Croton mentiens (S. Moore) Pax (1897: 211)

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 : 45-55

2.26. Croton sclerocalyx (Didr.) Müller Argoviensis (1865: 134) . — Podostachys sclerocalyx Didrichsen (1857: 134) .

Type:— BRAZIL. São Paulo: campis St. Carlos, January 1834, P. W. Lund s.n. (holotype: C 10011194!). ( Figs. 18 , L

and M; 19 , A–D)

Croton mentiens (S. Moore) Pax (1897: 211) , syn. nov. — Heterocroton mentiens Moore (1895: 461) . Type:— BRAZIL. Mato Grosso: Santa Cruz, 1891–1892, S. L. Moore s.n. (holotype BM 000939427!).

Subshrubs 0.4–1 m tall, erect, stems green-ferrugineous with simple or 2–5 radiate trichomes; latex clear. Stipules 4–6.5 × 0.7–1.2 mm, oblong to lanceolate, with ovoid or subulate glands at the base, persistent; petioles 0.5–1.8 cm long, with 2 sessile or shortly stipitate patelliform glands. Leaves alternate or opposite near to dichotomy of the branches, blades 4–8.7 × 2.2–5 cm, ovate, base obtuse, truncate or lightly cordate, apex acute, margin serrate or doubly serrate with sessile patelliform glands; upper surfaces with simple or 2 or 3 radiate trichomes and lower surfaces with stellate-2–9-radiate trichomes, membranaceous, venation semi-craspedodromous, with 4–6 pairs of secondary veins. Thyrses 1.8–4 cm long, terminal, staminate cymules with 2 or 3 flowers; inflorescence axis, both faces of sepals and bracts pistillate, dorsal surface of sepals and bracts staminate, ovary and styles with simple, 2–5 radiate or stellate-porrect trichomes; staminate bracts 1.1–2 × 0.5–1.2 mm, obovate, oblanceolate, elliptic or oblong, without glands; pistillate bracts 2.6–4.7 × 1.4–2.3 mm, ovate or lanceolate, without glands. Staminate flowers 3.8–4.2 mm long, pedicels 0.3–0.6 mm long; calyx 3–5-lobed, lobes 1.1–1.4 × 0.8–1.3 mm, united up to half, widely ovate, apex acute or obtuse; petals 3–5, 1.9–2.2 × 0.8–1.2 mm, obovate to oblanceolate, apex rounded, ventral surface villous basally; stamens 6–11, filaments glabrous. Pistillate flowers 4.9–6.2 mm long, pedicels 0.2–0.4 mm long; sepals 6 or 7, 3.5–5.2 × 1.7–3.2 mm, subequal, obovate, rarely oblong, apex acute, margins serrate, without glands; petals absent or 5, shaped gland or laminar, 1–1.3 × 0.2–0.4 mm, linear to lanceolate, disk 6-segmented; ovary 1.9–2.1 × 1.6–1.8 mm, oblongoid, styles 4-fid, pubescent. Capsules 5–5.5 mm long, globose or oblongoid, dark green, columella lacking prominent, smooth lobes; seeds 4–4.2 × 2.9–3.1 mm, oblongoid or ellipsoid, uniformly grayish.

Specimens examined:— BRAZIL. Goiás: Água Fria de Goiás, GO 118, ca. 10 km antes do município de São João d’Aliança, 27 October 2016, R. C. Sodré et al. 2681 ( UFG) ; Teresina de Goiás, GO 118 em direção a Alto Paraíso de Goiás, a 4 km de Teresina de Goiás, 05 April 2013, R. C. Sodré et al. 650 ( UFG) .

This species ocurrs in Brazil (DF, GO, MT, MG and SP) and Paraguay ( Caruzo & Cordeiro 2007). In was found in cerradão in the municipality of Água Fria de Goiás and Teresina de Goiás ( Fig. 16B ), flowering in October and fruiting in April. Its morphological relationship with C. odontadenius , has already been discussed in comments under that species.

This species shares the petioles with patelliform shortly stipitate glands, inflorescences with a space between the pistillate and staminate flowers, this latter shortly pedicellate and with a glabrous to glabrescent receptacle, as well as externally glabrous pistillate flowers with 6–8 ridges and serrate margins, as well as multifid styles. It belongs to C. section Geiseleria .

Taxonomic notes:— After studying the type collections from both species and their protologues, combined with the observation of morphological variation of the species in various localities in Goiás and Mato Grosso States, we consider C. mentiens to be a synonym of C. sclerocalyx , due the considerable overlap between their characters.

