspecific ranks in Habenaria, we have recognized H. paulistana as a species. As is common in the genus and in sect. Macroceratitae, the petals of this species can occasionally develop a third segment (Fig. 7). Habenaria hauman-mercki and H. spegazziniana were based mostly on the tripartite petals. However, this third projection is variable in size, ranging from a short tooth-like formation to the size of the posterior petal segment (Hauman, 1920, Fig. 1; Hoehne, 1940, table 13), and most probably represents a genetic abnormality during development since its appearance is occasional and shows no fixed pattern. This third segment can occur in a single flower among other normal flowers in an inflorescence or in one of the petals of a flower. Apparently, the same genetic program that results in the development of three segments in the lip is somehow turned on in the petals. Brazilian species excluded from sect. Macroceratitae Many of the Brazilian species placed in sect. Macroceratitae by Kraenzlin (1892) and Cogniaux (1893) do not show the features listed above and are sufficiently distinct to be removed from the section. Habenaria araneiflora Barb. Rodr.--The inclusion of this species in sect. Macroceratitae is unsupported since it does not show the long stigmas characteristic of the section. The Barbosa Rodrigues original illustration of the lateral view of the column of H. araneiflora (Spmnger, 1996) was deliberately modified by Cogniaux (1893) by the addition of long 36 BRITTONIA stigmas and long anther canals. It is not known why Cogniaux did this, since the stigmas of H. araneiflora are characteristically short and uncinate at the apex, as shown in the other figures in the same illustration (Cogniaux, 1893, table 3, Fig. 1). Also, the flowers of H. araneiflora are smaller than those in the other species of sect. Macroceratitae. Habenaria araneiflora is very similar to H. secunda Lindl., H. achalensis Kraenzl., H. brachyplectron Hoehne & Schltr., H. gustaviedwallii Hoehne, and H. paranaensis Barb. Rodr., which most likely form a natural group. Habenaria platantherae Rchb.f. probably belongs to the same group. Kraenzlin (1892) and Cogniaux (1893) placed species of this group in sects. Pentadactylae Kraenzl., Pratenses Kraenzl., Clypeatae Kraenzl., and Micranthae Kraenzl. Habenaria aricaensis Hoehne.--This species has never been formally placed in sect. Macroceratitae, but Hoehne (1940) and later Pabst and Dungs (1975) included H. aricaensis within their systems of infrageneric classification in the group corresponding to sect. Macroceratitae. However, apart from being a robust plant with large flowers and a long spur, there are few characters in common between H. aricaensis and the species in sect. Macroceratitae. In H. aricaensis, the lateral segments of the petals and lip are very short, and the spur is somewhat hook shaped and free from the bracts. The column structure of H. aricaensis is also very distinct, being remarkably similar to that of H. hamata Barb. Rodr., which may indicate a close relationship between the two species. If this assumption proves to be true, H. aricaensis would be better placed in sect. Seticaudae Kraenzl. Habenaria cryptophila