Utricularia tricolor Saint-Hilaire (1833: 418)

Baleeiro, Paulo C., Gonella, Paulo M., Sano, Paulo T. & Jobson, Richard W., 2022, Unveiling Utricularia amethystina’s true colours: a taxonomic revision of one of the largest species complexes (U. sect. Foliosa, Lentibulariaceae), Phytotaxa 576 (1), pp. 29-54 : 47-49

11. Utricularia tricolor Saint-Hilaire (1833: 418) — Fig. 4–I

Type: — BRAZIL. RIO DE JANEIRO: Lieux humides & sablonneux voisins de la mer près S . Joao de Barra, 1816–21, St.Hilaire B2-161 (holotype: P! (photo, M), Isotype: MPU, P) .

= Utricularia fontana Saint-Hilaire & Girard (1838: 870)

Type:— BRAZIL. SÃO PAULO: in quodam fonte prope Paulopolium, Saint-Hilaire C1-1147 (Holotype: P image!, Isotype: P image!) .

= Utricularia gomezii De Candolle (1844: 13)

Type:— BRAZIL. RIO DE JANEIRO: Gomez s.n. (Holotype: G, Isotype: K!, P image!) .

= Utricularia globulariifolia Martius ex Benjamin (1847: 241)

Type:— BRAZIL: RIO DE JANEIRO: Gardner 590 (Holotype: BM!, Isotype: G!, K!, NY!, P image!, S, W) .

= Utricularia fusiformis Warming (1874: 9)

Type:— BRAZIL. MINAS GERAIS: Lagoa Santa, Warming s.n. (Holotype: C image! (photo, M)) .

= Utricularia globulariifolia var. caudata Sylvén (1909: 23)

Type:— BRAZIL. MINAS GERAIS: Caldas, Mosén 4349 (Holotype: C image!) .

Illustrations: — Taylor (1989), fig. 79; Baleeiro et al. (2016), fig. 2m; Baleeiro et al. (2019), fig. 1c.

Description:—Terrestrial, or affixed sub-aquatic. Small to medium sized, annual herb. Rhizoids up to 90 mm long. Leaves usually 1(–3) from peduncle base, lamina obovate to circular, 5–55(–65) mm long, traps numerous on rhizoids and stolons, broadly ovoid, 1–2 mm long. Inflorescence simple (240–) 300–750 mm long, peduncle glabrous, 0.5– 1.4mm long, scales basifix, glabrous, deltoid, apex acute, margin entire 1.3–1.5 mm long, bract ovate to deltoid, apex acute c. 1.5 mm long, bracteoles smaller or similar length, lanceolate, connate near the base, or sometimes middle, glabrous. Flowers 1–8, pedicel (1–) 2–4 mm long, calyx lobes unequal, coriaceous, usually glabrous or papillose near margin, rarely with sparse sessile glands, inconspicuous simple nerves, upper lobe ovate-oblong or broadly elliptic to circular, concave, apex rounded, margin entire, hyaline 2.8–3.1 × 2.3–2.7 mm, lower lobe broadly elliptic, entire, margin not hyaline, apex crenate or emarginate, 1.5–2.0 × 2.5–3.1 mm, corolla lilac with yellow spot with white margin at base of lower lip, 10–25 mm long, with papillae and few glandular trichomes, upper lip broadly ovate, apex rounded, equal to about twice the length of calyx lobes, 2.0– 4.3 mm long, lower lip transversally elliptic or broadly obovate, base forming a bilobed swelling, apex entire or shallowly 3-lobed, 6–9 × 8–10 mm, spur slightly conical, apex obtuse, equal or slightly larger than the lower lip, 7.2–8.4 mm long, filaments curved, anther thecae sub-distinct, ovary globose with some glands on the outside wall. Capsule globose up to 4 mm diam. Seeds narrowly cylindrical c. 1 mm long, 0.15 mm wide.

Distribution and ecology: — Argentina, Brazil, Colombia, Uruguay, and Venezuela ( Fig. 1 ). In Brazil, in the states of Bahia, Espirito Santo, Goiás, Maranh„o, Mato Grosso, Minas Gerais, Pará, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina and S„o Paulo. Grasslands bordering creeks, lakes, gallery forests, veredas (a phytophysiognomy composed of the palm Mauritia flexuosa L.f.), restingas, and campo rupestre. Flowering August to May.

Etymology: —The epithet tricolor refers to the three colors of the corolla, lilac with yellow and white at base of the lower lip.