2.27. Croton siderophyllus Baillon (1864: 305) . Lectotype (designated by Medeiros et al. 2016):— BRAZIL. Minas Gerais: Serra da Caraça, June 1839, M. P. Claussen 110 (P 00634674!, isolectotypes: A 00277220!, G 00312522!, P 00634673!, SP 000666!). Syntypes:— BRAZIL. Minas Gerais: Serra da Caraça, 1841, M. P. Claussen 1573 (A 00277221!); without locality, no date, C. F. P. von Martius 1084 (NY 00262953!, M 0089083!, M 0089084!, G 00312520!). ( Figs. 18 , N–P; 19 , E–G)

Subshrubs 0.15–1.4 m tall, erect or subdecumbent, stems yellowish to brownish with sessile or stipitate stellate-porrect trichomes; latex clear. Stipules 0.3–1.5 × 0.1–0.3 mm, oblong or lanceolate, with ovoid glands at the base, persistent; petiole 0.3–1.8 cm long, without glands. Leaves alternate, blades 2.3–8.1 × 0.9–2.6 cm, elliptic, narrowly elliptic, lanceolate or oblong, base obtuse or discretely truncate, apex acute, margin entire or serrate without glands; upper and lower surfaces with subsessile stellate-porrect trichomes, membranaceous, venation brochidodromous, 4–6 pairs of secondary veins. Thyrses 1.2–3 cm long, terminal, staminate cymules with 1 or 2 flowers; inflorescence axis, dorsal surface of staminate sepals and bracts of both flowers, both faces of sepals pistillate, ovary and styles with stellate-porrect trichomes; staminate bracts 0.7–1.1 × 0.2–0.3 mm, ovate a lanceolate; pistillate bracts 1.7–1.8 × 0.2–0.3 mm, linear or narrowly lanceolate, both bracts with ovoid a ellipsoid glands. Staminate flowers 6–12 mm long, pedicel 2.5–8 mm long, sepals 5, 1.8–2.5 × 0.9–1.1 mm, ovate, apex acute, petals 5, 1.9–2.4 × 0.8–0.9 mm, oblong or narrowly elliptic, apex obtuse, ventral surface villous, stamens 10–11, filaments villous at the base. Pistillate flowers 6–7 mm long, pedicel 1–2 mm long, sepals 5, 2.8–5 × 1.2–2.5 mm, subequal, ovate to lanceolate, apex acute, margins entire, with ovoid or ellipsoid glands, petals absent, disk 5-lobed, ovary 1.5–2.2 mm long, globose, styles 2–4-fid, pubescent. Capsules 4–5 mm long, globose, brownish; seeds 3.2–3.7 × 2.3–2.5 mm, ellipsoid or widely ellipsoid.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, ca. 2 km N.W. of Veadeiros, road to Cavalcante , 14°S, 47°W, 1050 m, 22 December 1965, H. S. Irwin et al. 9485 ( UB) ; ca. 15 km W of Veadeiros , 14°S, 47°W, 1000 m, 8 February 1966, H. S. Irwin et al. 12352 ( UB) ; ca. 1 km W of Veadeiros , 14°S, 47°W, 900 m, 13 February 1966, H. S. Irwin et al. 12728 ( UB) ; 19 km N. of Alto Paraíso , ca. 1250 m, 20 March 1971, H. S. Irwin et al. 32794 ( UB) ; 21 km N. of Alto Paraíso , ca. 1250 m, 21 March 1971, H. S. Irwin et al. 32913 ( UB) ; 20 km by road N of Alto Paraíso , ca. 1600 m, March 1973, W. R. Anderson 6207 ( UB) ; 15, 8 km ao NE de Alto Paraíso de Goiás na rodovia para Teresina de Goiás, 14°04’S, 47°30’W, 5 March 1988, S. Ginzbarg & P. César 731 ( UB) ; a oeste do centro da cidade, 20 February 1991, D. Alvarenga et al. 723 ( CEN) ; 12 km de Alto Paraíso em direção a Colinas do Sul , 14°09’55”S, 47°35’58”W, 20 July 1994, M. Aparecida da Silva et al. 2175 ( IBGE) ; rodovia entre Alto Paraíso e Teresina de Goiás, 28 July 1994, M. Aparecida da Silva et al. 2108 ( IBGE) ; 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. 