Barb. R o d r . - Among the several unrelated species previously placed in sect. Macroceratitae, Habenaria cryptophila is the only one that has long stigmas relative to the size of the flowers, and thus the only species that could be tentatively assigned to sect. Macroceratitae. However, the stigmas are not involute and the flowers and spur are smaller than in the other species of the section. Habenaria cryptophila also differs by the broader, oblong-lanceolate, fiat, unkeeled leaves and [VOL. 58 by its habitat, it is a species that is typical of dry forests. Among the Brazilian Habenaria, Habenaria cryptophila seems rather unique, and its affinities are unknown. It is similar to H. alterosula Snuverink & Westra, known from Suriname and French Guiana, and to a group of Mexican and Central American species including H. jaliscana S. Wats, H. diffusa Rich. & Galeotti, H. costaricensis Schltr., H. ortiziana R. Gonz~lez, and H. agapitae R. Gonz~lez & Reynoso. Habenaria diffusa was placed by Kraenzlin (1892, 1901) in sect. Clypeatae. Habenaria distans Griseb.--This species, which also does not show the long stigmas characteristic of sect. Macroceratitae, can be distinguished from other species in the section by its small flowers and its leaves, which are basal, oval to orbicular, and broad. Habenaria distans is similar to H. glaucophylla Barb. Rodr. and possibly also to H. lindmaniana Kraenzl., H. schenckii Cogn., H. depressifolia Hoehne, and H. drepanopetala Pabst. Species of this group were placed by Cogniaux (1893) in sects. Micranthae and Pentadactylae. Habenaria hydrophila Barb. R o d r . - - I t is not clear why Kraenzlin (1892) placed this species in sect. Macroceratitae as it has short stigmas. The species is also distinguished by smaller plants and flowers, the very short lateral segments of the petals and lip, and the column structure. Also, contrary to the specific name, the species is typical of dry places. Habenaria hydrophila is similar to H. brevidens Lindl., H. aestivalis Barb. Rodr., H. guilleminii Rchb.f., H. loefgrenii Cogn., and H. graciliscapa Barb. Rodr. According to the current infrageneric classification, these species would be best placed in sect. Micranthae Kraenzl. Habenaria melvilii Ridl.--Based on the original description of this species, Hoehne (1940) considered H. melvilii to be a mixture of the vegetative part of a Cyclopogon species and the stem and flowers of a Habenaria from sect. Macroceratitae. The examination of an image of the species holotype, located at BM, confirmed Hoehne's supposition and revealed that the type material is composed of the vegetative part of a Spiranthoideae species (probably a Cyclo- 2006] BATISTA ET AL.: HABENARIA SECT. MACROCERATITAE (ORCHIDACEAE) p o g o n or Prescottia) and two loose flowers of a species from sect. Macroceratitae. T h e flowers are p r o b a b l y referable to H. gourlieana or H. nabucoi, but based on the i m a g e alone we have not been able to reach a definite conclusion. H a b e n a r i a rodeiensis Barb. R o d r . - - I n contrast to other species included in sect. Macroceratitae, H. rodeiensis has smaller flowers, a long-pedicellate ovary, and stigmas that are shorter, not involute