Research Article |
Corresponding author: Marcos José da Silva ( marcos_agrorural@hotmail.com ) Academic editor: Geoffrey Levin
© 2017 Marcos José da Silva, Laís de Souza Inocencio, Rodolfo Carneiro Sodré, Alexandre Antonio Alonso.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Silva MJ, Inocencio LS, Sodré RC, Alonso AA (2017) Morphological and anatomical evidence support a new wild cassava: Manihot fallax (Crotonoideae, Euphorbiaceae), from Mato Grosso, Brazil. PhytoKeys 91: 139-156. https://doi.org/10.3897/phytokeys.91.21465
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During the preparation of the taxonomic treatment of Manihot in the Midwest Region of Brazil, a new species was found. Manihot fallax M.J. Silva & L.S. Inocencio is described, illustrated and morphologically compared with similar simple-leaved species. The conservation status, geographic distribution (including map), ecology, phenology and notes about leaf anatomy of the new species are given. The synonymisation of M. robusta M. Mend. & T. B. Cavalc. under M. attenuata Müll. Arg. and lectotypes for M. attenuata and M. brachystachys Pax & K. Hoffm are also proposed. An emended description of M. attenuata is proposed as the original description is incomplete as it lacks information on the pistillate flowers, fruits and seeds.
Central Brazil, leaf anatomy, nomenclature, typification, Manihoteae
The taxonomy of Manihot was studied by
Despite the revision of Manihot (
Considering the species of Manihot present in the Midwest Region of Brazil, the knowledge about the genus is scarce mainly in the states of Mato Grosso and Mato Grosso do Sul, where
During the preparation of the taxonomic treatment of Manihot in the Midwest Region of Brazil, some collections were found from Serra Azul, Serra do Roncador, Serra do Taquaral and neighbouring areas in the state of Mato Grosso, which could not be assigned to any known species. One of these species, with entire and unlobed leaves and habit slender and virgate, is hereby illustrated, described as new and designated as M. fallax. The new species is compared with M. attenuata Müll. Arg. and M. weddelliana Baill., the taxa most morphologically similar to it and its phenology, conservation status and geographic distribution are also presented. The leaf anatomy of the new species and the species most morphologicaly similar to it were also compared because leaf anatomy constitutes an important tool for delimiting taxa in Manihot (e.g.
The morphological description of the new species is based on field observations, conducted by the authors during expeditions to the state of Mato Grosso and on morphological analyses of 34 collections from the authors and 10 collections from herbaria (UFMT/ICLMA and IAC). The emended description of Manihot attenuata and the nomenclatural revision for all taxa associated with it, resulted from analysis of all protologues, type and historical collections (17 exsiccatae) of the same and also of extensive field work in their areas of occurrence, as well as analyses of 30 collections from herbaria (BR, CEN, F, G, HRCB, HUEFS, IBGE, K, MG, MO, NY, P, RB, S, SP, UB and UFG). The terminology used in the description of both species is based on specific literature such as
The conservation assessment of both the species was based on field observations and applying the IUCN Red List Categories and Criteria (
Leaves of the type collection of the new species (i.e. L. S. Inocencio, A. O. Souza, L. L. C. Antunes and C. C. Oliveira 293) and M. attenuata (M. J. Silva 4063/UFG) with fully expanded leaf blades were fixed in FAA (formaldehyde/glacial acetic acid/ethanol 50% at 1:1:18 v/v/v) for 48h and transferred to 70% ethanol (
BRAZIL. Mato Grosso: Município de Barra do Garça, BR-158, na altura do km 726, margem esquerda da estrada no sentido Nova Xavantina, campo cerrado em encosta, 15°17'9.7"S, 52°11'15.5"W, 385 m a.s.l., 26 Jan 2014, fl., L. S. Inocencio, A. O. Souza, L. L. C. Antunes, and C. C. Oliveira 293 (holotype: UFG; isotypes: K, MO, NY).
Shrubs slender and virgate, up to 1.9 m tall; leaves light green, entire, unlobed, with secondary veins perpendicular to the midvein. Stipules 1–3 × 0.2–0.3 mm, setaceous, entire, early caducous; racemes 2.5–6 cm long, solitary, subspicate, staminate buds obovoid, stamens with filaments pubescent, pistillate calyx dialisepalous and fruits light green, without wings.