4069 ( UFG), 4071 ( UFG) ; próximo à GO 118, entre Alto Paraíso e a Serra do Pouso Alto , 14°03’14.8”S, 47°30’29.5”W, 1083 m, 09 March 2012, R. C. Sodré et al. 43 ( UFG), 44 ( UFG) ; ibd., imediações da estrada para o cruzeiro 13°58’13.7”S, 47°29’15.8”W, 1542 m, 21 January 2012, M. J. Silva et al. 4073 ( UFG), 4074 ( UFG), 4075 ( UFG), 4076 ( UFG), 4080 ( UFG) ; ibd., R. C. Sodré et al. 24 ( UFG), 25 ( UFG) ; Região da Serra de Pouso Alto, ca. 5 km ao norte do Morro do Japonês, 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 et al. 212 ( UFG), 213 ( UFG), 214 ( UFG), 215 ( UFG), 216 ( UFG), 221 ( UFG), 222 ( UFG), 223 ( UFG), 224 ( UFG), 225 ( UFG), 226 ( UFG), 227 ( UFG), 228 ( UFG), 229 ( UFG), 235 ( UFG) ; ibd., M. J. Silva et al. 4486 ( UFG), 4487 ( UFG) ; GO 118 , próximo à placa que indica a Serra do Pouso alto, entre Alto Paraíso de Goiás e a Serra , 14°02’44.8”S, 47°31’25.7”W, 1083 m, 09 March 2012, R. C. Sodré et al. 57 ( UFG) ; GO 239 , cerca de 20 Km após Alto Paraíso de Goiás em direção ao povoado de São Jorge, 28 September 2012, R. C. Sodré et al. 155 ( UFG) ; região da Serra de Pouso Alto, ca. 200 m à esquerda da GO 118, sentido Alto Paraíso de Goiás a Teresina de Goiás, 14°1’30.9”S, 47°31’56.9”W, 1150 m, 26 October 2012, R. C. Sodré et al. 244 ( UFG), 245 ( UFG) ; a esquerda da GO 118, sentido Alto Paraíso a Teresina de Goiás, no final da cerca que limita o parque na sua porção leste, 13°55’37”S, 47°25’56”W, 1168 m, 14 December 2012, R. C. Sodré et al. 383 ( UFG), 386 ( UFG) ; trilha em direção 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. 503 ( UFG), 504 ( UFG), 505 ( UFG), 506 ( UFG) ; imediações do posto Vale da Lua , 14°8’19.3”S, 47°31’23.8”W, 1241 m, 16 February 2013, R. C. Sodré et al. 546 ( UFG) ; imediações do alojamento do Parque , 14°9’26”S, 47°47’27.9”W, 1080 m, 13 December 2014, R. C. Sodré et al. 1528 ( UFG) ; cerca de 30 km a esquerda da GO 118 de São João d’Aliança à Alto Paraíso de Goiás), na estrada para catarata do rio dos Couros , 14°17’35”S, 47°42’32”W, 1028 m, 02 January 2015, R. C. Sodré et al. 1586 ( UFG) ; ca. 1 km após a casa dos pesquisadores, na continuidade das estrada que leva aos alojamentos do Parque , 14°8’27.4”S, 47°46’2.8”W, 1115 m, 04 January 2015, R. C. Sodré et al. 1629 ( UFG). Cavalcante, 20 km by Road S of Teresina, 1200 m, 18 March 1973, W. R. Anderson et al. 7433 ( UB) ; povoado Kalunga, em direção a cachoeira Santa Bárbara , 07 March 2003, J. F. B. Pastore & E. Suganuma 387 ( CEN) ; 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. 4073 ( UFG), 4075 ( UFG), 4076 ( UFG), 4080 ( UFG) ; ibd., 13°56’39.3”S, 47°29’52.8”W, 1084 m, 09 March 2012, R. C. Sodré et al. 49 ( UFG) ; estrada para o povoado Kalunga , Engenho II, 13°42’1.4”S, 47°27’50.3”W, 898 m, 05 April 2013, R. C. Sodré et al. 657 ( UFG), 661 ( UFG) ; a esquerda do km 204 da GO 118, de Alto Paraíso para 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 1227 ( UFG). Without municipality, Serra do Pouso Alto , 21 December 1968, Graziela, M. José & Ana Lima 817 ( UB) .