and closely parallel, thus forming a single receptive surface. H a b e n a r i a rodeiensis is very similar to H. santensis Barb. Rodr., H. tamanduensis Schltr., H. longipedicellata Hoehne, H. dusenii Schltr., and H. sylvicultrix Lindl. Species of this group were placed by Kraenzlin (1892) and C o g n i a u x (1893) in sects. Microdactylae Kraenzl. and Pentadactylae Kraenzl. Acknowledgments The authors thank the curators of A M E S , B H C B , B H M H , B O T U , CEN, CEPEC, CESJ, CH; C V R D , ESA, FUEL, HB, H U E F S , H U F U , IAC, IAN, IBGE, INPA, M B M , M B M L , M G , NY, OUPR, OXF, PACA, PAMG, R, RB, S E S P E S P S E UB, UEC, U F G , U P C B , and VIC for the loan of material or for providing access to their collections. We also t h a n k Jorge Fontella and Fabio de Barros for helpful suggestions, E m e r s o n R. Pansarin for his assistance in locating H. paulistana in the field and for providing fixed flowers of H. j o h a n e n n s i s , Eduardo Borba and Cassio van den Berg for providing fixed flowers of H. macronectar, Walter Rossi for p e r m i s s i o n to use the photograph of H. bractescens, Eduardo Gonqalves for locating the p o p u l a t i o n of H. longicauda at Barreiras, Stephen Hyslop for corrections in the English, Claudio N. Fraga for some of the references, Gustavo Romero, T h o m a s A. Z a n o n i , Lawrence M. Kelly, and an a n o n y m o u s reviewer, for i m p r o v e m e n t s in the m a n uscript. We are also particularly thankful to the following researchers, curators, and herbarium staff for the images of H a b e n a r i a used in this work: E l m a r Robbrecht and A l a i n V. Velde (BR), O l o f R y d i n g (C), Clare Drinkell (K), Mia E h n (S), Victoria N o b l e 37 (BM), Gustavo R o m e r o (AMES), T h o m a s Z a n o n i (NY), Ernst Vitek and H e i m o Rainer (W), and Toscano de Britto. Literature Cited Alves, R.J.V. 1991 9Field guide to the orchids of the Serra de S~o Josr, Minas Gerais, Brazil. Tropicaleaf, Prague 9 Angely, J. 1965. Orchidaceae. Habenaria, Pp. 234-237. ht: J. Angely (ed.), Flora Anal/tica do Paran~i.Vol. 7. Cole~o Saint-Hilaire. Ediqres Phyton, Curitiba, Paran~, Brasil. Artueio, P.I. 1984. Orchidaceae. Habenaria. In: A. Lombardo (ed.). Flora Montevidensis. 3. Monocotiledonras: 402-410. Intendencia Municipal de Montevideo, Montevideo, Uruguay. Barbosa Rodrigues, J. 1881. Resultado botfinico de uma breve excursao a S. Joao d'el-Rei, Minas Gerais. Revista de Engenharia 3: 73-75. Bateman, R.M., P.M. Hollingsworth, J. Preston, L. Yi-Bo, A.M. Pridgeon & M.W. Chase. 2003. 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Beitr~ige zu einer monographie der tions. Friedrich Reinhardt Verlag, Basle, Switzerland. gattung Habenaria Wilt& Botanisch Jahrbficher f~r Steam, W.T. I983. Botanical Latin, Ed. 3 9 Newton Systematik 16: 52-223. Abbot, Devon, England 9 91901. Orchidacearum genera et species. Vol. I. Szlaehetko, D . L . 2004. Mat6riaux. pour la r6vision des Habenaria. Mayer and Miiller, Berlin. Habenadinae (Orchidaceae, Orchidoideae)- 4. 9 191 t. Beitrage zur Orchideenflora SuRichardiana 4: 52~55. damerikas. Habenaria. Kongl. Svenska VetenskapToscano de Brito, A . L . 