Croton longinervius var. major Müller Argoviensis (1873: 215),

Sodré, Rodolfo C., Berry, Paul E. & Da Silva, Marcos J., 2017, The tribe Crotoneae (Euphorbiaceae, Crotonoideae) in the Chapada dos Veadeiros, Goiás, Brazil, Phytotaxa 321 (1), pp. 1-59 : 9-45

2.1. Croton abaitensis Baillon (1864: 337) . Type:— BRAZIL. Minas Gerais: Abaité, 1844, M. Weddell 1818 (holotype

P 00623674!, isotypes A 00047215!, F V0056098F!). ( Figs. 4 , A–E; 5 , A–C)

Croton longinervius Müller Argoviensis (1873: 215) — Croton longinervius var. major Müller Argoviensis (1873: 215) , syn. nov. Lectotype (designated here):— BRAZIL. Goiás: Serra de Christaës, no date, J. B. E. Pohl 826 (W 0051292!, isolectotype G 00434553!).

Croton longinervius var. minor Müller Argoviensis (1873: 216) , syn. nov. Lectotype (designated here):— BRAZIL. Goiás: ad Rio S. Marcos prope S. Luzia, no date, J. B. E. Pohl 1637 (W 0051293!, isolectotypes A 00257952!, G 00434554!, K 000186104!, K 000252628!, W 0051294!).

Croton megaponticus Müller Argoviensis (1873: 210) , syn. nov. Lectotype (designated here):— BRAZIL. Goiás: prope Megaponte, no date, J. B. E. Pohl 1110 (W 0004076!, isolectotypes G 00434616!, F 24501!).

Subshrubs 0.3–1.2 m tall, erect, stems greenish, yellowish or ferrugineous covered by stellate-porrect trichomes; latex clear. Stipules 0.7–3.5 × 0.2–0.8 mm, lanceolate to linear, with ovoid to ellipsoid glands at the base, persistent; petioles 0.4–1 cm long, with 2(–4) shortly stipitate patelliform yellowish glands. Leaves alternate or whorled near to dichotomy of the branches, blades 1.8–8 × 1.3–4.3 cm, oval or sometimes elliptic, base obtuse or cordate, apex acute, acuminate or obtuse, margin serrate with shortly stipitate patelliform glands in the sinuses; upper and lower surfaces with stellate-porrect trichomes, membranaceous, venation actinodromous-brochidodromous, 2 or 3 (4) pairs of secondary veins, the basal pair of veins surpassing half the length of the blade. Thyrses 4.5–14.5 cm long, terminal, staminate cymules with 1–3 flowers, inflorescence axis, both faces of pistillate sepals and bracts of both flowers, dorsal surface of staminate sepals, ovary and styles with stellate-porrect trichomes; staminate and pistillate bracts 2–5 × 0.2–0.8 mm, linear to lanceolate, without glands. Staminate flowers 5–8.5 mm long, pedicels 2–4.5 mm long; sepals 5, 2–2.3 × 1.2–1.8 mm, ovate, apex acute or obtuse; petals 5, 2.4–2.9 × 0.8–1.2 mm, oblanceolate to spatulate, apex obtuse, villous on both surfaces; stamens 11, filaments villous. Pistillate flowers 5–8 mm long, pedicels 1–2 mm long; sepals 5, 2.5–4.8 × 0.7–5 mm, unequal, obovate, widely elliptic, ovate or less often oblong or lanceolate, apex obtuse, acuminate or acute, margins entire without glands; petals 5, 0.4–0.8 × 0.05–0.1 mm, linear, disk 5-lobed; ovary 1.2–1.7 mm long, globose, styles 2-fid, 3.5–5 mm long, pubescent. Capsules 3.5–4 mm long, globose, brownish, columella lacking prominent, smooth lobes; seeds 3–3.3 × 2–2.2 mm, ellipsoid, uniformly dark brown.

Specimens examined:— BRAZIL. Goiás: Água Fria de Goiás, a direita do km 61 da GO 118, de São Gabriel a São João d’Aliança , 14°59’9.2”S, 47°36’12.3”W, 1108 m, 11 July 2013, M. J. Silva et al. 5079 ( UFG). Alto Paraíso de Goiás, acima dos alojamentos do Parque, 14°9’32.2”S, 47°47’27”W, 1093 m, 10 March 2012, R. C. Sodré et al. 78 ( UFG), 79 ( UFG), 80 ( UFG), 81 ( UFG), 86 ( UFG) ; ibd., M. J. Silva et al. 4161 ( UFG), 4162 ( UFG), 4163 ( UFG) ; ibd., 23 November 2012, M. J. Silva et al. 4543 ( UFG), 4544 ( UFG) ; ibd., 14 December 2012, R. C. Sodré et al. 403 ( UFG), 404 ( UFG), 405 ( UFG), 406 ( UFG), 407 ( UFG), 408 ( UFG) ; ibd., M. J. Silva et al. 4642 ( UFG), 4643 ( UFG), 4645 ( UFG), 4646 ( UFG), 4647 ( UFG), 4648 ( UFG), 4649 ( UFG) ; ibd., 08 February 2013, M. J. Silva et al. 4681 ( UFG), 4682 ( UFG) ; imediações do Posto de Combustível Vale da Lua , 14°8’19.3”S, 47°31’23.8”W, 1241 m, 16 February 2013, R. C. Sodré & M. T. Faria 553 ( UFG), 555 ( UFG), 556 ( UFG) ; GO 118 , km 155, entre São João d’Aliança e Alto Paraíso de Goiás, 14°12’49.4”S, 47°29’15.9”W, 1122 m, 10 October 2013, M. J. Silva et al. 5408 ( UFG) ; a esquerda do km 156 da GO 118, de Alto Paraíso de Goiás para São João d’Aliança , 14°12’49.4”S, 47°29’15.9”W, 1182 m, 03 November 2013, R. C. Sodré et al. 985 ( UFG), 990 ( UFG), 991 ( UFG), 997 ( UFG), 998 ( UFG), 999 ( UFG), 1000 ( UFG) ; GO 118 , km 153, entre São João d’Aliança e Alto Paraíso de Goiás, 14°14’3.8”S, 47°29’12.2”W, 31 October 2013, M. J. Silva et al. 5479 ( UFG), 5482 ( UFG), 5484 ( UFG), 5485 ( UFG), 5486 ( UFG), 5488 ( UFG), 5489 ( UFG), 5490 ( UFG) ; a esquerda do km 131 da GO 118, de São João d’Aliança a Alto Paraíso de Goiás, próximo a divisa destes municípios, 14°25’20.7”S, 47°30’35”W, 1000 m, 20 February 2014, R. C. Sodré et al. 1220 ( UFG) ; a direita do km 131 da GO 118, de São João d’Aliança a Alto Paraíso de Goiás, 14°25’18.3”S, 47°30’27.4”W, 1005 m, 20 February 2014, M. J. Silva et al. 5854 ( UFG) ; cerca de 30 km a esquerda da GO 118, de São João d’Aliança a Alto Paraíso de Goiás na estrada para a Catarata do rio dos Couros , 14°17’35”S, 47°42’32”W, 1028 m, 02 January 2015, R. C. Sodré et al. 1575 ( UFG), 1577 ( UFG), 1581 ( UFG), 1584 ( UFG) ; ca. 1 km após a casa dos pesquisadores na estrada que leva aos alojamentos do Parque , 14°8’37.2”S, 47°46’6.5”W, 1141 m, 04 January 2015, R. C. Sodré et al. 1624 ( UFG). São João d’Aliança, a direita da GO 118, a 300 metros do Atos Hotel, sentido Alto Paraíso de Goiás, 14°41’35.8”S, 47°31’17.3”W, 1047 m, 28 January 2014, M. J. Silva et al. 5722 ( UFG), 5723 ( UFG) ; a direita da GO 118, após o hotel Chapéu de Sol , 14°41’35.3”S, 47°31’19.1”W, 1005 m, 28 January 2014, R. C. Sodré et al. 1166 ( UFG), 1167 ( UFG) .

Croton abaitensis occurs in cerrado sensu stricto and grasslands in the Federal District, and in Goiás, Mato Grosso and Minas Gerais states. In Chapada dos Veadeiros it was found growing in cerrado sensu stricto, grassland and cerrado rupestre in the municipalities of Água Fria de Goiás, Alto Paraíso de Goiás and São João d’Aliança ( Fig. 3B ), flowering from October to February and fruiting from January to March.

This species can be recognized by its leaves with actinodromous-brochidodromous venation with two to four pairs of veins, the first pair reaching more than half the blade, subsessile acropetiolar glands and also along the leaf margins, pistillate sepals with entire margins, and staminate flowers with oblanceolate-spatulate petals and villous stamens. It resembles C. spica Baillon (1864: 362) by the leaf blade with crenate or serrate margins, long staminate bracts (2.6 to 6 mm), linear or narrowly lanceolate pistillate flowers with at least two foliaceous obovate or widely elliptic sepals pubescent on both surfaces. However, C. spica has brochidodromous venation with 4–10 pairs of secondary veins, the first pair never reaching more than half the blade, sessile acropetiolar and marginal glands, serrate pistillate sepals, oblong-elliptic staminate petals and glabrous stamens.

Taxonomic notes:— The W sheets of C. megaponticus and the varieties of C. longinervius were chosen as lectotypes because the original Pohl collections are housed at W, and the G specimens appear to be fragments removed from the W sheets. In each case, the lectotype was seen and annotated by Müller.

Croton abaitensis , the two varieties of C. longinervius , and C. megaponticus share leaf blades with actinodromous-brochidodromous venation, with the first pair of secondary veins reaching more than half of the blade (Müller called this “limbo palmatinervio”) as well as the acropetiolar, patelliform, subsessile glands, linear-lanceolate bracts, staminate petals and the pubescent filaments, and the shortly pedicellate pistillate flowers with ovate, equal or rarely unequal sepals and 2-fid styles. Croton abaitensis belongs to C. section Geiseleria .

2.2. Croton agoensis Baillon (1864: 348) . Type:— BRAZIL. Minas Gerais: Brasilia Meridionalis, prope Olho d’Agua, 1816–1821, A. F. C. P. Saint-Hilaire s.n. (holotype P 00623665!, isotypes A 00047218!, P 00623664!). ( Fig. 5 , D and E)

Subshrubs 0.4–0.9 m tall, erect, stems yellowish to ferrugineous with sessile or stipitate, multiradiate-stellate-porrect trichomes; latex clear. Stipules 0.1–1 × 0.2–0.4 mm, ovate, linear or subulate, without glands, persistent; petioles 0.1– 0.7 cm long, with (2)4 stipitate-patelliform glands at the apex. Leaves alternate, blades 3.4–4.4 × 1.8–2.9 cm, ovate or oval-lanceolate, base lightly cordate, apex apiculate, margin crenate or serrate with stipitate-capitate glands, upper surface with stellate-porrect trichomes and lower surface with multiradiate-stellate trichomes, chartaceous, venation brochidodromous, with 3–5 pairs of secondary veins. Thyrses 1.9–6.5 cm long, terminal, staminate cymules with 3–6 flowers, inflorescence axis, both surfaces of pistillate sepals and bracts of both flowers, dorsal surface of staminate sepals, pistillate petals, ovary and styles with multiradiate-stellate-porrect trichomes, sessile or stipitate; bracts of both flowers 1.7–2.9 × 0.2–0.4 mm, linear or linear-lanceolate or lanceolate, without glands. Staminate flowers 5.5–6.5 mm long, pedicels 2–2.9 mm long; sepals 5, 1.9–2.2 × 1–1.3 mm, ovate, apex acute or obtuse; petals 5, 2.2–2.8 × 0.8–1.5 mm, oblong, oblong-oblanceolate or obovate, apex obtuse or rounded, ventral surface villous, dorsal surface glabrous, ciliate on the margin and barbellate near the base; stamens 11, filaments villous. Pistillate flowers 3.7–5 mm long, pedicels 0.4–1.2 mm long; sepals 5(6), 2.1–3.5 × 0.8–1.1 mm, subequal, narrowly triangular, lanceolate or linear-lanceolate, apex apiculate, margins entire with stipitate-capitate glands; petals 5, 0.5–2.1 × 0.1–0.3 mm, linear or linear-lanceolate, disk 5-segmented; ovary 1.2–1.4 × 1.5–1.6 mm long, oblong, styles 2-fid, pubescent. Capsules 5.8–6.1 mm long, oblong, yellow-brownish; seeds 3.7–4 × 2.8–3 mm, ovoid, uniformly brownish.

Specimens examined:— BRAZIL. Goiás: Alvorada do Norte, cerrado ao lado do Parque de Exposições, saída para Guarani , 12 October 2012, J. P. Santos & M. Y. C. Sena 609 ( UFG) ; cerrado logo depois da entrada da estrada de chão ao rancho Hous Bear , 12 October 2012, J. P. Santos & M. Y. C. Sena 622 ( UFG) ; topo do morro próximo ao loteamento, na entrada da cidade, 12 October 2012, J. P. Santos & M. Y. C. Sena 659 ( UFG). Cavalcante, estrada em direção à Reserva Natural Serra do Tombador, ca. 33 km do município de Cavalcante , 13°34’3.2”S, 47°31’26.6”W, 1173 m, 06 September 2016, J. A. Oliveira, T. P. Mendes & A. O. Souza 144 ( UFG), 145 ( UFG) .

According to Carneiro-Torres (2009), Croton agoensis occurs in cerrado in the states of Bahia, Goiás, Maranhão, Mato Grosso, Minas Gerais, Piauí, and Tocantins. In the Chapada dos Veadeiros region it is known from a single collection from the Natural Reserve of Serra do Tombador ( Fig. 3B ), flowering in September.

This species is similar to C. catariae Baillon (1864: 349) and C. gracilirameus M. J. Silva, Sodré & P. E. Berry in Silva et al. (2015: 162) in its leaf blades with serrate or crenate margins and stipitate-capitate acropetiolar glands, cordate or lightly cordate base, and usually lanceolate pistillate sepals with stipitate-capitate glands. However, C. agoensis differs from them by the lack of glands on the stipules and bracts and the pistillate pedicels 0.4–1.2 mm long (vs. stipules and bracts of both flowers with stipitate-capitate glands and pistillate pedicels 5–9 mm long in C. catariae and C. gracilirameus ). Croton agoensis belongs to C. section Geiseleria .

2.3. Croton agrarius Baillon (1864: 319) . Lectotype (designated here):— BRAZIL. Minas Gerais: without locality, no date, 1838, M. P. Claussen 764 (P 00623663!). Syntype:— BRAZIL. Minas Gerais: without locality, 1816–1821, A. F. C. P. Saint-Hilaire cat. C 1, 210bis (P 00623661!, P 00623662!). ( Figs. 4 , F–H; 5 , F and G)

Subshrubs 0.6–1.1 m tall, erect, stems yellowish to ferrugineous with subsessile stellate-porrect trichomes; latex clear to orange. Stipules 0.3–2.8 × 0.1–0.6 mm, linear or ovate, with globose or ovoid glands at the base, persistent; petioles 0.3–1.1 cm long, without glands. Leaves alternate or opposite near the dichotomy of the branches, blades 4.6–11.7 × 2.1–5.8 cm, ovate to elliptic, sometimes obovate, oblanceolate or lanceolate, base lightly cordate or obtuse, apex mucronate or obtuse, margin entire without glands; upper and lower surfaces with stellate-porrect trichomes, membranaceous, venation brochidodromous, with 6–14 pairs of secondary veins. Thyrses 2–18.3 cm long, terminal, staminate cymules with 2–4 flowers; inflorescence axis, both surfaces of pistillate sepals, bracts of both flowers, dorsal surface of staminate sepals, pistillate petals, ovary and styles with stellate-porrect trichomes; bracts of both flowers 3.5–5 × 0.9–2.2 mm, oval-lanceolate or lanceolate with globose glands. Staminate flowers 5–7 mm long, pedicels 1–2.2 mm long; sepals 5, 2.4–3 × 1.3–1.8 mm, ovate, apex acute or obtuse; petals 5, 2.6–3 × 0.7–1.1 mm, oblanceolate or obovate, apex obtuse, ventral surface villous at the base; stamens 15–16, filaments glabrous. Pistillate flowers 5.5–7 mm long, pedicels 0.3–1 mm long; sepals 5, 2.7–3.3 × 0.7–1.6 mm, subequal, ovate to lanceolate, apex acute, margins entire without glands; petals 5, 0.3–1.2 × 0.1–0.2 mm, linear, disk subentire or 5-lobed; ovary 1.7–2.2 mm long, globose, styles 2-fid, ascending, pubescent. Capsules 5–5.5 mm long, globose, ferrugineous-green, covered by subsessile trichomes, columella with three prominent, smooth lobes; seeds 3.9–4.1 × 2.5–2.7 mm, oblong-ellipsoid, uniformly gray.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, próximo ao Vale da Lua , 14°10’47.9”S, 47°47’38.1”W, 969 m, 20 January 2012, R. C. Sodré et al. 14 ( UFG) ; ibd., 14°10’35.7”S, 47°47’18.1”W, 999 m, 20 January 2012, R. C. Sodré et al. 16, 17 ( UFG) ; ibd., M. J. Silva et al. 4056 ( UFG), 4057 ( UFG), 4058 ( UFG). Cavalcante, 1 a vertente do rio Carmo , 13°24’17”S, 48°6’54”W, 320 m, 12 December 2000, G. Pereira-Silva & J. B. Pereira 4437 ( CEN) ; ca. 8 km south of Cavalcante , 07 March 1969, H. S. Irwin et al. 24007 ( UB) ; imediações da estrada que leva a Colinas do Sul, ca. 18 km de Cavalcante, 29 January 2014, R. C. Sodré et al. 1187 ( UFG). Colinas do Sul , estrada UHE Serra da Mesa / Colinas do Sul , 13°53’S, 48°18’W, 410 m, 09 December 1991, B. M. T. Walter et al. 951 ( CEN) ; 2 Km da estrada Serra da Mesa / Colinas do Sul , 13°50’S, 48°17’W, 410 m, 12 December 1991, B. M. T. Walter et al. 1082 ( CEN) ; reservatório do AHE Serra da Mesa, 13°56’S, 48°17’W, 420 m, 28 January 1997, B. M. T. Walter et al. 3653 ( CEN). Monte Alegre de Goiás, 33 Km a SW de Campos Belos rumo a Monte Alegre, 13°12’S, 47°8’W, 565 m, 21 November 1984, A. C. Allem et al. 3073 ( CEN). Niquelândia, encosta da Serra Negra, 14°18’S, 48°27’W, 460 m, 23 November 1992, R. F. Vieira et al. 1356 ( CEN). Nova Roma, imediações do povoado de Aurominas, 13°37’2.4”S, 47°3’22.7”W, 404 m, 01 November 2014, R. C. Sodré et al. 1516 ( UFG), 1518 ( UFG). Teresina de Goiás, 7 Km by road S of Teresina, 17 Mar 1973, W. R. Anderson et al. 7313 ( UB) ; ca. 10 km do município, em direção a Alto Paraíso de Goiás, 21 February 2014, R. C. Sodré et al. 1237, 1238, 1240 ( UFG). Sem município, 41 Km North of Veadeiros , 17 Mar 1969, H. S. Irwin et al. 24547 ( UB) .

Croton agrarius was reported by BFG (2015) to occur in the Federal District and in Goiás, Mato Grosso and Minas Gerais states. We found it in cerrado sensu stricto, campos sujos or degraded pastures in all municipalities except Campos Belos and São João d’Aliança ( Fig. 3B ), flowering and fruiting from November to March.

This species was identified by Sodré et al. (2014) as C. heliotropiifolius Kunth (1817: 83) and has been often misidentified as C. campestris Saint-Hilaire (1827 : tab. 60) in herbaria from midwestern Brazil. However, after consulting the type collections and diagnoses of both species, we conclude that C. agrarius differs from these species in having a subshrubby habit up to 1 m tall, yellowish and velutinous branches and leaves with stellate-porrect and subsessile trichomes, short petiolate leaves (the petiole up to 1.1 mm long), leaf blades with an obtuse or slightly cordate base, inflorescences congested, and pistillate sepals oval to lanceolate with inner surface glabrescent. Croton agrarius belongs to C. section Adenophylli .

