Victoria amazonica (Poepp.) Klotzsch

Smith, Lucy T., Magdalena, Carlos, Przelomska, Natalia A. S., Pérez-Escobar, Oscar A., Melgar-Gómez, Darío G., Beck, Stephan, Negrão, Raquel, Mian, Sahr, Leitch, Ilia J., Dodsworth, Steven, Maurin, Olivier, Ribero-Guardia, Gaston, Salazar, César D., Gutierrez-Sibauty, Gloria, Antonelli, Alexandre & Monro, Alexandre K., 2022, Revised Species Delimitation in the Giant Water Lily Genus Victoria (Nymphaeaceae) Confirms a New Species and Has Implications for Its Conservation, Frontiers in Plant Science 13, pp. 1-31 : 20-22

Victoria amazonica (Poepp.) Klotzsch ,

Bot. Zeitung (Berlin) 5: 245 (1847). Euryale amazonica Poepp. Froriep’s Not. Natur- Heilk. 35: 131 (1832).

Type: Poeppig s.n. (holotype W -presumed destroyed in WWII); Brazil, Amazonas, Careiro da Várzea [ Teresina ], Ilha de Careiro , 25 Sept. 1974, G.T. Prance 22745 (neotype (selected by de Lima et al., 2021): INPA ( INPA 46745 ); isoneotypes: K ( K 000837777 !), NY ( NY 2269910 , NY 2269911 , NY 2269928 ) , MO ( MO 3414212 ) , US ( US 01341606 )) .

Vernacular names: Forno de Jaçanã, Auapé yapóna, Victoria regia , Giant Amazonian Waterlily. Figures 1A , 2 , 3A–C , 4 , 9 .

Victoria regina R.H.Schomb. Athenaeum (London) 515: 661 (September 9, 1837).

Victoria regia Lindl., Monograph : 3 (October 16, 1837). Type: Victoria Regia: 3, Plate 1 (October 16, 1837). nom. superfl.

Victoria regina J. E. Gray, Mag. Zool. and Bot. 2(11): 440 (December 1, 1837). nom. superfl. Victoria reginae Hook. Hooker’s J. Bot. Kew Gard. Misc. 2: 314 (1850). orth. var.

Leaves up to 2.3 m broad, adaxial surface of lamina green, occasionally tinged bronze in younger leaves; abaxial surface of lamina maroon or green, radial and reticulate ribs maroon, yellow or green; leaf margins form a low to moderate rim c. 4–7% of the lamina length (higher in crowded habitats), rim curved at its base then ± perpendicular to adaxial surface, abaxial surface of rim maroon or green; hairs 0.3–12 mm, simple, multicellular, 3–12 segmented. Flowers up to 28 cm in diameter at second-night anthesis. Ovary 8–12 cm diameter, outer surface covered in prickles 1– 18 mm (dried), prickles gradually tapering to a sharp point, hairs absent or present, where present simple, 0.1–0.4 mm, inner surface of ovary with deeply concave stigmatic surface, rounded to triangular in longitudinal profile, ridged with lines corresponding with 25–36 radially arranged locules, each containing 25–28 ovules, 1–1.5 mm diameter (fresh). Outer tepals 4.9–12 × 4–8 cm when fresh; abaxial surface predominantly brown/maroon, bearing 55–330 prickles per tepal, prickles tapering gradually to a sharp point, ranging from 1–14 mm (dried), spaced regularly, irregularly, or clustering more densely toward the base over entire surface, hairs absent or present on abaxial surface, where present 0.1–0.2 mm. Inner tepals 7–15 × 2–6 cm (fresh), innermost deep maroon in bud; all others remaining white or turning pink to dark pink at second-night anthesis; outer staminodia> 25, 5–6 × 1–1.5 cm thick, rigid, apiculate; stamens 2–4 × 0.5–1 cm; inner staminodia, 4–6 × 0.5–1 cm; base of lower parts of carpellary appendage auriculate/rounded in shape and hanging free from extension of stigmatic surface, length of upper parts not exceeding that of lower parts. Flower at first night of anthesis, inner tepals white, with innermost tepals dark maroon, outer staminodia tipped pink; second night anthesis, innermost tepals dark maroon, inner tepals remaining white or pink to dark pink or red, darkest at base, outer staminodia remaining white or dark pink for basal two thirds of their length, tipped pink, inner staminodia pink at base. Seeds 600–1000 per fruit, 7–8 × 9–10 mm, ellipsoid, green to brown, raphe faintly visible.