This species is endemic to Brazil and was reported by Lima & Pirani (2003) for Bahia, Goiás and Minas Gerais states. We found it in cerrado sensu stricto in the municipalities of Alto Paraíso de Goiás and Cavalcante ( Fig. 16B ), flowering and fruiting almost throughout the year.

Croton siderophyllus can be recognized by the cinereous-yellowish branches and leaves, elliptical to lanceolate leaf blades with entire to serrate margins, eglandular petioles, inflorescences up to 3 cm long with few flowers and pistillate sepals with ovoid or subgloboid and sessile glands. It belongs to section Barhamia subsection Medea .

2.28. Croton spica Baillon (1864: 362) . Type:— BRAZIL. Mato Grosso: without locality, 1831–1833, C. Gaudichaud-Beaupré 249 (holotype P 00634647!, isotype F V0056198!). ( Figs. 19 , H–K; 20 , A–C)

Subshrubs 0.4–0.9 m tall, erect, stems yellowish with subsessile or stipitate stellate-porrect trichomes; latex clear. Stipules 0.5–1.3 × 0.3–0.6 mm, oblong, with globose glands at the base, persistent; petioles 0.6–2.2 cm long, with 2 sessile patelliform glands, yellowish. Leaf blades 4.3–8.3 × 2.3–5.2 cm, oblong, elliptic or ovate, base discretely cordate or obtuse, apex obtuse or mucronate, margin crenate or serrate with sessile patelliform glands; upper and lower surfaces with subsessile stellate-porrect trichomes, membranaceous, venation brochidodromous, with 5–10 pairs of secondary veins, the basal pair of veins never surpassing half the length of the blade. Thyrses 4–7.3 cm long, terminal, staminate cymules with 1 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; bracts of both flowers 4–7 × 0.15–0.5 mm, linear, without glands. Staminate flowers 5–6.5 mm long, pedicel 1–2.2 mm long, sepals 5, 2.2–2.5 × 1.2–1.5 mm, ovate, apex acute, petals 5, 2.7–3.2 × 0.7–0.9 mm, lanceolate or oblong, apex obtuse or acute, both surface villous, stamens 11, filaments glabrous. Pistillate flowers 5.5–7.2 mm long, pedicels 0.9–1.6 mm long; sepals 5, 4.5–8 × 1.1–7.5 mm, unequal, obovate, ovate, widely elliptic or lanceolate, apex obtuse, acuminate or acute, margins serrate, without glands; petals 5, 0.2–3 × 0.1–0.2 mm, linear or lanceolate, disk 5-lobed; ovary 1.3–2 mm long, globose, styles 2-fid, 3.5–5 mm long, pubescent. Capsules 5–5.5 mm long, globose or oblongoid, canescent-green; seeds 4–4.3 × 2.6–2.8 mm, oblongoid or ovoid, brownish, maculate.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, cerca de 3,5 Km a noroeste do Morro do Buracão , 14°07’25.4”S, 47°44’12.1”W, 1189 m, 10 May 2013, R. C. Sodré et al. 719 ( UFG), 720 ( UFG), 721 ( UFG) ; próximo a bifurcação da estrada de terra que leva a Cachoeira das Sete Quedas , 14°07’25.4”S, 47°44’12.1”W, 1189 m, 14 December 2013, M. J. Silva et al. 5660 ( UFG), 5661 ( UFG), 5662 ( UFG), 5663 ( UFG), 5664 ( UFG) ; ibd., 1 km a partir da bifurcação que leva a Cachoeira das Sete Quedas , 14°07’30”S, 47°44’24.6”W, 1182 m, 14 December 2013, M. J. Silva et al. 5666 ( UFG), 5667 ( UFG) ; ibd. R. C. Sodré et al. 1069 ( UFG), 1070 ( UFG), 1071 ( UFG), 1072 ( UFG), 1074 ( UFG) ; ibd., 14°07’50.1”S, 47°44’38.3”W, 1182 m, R. C. Sodré et al. 1077 ( UFG) ; ibd., 2.5 km a noroeste do Morro do Buracão , 14°07’39.2”S, 47°44’11.6”W, 1191 m, 14 December 2013, M. J. Silva & R. C. Sodré 5676 ( UFG) .

According to Silva et al. (2015b), Croton spica occurs in the Federal District and in Goiás and Mato Grosso states, growing in cerrado sensu stricto. It was collected in the municipality of Alto Paraíso de Goiás ( Fig. 16C ), growing in cerrado sensu stricto, cerrado rupestre and campos. It flowers in December and May, and fruits in May. The morphological relationships of this species with C. abaitensis , which it most closely resembles, have already been mentioned in the comments under C. abaitensis . It belongs to C. section Geiseleria ( Silva et al. 2015b) .

2.29. Croton subacutus (Baill.) Müller Argoviensis (1873: 167) . — Croton campestris var. subacutus Baillon (1864: 317) . Lectotype (designated here):— BRAZIL. Minas Gerais: without locality, 1816–1821, A. F. C. P. Saint-Hilaire cat. B 1 1323 (P 00635486!, isolectotypes A 00257897!, P 00623088!). Syntypes:— BRAZIL. Minas Gerais: without locality, 1816–1821, A. F. C. P. Saint-Hilaire cat. B 1 1971 (A 00257896!, P 00623089!). ( Figs. 20 , D–G; 21 , A–C)

Subshrubs or shrubs 1–1.8 m tall, erect, stems yellowish or greenish with stipitate stellate-porrect trichomes; latex clear. Stipules 0.5–1 mm long, lanceolate, with globose glands; petiole 0.7–1.8 cm long, without glands. Leaves alternate or whorled near to dichotomy of the branches, blades 3.3–10 × 1.3–3.6 cm, lanceolate or less commonly oval-lanceolate, oblong or elliptic, base obtuse or sometimes lightly cordate, apex acute or acuminate, margin entire without glands; upper and lower surfaces with stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, with 7–15 pairs of secondary veins. Thyrses 1.8–10 cm long, terminal, staminate cymules with 3 flowers; inflorescence axis, both faces of sepals pistillate and of both flowers, dorsal surface of staminate sepals, ovary and styles with shortly stipitate stellate-porrect trichomes; bracts of both flowers 0.5–1.1 × 0.1–0.3 mm, linear to lanceolate, without glands. Staminate flowers 2.5–4 mm long, pedicels 0.7–1.5 mm long; sepals 5, 1.4–1.9 × 0.7–1.2 mm, ovate, elliptic or less commonly oval-lanceolate, apex acute or obtuse; petals 5, 1.4–2 × 0.4–0.8 mm, oblanceolate to elliptic, apex obtuse, ventral surface villous at the base; stamens (14)16(17), filaments glabrous or sometimes pubescent. Pistillate flowers 2.5–3.5 mm long, pedicels 0.1–0.5 mm long; sepals 5, 1.1–2.3 × 0.3–0.6 mm, lanceolate, oblong or linear, apex acute, margins entire without glands; petals 5, 0.1–0.3 mm, globose or sometimes linear, disk 5-segmented; ovary 1–1.7 mm long, globose, styles 2-fid, pubescent. Capsules 5–5.5 × 4–4.5 mm, elliptic-oblongoid, green-grayish, columella with three prominent, smooth lobes; seeds 3.7–3.8 × 2.2–2.3 mm, oblongoid, blackish, with clear spots.

