Sacoila espinhacensis J.A.N.Bat. & Meneguzzo, 2025

Sacoila espinhacensis J.A.N.Bat. & Meneguzzo , sp. nov.

( Figures 2–4 View FIGURE 2 View FIGURE 3 View FIGURE 4 )

Type: — BRAZIL. Minas Gerais: Diamantina, Guinda , 1378 m, 22 Jan 2020 (fl.), J.A.N. Batista, G.M. Cruz-Lustre & E.L.F Menezes 3666 (holotype: BHCB barcode BHCB204978 !; isotypes: CEN, MBM, RB, SP) .

Sacoila espinhacensis is similar to S. lanceolata in general morphology, but distinguished by leaves present at anthesis, with prominent hyaline margins, pseudopetiolate, and proportionally wider (length/width ratio 2.5–6.8), flowering during the rainy season, rachis short, congested, often pyramidal, and flowers orange. It is also vegetatively similar to S. cerradicola , but differs in larger flowers (e.g. lip 16.5–25 mm long), short rachis, pedicel evident, flowers fully orange, and geographical distribution restricted to the Espinhaço range in the states of Minas Gerais and Bahia.

Description: — Herb terrestrial. Roots mANy, cA. 5–6 mm IN dIAmEtEr, fAScIculAtE, tubErouS, flEShy, cylINdrIcAl, glabrous, whitish. Stem inconspicuous, underground. Leaves 1–3(–7), 6–16(–25) × 1.2–4.5 cm, length to width ratio 2.5–6.8, forming a basal rosette, present at anthesis, erect, rigid, dark green, elliptical, broadly elliptical or lanceolate, attenuate at base into a narrowly distinct pseudopetiole 2–6 cm long, apex usually acute, occasionally rounded or obtuse, margins entire, hyaline. Inflorescence 18–75 cm tall, erect, racemose; peduncle greenish to brownish, glabrous below, gradually becoming pubescent towards the apex, provided with four to seven sheaths, sheaths 2.1–6.0 × 0.5– 1. 2 cm, membranous, imbricating, light green to brownish; rachis 2.5–11.0 cm long, pubescent, flowers forming a triangular or cylindrical cluster, congested; floral bracts 11–17(–24) × 4–7(–12) mm, pubescent, orange, becoming greenish or brownish during aging, lance-ovate, acuminate, approximately the same size as the pedicellate ovary and clasping them on one side, decreasing in size and width towards the apex of the rachis. Flowers (3–)6–18(– 23), resupinate, slightly ascending, showy, fleshy, odorless, orange; pedicellate ovary 11–18 mm, orange, becoming greenish or brownish during development of the fruit, ascending, obliquely fusiform, slightly arcuate, slightly twisted, pubescent throughout, except ventrally in the part comprising the extension of the column, ovary 9.5–14.0 mm long, pedicel 1.5–5.0 mm long, slender. Sepals pubescent externally, glabrous internally, orange, trichomes elongated, linear, apically dilated; dorsal sepal 15–21 × 5–6 mm, concave, lance-ovate, acute; lateral sepals 18–24(–29) × 4–5(–6) mm, basally fused 3–5(–8) mm along ovary margins, obliquely lanceolate, acute. Petals 15–18 × 3.0– 5.5 mm, mostly glabrous, inner half longitudinally adherent to the dorsal sepal, the free part externally slightly pubescent, embracing the tunnel formed by the labellum and column, oblong-lanceolate, slightly falcate, acute to subacute, light orange. Labellum 16.5–25 mm long, when spread out 5.5–8 mm wide between the margins of the widest portion, sessile, fleshy, concave–cymbiform, forming a tunnel with the column, except for the reclined apical lobe, light orange; base clawed, claw ca. 2 mm long; hypochile 6–9 × 3–5(–7) mm, 0.33–0.47 of total lip length, externally densely pubescent, channelled, forming a central groove, when spread out oblong, provided at each side near the margin with a linear thickening (nectary?), thickening 3.5–6 mm long, internally glabrous but externally prominently pubescent, between the thickening and the epichile with a semicircular fold turned inwards, densely papillose on both sides; epichile 6–8.5 mm long, 0.28–0.40 of total lip length, base forming a slight depression, mostly glabrous, occasionally externally slightly pubescent, adherent marginally to the sides of the column, base when spread out rhombic, apex triangular, reclined, sometimes perpendicular to the rest of the labellum, slightly papillose. Column 7.5–9.5 mm long, stout, glabrous, with narrow membranaceous clinandrium margins, clinandrium 0.8–1.5 mm high, column foot free and incurved at apex; anther 5.5–7.5 mm long straight or apex slightly curved, articulate, lanceolate, acuminate, with filament completely adnate; pollinia whitish, narrowly clavate, cleft, with ventral-apical, ligulate, sheathing viscidium; rostellum, including the rostellum remnant, 3.2–4.1 mm long, narrowly triangular, stiff; stigma 2.2–2.4 × 1.9–2.5 mm, broadly obovate, oriented on an oblique plane toward the apex of the column, lower border prominent, slightly convex and with a slight depression near the lower border. Capsules not examined.