Distribution and Conservation Status — Victoria amazonica is restricted to the Amazon river basin, from Northern Brazil, Bolivia, Colombia, Guyana and Peru. Its EOO is estimated to be 2,640,795 km 2, exceeding the threshold for an IUCN threat category under criterion B, whilst its AOO is estimated as 476 km 2, falling into the Endangered category. We believe that our calculation of the AOO is likely an underestimate, resulting from difficulties in observing the species in the field and under-representation of the genus in biological collections.

There are more than 10 locations for which threats have been assessed, but there is not the information on population fragmentation or fluctuation available in order to be able to assess the ‘severely fragmented’ and ‘extreme fluctuations’ subcriteria. A continuing decline in habitat quality is inferred due to the presence of hydroelectric dams, mining, and deforestation of the river systems from where V. amazonica is documented. For example, in Peru, localities along the Marañón river to the headwaters of the Amazon and the Ucayali river have been heavily deforested by gold mining ( i-Terra, 2021). Gold mining is associated with profound mercury contamination of rivers and aquatic species ( USGS Environmental Health Program, 2019) and increases in Peruvian mercury imports suggest that contamination must be increasing ( Swenson et al., 2011). In Brazil, as in Bolivia there have also been reports of mining activities in indigenous lands adjacent to several Victoria populations ( Hutukara Associação Yanomami and Associação Wanasseduume Ye’kwana, 2020; INPE, 2021; MapBiomas, 2021; Mercado, 2021).

Victoria amazonica is here assessed as Least Concern (LC) considering that its range stretches across Amazonia and currently exceeds the parameters for a threatened category under criterion B. We note, however, a moderate number of locations where populations are under threat and there is a continuing decline in habitat quality. Further investigation and surveys are needed to better understand trends in population size, fragmentation, distribution and the impact of climate change.

Notes. Victoria amazonica is the only species whose leaves do not always form an upturned rim, and when they do, it is usually low and vertical in profile rather than recurving over the flat part of the lamina. Its flowers are distinguished from V. cruziana and V. boliviana both in bud and on first-night opening, as the innermost tepals are dark maroon rather than white. Carpellary appendages are curved at the base of the lower part and hang freely away from the attachment point. The prickles covering both outer tepal abaxial surface and outer ovary are uniquely gradually tapering to a sharp point (not abruptly tapering as in other species), and always cover the entire abaxial surface of the outer tepals. Its seeds are ellipsoid rather than globose.