Specimens examined:— BRAZIL. Goiás: Água Fria de Goiás, GO 118, km 82, de Água Fria para São João d’Aliança , 14°47’49.1”S, 47°46’22.1”W, 999 m, 01 August 2013, M. J. Silva et al. 5166 ( UFG) ; GO 118 , km 83, entre São Gabriel e São João d’Aliança , 14°47’48.87”S, 47°33’26.34”W, 1073 m, 01 August 2013, R. C. Sodré et al. 796 ( UFG), 797 ( UFG), 798 ( UFG), 799 ( UFG), 800 ( UFG) ; ibd., 05 September 2013, R. C. Sodré et al. 840 ( UFG), 843 ( UFG), 844 ( UFG) ; ibd., 03 November 2013, R. C. Sodré, P. H. B. Santos & M. J. Silva 1003 ( UFG), 1004 ( UFG), 1005 ( UFG). Alto Paraíso de Goiás, cerca de 2 km a nordeste da casa dos pesquisadores, 14°7’38.9”S, 47°46’30.2”W, 1057 m, 21 June 2013, R. C. Sodré et al. 786 ( UFG), 787 ( UFG), 788 ( UFG) ; trilha para a cachoeira das Sete Quedas , ca. 1 km da torre, 14°6’33.4”S, 47°44’55.2”W, 1157 m, 02 August 2013, R. C. Sodré et al. 812 ( UFG) ; estrada Alto Paraíso de Goiás a Campos Belos, km 41, 28 November 1976, G. J. Shepherd et al. 3739 ( IBGE). Cavalcante , região da cachoeira das Sete Quedas, 14°5’37.1”S, 47°44’56.9”W, 1065 m, 02 August 2013, R. C. Sodré et al. 815 ( UFG) ; cerca de 1 km da GO 118, km 195, entre Alto Paraíso de Goiás e Teresina de Goiás, 13°55’40”S, 47°26’55”W, 06 September 2013, R. C. Sodré et al. 848 ( UFG), 849 ( UFG) ; estrada que leva ao povoado Kalunga, Engenho II, 05 April 2013, R. C. Sodré et al. 665 ( UFG), 666 ( UFG). Without municipality, ca. 65 km due North of Brasilia , 22 December 1968, R. M. Harley et al. 11466 ( UB) ; 20 km by Road S of Teresina, 18 March 1973, W.R. Anderson et al. 7438 ( UB) ; without locality, G. Barroso et al. 595 ( UB) .

This species was recorded for Minas Gerais ( BFG 2015) and now in Goiás state. In Chapada dos Veadeiros it was found in the municipalities of Alto Paraíso de Goiás, Cavalcante and São João d’Aliança in campo sujo, cerrado sensu stricto near rocks ( Fig. 16C ), with flowers and fruits the whole year.

Croton subferrugineus resembles C. subacutus by the leaves with entire margin, staminate flowers and small bracts, pistillate flowers with vestigial and usually subgloboid petals and columella with trilobed apex. However, C. subacutus can be recognized by the petiole without glands (vs. petiole with glands in C. subferrugineus ), and staminate flowers with 16 stamens (vs. 11). It belongs to section Adenophylli .

2.30. Croton subferrugineus Müller Argoviensis (1865: 135) . Lectotype (designated here):— BRAZIL. Goiás?: Ad Fazendam Almas, no date, J. B. E. Pohl 1622 (W 0004092!, isolectotypes B†, K 000186090!, K 000186089!, G 00434678!, W 0004091!, W 0004093!). ( Figs. 20 , H and I; 21 , D–G; 22A )

Subshrubs 0.3–0.9 m tall, erect, stems ferruginous with shortly stipitate stellate-porrect trichomes; latex clear, orange after oxidation. Stipules inconspicuous, with globose glands; petiole 0.2–0.7 cm long, with 2 sessile patelliform glands. Leaves alternate, blades 2.5–5.5 × 1.1–2 cm, ovate, oval-lanceolate or elliptic, base acute or obtuse, apex acute, margin entire without glands; upper surfaces with sessile stellate-porrect trichomes and lower surfaces with stipitate stellate-porrect trichomes, membranaceous, venation brochidodromous, 4–8 pairs of secondary veins. Thyrses 1–2.5 cm long, terminal, staminate cymules with 2–3 flowers; inflorescence axis, dorsal surface of bracts and sepals of both flowers, ovary and styles with sessile stellate-porrect trichomes; bracts of both flowers 0.3–1 × 0.2–0.4 mm, triangular or oval-lanceolate, with globose glands at the base. Staminate flowers 2.6–3.8 mm long, pedicels 0.3–1 mm long; sepals 5, 1.1–1.7 × 0.7–1.1 mm, ovate, apex acute or obtuse; petals 5, 1.5–1.9 × 0.4–0.5 mm, oblanceolate or narrowly oblanceolate, apex obtuse, glabrescent on both surfaces, ciliate at the base; stamens 11, filaments glabrous. Pistillate flowers 3–4.5 mm long, pedicels 0.5–0.9 mm long; sepals 5, 0.9–1.8 × 0.4–0.7 mm, ovate or oval-lanceolate, apex acute, margins entire without glands; petals 5, globose, disk 5-lobed; ovary 0.9–1.2 mm long, globose, styles 2-fid, pubescent. Capsules 5–5.5 mm long, oblongoid, green-ferruginous, columella with three prominent, smooth lobes; seeds 4.2–4.5 × 2.4–2.7 mm, ellipsoid, uniformly grayish.