2.4. Croton antisyphiliticus Martius in Spix & Martius (1823: 282). Type:— BRAZIL. São Paulo: campis editis e.g. ad Ypanema , January 1818, C. F. P. von Martius s.n. (holotype M 0086128!). ( Figs. 4 , I–M; 5 , H and I)

Subshrubs 15–50 cm tall, erect, barely branched, stems brownish with stellate-porrect trichomes; latex clear. Stipules 3.5–5 × 0.8–2 mm, oval-lanceolate or oblong, without glands, persistent; petioles 0.3–1.5 cm long, with 2(4) sessile, patelliform glands at the apex. Leaves alternate, blades 5–13.2 × 1.3–6.5 cm, oval-lanceolate, oblong or elliptic, base acute, obtuse or truncate, apex acute, margin doubly serrate with sessile patelliform glands; upper and lower surfaces with fasciculate and stellate-porrect trichomes respectively, chartaceous, venation brochidodromous, with 4–5 pairs of secondary veins. Thyrses 3.5–16.5 cm long, terminal, staminate cymules with 1–3 flowers; inflorescence axis, dorsal surface of bracts, sepals of both flowers and ovary with stellate-porrect trichomes; staminate bracts 1.5–2.5 × 0.3–0.7 mm, oblanceolate or narrowly elliptic; pistillate bracts 1.4–3.5 × 0.4–0.9 mm, lanceolate or oblong. Staminate flowers 4–5.5 mm long, pedicels 2–3 mm long; sepals 5, 2.3–2.8 × 1.3–1.6 mm, ovate, apex acute; petals 5, 2–2.5 × 0.7–0.8 mm, oblong to oblanceolate, apex obtuse, ventral surface villous; stamens 11, filaments glabrous. Pistillate flowers 4–7 mm long, pedicels 0.5–2 mm long; sepals 5, 2.8–4 × 1.2–2 mm, subequal, ovate to oblong, apex acute, margins entire without glands; petals absent, disk 5-lobed; ovary 1.3–2 mm long, globose, hispid, styles 4(2)-fid, glabrous. Capsules 4.7–5.5 mm long, globose, brownish; seeds 3.5–4 × 2.2–2.5 mm, ellipsoid, uniformly dark brown.

Specimens examined:— BRAZIL. Distrito Federal: Brasília , 16°36’01”S, 49°15’38”W, 1108 m, 20 October 2011, M. J. Silva et al. 3795 ( UFG). Goiás: Abadiânia, a 200 m do Restaurante Jerivá sentido Goiânia-Brasília, 16°11’1.5’’S, 48°41’27.1’’W, 1038 m, 23 October 2011, M. J. Silva et al. 3893 ( UFG). Água Fria de Goiás, GO 118, km 83, entre São Gabriel e São João d’Aliança, 14°47’48.8”S, 47°33’26.3”W, 1073 m, 05 September 2013, R. C. Sodré et al. 845 ( UFG), 846 ( UFG) ; a esquerda do Km 82 da GO 118, entre São Gabriel e São João d’Aliança , 03 November 2013, R. C. Sodré et al. 1002 ( UFG). Alto Paraíso de Goiás, 14°09’48”S, 47°35’35”W, 1155 m, 18 December 2004, E. Chaves et al. 160 ( UB) ; trilha de acesso ao Morro da Baleia , 14°07’47.7”S, 47°38’21.9”W, 1220 m, 28 September 2012, R. C. Sodré et al. 165 ( UFG), 166 ( UFG), 167 ( UFG), 168 ( UFG), 169 ( UFG), 170 ( UFG), 171 ( UFG), 172 ( UFG), 173 ( UFG) ; cerca de 15 Km de Alto Paraíso de Goiás, a direita da GO 239, em direção a Vila de São Jorge , 14°09’20.1”S, 47°38’7.5”W, 1145 m, 25 October 2012, R. C. Sodré et al. 206 ( UFG), 207 ( UFG), 208 ( UFG) ; ibd., 14°09’29.64”S, 47°37’54.9”W, 1168 m, 13 December 2012, R. C. Sodré et al. 371 ( UFG), 372 ( UFG), 374 ( UFG) ; na entrada da fazenda Primavera, 14°23’26.04”S, 47°31’33.18”W, 1202 m, 13 December 2012, R. C. Sodré et al. 365 ( UFG). Cavalcante , estrada do “Porto dos Paulistas”, 13°28’06”S, 48°7’8”W, 410 m, 07 November 2000, B. M. T. Walter et al. 4541 ( CEN) ; canteiro de obras da Hidrelétrica Cana Brava, 13°23’45”S, 48°7’45”W, 370 m, 23 January 2001, G. Pereira-Silva et al. 4583 ( UFG, CEN). São João d’Aliança, GO 118, cerca de 15 Km após São João d’Aliança, em direção a Alto Paraíso de Goiás, 27 September 2012, R. C. Sodré et al. 150 ( UFG) ; GO 118 , 14°41’36.1”S, 47°31’17”W, 1160 m, 25 October 2012, R. C. Sodré et al. 197 ( UFG), 198 ( UFG), 200 ( UFG), 203 ( UFG) .

This species is widely distributed in Brazil, Bolivia, and Paraguay ( Carneiro-Torres 2009, BFG 2015). In this study, it was collected in campos sujos, cerrado sensu stricto in the municipalities of Água Fria de Goiás, Alto Paraíso de Goiás, Cavalcante and São João d’Aliança ( Fig. 3C ), flowering and fruiting from September to January.

The short habit (up to 50 cm tall), brownish branches, usually dark green leaves with the upper surface rough to the touch and pistillate flowers with hirsute ovary and 4(–2)-fid styles make C. antisyphiliticus easily distinguished from C. gracilescens Müller Argoviensis (1865: 34) and C. desertorum Müller Argoviensis (1873: 265) , which it resembles by the short petioles (up to 1.5 cm long) with a pair of sessile glands, small staminate bracts (up to 2.5 cm long), pistillate flowers with equal or subequal sepals and usually ovate and pubescent externally and glabrous to glabrescent styles. Croton antisyphiliticus belongs to C. section Geiseleria .

2.5. Croton arirambae Huber (1914: 182) . Type:— BRAZIL. Pará: Alto Ariramba, 20 December 1906, A. Ducke s.n. (holotype MG 008027!, isotype G). ( Fig. 6 , A–D)

Subshrubs 0.5–1 m tall, erect, stems ferrugineous with stipitate stellate-porrect trichomes; latex clear. Stipules 4– 5.5 × 0.3–0.5 mm, linear, with stipitate-capitate glands, persistent; petioles 0.7–1.8 cm long, without glands. Leaves alternate, blades 5.4–8.2 × 1.4–2.3 cm, oblong, lanceolate or narrowly elliptic, base acute, obtuse or asymmetric, apex acute, margin entire with stipitate-capitate glands; upper and lower surfaces with stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, with 5–7 pairs of secondary veins. Thyrses 6–11 cm long, terminal, staminate cymules with 1 flower; inflorescence axis, dorsal surface of bracts and sepals of both flowers, ovary and styles with stellate-porrect trichomes; staminate bracts 2.8–3 × 0.3–0.4 mm, narrowly lanceolate, with stipitate-capitate glands; pistillate bracts 5–6.5 × 0.5–1.1 mm, linear to narrowly lanceolate. Staminate flowers 6.8–8.5 mm long, pedicels 1.8–4.2 mm long; sepals 5, 3–3.5 × 1.3–1.7 mm, ovate, apex acute, villous on ventral surface; petals 5, 2.9–3.1 × 0.8–1.1 mm, elliptic, apex acute, villous on both surfaces; stamens 11, filaments villous. Pistillate flowers 10–15 mm long, pedicels 2–3.5 mm long; sepals 5, 5.5–10.2 × 1.4–3 mm, subequal, lanceolate, apex acute, margins entire with stipitate-capitate glands; petals 5, 2.5–3 × 0.15–0.2 mm, linear, disk 5-lobed, ovary 1.8–2 mm long, globose, styles 2-fid, pubescent. Capsules 6–6.2 mm diam., globose, brownish; seeds not seen.

Specimens examined:— BRAZIL. Goiás: Cavalcante, caminho para o povoado Kalunga, de fronte a entrada da fazenda Vicente , 03 February 2004, J. F.B. Pastore et al. 813 ( CEN, UFG) ; estrada para Prata (Kalungas), 13°23’21.2’’S, 47°42’27.2’’W, 1105 m, 14 April 2004, M. L. Fonseca et al. 5108 ( IBGE) .

Croton arirambae is endemic to Brazil and recorded from Goiás, Pará and Rondônia states ( Silva et al. 2015a). It was found in campo sujo in the municipality of Cavalcante ( Fig. 3C ), flowering and fruiting from February to April. It shares with C. catariae and C. gracilirameus the stipules, leaf margin, bracts of both flowers and pistillate sepals with stipitate-capitate glands. However, C. arirambae differs from both by the oblong leaf blades with obtuse base and entire margins (vs. ovate with cordate base and crenate to serrate margins in C. catariae and C. gracilirameus ), petiole without glands (vs. with 2–10 patelliform glands on lower surface), pistillate pedicels 2-3.5 mm long (vs. 5–9 mm) and rigid pistillate sepals (vs. membranaceous). Croton arirambae belongs to C. section Barhamia subsection Medea .

2.6. Croton betaceus Baillon (1864: 341) . Lectotype (designated here):— BRAZIL. Ceará: without locality, 1838, G. Gardner 1840 (G 00312269!, isolectotypes K 000186107!, K 000186106!, NY 00246514!, P 00623620!, P 00623621!, P 00623622!, S 07-12800!). ( Fig. 6 , E–H)

Subshrubs or shrubs 1–2 m tall, erect, stems greenish with subsessile stellate-porrect trichomes; latex clear. Stipules 0.4–1.3 × 0.3–0.6 mm, triangular to lanceolate, with subglobose glands at the base and apex, persistent; petioles 0.7–1.7 cm long, with two sessile acropetiolar patelliform glands. Leaves alternate or whorled near to dichotomy of the branches, blades 5.8–10 × 2.2–5.2 cm, ovate or rarely elliptic, base obtuse, sometimes acute, apex acute, margin entire without glands; upper surface glabrous or glabrescent, lower surfaces with stellate-porrect trichomes, membranaceous, venation brochidodromous, 9–14 pairs of secondary veins. Thyrses 7–19 cm long, terminal or in the dichotomy of branches, cymules bisexual or unisexual, staminate cymules with 1–3 flower; inflorescence axis, dorsal surface of bracts and sepals of both flowers, ovaries, and styles with stellate-porrect trichomes; bracts of both flowers 0.6–2 × 0.3–0.7 mm, ovate, lanceolate or linear, without glands. Staminate flowers 3–4 mm long, pedicels 0.7–1.2 mm long; sepals 5, 1.5–2.1 × 1.1–1.3 mm, ovate, apex acute; petals 5, 2.2–2.7 × 0.5–0.7 mm, oblanceolate, apex acute or obtuse, glabrous on both surfaces, ciliate at the base; stamens 10–11, glabrous. Pistillate flowers 2.5–3.5 mm long, pedicels 0.1–0.2 mm long; sepals 5, 1.7–2.1 × 0.6–1.1 mm, subequal, oval-lanceolate, apex acute, margins entire without glands; petals glanduliform, subglobose, disk 5-lobed; ovary 1.3–1.5 × 1.5–2 mm, globose, styles 2-fid, pubescent. Capsules 5.5–6 × 5.5–6 mm, oblongoid, greenish, columella with three prominent, smooth lobes; seeds 4.1–4.5 × 2.4–2.7 mm, oblongoid, brownish, with darker spots.

Specimens examined:— BRAZIL. Goiás: Cavalcante, estrada entre Cavalcante e Colinas do Sul , 29 January 2014, M. J. Silva et al. 5755 ( UFG), 5754 ( UFG), 5756 ( UFG) ; Colinas do Sul, Canteiro de obras da Usina de Serra da Mesa , 24 February 1991, D. Alvarenga et al. 790 ( IBGE) ; GO 239 , entre o povoado de São Jorge e Colinas do Sul , 14°13’27.4”S, 47°55’22.2”W, 550 m, 26 February 2016, R. C. Sodré & G. L. Soares-Feitosa 2018 ( UFG), 2019 ( UFG), 2020 ( UFG), 2021 ( UFG), 2022 ( UFG) ; Nova Roma, imediações do povoado de Aurominas, após o rio das Pedras , 13°37’2.4”S, 47°3’22.7”W, 404 m, 01 November 2014, R. C. Sodré et al. 1517 ( UFG) .

This species is recorded from Bolivia, Brazil (BA, CE, DF, GO, MA, MG, PA, PI) and Paraguay ( BFG 2015). It was collected at the edges of cerradão in the municipalities of Cavalcante, Colinas do Sul and Nova Roma ( Fig. 3C ), flowering and fruiting from November to February.

Croton betaceus differs from the other species in the region by its short petioles with two sessile patelliform glands, entire leaf margins without glands and upper surface usually with trichomes only along the veins, thyrses 7–19 cm long with laxly distributed flowers, bisexual or unisexual cymules and small flowers (2.5–4 mm long). Croton betaceus belongs to C. section Adenophylli .

2.7. Croton campestris Saint-Hilaire (1827 : tab. 60). Lectotype (designated by van Ee & Berry 2010):— BRAZIL. Minas Gerais: champs artificiel près Congonhas da Serra, 1816–1821, A. F.C.P. Saint-Hilaire 2155bis (P 00623075!, isolectotypes P 00623073!, P 00623074!, MPU 015220!). ( Figs. 5 , J–L; 6 , I–K)

Subshrubs 0.1–0.5 m tall, erect, stems ferrugineous, tomentose with subsessile stellate-porrect trichomes; latex clear or orange after oxidation. Stipules 0.4–1 mm, linear, with subglobose glands at the base, persistent; petioles 0.3–0.9 (1.6) cm long, without glands. Leaves alternate, blades 7.3–16.5 × 2.4–5.2 cm, oblong, oblanceolate or narrowly elliptic, base acute or attenuate, apex acute or obtuse, margin entire without glands; upper and lower surfaces tomentose with stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, 5–10 pairs of secondary veins. Thyrses 5.5–12.2 (36) cm long, terminal, flowers densely distributed in the inflorescence axis, staminate cymules with 1–3 flowers; inflorescence axis, dorsal surface of bracts and sepals of both flowers and ovary tomentose with stellate-porrect trichomes; staminate bracts 1.1–2 × 0.4–0.7 mm, ovate to lanceolate; pistillate bracts 1.5–2.5 × 0.2–0.4 mm, lanceolate. Staminate flowers 6.5–9.5 mm long, pedicels 2.5–4.2 mm long; sepals 5, 2.6–3.2 × 1.3–1.8 mm, ovate, apex obtuse, pubescent internally; petals 5, 2.9–4.1 × 0.7–1.8 mm, obovate, elliptic-obovate or oblanceolate, apex obtuse, ventral surface villous at the base; stamens 15–16 (20), filaments glabrous. Pistillate flowers 5–7.5 mm long, pedicels 0.5–2 mm long; sepals 5, 2.8–4 × 0.6–1.3 mm, subequal, ovate to lanceolate, glabrescent internally, apex acute, margins entire without glands; petals 5, unequal, 0.5–2.4 × 0.1–0.2 (0.4) mm, linear to lanceolate, disk subentire; ovary 1.8–2.5 × 2.3–3 mm, globose, styles 2-fid, pubescent. Capsules 5.5–6 × 5.8–6.5 mm, globose, brownish, columella with three prominent, smooth lobes; seeds 4.8–4.9 × 3.2–3.3 mm, ellipsoid, brownish.

Specimens examined:— BRAZIL. Distrito Federal: Brasília, Bacia do rio São Bartolomeu , quadrícula n° 141, 17 March 1980, E. P. Heringer et al. 4054 ( UFG). Goiás: Alto Paraíso de Goiás, without locality, 18 September 1977, A. Allem & G. Vieira 1062 ( UFG) ; Parque Nacional da Chapada dos Veadeiros, descida para as cachoeiras de 80 e 120 metros no rio Preto , 14°9’47”S, 47°50’2”W, 10 September 1996, R. C. Mendonça et al. 2693 ( IBGE) ; estrada que leva ao Vale da Lua , 14°10’17.9”S, 47°46’48.2”W, 1020 m, 15 October 2010, M. J. Silva et al. 3053 ( UFG) ; GO 118 , km 153, entre São João d’Aliança e Alto Paraíso de Goiás, 14°14’3.8”S, 47°29’12.2”W, 10 October 2013, R. C. Sodré et al. 872 ( UFG), 874 ( UFG), 875 ( UFG), 876 ( UFG) ; GO 118 , km 155, São João d’Aliança / Alto Paraíso de Goiás, 14°12’49.4”S, 47°29’15.9”W, 1122 m, 10 October 2013, M. J. Silva et al. 5425 ( UFG), 5437 ( UFG) ; GO 239 , entre o povoado de São Jorge e Colinas do Sul , 13 December 2013, R. C. Sodré et al. 1062 ( UFG), 1063 ( UFG), 1064 ( UFG) ; a 1.2 km da entrada para o Vale da Lua , 12 October 2013, R. C. Sodré et al. 908 ( UFG), 911 ( UFG) ; cerca de 30 km a esquerda da GO 118 (sentido São João d’Aliança a Alto Paraíso de Goiás), na estrada que leva a catarata do rio dos Couros , 14°17’23.6”S, 47°42’33.2”W, 1049 m, 31 October 2014, R. C. Sodré et al. 1492 ( UFG) ; ibd., 14°17’35”S, 47°42’32”W, 1028 m, 31 October 2014, R. C. Sodré et al. 1502 ( UFG) ; ibd., 02 January 2015, R. C. Sodré, M. J. Silva & C. H. G. Machado-Filho 1580 ( UFG) ; imediações da fazenda Campo Seco, cerca de 15 km após a vila São Jorge, em direção a Colinas do Sul , 14°13’39.9”S, 47°53’41.2”W, 700 m, 22 February 2014, R. C. Sodré, T. M. S. Melo & M. J. Silva 1247 ( UFG) ; ibd., 14°13’37.5”S, 47°53’39.3”W, 705 m, 14 December 2014, R. C. Sodré et al. 1561 ( UFG), 1562 ( UFG) ; a esquerda do km 131 da GO 118, sentido São João d’Aliança a Alto Paraíso de Goiás, próximo à divisa dos municípios, 14°25’20.7”S, 47°30’35”W, 1000 m, 20 February 2014, R. C. Sodré, T. M. S. Melo & M. J. Silva 1215 ( UFG) ; Parque Nacional da Chapada dos Veadeiros, trilha para os Saltos , a 300 metros da bifurcação para as corredeiras do rio Preto , 14°9’42.3”S, 47°50’12.6”W, 863 m, 31 October 2014, R. C. Sodré et al. 1474 ( UFG) ; ca. 1 km após a casa dos pesquisadores, 14°8’27.4”S, 47°46’2.8”W, 1115 m, 04 January 2015, R. C. Sodré, M. J. Silva & C. H. G. Machado-Filho 1631 ( UFG) ; estrada Alto Paraíso – Teresina, 11 October 1979, E. P. Heringer et al. 2490 ( IBGE). Cavalcante, reserva da fazenda Gavião , 14°8’37”S, 47°52’45.5”W, 01 November 2013, R. C. Sodré, M. J. Silva & P. H. B. Santos 947 ( UFG), 948 ( UFG), 949 ( UFG), 958 ( UFG), 960 ( UFG). Colinas do Sul , estrada de acesso a fazenda Gavião, cerca de 3 km a direita da GO 239, sentido Alto Paraíso de Goiás a Colinas do Sul , 01 November 2013, R. C. Sodré, M. J. Silva & P. H. B. Santos 925 ( UFG), 930 ( UFG), 934 ( UFG) ; ibd., 13 December 2013, R. C. Sodré et al. 1054 ( UFG) ; estrada para a fazenda Gavião, cerca de 4 km da GO 239, 14°10’1”S, 47°58’8.5”W, 650 m, 29 January 2014, R. C. Sodré et al. 1199 ( UFG). Niquelândia, estrada para o DNPM, Placa do IBAMA, 14°29’46”S, 48°27’0”W, 830 m, 19 October 1996, R. Marquete et al. 2679 ( IBGE) ; ibd., A. F. Vaz et al. 1109 ( IBGE), 1112 ( IBGE) ; estrada de chão em direção a reserva do IBAMA, ca. 5 km de Niquelândia , 14°29’46”S, 48°27’0”W, 635 m, 19 October 1996, M. L. Fonseca et al. 1236 ( IBGE) ; fazenda Engenho, ca. 11 km de Niquelândia / Dois Irmãos , cabeceira do rio Traíras , 14°40’28”S, 48°25’25”W, 680 m, 14 August 1997, M. L. Fonseca et al. 1549 ( IBGE) ; Companhia de Níquel Tocantins – CNT, estrada à esquerda da mina de níquel, ca. 2 km da entrada, 14°20’37”S, 48°23’57”W, 1040 m, 01 October 1997, R. C. Mendonça et al. 3099 ( IBGE), ibd., M. L. Fonseca et al. 1578 ( IBGE) ; próximo ao povoado de Macedo, 14°23’48”S, 48°25’59”W, 17 September 1996, M. Aparecida da Silva & C. C. S. Ferreira 3091 ( IBGE) ; morro do Cristo , 14°27’8”S, 48°27’26”W, 19 September 1996, M. L. Fonseca et al. 1195 ( IBGE) .

