Habenaria leprieurii Reichenbach (1846: 376)

Batista, João A. N., Reis, André F. De S., Leite Junior, Josefran L. & Bianchetti, Luciano De Bem, 2018, Phylogeny and taxonomy of Habenaria leprieurii, H. alpestris and H. sect. Microdactylae (Orchidaceae), with descriptions of two new taxa, Phytotaxa 373 (4), pp. 241-271 : 252-260

Habenaria leprieurii Reichenbach (1846: 376) . Type:— FRENCH GUIANA. Cayenne, 1839, F.R. Leprieur s.n.

(holotype G! [G00169023], RENZ! [13725] holotype photo, isotype G! [006566/000008]). ( Fig. 3 )

Habenaria schwackei Barbosa Rodrigues (1882: 254) , as H. schwackii . Type:— BRAZIL. Paraná: Ponta Grossa, 13 January 1880, C.A.W. Schwacke 191 (2508) (holotype R! [R000002481], isotypes BR! [642686], R! [R000036268], RB! [36995]).

Habenaria platydactyla Kränzlin (1911: 9) . Type:— BRAZIL. Paraná: Capão Grande, Turma 23, próximo Rio Guavirova, 24 January 1910, P.K.H. Dusén 9089 (lectotype, designated here: S! [R-2727], isotypes HB! [57155] fragment, HBG! [500131], K! [000363801], MBM! [253385, 256036], MO! [1578703], NY [00687980] photo, NY! [00688020], P! [00409012], S! [07-7182, S14-50090]).

Habenaria amazonica Schlechter (1925a: 69) . Type:— BRAZIL. Rio Branco [Roraima]: July 1913, J.G. Kuhlmann 787 (holotype B, destroyed; lectotype designated by Batista et al. 2011b: RB! 3345 [00542614], isotypes AMES! [00287151], SP! [31798]).

Herb terrestrial, caulescent, sympodial, geophyte. Roots few, short, 0.6–1.0 mm wide. Tuberoid ellipsoid or globose, 9–26 × 3–10 mm. Stem erect, slightly sinouse, (13–)17–55(–61) cm long, including the inflorescence, 1–3 mm wide. Leaves 3–9, base sheathing, sheath closed, blade appressed to the stem or ascending, membranaceous, lanceolate, narrow, acuminate, 1.7–9.6 × 0.35–0.45 cm. Inflorescence 3–13(–20) cm, few to many-flowered, lax, secund to subsecund, 0.7–1.4(–2.5) flowers/cm inflorescence; floral bracts green, lanceolate or lanceolate-ovate, caudate, 7.5–15.6(–30.5) × (1.8–)4.0– 7.1 mm, smaller, equal or larger than the pedicellate ovary. Flowers (1–)2–15(–24), resupinate, ascending, glabrous; pedicellate ovary 19–24(–34) mm long, green; ovary arched, 10–18 mm long, 1.1–2.1 times as long as the pedicel; pedicel (6.5–)8.0–10.0(–15.0) mm long. Sepals green, margin whitish, smooth, rounded to truncate, mucronulate; dorsal sepal slightly concave, ovate or elliptical when flattened, 3.6–5.2(–6.7) × 2.6–4.6 mm; lateral sepals concave, obliquely ovate or lanceo-ovate, deflexed, acute, 4.4–7.2(–8.4) × 1.8–3.4 mm. Corolla white. Petals bipartite; posterior segment oblong to narrowly elliptical, falcate, acute, 3.6–6.4(–7.2) × 1.4–2.9 mm, adherent to the dorsal sepal; anterior segment linear to spatulate, erect, reflexed, usually embracing from behind the dorsal sepal, inserted at the base of the posterior segment, (4.0–)5.2–6.4(–8.6) × 0.8–2.1 mm long, 0.9–1.4(–1.7) times as long as the posterior segment. Labellum tripartite; undivided basal part short to long, 0.4–1.6(–2.7) × (1.1–) 1.6–2.6 mm; lateral segments linear to spatulate, base perpendicular to the median segment, erect and reflexed towards the apices, (3.6–)4.5–7.7(–8.6) × 0.8–2.1 mm long, 0.8–1.2 times as long as the median segment; median segment linear, straight to slightly curved, rounded, 3.8–7.7(–9.6) × 1.4–2.9 mm; spur deflexed, apex enclosed by the bract, arched forwards when free, clavate to slightly clavate, rarely linear, 11.2–27.1 × 0.9–1.8(–2.2) mm, 0.8–0.9 times as long as the pedicellate ovary, whitish to greenish. Gynostemium erect, 2.6–3.1 mm high; connective emarginate, white; lateral appendages (auricles) white, fleshy, verrucose, obtuse, 0.7–0.9 mm × 0.4–0.6 mm. Anther bilocular, loculi parallel or slightly convergent, 1.6–1.9 mm high; canals short, about 0.5 mm long; hemipollinaria united by the viscidia; viscidia ellipsoid, 0.4 × 0.2 mm; caudicles 0.4–0.6 mm long. Stigmatophores (stigma lobes) 2, white, closely parallel, 3.2–3.5 mm long, receptive surface straight, facing upwards, almost perpendicular to the anther loculi, 1.7–2.0 × 0.9–1.1 mm, apex rounded, inner margin slightly thickened. Rostellum white, 1.7–2.6 mm long; mid-lobe triangular, fleshy, erect, obtuse, completely placed between the anther loculi, 1.0– 1.2 mm high; side-lobes converging towards the apices, ca. 0.9 mm long, base 0.4 mm wide.

Distribution and Conservation Status:— Habenaria leprieurii is widely distributed, occurring in Colombia, Venezuela, Guyana, French Guiana, Suriname, Brazil (BA, GO, MA, MG, MS, MT, PA, PR, RR, SP, TO), and Paraguay. Based on the IUCN Red List Categories and Criteria and their guidelines ( IUCN 2012, 2016), H. leprieurii can be tentatively classified as of least concern (LC), due to its wide geographic distribution and large number of known populations, many of which occur inside protected areas.

Habitat, phenology, and ecology:— Habenaria leprieurii occurs in permanently to seasonally wet grasslands. Flowering occurs during the rainy season, varying according to the region. In the southeastern and central-western regions of Brazil, flowering occurs from November to March (but is concentrated during December and January), while in Roraima State in northern Brazil flowering occurs from July to September.

Notes:—The identity of H. leprieurii has often been confused due to ambiguous interpretations by various authors (or even the same author). This confusion began with Reichenbach, the author of the species, who identified specimens of H. cruegeri (Leprieur s.n. [W-R! 51238]) and H. omissa (Warming 149 [W-R! 30472]) as H. leprieurii . Most later authors were able to morphologically differentiate H. leprieurii , H. heptadactyla , and H. cruegeri , but they erred in the applications of those names, in the circumscription of the taxa, by synonymizing distinct species, or in its separation from other graminiform-leaved Habenaria species such as H. omissa , H. mesodactyla , H. secundiflora , H. sprucei Cogniaux (1893: 40) , H. ayangannensis Renz (1992: 490) , and H. paxamorque Léotard & Galliffet in Léotard et al. (2017: 2). The name H. leprieurii , for example, has been applied to specimens of H. mesodactyla and H. crugeri ( Schultes 1960) , H. heptadactyla ( Ames 1910, Dunsterville & Garay 1972, Foldats 1969, Snuverink & Westra 1983, Kenny 1992) and, more recently, to H. cruegeri ( Renz 1992, Batista & Bianchetti 2002, Foldats et al. 2003, Batista et al. 2008, 2011 a, Carvalho et al. 2013, Léotard et al. 2017). Some authors ( Maguire 1948, Schweinfurth 1967, Foldats 1969, Snuverink & Westra 1983) correctly identified specimens of H. leprieurii , but included H. heptadactyla as a variety or a synonym. Habenaria leprieurii has most recently been identified as H. platydactyla ( Renz 1992) or H. schwackei ( Batista et al. 2008, Romero-González & Batista 2009, Léotard et al. 2017), which are now treated as synonyms. The collection Ule 3121 (= 498) (AMES!, BR! [697072], R! [2483], SP! [42473]) from Serra Dourada, Goiás, identified as H. leprieurii by Cogniaux (1906) and as H. alpestris by Hoehne (1940) is, in fact, H. cf. pseudoculicina J.A.N.Bat. & B.M.Carvalho in Batista et al. (2010: 32).