Manihot fallax. A Habit B Staminate bracts C Staminate bracteole D Staminate bud E Staminate flower F Staminate flower with calyx split and open G Inner surface of the staminate calyx showing the trichomes H Pistillate bracts I Pistillate bracteole J Pistillate flower K Inner surface of the pistillate calyx showing the trichomes L Fruit M Seed, ventral side N Seed, dorsal side. Drawn by Cristiano Gualberto from the holotype.
Shrubs 1–1.9 m tall, slender and virgate, monoecious, glabrous. Stem solid in cross section, greenish to reddish, glossy and waxy in young parts; latex clear, abundant; roots fibrous. Stipules 1–3 × 0.2–0.3 mm, setaceous, margins entire, early caducous; petiole 0.3–0.7 mm long, angulate, greenish. Leaves alternate spiral, sparsely distributed along the branches, but more concentrated near apex, blades 8–14 × 1–2 cm, lanceolate, narrowly elliptic or linear, entire, unlobed, non-peltate, base obtuse, apex obtuse with a short mucro, firmly membranaceous to chartaceous, glabrous on both surfaces, adaxial surface dark green, abaxial surface glaucous, the latter with a smooth wax pattern, venation camptodromous-brochidodromous, primary and secondary veins impressed on both surfaces, the secondary veins perpendicularly arranged in relation to the midvein, all of them pinkish to yellowish. Racemes 2.5–6 cm long, staminate or bisexual, solitary, terminal or arising from dichotomy of the branches, glabrous, angulate, glaucous to cinereous, waxy. Staminate flowers: buds 5.8–6 × 3.9–4 mm, obovoid, yellowish-green, without purplish pigmentation; bracts 6–6.1 × 3.9–4 mm, widely elliptic, foliaceous, apex acuminate, persistent; bracteoles situated along the lower third up to half of the pedicel, 4–4.1 × 0.4–0.5 mm, lanceolate, foliaceous, persistent, subopposite; pedicels 0.5–0.7 mm long, cylindrical, glabrous, light green; calyx 13.2–14 × 5.9–6 mm, campanulate, yellowish, without purplish pigmentation, shortly tomentose internally, lobes widely triangular, ovate to oblong, apex obtuse, base truncate; stamens 10, in two whorls of five, filaments pubescent near apex, the longer 12.5–12.6 mm long, the shorter 8.4–8.5 mm long, both thickened; anthers 5–7 mm long, oblong, dorsifixed, bright yellow; disc 10-lobed, intrastaminal, dark yellow. Pistillate flowers: buds 6–9 × 4–6 mm, ovoid, green-yellowish, without purplish pigmentation; bracts 6.4–7 × 2.9–3 mm long, widely ovate, margin entire, glabrous; bracteoles 3.7–3.8 × 0.8–0.9 mm, lanceolate, foliaceous, persistent, margin entire, apex acuminate, opposite along the lower third of the pedicel, glabrous; pedicels 4–6 mm long, cylindrical-clavate, glabrous, green; sepals 10–12 × 5–7 mm, lanceolate, apex acute, shortly tomentose externally, yellowish, without purplish pigmentation; ovary 4–6 × 3–4 mm, globose to ovoid, glabrous, green, disc patelliform, lobed, yellow; styles 3, shortly united at the base, stigmatic surface 2–3 mm long, densely papillose. Capsules 0.9–1.5 × 0.8–1.2 cm, oblong, light green, smooth, glabrous, without wings, dehiscence septicidal and loculicidal; columella (carpophore) persistent, 0.9 × 2–2.2 mm (width at narrowest point in middle), narrowly alate. Seeds 0.7–1 × 0.39–0.4 cm, oblong-ellipsoid, dark grey, with black spots; caruncle triangular, prominent, apex bilobed, cream to yellowish.
The species has been collected with flowers and fruits from October to March.
The specific Latin epithet “fallax” refers to deceptive and was chosen due to the false similarity of the new species to M. attenuata and M. weddelliana.