This species is known from Argentina, Bolivia, Brazil (AL, BA, CE, DF, ES, GO, MG, MS, PA, PB, PE, PI, PR, RJ, RN, RS, SP, TO) and Paraguay ( Carneiro-Torres 2009, Lima & Pirani 2003). However, the specimens identified as C. campestris in herbaria consulted in this study and several floristic works carried out in Brazil (e.g., Carneiro-Torres 2009, Lima & Pirani 2003, Silva et al. 2010) correspond to different species belonging to C. sect. Adenophylli such as C. agrarius , C. subferrugineus Müller Argoviensis (1865: 135) and C. subvillosus Müller Argoviensis (1873: 168) . These species needs to be carefully studied to be better defined. In this study, C. campestris was found in campo limpo, cerrado sensu stricto and cerradão in the municipalities of Alto Paraíso de Goiás, Cavalcante and Colinas do Sul ( Fig. 3D ), flowering September to January and fruiting from October to February.

Croton campestris is morphologically similar to C. subvillosus by the combination of the poorly branched, subshrubby habit up to 50 cm tall, leaves without glands and, with an acute or attenuate base, and upper surface with trichomes. Also, both species have pistillate flowers with pedicels 0.5–2.8 mm long, ovate to lanceolate sepals, developed linear petals 0.5–2.4 mm long, a subentire disk and a hirsute ovary. However, C. subvillosus differs from C. campestris by leaves with the upper surface dark green, glossy, and glabrous or sparsely pubescent; pedicels of both flowers glabrous or glabrescent; and patent styles. Croton campestris has leaves with the upper surface greenish but never glossy, and densely pubescent; the pedicels of both flower sexes are shortly tomentose, and the styles are ascending. It belongs to C. section Adenophylli .

2.8. Croton catariae Baillon (1864: 349) . Type:— BRAZIL. Mato Grosso: without locality, 1831–1833, C. Gaudichaud-Beaupré 247 (holotype: P 00623092!, isotype: A 00047263!). ( Figs. 7A–E , 8A–D ) Croton chaetocalyx Müller Argoviensis (1865: 133) . Lectotype (designated here):— BRAZIL. Goiás: ‘prope Megaponte et Villa Boa de

Goyaz’, 1837 or 1839, J. B. E. Pohl 1626 (W 0051197!, isolectotypes A 00063009!, B†, BR 8764560!, G 00434439! K 000186108!

K 000186109!, K 000186110!, M 0086112!, W 0051198!)

Subshrubs 0.4–1.1 m tall, erect, stems canescent-green to dark yellowish with stellate-porrect trichomes; latex clear. Stipules 1–4.5 × 0.3–0.8 mm, linear-lanceolate, with stipitate-capitate glands, persistent; petioles 0.9–3 cm long, with 6–10 shortly stipitate patelliform glands. Leaves alternate or opposite near to dichotomy of the branches, blades 4.7–7.8 × 3.7–5.7 cm, ovate, base cordate, apex acute or obtuse, margin irregularly crenate-serrate with stipitate-capitate glands; upper and lower surfaces with stellate-porrect trichomes, membranaceous, venation actinodromous-brochidodromous, with 4–5 pairs of secondary veins. Thyrses 3–14 cm long, terminal, staminate cymules with 1–4 flowers; inflorescence axis, both sides of bracts and pistillate sepals, dorsal surface of bracts and staminate sepals, ovary and styles with stellate-porrect trichomes; staminate bracts 1.8–2.6 × 0.3–0.7 mm, lanceolate or narrowly lanceolate; pistillate bracts 3–5 × 0.6–1.2 mm, lanceolate or narrowly lanceolate, both bracts with stipitate-capitate glands. Staminate flowers 7– 10 mm long, pedicels 4–6 mm long; sepals 5, 2.1–2.4 × 0.7–1.6 mm, ovate, apex acute; petals 5, 2–2.5 × 0.8–1.3 mm, elliptic, apex acute or obtuse, ventral surface villous; stamens 9–11, filaments glabrous. Pistillate flowers 10–19 mm long, pedicels 5.5–9 mm long; sepals 5, 5.5–9.5 × 1–1.7 mm, subequal, oblong-lanceolate, apex acute, margins entire with stipitate-capitate glands; petals absent, disk 5-lobed; ovary 1.8–2.2 mm long, globose, styles 2-fid, pubescent or glabrous. Capsules 5.5–6 mm long, oblongoid or globose, greenish or brownish; seeds 4.2–4.5 × 2.5–2.9 mm, oblongoid, brownish, spotted.

Specimens examined:— BRAZIL. Goiás: Água Fria de Goiás, a esquerda do km 56 da GO 118, sentido povoado de São Gabriel à São João d’Aliança , 15°01’42.3”S, 47°36’1.5”W, 1204 m, 19 January 2012, M. J. Silva et al. 4029 ( UFG) ; ibd., 08 March 2012, R. C. Sodré et al. 34, 37, 38 ( UFG) ; ibd., 06 March 2013, R. C. Sodré et al. 557 ( UFG), 559 ( UFG), 560 ( UFG) ; ibd., 28 January 2014, M. J. Silva et al. 5715 ( UFG) ; ibd., R. C. Sodré et al. 1164 ( UFG) ; ibd., 15°01’56.3”S, 47°36’4.5”W, 1167 m, 20 June 2013, R. C. Sodré et al. 773 ( UFG) ; a direita do km 61 da GO 118, nas imediações do Sítio Parceiros de Deus, sentido São Gabriel a São João d’Aliança , cerca de 30 km deste último, 14°59’14.5”S, 47°36’16.1”W, 1131 m, 09 May 2013, R. C. Sodré & M. J. Silva 713 ( UFG) ; ibd., 12 December 2013, M. J. Silva et al. 5631 ( UFG), 5632 ( UFG) ; ibd., R. C. Sodré et al. 1032 ( UFG), 1034 ( UFG), 1035 ( UFG), 1036 ( UFG), 1037 ( UFG), 1038 ( UFG) ; margem da GO 118, próximo a fazenda São Paulo, 06 March 2013, R. C. Sodré et al. 4744 ( UFG). Alto Paraíso de Goiás, a oeste da cidade, 20 February 1991, D. Alvarenga, B. A. S. Pereira, B. M. T. Walter & F. C. A. Oliveira 725 ( CEN) ; a esquerda do km 131 da GO 118, de São João d’Aliança a Alto Paraíso de Goiás, 14°25’20.7”S, 47°30’35”W, 1000 m, 20 February 2014, R. C. Sodré et al. 1211 ( UFG) ; ibd., 14°25’18.3”S, 47°30’27.4”W, 1005 m, 20 February 2014, M. J. Silva et al. 5852 ( UFG), 5853 ( UFG) ; próximo a fazenda Campo Seco, cerca de 15 km após a Vila São Jorge, 14°13’39.9”S, 47°53’41.2”W, 700 m, 22 February 2014, R. C. Sodré et al. 1250 ( UFG). Cavalcante , 11 km by road E of Cavalcante, 17 March 1978, W. R. Anderson 7304 ( UB) ; estrada da balsa do “Porto dos Paulistas” (no rio Tocantins) para o Buracão , ca. 800 m do rio, 13°29’44”S, 48°8’34”W, 340 m, 07 November 2000, B. M. T. Walter et al. 4529 ( CEN) ; balsa da COTERRA (rio Tocantins) vila Veneno , 13°34’16”S, 48°6’4”W, 300 m, 13 December 2000, G. Pereira-Silva & J. B. Pereira 4471 ( CEN, UFG) ; canteiro de obras da Hidrelétrica Cana Brava , 13°23’45”S, 48°7’45”W, 370 m, 23 January 2001, G. Pereira-Silva et al. 4580 ( CEN, UFG) ; estrada Vila Veneno – Serra Branca, Km 3,6, 13°34’25”S, 48°4’20”W, 380 m, 20 February 2002, G. Pereira-Silva et al. 5896 ( CEN, UFG) ; estrada Rio Traíras / Rio Custódio, Km 2, 13°16’50”S, 48°8’39”W, 280 m, 28 November 2006, G. Pereira-Silva & G. A. Moreira 11143 ( CEN, UFG) ; imediações da cidade, em direção ao Lava Pés , 13°48’12.1”S, 47°28’0.2”W, 820 m, 16 February 2013, R. C. Sodré & M. T. Faria 525 ( UFG), 526 ( UFG), 527 ( UFG) ; estrada para o povoado Kalunga , 13°45’49”S, 47°27’18.9”W, 804 m, 07 March 2013, R. C. Sodré et al. 563 ( UFG), 570 ( UFG), 571 ( UFG), 572 ( UFG) ; próximo a fazenda Gavião , 14°8’25.8”S, 47°52’21.8”W, 574 m, 12 July 2013, M. J. Silva et al. 5084-B ( UFG) ; ibd., 14°8’S, 47°53’W, 01 November 2013, R. C. Sodré et al. 954 ( UFG) ; a esquerda da GO 241 em direção a Cavalcante, na entrada da Fazenda Siriema , 13°47’10.5”S, 47°23’51”W, 802 m, 29 January 2014, R. C. Sodré et al. 1168 ( UFG) ; a direita da estrada rumo a Colinas do Sul, ca. 9 km de Cavalcante , 13°51’13”S, 47°29’31.3”W, 843 m, 29 January 2014, R. C. Sodré et al. 1174 ( UFG) ; cerca 18 km de Cavalcante, 29 January 2014, R. C. Sodré et al. 1184 ( UFG) ; imediações da Serra do Ministro, a aproximadamente 500 metros da estrada entre Cavalcante e Colinas do Sul , 13°54’34”S, 47°38’55.4”W, 29 January 2014, R. C. Sodré et al. 1194 ( UFG) ; 100 metros a direita da entrada do município de Cavalcante, vindo de Teresina de Goiás, 26 July 2014, R. C. Sodré et al. 1371 ( UFG). Colinas do Sul , cerca de 6 km da estrada em direção a Fazenda Gavião, 13 December 2013, M. J. Silva et al. 5653 ( UFG) ; estrada para fazenda Gavião , cerca de 3 km a direita da GO 239, 04 August 2013, R. C. Sodré et al. 838 ( UFG) ; ibd., 13 December 2013, R. C. Sodré et al. 1052 ( UFG) .

Croton catariae is a South American species recorded from Bolivia, Brazil (DF, GO, MT, PA) and Suriname ( Silva et al. 2015a). It was collected growing in cerrado sensu stricto, campo sujo, margin of seasonal forest and in disturbed areas in Alto Paraíso de Goiás, Cavalcante, Colinas do Sul, and Água Fria de Goiás ( Fig. 3D ). It flowers and fruits throughout the year, but especially from November to March.

This species shares with C. gracilirameus the stipitate-capitate glands on the stipules, lower surface of the leaf blade and along the leaf margins, and bracts of both flower sexes and the pistillate sepals; petioles with stipitate-patelliform glands, ovate leaf blades with cordate base and crenate to serrate margins, long pedicellate staminate and pistillate flowers (pedicels 5–9 mm and 2.2–5 mm, respectively), stamens with glabrous filaments and oblongoid capsules. However, C. gracilirameus differs from C. catariae by its slender branches, leaf blades 2.4–4 × 1.4–3.7 cm, shortly tomentose on the upper surface with tertiary veins impressed and inconspicuous, as well as branches and leaves with stellate-porrect trichomes, with a porrect ray equal to more than twice the length of the other rays, and inflorescences with one or two pistillate flowers. Croton catariae has robust branches, leaf blades 4.7–7.8 × 3.7–5.7 cm that are pannose on upper surface, with tertiary veins prominent and conspicuous on the abaxial surface, branches and leaves with stellate-porrect trichomes having a porrect ray at least five to eight times longer than the lateral rays, and inflorescences with four or more pistillate flowers. Croton catariae belongs to C. section Barhamia subsection Barhamia .

2.9. Croton cinerascens Radcliffe-Smith & Govaerts (1997: 186) . — Croton villosus Müller Argoviensis (1873: 169) ; nom. illeg. — Croton villosus var. cinerascens Müller Argoviensis (1873: 169) . Lectotype (designated here):— BRAZIL. Without locality, no date, H. W. Schott 3913 (W 0051328!, isolectotype G 00434507!). ( Fig. 7 , F–H) Croton villosus var. rufescens Muller Argoviensis (1873: 169) . Lectotype (designated here):— BRAZIL. Goiás: prope S. Pedro, no date, J.

B. E. Pohl 490, (W 0051329!, isolectotype G 00434506!). Syntype:— BRAZIL. Goiás: prope Catalão, August 1834, L. Riedel 2473*

(LE!).

Subshrubs 0.2–0.35 m tall, erect, stems brownish to ferrugineous with sessile or shortly stipitate stellate-porrect trichomes, hirsute to the touch, with a well developed porrect central ray; latex clear or orange after exposure to the air. Stipules 0.2–0.3 mm long, glanduliform, ovoid or subglobose, persistent; petioles 0.5–2.5 mm long, without glands. Leaves alternate, blades 4.7–7.8 × 2–3.4 cm, elliptic or oblong-elliptic, base acute or sometimes obtuse, apex obtuse or acute, margin entire without glands; upper and lower surfaces hirsute covered by sessile or shortly stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, with 7–8 pairs of secondary veins. Thyrses 0.9–4.2 cm long, terminal, staminate cymules with 1 or 2 flowers; inflorescence axis, dorsal surface of bracts and sepals of both flowers, ovary and styles hirsute with sessile stellate-porrect trichomes; staminate bracts 0.8–1.4 × 0.2–0.7 mm, ovate to lanceolate, with globose glands at the base; pistillate bracts 2–2.1 × 0.4–0.5 mm, lanceolate, with ovoid or filiform glands at the base. Staminate flowers 5.5–6 mm long, pedicels 1.3–1.5 mm long; sepals 5, 2.1–2.3 × 1–1.1 mm, ovate, apex acute; petals 5, 2.9–3.1 × 0.8–0.9 mm, oblanceolate, apex obtuse, ventral surface villous at the base; stamens 16, filaments glabrous. Pistillate flowers 5–6.5 mm long, pedicels 0.4–1.1 mm long; sepals 5(6), 1.6–2 × 0.2–0.8 mm, subequal, narrowly triangular, oval-lanceolate, or sometimes linear-lanceolate, apex acute, margins entire without glands; petals 5, 0.3–1.3 × 0.1–0.3 mm, linear to lanceolate, usually with ellipsoid gland at the apex, disk 5-segmented; ovary 1.5–1.6 × 1.7–1.9 mm, subglobose, styles 2-fid, pubescent. Capsules 5.7–6.3 × 6–6.4 mm, subglobose, brownish, columella with three prominent, smooth lobes; seeds 3.8–4 × 2.6–2.8 mm, ellipsoid, uniformly greyish.

Specimens examined:— BRAZIL. Goiás: Água Fria de Goiás, km 56 da GO 118, 15°2’3.3”S, 47°36’4.1”W, 1085 m, 28 January 2014, R. C. Sodré et al. 1159 ( UFG), 1160 ( UFG), 1161 ( UFG), 1162 ( UFG) ; GO 118 , entre Água Fria de Goiás e São João d’Aliança, a 14 km da fazenda Serra Verde , 19 January 2012, M. J. Silva et al. 4034 ( UFG) .

Croton cinerascens was a name proposed by Radcliffe-Smith & Govaerts (1997) to replace C. villosus Müller Argoviensis (1873: 169) , a homonym of C. villosus Forsskål (1775: 163) . Here, we confirm the occurrence of C. cinerascens in Goiás growing in campo sujo and cerrado rupestre in the municipality of Água Fria de Goiás ( Fig. 3D ), flowering and fruiting in January. Croton cinerascens is characterized by having branches, leaves and flowers with stellate-porrect trichomes with a well developed porrect ray, shortly petiolate leaves with elliptical to oblong-elliptic leaf blades and staminate flowers with pedicels 1.3–1.5 mm long. It belongs to C. section Adenophylli .

2.10. Croton desertorum Müller Argoviensis (1873: 265) . Lectotype (designated here):— BRAZIL. Minas Gerais: in deserto, no date, C. F. P. von Martius s.n. (M 0086104!, isolectotypes F V006600!, G [G00434490]). ( Figs. 7 , I–L; 8 , E–H)

Subshrubs 0.2–1.2 m tall, erect, much branched, stems yellowish to ferrugineous with shortly stipitate stellate-porrect trichomes; latex clear. Stipules 1.5–6 × 0.4–1 mm, triangular to lanceolate, with subglobose glands at the base, persistent; petioles 0.1–0.7(1.1) cm long, with 2 sessile, acropetiolar patelliform glands. Leaves alternate, blades 3.1–9.7(12.8) × 1.2–3.2(7) cm, oval-lanceolate or oblong-elliptic, base obtuse, apex acute, margins serrate with sessile patelliform glands, both surfaces with subsessile stellate-porrect trichomes, membranaceous, venation brochidodromous, 4–6 pairs of secondary veins. Thyrses 1.2–5(14.5) cm long, terminal, staminate cymules with 1(3) flowers; inflorescence axis, external surface of bracts and sepals of both flowers, and ovary stellate-porrect; staminate bracts 1–1.5 × 0.4–0.6 mm, oval to lanceolate, without glands; pistillate bracts 1.7–3.2 × 0.5–1.3 mm, oval to lanceolate, without glands. Staminate flowers 4.5–6 mm long, pedicels 0.8–1.9 mm long; sepals 5, 1.3–2.2 × 1.2–1.6 mm, ovate to widely ovate, apex acute or obtuse; petals 5, 2.3–2.8 × 0.7–1 mm, oblanceolate to elliptic, apex acute or obtuse, glabrous on both surfaces, ciliate at the base; stamens 11, glabrous. Pistillate flowers 3.5–5.5 mm long, pedicels 0.3–1 mm long; sepals 5, 2.3–3 × 1.2–2.1 mm, subequal, widely ovate, oval or oval-lanceolate, apex acute, margins entire without glands; petals ovoid (ca. 0.1 mm), filiform (ca. 0.3 mm) or laminar 1.2–2.2 × 0.2–0.3, linear, villous, disk 5-segmented; ovary 1.3–2 × 1.8–2.3 mm, subglobose, styles 2-fid, glabrous or glabrescent. Capsules 5.5–6 × 5.5–6 mm, subgloboid, green-ferrugineous; seeds 3.3–3.4 × 2.9–3.1 mm, widely ellipsoid, uniformly brownish.