The main distinctive characteristics of H. leprieurii are its petals and lip invariably pure white, and the length of the pedicel [(6.5–)8.0–10.0(–15.0) mm], being longer than other species of the section ( Table 2). In addition to the characteristics cited above, H. leprieurii is easily distinguished from H. cruegeri by the shape and position of the pedicellate ovary: more or less curved throughout its length in H. leprieurii vs mostly straight and parallel to the inflorescence axis, and curved only at the apex in H. cruegeri . H. omissa is the most similar species morphologically as it also has whitish petals and lip, although it is distinct by having smaller flowers and shorter pedicels (2.0–5.0 mm long). The direction of the anterior segment of the petals, curved backwards, as mentioned by Renz (1992), is not particularly distinctive as it is also frequently observed in H. cruegeri .

As accepted here, H. leprieurii is a morphologically variable species, particularly in terms of the sizes of its flowers and the widths of its petals and lip lateral segments. In some populations or specimens the widths of the lateral segments of the petals and the lip are greater than the width of the posterior segment of the petals and the median segment of the lip respectively, which is an unusual feature among the Neotropical species of the genus. Specimens from Paraná State, Brazil, near the southern limit of its distribution and near the type locality of H. schwackei and H. platydactyla , have large flowers, while specimens from northern South America (near the northern limit of its distribution) and near the type locality of H. leprieurii and H. amazonica , have small flowers and wide lateral segments. However, both characteristics vary within the geographic distribution of the species and within populations, and intrapopulational variations in perianth morphology are illustrated here in Figure 3 . Flowers in figures 3D, 3E and 3F are from Chapada dos Veadeiros, Goiás (3D and 3F of the same population), flowers in figures 3G and 3H are of the same population from Ponta Grossa, Paraná, and flowers in figures 3I and 3J are of the same population from Pirenópolis, Goiás. Therefore, as separation into two or more taxa would be very difficult we prefer to recognize H. leprieurii here as a single species showing morphological variability. In at least one locality, Serra dos Pirineus, in Goiás State, Brazil, H. leprieurii and H. omissa are sympatric and flower at the same time. While the local specimens of H. omissa are in agreement with the morphological patterns of that species, the floral morphology of the local population of H. leprieurii is quite variable, with some specimens being difficult to differentiate between the two species, suggesting hybridization, probably with H. omissa as the pollen donor.

Szlachetko et al. (2016, 2017) accepted H.platydactyla , included H. amazonica as a synonym, and used the size of the spur and position and form of the lip lateral segments to distinghish it from H. schwackei : spur 11–19 mm long and lip lateral segments linear, attenuate towards apex in H. schwackei vs 18–26 mm and upcurved, linear-oblanceolate, obliquely truncate to obtuse, widely divergent, in H. platydactyla . However, these characters are not consistent with the variability observed in these taxa as well as with the species types. In the type material of H. schwackei (Schwacke 191 [R!]), for example, the lip lateral segments are upcurved, range from linear to linear-oblanceolate, and the spur is about 22 mm long. In two of our collections of the species (Batista et al. 2524, from Ponta Grossa, Paraná, the same locality as the types of H. schwackei and H. platydactyla ; and Batista 815, from Chapada dos Veadeiros, Goiás) the spur ranges from 16 to 26 mm long, while lateral segments of the lip range from linear to oblanceolate within the same population.As stated above, the flower size and width of the lateral segments of the petals and the lip are quite variable in the group, and extreme forms appear distinct from each other. However, the existence of intermediate forms and variations within the populations make the distinction of more than one taxon impractical. In this way, we recognize a single species in the group and apply the oldest name, here recognized as H. leprieurii , including H. schwackei , H. platydactyla and H. amazonica as synonyms.

In the protologue of H. leprieurii , the type is indicated as: Cayenne, Leprieur, 1839, Delessert Herbarium. There are currently two Leprieur collections at G with the number 1839 (G00169023! and 006566/000008!), which refers to the collection year. A third collection is labelled as Guyane Française, 1834, Leprieur 231 (G00169196!), with a duplicate at P (P00408673!). We assume that the collection G00169023 is the holotype, because it is located in G, is the one that best matches the protologue data, and the only one that has a label with Reichenbach’s handwriting identifying it as H. leprieurii , whereas G 006566/000008 is a duplicate. G00169023 has a second label added over the upper left corner of the collection label with the number 231, while G 006566/000008 bears the number 244, added in a different handwriting in the lower right corner, but both numbers were certainly added afterwards. G00169196 and P00408673 (Leprieur 231) were collected in 1834, and although they are H. leprieurii , they are not related to the protologue.

In the protologue of H. schwackei , the type is indicated as: Paraná, Ponta Grossa, January, Schwacke 191, Musée de Rio de Janeiro. The only specimen that has exactly that data is R000002481!, from the National Museum of Rio de Janeiro. This sample is also the only one that has a label with Barbosa Rodrigues’ handwriting identifying it as H. schwackii —and we therefore assume that it is the holotype of H. schwackei . Other specimens (R000036268!, BR 642686!, RB 36995!) have the collection number Schwacke 2508, but the collector, date (13 January 1880), and locality (Paraná, Ponta Grossa) are exactly the same as that of number 191, and we assume that they are all from the same collection. Another specimen located in P (P00409022!) of imprecise origin from Brazil, also has an additional label with the number 191 and is quite likely a duplicate of the type. Kränzlin did not indicate the holotype of H. platydactyla . Among the various herbaria with specimens of the type collection (Dusén 9089), S (R-2727!) and HBG (500131!) are the only ones that have labels in Kränzlin’s handwriting indicating them as types. We chose the specimen at S as the lectotype, since S is the main herbarium with Dusén collections and types.

Illustrations: — Cogniaux (1893, fig. 7.1, based on Barbosa Rodrigues original illustration of H. schwackei ), Kränzlin (1911, taf. 2, fig. 5, original illustration of H. platydactyla ), Mansfeld (1930, t. 1, fig. 2, Schlechter original drawing of H. amazonica ), Foldats (1969, fig. 19), Pabst & Dungs (1975, pg. 252, fig. 111, as H. amazonica , based on Lima 76 [IAN!]; pg. 253, fig. 129, as H. platydactyla , based on Hatschbach 2964 [HB!]; pg. 254, fig. 138, as H. schwackei , based on Barbosa Rodrigues original illustration of H. schwackei ), Renz (1992, fig. 2, as H. platydactyla ), Sprunger (1996, vol. I, Fig. 10C, Barbosa Rodrigues original illustration of H. schwackei ), Batista et al. (2008, fig. 7 A-C, as H. schwackei ), Romero-González & Batista (2009, figs. 1D, 5, as H. schwackei ), Szlachetko et al. (2016, fig. 59, based on Kramer & Hekking 2941 [AMES!] fig. 60, based on Sastre 14 [P!], both as H. schwackei ; fig. 62, based on Dusén 9089 [P!] isotype of H. platydactyla ; fig. 63, based on Lindeman et al. 173 [NY!], as H. platydactyla ), Léotard et al. (2017, fig. 10, as H. schwackei ), Szlachetko et al. (2017, fig. 199, based on Kramer & Hekking 2941 [AMES!], as H. schwackei ; fig. 200, based on Leprieur 104 [K!], as H. schwackei ; fig. 201, based on Silva et al. 1833 [MO, G!, HBG!, HUEFS!, MBM!], as H. schwackei ; fig. 203, based on Dusén 9089 [P!] isotype of H. platydactyla ; fig. 204, based on Hatschbach 13509 [ US, AMES!, HB!, K!, MBM!, NY!, RFA, SPF!, UPCB!, W-R!], as H. platydactyla ; fig. 205, based on Lindeman et al. 173 [NY!], as H. platydactyla ; fig. 206, based on Sagot s.n. [BM!], as H. platydactyla ; fig. 208, based on Sastre 14 [P!], as H. platydactyla ,).