Specimens examined:— BRAZIL. Goiás: Água Fria de Goiás, a direita do km 61 da GO 118, sentido São Gabriel a São João d’Aliança , cerca de 30 km deste último, 09 May 2013, R. C. Sodré & M. J. Silva 711 ( UFG), 715 ( UFG), 717 ( UFG) ; GO 118 , Km 83, entre São Gabriel e São João d’Aliança , 14°47’48.87”S, 47°33’26.34”W, 1073 m, 01 August 2013, R. C. Sodré et al. 801 ( UFG), 802 ( UFG), 803 ( UFG), 804 ( UFG), 805 ( UFG), 806 ( UFG) ; ibd., 05 September 2013, R. C. Sodré et al. 841 ( 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 et al. 1316 ( UFG), 1317. Alto Paraíso de Goiás, without locality, 21 March 1969, H. S. Irwin et al. 24797 ( UB) ; GO 118 , Km 156, entre São João d’Aliança e Alto Paraíso de Goiás, 14°12’49.4”S, 47°29’15.9”W, 1182 m, 22 June 2013, R. C. Sodré et al. 793 ( UFG), 794 ( UFG) ; a esquerda do km 131 da GO 118, entre São João d’Aliança e Alto Paraíso de Goiás, 14°25’20.7”S, 47°30’35”W, 1000 m, 20 February 2014, R. C. Sodré et al. 1209 ( UFG), 1212 ( UFG) ; ca. 2 km de Alto Paraíso de Goiás em direção a São João d’Aliança, 21 June 2014, R. C. Sodré et al. 1310 ( UFG). Cavalcante , 5 km antes de Araí, 13°32’47”S, 47°35’31”W, 1120 m, 13 April 2004, R. C. Mendonça et al. 5520 ( IBGE) ; a esquerda da GO 118, de Alto Paraíso para Teresina de Goiás, no final da cerca a leste do PNCV, 13°55’37”S, 47°25’56”W, 1168 m, 14 December 2012, R. C. Sodré et al. 382 ( UFG), 384 ( UFG), 387 ( UFG), 388 ( UFG) ; ibd., M. J. Silva et al. 4618 ( UFG), 4619 ( UFG), 4622 ( UFG), 4623 ( UFG), 4626 ( UFG), 4627 ( UFG) ; antes do córrego Quebra Coco em direção a Comunidade dos Kalungas , 13°36’27.5”S, 47°28’24.2”W, 1066 m, 07 March 2013, M. J. Silva et al. 4756 ( UFG), 4757 ( UFG), 4758 ( UFG) ; estrada que leva ao povoado Kalunga , Engenho II, 13°45’49”S, 47°27’18.9”W, 1066 m, 07 March 2013, R. C. Sodré et al. 573 ( UFG), 574 ( UFG), 575 ( UFG) ; próximo a reserva da fazenda Gavião , a sudoeste do PNCV, 14°8’S, 47°53’W, 01 November 2013, R. C. Sodré et al. 940 ( UFG), 941 ( UFG) ; a esquerda do km 204 da GO 118, sentido Alto Paraíso a Teresina de Goiás, 13°54’16”S, 47°22’12.9”W, 1299 m, 21 February 2014, R. C. Sodré et al. 1224 ( UFG) ; ibd., 13°54’22”S, 47°22’11”W, 1267 m, 24 July 2014, R. C. Sodré et al. 1324 ( UFG) ; em direção ao Engenho II, região dos Kalungas, ca 18 km após Cavalcante , 13°39’29.4”S, 47°28’31.4”W, 1108 m, 25 July 2014, R. C. Sodré et al. 1327 ( UFG), 1334 ( UFG), 1337 ( UFG), 1338 ( UFG) .

Lima & Pirani (2003) cited this species for Bahia and Minas Gerais States, and we confirm its occurrence here in Goiás, where its occurrence was doubtful according to Müller Argoviensis (1873). It grows in campo sujo and cerrado sensu stricto in the municipalities of Alto Paraíso de Goiás and Cavalcante ( Fig. 16C ), flowering and fruiting during most of the year. Croton subferrugineus can be confused with C. subacutus , except by the characters already noted in the comments of that species. Croton subferrugineus belongs to section Adenophylli .

