Utricularia amethystina Salzm. ex Saint-Hilaire & Girard (1838: 870)

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 : 34-37

1. Utricularia amethystina Salzm. ex Saint-Hilaire & Girard (1838: 870) — Figs. 2 , 3A–C, E–F

Type: — BRAZIL. Bahia: no date, Salzmann s.n. (Holotype: P!; Isotypes: F!, G image!, K!, LE, MEL!, MO!, MPU, P!, R!, W) .

≡ Calpidisca amethystina (Salzm. ex A.St.-Hil & Girard) Barnhart (1931: 468)

= Utricularia modesta De Candolle (1844: 17)

= Calpidisca modesta (A.DC.) Barnhart (1931: 469)

Type:— BRAZIL: Pará , no date, Poeppig 3017 (Holotype: G image!; Isotypes: B (destroyed; photo, M), LE, P image! (photo, M), W) .

= Utricularia obovata Miquel (1847: 144)

Type:— SURINAME: Onoribo , no date, Focke 1051 (Holotype: U3678 ) .

= Utricularia versicolor Benjamin (1847: 488)

Lectotype (designated here):— VENEZUELA: Monegas : Maturin, Otto 946 ( P!).

= Utricularia spatulifolia Pilger (1914: 190)

Type:— VENEZUELA: Roraima , no date, Ule 8757 (Lectotype designated here: K!; Isolectotype: F!, G image!) .

= Calpidisca standleyae Barnhart (1933: 1234)

= Utricularia standleyae (Barnhart) Rickett (1967: 518)

Type:— USA: Florida : Lee County, “Vicinity of Fort Meyers” Among (c.f.) Palmetos ; Standley 406 (Holotype: NY!; Isotype: US!).

= Utricularia williamsii Steyermark (1950: 126)

Type:— HONDURAS: oak-pine forest near Hoya-Grande , Río Yeguare drainage, Williams 10639 (Holotype: F!; Isotypes: BM!, EAP) .

= Utricularia turumiquirensis Steyermark (1953: 552)

Type:— VENEZUELA: Sucre, Cerro Turumiquire, Steyermark 62752 (Holotype: F!; Isotype: VEN) .

Illustrations: — Taylor (1989), figs. 77a, 77b, showing a mix of specimens from different localities; Baleeiro et al. (2016), figs. 2a,g; Baleeiro et al. (2019), fig. 1a.

Description:—Terrestrial or affixed sub-aquatic. Small to medium sized annual herb. Rhizoids 1–4, up to 14 mm long, branches ramified. Stolons numerous. Leaves 1–7, rosulate from peduncle base, lamina normally spathulate to circular (15–)30–40 × 1.5–8.5 mm; traps numerous on rhizoids and stolons, c. 1 mm long, globular, the trap door basal, stalk deltoid 0.5mm long. Inflorescence simple or loosely ramified (30–) 40–300 (–400) mm long, peduncle glabrous, capillary or sometimes thicker near the base, 0.35–1.3 mm diam.; scales basifix, glabrous, ovate, margin entire (3–6) c. 1.1 mm long; bracts and bracteoles unequal, basifix, often connate near base, glabrous without nerves, bract broadly ovate, apex acute, bracteoles linear, apex obtuse 1.2–2.0 mm long. Flowers 1–9 (usually single flowered); pedicel erect, filiform (–1) 2–11 (–18) mm long, 0.4 mm diam.; calyx lobes sub-similar, cartaceous, concave, glabrous, with inconspicuous simple nerves, upper lobe broadly ovate or oblong, apex rounded, entire margin hyaline (1.5–) 2.5–3.0 (–3.9) mm long.; lower lobe ± broadly elliptic, ± entire margin not hyaline, apex rounded or emarginate, 1.3–2.8 mm long; corolla lilac with single yellow spot bordered by white at base of lower lip, spur white at base with apex lilac, 7–12 mm long; upper lip glabrous, oblong, apex rounded 1.35–4.37 × 1.41–4.13 mm; lower lip broadly obovate, base forming bilobed swelling, apex shallowly 3-lobed, 1.4–5 × 2.22–6.73 mm; spur conical from the base, apex obtuse 3.2–7.0 mm long, sparsely glandular; filaments slightly curved, anther thecae 1/3 the size of the filament; ovary globose, some glands present, style very short, stigma lower lip semicircular, upper lip absent. Capsule globose 2–3mm diam., calyx lobe not embracing the stigma lips. Seeds obliquely ovoid, c. 0.35 mm long 0.15 mm wide.