Misidentified illustrations: — Ames (1910, plate 70, fig. 2 = H. heptadactyla ), Dunsterville & Garay (1972, pg. 135 = H. heptadactyla ), Snuverink & Westra (1983, fig. 8, based on Oldenburger et al. 927 [U!] = H. heptadactyla ), Werkhoven (1986, pg. 141 = H.sprucei ), Kenny 1992 (pg. 17 = H. heptadactyla ; pg. 18, as H.leprieurii var. heptadactyla = H. heptadactyla ), Szlachetko et al. (2016, fig. 64) and Szlachetko et al. (2017, fig. 198), both based on Hatschbach 37932 [MO, MBM!], as H. platydactyla = Habenaria sp.

Additional specimens examined:— BRAZIL. Bahia: Cocos, Fazendas Trijunção, estrada velha da sede para o Guará, 14º46’35”S, 45º56’32”W, 11 December 2001, Fonseca et al. 3042 (CEN!, HUEFS!, IBGE!); Cocos, Fazenda Trijunção, estrada velha da sede administrativa para a sede do Guará, 14 o 40’55”S, 45 o 50’39”W, 830 m, 12 December 2001, Walter et al. 4940 (CEN!, IBGE!); Cocos, Fazendas Trijunção, região de Santa Luzia, 14º55’57”S, 45º56’57”W, 13 December 2001, Walter et al. 4984 (CEN!, IBGE!). Goiás: Pirenópolis, Morro do Cabeludo, 17 January 1992, Batista 241 (CEN!); Alto Paraíso, Parque Nacional da Chapada dos Veadeiros, 28 January 1993, Batista & Bianchetti 399 (CEN!); Pirenópolis, Santuário de Vida Silvestre Morro do Cabeludo, 22 January 1995, Batista & Lemos 500 (CEN!); Pirenópolis, Morro do Cabeludo, 19 February 1995, Batista & Bianchetti 533 (CEN!); Cavalcante, 32 km de Cavalcante no alto da Chapada, em direção a região conhecida como São Domingos , 31 December 1998, Batista 815 (BHCB!, CEN!, EAN!, HB!, MBM!, SP!); Cavalcante, cerca de 28 km de Cavalcante, no alto da Chapada, em direção a região conhecida como São Domingos , 1 January 1999, Batista 840 (CEN!, ESA!, HUFU!, SP!, UEC!); Teresina de Goiás, Chapada dos Veadeiros, GO-118, em direção a Alto Paraíso, ca. de 38 km ao N e antes de Alto Paraíso, na altura do km 204, 9 January 2001, Batista & Pansarin 1147 (CEN!); Alto Paraíso de Goiás, Chapada dos Veadeiros, GO-118, em direção a Teresina de Goiás, ca. de 19 km após e ao N de Alto Paraíso, na altura e entre os km 185–186, ca. de 2 km antes do morro com o cruzeiro, 10 January 2001, Batista & Pansarin 1155 (CEN!, HUEFS!, UEC!); Alto Paraíso de Goiás, Chapada dos Veadeiros, área a direita da pista na GO-118, em direção a Teresina de Goiás, ca. de 19 km após e ao N de Alto Paraíso, na altura e entre os km 185-186, ca. de 2 km antes do morro com o cruzeiro, 10 January 2001, Batista & Pansarin 1162 (CEN!); Cavalcante, cerca de 28 km da cidade, Engenho II, área dos Calungas, margem da estrada de terra entre o entroncamento de acesso para o Engenho II e o povoado Calunga, 26 November 2004, Batista et al. 1524 (BHCB!); Teresina de Goiás, Chapada dos Veadeiros, GO-118, cerca de 36 km N de Alto Paraíso, 13º54’9.5”S, 47º23’18.2”W, 1268 m, 4 January 2008, Batista 2384 (BHCB!); Alto Paraíso, Chapada dos Veadeiros, ca. 15, 4 km N de AltoParaíso, 14º0’52”S, 47º31’31”W, 1512 m, 30 December 2010, Batista & Bianchetti 3105 (BHCB!); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, entrada do parque, 13 December 1991, Bianchetti & Pereira-Silva 1234 (CEN!); Pirenópolis, 20 km N.W. of Corumbá de Goiás, Serra dos Pirineus, 26 January 1968, Irwin et al. 19246 (UB!, mixed with H. egleriana ); Pirenópolis, 20 km NW of Corumbá de Goiás, near Pico dos Pirineus, 27 January 1968, Irwin et al. 19326 (HB!, NY!, UB!); Alto Paraíso de Goiás, 8 km de Alto Paraíso, estrada Alto Paraíso/Colinas, 14º9’34”S, 47º35’33”W, 9 December 1988, Mendonça & Furtado 1146 (IBGE!); Alto Paraíso de Goiás, 8 km de Alto Paraíso, estrada Alto Paraíso/Colinas, 14º9’34”S, 47º35’33”W, 9 December 1988, Mendonça & Furtado 1159 (IBGE!); Alto Paraíso de Goiás, Buriti Sozinho, ca. 8 km a esquerda de Alto Paraíso para São Jorge, Chapada dos Veadeiros , 14º9’78”S, 47º36’79”W, 22 November 1997, Munhoz et al. 578 (HEPH!, UB!); Alto Paraíso de Goiás, Fazenda Água Fria, ca. 10 km em direção a Teresina de Goiás, 1448 m, 14 o 4’21.7”S, 47 o 30’33.6”W, 1448 m, 20 December 2000, Munhoz et al. 2326 (UB!); Terezina de Goiás, Rodovia GO 118, Km 189, Sentido Alto Paraíso de Goiás - Terezina de Goiás, próximo ao cruzeiro, 13º57’S, 47º28’W, 29 December 2001, Pansarin & Mickeliunas 866 (UEC!); Alto Paraíso de Goiás, estrada de Alto Paraíso para São Jorge , próximo ao “Portal da Chapada”, em frente ao “Jardim de Maytreia”, 4 January 2003, Pellizzaro et al. 43 (CEN!); Alto Paraíso de Goiás, 13 km de Alto Paraíso, estrada Alto Paraíso-Colinas, 14º10’00”S, 47º37’10”W, 6 December 1988, Pereira Neto 112 (IBGE!); [Pirenópolis] Serra dos Pirineus, December 1892, Ule 446 (P!); [Pirenópolis] Serra dos Pirineus, December 1892, Ule 496 (P!, R!, SP!, SPF!); [Pirenópolis] Serra dos Pirineus, December 1892, Ule 3119 (BR!); Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada entre Alto Paraíso de Goiás e São Jorge , próximo a São Jorge , 14º10’1”S, 47º47’10”W, 1013 m, 17 December 2012, Vale & Vale 176 (BHCB!); Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada entre Alto Paraíso de Goiás e São Jorge , 14º9’44”S, 47º36’50”W, 1154 m, 30 January 2013, Vale et al. 204 (BHCB!); Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada entre Alto Paraíso de Goiás e Teresina de Goiás, 13º54’7”S, 47º22’59”W, 1263 m, 31 January 2013, Vale et al. 221 (BHCB!); Alto Paraiso de Goiás, Estrada para o Parque Nacional da Chapada dos Veadeiros, 29 November 1988, Wanderley et al. 1695 (SP!); Alto Paraíso de Goiás, ca. 5 km N of Alto Paraíso, 3 December 1988, Wanderley et al. 1830 (BHCB!, SP!). Maranhão: Carolina, 20 km de Carolina, na estrada para Estreito, 16 January 1998, Oliveira & Gonçalves 269 (CEN!, UB!); Balsas, projeto geral de Balsas, lote 16, 12 November 1996, Oliveira & Pereira-Silva 464 (CEN!, HEPH!). Mato Grosso: 12 km SW of base camp (12 o 49’S, 51 o 46”W), 5 October 1968, Harley 10493 (K!, UB!); Chapada dos Guimarães, Rio da Casca, 15 February 1975, Hatschbach 37601 (MBM!); c. 