Specimens examined:— BRAZIL. Goiás. Água Fria de Goiás, a direita do km 61 da GO 118, sentido São Gabriel a São João d’Aliança , 14°59’23.4”S, 47°36’16.4”W, 1119 m, 30 October 2014, R. C. Sodré, A. O. Souza & L. S. Inocencio 1470 ( UFG) ; a direita do km 60 da GO 118, sentido povoado de São Gabriel a São João d’Aliança, 14°59’14.5”S, 47°36’16.1”W, 1131 m, 24 July 2014, R. C. Sodré, R. D. Sartin & M. J. Silva 1313 ( UFG), 1314 ( UFG). Alto Paraíso de Goiás, Sítio Pedrão, 14°10’55.5”S, 47°50’45.6”W, 982 m, 21 October 2011, M. J. Silva et al. 3816 ( UFG), 3817 ( UFG) ; próximo ao alojamento do Parque , 14°09’30.6”S, 47°47’38.3”W, 1086 m, 16 October 2010, M. J. Silva et al. 3070 ( UFG) ; ibd., 21 October 2011, M. J. Silva et al. 3841 ( UFG) ; ibd., 20 January 2012, R. C. Sodré et al. 19 ( UFG) ; ibd., 10 March 2012, R. C. Sodré et al. 71 ( UFG), 74 ( UFG), 75 ( UFG), 77 ( UFG), 83 ( UFG), 84 ( UFG), 85 ( UFG), 88 ( UFG), 89 ( UFG) ; Vale da Lua , 14°10’47.9”S, 47°47’38.1”W, 969 m, 15 October 2010, M. J. Silva et al. 3060 ( UFG) ; estrada para ao vale da lua, 14°10’17.6”S, 47°46’53”W, 1013 m, 20 January 2012, R. C. Sodré et al. 08 ( UFG), 09 ( UFG) ; ibd., 14°10’20.4”S, 47°47’43.7”W, 1023 m, 22 October 2011, M. J. Silva et al. 3865 ( UFG), 3879 ( UFG) ; a esquerda da estrada para o Vale da Lua , 05 September 2014, R. C. Sodré et al. 1415 ( UFG), 1416 ( UFG), 1418 ( UFG), 1419 ( UFG), 1421 ( UFG) ; estrada para a sede do PNCV, 15 October 2010, M. J. Silva et al. 3031 ( UFG), 3033 ( UFG) ; imediações do Morro do Buracão , 14°07’46.9”S, 47°41’12.8”W, 20 January 2012, M. J. Silva et al. 4037 ( UFG), 4038 ( UFG), 4040 ( UFG) ; ibd., 08 March 2012, R. C. Sodré et al. 42 ( UFG) ; ibd., 26 May 2012, R. C. Sodré et al. 123 ( UFG) ; entrada a esquerda após o alojamento do Parque em direção à vila São Jorge , 14°10’10.5”S, 47°47’29.4”W, 20 January 2012, R. C. Sodré et al. 04 ( UFG) ; próximo a GO 118, entre Alto Paraíso de Goiás e a Serra do Pouso Alto , 14°3’43”S, 47°30’14”W, 25 May 2012, R. C. Sodré et al. 97 ( UFG), 98 ( UFG), 99 ( UFG), 100 ( UFG) ; cerca de 11 km da cidade em direção a Teresina de Goiás, após o morro do Japonês , 14°2’32”S, 47°31’40.3”W, 1540 m, 24 November 2012, M. J. Silva et al. 4568 ( UFG) ; GO 118 , a aproximadamente 10 Km de Alto Paraíso em direção a Teresina de Goiás, a esquerda do asfalto, 26 May 2012 R. C. Sodré 113 ( UFG), 114 ( UFG), 115 ( UFG) ; imediações do Morro da Baleia , 14°7.092’S, 47°38.776’W, 1330 m, 28 September 2012, R. C. Sodré et al. 158 ( UFG) ; proximidades da GO 239, acima da Mata do Buracão entre esta última e a Vila São Jorge , 14°8.259’S, 47°43.895’W, 1221 m, 28 September 2012, R. C. Sodré et al. 174 ( UFG), 175 ( UFG) ; ca. 500 metros do alojamento, 29 September 2012, R. C. Sodré et al. 176 ( UFG) ; ca. 2 Km do alojamento, 29 September 2012, R. C. Sodré et al. 179 ( UFG) ; arredores da vila de São Jorge, em direção ao Cânion 1, 14°9.943’S, 47°48.625’W, 1082 m, 29 September 2012, R. C. Sodré et al. 186 ( UFG), 187 ( UFG), 188 ( UFG) ; ao norte do morro da Baleia, na estrada para o rio Preto , 14°3’56.6”S, 47°38’21.4”W, 1193 m, 05 September 2014, R. C. Sodré et al. 1386 ( UFG) ; imediações da Reserva Ecológica dos Cristais , 14°5’56.4”S, 47°29’51.6”W, 1243 m, 06 September 2014, R. C. Sodré et al. 1425 ( UFG) ; a direita do km 131 da GO 118, sentido São João d’Aliança a Alto Paraíso de Goiás, 14°25’17.3”S, 47°30’31.5”W, 1000 m, 12 December 2014, R. C. Sodré et al. 1525 ( UFG) ; a esquerda do km 131 da GO 118, sentido São João d’Aliança a Alto Paraíso de Goiás, 14°25’20.7”S, 47°30’35”W, 1000 m, 20 January 2014, R. C. Sodré, T. M. S. Melo & M. J. Silva 1216 ( UFG) ; a aproximadamente 2 km de Alto Paraíso de Goiás em direção a São João d’Aliança , 21 June 2014, R. C. Sodré et al. 1305 ( UFG) ; ca. 1 km após a casa dos pesquisadores do PNCV, 14°8’31.9”S, 47°45’55.7”W, 1121 m, 04 January 2015, R. C. Sodré, M. J. Silva & C. H. G. Machado-Filho 1628 ( UFG) ; cerca 30 km a esquerda da GO 118 de São João d’Aliança a Alto Paraíso de Goiás, na estrada para catarata do rio dos Couros, 14°17’23.6”S, 47°42’33.2”W, 1049 m, 31 October 2014, R. C. Sodré et al. 1494 ( UFG), 1496 ( UFG), 1497 ( UFG). São João d’Aliança, 10 km de São João d’Aliança / Alto Paraíso, margem da rodovia GO – 118, 25 November 1994, M. Aparecida da Silva , F. C. A. Oliveira & T. S. Filgueiras 2434 ( UFG). Without municipalities, ca. 12 km N.W. of Veadeiros , road to Cavalcante, 21 October 1965, H. S. Irwin et al. 9440 ( UB) .

Croton desertorum is endemic to Brazil and was described from Minas Gerais by Müller Argoviensis (1873). In Goiás, it is frequently found in campo sujo and cerrado sensu stricto the municipalities of Alto Paraíso de Goiás, São João d’Aliança and Água Fria de Goiás ( Fig. 9A ), flowering and fruiting throughout the year, except in July and August. It can be confused with C. gracilescens , as both have short petioles (up to 3 mm long) with two patelliform sessile glands at apex, flowers of both sexes are subsessile (pedicels 0.5–2 mm long), pistillate sepals slightly unequal, ovate, and externally pubescent, as well as glabrous, 2-fid styles. However, C. desertorum has striate, rusty young branches (vs. smooth and green-white in C. gracilescens ), elliptic leaves with rare marginal glands (vs. linear, the margin always with glands) and thyrses with congested flowers (vs. lax flowers) and hirsute ovaries (vs. tomentose). Croton desertorum belongs to C. section Geiseleria .

2.11. Croton didrichsenii Webster (1992: 271) . — Julocroton humilis Didrichsen (1857: 132) . Type:— BRAZIL. São Paulo: campis Mugi, November 1833, P. W. Lund s.n. (holotype C 10011298!, isotypes A 00277247!, C 10011297!, G 00312779!, K). ( Figs. 8 , I–L; 10 , A–C)

Subshrubs 0.1–1 m tall, erect, stems yellow to rusty with stipitate stellate-porrect trichomes; latex clear. Stipules 4–15 × 0.4–0.8 mm, linear to linear-lanceolate, without glands, persistent; petioles 0.4–2.5 cm long, without glands. Leaves alternate or opposite near to dichotomy of the branches, blades 4–15.7 × 1.1–7.8 cm, ovate, oval-lanceolate, linear-lanceolate, base obtuse or lightly decurrent, apex acute, margin entire or serrate without glands; upper and lower surfaces with stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, with 4–7 pairs of secondary veins. Thyrses 1.5–4.5 cm long, terminal, staminate cymules with 1 flower; inflorescence axis, both faces of pistillate sepals and bracts of both flowers, dorsal surface of staminate sepals, ovary and styles with stipitate stellate-porrect trichomes; bracts of both flowers 7–14 × 0.4–0.9 mm, linear, with stipitate-capitate glands. Staminate flowers 5.5–7 mm long, pedicels 1.5–2.2 mm long; sepals 5, 1.8–2.3 × 1–2 mm, ovate to triangular, apex acute or obtuse; petals 5, 3–3.6 × 0.3–0.5 mm, linear-oblanceolate, apex acute, dorsal surface pubescent, ventral surface villous; stamens 11, villous. Pistillate flowers 8–9.5 mm long, pedicels 0.6–1.2 mm long; sepals 3, 5 or 7, 3(4) developed, 6.5–8 × 1–2.5 mm, subequal, linear-lanceolate to oval-lanceolate, apex acuminate, margins laciniate without glands, and 2 or 4 small sepals, 2–4 × 0.6–1.5 mm, lanceolate, margins entire or laciniate; petals absent; disk 3-segmented; ovary 1.9–2.7 × 1.6–1.8 mm, subglobose, styles 4-fid. Capsules 4.5–5 × 4–4.5 mm, subglobose, yellow-ferrugineous; seeds 4–4.2 × 2.7–3.2 mm, oblongoid or oval-ellipsoid, grayish.

Specimens examined:— BRAZIL. Goiás: Água Fria de Goiás, GO 118, km 83, entre São Gabriel e São João d’Aliança , 14°47’48.8”S, 47°33’26.3”W, 1073 m, 05 September 2013, R. C. Sodré et al. 842 ( UFG). Alto Paraíso de Goiás, ca. 20 km W. of Veadeiros , 10 February 1966, H. S. Irwin et al. 12467 ( UB) ; 19 km N. of Alto Paraíso de Goiás, 20 March 1971, H. S. Irwin et al. 32795 ( UB) ; ca. 18 km N. of Alto Paraíso de Goiás, 21 March 1971, H. S. Irwin et al. 32881 ( UB) ; 7 km by Road N of Alto Paraíso, 4 March 1973, W. R. Anderson et al. 6270 ( UB) ; área do aeroporto, 14°8.95’S, 47°46.78’W, 1155 m, 27 September 1995, M. L. Fonseca et al. 607 ( IBGE) ; estrada para o Vale da Lua , 14°10’28.5”S, 47°47’6.8”W, 1024 m, 15 October 2010, M. J. Silva 3061 ( UFG) ; GO 118 , cerca de 17 km de Alto Paraíso de Goiás, sentido Teresina de Goiás, 14°2’31.4”S, 47°31’42.7”W, 1520 m, 21 January 2012, M. J. Silva et al. 4068 ( UFG), 4070 ( UFG) ; ibd., R. C. Sodré et al. 23 ( UFG) ; ibd., J. E. C. Júnior et al. 160 ( UFG) ; imediações do morro do Japonês, a esquerda da GO 118, cerca de 15 km após Alto Paraíso de Goiás, em direção a Teresina de Goiás, 09 March 2012, M. J. Silva et al. 4134 ( UFG), 4135 ( UFG), 4136 ( UFG) ; ibd., 14°2’14.8”S, 47°31’46.9”W, 1540 m, 06 April 2013, R. C. Sodré et al. 677 ( UFG) ; ibd., 14°2’36.5”S, 47°31’43.1”W, 1482 m, 11 May 2013, R. C. Sodré & M. J. Silva 735 ( UFG), 736 ( UFG), 740 ( UFG) ; ibd., 14°3’7.9”S, 47°32’41.3”W, 1489 m, 11 May 2013, R. C. Sodré & M. J. Silva 751 ( UFG), 752 ( UFG) ; GO 118 , próximo a placa indicatica da Serra do Pouso Alto , 14°2’44.8”S, 47°31’25.7”W, 1083 m, 09 March 2012, R. C. Sodré et al. 51 ( UFG), 52 ( UFG), 53 ( UFG) ; PNCV, ca. 5 km ao norte do Morro do Japonês, ca. 200 m a esquerda da GO 118, sentido Alto Paraíso de Goiás a Teresina de Goiás, 14°1’30.9”S, 47°31’56.9”W, 26 October 2012, R. C. Sodré & M. J. Silva 242 ( UFG), 243 ( UFG) ; PNCV, região da Serra de Pouso Alto, ca. 1 km a esquerda da GO 118, sentido Alto Paraíso a Teresina de Goiás, 14°1’27”S, 47°32’13.2”W, 1150 m, 26 October 2012, R. C. Sodré & M. J. Silva 219 ( UFG), 220 ( UFG), 231 ( UFG), 232 ( UFG) cerca de 11 km da cidade em direção a Teresina de Goiás, após o morro do Japonês , 14°2’32”S, 47°31’40.3”W, 1540 m, 24 November 2012, M. J. Silva et al. 4566 ( UFG), 4567 ( UFG), 4569 ( UFG) ; a direita da GO 239, de Alto Paraíso de Goiás ao povoado de São Jorge, próximo ao morro da Baleia , 14°9’29.64”S, 47°37’54.9”W, 1168 m, 13 December 2012, R. C. Sodré et al. 376 ( UFG), 377 ( UFG), 379 ( UFG) ; imediações do morro do Japonês , 14 December 2012, R. C. Sodré et al. 395 ( UFG), 396 ( UFG), 397 ( UFG), 398 ( UFG) ; acima dos alojamentos do Parque , 08 February 2013, R. C. Sodré et al. 463 ( UFG) ; PNCV, cerca de 2 km a NE do morro do Japonês , 14°2’33.6”S, 47°32’28.4”W, 1455 m, 11 May 2013, M. J. Silva et al. 4920 ( UFG) ; GO 118 , km 155, no lado esquerdo da rodagem, 14°12’49.4”S, 47°29’15.9”W, 1122 m, 10 October 2013, M. J. Silva et al. 5409 ( UFG) ; cerca de 10 km ao norte de Alto Paraíso de Goiás, entre a rodovia (GO 118) e o morro do Japonês , 14°2’44.7”S, 47°31’24.6”W, 1468 m, 02 November 2013, R. C. Sodré et al. 964 ( UFG), 966 ( UFG), 970 ( UFG) ; a esquerda do km 131 da GO 118, sentido São João d’Aliança a Alto Paraíso de Goiás, próximo a divisa destes dois municípios, 14°25’17.9”S, 47°30’28.6”W, 1004 m, 30 October 2014, R. C. Sodré et al. 1471 ( UFG), 1472 ( UFG) ; a direita do km 131 da GO 118, sentido São João d’Aliança a Alto Paraíso de Goiás, 14°25’18.3”S, 47°30’27.4”W, 1005 m, 20 February 2014, M. J. Silva et al. 5845 ( UFG) ; imediações da fazenda Campo Seco, cerca de 15 km após a vila São Jorge, em direção ao Município de Colinas do Sul pela GO 239, 14°13’37.5”S, 47°53’39.3”W, 705 m, 14 December 2014, R. C. Sodré et al. 1560 ( UFG) ; cerca de 30 km a esquerda da GO 118 de São João d’Aliança para Alto Paraíso de Goiás, na estrada que leva a catarata do rio dos Couros , 14°17’35”S, 47°42’32”W, 1028 m, 02 January 2015, R. C. Sodré et al. 1589 ( UFG) ; 26 km de Alto Paraíso de Goiás para São João d’Aliança, lado esquerdo da rodovia GO – 118, km 141, fazenda Mata da Toca , 23 November 1994, M. Aparecida da Silva & F. C. A. Oliveira 2379 ( UFG) ; without locality, 18 September 1977, A. Allem & G. Vieira 1060 ( UFG), 1061 ( CEN). Campos Belos, estrada velha de São Domingos para Campos Belos , entrada da Fazenda Barrigudinha, 13°2’37”S, 46°47’32”W, 660 m, 22 October 2001, M. L. Fonseca et al. 3010 ( IBGE). Cavalcante, fazenda Horta, 13°49’00”S, 47°48’7”W, 22 October 1997, C. D. Costa et al. 59 ( IBGE) ; estrada balsa “Porto dos Paulistas” (rio Tocantins) para Buracão a cerca de 4, 9 km do rio, 13°28’6”S, 48°7’8”W, 410 m, 07 November 2000, G. Pereira-Silva et al. 4374 ( CEN) ; reserva da fazenda Gavião , 14°8’25.8”S, 47°52’21.8”W, 574 m, 12 July 2013, M. J. Silva et al. 5088-A ( UFG), 5089-A ( UFG) ; ibd., 14°8’37”S, 47°52’45.5”W, 01 November 2013, R. C. Sodré et al. 942 ( UFG), 943 ( UFG), 944 ( UFG), 945 ( UFG) ; sudoeste do PNCV acessado a partir da fazenda Gavião , 14°13’18”S, 47°53’29.9”W, 577 m, 01 November 2013, M. J. Silva et al. 5518 ( UFG) ; imediações da Serra do Ministro, cerca de 500 metros da estrada entre Cavalcante e Colinas do Sul , 13°54’34”S, 47°38’55.4”W, 29 January 2014, R. C. Sodré et al. 1192 ( UFG) ; ibd., 13°54’38.4”S, 47°38’56.6”W, 537 m, 29 January 2014, M. J. Silva et al. 5750 ( UFG), 5751 ( UFG) ; estrada que leva a Colinas do Sul , ca. 18 km de Cavalcante, 29 January 2014, R. C. Sodré et al. 1185 ( UFG), 1186 ( UFG) ; em direção ao Engenho II, região dos Kalungas, cerca de 18 km após Cavalcante, 13°39’29.4”S, 47°28’31.4”W, 1108 m, 25 July 2014, R. C. Sodré et al. 1335 ( UFG), 1336 ( UFG). Colinas do Sul , estrada de acesso a fazenda Gavião, ca. 3 km da GO 239, 04 August 2013, R. C. Sodré et al. 833 ( UFG) ; ibd., 01 November 2013, R. C. Sodré et al. 927 ( UFG) ; ibd., 13 December 2013, R. C. Sodré et al. 1055 ( UFG), 1056 ( UFG). Niquelândia, estrada de terra 07 km da cidade em direção a Colinas, 14°27’3”S, 48°19’37”W, 19 September 1996, M. Aparecida da Silva et al. 3132 ( IBGE) ; estrada para o DNPM, km 5 de Niquelândia , 14°29’46”S, 48°27’0”W, 645 m, 19 October 1996, A. F. Vaz et al. 1113 ( IBGE) ; ca. 5, 5 km a direita da mina de níquel, 14°24’2”S, 48°25’20”W, 1045 m, 28 November 1996, M. L. Fonseca et al. 1379 ( IBGE) ; proximidades do condomínio “ Vista do Lago ”, 14°11’40.6”S, 48°18’43.4”W, 468 m, 20 June 2014, R. C. Sodré et al. 1284 ( UFG) ; região da Pedra Encantada, cerca de 8 km de Colinas do Sul, na estrada que leva a Niquelândia, 14°11’31.8”S, 48°7’22.5”W, 536 m, 21 June 2014, R. C. Sodré et al. 1302 ( UFG). Nova Roma, de Teresina de Goiás para Monte Alegre de Goiás, na GO 576 para Nova Roma, 13°28’20”S, 47°7’52.2”W, 428 m, 01 November 2014, R. C. Sodré et al. 1503 ( UFG), 1504 ( UFG), 1506 ( UFG). São João d’Aliança, a direita da GO 118, 300 metros do Atos Hotel, 14°41’35.8”S, 47°31’17.3”W, 1047 m, 28 January 2014, M. J. Silva et al. 5717 ( UFG), 5718 ( UFG), 5719 ( UFG), 5720 ( UFG), 5721 ( UFG) .

This is a South American species distributed in Argentina, Bolivia, Brazil (BA, DF, GO, MG, MS, MT, PR, SC, SP, RS), Paraguay and Uruguay in savanna vegetation ( Caruzo & Cordeiro 2007). We found it in grassland, cerrado rupestre, campo sujo in all municipalities of the Chapada dos Veadeiros except Teresina de Goiás ( Fig. 9A ), flowering and fruiting throughout the year. Among the species studied, C. didrichsenii is unique among the studied species in having two vestigial pistillate sepals and three conspicuously laciniate ones. This characteristic together with the yellow-rustiness of the branches with long stipitate stellate trichomes and deltoid inflorescences with long bracts (7–14 mm long), makes it easily recognizable. Croton didrichsenii belongs to C. section Julocroton .

2.12. Croton glandulosus Linnaeus (1759: 1275) . Lectotype (designated by Fawcett & Rendle 1920): — JAMAICA. Without locality, no date, P. Browne 1 (LINN HL1140-7!, isolectotypes S 09-18941!, S 09-18929!). ( Fig. 10 , D − H)

Subshrubs 0.2–1 m tall, erect, stems canescent-green with sessile stellate-porrect trichomes; latex clear. Stipules 1.7–4 × 0.3–0.4 mm, linear, without glands, persistent; petioles 0.7–2.4 cm long, with 2 shortly stipitate patelliform glands. Leaves alternate or opposite near the dichotomies of the branches, blades 1.6–4.5 × 0.9–3 cm, ovate or oval-lanceolate, base obtuse, apex acute, margin serrate without glands or rarely with patelliform glands; upper and lower surfaces with sessile, stellate-porrect trichomes, membranaceous, venation brochidodromous, with 4–6 pairs of secondary veins. Thyrses 0.8–2.7 cm long, terminal; inflorescence axis, dorsal surface of bracts and sepals of all flowers, ovary and styles with stellate-porrect trichomes; bracts of flowers 1.2–2 × 0.2–0.5 mm, linear or linear-lanceolate, with sessile pyriform glands. Staminate flowers 3.2–3.8 mm long, pedicels 1.2–1.6 mm long; sepals 5, 1.6–2.2 × 0.7–1 mm, ovate, apex acute; petals 5, 1.8–2.4 × 0.6–0.9 mm, elliptic or often obovate, apex obtuse, ventral surface villous, dorsal surface glabrous; stamens 9–11, filaments glabrous. Pistillate flowers 3–4 mm long, pedicels 0.2–0.5 mm long; sepals 5, 1.8–2.5 × 0.5–1 mm, subequal, obovate, spatulate, oblanceolate or oblong, apex acute or obtuse, margins entire without glands; petals 5, shaped gland or laminar, 0,8–1,5 mm long, linear or triangular, villous; disk 5-segmented, ovary 1.5–2 × 1.6–2.2 mm, subglobose, styles 2-fid. Capsules 5–5.4 × 4.8–5.2 mm, subglobose, greenish; seeds 3.8– 4.2 × 2.2–2.4 mm, oblong-ellipsoid, brownish, with spots.