According to the
Brazil. Mato Grosso. Barra do Garças, Serra Azul, trilha as usina, próximo a sede da Moçonaria, 17 Mar 1990, fl., fr., L. Santos s.n. (UFMT/ICLMA 00856); próxima da cidade, nas imediações da antena do Cristo, 09 Mar 1991, fl., fr., L. Santos s.n. (UFMT/ICLMA 00866); Serra do Taquaral, Cerrado, 23 Nov 1997, fl., fr., L. C. Bernacci & G. Arborcz 2525 (UFMT, IAC); cerrado do Marcello, 15°50'50.9"S, 52°15'54.7"W, 14 Nov 2005, fl., fr., I. Faria & M. L. Mayer 339 (UFMT/ICLMA); gruta do pezinho, 15°50'29.6"S, 52°16'50"W, 28 Dec 2005, fl., I. Faria, M. L. Mayer & Moisés 517 (UFMT/ICLMA); Parque Estadual da Serra Azul, 15°51'S, 52°16'W, 21 Oct 2005, fl., M. Sanchez 2216 (UFMT/ICLMA); ib., 21 Feb 2006, fl., fr., I. Faria, M. L. Mayer & M. S. M. Ramos 787 (UFMT/ICLMA); ib., 15°52'S, 52°16'W, 04 Oct 2006, fl., R. Freitas & M. L. Mayer 145 (UFMT/ICLMA); ib., 13 Feb 2007, fl., R. Freitas & M. L. Mayer 404 (UFMT/ICLMA); ib., 1 Mar 2007, fl., fr., R. Freitas & M. L. Mayer 445 (UFMT/ICLMA); BR 158 na altura do km 726, margem esquerda, sentido Nova Xavantina, adjacências da Serra do Roncador, 15°17'21.8"S, 52°11'38.7"W, 389 m a.s.l., 28 Jan 2014, L. S. Inocencio, A. O. Souza & G. H. Silva 68, 69, 70, 71, 72 and 73 (UFG); ib., 15°17'9.7"S, 52°11'15.5"W, 385 m a.s.l., 26 Jan 2015, fl. fr., L. S. Inocencio, A. O. Souza, L. L. C. Antunes & C. C. Oliveira, 290, 291, 292 and 294 (UFG); BR 158, na altura do km 726, margem esquerda, sentido Nova Xavantina, 15°17'9.8"S, 52°11'22.3"W, 422 m a.s.l., 1 Feb 2015, fl. fr., L. S. Inocencio, A. O. Souza, L. L. C. Antunes & C. C. Oliveira 363, 364, 365, 366, 367, 368, 369, 370, 371, 372, 373, 374, 375, 376, 377, 378, 379, 380, 381, 382, 383, 384, 385, 386, 387 and 388 (UFG); Serra do Roncador, ca. 60 km a norte da cidade pela BR-158 e 5 km a esquerda desta rodovia em direção a Xavantina a partir do povoado Vale dos Sonhos, 52°13'54"S, 15°20'54"W, 642 m a.s.l., 24 Mar 2016, R. C. Sodré, T. P. Mendes & J. A. Oliveira 2174 (UFG).
Manihot fallax stands out from the other species of the genus with entire and unlobed leaves (
Regarding leaf anatomical features, M. fallax differs from M. attenuata in having the vascular cylinder with arch-shaped collateral vascular bundles surrounded by pericyclic fibres in the midvein (Fig.
Leaf anatomy of species. A–J Cross section of the leaf blade. Manihot fallax (A–C): A Median portion of the midvein B Median portion of the leaf blade; abaxial epidermis papilose and stomata indicated by arrow C Edge. Manihot attenuata (D–J): D Median portion of the midvein E Median portion of the leaf blade; note stomata (arrow) and epidermal idioblast with druse (arrowhead) F Detail of epidermal idioblast with druse (arrowhead) G Laticifers in the midvein phloem (arrowheads) H Detail of the epidermis and collenchyma in the midvein I Vascular bundles with parenchymatical sheath in the median portion of the leaf blade (arrowheads correspond to laticiferous) J Vascular bundles in the median portion of the leaf blade (arrowheads correspond to laticifers) K–M Dissociated epidermis of both species. Manihot fallax. K. Paracytic stomata. Manihot attenuata (L–M): L Stomata distribution in the abaxial surface of the leaf blade M Stomata distribution in the adaxial surface of the leaf blade lateral to midvein N and O Schematic representation of the petiole in cross section N Manihot fallax O Manihot attenuata. CT: cortex; VC: vascular cylinder; co: collenchyma; ep: epidermis; pp: palisade parenchyma; sp: spongy parenchyma; pa: medullar parenchymatic.
Both M. fallax and M. attenuata have midveins with angular collenchyma on both surfaces, ground parenchyma surrounding the vascular cylinder (Fig.