Distribution and ecology: — Sacoila espinhacensis occurs in campos rupestres ( Figure 3A–B View FIGURE 3 ) and associated vegetation as sandbanks ( Figure 3C View FIGURE 3 ) or seasonally wet grasslands ( Figure 3D View FIGURE 3 ). The soil is generally sandy, mixed with dark organic matter or peat, and is seasonally moist during the rainy season, drying out completely during the dry season. Plants grow and flower during the rainy season, when there is greater water availability, and lose their leaves and enter vegetative dormancy during the dry season, surviving on the basis of fleshy roots that are completely buried in the soil. Flowering in Minas Gerais is concentrated during the rainy season, between January and March ( Figure 5A View FIGURE 5 ), but see the taxonomic discussion of flowering time below. In the Espinhaço range of Bahia, which has a more heterogeneous rainfall regime ( Harley 1995), flowering occurs almost throughout the year, but is concentrated between June and September ( Figure 5B View FIGURE 5 ), corresponding to the months with the lowest rainfall index. However, S. espinhacensis usually occurs in the higher elevations of the Espinhaço range of Bahia, regions that have a second annual rainfall peak between May and July ( Zappi et al. 2003), which may explain this apparent difference with respect to the Espinhaço of Minas Gerais.

The new species is restricted to the Espinhaço range in the states of Minas Gerais and Bahia, Brazil ( Figure 6 View FIGURE 6 ). Of the total known records, 84 (88%) are from Minas Gerais and only 12 (12%) from Bahia, suggesting that the species is more common in the Espinhaço range of Minas Gerais than in Bahia. Similar to other orchid species restricted to the Espinhaço range in the states of Minas Gerais and Bahia, there is an apparent discontinuity in distribution between the two states ( Batista et al. 2021, Cruz-Lustre et al. 2022). It is not yet clear whether there are specific conditions favouring the occurrence of these taxa in the core areas of the Espinhaço in each state, or whether this discontinuity is due to a sampling bias or gap. The intermediate region of the Espinhaço in northern Minas Gerais and southern Bahia is certainly less well sampled. However, the Cabral Range (Serra do Cabral) ( Figure 6 View FIGURE 6 ), a discontinuous part of the Espinhaço range and separated from the Diamantina Plateau by just 30 km, has no records of the species. Although the Cabral Range is fairly well sampled, it is drier than the Diamantina Plateau, the latter with numerous records of S. espinhacensis , suggesting that climatic factors also influence the distribution of the species.

Etymology: —In reference to the Espinhaço mountain range in Brazil, Serra do Espinhaço or Cadeia do Espinhaço in the local language, which stretches about 1,100 km from north to south through the states of Minas Gerais and Bahia, where the species is restricted to.

Conservation assessment:— Sacoila espinhacensis is currently known from 96 records from 40 localities and has an estimated EOO of 70,046 km 2 and an AOO of 172 km 2. Based on the IUCN Red List Categories and Criteria and their guidelines ( IUCN 2012, 2016), the new species can be provisionally classified as Least Concern (LC) due to its wide geographic distribution and known occurrences at more than 10 localities, many of which are within legally protected areas.

Illustrations:— Toscano de Brito & Cribb (2005: 248 and 366, as S. lanceolata ), Salazar et al. (2018: Figure 1A View FIGURE 1 , as S. hassleri ).