Specimens examined:— BRAZIL. Goiás. Água Fria de Goiás, em frente à chácara Cantinho dos Batistas, 14°54’4.8”S, 47°35’3.7”W, 1132 m, 08 March 2012, R. C. Sodré et al. 40 ( UFG), 41 ( UFG). Alto Paraíso de Goiás, estrada para o vale da lua, 14°10’20.4”S, 47°47’43.7”W, 1023 m, 22 December 2011, M. J. Silva et al. 3886 ( UFG) ; ibd., 10 March 2012, R. C. Sodré et al. 62 ( UFG) ; sítio Pedrão , 14°11’0.8”S, 47°51’8.6”W, 950 m, 21 December 2011, M. J. Silva et al. 3827 ( UFG) ; ibd., 10 March 2012, R. C. Sodré et al. 72 ( UFG) ; porteira que dá acesso a sede, 14°10’10.5”S, 47°47’29.4”W, 1013 m, 19 January 2012, R. C. Sodré et al. 02 ( UFG). Cavalcante, a esquerda da GO 241, de Teresina de Goiás para a Cavalcante, na entrada da fazenda Siriema, 13°47’10.5”S, 47°23’51”W, 802 m, 29 January 2014, R. C. Sodré et al. 1170 ( UFG), imediações da estrada que leva a Colinas do Sul , ca. 18 km de Cavalcante em campo sujo, 29 January 2014, R. C. Sodré et al. 1182 ( UFG) .

Croton glandulosus is a widespread Neotropical weedy species that has also been introduced in Africa and Australia (van Ee et al. 2011). We found it in anthropic environments, flowering and fruiting from December to March. It resembles C. hirtus L’Héritier (1785: 17) by having petioles with stipitate patelliform glands, inflorescences with flowers evenly distributed along the axis, bracts with pyriform glands, and usually obovate to spatulate pistillate sepals. In C. glandulosus the upper surface of the leaves has stellate-porrect trichomes, the glands of the bracts are sessile, and the pistillate sepals are equal or discretely unequal in size. Croton hirtus has the upper surface of the leaves hispid, glands of the bracts long stipitate and pistillate sepals conspicuously unequal. Croton glandulosus belongs to C. section Geiseleria .

2.13. Croton goyazensis Müller Argoviensis (1865: 120) . Lectotype (designated here):— BRAZIL. Goiás: in Campis Petrosis Catalão, August 1834, L. Riedel 2473 (LE! [the sheet annotated with Müller’s handwriting], isolectotypes B [destroyed, photo at F F0BN005103], LE! [3 more sheets, none with annotations by Müller], K 000186100!, G 00434536!, GH 00047305!. Syntypes:— BRAZIL. Goiás: in Campis Catalão , September 1834, P. W. Lund s.n (G 00312494!, F V0093325F!) ; without locality, no date, J. B. E. Pohl 1618 (BR 8762290!, K 000913002!, K 000913003!). ( Figs. 8 , M–O; 10 , I–M) Croton luzianus Müller Argoviensis (1873: 218) , syn. nov. Lectotype (designated here):— BRAZIL. Goiás: ad Ponto Alto prope S. Luzia ,

no date, J. B. E. Pohl 1618 (Catalog number: 1034) (W 0051288! [annotated by Müller as C. luzianus ], isolectotype: W 0051287!, F

V0056157!, G 00434543!). Syntype:— BRAZIL. Goiás: ad Ponto Alto, no date, J. B. E. Pohl 1037 (W 0051286!).

Shrubs 0.35–0.7 m tall, erect, stems canescent-yellow with stipitate stellate multiradiate trichomes; latex clear. Stipules 0.4–0.9 × 0.4–0.6 mm, triangular, without glands, persistent; petioles 0.5–1.4 cm long, with 2 subcylindrical glands. Leaves alternate, blades 3.2–8.1 × 2.4–5.8 cm, oval, base obtuse or slightly cordate, apex acute, margin serrate, crenate or rarely subentire with or without stipitate-capitate glands; both surfaces with shortly stipitate stellate-porrect trichomes, membranaceous, venation semi-craspedodromous, with 4–6 pairs of secondary veins. Thyrses 1.2–14 cm long, terminal, staminate cymules with 1–4 flowers; inflorescence axis, dorsal surface of bracts and sepals of both flowers, ovary and style with stellate-porrect trichomes; staminate bracts 0.2–1 × 0.1–0.2 mm, linear, without glands; pistillate bracts 1.1–2 × 0.3–0.5 mm, linear-lanceolate, without glands. Staminate flowers 6.5–9 mm long, pedicels 2.5–5 mm long; sepals 5, 2–2.3 × 1.1–1.6 mm, ovate, apex acute or obtuse; petals 5, 2.3–3 × 0.7–1.1 mm, oblanceolate, obovate or elliptic, apex acute or obtuse, dorsal surface glabrous, ventral surface villous at the base; stamens 11, filaments villous. Pistillate flowers 8–11 mm long, pedicels 3.5–6.7 mm long; sepals 5, 2.3–3.5 × 0.7–1.2 mm, subequal, oblong-oblanceolate, oblong-ovate or oblong-oblanceolate, apex acute, margins entire without glands; petals 5, 0.5–1.7 × 0.05–0.1, linear or linear-lanceolate, villous, disk 5-segmented; ovary 1.8–2.3 × 1.7–2 mm, oblongoid, styles 2-fid. Capsules 5.5–6 × 5–5.5 mm, oblongoid, grayish; seeds 3.9–4 × 2.6–2.8 mm, ellipsoid, uniformly brownish.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, a oeste do Morro da Baleia , 09 February 2013, R. C. Sodré et al. 476 ( UFG), 477 ( UFG), 478 ( UFG), 479 ( UFG) ; imediações da cidade, próximo a estação da Saneago 14°07’50”S, 47°32’21.1”W, 1343 m, 06 September 2013, R. C. Sodré et al. 853 ( UFG), 854 ( UFG), 855 ( UFG), 856 ( UFG), 857 ( UFG), 858 ( UFG), 859 ( UFG), 860 ( UFG), 862 ( UFG), ibd., M. J. Silva et al. 5358 ( UFG), 5365 ( UFG), 5371 ( UFG) ; 100 metros a direita da GO 118 na entrada para a Reserva Ecológica dos Cristais , 14°05’12.8”S, 47°31’5.5”W, 1377 m, 06 September 2014, R. C. Sodré et al. 1422 ( UFG), 1423 ( UFG) ; cerca 1 km a esquerda da GO 239, sentido Alto Paraíso de Goiás ao povoado de São Jorge, em estrada oposta a entrada para o Bona Espero , 14°10’2.4”S, 47°37’55.8”W, 06 September 2014, R. C. Sodré et al. 1439 ( UFG) ; cerca 30 km a esquerda da GO 118 na estrada para o rio dos Couros , 14°17’23.6”S, 47°42’33.2”W, 1049 m, 31 October 2014, R. C. Sodré et al. 1493 ( UFG) .

Croton goyazensis occurs in areas of savanna and seasonally dry forests in Bolivia, Brazil (AM, BA, DF, GO, MG, PA, PI), and Paraguay ( Alves 2012). In Chapada dos Veadeiros it was collected in Alto Paraíso de Goiás growing in cerrado sensu stricto and campos sujos close to PNCV ( Fig. 9B ), flowering in September, October and February and fruiting in September.

Croton goyazensis is easily recognized by the cinereous-whitish branches and leaves, cylindrical acropetiolar glands, leaf margin usually without glands or else rarely with stipitate-capitate glands, semi-craspedodromous leaf venation, and long-pedicellate, oblongoid capsules. Croton goyazensis belongs to C. section Geiseleria .

Taxonomic notes:— Müller Argoviensis (1873) related C. luzianus to C. goyazensis , stating that the former differed from the latter because the lack of glands on the petiole (vs. presence of glands in C. goyazensis ), clavate staminate pedicel (vs. not clavate), obovate-oblanceolate pistillate sepals (vs. linear-lanceolate), stamens with villous filaments (vs. glabrous) and styles shortly 2-fid (vs. deeply bipartite). However, field observations during this study led us to conclude that the characters cited by Müller to differentiate C. luzianus from C. goyazensis are variable in the same population, which is why we choose to treat C. luzianus as a synonym of C. goyazensis .

2.14. Croton gracilescens Müller Argoviensis (1865: 110) . Type:— BRAZIL. Goiás: prope Cavalcante , no date, J. B. E. Pohl 1619 (holotype B 10-0242753!, isotypes A 00257930!, BR 8762627!, G 00434535!, K 000254366!). ( Figs. 11 , A–D; 12 , A–D)

Subshrubs 0.2–0.8 (1.3) m tall, erect, young stems greenish with sessile stellate-porrect trichomes; latex clear. Stipules 1.5–5 × 0.4–1 mm, oval to lanceolate, without glands, persistent; petioles 1–4 mm long, with 2 sessile patelliform glands. Leaves alternate, blades 8.8–13.8 × 0.7–4.3 cm, linear to narrowly elliptic, base obtuse, apex acute, margin serrate with patelliform glands; upper and lower surfaces with sessile stellate-porrect trichomes, membranaceous, venation brochidodromous, with 11–22 pairs of secondary veins. Thyrses 1.7–9.5(–20.2) cm long, terminal, staminate cymules with 1 or 3 flowers; inflorescence axis, dorsal surface of bracts and sepals of both flowers and ovary with stellate trichomes; staminate bracts 1.0–1.2 × 0.4–0.6 mm, ovate; pistillate bracts 1.2–1.9 × 0.6–1 mm, ovate to elliptic. Staminate flowers 4–5 mm long, pedicels 1–2(3.5) mm long; sepals 5, 1.2–2 × 0.7–1.1 mm, ovate, apex acute; petals 5, 2–2.3 × 0.4–0.7 mm, elliptic or sometimes oblong or obovate, apex obtuse, dorsal surface glabrous, ventral surface villous; stamens 11, filaments glabrous. Pistillate flowers 3.5–5.3 mm long, pedicel 0.6–1.2 mm long; sepals 5, 1.3–2.8 × 1.2–1.7 mm, equal or subequal, ovate, apex acute or obtuse, margins entire without glands; petals 5, 0.2–1 mm long, linear or glanduliform, globose, disk 5-segmented; ovary 1.1–1.6 × 1.5–2 mm, globose, shortly tomentose, styles 2-fid, glabrous. Capsules 3.8–4 × 4.6–4.8 mm long, globose, greenish; seeds 3.6–3.7 × 2.4–2.7 mm, oblong-elliptic, grayish.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, estrada para o alojamento do Parque , 15 October 2010, M. J. Silva 3049 ( UFG) ; estrada para o Vale da Lua , 14°10’25.4”S, 47°47’5.1”W, 1040 m, 15 October 2010, M. J. Silva 3057 ( UFG) ; ibd., 22 October 2011, M. J. Silva et al. 3856 ( UFG), 3878 ( UFG) ; ibd., 31 October 2014, R. C. Sodré et al. 1477 ( UFG), 1478 ( UFG) ; imediações do alojamento do PNCV, 16 October 2010, M. J. Silva 3080 ( UFG), 3096 ( UFG) ; na estrada para o Raizama , 14°11’37.7”S, 47°49’36.2”W, 902 m, 16 October 2010, M. J. Silva 3089 ( UFG) ; saindo do povoado São Jorge em direção a Colinas do Sul, ca. 2 km após a entrada para o Raizama , 13 December 2013, R. C. Sodré et al. 1043 ( UFG) ; imediações da fazenda Campo Seco, cerca de 15 km após a vila São Jorge pela GO 239, 14°13’39.9”S, 47°53’41.2”W, 700 m, 22 February 2014, R. C. Sodré, T. M. S. Melo & M. J. Silva 1242 ( UFG) ; a esquerda da estrada para o Vale da Lua , cerca de 100 metros da GO 239, 05 September 2014, R. C. Sodré et al. 1413 ( UFG), 1414 ( UFG), 1417 ( UFG), 1420 ( UFG) ; parcelas 7 e 8, próximo à sede, 14°9’65”S, 47°47’50”W, 1477 m, 26 September 1995, M. L. Fonseca & M. P. Neto 570 ( IBGE). Cavalcante , estrada balsa dos Paulistas (rio Tocantins) 1, 13°27’49”S, 48°07’17”W, 370 m, 16 October 2001, G. Pereira-Silva et al. 5606 ( CEN) ; RPPN Serra do Tombador , 13°38’04”S, 47°49’06”W, 857 m, 29 October 2011, A. R. O. Ribeiro et al. 262 ( UB) ; próximo a Serra do Ministro, a 500 metros da estrada entre Cavalcante e Colinas do Sul , 13°54’34”S, 47°38’55.4”W, 29 January 2014, R. C. Sodré 1188 ( UFG), 1189 ( UFG), 1190 ( UFG), 1191 ( UFG) ; próximo da estrada que leva a Colinas do Sul, ca. 18 km de Cavalcante, 29 January 2014, R. C. Sodré et al. 1179 ( UFG), 1183 ( UFG). Colinas do Sul , estrada que dá acesso à fazenda Gavião , ca. 3 km a direita da GO 239, de Alto Paraíso à Colinas do Sul , 13 December 2013, R. C. Sodré et al. 1058 ( UFG). Niquelândia , margem da GO 132, entre Colinas do Sul e Niquelândia , 14°26’34.4”S, 48°8’44.9”W, 498 m, 30 January 2014, R. C. Sodré 1202 ( UFG) ; a direita da fazenda Pai-Herói , 14°27’23.4”S, 48°13’21.8”W, 512 m, 20 June 2014, R. C. Sodré et al. 1286 ( UFG) ; próximo ao Recanto Sousa Mendes , 14°25’1.9”S, 48°6’5.9”W, 480 m, 21 June 2014, R. C. Sodré 1297 ( UFG), 1298 ( UFG), 1299 ( UFG) .

This species is apparently endemic to Goiás State ( Sodre et al. 2014). It was found in PNCV growing in campo sujo and cerrado sensu stricto in the municipalities of Alto Paraíso de Goiás, Cavalcante and Colnas do Sul ( Fig. 9B ), flowering and fruiting from June to February. Morphologically, Croton gracilescens resembles C. antisyphiliticus and C. desertorum , as discussed in the comments under the C. desertorum . It is distinguished by the linear to narrowly elliptical leaves, laxly flowered inflorescences and bifid styles. This species also has apparently glabrous stems and flowers, whereas in C. desertorum the stems and flowers are conspicuously pubescent. Croton gracilescens belongs to C. section Geiseleria .

2.15. Croton gracilipes Baillon (1864: 333) . Lectotype (first step designated by Caruzo & Cordeiro 2007, second step designated here):— BRAZIL. Minas Gerais: without locality, 1838, M. P. Claussen 773 (P 00493373!), isolectotypes G 00311426!, P 00493372!). Syntype:— BRAZIL. Mato Grosso: without locality, 1831–1833, C. Gaudichaud-Beaupré 250 (P 00493374!). ( Figs. 11 , E–H; 12 , E–G)

Shrubs or trees 2.5–5 m tall, erect, stems greenish with stellate-porrect trichomes; latex turbid. Stipules 2.5–5 × 0.2–0.4 mm, linear, with 2 globose glands basally, persistent; petiole 4–19 cm long, with 2 patelliform glands. Leaves alternate or whorled near to dichotomy of the branches, blades 11–28 × 3–13.2 cm, ovate to lanceolate, base obtuse or sometimes lightly cordate, apex acuminate, margin subentire without glands; upper and lower surfaces with sessile stellate-porrect trichomes respectively, membranaceous, venation brochidodromous, with 6–10 pairs of secondary veins. Thyrses 16– 78 cm long, terminal, bisexual cymules, at the base, and staminate cymules with 2–4 staminate flowers, inflorescence axis, dorsal surface of bracts and sepals of both flowers, ovary and styles with stellate-porrect trichomes; staminate bracts 0.8–1.5 × 0.5–1 mm, ovate; pistillate bracts 1–1.3 × 0.5–0.7 mm, oval-lanceolate. Staminate flowers 6–10 mm long, pedicels 3.5–7 mm long; sepals 5, 2–2.5 × 1.5–2 mm, ovate or elliptic, apex acute; petals 5, 2.2–2.6 × 0.5–0.8 mm, oblong or oblanceolate, apex obtuse or acute, ventral surface villous at the base; stamens 16–19, filaments glabrous. Pistillate flowers 3–6 mm long, pedicels 0.3–0.8 mm long; sepals 5, 1.6–2 × 0.9–1.5 mm, subequal, narrowly triangular or anceolate, apex acute, margins entire without glands; petals 5, globose (ca. 0.3 mm), or laminar 0.8–1.5 × 0.2–0.3 mm, linear or narrowly lanceolate, disk subentire; ovary 1.5–2.5 mm long, globose, styles 2-fid, pubescent. Capsules 5–6 mm long, globose, greenish, columella with three prominent, smooth lobes; seeds 4–4.5 × 2.5–3 mm, ellipsoid, grayish, spotted.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, trilha perpendicular ao espaço destinado para estacionamento do Vale da Lua , 14°10’20.7”S, 47°46’45.4”W, 929 m, 20 January 2012, M. J. Silva et al. 4049 ( UFG), 4050 ( UFG), 4051 ( UFG), 4052 ( UFG) ; ibd., 11 February 2013, R. C. Sodré et al. 516 ( UFG), 517 ( UFG) ; ibd., M. J. Silva et al. 4734 ( UFG), 4735 ( UFG), 4736 ( UFG). Teresina de Goiás, 7 Km by road S of Teresina, 1080 m, 17 March 1973, W.R. Anderson et al. 7316 ( UB) ; 4 Km by road S of Teresina, 1000 m, 18 March 1973, W.R. Anderson et al. 7372 ( UB). Without municipality, 40 Km north of Veadeiros , 1000 m, 15 March 1969, H.S. Irwin et al. 24435 ( UB) .

This species was reported by Caruzo & Cordeiro (2007) in mesophilic forests of Argentina, Bolivia, Brazil (GO, MG, MS, MT, PR, SP) and Paraguay. It was collected in the municipalities of Alto Paraíso de Goiás and Teresina de Goiás ( Fig. 9B ), in ombrophilous low montane forests, flowering and fruiting from January to March.

Croton gracilipes and C. urucurana Baillon (1864: 335) share the arboreal habit, the long petioles (4–19 cm long) with a pair of patelliform glands, the leaf blades ovate commonly cordate with subentire margin, and long thyrses (15– 78 cm long) with bisexual and staminate cymules. However, C. gracilipes has branches with stellate-porrect trichomes (vs. dendritic in C. urucurana ), linear stipules (vs. lanceolate), triangular or lanceolate pistillate sepals (vs. oblong to ovate), and pubescent styles (vs. glabrous). Croton gracilipes belongs to C. section Adenophylli .