1.5 km E. of base camp (12º49’S, 51º46’W), 26 November 1967, Philcox et al. 3276 (K!); Xavantina, 15 km S. na estrada Xavantina-São Felix, 6 October 1968, Sidney 1165 & Onishi 386 (UB!). Mato Grosso do Sul: Sidrolândia, Santa Fé, 23 January 1971, Hatschbach 26061 (HB!, MBM!). Minas Gerais: Formoso, Parque Nacional Grande Sertão Veredas, próximo a barra do Rio Preto com o Riacho Santa Rita, 15 o 09’40.5”S 45 o 45’42”W, 690 m, 6 December 1997, Mendonça et al. 3389 (CEN!, IBGE!, SP!). Pará: Ilha de Marajó, Rio Camará, fazenda Santa Rita, retiro Pau Grande, March 1950, Lima 76 (IAN!). Paraná: Ponta Grossa, Itaiacoca, Cachoeira da Mariquinha, 25º11’55.6”S, 49º57’1”W, 921 m, 6 February 2008, Batista et al. 2524 (BHCB!); Tibagi, Guartelá, 23 December 1992, Cervi 3996 (MBM!); Palmeira, Rio dos Papagaios, 15 January 1987, Cordeiro & Hatschbach 503 (MBM!); Palmeira, Fazenda Santa Rita, 14 January 1981, Dombrowski & Scherer Neto 12377 (MBM!); Ponta Grossa, 10 December 1903, Dusén 3206 (R!, SP!); Serrinha, 3 January 1915, Dusén 16264 (AMES!, MBM!, MO, S!); Ventania, Fazenda Santa Inês, 14 December 2004, Estevan et al. 601 (FUEL!, SP!); Jaguariaíva, Parque Estadual do Cerrado, trilha para mirante do Rio Jaguariaíva, 24°09’40.5”S, 49°39’20.3”W, 762 m, 3 February 2011, Fernandes et al. 698 (BHCB!); Tibagi, Parque Estadual do Guartelá, trilha para o mirante do Rio Iapó, 4 February 2011, Fernandes et al. 719 (BHCB!); Ponta Grossa, Vila Velha, 21 December 1952, Hatschbach 2964 (HB!, MBM!, RENZ!, SI!); Jaguariaíva, estrada para Cerro Alto, 21 December 1961, Hatschbach 8687 (HB!, MBM!); Ponta Grossa, Vila Velha, 21 December 1962, Hatschbach 9580 (HB!, MBM, U!); Castro, Carambeí, Rio São João , 15 January 1965, Hatschbach et al. 12119 (MBM!); Piraí do Sul, Serra das Furnas, Campo das Cinzas, 16 January 1965, Hatschbach et al. 12168 (MBM!); Palmeira, Serra das Almas, 12 January 1966, Hatschbach 13509 (AMES!, HB!, K!, MBM!, NY!, RFA, SPF!, UPCB!, US, W-R!); Jaguariaíva, Fazenda Chapada Santo Antônio, 27 November 1968, Hatschbach 20398 (MBM!); Arapoti, Rio das Perdizes, 6 April 1970, Hatschbach 24114 (MBM!); Palmeira, Córrego da Anta, Hatschbach & Pedersen 35866 (MBM!); Ponta Grossa, 1 January 1977, Krieger 15585 (CESJ!, HB!); Ponta Grossa, 1 January 1980, Krieger 16908 (HB!); Jaguariaíva, Rio das Mortes, 12 February 1997, Ribas & Pereira 1804 (MBM! mixed with H. secundiflora ); Palmeira, Recanto dos Papagaios, 15 January 1993, Silva & Santos 1218 (G!, HBG!, MBM!); Tibagi, canyon Guartelá, 13 December 1996, J.M. Silva et al. 1833 (G!, HBG!, HUEFS!, MBM!, MO); Jaguariaíva, Parque Estadual do Cerrado, 10 October 2000, von Linsingen & Sonehara 578 (ESA!). Roraima: Caracaraí, Parque Nacional do Viruá, na margem da Estrada Perdida, 1º25’35”N, 60º59’0”W, 62 m, 17 July 2010, Barbosa & Costa 1141 (UEC!); Caracaraí, Parque Nacional do Viruá, 1º28’6”N, 61º0’25”W, 67 m, 24 July 2010, Barbosa & Costa 1309 (UEC!); Caracaraí, Parque Nacional do Viruá, grade do PPBio, 1º20’50”N, 6º5’10”W, 23 July 2010, Pessoa et al. 338 (RB!). São Paulo : Angatuba, estrada Angatuba - Itatinga, km 30, 23 December 1997, Pansarin et al. 113 (UEC!); 1816–1821, Saint-Hilaire C2 1399 (P!). Tocantins: Palmas, Próximo a Palmas, entre a pista em direção a Tocantinia e o Rio Tocantins, Fazenda São Paulo , 17 February 1997, Batista et al. 689 (CEN!); 10 km antes de Filadelfia, na estrada para Araguaína, 15 January 1998, Oliveira & Gonçalves 262 (CEN!, UB!); Goiatins, balsa do Rio Tocantins (Palmeirante) Goiatins, km 11, 7º55’30”S, 47º50’6”W, 230 m, 21 January 2008, Pereira-Silva & Moreira 12850 (CEN!); Dianópolis, 11°37’03”S, 46°24’55”W, 635 m, 27 September 2003, Scariot et al. 922 (CEN!). Without locality, Muller (?) s.n. (P00409022!). COLOMBIA: upper Apaporis Basin, rio Macaya, cerro Chiribiqueta, 24 July 1943, Schultes 5627 (AMES!). FRENCH GUIANA: 27 December 1986, Cremers 9548 (RENZ!); Savane Bordelaise, 25 January 1983, Labat 181 (P!); 1840, Leprieur s.n. (P00408671!); 1850, Leprieur 104 (K000363785!, P00408668!, P00408669!, P00408670!); Leprieur 140 (P00409019!); 1834, Leprieur 231 (G00169196!, P00408673!); Leprieur 233 (P00408674!); Cayenne, 1859, Leprieur & Sagot s.n. (BM000032656!, P00408666!); Savane de Corossony, sur la piste de St Elie, 4 à 7 km à l’WSW de Sinnamary, 22 December 1977, Raynal-Roques 19692 (P!); 15 March 1985, Renz 14000 (RENZ!); Cayenne, 1859, Sagot s.n. (BM000032656!); Savane route de Stoupan, 1 km avant Crique Macrabo, 24 February 1968, Sastre 14 (AMES! mixed with H. sprucei , P!, RENZ!). GUYANA: Soesdyke, September 1973, Coope 299 (U!); Savannah, camp river, 1884, Jenman 1881 (K!); Potaro-Siparuni, Annai-Karupukari road, 45 km N of Surama village, 4º20’N, 58º50’W, 24 April 1992, Hoffman et al. 1432 (U!); Kaietur Plateau, 7 May 1944, Maguire & Fanshawe 23288 (NY!). PARAGUAY: San Luis, 16 January, Fiebrig 5303b (G!). SURINAME: Palaime savanna, Sipaliwini, 2º8’N, 56º12’W, 24 February 1963, Boer 795 (U!); Coesewijne savanna, 5 March 1959, Donselaar 592 (RENZ!, U!); Saramacca, Tafelberg, Kappel-Savanna, 19 February 1961, Kramer & Hekking 2936 (RENZ!, U!); Saramacca, Tafelberg, Kappel-Savanna, 19 February 1961, Kramer & Hekking 2941 (AMES! mixed with H. cruegeri , U!); Coesewijne Savanna, 28 February 1977, Lindeman et al. 173 (NY!, RENZ!, U!); Saparakreek savanna, 10 May 1966, Maas 3255 (U!); Sipaliwini savanna area on Brazilian border, 14 January 1969, Oldenburger et al. 927 (U!); Boven Sipaliwini, kamp B., 20 January 1936, Rombouts 387 (U!); Zanderij II, March 1943, Stahel s.n. (U 0205499!); Marowijne, Buffelsavanne along Wanekreek, 29 January 1972, Teunissen 1170 (U!). VENEZUELA: Ter. Amazonas: Sabana Caname, rio Atabapo, 5 km below Guarinumo, 8 June 1959, Wurdack & Adderley 42859 (AMES!, NY!); Sabana Manacal, Rio Atabapo, 12 June 1959, Wurdack & Adderley 42953 (NY!); Rio Casiquiare, above Solano, 10 July 1959, Wurdack & Adderley 43390 (NY!).

