THE VIOLETS OF THE BRAZILIAN SAVANNA: A REVISION OF THE POMBALIA LANATA COMPLEX (VIOLACEAE), WITH DESCRIPTIONS OF TWO NEW SPECIES Author Paula-Souza, J. Author Lima, A. G. Author Souza, V. C. text Edinburgh Journal of Botany 2021 2021-04-07 78 357 1 32 http://dx.doi.org/10.24823/ejb.2021.357 journal article 285207 10.24823/EJB.2021.357 79160911-6a72-466a-b594-3a50c8a4341c 1747-0036 10513850 3. Pombalia lanata (A.St.-Hil.) Paula-Souza, Phytotaxa 183(1): 9 (2014) . – Ionidium lanatum A.St.- Hil., Ann. Sci. Nat., Ser. 1, 2: 254 (1824) . Type: Brazil , Minas Gerais , “Paturages herbeus pr. Tejuco dans le District des diamans, campis”, 1817, St. Hilaire Catal. B1 no. 2080 ( lectotype P ! [ P02141267 ], designated by Paula-Souza in Paula-Souza & Ballard, 2014 ) – Solea lanata (A.St.-Hil.) Spreng., Syst. Veg. 4(2): 97 (1827) . – Hybanthus lanatus (A.St.-Hil.) Baill. , Traité Bot. Méd. Phan. 2: 841 (1884). – Calceolaria lanata (A.St.-Hil.) Kuntze , Revis. Gen. Pl. 1: 41 (1891) . – Hybanthus lanatus (A.St.-Hil.) Taub. , Nat. PflanZenfam. 3(6): 333 (1895). Figures 1A,B , 5A–D , 6A–F . Figure 5 . A–D, Pombalia lanata : A, habit; B–D, details of flowering branches, showing colour variation; E and F, Pombalia poaya : E, detail of flowering branch; F, habit. Photographs: A, C, D and F, Henrique Moreira; B, Jair E. Q. Faria. Figure 6 . A–F, Pombalia lanata : A, habit; B, variation of size and shape of the corolla; C, gynoecium on the pedicel; D, calyx; E 1 –E 3 , variation of size, indument and shape of connective appendage of posterior stamens; E 4 –E 8 , variation of size, indument and shape of connective appendage of anterior stamens; F, typical configuration of the androecium in the Pombalia lanata complex. G, Pombalia atropurpurea : typical configuration of the androecium in the remaining hybanthoid lineages ( an , anther; ca , connective appendage; fl , filament; gl , nectary gland; st , style). A–F, modified from Paula-SouZa et al. , 2011 , and reproduced with the permission of the Conservatoire et Jardin botaniques de Genève; G, drawn from Souza 3545. Illustration by by Rogério Lupo. Ionidium lanatum A.St. -Hil. var. dentatum A.St. -Hil. (“ dentata ”), Mém. Mus. Hist. Nat. 11: 484 (1824) . – Type : BraZil , “Retiro pr. pago Contendas in deserto flum, S. Francisco”, Saint Hilaire s.n. ( lectotype P! [ P2141270 ], designated here; isolectotype P! [ P21412701 ]), syn. nov. Ionidium lacteum S.Moore, Trans. Linn. Soc. London, Bot., Ser. 2, 4: 307 (1895) . – Type: Brazil , Mato Grosso : “In campis ad Serra da Chapada alt. circa 1800 ped.”, 1901–1902, Moore 187 ( holotype BM! [ BM000626822 ]), syn. nov. Hybanthus subpoaya Schulze-Menz, Notizbl. Bot. Gart. Berlin-Dahlem 12(111): 112 (1934). – Type: Brazil , “Goyaz [currently Tocantins ]. Sandy campos banks of the Rio Manoel Alvez, near the Miguari do Ouro, 1842”, Gardner 3006 ( holotype B †; lectotype BM ! [ BM000617712 ], designated here; isolectotype E !, F !, G !, K !, P !), syn. nov . Suffrutescent herb, 15–20 cm high, branched from a well-developed, bud-bearing underground root-like system, erect; internodes 5–18 mm . Stipule 3.5–6 × 0.5–1 mm , margin entire; leaves alternate, subsessile; blade 15–36 × (5.5) 7–15 mm , oblong, ovate, elliptic, obovate to oblanceolate, less frequently lanceolate or suborbicular, apex acute to obtuse or less frequently rounded, apiculate, margin entire or irregularly serrate at the apex, base acute to attenuate, tomentose to densely villous, less frequently pubescent, hairs all stellate or frequently simple with sparse or rare stellate hairs. Flowers deep violet, blue, light blue, lilac, white, greenish or yellowish, with a yellow spot at the base of the anterior petal, solitary, axillary; pedicel 11–21 mm , tomentose to villous; bracteoles 1–1.5 × c. 0.5 mm ; sepals equal, 5–8(10) × 1.5–3 mm , lanceolate to ovate, frequently subfalcate, margin entire, base truncate, tomentose to villous; posterior petals 5.5–8 × 1–3 mm , ovate or oblong, subfalcate; lateral petals 10.5–16.3 × 2–5(12) mm, spatulate to oblong, frequently constricted at the central portion, falcate, rarely clawed, tomentose to villous at apex; anterior petal 16–18.5 × 13–22 mm including a claw 7–9 mm long, oblate, deltoid, transversely oblong or elliptic, or widely transversely rhombic, apex truncate, emarginate or obtuse, smooth, tomentose to villous; filaments 1–2 mm , connective appendages hyaline, 0.3–0.5 mm long, nectar glands noduliform, frequently much reduced, inserted on the filaments usually near the base of the anthers, subglabrous to villous; style 3–4 mm , glabrous or rarely tomentose at the base, straight or less frequently subsigmoid, ovary 2–2.5 mm , villous. Capsule 8.5–10 × 6–7.5 mm , ellipsoid to ovoid; seeds c.3 × 2–2.5 mm . Vernacular name . Poaia ( Brazil ). Etymology . The epithet lanata (‘woolly’) refers to the dense indument typical of this species. Distribution and habitat . Pombalia lanata occurs in savannas, grasslands and campos rupestres of the Brazilian Plateau from Tocantins to Mato Grosso do Sul, also reaching Espinhaço Range in Minas Gerais, with a disjunction in Misiones province, Argentina ( Figure 7 ; see also Paula-Souza et al. , 2011 ). The collection Gardner 2397 at K registered for Ceará state is probably a case of mixed labels, because there is a gathering of Pombalia arenaria by Gardner with the same number from the same location (“Cachoeira”), which is situated in a typical xerophytic, deciduous Caatinga vegetation quite distant from the known geographical limits of P. lanata . Figure 7 . Distribution of Pombalia lanata . Phenology . The species has been collected with flowers and fruits throughout the year, but most specimens were collected from July to November, peaking in September and October. Conservation status . This is the most widely distributed species of the Pombalia lanata complex (EOO, 1,554,464 km 2 ; AOO, 152 km 2 ), with abundant records in conservation units. Pombalia lanata is thus assigned a conservation status of Least Concern (LC). Selected specimens examined . ARGENTINA . Misiones : San Ignácio , Teyucuaré , 27°16′S , 55°33′W , 150 m a.s.l. , 19 ix 2000 , Múlgura de Romero et al. 2065 ( MO , NY , SI ) . BRAZIL . Distrito Federal : Brasília , APA Gama-Cabeça de Veado , região administrativa do Núcleo Bandeirante , ARIE do Córrego do Cedro , SMPW-Q .26 Conj. 3, 15°54′49′′S , 47°57′50′′W , 1050 m a.s.l. , 14 x 