2.16. Croton gracilirameus Silva et al. (2015: 162) . Type:— BRAZIL. Goiás: Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, 3 Km a noroeste do Morro do Buracão , 14°6’33.6”S, 47°43’32.7”W, 1181 m, 10 February 2013, M. J. Silva et al. 4716 (holotype UFG!, isotypes MICH!, NY!). ( Figs. 11 , I–L; 12 , H–J)

Subshrubs 0.5–0.9 m tall, erect, stems canescent-green with sessile stellate-porrect trichomes; latex clear. Stipules 1–2 × 0.2–0.3 mm, triangular, oval-lanceolate or linear, with stipitate-capitate glands, persistent; petioles 0.5–2 cm long, with 2–6 shortly stipitate patelliform glands. Leaves alternate, blades 2.4–4 × 1.4–3.7 cm, ovate or widely ovate, base lightly cordate, apex acute, margin serrate or crenate with stipitate-capitate glands; upper and lower surfaces with subsessile stellate-porrect trichomes, membranaceous, venation brochidodromous, with 4 or 5 pairs of secondary veins. Thyrses 2–8 cm long, terminal, staminate cymules with 1 or 2 flowers; inflorescence axis, both faces of sepals and bracts pistillate, dorsal surface of sepals and bracts staminate, ovary and styles with stellate-porrect trichomes; bracts of both flowers 1.3–2.2 × 0.3–0.5 mm, lanceolate or linear, with stipitate-capitate glands. Staminate flowers 5– 7.5 mm long, pedicels 2.2–5 mm long; sepals 5, 1.7–1.9 × 0.6–0.8 mm, ovate or oval-lanceolate, apex acute; petals 5, 1.7–2.1 × 0.7–0.9 mm, widely elliptic, apex obtuse, ventral surface villous; stamens 9–11, filaments glabrous. Pistillate flowers 11–13 mm long, pedicels 5–9 mm long; sepals 5, 3.2–6.7 × 0.6–1.1 mm, subequal, narrowly lanceolate, apex acute, margins entire, with stipitate-capitate glands; petals absent, disk 5-lobed; ovary 1.5–2 × 1–1.7 mm, oblongoid, styles 2-fid, pubescent. Capsules 5–5.5 × 4.3–4.8 mm, oblongoid, light green; seeds 4–4.2 × 2–2.2 mm, ellipsoid, brownish, with spots.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, próximo a torre de transmissão da cidade, por trás do “Hotel Novo Paraíso”, 21 November 1991, B. M. T. Walter et al. 592 ( IBGE) ; ibd., a aproximadamente 1 Km do alojamento, em direção ao Rio Preto , 14°08’S, 47°47’W, 27 October 2012, R. C. Sodré et al. 257 ( UFG) ; ibd., a aproximadamente 1,5 Km do alojamento, em direção ao Rio Preto , 14°08’30.6”S, 47°47’56.9”W, 1044 m, 27 October 2012, R. C. Sodré et al. 268 ( UFG) ; ibd., imediações do Rio Preto , 14°08’38.3”S, 47°47’26.5”W, 970 m, 27 October 2012, R. C. Sodré et al. 270 ( UFG) ; ibd., cerca de 1 Km do Rio Preto , sentido alojamento, 27 October 2012, R. C. Sodré et al. 275 ( UFG) ; ibd., Rio Preto, proximidades do Cânion 1, 14°08.645’S, 47°48.319’W, 949 m, 15 December 2012, R. C. Sodré, M. J. Silva & A. O. Souza 423 ( UFG), 427 ( UFG), 429 ( UFG), 431 ( UFG) ; ibd., M. J. Silva, R. C. Sodré & A. O. Souza 4673 ( UFG), 4674 ( UFG), 4675 ( UFG), 4676 ( UFG) ; ibd., 3 Km a noroeste do Morro do Buracão , 14°06’33.6”S, 47°43’32.7”W, 1181 m, 10 February 2013, R. C. Sodré et al. 493 ( UFG), 494 ( UFG), 495 ( UFG), 496 ( UFG) ; ibd., Rio Preto, região das Corredeiras , 14°09’17.4”S, 47°50’8.4”W, 1188 m, 04 March 2013, R. C. Sodré et al. 582 ( UFG), 583 ( UFG), 589 ( UFG), 590 ( UFG), 591 ( UFG) ; ibd., GO 118, a direita do Km 156, sentido São João d’Aliança a Alto Paraíso de Goiás, 04 April 2013, R. C. Sodré et al. 638 ( UFG), 641 ( UFG), 643 ( UFG), 644 ( UFG), 645 ( UFG) ; ibd., M. J. Silva et al. 4793 ( UFG) ; ibd., Serra do Pouso Alto, aproximadamente 3 km a direita da GO 118, sentido Alto Paraíso de Goiás a Teresina de Goiás, entrando no cruzeiro, 13°50’7.3”S, 47°15’22.5”W, 953 m, 06 April 2013, R. C. Sodré et al. 680 ( UFG), 681 ( UFG), 684 ( UFG), 686 ( UFG), 687 ( UFG) ; ibd., a 300 m da casa abandonada após o córrego, em direção a Serra do Pouso Alto , 13°57’42.4”S, 47°28’49.9”W, 1405 m, 06 April 2013, M. J. Silva et al. 4830 ( UFG), 4832 ( UFG) ; ibd., a nordeste do Morro do Buracão, 14°06’42.1”S, 47°43’56.4”W, 1162 m, 10 May 2013, R. C. Sodré et al. 732 ( UFG). Cavalcante, Parque Nacional da Chapada dos Veadeiros , base da Serra de Pouso Alto , 09 March 2012, R. C. Sodré 47 ( UFG), 48 ( UFG) .

This species is endemic to Goiás state and it has been recorded from Serra Dourada and the Chapada dos Veadeiros ( Silva et al. 2015a). We found it in the municipalities of Alto Paraíso de Goiás and Cavalcante ( Fig. 9C ), especially inside the PNCV in cerrado sensu stricto near rock outcrops, flowering and fruiting from October to March. It is morphologically most similar to C. catariae . Croton gracilirameus belongs to C. section Geiseleria .

2.17. Croton grandivelus Baillon (1864: 322) Lectotype (first step designated by Caruzo & Cordeiro 2007, second step designated here):— BRAZIL. São Paulo: without locality, 1816–1821, A. F. C. P. Saint-Hilaire cat. C 2 1358 (P 00623596!, isolectotypes A 00047307!, P 00623597!). ( Figs. 12 , K–O; 13 , A–C)

Subshrubs 0.25–1 m tall, erect, stems yellowish to ferrugineous with stipitate stellate-porrect trichomes; latex clear to yellowish, reddish after exposure to air. Stipules 2.5–8 × 0.3–1.1 mm, linear to lanceolate, sometimes lobed at the base, with papilliform glands basally or on the margin, persistent; petioles 0.2–1.1 cm long, without glands. Leaves alternate or opposite near the dichotomies of the branches, blades 6.7–14.1 × 2.3–7.4 cm, oboval, oblanceolate, elliptic or oblong, base acute, apex obtuse, mucronate or emarginate, margin entire without glands; upper and lower surfaces with stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, with (6–)8–11 pairs of secondary veins. Thyrses 3.5–17.3 cm long, terminal, staminate cymules with 1–6 flowers; inflorescence axis, both faces of sepals pistillate and bracts of both flowers, dorsal surface of staminate sepals, ovary and styles with stellate-porrect trichomes; staminate bracts 4.2–6 × 0.7–1.1 mm, narrowly lanceolate; pistillate bracts 6–13 × 0.5–1.7 mm, linear to lanceolate. Staminate flowers 6.5–9 mm long, pedicels 2.3–4.3 mm long; sepals 5, 2.5–3.2 × 1.7–2.3 mm, ovate, apex acute; petals 5, 3.2–4.3 × 0.7–1.3 mm, oblanceolate, oblong or elliptic, apex obtuse or acute, ventral surface villous basally; stamens 15–17, filaments villous basally. Pistillate flowers 6–8 mm long, pedicels 0.1–0.2 mm long; sepals 5, 5–10 × 0.3–1 mm, subequal, linear or narrowly lanceolate, apex acute, margins entire without glands; petals 5, 1–4 × 0.1–0.2 mm, linear, disk 5-lobed; ovary 3.5–5 mm long, globose, styles 2-fid, ascending, pubescent. Capsules 7–9 mm long, globose, yellow-green to yellow-brown, covered by stipitate stellate trichomes, columella with three prominent, smooth lobes; seeds 4.2–4.7 × 2.5–3 mm, widely ellipsoid, dark brown to uniformly grayish.

Specimens examined:— BRAZIL. Goiás: Água Fria de Goiás, GO 118, a esquerda do km 56, em direção a São João d’Aliança , 15°01’42.3”S, 47°36’1.5”W, 1204 m, 08 March 2012, R. C. Sodré et al. 35 ( UFG), 36 ( UFG), 39 ( UFG) ; ibd., 15°01’59.4”S, 47°36’1.7”W, 1176 m, 28 January 2014, M. J. Silva et al. 5714 ( UFG) ; a direita do km 61 da GO 118, sentido povoado de São Gabriel da Aliança ao município de São João d’Aliança, 14°59’14.5”S, 47°36’16.1”W, 1131 m, 12 December 2013, R. C. Sodré et al. 1040 ( UFG), 1041 ( UFG). Alto Paraíso de Goiás, ca. 15 km W of Veadeiros, 14°S, 47°W, 1000 m, 8 February 1966, H. S. Irwin et al. 12332 ( UB) ; ca. 16 km south of Alto Paraíso , 20 March 1969, H. S. Irwin et al. 24756 ( UB) ; 2 km by Road N of Alto Paraíso, 1400 m, 6 March 1973, W. R. Anderson et al. 6411 ( UB) ; estrada para Nova Roma, ca. 6 km leste de Alto Paraíso de Goiás, 14°07’S, 47°26’W, 1200 m, 13 March 1995, T. B. Cavalcanti et al. 1278 ( CEN) ; GO 118 , próximo à placa indicativa da Serra do Pouso , 14°02’44.8”S, 47°31’25.7”W, 1083 m, 09 March 2012, R. C. Sodré et al. 56 ( UFG), 58 ( UFG), 59 ( UFG) ; ca. 10 km ao norte de Alto Paraíso de Goiás pela GO 118, entre a rodovia e o morro do Japonês , 26 May 2012, R.C. Sodré et al. 112 ( UFG), 117 ( UFG), 118 ( UFG), 119 ( UFG) ; ibd., 14°2’44.7”S, 47°31’24.6”W, 1468 m, 02 November 2013, R. C. Sodré et al. 962 ( UFG), 967 ( UFG) ; cerca de 11 km da cidade em direção a Teresina de Goiás, 14°02’32”S, 47°31’40.3”W, 1540 m, 24 November 2012, M. J. Silva et al. 4580 ( UFG) ; estrada para o Vale da Lua , 24 November 2012, M. J. Silva et al. 4581 ( UFG) ; a direita da GO 239, sentido Alto Paraíso ao povoado de São Jorge, próximo ao Morro da Baleia , 14°09’29.64”S, 47°37’54.9”W, 1168 m, 13 December 2012, R. C. Sodré et al. 375 ( UFG), 378 ( UFG) ; próximo ao morro do Japonês, ca. 15 km ao norte de Alto Paraíso pela GO 118, 14 December 2012, R. C. Sodré et al. 399 ( UFG), 400 ( UFG) ; ibd., M. J. Silva et al. 4636 ( UFG), 4638 ( UFG) ; ibd., 14°02’14.8”S, 47°31’46.9”W, 1540 m, 06 April 2013, R. C. Sodré et al. 668 ( UFG) ; GO 118 , km 153, entre São João d’Aliança e Alto Paraíso de Goiás, 14°14’3.8”S, 47°29’12.2”W, 31 October 2013, M. J. Silva et al. 5475 ( UFG), 5477 ( UFG), 5492 ( UFG) ; lateral direita do morro do Japonês, ca. 7–8 km de Alto Paraíso de Goiás, 14°1’13.7”S, 47°31’8.6”W, 1459 m, 02 November 2013, M. J. Silva et al. 5531 ( UFG), 5532 ( UFG), 5549 ( UFG) ; a direita do km 131 da GO 118, de São João d’Aliança para Alto Paraíso de Goiás, 14°25’18.3”S, 47°30’27.4”W, 1005 m, 20 February 2014, M. J. Silva et al. 5843 ( UFG), 5844 ( UFG) ; km 131 da GO 118, de São João d’Aliança a Alto Paraíso de Goiás, 14°25’20.7”S, 47°30’35”W, 1000 m, 20 February 2014, R. C. Sodré et al. 1217 ( UFG), 1219 ( UFG). Serra Geral de Goiás, rio Paranã, ca. 35 km N. of Formosa on Road to São Gabriel, 14°S, 46°W, 950 m, 28 March 1966, H. S. Irwin et al. 14185 ( UB) .

Caruzo & Cordeiro (2007) reported this species from Bolivia and Brazil (DF, MT, MS, MG, PR, RJ, and SP). It is reported here in Goiás state for the first time, where it grows in campo limpo and in cerrado rupestre in the municipalities of Água Fria de Goiás, Alto Paraíso de Goiás and São João d’Aliança ( Fig. 9C ), flowering and fruiting from October to May.

Croton grandivelus can be easily recognized by the yellowish to ferrugineous indument of long stipitate, stellate-porrect trichomes that densely cover both vegetative and reproductive parts, as well as by the usually obovate or oblanceolate leaves, pistillate flowers with large, linear to narrowly lanceolate sepals (5–10 mm long), linear petals, and large capsules (7–9 mm long) densely covered by stipitate stellate trichomes.

Taxonomic notes:— Since there are two sheets of the type at P, we selected as second-step lectotype the one referred to specifically by Baillon in the protologue as Glaziou’s annotation of “ C. campestre var. grandivelum ”. Croton grandivelus belongs to C. section Adenophylli .

2.18. Croton hirtus L’Héritier de Brutelle (1785: 17) . Type:— FRENCH GUIANA. Without locality, no date, L. C. Richard s.n. (holotype P 00623551!). ( Figs. 13 , D–H; 14 , A)

Subshrubs 0.15–0.45 m tall, erect, stems greenish with sessile or subsessile stellate-porrect trichomes; latex clear. Stipules 1.8–6 × 0.2–0.3 mm, linear, with glands ovoid or subglobose, persistent; petioles 4–8 mm long, with 2 stipitate patelliform glands. Leaves alternate or whorled near to dichotomy of the branches, blades 3.5–5.4 × 2–3.3 cm, ovate, base obtuse or asymmetric, apex acute, margin serrate without glands; upper surfaces with hispid trichomes and lower surfaces with sessile stellate-porrect trichomes, membranaceous, venation brochidodromous, with 4–6 pairs of secondary veins. Racemes 0.8–3.5 cm long, terminal; inflorescence axis, dorsal surface of bracts and sepals of both flowers, ovary and styles with hispid or stellate-porrect trichomes; bracts of both flowers 1.4–2.4 × 0.3–0.5 mm, narrowly lanceolate, with stipitate pyriform glands. Staminate flowers 2.4–3 mm long, pedicels 0.7–1.2 mm long; sepals 5, 0.9–1.1 × 0.7–0.8 mm, ovate, apex acute; petals 5, 1–1.4 × 0.5–0.6 mm, obovate, apex obtuse, ventral surface villous, dorsal surface glabrous; stamens 11, filaments glabrous. Pistillate flowers 2.4–3.3 mm long, pedicels 0.3–0.7 mm long; sepals 5, 0.5–2.9 × 0.2–1.2 mm, unequal, oblanceolate linear-oblanceolate, lanceolate or triangular, apex acute, margins entire without glands; petals absent, disk 5-segmented; ovary 0.5–0.9 × 0.7–1.1 mm, subglobose, styles 2-fid, pubescent. Capsules 4–4.2 × 3.8–4 mm, subglobose, greenish; seeds 3.5–3.7 × 2.5–2.6 mm, oblongoid, grayish, spotted.

Specimens examined:— BRAZIL. Goiás: Caldas Novas, Serra de Caldas Novas, termas do rio Quente , 04 January 1977, E. P. Heringer et al. 16669 ( IBGE). Cavalcante , imediações da estrada que leva a Colinas do Sul , a aproximadamente 18 km de Cavalcante , 29 January 2014, R. C. Sodré et al. 1178 ( UFG) ; porção central do povoado Kalunga Engenho II, 16 February 2013, R. C. Sodré & M. T. Faria 537 ( UFG) .

This is a weedy Neotropical species present in all regions of Brazil ( Caruzo & Cordeiro 2007), and introduced in the Old World. It was collected in anthropic environments and cerrado sensu stricto in the municipality of Cavalcante ( Fig. 9C ), flowering and fruiting in January and February. Its morphological relationships with C. glandulosus are discussed in the comments of that species. Both belong to C. section Geiseleria .

2.19. Croton horminum Baillon (1864: 315) . Type:— BRAZIL. Minas Gerais: without locality, 1838, M. P. Claussen 768 (holotype: P 00623543!). ( Figs. 13 , I–K; 14 , B–D)

Subshrubs 0.2–0.85 m tall, erect, stems canescent-green to ferrugineous with stipitate stellate-porrect trichomes; latex clear. Stipules 0.5–2 × 0.1–0.3 mm, oblong to narrowly lanceolate, with globose glands at the base an one gland at the apex, persistent; petioles 0.2–0.8(1.5) cm long, without glands. Leaves alternate or whorled near to dichotomy of the branches, blades 2.2–5.8 × 0.7–1.9 cm, lanceolate or narrowly elliptic, base obtuse, apex acute, margin entire without glands; upper and lower surfaces with shortly stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, with 4–8 pairs of secondary veins. Thyrses 0.6–2.5 cm long, terminal, staminate cymules with 1 flower; inflorescence axis, dorsal surface of bracts and sepals of both flowers, pistillate petals, ovary and styles with subsessile stellate-porrect trichomes; staminate bracts 0.6–0.8 × 0.2–0.3 mm, lanceolate, triangular or oblong; pistillate bracts 0.6–0.9 × 0.2–0.5 mm, lanceolate or triangular, both bracts with 2 stipitate-capitate glands at the base. Staminate flowers 3–4 mm long, pedicels 0.9–1.6 mm long; sepals (4)5, 1.1–1.4 × 0.7–1.2 mm, ovate, apex acute; petals 5, 2–2.4 × 0.4–0.5 mm, narrowly oblanceolate, apex obtuse, glabrescent on both surfaces, ciliate; stamens 11, filaments glabrous. Pistillate flowers 3.5–6 mm long, pedicels 1–2 mm long, typically recurved; sepals 5, 1.2–2.6 × 0.3–0.7 mm, oblong to narrowly elliptic, apex acute, margins entire without glands; petals 5, 0.4–1.2 × 0.05–0.1 mm, linear, disk 5- lobed; ovary 1–2 mm long, globose, styles 2-fid, pubescent. Capsules 4–5 mm long, globose to oblongoid, canescent-green to ferrugineous; seeds 2.8–3.9 × 1.7–2.3 mm, ellipsoid, grayish, spotted.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, ca. 48 km a oeste do município, na estrada para Niquelândia, 15 km a oeste da sede do PNCV, fazenda de Dom Manuel na trilha até as inscrições, 14°6’S, 47°54’W, 4 March 1988, S. Ginzbarg et al. 728 ( UB). Cavalcante , balsa da COTERRA (rio Tocantins) vila veneno km 1, área de influência da futura hidrelétrica de Cana Brava, 13°34’16”S, 48°6’4”W, 300 m, 13 December 2000, G. Pereira-Silva & J.B. Pereira 4469 ( CEN, UFG) ; estrada Cavalcante ao povoado Kalunga Engenho II, 13°45’49”S, 47°27’18.9”W, 804 m, 07 March 2013, R. C. Sodré 566, 567 ( UFG) ; ibd., M. J. Silva 4745, 4746, 4747, 4748, 4749, 4750 ( UFG) .

This is a Brazilian species described from Minas Gerais by Müller Argoviensis (1873) but apparently more common in Goiás state. We found it in cerrado sensu strito of Alto Paraíso de Goiás and Cavalcante municipalities ( Fig. 9D ), flowering and fruiting in December and March.

The combination of green-whitish to rusty branches, entire-margined, eglandular leaves, inflorescences up to 2.5 cm long, bracts 0.6–0.9 mm long, flowers of both sexes shortly pedicellate (pedicels up to 2 mm long), and oblongoid capsules with typically curved pedicels, make C. horminum easily recognizable and distinct from other congeners. It belongs to C. section Pedicellati .