Conservation status: —Least Concern. Based on its wide distribution (EOO = 8,284,102 km 2, AOO = 132 km 2) and frequent presence in protected/conservation areas, U. tricolor is here considered Least Concern, in accordance with Cross et al. (2020).

Notes: — Utricularia tricolor ( Fig.4I ) is easily distinguished from all other U. sect. Foliosa species by its robust specimens with tall inflorescence scapes and large flowers. The most similar species is U. bicolor (previously a synonym of U. amethystina ), which can be mistaken for U. tricolor from the Espinhaço Range in Minas Gerais, where both taxa are sympatric, and specimens of U. tricolor are often smaller. In some specimens, it is possible to observe a few trichomes and papillae near the calyx margin, one of the defining characters of U. bicolor . However, U. tricolor has a larger corolla with lower lip apex almost entirely or shallowly 3-lobed and glabrous calyx, except in some populations collected from the Espinhaço Range. At this site it is notable that U. bicolor does not possess a white margin surrounding the yellow spot at the base of lower lip, as does U. tricolor . Nevertheless, the material examined during this study suggests hybridisation between these two taxa. It is worth mentioning that these two species are not phylogenetically close, with U. tricolor sister to U. amethystina s.str., and U. bicolor sister to a clade containing the U. hirtella complex ( Baleeiro et al. 2019).

Additional specimens Examined: — ARGENTINA. Corrientes, Dep. San Tome, 4km E de Ruta Nac. n.14, caminho a colônia Garabi , 3 Dic. 1970, Krapovickas s/n ( RB 176498 ) .