Taxonomic discussion: —The first collections of S. espinhacensis date back to the beginning of the 19th century, around two centuries ago. August A. C. P. de Saint-Hilaire collected the species between 1816 and 1821, and C. F. P. von Martius in 1818, both in the state of Minas Gerais in south-eastern Brazil. Lindley (1840b) was probably the first orchid taxonomist to examine material of S. espinhacensis and identified it as Stenorrhynchos orchioides ( Swartz 1788: 118) Richard (1817: 37) , now a synonym of Sacoila lanceolata . Later, Cogniaux (1893 –1896) identified the species as Stenorrhynchos orchioides and Stenorrhynchos hassleri Cogniaux (1906: 534) , Hoehne (1945) identified it as Stenorrhynchos bradei Schlechter in Schlechter & Hoehne (1922: 30) (currently a synonym of Sacoila duseniana ( Kraenzlin 1911: 28) Garay (1982: 351)) , while G. F. J. Pabst identified it mainly as Stenorrhynchos hassleri , occasionally also as Stenorrhynchos lanceolatum or Stenorrhynchos bradei . In Flora e Funga do Brasil, the most recent treatment of the Brazilian flora, specimens of S. espinhacensis were included under Sacoila hassleri ( Guimarães 2025) . Lindley (1840b) was also the first taxonomist to make notes on S. espinhacensis , noting that in the Martius herbarium, now in the Munich herbarium (M), there were specimens from the province of Minas Gerais in flower with leaves, in contrast to S. lanceolata , which is notable for flowering earlier than the leaves appear. In the same work, Lindley presciently stated that there were probably two or three different species under Stenorrhynchos orchioides , but that he had no means of distinguishing between them ( Lindley 1840b).

After 190 years, the situation reported by Lindley (1840b) still exists and the differentiation of Sacoila species is difficult. A taxonomic revision of Sacoila is beyond the scope of this work, but our phylogenetic and taxonomic study of the genus allows some preliminary results, including the recognition as a new species of this taxon restricted to the Espinhaço range in Minas Gerais and Bahia. The main distinguishing characteristics of S. espinhacensis are leaves pseudopetiolate, flat, usually elliptic, length to width ratio 2.5–6.8, with a conspicuous hyaline margin, present at anthesis, rachis short (2.5–11 cm long), congested, and flowers orange ( Table 2 View TABLE 2 ). This species flowers during the rainy season and occurs exclusively in the Espinhaço Range, in rocky field vegetation between 800 and 1600 m, mostly between 1200 and 1500 m. Leaf size is quite variable ( Figure 4 View FIGURE 4 ) and is probably related to the age of the plants, the growth cycle and the microclimatic conditions in which they grow. We have no data to evaluate the first two parameters, but in relation to the last one, plants with more developed leaves are generally found in wetter, shadier locations and/or with a higher content of organic matter in the soil.

Among the species with which S. espinhacensis has been confused, S. hassleri also has leaves during flowering, and it was probably this characteristic that led to the misidentification of S. espinhacensis as S. hassleri by several authorities ( Cogniaux 1893 –1896, Pabst & Dungs 1975, Guimarães 2025). However, S. hassleri differs in vegetative morphology by having long-pseudopetiolate, lanceolate (length/width ratio 4.5–18.7), conduplicate leaves and generally elongated rachis, (6–)10–15(–17) cm long ( Table 2 View TABLE 2 ). Beyond the phylogenetic distance ( Figure 1 View FIGURE 1 ), S. lanceolata differs by flowering during the dry season throughout its geographical range, leaves lanceolate, proportionally narrower, length/width ratio 3.5–8.5(–10.4), absent during flowering, inflorescences usually elongated, flowers red, pink or salmon and geographical distribution throughout the Neotropical region (central Argentina to Florida in the USA), generally at altitudes below 1000 m ( Table 2 View TABLE 2 ). Finally, S. cerradicola is also similar to S. espinhacensis in vegetative morphology, habitat, and leaves with margin hyalin, but differs in the elongated inflorescence, (4.5–)7–12(–18) cm long, smaller flowers (e.g. lip 8.5–12 mm long vs. 16.5–25 mm long in S. espinhacensis ), flowers with pedicel subssessile (pedicel evident in S. espinhacensis ), sepals orange, petals and lip whitish (sepals, petals, and lip orange in S. espinhacensis ), and distribution restricted to Chapada dos Veadeiros in the state of Goiás (Serra do Espinhaço in S. espinhacensis ) ( Table 2 View TABLE 2 ). These two species are allopatric, separated by at least 500 km along in an east-west straight line.