2.20. Croton inaequilobus Steyermark (1958: 7) . Type:— BRAZIL. Goiás: Chapada dos Veadeiros, 14 Km South of Veadeiros , in sandstone rocky area on West bank of stream and above, 47°30’S, 14°20’W, 25 April 1956, E. Y. Dawson 14685 (holotype R 000104977!, isotypes F V0056145F!, RSA 0002666!). ( Figs. 14 , E–H; 15 , A–E)

Subshrubs to shrubs 0.6–2 m tall, erect, young stems canescent to pinkish with stipitate stellate-porrect trichomes; latex clear. Stipules 0.2–1.5 × 0.05–0.2 mm, ovate or linear, with globose glands at the apex, caducous; petioles 0.5–3.3 cm long, with 2 sessile patelliform glands, orange to reddish. Leaves alternate or opposite near the dichotomies of the branches, blades 3–9.8 × 1.3–4.5 cm, ovate or oval-lanceolate, base obtuse or sometimes truncate, apex acute or obtuse, margin crenate, serrate or triply serrate with shortly stipitate patelliform glands; upper and lower surfaces with stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, with (3)4–5 pairs of secondary veins. Thyrses 1–2.9 cm long, terminal, staminate cymules with 1 flowers; inflorescence axis, both faces of sepals pistillate and bracts of both flowers, dorsal surface of staminate sepals, ovary and styles with stellate-porrect trichomes; staminate bracts 1.6–2.5 × 0.2–0.3 mm, narrowly lanceolate, with globose glands at the base; pistillate bracts 1.7–2.2 × 0.3–0.4 mm, narrowly lanceolate. Staminate flowers 3.5–5 mm long, pedicels 1–2 mm long; sepals 5, 1.5–2 × 0.8–1.3 mm, ovate, apex acute; petals 5, 1.8–2.2 × 0.6–0.8 mm, elliptic to oboval-oblanceolate, apex acute, villous on both surfaces; stamens 11, filaments glabrescent. Pistillate flowers 3.3–6.5 mm long, pedicels 0.7–1 mm long; sepals 5, 0.9–4.2 × 0.2–2.2 mm, unequal, obovate, oblanceolate, lanceolate, narrowly lanceolate or oblong, apex obtuse or acute, margins entire without glands; petals 5, 0.5–1 × 0.1–0.15 mm, narrowly lanceolate, disk 5-lobed; ovary 1.1–1.9 mm long, globose, styles 2-fid, ca. 2 mm long, pubescent. Capsules 4–4.5 mm long, globose, canescent-green; seeds 3–3.2 × 1.8–2.2 mm, ovoid or oblongoid, light brown, spotted.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros , 31 December 1990, M. Armando & F. Leite 1107 ( CEN) ; imediações do Cânion 1 do rio Preto , próximo a garganta dele, em área ilhada 14°8’38.7”S, 47°48’19.14”W, 949 m, 15 December 2012, M. J. Silva et al. 4666 ( UFG), 4667 ( UFG), 4668 ( UFG), 4669 ( UFG), 4670 ( UFG), 4671 ( UFG) ; ibd., R. C. Sodré et al. 418 ( UFG), 419 ( UFG), 420 ( UFG), 421 ( UFG), 422 ( UFG), 425 ( UFG) ; trilha em direção às Sete Quedas do Rio Preto, cerca de 700 m do rio Preto, a noroeste do Morro do Buracão , 14°6’7”S, 47°43’32.7”W, 1158 m, 10 February 2013, M. J. Silva et al. 4722 ( UFG), 4724 ( UFG), 4726 ( UFG) ; ibd., R. C. Sodré et al. 499 ( UFG), 500 ( UFG), 501 ( UFG), 502 ( UFG) ; trilha para às Sete Quedas do Rio Preto , 14°6’43.9”S, 47°43’20.9”W, 1192 m, 10 February 2013, R. C. Sodré et al. 507 ( UFG) ; trilha para as Corredeiras do rio Preto , 14°9’54.2”S, 47°50’29.4”W, 854 m, 8 March 2013, R. C. Sodré et al. 585 ( UFG), 586 ( UFG), 587 ( UFG) ; ibd., M. J. Silva et al. 4777 ( UFG), 4778 ( UFG) ; imediações das Sete Quedas do rio Preto , 14°6’12.3”S, 47°44’10.5”W, 1128 m, 10 May 2013, R. C. Sodré et al. 730 ( UFG), 731 ( UFG), 734 ( UFG) ; cerca de 2 km a NO do morro do Buracão , 10 May 2013, M. J. Silva et al. 4888 ( UFG) ; ca. 2 km a norte do alojamento principal do Parque, 14°8’37”S, 47°47’37”W, 978 m, 05 June 2015, M. J. Silva et al. 6748 ( UFG). Cavalcante, após às Corredeiras do rio Preto , 14°9’17.4”S, 47°50’8.5”W, 876 m, 8 March 2013, R. C. Sodré et al. 593 ( UFG), 594 ( UFG) ; ibd., M. J. Silva et al. 4784 ( UFG) .

This species is apparently endemic to the PNCV, where it grows in cerrado rupestre near Rio Preto in the municipalities of Alto Paraíso de Goiás and Cavalcante ( Fig. 9D ). It flowers and fruits throughout the year, but especially from December to May.

Croton inaequilobus can be recognized by having a subshrubby to shrubby habit up to 2.4 m tall, cinereous leaf blades on abaxial surface, patelliform, orange, sessile acropeciolar glands, unequal pistillate sepals, and pubescent 2- fid styles. It belongs to C. section Geiseleria .

2.21. Croton leptobotryus Müller Argoviensis (1873: 155) . Type:— BRAZIL. Minas Gerais: ad Lagoa Santa super rupes calcareas et in virgultis ad Sumidouro, 02 December 1864, J. E. B. Warming 1628 (holotype C 10011177!, isotypes G 00434560!, A 00257981!). ( Fig. 15 , F–J)

Subshrubs 0.4–1 m tall, erect, stems greenish with sessile stellate-porrect trichomes; latex clear. Stipules 0.3–1 × 0.1–0.3 mm, lanceolate, with 2 globose glands at the base and one at the apex, persistent; petioles 2.6–7.1 cm long, with 2 obconical glands. Leaves alternate or whorled near to dichotomy of the branches, blades 8.5–13.5 × 4.8–8 cm, ovate, base obtuse, apex acuminate, margin serrate without glands; upper and lower surfaces with stellate-porrect trichomes, membranaceous, venation brochidodromous, 5–7 pairs of secondary veins. Thyrses 1.2–7.6 cm long, terminal, staminate cymules with (1–)3 flowers; inflorescence axis, both faces of sepals pistillate and bracts of both flowers, dorsal surface of staminate sepals, pistillate petals, ovary and styles with stellate-porrect trichomes; bracts of both flowers 0.8–1.1 × 0.3–0.4 mm, lanceolate or triangular, with globose glands. Staminate flowers 3.7–4.2 mm long, pedicels 1.2–2.1 mm long; sepals 5, 1.4–2 × 0.8–1.1 mm, ovate, oblong or oval-elliptic, apex obtuse or acute; petals 5, 1.6–2 × 0.4–0.7 mm, oblanceolate or narrowly oblanceolate, apex obtuse, ventral surface villous at the base; stamens 13–16, filaments glabrous. Pistillate flowers 3.4–4.2 mm long, pedicels 1.1–1.5 mm long; sepals 5, 2–2.2 × 0.7–1.1 mm, subequal, oblong or lanceolate, apex acute, margins entire without glands; petals glanduliform, globose, disk 5- segmented; ovary 0.9–1 × 1–1.1 mm, subglobose, styles 2-fid, pubescent. Capsules and seeds not seen.

Specimens examined:— BRAZIL. Goiás: Colinas do Sul, cerca de 2 km da ponte do rio Bagagem, 14°9’S, 48°04’W, 450 m, 23 November 1992, R. F. Vieira et al. 1380 (CEN). Nova Roma, fazenda Santa Clara, localidade Sucuri ou Pinga, 13°45’13”S, 46°51’31”W, 450 m, 29 February 2000, M. Aparecida da Silva 4273 (IBGE). São Domingos, Parque Estadual de Terra Ronca, 13°44’08”S, 46°21’27”W, 575 m, 29 October 2000, M. Aparecida da Silva 4588 (IBGE).

This species was reported by Caruzo & Cordeiro (2007) along the edges of mesophilic forests in Goiás, Minas Gerais and São Paulo States. In this study, it was collected in these same environments in Colinas do Sul and Nova Roma ( Fig. 9D ), flowering in October, November and February.

Croton leptobotryus stands out among its other congeners by having petioles 2.6 to 7.1 cm long with obconical glands at the apex, oval to oval-lanceolate leaf blades with serrate margins and an acuminate apex, inflorescences with laxly distributed flowers, diminute bracts (0.8–1.1 mm long), and staminate flowers with 13–16 stamens. It belongs to C. section Geiseleria .

2.22. Croton longifolius Müller Argoviensis (1873: 170) . Lectotype (designated here):— BRAZIL. Goiás: in Serra de Montes Claros et Corumba, no date, J. B. E. Pohl 1609 (Catalog number 1038) (W 0051291!, isolectotypes: G 00434552!, K 000186133!). ( Figs. 14 , I–K; 15 , K–M)

Subshrubs 0.3–0.5 m tall, erect, stems ferrugineous, tomentose with shortly stipitate stellate-porrect trichomes; latex clear or orange after oxidized. Stipules 0.1–0.2 mm long, glanduliform, ovoid or subglobose, persistent; petiole 0.5–1 (1.5) mm long, without glands. Leaves alternate, blades 5.5–13.5 × 1.7–4.2 cm, lanceolate, or sometimes narrowly elliptic, base obtuse, apex acute, margin entire without glands; upper and lower surfaces tomentose with shortly stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, 9–13 pairs of secondary veins. Thyrses (2)4.5–11(20) cm long, terminal, flowers sparsely distributed along the axis, staminate cymules with 1–3 flowers; inflorescence axis, dorsal surface of sepals and bracts of both flowers, ovary and styles tomentose with sessile stellate-porrect trichomes; bracts of both flowers 0.8–2.2 × 0.2–0.6 mm, lanceolate, with ovoid or subglobose glands. Staminate flowers 4–4.5 mm long, pedicels 1–1.3 mm long; sepals 5, 2.2–2.5 × 1.1–1.4 mm, ovate, apex acute; petals 5, 2.3–2.8 × 0.8–0.9 mm, oblanceolate, or sometimes elliptic, apex acute or obtuse, ventral surface villous basally; stamens 16, glabrous. Pistillate flowers 5–6.2 mm long, pedicels 0.2–0.5 mm long; sepals 5, 1.8–2.1 × 0.8–1 mm, subequal, triangular, apex acute, margin entire, without glands, petals 5, 0.3–0.8 mm long, linear, with hirsute trichomes, disk 5-segmented; ovary 1.4–1.8 × 1.8–2 mm, subglobose, styles 2-fid, pubescent. Capsules 6.2–6.6 × 6.4–6.7 mm, globose, brownish, columella with three prominent, smooth lobes; seeds 3.9–4.8 × 2.3–3 mm, ellipsoid, grayish, lightly maculate.

Specimens examined:— BRAZIL. Distrito Federal: Gama, a esquerda do km 19 da BR 060, sentido Brasília , 15°58’28.6’’S, 48°11’27.3’’W, 984 m, 28 January 2014, M. J. Silva et al. 5698, 5699 ( UFG) ; ibd., R. C. Sodré et al. 1156 ( UFG). Goiás: Alto Paraíso de Goiás, estrada para o Vale da Lua , 14°10’20.4’’S, 47°47’43.7’’W, 1023 m, 10 March 2012, R. C. Sodré et al. 60 ( UFG) ; ibd., 03 January 2015, R. C. Sodré, M. J. Silva & C. H. G. Machado Filho 1605 ( UFG) ; estrada para o Vale da Lua , abaixo do portão da SANEAGO, 14 December 2013, M. J. Silva et al. 5677 ( UFG) ; acima da primeira baixada que leva ao Vale da Lua , 14 December 2013, M. J. Silva et al. 5685 ( UFG) ; início da estrada para o Vale da Lua , 14 December 2013, R. C. Sodré et al. 1081 ( UFG), 1082 ( UFG) ; a direita do km 131 da GO 118, entre São João d’Aliança e Alto Paraíso de Goiás, 14°25’18.3’’S, 47°30’27.4’’W, 1024 m, 20 February 2014, M. J. Silva, T. M. S. Melo & R. C. Sodré 5856 ( UFG), 5863 ( UFG) ; ibd., 14°25’20.7’’S, 47°30’35’’W, 1000 m, 20 February 2014, R. C. Sodré, M. J. Silva & T. M. S. Melo 1213 ( UFG), 1214 ( UFG) ; próximo ao Posto Vale da Lua, 14°8’19.3’’S, 47°31’23.8’’W, 1241 m, 16 February 2013, R. C. Sodré & M. T. Faria 548 ( UFG). Cavalcante, a esquerda do km 204 da GO 118, entre Alto Paraíso de Goiás e Teresina de Goiás, 13°54’16’’S, 47°22’12.9’’W, 1299 m, 21 February 2014, R. C. Sodré, T. M. S. Melo & M. J. Silva 1222 ( UFG), 1223 ( UFG). Santo Antônio do Descoberto , rio Brechó , 01 December 1976, A. Allem 622 ( CEN) .

According to BFG (2015), Croton longifolius is endemic to Goiás State and known then only from the type collection. In the Chapada dos Veadeiros it has been collected in cerrado sensu stricto and campos sujos of Alto Paraíso de Goiás and Cavalcante ( Fig. 16A ), flowering and fruiting from October to January. It is characterized by its ferrugineous branches, leaves and inflorescences, the striated branches and axes of the inflorescence, the subsessile leaves with lanceolate to linear-lanceolate leaf blades, and the small bracts (0.8–1.1 mm long), and flowers laxly distributed on the inflorescence. It belongs to C. section Adenophylli .

2.23. Croton megalocalyx Müller Argoviensis (1873: 220) Lectotype (designated here):— BRAZIL. Goiás: in campis lapidosis petrosisque in Serra S. Marcos prope Chapadão , August 1834, L. Riedel 2501 (LE!, isolectotypes G 00434617!, K 000186097!, P 00633432!). ( Figs.14 , L–N; 17 , A–E)

Subshrubs or shrubs 1–1.7m tall, erect,stems yellowish-ferrugineous or blackish with stipitate stellate-porrect trichomes; latex clear. Stipules 7–15 × 0.4–2 mm, linear or narrowly lanceolate, with papilliform glands basally or on the margin, persistent; petioles 0.6–2 cm long, with 5–10 shortly stipitate patelliform glands. Leaves alternate or opposite near to dichotomy of the branches, blades 4.1–8 × 2.8–5.2 cm, ovate, oval-lanceolate or sometimes elliptic, base obtuse, asymmetric or hardly ever cordate, apex acuminate, margin entire or serrate with shortly stipitate patelliform glands; upper and lower surfaces with shortly stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, with 4–6 pairs of secondary veins. Thyrses 2–4.8 cm long, terminal, staminate cymules with 1 flowers; inflorescence axis, both faces of sepals pistillate and bracts of both flowers, dorsal surface of staminate sepals and petals, filaments, ovary and styles with stellate-porrect trichomes; staminate bracts 7–10 × 0.4–0.7 mm, linear; pistillate bracts 8–16 × 1–1.5 mm, narrowly lanceolate, both bracts with papilliform glands. Staminate flowers 10–12 mm long, pedicels 2.3–4 mm long; sepals 5, 3–3.5 × 1.5–2.4 mm, ovate, apex acute; petals 5, 4–5.2 × 0.5–0.7 mm, linear or narrowly lanceolate, apex acute, ventral surface glabrescent; stamens 11, filaments with stellate-porrect trichomes. Pistillate flowers 9–12 mm long, pedicels 1.5–2 mm long; sepals 5–7, 4–8.5 × 0.7–5 mm, unequal, ovate to lanceolate, or less frequently narrowly lanceolate, apex acute or acuminate, margins entire or laciniate, without glands; petals 5–7, 2–4.5 × 0.2–0.4 mm, linear, disk 3-lobed; ovary 1.5–2 mm long, globose, styles 2–4-fid, pubescent. Capsules 5–5.5 mm long, globose, grayish-green; seeds 3.9–4.5 × 2.1–2.3 mm, ellipsoid, grayish, maculate.

Specimens examined:— BRAZIL. Goiás: Cavalcante, caminho para a cachoeira Santa Bárbara , 07 March 2003, J. F. B. Pastore & E. Suganuma 385 ( CEN, UFG) ; em direção ao Engenho II, região dos kalungas, 13°42’1.4”S, 47°27’50.4”W, 1122 m, 05 April 2013, R. C. Sodré et al. 654 ( UFG), 655 ( UFG), 656 ( UFG), 658 ( UFG), 660 ( UFG) ; ibd., M. J. Silva et al. 4807 ( UFG), 4808 ( UFG), 4809 ( UFG), 4810 ( UFG), 4811 ( UFG), 4813 ( UFG). Teresina de Goiás, Km 12 da Rod. GO 118, Teresina ( Distrito de Cavalcante ), 30 Apr 2012, A. Allem & G. Vieira 2742 ( CEN) .

This species is probably endemic to Goiás state ( BFG 2015). It was collected in cerrado sensu stricto in Cavalcante and Teresina de Goiás municipalities ( Fig. 16A ), with flowers in March and flowers and fruits in April and July.

Croton megalocalyx has a shrubby habit 1–1.7 m tall, blackish stems, greenish-yellowish young stems and leaves, and a deltoid, ferrugineous inflorescence. In addition, it has several glands at the apex of the petiole and leaf margins, stipules 0.7–1.5 cm long with papilliform glands at the base, large bracts (7–16 mm long) and flowers of both sexes more than 1 cm long. These characters make this species easily to diagnose. It belongs to section Barhamia subsection Medea .

Taxonomic notes:— There are two duplicates of Riedel 2501 at LE; the lectotype is the sheet with two labels at the bottom, as well as an annotation of the species name by Müller.

2.24. Croton odontadenius Müller Argoviensis (1873: 227) . Lectotype (designated here):— BRAZIL. Minas Gerais: in campis aridis petrosis prope Lagoa Santa, 05 October 1863, J. E. B. Warming 1642 (C 10011184!, isolectotype G 00434597!). Syntypes:— BRAZIL. Minas Gerais: without locality, 1845, J. F. Widgren s.n. (S R-10551!) ; in campis aridis petrosis prope Lagoa Santa, 05 October 1863, J. E. B. Warming 1641 (C 10011185!). ( Figs. 17 , F–J; 18 , A–G)