2002 , Fonseca et al. 3671 ( IBGE , RB ) ; Planaltina , Área de Chapadão BR-020, Sobradinho-Planaltina , km 15, à direita da rodovia, 15°37′00′′S , 47°42′30′′W , 900 m a.s.l. , 1 ix 1994 , J . C . Silva 739 ( CEN ) ; Vargem Bonita , APA do Gama – Cabeça de Veado ( RA ) Núcleo Bandeirante , Park Way , Lagoa do Córrego do Cedro na entrada para a Vargem Bonita , 15°53′51.8′′S , 47°56′39.8′′W , 990 m a.s.l. , 11 ix 2002 , Mendonça et al. 5018 ( IBGE , RB ) . Goiás : Alto Paraíso de Goiás , Parque Nacional da Chapada dos Veadeiros , Pouso Alto , 13°58′26′′S , 47°29′59′′W , 1546 m a.s.l. , 26 xi 2014 , Verdi et al. 7311 ( CEN , RB ) ; Alvorada do Norte , Rod. BR 020 , 25 km S de Alvorada do Norte , 11 x 1976 , Hatschbach 39133 ( MBM ) ; Barro Alto, área da mineradora Anglo American Brasil , 15°06′06.2′′S , 49°00′43′′W , 7 vii 2017 , Pachêco et al. 68 ( CEN ) ; Caiapônia , Serra do Caiapó , c. 33 km S of Caiapônia on road to Jataí , 17°12′S , 51°47′W , 800–1000 m a.s.l. , 18 x 1964 , Irwin & Soderstrom 6995 ( COL , MO , NY ) ; Campinaçu , depois de Buriti , km 18, estrada vicinal, 13°31′08′′S , 48°11′14′′W , 330 m a.s.l. , 5 x 2000 , Cavalcanti et al. 2669 ( CEN ) ; Cavalcante , estrada Balsa Porto dos Paulistas (rio Tocantins ) para HU , c. 32 km da balsa, 13°22′52′′S , 48°00′00′′W , 760 m a.s.l. , 9 xi 2000 , G . P . Silva et al. 4408 ( CEN ) ; Chapadão do Céu / Mineiros , Parque Nacional das Emas , próxima ao ponto J , 17°49′– 18°28′S , 52°39′– 53°10′W , 10 ix 1999 , Batalha 3885 ( ESA ) ; Cocalzinho , Serra dos Pireneus , Parque Estadual da Serra dos Pireneus , após a 2a. porteira, saindo da Faz. Capitão do Mato , sentido para o Morro do Cabeludo , 15°48′S , 48°48′W , 24 x 2004 , Delprete et al. 8981 ( NY , SPF ) ; Corumbá de Goiás , Serra do Bicame , estrada para Pirenópolis , 1 viii 1984 , Pereira et al. 1093 ( IBGE , US ) ; Cristalina, Rod. BR 050, km 70, 16.6056°S , 47.7684°W [16°36′20.2′′, 47°46′06.2′′], 3 x 2007 , Paula-Souza et al. 8753 ( SPF ) ; Formosa , 19 x 1965 , A . P . Duarte 9379 ( RB ) ; Goiânia, na GOM-9, p/ Nerópolis, 2 km da Esc. Agron. e Veterinária , 1 xii 1968 , Rizzo & Barbosa 2841 ( UFG ) ; Hidrolândia , Morro Feio , 8 xi 2007 , Pastore 2257 ( HUEFS ) ; Jataí , estrada de Jataí para Serranópolis , a 20 km do Ribeirão Ariranha , 18 x 1972 , Rizzo 8474 ( ESA , UFG ) ; Mineiros , Rod. BR 364 , prox. Córrego Alegre , 20 ix 1974 , Hatschbach et al. 35015 ( MBM ) ; Mossâmedes / Goiás , Serra Dourada , área da UFG , s.d. , Rizzo 4484 ( RB , UFG ) ; Niquelândia , Companhia de Níquel de Tocantins – CNT ; estrada à esquerda da mina de níquel, c. 2 km da entrada, 1040 m a.s.l. , 14°20′37′′S, 48°23′57′′SW, s.d. , Mendonça et al. 3097 ( IBGE , RB ) ; Pirenópolis , Parque Estadual da Serra dos Pireneus , estrada para o Morro do Cabeludo , c. 500 m da estrada principal, 15°48′00.4′′, 48°49′41.6′′, 1324 m a.s.l. , 25 x 2012 , Paula-Souza et al. 11354/728 ( ESA ) ; Planaltina , GO-118, próximo ao km 0 (divisa GO-DF ), 33 km S de São Gabriel , 15.5116°S , 47.5283°W [15°30′41.8′′S, 47°31′41.9′′W], 3 x 2007 , Paula-Souza et al. 8792 ( SPF ) ; Teresina de Goiás , Faz. Hotel Ecológico Alpes Goianos , Rod. GO-118, km 202, 13°53′59.1′′S , 47°23′48.9′′W , 31 vii 