In the southern part of the Espinhaço range in the state of Minas Gerais, in the Serra do Cipó region, there are several herbarium specimens (Batista et al. 2103— BHCB; Duarte 11076 —BHCB, ESA; Joly & Semir in CFSC 3133 —SP and 3163 —SP; Mello Barreto 4851— BHCB and 4878 —BHCB, SP; Menezes 1354 —SPF; Menezes in CFSC 3483 — SP and 6556 —SP; Sazima 13411 —IBGE, UEC; Semir 13426 —IBGE, UEC) that are similar to S. espinhacensis , with short, congested inflorescences, orange flowers and from rupestrian fields. However, these specimens differ in that they have no leaves during flowering and flower during the dry season ( Figures 3E View FIGURE 3 , 4D View FIGURE 4 ). These specimens correspond to the August and September flowering records in Figure 5A View FIGURE 5 . We examined the flowers of some of these specimens and found no differences from S. espinhacensis , suggesting that they belong to this species. If this conclusion is correct, it shows that some specimens can occasionally deviate from the species-typical pattern of flowering during the rainy season and with leaves present at anthesis. At least one of these specimens (Batista et al. 2103 —BHCB, Figure 4D View FIGURE 4 ) was collected in an area that had previously been burned ( Figure 3E View FIGURE 3 ), suggesting that flowering may have been stimulated by bushfires during the dry season, as is characteristic of other species in the genus, such as S. lanceolata and S. pedicellata ( Cogniaux 1903: 932) Garay (1982: 352) . Unfortunately, we were unable to obtain DNA from any of these specimens and a definitive comparison with S. espinhacensis will have to await the availability of suitable material for DNA extraction and inclusion in molecular phylogenetic analyses.

Although the taxonomic circumscription of Sacoila remains unresolved due to the paraphyly of the genus ( Salazar et al. 2018, and this paper), we follow the concept of Garay (1982) here to assign the genus of the new species. A reassessment of the generic circumscription of Sacoila is in progress and will be published elsewhere.