Subshrubs 0.2–0.55 m tall, subdecumbent or rarely erect, stems brownish with simple or sessile with 2–7 radiate trichomes; latex clear. Stipules 4–10 × 0.6–1.3 mm, linear or narrowly lanceolate, with ovoid to subulate glands at the base, persistent; petioles 0.4–1.6 cm long, with or without 2 shortly stipitate patelliform glands. Leaves alternate or opposite near to dichotomy of the branches, blades 5.3–12.5 × 1.8–4 cm, oval-lanceolate, elliptic, or less frequently oblong, base acute or asymmetric, apex acute, margin triply serrate with shortly stipitate patelliform glands; upper and lower surfaces with stellate-radiate and stellate-porrect trichomes respectively, membranaceous, venation semi-craspedodromous, with 5–6(7) pairs of secondary veins. Thyrses 1.8–3 cm long, terminal, staminate cymules with 2 or 3 flowers; inflorescence axis, both faces of sepals pistillate and bracts of both flowers, dorsal surface of staminate sepals, ovary and styles with stellate-radiate and stellate-porrect trichomes; staminate bracts 2.5–3.5 × 0.5–1 mm, linear to lanceolate; pistillate bracts 2.8–6 × 0.4–0.8 mm, linear or narrowly lanceolate, both bracts with ovoid glands at the base. Staminate flowers 6.5–8 mm long, pedicels 1.2–2.3 mm long; calyx 5,6–lobed, lobes 2.3–3.2 × 1–1.2 mm, united up to half, ovate, apex acute or obtuse; petals 6(5), 2.8–3.7 × 1–1.8 mm, obovate to oblanceolate, apex obtuse, ventral surface villous; stamens (13–)17–24, filaments glabrous or glabrescent. Pistillate flowers 7–9 mm long, pedicels 0.5–1.7 mm long; sepals (6)7–8, 4–6 × 0.9–2.1 mm, subequal, oblong or lanceolate, apex acute, margins entire, with globose glands; petals 5, 0.2–1 × 0.05–0.1 mm, linear, disk 7–8-lobed; ovary 1.4–1.7 mm long, oblong-globose, styles 4-fid, pubescent. Capsules 5–5.5 mm long, globose, green glossy; seeds 3.9–4 × 3–3.4 mm, broadly ellipsoid.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, ca. 20 km W. of Veadeiros , 14°S, 47°W, 1000 m, 16 February 1966, H. S. Irwin et al. 12915 ( UB) ; ibd., 17 February 1966, H. S. Irwin et al. 12546 ( UB) ; ca. 18 km N. of Alto Paraíso , ca. 1250 m, 21 March 1971, H. S. Irwin, R. M. Harley & G. L. Smith 32878 ( UB) ; 4 Km NE of road, 16 Km by road N of Alto Paraíso, 14°S, 47°W, 1600 m, 03 February 1979, H. Gates & H. Estabrook 97 ( UB) ; Serra do Pouso Alto , imediações da estrada que se inicia no cruzeiro (GO 118), 13°58’13.7”S, 47°29’15.8”W, 1542 m, 21 January 2012, R. C. Sodré et al. 26 ( UFG), 29 ( UFG) ; a esquerda da GO 118, cerca de 15 km após Alto Paraíso de Goiás, em direção a Teresina de Goiás, 09 March 2012, M. J. Silva et al. 4132 ( UFG), 4133 ( UFG) ; GO 118 , próximo a placa indicatica a Serra do Pouso Alto , 14°02’44.8”S, 47°31’25.7”W, 1083 m, 09 March 2012, R. C. Sodré et al. 54 ( UFG), 55 ( UFG) ; ca. 10 km ao norte de Alto Paraíso de Goiás pela GO 118, 14°03’43”S, 47°30’14”W, 25 May 2012, R. C. Sodré 101 ( UFG), 102 ( UFG) ; ca. 7 km da cidade em direção a Teresina de Goiás, na borda da GO 118, 25 May 2012, M. J. Silva et al. 4201 ( UFG) ; ca. 15 km do município, a direita da GO 239, de Alto Paraíso de Goiás para o povoado de São Jorge, próximo ao Morro da Baleia , 14°9’29.64”S, 47°37’54.9”W, 1168 m, 13 December 2012, R. C. Sodré et al. 366 ( UFG), 367 ( UFG), 368 ( UFG), 373 ( UFG) ; ibd., 08 February 2013, R. C. Sodré et al. 461 ( UFG) ; imediações do morro do Japonês , 14 December 2012, R. C. Sodré et al. 394 ( UFG) ; imediações do Posto de Combustível Vale da Lua , 14°8’19.3”S, 47°31’23.8”W, 1241 m, 16 February 2013, R. C. Sodré & M. T. Faria 547 ( UFG) ; Serra do Pouso Alto , ca. 4 km da estrada do inicia no cruzeiro (GO 118), 13°57’44.1”S, 47°28’54”W, 1527 m, 05 April 2013, R. C. Sodré et al. 667 ( UFG) ; GO 118 , entre Alto Paraíso de Goiás e Teresina de Goiás, 05 April 2013, M. J. Silva et al. 4821 ( UFG), 4822 ( UFG) ; próximo do Morro do Japonês, ca. 15 km ao norte de Alto Paraíso pela GO 118, 14°2’14.8”S, 47°31’46.9”W, 1540 m, 06 April 2013, R. C. Sodré et al. 676 ( UFG) ; ibd., 14°2’36.5”S, 47°31’43.1”W, 1482 m, 11 May 2013, R. C. Sodré et al. 737 ( UFG) ; ibd., 14°3’15.2”S, 47°32’19.5”W, 1513 m, 11 May 2013, R. C. Sodré et al. 746 ( UFG) ; ibd., 14°3’7.9”S, 47°32’41.3”W, 1489 m, 11 May 2013, R. C. Sodré et al. 753 ( UFG) ; ibd., 14°2’33.6”S, 47°32’28.4”W, 1455 m, 11 May 2013, R. C. Sodré et al. 758 ( UFG) ; ibd., 14°2’38.1”S, 47°31’45.7”W, 1528 m, 03 January 2015, R. C. Sodré et al. 1602 ( UFG) ; ca. 4 km ao norte do morro da Baleia em direção ao rio Preto , 14°5’12.3”S, 47°38’12.4”W, 1175 m, 03 August 2013, R. C. Sodré et al. 819 ( UFG) ; ca. 30 km a esquerda da GO 118, de São João d’Aliança a Alto Paraíso de Goiás, na estrada para o rio dos Couros, 14°17’35”S, 47°42’32”W, 1028 m, 02 January 2015, R. C. Sodré et al. 1576 ( UFG), 1585 ( UFG), 1588 ( UFG). Cavalcante, serra do Pouso Alto, cerca de 3 km a esquerda do cruzeiro na GO 118, de Alto Paraíso a Teresina de Goiás, 13°56’31.3”S, 47°29’54.8”W, 1078 m, 21 January 2012, M. J. Silva et al. 4074 ( UFG), 4084 ( UFG), 4087 ( UFG) ; ibd., 13°56’39.3”S, 47°29’52.8”W, 1084 m, 09 March 2012, R. C. Sodré et al. 50 ( UFG) ; Serra do Pouso Alto, cerca de 4 km após o cruzeiro, sentido Alto Paraíso a Teresina de Goiás, 20 April 2012, M. J. Silva et al. 4186 ( UFG) ; imediações do km 205 da GO 118 de Alto Paraíso de Goiás à Teresina de Goiás, 26 July 2014, R. C. Sodré et al. 1361 ( UFG) .

This is a Brazilian species reported so far in Minas Gerais ( BFG 2015). We found it in the municipalities of Alto Paraíso de Goiás and Cavalcante ( Fig. 16A ), growing in campo limpo and cerrado rupestre, especially inside the PNCV and surrounding areas. It flowers and fruits from December to May.

This species resembles C. sclerocalyx ( Didrichsen 1857: 134) Müller Argoviensis (1865: 134) by having a similar petiole with subsessile patelliform glands, the staminate calyx with sepals united up to half their length, pistillate flowers with six to eight sepals, and multifid styles. However, C. odontadenius has staminate flowers with 17–24 stamens (vs. 9–11 in C. sclerocalyx ) and entire pistillate sepals (vs. serrate), and usually decumbent habit (vs. erect). It belongs to section Geiseleria .

2.25. Croton planaltoanus Silva et al. (2014: 89) . Type:— BRAZIL. Goiás: Alto Paraíso de Goiás, cerca de 11 km da cidade em direção a Teresina de Goiás após o Morro do Japonês , 14°02’00”S 47°3140.3”W, 1540 m, 24 November 2012, M. J. Silva & J. P. Santos 4577 (holotype UFG!, isotype UB!). ( Figs. 17 , K–N; 18 , H–K)

Subshrubs 0.15–0.6 m tall, erect or subdecumbent, stems canescent-green with subsessile stellate-porrect trichomes; latex clear. Stipules 0.4–4 × 0.2–0.7 mm, oblong or lanceolate, with globose glands at the apex, caducous; petioles 0.3–1.2 cm long, with or without 1 shortly stipitate patelliform gland, yellowish. Leaves alternate or opposite near to dichotomy of the branches, blades 2.7–8.5 × 1.5–4.3 cm, elliptic or discretely ovate, base acute, obtuse or attenuate, apex acute or acuminate, margin serrate or doubly serrate with sessile patelliform glands; upper and lower surfaces with stellate-porrect trichomes respectively, membranaceous, venation brochidodromous, with 4–6 pairs of secondary veins, the basal ones never surpassing half the length of the blade. Thyrses 4.2–7(–15) cm long, terminal, staminate cymules with 1 flowers; inflorescence axis, both faces of sepals pistillate and bracts of both flowers, dorsal surface of staminate sepals, ovary and styles with stellate-porrect trichomes; staminate bracts 2.1–2.9 × 0.2–0.4 mm, linear-oblanceolate; pistillate bracts 3–5 × 0.25–0.9 mm, linear or lanceolate, both bracts without glands. Staminate flowers 4.5–5.5 mm long, pedicels 1.8–2.3 mm long; sepals 5, 1.6–2 × 0.8–1.1 mm, ovate, apex acute; petals 5, 2–2.3 × 0.6–0.8 mm, elliptic or oblanceolate, apex obtuse, villous on both surfaces; stamens 11, filaments villous. Pistillate flowers 5–9 mm long, pedicels 0.5–1 mm long; sepals 5, 1.6–4.5 × 0.4–2.5 mm, unequal, obovate, oblong or ovate, apex acute or obtuse, margins entire, without glands; petals absent or 5, 2.2–2.8 × 0.05–0.1 mm, oblong or linear, disk 5-lobed; ovary 1–1.8 mm long, globose, styles 2-fid, 3.5–5 mm long, pubescent. Capsules 5–5.5 mm long, globose, greenish; seeds 3.7–4.2 × 2.7–3 mm, ovoid or ellipsoid, spotted.

Specimens examined:— BRAZIL. Goiás: Água Fria de Goiás, a direita do Km 61 da GO 118, sentido São Gabriel a São João d’Aliança , 14°59’23.4”S, 47°36’16.4”W, 1119 m, 30 October 2014, R. C. Sodré, A. O. Souza & L. S. Inocencio 1466 ( UFG), 1469 ( UFG). Alto Paraíso de Goiás, ca. 12 km N.W. de Veadeiros, estrada para Cavalcante, 14°S, 47°W, 1,200 m, 21 October 1965, H. S. Irwin, R. Souza & Reis dos Santos 9439 ( UB) ; estrada para Colinas do Sul a 14 km de Alto Paraíso de Goiás, 24 November 1994, F. C. A. Oliveira 224 ( IBGE) ; Região de Pouso Alto, ca. 20 km de Alto Paraíso de Goiás em direção a Teresina de Goiás, 14°01’10.7”S, 47°32’41.6”W, 1,150 m, 26 October 2012, M. J. Silva & R. C. Sodré 4488 ( UFG), 4489 ( UFG), 4490 ( UFG), 4491 ( UFG), 4492 ( UFG), 4493 ( UFG) ; ibd., R. C. Sodré & M. J. Silva 236 ( UFG), 237 ( UFG), 238 ( UFG), 239 ( UFG), 240 ( UFG), 241 ( UFG) ; ibd., cerca de 11 km de Alto Paraíso de Goiás para Teresina de Goiás 14°02’00”S, 47°31’40.3”W, 24 November 2013, M. J. Silva et al. 4570 ( UFG), 4571 ( UFG), 4572 ( UFG), 4573 ( UFG), 4579 ( UFG) ; entrada para o Hotel Água Fria, região do Pouso Alto , 14°04’22.4”S, 47°30’35.3”W, 1355 m, 16 February 2013, R. C. Sodré & M. T. Faria 544 ( UFG) ; cerca de 1 km a esquerda da GO 239, de Alto Paraíso ao povoado São Jorge , 14°10’2.4”S, 47°37’55.8”W, 06 September 2014, R. C. Sodré et al. 1446 ( UFG) ; cerca de 12 km após Alto Paraíso, a direita da GO 239, em direção ao povoado São Jorge , 31 October 2014, R. C. Sodré et al. 1480 ( UFG), 1481 ( UFG), 1482 ( UFG) ; a noroeste de Alto Paraíso, ca. 1 km a direita da entrada da cidade, 14°7’59”S, 47°31’47.9”W, 1278 m, 31 October 2014, R. C. Sodré et al. 1485 ( UFG). Cavalcante, estrada para povoado Kalunga , Engenho II, 13°42’1.4”S, 47°27’50.3”W, 898 m, 05 April 2013, R. C. Sodré et al. 653 ( UFG) .

This species is endemic to Goiás state, where it grows in campos limpos and in cerrado rupestre in the municipalities of Água Fria de Goiás, Alto Paraíso de Goiás and Cavalcante ( Fig. 16B ) ( Silva et al. 2014). It flowers and fruits from September to April.

Croton planaltoanus is an erect or decumbent, cespitose subshrub up to 50 cm tall, with greenish to grayish branches. It typically has elliptical leaf blades that are concolorous, with serrate margins, an acute to shortly acuminate apex, pistillate flowers with unequal sepals and entire margins with trichomes on both surfaces, and 2-fid, pubescent styles and villous stamens that disitnguish it from others species in the area. It belongs to C. section Geiseleria .

Figures

FIGURE 3. Map of geographic distributions of the Crotoneae species in the Chapada dos Veadeiros, Goiás, Brazil. A. Astraea gardneri and A. lobata. B. Croton abaitensis, C. agoensis and C. agrarius. C. C. antisyphiliticus, C. arirambae and C. betaceus. D. C. campestris, C. catariae and C. cinerascens.

FIGURE 4. A−E. Croton abaitensis. A. Flowering branch. B. Detail of the leaf base showing the acropetiolar glands. C. Detail of the margin of the lower side of the leaf showing glands in the sinus and teeth apex. D. Staminate flower. E. Pistillate flower. (A: R. C. Sodré et al. 403; B−D: R. C. Sodré et al. 1000; E: R. C. Sodré et al. 81). F−H. Croton agrarius. F. Flowering branch. G. Staminate flower. H. Pistillate flower. (F−H: R. C. Sodré et al. 1187). I−M. Croton antisyphiliticus. I. Flowering branch. J. Detail of the base of leaf showing the glands. K. Detail of the margin of the lower side of the leaf showing a gland in the sinus. L. Staminate flower. M. Pistillate flower with a sepal removed, showing the ovary and two petals. (I−M: R. C. Sodré et al. 206).

FIGURE 5. A−C. Croton abaitensis. A. Habit. B. Inflorescence showing staminate flowers. C. Capsules. D and E. C. agoensis D. Habit. E. Inflorescence showing pistillate flowers. F and G. C. agrarius. F. Habit. G. Inflorescence showing young fruit and staminate flowers. H and I. Habit of C. antisyphiliticus. J−L. C. campestris. J. Habit. K. Inflorescence showing staminate flowers. L. Inflorescence showing staminate and pistillate flowers.

FIGURE 6. A−D. Croton arirambae. A. Flowering branch. B. Detail of the upper side of a leaf showing stellate trichomes and marginal glands. C. Staminate flower.D. Pistillate flower with a sepal removed to show the ovary and bifid styles. (A−D: J. F. B. Pastore et al. 813). E−H. Croton betaceus. E. Flowering branch. F. Detail of a leaf base showing the acropetiolar glands. G. Staminate flower. H. Pistillate flower. (E−H: M. J. Silva et al. 5755). I−K. Croton campestris. I. Flowering branch. J. Staminate flower. K. Pistillate flower. (I−K: R. C. Sodré et al. 930).

FIGURE 7. A−E. Croton catariae. A. Flowering branch. B. Detail of a leaf base with lobes folded back to show the acropetiolar glands. C. Detail of the upper side of a leaf showing the marginal glands. D. Staminate flower. E. Pistillate flower. (A−E: M. J. Silva et al. 5852). F−H. Croton cinerascens. F. Flowering branch. G. Staminate flower. H. Pistillate flower. (F−H: R. C. Sodré et al. 1159). I−L. Croton desertorum. I. Flowering branch. J. Detail of the lower side of a leaf base showing the basilaminar glands. K. Staminate flower. L. Pistillate flower. (I−L: M. J. Silva et al. 4568).

FIGURE 8. A−D. Croton catariae. A. Habit. B. Flowering branch. C. Inflorescence showing staminate flowers. D. Capsules. E−H. C. desertorum. E. Habit. F. Flowering branch. G. Inflorescence showing young capsules and staminate flowers. H. Inflorescence showing young capsules and staminate buds. I−L. C. didrichsenii. I. Habit. J. Inflorescence showing staminate flowers. K and L. Inflorescence showing pistillate flowers. M−O. C. goyazensis. M. Flowering branch. N. Inflorescence showing staminate flowers. O. Inflorescence showing staminate and pistillate flowers.

FIGURE 9. Map of geographic distributions of the Crotoneae species in the Chapada dos Veadeiros, Goiás, Brazil.A. Croton desertorum, C. didrichsenii and C. glandulosus. B. C. goyazensis, C. gracilescens and C. gracilipes. C. C. gracilirameus, C. grandivelus and C. hirtus. D. C. horminum, C. inaequilobus and C. leptobotryus.

FIGURE 10. A−C. Croton didrichsenii. A. Flowering branch. B. Staminate flower. C. Pistillate flower, showing the laciniate sepals. (A−C: M. J. Silva et al. 4134). D−H. C. glandulosus. D. Flowering branch. E. Detail of a leaf base showing the stipitate, acropetiolar glands. F. Staminate bract with two sessile glands. G. Staminate flower. H. Pistillate flower with one sepal removed. (D−H: M. J. Silva et al. 3886). I−M. C. goyazensis. I. Flowering branch. J. Detail of a leaf base showing the cylindrical acropetiolar glands. K. Detail of a leaf margin showing a narrow gland at the tip of a tooth. L. Staminate flower. M. Pistillate flower. (I−M: M. J. Silva et al. 5358).

FIGURE 11. A−D. Croton gracilescens. A. Flowering branch. B. Detail of the lower surface of a leaf showing a large cupular gland in a sinus of the leaf margin. C. Staminate flower. D. Pistillate flower. (A−D: M. J. Silva et al. 3057). E−H. C. gracilipes. E. Flowering branch. F. Detail of the lower surface of a leaf showing the saucer-shaped basilaminar glands. G. Staminate flower. H. Pistillate flower.(E−H: M. J. Silva et al. 4052). I−L. C. gracilirameus. I. Flowering branch. J. Detail of the lower side of a leaf base showing multiple glands at the apex of the petiole, along the leaf margin and on the surface of the blade. K. Staminate flower. L. Pistillate flower with one of the sepals removed; note the glandular sepals. (I−L: R. C. Sodré et al. 494).

FIGURE 12. A−D. Croton gracilescens. A. Habit. B. Inflorescence showing pistillate flowers. C. Inflorescence showing young capsules. D. Capsules. E−G. C. gracilipes. E. Habit. F. Flowering branch. G. Inflorescence showing young capsules and staminate flowers. H−J. C. gracilirameus. H. Habit. I. Flowering branch. J. Capsules. K−O. C. grandivelus. K. Habit. L. Inflorescence showing staminate flowers. M. Inflorescence showing pistillate flowers and staminate buds. N. Fruiting branch. O. Capsules.

FIGURE 13. A−C. Croton grandivelus. A. Flowering branch. B. Staminate flower. C. Pistillate flower; note the laciniate petals and bifurcate stigmas. (A−C: M. J. Silva et al. 4580). D−H. C. hirtus. D. Flowering branch. E. Detail of a leaf base showing the stipitate acropetiolar glands and long stiff trichomes. F. Staminate bract with stipitate glands. G. Staminate flower. H. Pistillate flower; note the one much smaller sepal. (D−H: R. C. Sodré et al. 535). I−K. C. horminum. I. Flowering branch. J. Staminate flower. K. Pistillate flower; note the filaments in the petal positions. (I−K: M. J. Silva et al. 4749).

FIGURE 14. A. Flowering branch of Croton hirtus. B−D. C. horminum. B. Flowering branch. C. Inflorescence showing young capsules and staminate buds. D. Inflorescence showing pistillate flowers. E−H. C. inaequilobus. E. Habit. F. Flowering branch. G. Petiole showing orange glands. H. Inflorescence showing young capsules and staminate flowers. I−K. C. longifolius. I. Flowering branch. J. Inflorescence showing staminate flowers and buds. K. Inflorescence showing pistillate flowers and staminate buds. L−N. C. megalocalyx. L. Habit. M. Flowering branch showing staminate flowers. N. Flowering branch showing pistillate flowers.

FIGURE 15. A−E. Croton inaequilobus. A. Flowering branch. B. Detail of a leaf base showing the sessile acropetiolar glands. C. Detail of the upper side of a leaf blade showing a shortly stipitate marginal gland in the sinus. D. Staminate flower. E. Pistillate flower, with a markedly smaller sepal. (A−E: M. J. Silva et al. 4777). F−J. C. leptobotryus. F. Flowering branch. G. Detail of a leaf base showing the conical acropetiolar glands. H. Staminate bud showing the inflexed stamens. I. Staminate flower. J. Pistillate flower with one sepal removed. (F−J: R. F.Vieira et al. 1380). K−M. C. longifolius. K. Flowering branch.L. Staminate flower.M. Pistillate flower, with vestigial petals. (K−M: M. J. Silva et al. 5677).

FIGURE 16. Map of geografic distribution of the Crotoneae species in the Chapada dos Veadeiros, Goiás, Brazil.A. Croton longifolius, C. megalocalyx andC. odontadenius. B.C. planaltoanus, C.sclerocalyx and C. siderophyllus. C. C.spica, C.subacutus and C.subferrugineus. D. C. subvillosus, C. urucurana and Croton sp.

FIGURE 17. A−E. C. megalocalyx. A. Flowering branch. B. Detail of the lower side of a leaf base showing the cylindrical basilaminar glands. C. Detail of a leaf margin showing a cylindrical gland. D. Staminate flower. E. Pistillate flower with one sepal removed; note the lanceolate petals. (A−E: M. J. Silva et al. 4807). F−J. C. odontadenius. F. Flowering branch. G. Detail of the lower side of an unequal leaf base showing the basilaminar glands. H. Detail of the upper side of a leaf base showing a marginal gland in the sinus. I. Staminate flower. J. Pistillate flower with two sepals removed (but six still remaining). (F−J: M. J. Silva et al. 4087). K−N. Croton planaltoanus. K. Flowering branch. L. Detail of the margin of the lower side of a leaf showing a cupular gland in the sinus. M. Staminate flower. N. Pistillate flower, with unequal sepals. (K−N: M. J. Silva & J. P. Santos 4577).

FIGURE 18. A−G. Croton odontadenius. A and B. Habit. C and D. Inflorescence showing staminate flowers. E. Inflorescence showing pistillate flowers and staminate buds. F and G. Capsules. H−K. C. planaltoanus. H. Habit. I. Inflorescence showing staminate flowers. J. Inflorescence showing pistillate flowers and staminate buds. K. Capsules. L and M. C. sclerocalyx. L. Habit. M. Inflorescence showing pistillate and staminate flowers. N−P.C. siderophyllus. N. Habit. O. Inflorescences showing pistillate and staminate flowers. P. Capsule.