Habenaria cruegeri Cogniaux (1909: 302) . Syntypes:— TRINIDAD. Aripo, 26 May 1861, H. Crueger 49 (GOET! [GOET008447] mixed with H. mesodactyla, BR ! mixed with H. mesodactyla, TRIN ! [1827]); Piarco, 12 May 1858, H. Crueger 67 (lectotype, designated here: GOET! [GOET008449], isolectotype GOET! [GOET008448]). ( Fig. 4 ) Habenaria culmiformis Schlechter (1925a: 70) . Type:— BRAZIL. Rio Branco [Roraima]: August 1913, J.G. Kuhlmann 779 (holotype: B,

destroyed; lectotype designated by Batista et al. 2011a: RB! 3343 [00542617], isotypes AMES!, SP! 31796 [ SP002110 ] ).

Herb terrestrial, caulescent, sympodial, geophyte. Roots few, up to 6 cm long, usually shorter, 0.5–0.6 mm wide, hairy. Tuberoid ovoid or ellipsoid, 0.9–1.7 × 0.3–0.5 mm. Stem erect, sometimes slightly sinouse, 9–30(–40) cm long, including the inflorescence, 0.7–1.3 mm wide. Leaves 4–6, base sheathing, sheath closed, blade appressed to the stem or ascending, membranaceous, linear, narrow, acuminate, 2.6–10.0 × 0.2–0.35 cm. Inflorescence 1.1–9.2 cm, few flowered, lax, secund to subsecund, 0.7–1.1(–1.8) flowers/cm inflorescence; floral bracts green, ovate, caudate, 10–18.0 × 4.0– 5.5 mm, about the same size as the pedicellate ovary, decreasing in size towards the inflorescence apex. Flowers (1–)2–6(–9), resupinate, ascending, glabrous; pedicellate ovary (11–) 13–16 mm long; ovary straight, mostly parallel to the inflorescence axis, apex arched, (10.0–) 11.5–14.5 mm long, 3.0–13.0 times as long as the pedicel; pedicel 1.0–2.5(–4.0) mm long. Sepals green, smooth, discreetly mucronate; dorsal sepal concave, ovate when flattened, 2.7–3.7 × 2.0– 2.6 mm; lateral sepals slightly concave, obliquely lanceo-ovate, reflexed, 3.3–4.1(–4.5) × 1.1–1.3(–1.6) mm. Corolla green or greenish-brown, lighter at the base. Petals bipartite; posterior segment narrowly oblong, falcate, acute, 2.5–3.3 × 0.7–0.9 mm, adherent to the dorsal sepal; anterior segment filiform, erect, usually reflexed, touching each other behind the dorsal sepal, inserted at the base of the posterior segment, 2.2–3.2 mm long, 0.8–1.0(–1.1) times as long as the posterior segment. Labellum tripartite; undivided basal part short, 0.6–1.1 × 0.8–1.1 mm; lateral segments, perpendicular to the median segment, 2.0–3.0 mm long, 0.7–1.0 times as long as the median segment; median segment linear, 2.5–4.0 × 0.7–0.9 mm; spur deflexed, mostly covered by the bract, linear to slightly clavate, (7.0–)8.0–11.0 mm, (0.5–)0.6–0.8(–1.0) times as long as the pedicellate ovary, green. Gynostemium erect, about 2.0 mm high; connective emarginate, light green; lateral appendages (auricles) whitish, fleshy, verrucose, about 0.5 × 0.7 mm. Anther bilocular, loculi parallel, 1.1–1.3 mm high; canals short, about 0.3–0.4 mm long; hemipollinaria united by the viscidia; viscidia ellipsoidal, about 0.5 × 0.15 mm; caudicles 0.5 mm long. Stigmatophores (stigma lobes) 2, green, closely parallel, 1.6–1.8 mm long, receptive surface straight, facing upwards, almost perpendicular to the anther loculi, 1.0– 1.2 mm long, apex about 0.6–0.7 mm wide, obtuse. Rostellum green, about 1.1–1.3 mm long; mid-lobe triangular, fleshy, erect, obtuse, completely placed between the anther loculi, 0.6 mm high; side-lobes converging towards the apices, about 0.5 × 0.2 mm.

Distribution and Conservation Status:— Habenaria cruegeri is widely distributed, occurring in Venezuela, Trinidad, Guyana, French Guiana, Suriname, and Brazil (AP, DF, GO, MA, MG, MT, PA, PB, RN, RR, SP, TO). Based on the IUCN Red List Categories and Criteria and their guidelines ( IUCN 2012, 2016), H. cruegeri can be classified as of least concern (LC), due to its wide geographic distribution and large numbers of known populations, many of them occurring inside protected areas.

Habitat, phenology and ecology:— Habenaria cruegeri grows in seasonally to permanently wet grassland (campo limpo estacionalmente ou permanentemente úmido), in dark sandy-clay, hydromorphic soils, at elevations from sea level to 1,300 m a.s.l. Those fields usually accumulate surface water during the rainy season, but become drier to greater or lesser extents during the dry season. In central Brazil, those fields frequently occur associated with irregularly spaced earthen mounds known as murundus. Flowering occurs during the rainy season.