2.31. Croton subvillosus Müller Argoviensis (1873: 168) . Lectotype (designated here):— BRAZIL. Goiás: prope Gamelleira, no date, J. B. E. Pohl 1617 (Catalog Number: 2164) (W 0051217!, isolectotypes G 00434673!, M 0089120!, K 000186135!, K 000186136!). ( Figs. 21 , H–J; 22 , B–D)

Subshrubs 0.05–0.17 m tall, erect, stems greenish-ferruginous with sessile or shortly stipitate stellate-porrect trichomes; latex clear. Stipules inconspicuous, with ovoid to subulate glands; petioles 4–6 mm long, without glands. Leaves alternate, blades 4.5–10 × 2.2–4.3 cm, elliptic, obovate, or oboval-oblanceolate, base acute, apex obtuse or rounded, margin entire without glands; upper surfaces dark green and glossy, glabrous or with shortly stipitate stellate-porrect trichomes and lower surfaces always pubescent with the same trichomes, membranaceous, venation brochidodromous, with 6–9 pairs of secondary veins. Thyrses 2–11 cm long, terminal, staminate cymules with 2 flowers; inflorescence axis, dorsal surface of bracts and sepals of both flowers, ovary and styles with subsessile stellate-porrect trichomes; bracts of both flowers 1–2.7 × 0.6–0.8 mm, lanceolate to triangular, without glands. Staminate flowers 5–8 mm long, pedicels 2.3–4.8 mm long; sepals 5, 2.1–3.2 × 1.2–1.9 mm, ovate, apex acute or obtuse; petals 5, 2.5–3.3 × 1–1.5 mm, oblanceolate or elliptic, apex obtuse or sometimes acute, ventral surface villous at the base; stamens (15)16(–18), filaments glabrous. Pistillate flowers 3.5–7.5 mm long, pedicels 0.7–2.8 mm long; sepals 5, 2.3–2.9 × 0.5–1.1 mm, lanceolate, apex acute, margins entire without glands; petals 5, 1.2–2 × 0.1–0.2 mm, linear, apex acute, globose, disk entire; ovary 1.6–2 mm long, globose, styles 2-fid, patent, pubescent. Capsules 5.5–6.5 × 6–6.5 mm, globose, greenish, pedicels 3–4 mm long, columella with three prominent, smooth lobes; seeds 4–4.2 × 2.2–2.4 mm, widely ellipsoid, uniformly dark brown.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, PNCV, campo limpo, 1.8 km a partir do rio Preto em direção ao Morro Peito de Moça , 03 August 2013, M. J. Silva et al. 5230 ( UFG) ; cerca de 5 km do morro da Baleia, na estrada que leva ao rio Preto, 13 July 2013, M. J. Silva, A. O. Souza & P. H. B. Santos 5122 ( UFG), 5124 ( UFG), 5128 ( UFG), 5129 ( UFG). Cavalcante , PNCV, ca. 23 km a norte de Alto Paraíso de Goiás, cruzeiro, 13°56’39”S, 47°29’38”W, 1410 m, 12 November 1996, M. L. Fonseca, A. J. Santos & F. C. A. Oliveira 1315 ( IBGE) ; ibd., M. Aparecida da Silva & A. J. V. dos Santos 3234 ( IBGE) ; RPPN Serra do Tombador , 13°38’4”S, 47°49’6”W, 857 m, 29 October 2011, A. R. O. Ribeiro et al. 283 ( UB) .

This species was recorded for Goiás and Tocantins states ( BFG 2015). It was collected in campo sujo in the municipalities of Alto Paraíso de Goiás and Cavalcante ( Fig. 16D ), flowering from July to November, and fruiting in November. It resembles C. campestris morphologically as already discussed. Croton subvillosus belongs to section Adenophylli .

2.32. Croton urucurana Baillon (1864: 335) . Lectotype (first step designated by Smith et al. 1988, second step designated here):— BRAZIL. Without locality, 1816–1821, A. F. C. P. Saint-Hilaire s.n. (P 00634745!; isolectotype: P 00634744!). Syntypes:— BRAZIL. Rio de Janeiro or São Paulo, without locality, 1842, Pissis 9 (P 00634748!, P 00634749!, SP 000668!); Minas Gerais: Sabara, 1844, M. Weddell 1353 (P 00634750!, P 00634751!, P 00634752!); Minas Gerais: without locality, 1838, M. P. Claussen 770 (P 00634746!, P 00634747!, G 00311423!, SP 000667!). ( Figs. 22 , E–G; 23 , A–E)

Trees 3–8 m tall, erect, stems canescent with sessile or stipitate dendritic trichomes; latex clear to reddish. Stipules 7–12 × 2–5 mm, lanceolate, without glands, persistent; petioles 7.2–17 cm long, with 2 patelliform glands. Leaves alternate, blades 13.2–24.5 × 7.6–18 cm, ovate, base cordate, apex acuminate, margin entire without glands; upper and lower surfaces with stellate-porrect trichomes, membranaceous, venation actinodromous-brochidodromous, with 6–8 pairs of secondary veins. Thyrses 15–42 cm long, terminal, bisexual cymules, at the base, and staminate cymules with 4–8 staminate flowers; inflorescence axis, dorsal surface of bracts and sepals of both flowers, ovary and styles with stellate-porrect trichomes; bracts of both flowers 1.7–3 × 0.8–2.3 mm, ovate to lanceolate, without glands. Staminate flowers 6.5–8.5 mm long, pedicels 2–4 mm long; sepals 5, 2.5–3 × 1.2–1.9 mm, ovate, apex acute; petals 5, 2.6–3.2 × 0.6–1.2 mm, oblanceolate or oblong, apex obtuse, villous on both surfaces; stamens 15–17, filaments glabrous or glabrescent. Pistillate flowers 7–8.5 mm long, pedicels 1.3–2 mm long; sepals 5, 3–4 × 1.5–1.9 mm, subequal, oblong to ovate, apex acute or obtuse, margins entire, without glands; petals 5, 0.8–2 × 0.1–0.2 mm, linear, disk 5-lobed; ovary 1.3–1.8 mm long, globose, styles 2-fid, glabrous. Capsules 4.5–5 mm long, globose, brownish; seeds 3.3–3.8 × 2.5–2.7 mm, ellipsoid or widely ellipsoid, uniformly grayish.