Additional specimens examined (Paratypes): — BRAZIL. Bahia: Ibicoara, Pau Ferrada ( Batava )/ Baixão , 12 January 2005 (fl.), R. Funch 469 ( HUEFS!); Lençóis, Serra da Chapadinha , 900 m, 31 August 1994 (fl.), M.L. Guedes et al. in PCD 710 ( ALCB!); ibidem, 915 m, 29 July 1994 (fl.), H.P. Bautista et al. in PCD 325 ( ALCB!); Mucugê, Morro do Beco , Chapada Diamantina , 15 September 2006 (fl.), A.A. Conceição et al. 1829 ( HUEFS!); Mucugê, Guiné, Serra do Esbarrancado , 1415 m, 11 April 2008 (fl.), A.A. Conceição 2855 ( HUEFS!); Mucugê, Chapadinha , 6 February 2005 (fl.), R. Funch 669 ( HUEFS!); Mucugê, Chapada Diamantina , posto 2, 19 September 1998 (fl.), M.L. Guedes et al. 6121 ( ALCB!); Mucugê, estrada para Andarai , 22 June 1996 (fl.), M. Luceño et al. 328 ( UFP!); [Mucugê e Andaraí ], Chapada da Calabocaria , 1300 m, 16 June 1976 (fl.), R. Windisch & A. Ghillány 557 ( HB!); Piatã, Serra de Santana , 1309 m, 3 November 1996 (fl.), L.P. de Queiroz et al. in PCD 3995 ( ALCB!); Serra do Sincorá, s.d., s.c. (P00366513!). MINAS GERAIS: Botumirim , Campina do Bananal , 1 March 2000 (fl.), M.F. Vasconcelos & S.A. Neto s.n. ( BHCB 50873 !); Cocais [ Barão de Cocais ], Serra do Brucutú , 6 February 1943 (fl.), M. Magalhães 2703 ( BHCB!); [ Barão de Cocais ], lower slopes of Serra da Caraça , ca. 10 km W. of Barão de Cocais , 1500 m, 27 January 1971 (fl.), H.S. Irwin et al. 29246 ( NY!, UB!); Caraça, January 1885 (fl.), Jobert 80 ( P!); Carassa [ Caraça ], April 1818 (fl.), C.F.P. Martius s.n. ( M!); Serra do Caraça , March 1892 (fl.), E. Ule 4015 ( HBG!); Catas Altas, Serra do Caraça , Campo de Fora , 21 February 2006 (fl.), J.A.N. Batista & R.C. Mota 1607 ( BHCB!); ibidem, 1478 m, 18 February 2009 (fl.), B.M. Carvalho & J.A.N. Batista 16 ( BHCB!); ibidem, 1444 m, 10 February 2010 (fl.), B.M. Carvalho et al. 46 ( BHCB!); Catas Altas, Serra do Caraça , trilha para o Pico do Sol , 1724 m, 21 March 2009 (fl.), B.M. Carvalho 32 ( BHCB!); Catas Altas, Serra do Caraça , 2 April 2000 (fl.), R.C. Mota 771 ( BHCB!); ibidem, 1800 m, 20 March 2005 (fl.), R.C. Mota 2814 ( BHCB!); Catas Altas, Serra do Caraça , Campo de Fora , 2 April 2000 (fl.), J. Ordones 96 ( BHZB!); ibidem, J. Ordones 103 ( BHZB!); ibidem, 17 February 2001 (fl.), J. Ordones 729 ( BHZB!); Catas Altas, Serra do Caraça , 15 March 1998 (fl.), M.F. Vasconcelos s.n ( BHCB 41413 !); Catas Altas, Serra do Caraça , 1350 m, 17 February 2000 (fl.), M.F. Vasconcelos s.n ( BHCB 52820 !); Cocaes, Serra do Garimpo , 13 January 1921 (fl.), F.C. Hoehne s.n. (SP 4971 !); encosta norte do Monte Boa Vista, Serra da Conceição ( Serra B. Vista ), 6 March 1982 (fl.), N. Hensold et al. in CFCR 2887 (SP!); Conceição do Mato Dentro, Parque Natural Municipal do Ribeirão do Campo , 28 December 2002 (fl.), R.C. Mota et al. 2535 ( BHCB!); Conceição do Mato Dentro, Parque Natural Municipal do Ribeirão do Campo , 8 November 2002 (fl.), R.C. Mota et al. 2569 ( BHCB!); [Conceição do Mato Dentro], estrada de Conceição , 7 August 1933 (fl.), H.L. Mello Barreto 4851 ( BHCB!); Conceição do Mato Dentro, Parque Estadual da Serra do Intendente , na parte de cima do parque, 1378 m, 7 August 2007 (fl.), J.A.N. Batista et al. 2103 ( BHCB!); Congonhas do Norte, Serra Talhada ( Braço Norte da Serra do Cipó ), 9,6 Km SW da estrada de Congonhas do Norte-Gouveia , 19 January 2004 (fl.), J.R. Pirani