Notes:—As far as we could determine, H. cruegeri was first collected by Leprieur in French Guiana in 1833. At some point, Reichenbach examined some of the specimens collected by Leprieur, but misidentified them as H. leprieurii . Grisebach (1864) was the first taxonomist to produce a specific description of the species, based on specimens collected by Purdie and Crueger in Trinidad. He noticed the specimens were distinct from H. mesoadactyla , which he described in the same work—but misidentified them as H. setacea . Cogniaux (1909) noted Grisebach’ mistake and described the species as H. cruegeri , based on collections by Crueger in Trinidad in 1858 and 1861. Cogniaux’s name went largely unnoticed, however, and was rarely applied and, starting with Reichenbach, H. cruegeri was repeatedly misidentified as H. leprieurii . Renz (1992) in his treatment of the Habenaria of the Guianas, provided a good color photograph of the species, but misidentified it as H. leprieurii , which was followed by subsequent authors ( Batista & Bianchetti 2003, Foldats et al. 2003, Batista et al. 2008, 2011 a, 2013, Bianchetti et al. 2006, Carvalho et al. 2013, Léotard et al. 2017).

Among the species in H. sect. microdactylae, H. cruegeri is the most distinct. Phylogenetically, it is sister to, and well separated from, the remaining species of the clade, which are mostly placed in a polytomy. Morphologically, it is the only species in the clade with consistently green flowers. Habenaria leprieurii and H. omissa have white or greenish-white flowers, while H. heptadactyla has yellow flowers; some specimens of the latter species, however, can occasionally present green flowers. Habenaria cruegeri is also distinct by the shape and orientation of its pedicellate ovary, which is mostly straight, parallel to the inflorescence axis, and curved only at the apex—whereas it is more or less curved throughout its length in all other species of the clade. Another difference can be observed in the width of the lateral sepals: in H. cruegeri they are very narrow (1.1–1.3(–1.6) mm wide), while in the other species of the clade they are broader (1.8–3.4 mm wide in H. leprieurii , 1.4–2.1 mm wide in H. omissa , and 1.9–2.1 mm wide in H. heptadactyla ) ( Table 2).

Szlachetko et al. (2016) accepted H. culmiformis and used the spur length to distinguish it from H. leprieurii : equal to the ovary in H. culmiformis and distinctly shorter than the ovary in H. leprieurii . However, their concept of these species ( Szlachetko et al. 2016, 2017) is based on misidentifications. Between the listed material of H. culmiformis there are specimens of H. mesodactyla (Jansen-Jacobs et al. 4790 [P!]), H. cruegeri (Lanjouw & Lindeman 3053 [AMES!, K!, NY]) and a third unidentified species (Maguirre et al. 43864A [MO]), while between the cited material of H. leprieurii there are specimens of H. cruegeri (Maas & Westra 3761 [K!, P!], Maas & Westra 4030 [K!, P!], Leprieur 132 [K!, P!], Raynal & Tirel 18422 [P!], Raynal-Roques & Jérémie 21221 [P!], Renz 14004 [RENZ!]; Sastre 4781a [P!, RENZ!], Crueger 34 [W-R!]), H. leprieurii (Leprieur 231 [G!, P!]), H.omissa (Warming 149 [W-R!]) and H. heptadactyla (Langenheim 3255 [COL!]). The corresponding illustrations of both species are also misidentified: the illustrated specimens identified as H. leprieurii are H. omissa (Warming 149 [W-R!]) and H. heptadactyla (Langenheim 3255 [COL!]), while one of the specimens illustrated as H. culmiformis (Jansen-Jacobs et al. 4790 [P!]) is H. mesodactyla .

Furthermore, the examination of the types of H. cruegeri (Crueger 49 [GOET!, BR! TRIN!], Crueger 67 [GOET!]) and H. culmiformis (Kuhlmann 779 [AMES!, RB!, SP!]) confirmed that both are conspecific, as first suggested by Renz (1992). The comparison between H. culmiformis and H. goyazensis Cogniaux (1893: 77) ( Szlachetko et al. 2016) is unwarranted because the two species are very different. The type material of Habenaria goyazensis (Gardner 3995 [K!, BR!, BM!, G!, K-L!, OXF!, W-R!, RENZ!]) has lanceolate leaves up to 26 cm long and 2 cm wide, larger flowers (dorsal sepal about 7 mm long), and the spur 4 cm long, free from the bracts, among other differences. As we showed before ( Batista et al. 2014), H goyazensis is conspecific with H. bicornis Lindley (1835: 309) , which is morphologically and phylogenetically unrelated to H. cruegeri in the Neotropical clade of the genus.

Illustrations: — Mansfeld (1930, t. 2, fig. 5, as H. culmiformis , Schlechter’ original drawing of the species), Pabst & Dungs (1975, pg. 252, fig. 117, as H. culmiformis , based on Black & Lobato 50-9511 [IAN!]), Kenny 1992 (pg. 18, as H. mesodactyla ), Renz (1992, fig. 1a, as H. leprieurii ), Batista et al. (2008, fig. 4 E-G, as H. leprieurii ), Carvalho et al. (2013, figs. 5E, 6B, as H. leprieurii ), Léotard et al. (2017, fig. 9, as H. leprieurii ).

Misidentified illustrations: — Snuverink & Westra (1983, fig. 4 [based on M. &. P. Teunissen 1259 [U!], as H. culmiformis , = H. paxamorque ] and fig. 5 [based on M. &. P. Teunissen 1261 [U!], as H. culmiformis , = H. subfiliformis Cogniaux (1906: 272) ], Szlachetko et al. (2016, fig. 57, based on Jansen-Jacobs et al. 4790 [P!], as H. culmiformis = H. mesodactyla ; fig. 58, based on Maguirre et al. 43864A [MO], as H. culmiformis = Habenaria sp. ).