BRAZIL. BAHIA: Abaíra, Catolés, subindo riacho que desce da serra do Guarda-mor , 26 Jul. 1995, Rivadavia 479 (SPF); ibid, em campos rupestres beirando a trilha para o campo da Mutuca, Serra da Mesa , Vila de Catolés , 15 Jul. 2005, Rivadavia et al. 2020 (SPF). Mucugê, Serra de S„o Pedro, 17 Dez. 1984, Lewis et al. CFCR 7078 (SPF). Piat„, campos rupestres ao lado da capelinha na trilha subindo a Serra do Santana, 13 Jul. 2005, Rivadavia et al. 2001 (SPF). Rio de Contas, campos rupestres próximo a trilha para o alto do Pico do Itobira , Serra da Mesa , 18 Jul. 2005, Rivadavia et al. 2077 (SPF). GOIÁS: Alto Paraiso de Goiás, Parque Nacional dos Veadeiros, córrego dos Ingleses, 19 Jun. 2013, Baleeiro & Carneiro 335 (SPF); ibid, Parque Nacional dos Veadeiros, cachoeiras Almecegas I e II, 21 Jun. 2013, Baleeiro & Carneiro 341 & 344 (SPF); ibid, Chapada dos Veadeiros, km 200 da estrada Brasília—Alto Paraíso , 13 Abr. 1995, Rivadavia & R.C. Ogassavara 373 (SPF). Barro Alto , GO-080, 6 Jan. 2001, Rivadavia 1229 (SPF). Cristalina, BR-060. RPPN Linda Serra dos Topázios, 9 Jun. 2009, Baleeiro et al. 72 (R). Mineiros , estrada para o Parque Nacional das Emas, 3 Mai. 1999, Rivadavia & R.K. Sano 1041 (SPF). Rio Verde , BR 060 , 16 Jan. 2004, C.P. Bove et al. 1332 (R). DISTRITO FEDERAL: Brasília , Reserva Ecológica do IBGE, brejo entre o Guará e o setor da indústria, 21 Fev. 1981, F. Chagas & Silva 390 (IBGE). Gama, estrada para o Santuário Ecológico , 26 Abr. 1999, Rivadavia & R.K. Sano 944 (SPF). MATO GROSSO DO SUL: Rio Verde do Mato Grosso, estrada para Morro do Padre , vereda, 3 Nov. 20112, A.V. Scatigna et al. 281 (UEC). MINAS GERAIS: Chapada Gaúcha, nascente brejosa com buritis no lado E do Parque Nacional Grande Sert „o Veredas, 28 Abr. 1999, Rivadavia & R.K. Sato 960 (SPF). Diamantina, Parque Nacional Sempre Vivas, 8 Mar. 2013, Baleeiro et al. 307 (SPF); ibid, campo em frente ao entroncamento para Conselheiro Mata, 9 Mar. 2013, Baleeiro et al. 310 (SPF). Jaboticatubas, Serra do Cipó, 11 Set. 1994, Rivadavia 307 (SPF). Prata, c. 2.5km ao noroeste da cidade, brejo beirando a BR497 , 28 Jul. 2007, Gonella et al. 19 (SPF). Rio Vermelho, Pedra Menina, Serra do Ambrósio, Espig„o do Meio Faz. Vargem do Anjo , 8 Set. 1986, Mello-Silva et al. CFCR 10208 (SPF). Santana do Riacho , Serra do Cipó , em nascente brejosa antes da bifurcaç„o da BR10 para Conceiç„o do Mato Dentro e Morro do Pilar , 12 Nov. 2007, Gonella et al. 43 (SPF); ibid, trilha para cachoeira Capivara, 5 Jul. 2012, Baleeiro et al. 204 (SPF). S„o Roque de Minas, 13 Mar. 1990, Souza, H.C. s/n (BHCB); ibid, caminho do Abrigo de Pedra, Parque Nacional Serra da Canastra , 22 Dez. 1991, Rivadavia 42 (SPF). Uberaba , em brejo c. 30km ao noroeste da cidade, no lado leste da MG 190 , 27 Jul. 2007, Gonella et al. 8 (SPF). PARAÍBA: Mamanguape , 1 Jan. 1900, Felix 2104 (JPB). PARANÁ: Palmeira, Rio das Pombas , beirando o km 522 da BR376 , 25 Nov. 2006, Rivadavia & Rohrbacher 2454 (SPF). RIO DE JANEIRO: Cabo Frio, entre Lagoa de Araruama e Praia de Massambaba , brejo dos Espinhos, 20 Dez. 1982, fl., Pedrosa, D.S. et al. 742 (GUA). Carapebus , praia entre Lagoa Paulista e Lagoa de Carapebus , C.P.R. Batista 35 et al. (R). Saquarema , Praia do Sossego , C.B. Moreira 53 et al. (R). Teresópolis , descendo a estrada Petrópolis-Teresópolis ( BR 495 ) em direç„o a Teresópolis, 19 Nov. 2000, Rivadavia 1220 (SPF). SÃO PAULO: Bocaina, 10 Fev 1870, Glaziou 8218 (R). Botucatu, c. 1km pela estrada de ferro da FEPASA a partir da rodovia Marechal Rondon, em nascentes brejosas em encosta de morro acima dos trilhos, 21 Mar 1999, Rivadavia et al. 860 (SPF). Cananéia, Parque Estadual Ilha do Cardoso , 9 dez. 2002, Foster et al. 978 (SPF). Itararé, descendo trilho de trem em direç„o a Gruta Santa, próximo à divisa SP/ PR, 25 Jan. 1993, Rivadavia 167 (SPF). Ribeir„o Pires, Velha estrada para Santos, próximo ao início da decida da Serra, 11 Jan. 1992, Rivadavia & Cardoso 53 (SPF). S„o Jose do Barreiro, Parque Nacional da Bocaina, cachoeira de Santo Izidro, 22 Out. 2002, Rivadavia & Morais 1414 (SPF). Vargem Grande do Sul , brejo na estrada para Casa Branca , antes do Rio Verdinho , 14 Nov. 1997, J.H.A. Dutilh & W. Marcondes-Ferreira s/n ( UEC 94951 ). SANTA CATARINA: Itapiruba, sul da cidade, a alguns km da BR 101 , 7 Set. 2004, Rivadavia 1861 (SPF). Tocantins, Parque Estadual do Jalap„o, Brejo do Porco Podre , local de colheita do capim dourado, 19 Jun. 2002, Barbosa Cavalcanti 3005 (CEN).

COLOMBIA. SANTANDER: Charala, corregimento de Virolin: em zonas pantanosas cerca de Virolin, 21 Mar. 2008, Vieira, S . 4 (SPF).

VENEZUELA. ESTADO AMAZONAS: Cerro Duidia , pico ao extremo NW do maciço, vizinho a aldeia indígena de Culebra, 5 Jan. 2003, Rivadavia & HoogenstriJd 1515 ( SPF) .

Figures

FIGURE 1. Geographic distribution map of Utricularia sect. Foliosa species. A. Distribution of all 14 species in South and Central Americas. B. Map of the species distribution in the Espinhaco range and Chapada dos Veadeiros National park. C. Map of the species distribution at the Guiana Shield.

FIGURE 4. A–D. U. hirtella s.l., E–G. U. velascoensis, H. U. tridentata, I. U. tricolor, J. U. biceps, K. U. bicolor, L. U. chapadensis, M–N. U. pantaneira, O. U. trinervia. Images C. Rivadavia, D. Rohrbacher, E–G. by Marcos Cardoso, J., K. Gonella.