Distribution and ecology:—Widely distributed across northern South America, from the Brazilian Northeast coast (i.e. restingas and tabuleiro ecoregions) ( Fig.1A,C ), through to the Guiana Shield ( Brazil, Guyana; French Guiana; Surinam, Venezuela), Amazonia lowland savannahs ( Brazil and Peru), and on Trinidad and Tobago ( Taylor 1989). Grows in water-soaked sand or clay substrate. On the Brazilian Northeast coast, it is distributed across the states of Bahia, from Maraú Peninsula’s restinga and further north; Sergipe, from Itabaiana mountain and coastal restinga; Paraíba, at Guaribas Biological Reserve; Rio Grande do Norte, Piauí and Maranh„o. In North Brazil it is found in the states of Pará (at Restinga or Mangrove), Amapá, Amazonas, and Roraima (Viruá National Park to Tapequém Mountain at northwest of Roraima state). In Venezuela it is found at Gran Sabana and some Tepuis, Guyana to French Guiana; and far west of Amazonia in Peru, a savannah site of Santuario Nacional de las Pampas del Heath. Flowering all year round, specially from Jun to Oct. Refer to species examined.

Etymology:—From the Greek amethystos meaning violet, referring to the lilac corolla.

Conservation status: —Least Concern. Based on its broad distribution range (EOO = 7,693,931 km 2; AOO = 228 km 2) and frequent presence in protected/conservation areas ( Fig. 1 ), U. amethystina is here considered Least Concern under ( IUCN, 2012) categories and criteria.

Notes: —After Taylor’s (1989) monograph, U. amethystina was reported to be a polymorphic species. As a result, the whole U. section Foliosa became a large species complex, with all specimens not fitting the narrowly circumscribed U. tricolor and U. tridentata being placed in a broadly circumscribed U. amethystina by exclusion. Utricularia amethystina s.str., as defined here, is the most common and widespread species of the complex in northern South America. The common morphotype from the Brazilian Cerrado domain, with deeply 3-lobed lower corolla lip, is here recognized as U. damazioi . A typical character that differentiates U. amethystina from the others is the lower lip of corolla being slightly 3-lobed, with half to 2/3 of corolla lower lip base swollen and given the name gibba; and the leaves usually forming a rosette. For some, U. amethystina s.str. used to be mistaken as a stunted form of U. tricolor , a similarity that is supported by their closer position in the phylogenetic hypothesis of Baleeiro et al. (2019).

As described by E. Ule (n. 8757) in the original publication of U. spatulifolia from Mt. Roraima’s lower campos (1600 m) “ The species is related to U. roraimensis N.E.Br. , but it is distinguished by its larger growth, longer spur etc. ”. However, here we list this name as synonym of U. amethystina s.str. and not U. roraimensis , due to morphological affinities with the former. Besides that, the collections of Steyemark 59648 (F, NY), 59742 ( US), 60154 ( US) under the name U. spatulifolia and original from Ptari Tepui is here designated as U. amethystina . Future molecular studies of well-sampled populations from across the range might shed light and provide more evidence for elevating these populations to associated names under synonymy or new names. As mentioned before, studies suggest that the tepuis from the Guiana Shield harbour high levels of endemicity and that these regions are still poorly explored for many group of plants.

The USA population collected on 14 Jan 1946 in Florida (L.J.Brass 15873, FSU 469) is described to have pale yellow flowers, but due to lack of sufficient material and since its natural population hasn’t been re-collected, this could not be placed under one of the names in the current monograph and the name U. standleyae remains as synonym. The Mexican population from Guerrero have yellow or white flowers and is found in savanna habitats around 1230m above sea level ( Rendón and Ruiz, 2017). These populations are shown with a star in the map of distribution of species ( Fig.1 ).