Additional specimens examined:— BRAZIL: Amapá. Macapá, fazenda Tucunaré, 19 April 1950, Black & Lobato 50-9511 (IAN!). Distrito Federal. Sobradinho, Colorado, 10 February 1991, Batista 182 (CEN!); Sobradinho, Chapada da Contagem, 31 March 1995, Batista & Bianchetti 558, (CEN!); Sobradinho, Chapada da Contagem, Cachoeiras de Mumunhas, 31 March 1995, Batista & Bianchetti 567 (CEN!); Brasília, Lago Sul, após a QI-17, atrás do Aeroporto, campo as margens do Córrego do Gama, 14 March 1996, Batista & Bianchetti 609 (CEN!); Samambaia, DF-280, na direção de Santo Antônio do Descoberto, dentro da área de Furnas, próximo ao limite final da cerca, ao lado da rede de condução elétrica, 7 March 1998, Batista 778 (CEN!); Brasília, área entre a QI-17 do Lago Sul e a quadra 25 do Setor de Mansões Park Way, um pouco acima do Córrego do Gama, possivelmente dentro da Reserva Ecológica do IBGE, 3 March 2001, Batista & Bianchetti 1195 (CEN!, EAN!); Sobradinho, Chapada da Contagem, Fazenda Dois Irmãos, 8 February 1999, Bertioli 32 (CEN!); Brasília, Setor de Mansões do Lago Norte, 17 January 1990, Bianchetti & Batista 843 (CEN!); Brasília, Estação Ecológica Jardim Botânico de Brasília, Área do Cristo Redentor, 3 February 2012, Chacon 900 (HEPH!); Brasília, Estação Ecológica Jardim Botânico de Brasília, Área do Cristo Redentor, 13 February 2012, Chacon 908 (HEPH!); Núcleo Bandeirante, lado direito da estrada que liga o Riacho Fundo ao restaurante da Fazenda Sucupira, 18 February 2000, Guarino & Pereira 24 (CEN!, SP!); Núcleo Bandeirante, Fazenda Sucupira, 18 April 2000, Guarino et al. 116 (CEN!, SP!); Núcleo Bandeirante, campos à direita da estrada que liga o Riacho Fundo ao restaurante da Fazenda Sucupira, 9 March 2001, Guarino et al. 656 (CEN!, HUEFS!); Núcleo Bandeirante, lado direito da estrada que liga o Riacho Fundo ao restaurante da Fazenda Sucupira, 8 March 2001, Guarino & Pereira 658 (CEN!); Gama, Saia Velha, 31 April 1964, Heringer 9655 (UB!); Sobradinho, NE of Lago Paranoá, near Ribeirão do Torto, 26 April 1966, Irwin et al. 15364 (UB!); Taguatinga, Taguatinga Norte, próximo ao Setor de Oficinas, as margens da BR-070 na saída para Cocalzinho, 15 February 2001, Miranda 78 (CEN!, EAN!, UEC!); Country Club, 12 February 2012, Pastore 4055 (HUEFS!); Brasília, Estação Ecológica Jardim Botânico de Brasília, 10 February 2006, Salles 3989 (HEPH!); Brasília, Estação Ecológica Jardim Botânico de Brasília, Cristo Redentor, 24 February 2006, Salles 4012 (HEPH!); Núcleo Bandeirante, fazenda Sucupira, região do vale do Córrego do Riacho Fundo, próximo a estrada na entrada sul, 3 April 2000, Walter et al. 4433 (CEN!, SP!). Goias. Pirenópolis, 31 March 1990, Batista & Salles 69 (CEN!); Cavalcante, noroeste de Cavalcante, 21 February 1997, Batista et al. 706 (CEN!, BHCB!); Cavalcante, Área a noroeste de Cavalcante, cerca de 20–25 km na margem da estrada para a região conhecida localmente como São Domingos , não a cidade de São Domingos , já na parte de cima da chapada, 21 February 1997, Batista 713 (CEN!, SP!); Cristalina, campo à esquerda da BR-040, cerca de 5.2 km N da entrada para Cristalina em direção a Brasília, 14 March 1999, Batista & Bianchetti 922 (UEC!); Alto Paraíso de Goiás, Chapada dos Veadeiros, área à direita da GO-118, na altura do km 157–158, ca. de 9 km ao S e antes de Alto Paraíso, 25 March 2001, Batista et al. 1207 (CEN!); Teresina de Goiás, Chapada dos Veadeiros, área à esquerda da GO-118, na altura do km 204, ca. de 37 km ao N e após Alto Paraíso, 25 March 2001, Batista et al. 1222 (BHCB!, CEN!, UEC!); Alto Paraíso de Goiás, Chapada dos Veadeiros, 5 February 1979, Gates & Estabrok 156 (UB!); Alto Paraíso, GO-12, Chapada dos Veadeiros, 20 February 1975, Hatschbach et al. 1066 (MBM!); Alto Paraíso de Goiás, Chapada dos Veadeiros, 1200–1500 m, 20 February 1975, Hatschbach et al. 36335 (MBM!); Cavalcante, área a noroeste de Cavalcante, cerca de 20–25 km na margem da estrada para a região conhecida localmente como São Domingos , não a cidade de São Domingos , já na parte de cima da chapada, 21 February 1997, Oliveira et al. s.n. (UB0018822!); Cavalcante, Caminho para a cachoeira Santa Bárbara, 7 March 2003, Pastore 393 (HUEFS!); Cavalcante, 5 May 2003, Pastore & Suganuma 578 (CEN!); Cristalina, entrada para a linda Serra dos Topázios, 8 February 2012, Pastore 3906 (HUEFS!); Cavalcante, estrada Vila Veneno-rio São Felix , km 3, 330 m, 22 February 2001, Pereira-Silva et al. 4789 (CEN!); Pirenópolis, Morro do Cabeludo, 31 March 1990, Salles 1658 (HEPH!); Cristalina, 11 April 1992, Salles 1846 (HEPH!); São Gabriel de Goiás, 1249 m, 1 March 2005, Salles et al. 3697 (HEPH!); Cristalina, 7 February 2002, Simões et al. 1194 (UEC!). Maranhão. Carolina, P.N. Chapada das Mesas , acesso E no km 612 da BR-230, 20 km em estrada vicinal em direção à localidade Buenos Aires, Ribeirão das Lages, 6 April 2016, Simon et al. 2726 (CEN!); Carolina, Parque Nacional da Chapada das Mesas , acesso N no km 519 da BR-230, 13 km N. Gleba 2, 9 April 2016, Simon et al. 2845 (CEN!). Mato Grosso. Parque Nacional, Casa de Pedra, Chapada, April 1911, Hoehne 4104 (R!); Parque Nacional, Casa de Pedra, Chapada, April 1911, Hoehne 4105 (R!); Vila Bela da Santíssima Trindade, Parque Estadual da Serra de Ricardo Franco, Fazenda Vale Verde, 8 March 2011, Mota 2171 (BHCB!). Minas Gerais. São Roque de Minas, Parque Nacional da Serra da Canastra, 12 March 2000, Batista & Lemos 1074 (CEN!, HUFU!, SP!); Conceição do Mato Dentro, Área de Proteção Ambiental Morro da Pedreira, 31 March 2003, Batista 1403 (CEN!); Presidente Kubitschek, 2 April 2003, Batista 1422 (CEN!); Presidente Kubitschek, 2 April 2003, Batista 1437 (CEN!, SP!, UEC!); Carrancas, Pedreira do Guilherme, 22 April 2006, Batista et al. 1624 (BHCB!); Presidente Kubitschek, 15 February 2007, Batista et al. 1937 (BHCB!); Serro, Milho Verde, 16 February 2007, Batista et al. 1954 (BHCB!); Joaquim Felício, Parque Estadual da Serra do Cabral, 3 March 2009, Batista 2810 (BHCB!); Diamantina, Parque Nacional das Sempre Vivas , 6 March 2009, Batista 2836 (BHCB!); Conceição do Mato Dentro, Parque Nacional da Serra do Cipó, 12 March 1999, Batista & Bianchetti 894 (CEN!); Carrancas, 20 January 2010, Borges 21 (BHCB!); São Sebastião do Paraíso, Baú, 26 April 1945, Brade & Barbosa 17837 (RB!); Jaboticatubas, Parque Nacional da Serra do Cipó, 5 March 1972, Joly et al. 1172 (UEC!); Santana do Riacho, Parque Nacional da Serra do Cipó, 29 January 2007, Mota 3363 (BHCB!); Patrocínio, 26 February 2013, Rocha & Fernandes 946 (BHCB!); Jaboticatubas, Parque Nacional da Serra do Cipó, 6 February 1972, Sazima 13398 (UEC!). Pará. Marapanim, vila de Marudá, praia do Crispim, 16 June 1991, Bastos et al. 1062 (MG!); Maracanã, Ilha de Maiandeua, 13 June 1994, Bastos et al. 1680 (MG!); Vigia, estrada para Vigia, 16 May 1952, Fróes 27840 (SP!); Vigia, Campina do Palha, próximo de Belém, 12 June 1966, Pabst 8963 (AMES!, K!, M!); Conceição do Araguaia, 20 km W of Redenção, near Córrego São João , troncamento Santa. Tereza, 10 February 1980, Plowman et al. 8661 (MG!). Paraíba: Mamanguape, cabeceira do Rio dos Patos, Sema I, 1 September 1981, Félix & Santana 2243 (JPB!, UFP!); Mamanguape, Capim Azul, Sema I, 13 June 1991, Félix et al. 3941 (JPB! mixed with Habenaria sp. ); Estação Ecológica Mamanguape, Capim Azul, 18 August 1988, Félix & Miranda s.n. (JPB 8441!). Rio Grande do Norte: Rio do Fogo, área militar de Rio do Fogo, 5º39’58”S, 35º37’92”W, 10 m, 28 August 2015, Garcia & Gonçalvez 4 (UFRN!); Rio do Fogo, área militar de Rio do Fogo, próximo Lagoa da Cotia, 5º23’33”S, 35º22’07”W, 30 m, 24 April 2016, Garcia & Gonçalvez 155 (UFRN!); Maxaranguape, Pititinga, 5º22’49”S, 35º21’50”W, 14 m, 22 July 2014, Moura & Soares 189 (UFRN!). Roraima. Boa Vista, campo à margem direita da BR-174, no sentido de Boa Vista para Pacaraima, cerca de 2.5 km após a ponte sobre o rio Truarú, na altura aproximada do km 566, 8 October 2005, Batista 1591 (BHCB!, CEN!); Boa Vista, em campo de lavrado próximo à EMBRAPA, próximo à lagoas temporárias, 17 July 2002, Bianchetti 1719b (CEN!); Boa Vista, localidade Monte Cristo, chácara do Sr. Cleber Barroso, 12 October 1991, da Silva 246 (MG!). São Paulo. Pedregulho, Estreito, Usina de Estreito , acampamento de estreito, 18 March 2003, Sasaki et al. 243 (SPF!). Tocantins. 18 km após Natividade, na estrada para Conceição do Tocantins, 20 February 1997, Batista et al. 701 (CEN!); Serra do Lajeado, Fazenda Três Irmãos, 2 April 1998, Batista & Oliveira 790 (CEN!, UB!); Natividade, cerca de 18 km após Natividade, área à margem direita da Pista na estrada para Conceição do Tocantins, 20 February 1997, Oliveira et al. s.n. (UB 87305!); Natividade, 19 km de Natividade, estrada para Conceição do Tocantins, 20 February 1997, Santos et al. 844 (HTINS!). FRENCH GUIANA: 1834, Leprieur 132 (K000363791!, P00408638!); 1833, Leprieur 256 (G!); 1840, Leprieur s.n. (P00408672!); Leprieur s.n. (W-R! 51238 mounted on the same sheet with a drawing of Leprieur 132, RENZ! photocopy of W-R); Cayenne, 1859, Leprieur & Sagot s.n. (P00408667!); Savane du Gallion, 20 km SSE de Cayenne, 4 March 1976, Raynal & Tirel 18422 (P!); Savane de Corossony, 5 km à l’ W de Sinnamary, 18 April 1979, Raynal-Roques & Jérémie 21221 (P!); 15 March 1985, Renz 14004 (RENZ!); 13 March 1985, Renz 14005 (RENZ!); Savane Changement, 6 km de Sinamary, 16 April 1976, Sastre 4781a (P!, RENZ!). GUYANA: Rupununi, E Kanuku Mts, 20 July 1983, Determann 235 (SEL!); Saramacca, Tafelberg, Kappel-Savanna, 19 February 1961, Kramer & Hekking 2941 (AMES! mixed with H. leprieurii ); Rupununi, Manari, 3º28’N, 59º41’W, 22 October 1979, Maas & Westra 3761 (K!, MO, NY, P!, S!, U!); Rupununi, Chaakoitou, near mountain point, just S of Kanuku Mts, 2º56’N, 59º40’W, 25 October 1979, Maas & Westra 4030 (K!, MO, NY, P!, S!, U!); Ayanganna Plateau, Upper Mazaruni River basin, Haieka savannah, E side of Haiekariver, E of Chinowieng village, 740 m, 21 July 1960, Tillett et al. 44892 (NY! mixed with H. ayangannensis, RENZ ! photocopy). SURINAME: kappelsavannah, E of Tafelberg, 8 July 1981, Determann 174 (SEL!); Brokopondo District, 4 km E. of Village Brownsweg, 22 December 1965, Donselaar 2841 (U!); Brokopondo District, 4 km E. of Village Brownsweg, 8 March 1966, Donselaar 3221 (RENZ!, U!); savana at km 116.5, 23 April 1949, Lanjouw & Lindeman 3053 (AMES!, K!, NY, U!, RENZ!, SEL!); Brinckheuvel-Saban pasi savanne, 16 January 1968, Teunissen 12174 (U!). TRINIDAD: St. Andrew, Cumuto, Aripo Savanna, 2 km E. of Cumuto, 10 o 35’N, 61 o 12’W, 40 m, 28 July 1976, Barnard et al. 385 (K!); Aripo Savanna, 29 September 1964, Bhorai B.861 (TRIN!); Aripo Savanna, 10 September 1963, Carrick 907 (K!); Crueger 34 (W 51241!); Crueger s.n. (TRIN! 1629); Piarco, Dannouse s.n. (TRIN! 35079); Finlay s.n. (TRIN! 3299); Aripo Savanna, 18 February 1915, Freeman s.n. (TRIN! 7961); Aripo Savanna, 15 July 1963, Jermy 2284 (K!); Aripo Savanna, 16 August 1963, Kee 10567 (P!); Long Stretch Savanna, 26 July 1977, Philcox & Andrews 8057 (K!); Purdie 34 (W-R!); Aripo, Jul 1848, Purdie 118 (K000363870!, K000363871!); Purdie s.n. (K000363792!, K000363872!); Aripo Savanna, 17 July 1983, Quesnel s.n. (TRIN! 28646); Aripo Savanna, 14 August 1959, Richardson 682 (K!); Aripo Savanna, 10 August 1961, Richardson 968 (K!); Aripo Savanna, near Cumuto, 14 August 1959, Webster et al. 9655 (S! mixed with H. heptadactyla, TRIN !); Aripo Savanna, Williams s.n. (NY!, TRIN! 11317); without collector (TRIN! 883). VENEZUELA: Bolívar, Gran Sabana, Santa Elena de Uairén, 9 October 2005, Batista 1595 (BHCB!, CEN!). WHITHOUT LOCALITY: Hooker 1040 (K000363873!).