Specimens examined:— BRAZIL. Goiás: Alto Paraíso de Goiás, ca. 1,5 Km W of Veadeiros, 1000 m, 12 February 1966, H. S. Irwin et al. 12670 ( MO, NY, UB) ; Portal da Chapada , 10 December 2011, L. H. Soares-Silva 1570 ( INPA, UB) ; ibd., imediações do Morro da Baleia, próximo ao Bar do Valdomiro, 14°8,439’S, 47°39,640’W, 1155 m, 28 September 2012, R. C. Sodré 156 ( UFG), 157 ( UFG). Cavalcante , Barra do rio Santo Antônio , próximo à futura balsa, 13°31’35”S, 48°2’57”W, 320 m, 20 September 2001, G. Pereira-Silva et al. 5416 ( HUEFS, CEN, SP) ; without locality, 13°34’55”S, 47°28’15”W, 10 November 2008, N. P. Massarotto et al. 98 ( UB) ; estrada que leva ao Engenho II, região dos Kalungas, 07 March 2013, R. C. Sodré et al. 580 ( UFG). Colinas do Sul, fazenda Saracura, estrada de manutenção das torres da nova linha de transmissão Minaçu / Niquelândia , 14°0’S, 48°12’W, 450 m, 08 September 1995, B. M. T. Walter et al. 2597 ( CEN, SP). Teresina de Goiás, 4 km by road S of Teresina, 1000 m, 12 February 1973, W. R. Anderson et al. 7378 ( NY, UB) .

Croton urucurana occurs in Argentina, Bolivia, Brazil (AC, AL, AM, BA, DF, ES, GO, MA, MG, MS, MT, PA, PR, RJ, RS SC, SP, TO), Paraguay and Uruguay ( Guimarães & Secco 2010). We found it in gallery forests in the municipalities of Alto Paraíso de Goiás, Cavalcante, Colinas do Sul and Teresina de Goiás ( Fig. 16D ), flowering and fruiting from September to March. This species belongs to C. section Cyclostigma .

Figures

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 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.

FIGURE 19. A−D. Croton sclerocalyx. A. Flowering branch. B. Detail of the lower side of a leaf base showing the saucer-shaped basilaminar glands. C. Staminate flower. D. Pistillate flower with two sepals removed. (A−D: R. C. Sodré et al. 650). E−G. Croton siderophyllus. E. Flowering branch. F. Staminate flower. G. Pistillate flower. (E−G: R. C. Sodré et al. 103). H−K. C. spica. H. Flowering branch. I. Detail of the lower side of a leaf base showing the acropetiolar glands. J. Staminate flower. K. Pistillate flower. (H−J: M. J. Silva et al. 5660; K: R. C. Sodré et al. 1069).

FIGURE 20. A−C. Croton spica. A. Habit. B. Inflorescence showing pistillate flowers and staminate buds. C. Inflorescence showing pistillate flowers and staminate flowers and buds. D−G. C. subacutus. D. Habit. E. Flowering branch. F and G. Inflorescence showing young capsules and staminate flowers. H and I. C. subferrugineus. H. Habit. I. Flowering branch.

FIGURE 21. A−C. Croton subacutus. A. Flowering branch. B. Staminate flower. C. Pistillate flower. (A−C: R. C. Sodré et al. 665). D−G. Croton subferrugineus. D. Flowering branch. E. Detail of a leaf base showing the acropetiolar glands. F. Staminate flower. G. Pistillate flower. (D−G: R. C. Sodré et al. 382). H−J. C. subvillosus. H. Flowering branch. I. Staminate flower. J. Pistillate flower. (H−J: M. L. Fonseca 1315).

FIGURE 22. A. Croton subferrugineus. Inflorescence showing capsules and staminate flowers. B−D. C. subvillosus. B. Habit. C. Inflorescence showing staminate flowers. D. Inflorescence showing pistillate flowers. E−G. C. urucurana. E. Flowering branch. F. Inflorescence showing staminate flowers and buds. G. Inflorescence showing pistillate flowers.

FIGURE 23. A−E. Croton urucurana. A. Flowering branch. B. Detail of the lower side of a leaf base showing the basilaminar glands. C. Staminate flower. D. Pistillate flower. E. Capsule. (A−E: R. C. Sodré et al. 580). F−I. Croton sp. F. Flowering branch. G. Detail of a leaf base showing the acropetiolar gland. H. Staminate flower. I. Pistillate flower. (F−I: R. C. Sodré, M. J. Silva & L.L.C. Antunes 938).