Amongst the anatomical features that differ between M. fallax and M. attenuata, the presence the fibres in the vascular cylinder in the midvein, the papillae in the epidermal cells and the arrangement and number of collateral bundles in the petiole have been cited by
Manihot brachystachys Pax & K. Hoffm., Planzenr. (Engler) 4, Fam. 147, II: 97. 1910. Type. Brazil. Südbrasilianishe Provinz, Staat Goiaz, 8 Jan 1898, fl., Glaziou 22126 (lectotype, designated here: P [P04786133!]; isolectotypes: BR [BR000005101320!], G [G00441911!], K [000600417!], P [P04786134!, P04786135!], S [S-R-9076!]).
Manihot robusta M. Mend. & T. B. Cavalc. Arnaldoa 22(2): 297, 2015. Type. BRAZIL. Alto Paraíso de Goiás, Parque Nacional Chapada dos Veadeiros, ca. 0.3 km da GO-239, sentido sede alojamento do ICMBio (lado direito), 14°09'55.92"S, 47°47'25.62"W, 1046 m a.s.l., 31 Oct 2014, M. Mendoza, J.B.A. Bringel, A.A. Santos and T. Reis 4343, syn. nov. (holotype: CEN!; isotypes: HRCB!, HUEFS!, K!, MG, MO!, NY!, RB!, SP!, UB!).
BRAZIL. Goiás: inter Goyaz et Cavalcante, Burchell 7865 (lectotype, designated here: K [K000600418!]; isolectotypes: BR [BR510866!], G!).
Shrubs 0.5–1.2 m tall erect or decumbent, in which case the stem to 0.6 m tall with lateral branches to 1.3 m long, monoecious, glabrous. Stems and branches angulose, glossy, brownish, the bark exfoliating when adult and purplish, greenish or combinations of these when young; latex yellow or clear. Stipules 10–12 × 0.5–0.1 mm, lanceolate, sparsely serreate, persistent; petiole 3–4 mm long, greenish. Leaves alternate, spiral, sparsely distributed along the branches, but more concentrated near apex; blades 17.5–24 × 4–4.5 cm, elliptic, widely elliptic, ovate-lanceolate, oblong-elliptic or sometimes lanceolate, entire, unlobed, non-peltate, base obtuse or attenuate, apex acute and mucronulate, chartaceous, glabrous on both surfaces, adaxial surface dark purple or greenish purple, abaxial surface glaucous to cinereous, the latter with a smooth wax pattern, venation camptodromous-brochidodromous, primary veins prominent on both surface, secondary veins impressed on both surfaces, diagonally arranged towards the midvein, all pinkish to purplish. Racemes 4.5–10.5 cm long, staminate or bisexual, solitary, terminal or arising from dichotomy of the branches, erect or pendent, congested, glabrous, angulate, glaucous to cinereous, waxy. Staminate flowers: buds 4–7 × 3–6 mm, widely ovoid, yellowish, without purplish pigmentation; bracts 10–15 × 3–7 mm, widely ovate, foliaceous, apex acuminate, margins sparsely and irregularly denticulate, persistent; bracteoles situated along the lower third up to half of the pedicel, 4–4.6 × 1–1.5 mm, ovate or ovate-laceolate, foliaceous, persistent, subopposite; pedicels 3–3.2 mm long, cylindrical, glabrous, light green; calyx 7–15 × 5–10 mm, campanulate, green-yellowish, without purplish pigmentation, shortly tomentose internally, lobes ovate-oblong with apex obtuse or rounded, base truncate; stamens 10, in two whorls of five, glabrous, the longer 7.5–7.6 mm long, the shorter 4.5–4.6 mm long, both thickened, anthers 3–4 mm long, oblong, bright yellow, dorsifixed; disc 10-lobed, intrastaminal, dark yellow. Pistillate flowers: buds 6–8 × 4–6 mm, ovoid, green-yellowish, without purplish pigmentation; bracts 5–9 × 1.5–5 mm long, widely ovate, entire, apex acuminate, margin irregularly serrate, glabrous, persistent; bracteoles 4–6 × 1–3 mm, oblong-elliptic, foliaceous, persistent, margin denticulate, apex acuminate, opposite along the lower third of the pedicel, glabrous; pedicels 4–5 mm long, cylindrical-clavate, glabrous, green; calyx 7–9 × 5–7 mm, campanulate, yellowish green, without purplish pigmentation, glabrous externally, shortly tomentose internally, lobes triangular, base truncate; ovary 5–7 × 2–3 mm, ovoid to oblong, glabrous, green, disc patelliform, lobed, yellow; styles 3, shortly united at the base, densely papillose lobed. Capsules 0.8–1.8 × 0.7–1.2 cm, oblong, light green, smooth, glabrous, without wings, waxy, dehiscence septicidal and loculicidal; columella (carpophore) persistent, 0.78–0.8 × 1.2–1.3 mm (width at narrowest point in middle), dilating to 2–2.2 mm at both tip and base, narrowly alate. Seeds 7–10 × 5–6 mm, oblong, dark grey or cinereous, with black spots; caruncle prominent, widely triangular with entire apex, cream.