et al. 5162 ( SPF!); Congonhas do Norte, Serra Talhada ( Braço Norte da Serra do Cipó ), 6 Km SW da estrada de Congonhas do Norte-Gouveia , 1266 m, 4 February 2009 (fl.), J.R. Pirani et al. 5759 ( CTES!, SPF!); Datas , 23 February 1983 (fl., fr.), I. Cordeiro & E. Simonis in CFCR 4044 ( SPF!, U!); entre Datas e Gouveia, Serra do Sonho , 8 January 1978 (fl.), A. Ghillány s.n. ( HB 66805 !); Diamantina, cerca de 20-21 km do entroncamento com a BR-367, km 171, em direção a Conselheiro Mata , 12 January 2000 (fl.), J.A.N. Batista & K. Proite 1016 ( CEN!); Diamantina, MG-220, cerca de 20-21 km do entroncamento com a BR-397, 1233 m, 6 January 2011 (fl.), J.A.N. Batista et al. 3137 ( BHCB!); Diamantina, subida para a usina, 16 January 1963 (fl.), A.P. Duarte & Graziela 7950 ( RB!); Diamantina, Campus JK da UFVJM, 1308 m, 30 January 2011, I.M. Franco & T.Q. Araújo 671 ( DIAM); Diamantina, 40 km NW de Diamantina, 1200 m, 15 February 1972 (fl.), A. Ghillány s.n. ( HB 57917 !); Diamantina, 12 km SW of Diamantina, 1370 m, 18 January 1969 (fl.), H.S. Irwin et al. 22162 ( F!, HB!, IAN!, NY!, UB!); Diamantina, 12 km NE of Diamantina, road to Medanha , 1300 m, 27 January 1969 (fl.), H.S. Irwin et al. 22702 ( F!, HB!, IAN!, NY!, UB!); [Diamantina], 3 km N. of São João da Chapada, 1200 m, 24 March 1970 (fl.), H.S. Irwin et al. 28305 ( NY!, UB!); Diamantina, Distrito de Sopa (tromba d’anta), caminho para a caverna do salitre, 18 February 1996 (fl.), I. Koch 440 ( UEC!); Diamantina, estrada para Conselheiro Mata , Km 192, 2 April 198? (fl.), S.J. Mayo et al. in CFCR 10429 ( SPF!); Diamantina, rodovia BR-259 para Gouveia, 28 February 1998 (fl.), J.R. Pirani et al. 4042 ( SPF!); Diamantina, 7 km E de Diamantina na estrada para Canto de Magalhães de Minas , 1294 m, 8 January 2003 (fl.), L.P. de Queiroz et al. 7564 ( HUEFS!); Diamantina, km 184 da MG 220 na direção de Conselheiro Mata , 7 February 2009 (fl.), J.G. Rando et al. 787 ( CTES!, SPF); Diamantina, 15 January 1947 (fl.), D. Romariz 119 ( RB!); Diamantina, APA Pau de Fruta (COPASA), 12 February 2001 (fl.), J.R. Stehmann et al. 2841 ( BHCB!); Diamantina, estrada para Conselheiro da Mata , km 175 e 176, 10 February 1999 (fl.), F.A. Vitta & R. Belinello 588 ( UEC!); Diamantina, estrada Conselheiro Mata-Diamantina , km 182, 29 January 1986 (fl.), D.C. Zappi et al. in CFCR 9413 (SP!, SPF!); Gouveia, hidroelétrica de Paraúnas , January 2012 (fl.), D.Q. Domingos & Wilder 63 ( ESAL!); Gouveia, BR-259, 10 km S de Gouveia, 22 January 1978 (fl.), G. Hatschbach 40815 ( CTES!, MBM!, NY!, UEC!, UC!, US); Jaboticatubas, Serra da Catinga , 5 March 1963 (fl.), M. Magalhães 18964 ( HB!); Jaboticatubas, Serra do Cipó , Km 132-5, 10 September 1972 (fl.), N.L. Menezes in CFSC 3483 (SP!); Jaboticatubas, Serra do Cipó , Km 132, 21 August 1972 (fl.), A.B. Joly & J. Semir in CFSC 3133 (SP!); Jaboticatubas, Serra do Cipó , Km 142, 22 August 1972 (fl.), A.B. Joly & J. Semir in CFSC 3163 (SP!); Jaboticatubas, Km 129 da rodovia Lagoa Santa , 3 September 1972 (fl.), J. Semir 13426 ( IBGE!, UEC!); Jaboticatubas, Km 126 da rodovia Lagoa Santa a Conceição do Mato Dentro, 4 September 1972 (fl.), M. Sazima 13411 ( IBGE!, UEC!); Presidente Kubitschek , km 442 da BR-259, 15 February 2007 (fl.), J.A.N. Batista et al. 1930 ( BHCB!); Presidente Kubistschek, estrada Datas-Serro , 35 Km de