Specimens Examined: — BRAZIL. BAHIA: Camaçari, km 11 da estrada Salvador-Camaçari-Dias d`Avila, em grande área brejosa a direita, 1 Jul. 2002, Rivadavia 1458 ( SPF) . Ituberá, encostas de morros com nascentes rodeando braço de mangue, a direita do km 15 da estrada p/ a praia de Pratigi , 8 Out. 2005, Rivadavia 2109 ( SPF) . Maraú , ca. 8km na estrada Maraú/Ubaitaba, 4 Set. 1999, Carvalho et al. 6768 ( NY!, CEPEC!) ; ibid, 22 Jun. 2015, Baleeiro 384 ( SPF) ; ibid, estrada sentido Itacaré/Maraú, depois da travessia do Rio de Contas , ca. 2 km, restinga arbórea, 7 Ago. 2005, Paixão et al 560 ( CEPEC) ; ibid, estrada de Ubaitaba para a Barra Grande, na bifurcaç„o para Itacaré, 8 Out. 2005, Rivadavia 2115 ( SPF) ; ibid, in restinga area, 3.8km N of the Rio de Contas at Itacaré ferry crossing on road to Maraú, 05 Aug. 2002, W. W . Thomas et al. 13121 ( CEPEC). Mata de S „o Jo„o, brejo a direita no km 94 da rodovia do coco ( BA099 ), próximo a praia de Massarandupio, 18 Jan. 2003, Rivadavia 1523 ( SPF) . Santa Luzia , km 31 da estrada Camaca-Canavieiras, a 10 km de Hermilandia, 26 Mar. 2005, Rivadavia 1971 ( SPF) . MARANHÃO: Barreirinhas , restinga alagada próximo a Lagoa Azul, Lençóis Maranhenses, 30 Mar. 2007, Rivadavia 2499 ( SPF) . PARÁ: Maracan „, Ilha de Maiandeua , 13 Jun. 1994, M. N . Bastos et al. 1688 ( MG). Marapanim, estrada de Maruda para Praia do Crispim , 7 Ago. 2005, Rivadavia 2100 ( SPF). Santarém, estrada Santarém-Alter do Ch „o, em brejo com buritis ao lado direito da estrada, 4 Mar. 2000, Rivadavia & Pozza 1194 ( SPF). Vigia, km 16 da estrada de Vigia para S „o Caetano de Odivelas , 7 Ago. 2005, Rivadavia 2104 ( SPF) . PARAÍBA: Mamanguape, Reserva Biológica Guaribas , área 1 Capim Azul, 187 m, 19 ago. 2002, A. C . Sevilha & G . Pereira-Silva 2250 ( CEN); ibid. Reserva Biológica Guaribas, área I - Capim Azul , 22 Jul. 2012, Silva 4 ( JPB) . RORAIMA: estrada Porto Velho-Cuiabá, BR 364 , km 159, margem esquerda, vegetaç„o rupestre, 40m mais elevada que o resto da área, 9 Fev. 1983, Teixeira et al. 1400 ( INPA) . Caracaraí, ManausCaracaraí Road, km 130, Igarapé Lages, swampy ground on rocks, 16 May 1974, Prance et al. 21041 ( NY!, INPA!). Parque Nacional Viruá, na grade da PPBIO, L1 / N6 , campina, 12 Set. 2010, Costa & Barbosa 695 ( INPA) ; ibid, Estrada Perdida, 22 Jul. 2010, Cavalcati et al 187 ( INPA) ; ibid, campinarana gramíneo-lenhosa, com muitas Malpighiaceae , Clusiaceae , área alagável com solo arenoso, nas porçıes mais secas, areia branca, 12 Sep. 2010, El Ottra et al. 65 ( INPA) ; ibid, grade da PPBIO L3-500 , 04 Sep. 2012, Baleeiro 231 ( SPF) ; ibid, estrada Perdida, próximo à guarita lado direito em direç„o à primeira bueira, 06 Sep. 2012, Baleeiro 249, 250, 256 ( SPF). Boa Vista, Zona Urbana, 07 Sep. 2012, Baleeiro 263 ( SPF) ; ibid, beira da estrada em igarapé do lado direito entre Boa Vista e entrada para Tepequém, 08 Sep. 2012, Baleeiro 268 ( SPF). Serra do Tepequém , atrás da pousada da Gisele, 09 Sep. 2012, Baleeiro & R . Perdiz 270 ( SPF) . SERGIPE: Areia Branca, E . Ecológica Serra de Itabaiana , 16 Sep. 1995, Landim et al. 670 ( UEFS); ibid, 28 Sep. 2013, Carregosa-Silva & E . Santos 501 e 502 ( ASE). Pirambu, povoado samambaia, 09 Sep. 2013, CarregosaSilva et al. 500 ( ASE) .