2000 , J . P . Souza et al. 3543 ( ESA ). Mato Grosso : Chapada dos Guimarães , Sta. Anna da Chapada , 20 ix 1902 , Robert 547 ( BM , K ) ; Itiquira , c. 23 km leste de Itiquira na estrada MT-299 a Alto Araguaia , 17°14′12′′S , 53°47′50′′W , 550 m a.s.l. , 17 ix 1995 , Pott et al. 7376 ( CPAP ) ; Xavantina , c. 70 km N of Xavantina , 14°40′S , 52°20′W , 300–400 m a.s.l. , 10 x 1964 , Irwin & Soderstrom 6727 ( UB ). Mato Grosso do Sul : Campo Grande , Estaca , 10 ix 1936 , Archer & Gehrt 164 ( SP , US ). Minas Gerais : Abadia dos Dourados , arid upland campo N . d’Abadia , v 1840 , s.col. 4091 ( K ) ; Curvelo , início da Serra do Espinhaço , 50 km de Diamantina , 3 x 1980 , Ferreira et al. 1477 ( RB ) ; Delfinópolis , estrada para Gurita , 20°16′52′′S , 46°52′16′′W , 816 m a.s.l. , 21 x 2003 , Volpi et al. 730 ( HUFU , RB ) ; Diamantina , Rapadura , 1280 m a.s.l. , 29 iv 1931 , Mexia 5710a ( BM , F , G , GH , K , MO , NY , US ) ; Gouveia , Serra do Espinhaço , rodovia para Barão do Guaçuí , 1300 m a.s.l. , 24 x 1999 , Hatschbach et al. 69629 ( CTES , MBM , NY ) ; Itacambira , 17°00.57,2′′ S , 43°20.26,6′′ W [17°00′57.2′′, 43°20′26.6′′], 1300 m a.s.l. , 13 xi 2001 , Tozzi & Alencar 2001-478 ( UEC ) ; Monte Azul , c. 23 km SE de Monte Azul , 1331 m a.s.l. , 27 x 2010 , Queiroz et al. 14974 ( HUEFS ) ; Montes Claros , Serra do Catuny , Brejo das Almas , 1000 m a.s.l. , 10 xi 1938 , Markgraf et al. 3230 ( F , R , RB ) ; Patos de Minas , entre Patos de Minas e Pirapora , 16 ix 1963 , R . S . Silva s.n. ( HB [ HB28626 ]) ; Pirapora , 17 km de Pirapora , rumo BR-7, 13 x 1962 , Pabst 7112 ( COL , HB , NY ) ; Presidente Olegário , em direção a Coromandeo , 26 x 1966 , L . Duarte 876 ( HB ) ; Rio Pardo de Minas , Sítio do Sr. José , 15°31′57′′S , 47°27′32′′W , 900 m a.s.l. , 6 xi 2006 , Sevilha et al. 4713 ( CEN ) ; Sacramento , PARNA Serra da Canastra , próximo à guarita de Sacramento , 19 viii 1994 , Nakajima et al. 413 ( ESA , HUFU ) ; Salgado , Brasilia in herbidis umbrosis, s.d. , Martius s.n. ( M ) ; Santana do Riacho , Serra do Cipó , ao lado da Pensão Chapéu de Sol , 6 vii 2001 , J . P . Souza et al. 3583 ( ESA ) ; São Roque de Minas , Parque Nacional da Serra da Canastra , 53.8 km W do centro de visitantes e 15 km E da portaria de Sacramento , 20°08′17′′S , 46°47′07.7′′W , 1367 m a.s.l. , 3 ix 2015 , Pirani et al. 6612 ( SPF ) ; Sopa , près de Biribiry , 1892, Glaziou 18871 ( K , P ) ; Uberlândia , Estação Ecológica do Panga , 29 ix 1992 , Barbosa 872 ( ESA , HUFU ). Tocantins : Natividade , Serra de Natividade , xii 1839 , Gardner 3196 ( K ) . This is the most morphologically variable species within the Pombalia lanata complex, and this variation led to the description of many names that are here treated as synonyms ( Ionidium lanatum A.St. -Hil. var. dentatum A.St. -Hil., I. lacteum S.Moore and Hybanthus subpoaya Schulze-Menz ). Although specimens showing intermediate characteristics might impose some difficulty in identification, Pombalia lanata can be distinguished from the remaining species of the group by a set of features listed in the Table . Notes accompanying some herbarium specimens of Pombalia lanata report that the flowers are intensely fragrant.