Figures

FIGURE 3. Habenaria leprieurii. A. Habit. B. Leaf. C–J. Dissected perianth, bract, pedicellate ovary, gynostemium and spur. K. Inflorescence. L–N. Flowers, lateral view. M. Flowers, front view. O. Flowers, ¾ lateral view. P. Gynostemium, front view. A. Vale et al. 221 (BHCB). B, D, F, O. Batista 815 (CEN). C. Batista et al. 689 (CEN). E. Batista & Pansarin 1147 (CEN). G–H, N. Batista et al. 2524 (BHCB). I. Batista 241 (CEN). J. Batista & Lemos 500 (CEN). K. Batista & Bianchetti 399 (CEN). L. Batista & Bianchetti 3105 (BHCB). M, P. Batista 2384 (BHCB). Photographs by J.A.N. Batista.

FIGURE 4. Habenaria cruegerii. A. Habit. B. Leaf. C–G. Dissected perianth, bract, pedicellate ovary, gynostemium and spur. I. Inflorescence. J. Flower, lateral view. K–L. Flower, front view. M. Gynostemium, front view. Habenaria cruegeri var. flaviflora. H. Dissected perianth, bract, pedicellate ovary, gynostemium and spur. N. Flower, front view. A. Mota 3363 (BHCB). B. Batista et al. 701 (CEN). C, I, K. Bianchetti & Batista 843 (CEN). D.?. E. Batista 778 (CEN). F. Batista & Salles 69 (CEN). G. Batista 1595 (BHCB). H, N. Salles 1547 (HEPH). J, L, M. Batista 2810 (BHCB). Photographs by J.A.N. Batista.