Datas , lajeado ao lado na nascente do Jequitinhonha , 1 January 2000 (fl.), R.C. Forzza & R. Mello-Silva 1517 ( SPF!); Presidente Kubitschek, estrada BR 259 a Serro, 9,7 km de Datas , 1170 m, 18 February 2003 (fl.), F. França et al. 4558 ( HUEFS!); Santa Bárbara, Caraça , caminho para gruta do Padre Caio , 14 March 1990 (fl.), W. Marcondes-Ferreira et al. 216 ( UEC!); Santa Luzia, Serra do Cipó , km 134, estrada do Pilar , 24 August 1933 (fl.), H.L. Mello Barreto 4878 ( BHCB!, SP!); Santana do Riacho, Parque Nacional da Serra do Cipó , afloramento das Vellozia Gigantea, 21 February 2016 (fl.), J.A.N. Batista et al. 3378 ( BHCB!); Santana do Riacho, P.N. da Serra do Cipó , Alto Palacio , junto às Velozias gigantes, 3 February 1994 (fl.), E.L. Borba 97 ( BHCB!); Santana do Riacho, RPPN Brumas do Espinhaço e Ermo do Gerais , 1223 m, 30 January 2013 (fl.), C.A. Ferreira Junior et al. 934 ( BHZB); [Santana do Riacho], Serra do Cipó , km 120, 1200 m, 14 February 1968 (fl.), H.S. Irwin et al. 20036 ( F!, HB!, IAN!, NY!, RB!, UB!, US); [Santana do Riacho], Serra do Cipó , summit, km 145, 1500 m, 22 February 1968 (fl.), H.S. Irwin et al. 20633 ( AMES!, F!, HB!, IAN!, MO, NY!, RB!, UB!, US); [Santana do Riacho], entre posto Palácio e Morro do Pilar , February 1953 (fl.), J. Vidal 5999 ( R!); Santa Luzia [Santana do Riacho], Serra do Cipó , 2 February 1934 (fl.), A.J. Sampaio 6806 ( R!); Santa Luzia [Santana do Riacho], Serra do Cipó , km 136, estrada do Pilar , 3 February 1934 (fl.), A.J. Sampaio 6899 ( BHCB!, R!); Santana do Riacho, Serra do Cipó , Km 126, 7 September 1980 (fl.), N.L. Menezes in CFSC 6556 (SP!); [Santana do Riacho], Serra do Cipó , km 138, 29 August 1968 (fl.), A.P. Duarte 11076 ( BHCB!, ESA!); Santana do Riacho, rodovia Belo Horizonte — Conceição do Mato Dentro, antigo km 142, 15 September 1996 (fl.), N.L. Menezes 1354 ( SPF!); [Santo Antônio do Itambé], eastern slopes of Pico do Itambé , 1550 m, 12 February 1972 (fl.), W.R. Anderson et al. 35848 ( HB!, IAC, NY!, UB!); [Santo Antônio do Itambé], Itambé , 1816-1821, A. de Saint-Hilaire s.n. ( P00366518 !); Serro, estrada para Capivari , próxima ao Pico do Itambé , 1220 m, 22 February 2013 (fl.), M. Monge 1553 ( UEC!); São Gonçalo do Rio Preto, caminho a Pindaíba , 1601 m, 29 February 2008 (fl.), A.J. Ramalho et al. 30 ( BHCB!); Serro, Milho Verde , Lageado , 1099 m, 16 February 2007 (fl.), J.A.N. Batista et al. 1961 ( BHCB!); Serro, MG-2, entre Serro e Datas , 19 January 1972 (fl.), G. Hatschbach et al. 28907 ( HB!, MBM!, NY!, UC!, US); Serro, 18 km north of Serro on road to Diamantina, 1200 m, 23 February 1968 (fl.), H.S. Irwin et al. 20656 ( AMES!, F!, HB!, IAN!, M!, MO, NY!, UB!, US); Serro, Milho Verde , Serra do Ouro , 1 February 2003 (fl.), R.C. Mota 2610 ( BHCB!); [Serro], Milho Verde , a cerca de 10 km de Serro, 1200 m, 14 February 2001 (fl.), E.R. Pansarin & A.O. Simões 830 ( UEC!); Serro, ca. 11 km de serro em direção a milho verde, 1000 m, 11 March 1995 (fl.), V.C. Souza et al. 8336 ( ESA!); Serro, Capivari , 13 January 1998 (fl.), M.F. Vasconcelos s.n. ( BHCB 40158 !); Serro e Datas , 5 February 1980 (fl.), P.G. Windisch 2606 ( HB!, M!); without locality, 1816-1821, A. de Saint-Hilaire B1-925 ( P!); ibidem, 1816-1821, A. de Saint-Hilaire s.n. ( P00366519 !); ibidem, 29 February 1976 (fl.), R. Windisch & A. Ghillány 479 ( HB!).