Material Examined — BOLIVIA. Pando: Manuripi: pond north of Rio Madre de Dios ., –66.126667, –10.903611, 09/07/1997, Ritter, N., Crow, G. & Crow, C. 4170 ( LPB, MO). BRAZIL. ‘North Brasil’: 1898, Vaughan, G. 61 ( K). Acre: Río Moa, margem esquerda; lugar chamado Humaita , –72.895556, – 7.614444, 01/10/1984, Ferreira, C. A. 5123 ( NY). Amapá: Itaituba, Igarape no Rio Tapajos , – 55.961111, –4.229167, 16/12/2017, Brogim, R. 4 ( UPCB). Amazonas: Ipixuna, Margem do Rio Croa , – 72.556667, –7.745278, 15/02/2009, Quinet, A., Saraiva, B., Firmeza, T. 1582 ( K, SPF); Teresina, Ilha de Careiro , –59.81667, – 3.1, 25/09/1974, Prance, G. T. 22745 ( K, NY, US); Basin of Rio Purus area . Lago Preto , 3 km north of Labrea, – 64.813056, –7.229772, 29/10/1968, Prance, G. T., Ramos, J. F. and Farias, L. G. 8016 ( NY); Rio Solimões , south bank near Carreiro , –59.806667, –3.168889, 05/02/1974, Steward, W. C. And Ramos, J. F. P20211 ( K, NY, US); Rio Amazonas , from Manaus to 100 km lower reaches, –59.141944, –3.216111, 08/08/1987, Tsugaru, S. and Yotaro Sano B-769 ( MO, NY); Ilha do Cantagalo , –61.503889, –1.570833, 04/07/1995, Adalardo- Oliveira, A. 2645 ( NY, SPF); Riverside and small islets of Rio Solimões within 100 km upper-stream from Manaus, – 60.728056, 3.260278, 15/08/1987, Tsugaru, S. and Yotaro Sano B-1069 ( NY); Pará: Santarem, Igarape , Ilha Grande de Santarum , –54.706840, –2.450070, –/10/1849 and –/11/1849, Spruce, R. 441 and Spruce s.n., s.d., ( K, M, P) , – /04/1850( NY) 1849 ( P); Oriximina, Lago Uraria , SW of Orixima, across Rio Trombetas, – 55.9025, –1.812778, 11/06/1980, Davidson, C. and Martinelli, G. 10241 ( MO, NY, RB, US); Rio Cupari , Lago Curuca , 01/01/1948, Black, G. A. 48-2223 ( IAN); Rio Cupari , Lago Curuca , 02/01/1948, Black, G. A. 48-2253 ( IAN); Rio Cupari , Lago de Curuca , 02/01/1948, Black, G. A. 48-2254 ( IAN, NY, US); Oriximina, Rio Trombetas , Lago Ururia , 6 km SW de Oriximina, –55.910278, –1.803889, 08/06/1980, Martinelli, G. 6945 ( RB); Pacoval, Rio Curua , –55.083333, –1.833333, 6– 8/08/1981, Jangoux, J., and Riberio, B. G. S. 1647 ( NY); Santarem, 25/12/1938, Markgraf 3873 ( RB); Monte Alegre, Rio Gubatuba , proximo a vila Pare Sol, –54.043333, –2.011944, 17/07/2011, Lima, C. T. 503 ( HUEFS); Pacoval, Rio Curua , –55.083333, – 1.833333, 6–8/08/1981. Roraima: Rorainopolis, Rio Branco , Lago do Pirarucu , 25 km antes da boca com o Rio Negro, – 61.8525, –1.158333, 28/03/2012, Martinelli, G., Moraes, M. A., Benevides, P., Forzza, R. C., Nadruz, M., Gallucci, S., Costa, D. 17700 ( RB). COLOMBIA. Amazonas: Leticia, below Quebrada de Arara , –70.065278, –4.05944, 28/01 – 07/02 /1969, Plowman, T., Lockwood, T., Kennedy, H., Schultes, R. E. 2313 ( K). GUYANA. Berbice: Berbice, –58.2778, 4.394033, 1837, Schomburgk s.n. ( K). Upper Takutu-Upper Essequibo: Karanambo, Rupununi River , –59.3, –3.75, 27/09/1988, Maas, P. J. M., Koek-N, J., Lall, H., ter Welle, B. J. H., Westra, L. Y. 7727 ( K). PERU. Maynas: East of Puerto Alegria , –70.0625, –4.103611, 15/03/1977, Gentry, A. and Daly, D. 18351 ( MO); Isla Padre ( Cocha Paster ), –76.166667, – 3.75, 21/12/1982, Vasquez, R.., Grandez, C. and N. Jaramillo, N. 3684 ( MO); Padre Isla in Río Amazonas, and in the cato below Iquitos., –73.163611, –3.651944, 22/05/1978, Gentry, A., Jarmillo N. 22133 ( MO). See Supplementary Data.

Figures

FIGURE 1 | Wild populations of (A) Victoria amazonica (Peru, Loreto), (B) V. boliviana sp. nov. (Bolivia, Beni) and (C) V. cruziana (Argentina, Chaco). (Photo Credits: (A) Laurel Allen, iNaturalist ID 821386. (B) Carlos Magdalena. (C) Fernanda Alarcón, iNaturalist ID. 19516477).

FIGURE 2 | Flower morphology and terms, using a Victoria amazonica second-night flower in longitudinal section for reference (above) and a fully dissected flower (below). (A) Ovary, (B) stigmatic surface and stigmatic chamber, (C) locule and ovules, (D) floral apex, (E) outer tepals, (F) inner tepals, (G) outer staminodia, (H) stamens, (I) inner staminodia, (J) carpellary appendages. Illustration and photo: Lucy T. Smith.

FIGURE 3 | Line drawing comparing flower and seed morphology in: Victoria amazonica (A–C), V. boliviana sp. nov. (D–F), and V. cruziana (G–I); flower bud whole and in longitudinal section showing abaxial surface of outer tepals, bud profile, stigmatic chamber, carpellary appendage and prickle morphology; seeds. (cultivated at RBG Kew) Illustration: Lucy T Smith.

FIGURE 4 | Victoria amazonica (A) habit, (B) abaxial leaf, (C) leaf rim profiles, (D) bud, whole and LS, (E) flower prickles, (F) first night flower, (G) second night flower, (H) carpellary appendages and tepal, staminode attachments; (I) ovule, whole and LS, (J) stamens, (K) fruit, (L) seed. (All from material cultivated at RBG Kew). Illustration: Lucy T. Smith.

FIGURE 9 | Victoria amazonica (A) bud whole, (B) bud L.S., (C) bud from above, (D) first night flower, (E) first night flower L.S., (F) second night flower, (G) second night flower LTS, (H) habit, and (I) seed. (A–H) (LTS) and (I) (CM) cultivated RBG Kew.