1994. Notes on Brazilian Orchisakademiens Handlingar 4 6 : 5 - 1 6 . daceae III. Lindleyana 9: 55-57. Kurzweil, H, & A. Weber. 1992. Floral morphology of 9 1995. Orchidaceae. Pp.725-767 p. ilustr. In: southern African Orchideae. II. Habenariinae. Nordic B.L. Stannard (ed.), Flora of the Pico das Almas: Journal of Botany 12: 39-61. Chapada Diamantina - Bahia, Brazil. Royal Botanic Mansfcid, R. 1930. Bliitenanalysen neuer Orchideen 9 1 Gardens, Kew. == Sfidamerikanische Orchideen. Habenaria. ReperVellozo, J , M , C . 1831. Florae Fluminensis. Vol. 9. Tytorium Specierum Novarum Regni Vegetabilis (Beipographia Nacional, Rio de Janeiro. hefte) 58: 1-7. Warming, E. 1884. Symbolae ad floram Brasiliae cenMiller, D. & R. W a r r e n . 1994. Orchids of the high tralis cognoscendam. Part. 30. Orchideae 2. Videnskmountain Atlantic rain forest in southeastern Brazil. abelige Meddelelser fra den Naturhistoriske Forening Editora Salamandra, Rio de Janeiro. i Kjebenhavn 5-8: 86-99. 2006] BATISTA ET AL.: HABENARIA SECT. MACROCERATITAE Werkhoven, M . C . M . 1986. Orchideen van Suriname, Orchids of Suriname. Vaco Uitgeversmaatschappij, Paramaribo, Surinam. Index to Scientific N a m e s in Habenaria sect. Macroceratitae in Brazil (Accepted names indicated by boldface lettering.) Bertauxia vaupellii (Rchb. f. & Warm.) Szlachetko, 3 Habenaria braeteseens Lindl., 1 Habenaria bradei Schltr., 3 Habenaria burkartiana Hoehne, 2 Habenaria dolichosceras Barb. Rodr., 5 HabenariafastorWarm. ex Hoehne, 2, 6, 7 Habenaria gourlieana Gill. ex Lindl., 2 Habenaria gourlieana Gill. ex Lindl. f. magna Hoehne, 6 Habenaria hauman-mercki Hickens, 1 Habenaria helodes Rchb. f., 5 Habenaria johannensis Barb. Rodr., 3 Habenaria juergensii Schltr., 5 Habenaria kleyi Schltr., 1 Habenaria Habenarla Habenaria Habenaria longieauda Hook., 4 macroneetar (VEIL) Hoehne, 5 nabueoi Ruschi, 6 paulistana J.A.N. Bat. & Bianchetti, 7 Habenaria pontagrossensis Kraenzl., 1 Habenaria sartor Lindl., 5 Habenaria sartor Lindl. forma minor Cogn., 1 Habenaria sartoroides Schltr., 5 Habenaria vaupellii Rchb. f. & Warm., 3 Kusibabella bractescens (Lindl.) Szlachetko, 1 Kusibabella burkartiana (Hoehne) Szlachetko, 2 Kusibabella fastor (Warm. ex Hoehne) Szlachetko, 6 Kusibabella gourlieana (Gill. ex Lindl.) Szlachetko, 2 Kusibabella longicauda (Hook.) Szlachetko, 4 Kusibabella macronectar (Veil.) Szlachetko, 5 Kusibabella sartor (Lindl.) Szlachetko, 5 Macrocentrum mendocinum Phil., 2 Orchis macronectar Veil., 5 Exsiccatae Amaral, A., 1761 (3) Amaral, M.C.E. & V. Bittrich, 20/2000 (3) Amaral, M. C. E. et al., 2000/38 (6) Amarante, E., 08 (5) Arechavaleta, J., 2625 (2) Ashton, E. M., s.n. (6) Atldns, S. et al., CFCR 14807 (3) Barreto, M., 4867 (3); 4869 (6) Barros, E, 657 (7); 2231 (3) Barros, E & W. Marcondes-Ferreira, 12247A (3) Batista, J.A.N., 253, 1040 (6); 128, 215, 344, 948 (2); 1039 (4); 836 (3) Batista, J. A. N. & L.B. Bianchetti, 908 (3) Batista, J. A. N. & A. R. C. Lemos, 867 (4); 869 (6) Batista, J. A. N. & R. S. Oliveira, 848 (6) Batista, J.A.N. & E.R. Pansarin, 1089, 1089A (7) Batista, J. A. N. & K. E Pellizzaro, 1320 (2) Batista, J.A.N. et al., 1378 (5) (ORCHIDACEAE) 39 Berro, M., 1472 (1); 3793, 7299 (5); 7298, 7299 (2) Beyrich, H.K., 70 (5) Bianchetti, L. B., 