Manihot attenuata. A Habit B Details of the branch showing the persistent stipule C Stipule D Staminate bracts E Staminate bracteole F Staminate bud G Staminate flower H Staminate flower with calyx split and open I Pistillate bracts J Pistillate bracteole K Pistillate flower, note the gamosepalous calyx L Pistillate flower with calyx split and open M Fruit N Seed, ventral side O Seed, dorsal side. Drawn by Cristiano Gualberto from the holotype.
Manihot attenuata is easily recognised by its shrub habit; erect or decumbent habit; large leaves (17.5–24 × 4–4.5 cm) that are purplish; sparsely serrate, persistent stipules; bracts and bracteoles of both staminate and pistillate flowers widely ovate, pubescent externally and acuminate at the apex; usually pendent inflorescences; and pubescent stigmatic branches. It is morphologically similar to M. fallax. However, it differs by the set of characters cited in Table
A species endemic to the northern portion of the state of Goiás, Chapada dos Veadeiros and neighbouring regions (municipalities of Niquelândia and Minaçu) (Fig.
Morphological characters useful in separating M. fallax from M. attenuata.
Character | M. fallax | M. attenuata |
---|---|---|
Leaves | 8–14 × 1–2 cm, lanceolate, narrowly elliptic or linear, adaxial surface dark green, abaxial surface glaucous | 17.5–24 × 4–4.5 cm, widely elliptic, ovate-lanceolate, oblong-elliptic or sometimes lanceolate, adaxial surface dark purple or greenish purple, abaxial surface glaucous or cinereous |
Arrangement of secondary veins in relation to the midvein | Perpendicular | Diagonal |
Stipules | Margin entire, early caducous 2.9–3 mm long | Margin serrulate, persistent, 10–11 mm long |
Inflorescence | Racemes subspicate, 2.5–6 cm long | Racemes with well developed pedicels, 4.5–10.5 cm long |
Bracts on flowers of both sexes | Glabrous, widely ovate with entire margin | Pubescent, widely elliptic with denticulate margin |
Lobes of staminate calyx | Triangular with apex acuminate | Ovate-oblong with apex obtuse or rounded |
Staminate buds | Obovoid | Ovoid |
Filaments | Pubescent | Glabrous |
Pistillate bracts | 5–9 mm long, margin entire | 6.4–7 mm long, margin irregularly serrate |
Pistillate bracteoles | Lanceolate, margin entire | Oblong-elliptic, margin denticulate |
Pistillate calyx | Dialisepalous | Gamosepalous |
Caruncle shape | Triangular with bilobed apex | Widely triangular with entire apex |
The species has been collected with flowers from November to March, with the flowers more common from September to December and with fruits from December to March.
Manihot attenuata is here classified as Vulnerable (VUVU) according to the IUCN Red List Categories and Criteria (
Recently,
BRAZIL. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada GO 327, aproximadamente 1800 metros antes da Vila São Jorge, 14°09'00"S, 47°37'00"W, 1000 m a.s.l., 29 Mar 1988, I. R. S. Costa et al. 19 (UFG); entrada do Parque Nacional da Chapada dos Veadeiros, 14°10'5"S, 47°47'25"W, 27 Jan 1996, fl., fr., W. L. Werneck 706 (CEN); entrada do Parque Nacional da Chapada dos Veadeiros (a esquerda), estrada Alto Paraíso-Colinas, 34 km da GO 118, 14°09'49"S, 47°47'8"W, 1250 m a.s.l., 22 Jan 1997, fl., B. M. T. Walter et al. 3644 (CEN); Parque Nacional Chapada dos Veadeiros (PNCV) área queimada com relevo plano, 15°44'14"S, 47°54'54"W, 1477 m a.s.l., 25 Sep 1995, fl., M. L. Fonseca et al. 549 (IBGE); ib. Cerca de 1 km acima do alojamento