COLOMBIA. Upper Apaporis basin, Macaya river, Mount Chiribiquete, 24 Jul. 1943, Schultes 5618 ( US). Cerro Castilho, 27 Jul. 1943, Schultes 5668 ( US). Rio Kuduyari, Cerro Yaapoboda, 5–6 Oct. 1951, Schults & Cabrera 14378 ( US). Rio Negro, San Felipe and vicinity (below confluence of Rio Guainia and Rio Casiquiare), 22 Oct. 1952, Schultes et al. 18131 ( US). Rio Kubiy’u (tributary of Rio Vaupes), Cerro Kanenda, savannahs about 15 miles upstream from mouth. Quarzite base, 10 Nov. 1952, Schultes & Cabrera 18391 ( US). Rio Kuduyari (tributary of Rio Vaupes), Yapoboda, 18 November 1952, Schultes et al. 18542 ( US). Rio Parana Pichuna (tributary of Rio Vaupes), Jun. 1953, Schultes & Cabrera 19957 ( US).

FRENCH GUIANA. Wilhelmina Gebergte, Lucie Rivier, 2 km of the river, 2 km west of Oost Rivier, 225 m, 11 Jul. 1963, Maguire et al. 54135 ( US).

GUYANA. Cuyuni-Mazaruni, Imbaimadai, Savannah between Imbaimadai and Partang River crossing, 570 m, 14 Dec. 1989, Gillespie , L. J . & D.R. Smart 2667 ( US). Potaro-Siparuni, Kaietuer Falls National park, near airstrip, 500 m, 2 Apr. 1988, Hahn , W., E . Judziewicz, S . Taiwari, & D. Gopaul. 4127 ( US). Ttrail from abandoned balata bleeders’ camp at base of Mt. Makarapan to Rupununi river , 20 Apr. 1988, Maas , P. J. M ., Koek-N, J., Lall, H ., Ter Welle, B. J. H . & Westra L. Y .Th. 7567 (INPA).

PERU. MADRE DE DIOS: Rio Heath, Santuario nacional de las Pampas del Heath , 15 Jun. 1992, Castilho & Foster 6913 ( F) .

SURINAME. East Creek branch of Lisa creek, savana like, sandy soil, 26 Jun. 1998, Lohmann, L.G. & Peckham H. 153 (MO) . Tafelberg ( Table Mountain ), shallow bogs savannah 1, 3 Aug. 1944, Maguire, Basset 24197 ( F) ; ibid, 5 km above falls, 525 m, 13 Sept. 1944, Basset Maguire 24727 ( F) . Lucie Rivier, 2 km north of Rivier, 225 m, 11 Jul. 1963, Maguire et al. 54135 ( F) .

TRINIDAD AND TOBAGO. 1977–80, Fendler 551 ( NY). Long stretch road side, 26 Sept. 1951, Simmonds et al. 4 and 9 ( K) .

VENEZUELA. ESTADO BOLIVAR: Llanos between Cano Azul and rocky ridge 3 km. east cano Azul , 23 Oct. 1954, Wurdack & Guppy 167 ( K) . Vicinity of Salto Pacairao, bordering Rio Pacairao, about 3 km east to northeast off Santa Teresita de Kavanayen , 24 Nov. 1944, Steyemark 60483 ( F) . by Ven. 10, 8 km N of San Rafael just S of San Juan de Camoran, gravelly sandy seep in wet savanna, 29 Jul. 1983, Kral & Gonzales 70588 ( MO) . Piar, Gran Sabana. 25 km N de Kama-meru, Carretera El Dorado-Sta. Elena , 4 Abr. 1985, Holst et al. 2179 ( MO) . Kanavayen, estrada para Liworiwo, Salto Aponwao , em campos rupestres ao lado esquerdo da estrada, já próximo de Liworiwo , 20 Jan. 1999, Rivadavia & HoogenstriJd 815 ( SPF) . Luepa, em campos rupestres brejosos cheios de Stegolepis sp. , beirando o km 145 da estrada El Dorado - Sta. Elena , 21 Jan. 1999, Rivadavia & HoogenstriJd 833 ( 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 2. Utricularia amethystina. A. Habit showing leaves, stolons, bladder traps and rhizoids, B. Inflorescence, C. Calyx frontal view, D. Corolla frontal view, E. Corolla rear view, F. Bract and bracteoles, G. Bladder trap, H. Capsule, I. Stamens, J. Stamens and ovary showing dorsal stigma surface, K. Seeds.

FIGURE 3. A. U. amethystina s.str. Type locality Marau peninsula Bahia, B. U. amethystina s.str. Virua National Park, C. U. amethystina s.str. white-flowered specimen, D. U. roraimensis, E. U. amethystina s.str. pink morphotype, F. U. amethystina Type locality, G. U. lindmanii, H–I. U. lunaris, J–N. U. damazioi. J. from Chapada Diamantina, K., M., N. from Chapada dos Veadeiros, L. from Diamantina Plateau.