466 (6) Bidd, A. et al., 283 (5) Bittencourt, N. S., 00/60 (3) Borba, E.L., 174 (3) Borba, E.L. et al., s.n. (5) Brade, A. C., 6200 (3); 19677 (5) Brade, A.C. & M. Barreto, 12092 (2) Brandao, M., 13196 (3) Bresolin, A., t 119 (3) Brina, A. E., s.n. (5) Bruxel, A., s.n. (5) Buck, P., s.n. (5) Burkart, A., 946 (2); 3812, 4510 (1) Cabrera, A.L., 1623 (1); 2710 (2) Camargo, L., 18 (1) Camargo, O.R., 3110 (5) Carneiro, J., 290 (3) Cesar, R. et al., 226 (3) Chagas, E & Silva, 1630 (3) Colnago, E., s.n. (6) Colnago, E. & Kautsky, 395 (6) Cordeiro, J. & P.I. Oliveira, 435 (5) Cordeiro, J. & A. Souza, 417 (5) Cruz, J., s.n. (3) Cullen, L., s.n. (1) Cust6dio Filho, A., 2040, 2561, 2612, 2796 (7) Damazio, L., 130 (3) Davidse, G. & W.G. D'Arcy, 10530 (3) Davidse, G. & T. P. Ramamoorthy, 10545 (3) Dittrich, V.A.O. et al., 317 (5) Dombrowski, L. Th. & Y. S. Kuniyoshi, 2252 (3) Dombrowski, L. Th. & P. S. Neto, 10109 (1) Duarte, A. P., 6330 (5); 9376 (2); 10173 (3) Duarte, A. P. & E. Pereira, 10287 (2) Ducke, A., s.n. (4) Dusen, P.K.H., 3514, 14420, 14470, 14571, s.n. (5); 7837 (i); 11575 (3) Dutilh, J.H.A. & W.M. Ferreira, 014 (3) Dutra, J., 673, 1086, 1087 (5) Edwall, G., CGGSP 4057 (3) Edwall, G. & P.C. Porto, s.n. (3) Emmerich, M., 3175 (5) Emmerich, M. & R. Dressier, 2803 (3) Emrich, K., s.n. (5) Falkenberg, D.B. et al., 6390 (2) Filgueiras, T. S,. 2349 (2) Fontella, J., 1164 (1) Forzza, R.C., 151 (3) Fraga, C.N. de, 580 (6) Freire, C.V., 293 (5) Gardner, G., 676, s.n. (5) Gehrt, A., s.n. (7) Germeck & Paolieri, L., s.n. (7) Ghillany, A., s.n. (5) Gibbs, P.E. & H. F. Leit~o Filho, 4294 (3) Gilberto, M., 38 (3) Gillies, J., s.n. (2) Glaziou, A . E M . , 6916 (5) Gonqalves, E.G., 175 (6); 177 (4) Gottsberger, G. K., s.n. (3) Hagelund, K., 12252, 13787 (1) Handro, O., 745 (7) 40 BRITTONIA Harley, R.M. et al., 20099, 24332 (1); 16086, 18787, 19597 (3) Hassler, E., 5950, 6594 (1); 8703 (3) Hatschbach, G., 611, 3006, 3104, 3813, 3818, 12387, 13679, 18320, 21173, 26330, 26553, 48171, 49359 (5); 9900, 9332, 9611, 43974 (1); 5472, 16065, 18231, 26101, 40715 (3) Hatschbach, G. & L.Z. Ahumada, 31255 (3) Hatschbach G. & O. Guimaraes, 19094, 20530 (1) Hatschbach G. & Haas, 5495 (1) Hatschbach, G. & A. Manosso, 52678 (3) Hatschbacb, G. el al., 54669 (2); 13808, 58616 (3); 67492 (5) Hauman-Merck, L.L., s.n. (1) Henz, E., s.n. (5) Heringer, E. P., 6383, 10039 (6); 8158, 10025, 10041 (2) Herter, W., 18370, 17166 (1) Hickens, C.M., s.n. (1) Hoehne, F.C., 6420, 6421 (2); s.n. (3); s.n. (5); s.n. (7) Hoehne, W., s.n. (3); s.n. (7) Houk, W. G., s.n. (3) Ikusima, I. & J.G. Gentil, s.n. (6) Irwin, H.S. et al., 11431, 30637 (3) Jardim, J.G. et al., 768 (1) Joly, A. B., 699 (7) Juergens, C., 23 (5); 1044 (1) Justen, A., s.n. (5) Kautsky, R., 228 (5) Kley, U., 01 (I) Kley, U. in herb. J. Dutra, 994 (1) Kolbek, J., & R.J.V. Aires, 1000 (5) Krapovickas, A. & C.L. Crist6bal, 28649 (1); 39723 (5) Krieger, P.L., 14365, 15492 (1); 16873 (5) Kuhlmann, J.G., 124, 125 (6) Kuhlmann, M,. 