dos brigadistas, 20 Jan 2012, fl. fr., M. J. Silva et al. 4063, 4065 (UFG); ib., Proximidades do alojamento dos brigadistas, 25 May 2012, fr., L. C. S. Almeida et al. 42 (UFG); ib., final do acesso da trilha que leva ao Cânion I, entre fendas de rocha, 14°09'6.55"S, 47°48'0.74"W, 1114 m a.s.l., 29 Sep 2012, fl., M. J. Silva et al. 4421 (UFG); 2.5 km em direção ao rio Preto, 26 Oct 2012, fl., M. J. Silva et al. 4502 and 4506 (UFG); na subida para o alojamento dos brigadistas do PNCV, 22 Oct 2011, fl., M. J. Silva et al. 3885 (UFG); Parque Nacional Chapada dos Veadeiros, estrada que leva ao alojamento dos brigadistas, 14°9'53.6"S, 47°47'25.7"W, 1034 m a.s.l., 31 Oct 2014, fl., L. S. Inocencio et al. 180, 181, 182 and 183 (UFG); ib., segundo morro a nordeste do alojamento principal dos brigadistas do PNCV, 27 Oct 2012, fl., M. J. Silva et al. 4503 and 4507 (UFG); topo da serra à direita da estrada que leva ao Cânion I, 14°09'05.7"S, 47°48'00"W, 1130 m a.s.l., 23 Nov 2012, fl., L.C.S. Almeida & I. A. M. Watanabe 67 and 69 (UFG); estrada para Vale da Lua, 24 Nov 2012, fl., M. J. Silva et al. 4589 (UFG); trilha a partir do dos alojamentos do PNCV em direção ao Cânion I, 14°08'S, 47°43'W, 1086 m a.s.l., 29 Nov 2012, fl., L. C. S. Almeida et al. 50 (UFG); afloramento rochoso à nordeste do Cânion I, 14°9'S, 47°48'W, 1082 m a.s.l., 29 Nov 2012, fl., L. C. S. Almeida et al. 51, 52 and 56 (UFG); subida de acesso ao alojamento dos brigadistas do PNCV, 14°09'37.6"S, 47°57'2.3"W, 1034 m a.s.l., 22 Feb 2014, fl., fr., M. J. Silva et al. 5921 (UFG); cerca de 500 m do alojamento dos brigadistas, 14°9'22.4"S, 47°47'38.9"W, 1089 m a.s.l., 13 Dec 2014, fl., L. S. Inocencio et al. 278 (UFG); Parque Nacional da Chapada dos Veadeiros, estrada que leva ao alojamento principal do Parque, 04 Jan 2015, fl., R. C. Sodré, M. J. Silva & C. H. G. M. Filho 1636 (UFG). Cavalcante, exatos 8 km a noroeste de Cavalcante ao longo da estrada de terra que leva à Colinas do Sul, 13°50'S, 47°33'W, 12 Dec 1986, fl., fr., A. C. Allem et al. 11307 (CEN); estrada Minaçu-Cavalcante passando pela balsa Serra Branca (COTERRA), à cerca de 126 km do rio Tocantins, 13°41'07"S, 47°51'22"W, 740 m a.s.l., 10 Nov 2000, fl., G. P. Silva 4421 (UFG); cerca de 4 km da Vila Veneno, na direção do rio São Félix, cerca de 12 km da Balsa da Coterra, 13°32'10"S, 48°03'29"W, 380 m a.s.l., 25 Jan 2001, fl., B. M. T. Walter et al. 4807 (UFG). Chapada dos Veadeiros, imediações da Serra do Ministro, a aproximadamente 500 m da estrada entre Cavalcante e Colinas do Sul, 13°54'34"S, 47°38'55,4"W, 20 Jan 2014, fr., R. C. Sodré et al. 1193 (UFG). Minaçu, 7 km após a entrada norte do canteiro de obras, 13°28'00"S, 48°23'00"W, 920 m a.s.l., 10 Mar 1992, fl., fr., T. B. Cavalcanti et al. 1112 (CEN). Niquelândia, Reservatório em formação do AHE Serra da Mesa, região na margem direita do rio Maranhão, próximo ao rio das Almas, 14°34'50"S, 48°59'07"W, 440 m a.s.l., 16 Oct 1997, fl., B. M. T. Walter 3924 (CEN).
The first author thanks Conselho Nacional de Desenvolvimento Científico e Tecnológico for the productivity grant (CNPq, Proc. no. 302648/2016-0) to develop the project “Estudos taxonômicos sobre o gênero Manihot Mill. (Euphorbiaceae) para a Região Centro-Oeste do Brasil”, to Cristiano Gualberto for the illustration; and Suzana Oellers for English language editing. The second and third authors thank Cordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the Master’s grants.