33 t (3); 2782, s.n. (7) Kuhlmann, M. & A. Gehrt, s.n. (3) Kumrrtrow, R., 377, 1068, 2244 (5); 2567 (3) Kummrow, R. & J. G. Stutts, 1776 (3) Kummrow, R. et al., 2559 (1) Kuniyoshi, Y. S., 4289 (1) Lagasa, E., 162 (6) Lanstyak, L., s.n. (3) Leinig, M., 279 (5) Lemos, D., s.n. (7) Lewis, G.P. et at., 1390 (1) Lindeman, J.C. & J. H. de Haas, 4596 (5) Loefgren, A., 483 (7) Loefgren, A., CGGSP 3258 (7); CGGSP s.n. (6) Lutz, A., 366, 415 (5) Lutz, A, & B. Lutz, 273, 1615, 1738 (5) Lutz, B., 1019, s.n. (5) Macedo, A., 2631 (1) Magalhaes, E, s.n. (3) Markgraf, F. et al., 3350 (2) Martins, A. B. et al., 31456 (3) Mattos, J., 5008, 5062, 5067 (5) Mello-Filho, L. E., 758 (5) Mendonffa, R.C. et al., 3838 (3) Miers, J., 2248 (5) Miguel, G., s.n. (5) Mimura, I_, 303 (3) Moreira-Filho, H., s.n. (1) Mos6n, C. W. H., 4558 (5) [VOL. 5 8 Mota, R.C. & EL. Viana, 1709 (3) Moura, J. T., 68, 104 (5) Nauhs, J., s.n. (1) Novaes, J.C., 541 (3) Oliveira, J. A., s.n. (3) Oliveira, R. S., 258 (3) Oliveira, R. S. & E.G. Gon~alves, 282 (4) Orth, C., s.n. (5) Osten, C., 14723 (2) Pabst, G.EJ., 1314, 1339, 1355, 7305 (5); 1340, 9081, 3216 (3) Pabst, G. E J. & M. Emmerich, 9029 (1) Pansarin, E. R., 559, 726 (7); 763 (3) Pansarin, E.R. et al., 97/86 (7) Pansarin, E.R. & L. Mickeliunas, 962, 990 (3); 1006, 1128 (5) Parker, C. S., s.n. (4) Pedersen, T. M., 10271 (2) Pereira, E., s.n. (5) Pereira, E. & G.EJ_ Pabst. 3174, 7962, 8002 (5); 3331 (3) Pires, J.M. & G. A. Black, 547, 1538 (4) Pivetta, 1179 (5) Polo, L., 09, s.n. (5) Porto, RC., 1027 (3); 1522, 3192, 3207(5) Prance, G.T. et al., 8057, 13665 (6) Proenga, C. & A. Lake, 1874 (2) Queiroz, L. P de & V. L. E Fraga, 3303 (6) Rambo, B. s.n. (1); s.n. (3); s.n. (5) Ramos, A.E., 179 (6) Raspalho, J., s.n. (1) Regnell, A. E, III-1186 (5) Reitz, R., 1359 (3); 6462 (5) Reitz, R. & R.M. Klein, 8437, 12408 (5) Ribas, O. S. & L. B. S. Pereira, 1773 (3) Ribas, O. S. & R M. Ruas, 509 (5) Ribas, O. S. & J. M. Silva, 3278 (5) Ribas, O.S. et al., 1251 (5) Richter, E., s.n. (5) Rojas, T., 3961 (1); 9334, 9585 (3) Rombouts, J. E., s.n. (2) Rosengurtt, B., B2577 (1); BI071, 5854 (2) Rosengurtt, B. et al., PE-4763 (2) Roth, L., 16475 (5); 16479 (2) Sacco, J.C., 1288 (5) Saleh, J.E_, 108 (5) Santos, T. S., 2859 (6) Santos, T.S., 3167 & L.A.M. Silva, 120 (6) Santos, N., 1261 (5) Scariot, A.O. et al., 344 (5); 718 (4) Schiller, s.n. (3) Schinini, A., 27559 (1) Schroeder, J., 15752 (2) Schulz, A.G., 582 (1) Schwacke, C. A. W., 11472 (3) Sendulsky, T., 989 (7) Sevilha, A. C. et al., 1802 (6) Sidney & E. Onishi, 1560 (1) Singer, R.B. & Dittrich, 30, s.n. (5); s.n. (3) Silva, C.J., 399 (6) Silva, J. B. F., 476 (4) Smith, L.B. & R.M. Klein, 10989 (5) Sobral, M. et al., 6700 (6) Souza, J. R et al., 875 (7) 2006] BATISTA ET AL.: HABENARIA Souza, V.C. et al., 4998, 5063, 5100 (3) Spegazzini, C.L., 970 (1); 104753 (2) Sucre, D. et al., 2993, 10652 (5) Tameirao-Neto, E., 2991 (3) Theissen, F., 629, s.n. (5) Tofollo, D., s.n. (3) Tweedie, J., s.n (1) Ule, E. H. G., 40 (3) Vanni, R. et al., 2826 (I) Vidal, M. R. R., 293 (3) SECT. MACROCERATITAE (ORCHIDACEAE) 41 Vidal, W. N. et al., 609 (5) Vidal, J., 11-5942 (5) Xifreda, C.C. & S. Maldonado, 484 (1) Waechter, J.L. & L. Baptism, 687 (5) Wanderley, M. G. L., 284 (5) Warming, E., 131 (3) Welter, R.N., 199 (2) Widgren, J. F., 754 (3) Windisch, R.W. & A. Ghillany, 259, 279, 444 (3); 645 (6)

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