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Aim The aim of this study was to further advance our understanding of the species‐rich, and ecologically important angiosperm family Combretaceae to provide new insights into their evolutionary history. We assessed phylogenetic relationships in the family using target capture data and produced a dated phylogenetic tree to assess fruit dispersal modes and patterns of distribution. Location Tropical and subtropical regions. Time Period Cretaceous to present. Major Taxa Studied Family Combretaceae is a member of the rosid clade and comprises 10 genera and more than 500 species, predominantly assigned to generaCombretumandTerminalia, and occurring on all continents and in a wide range of ecosystems. Methods We use a target capture approach and the Angiosperms353 universal probes to reconstruct a robust dated phylogenetic tree for the family. This phylogenetic framework, combined with seed dispersal traits, biome data and biogeographic ranges, allows the reconstruction of the biogeographical history of the group. Results Ancestral range reconstructions suggest a Gondwanan origin (Africa/South America), with several intercontinental dispersals within the family and few transitions between biomes. Relative abundance of fruit dispersal types differed by both continent and biome. However, intercontinental colonizations were only significantly enhanced by water dispersal (drift fruit), and there was no evidence that seed dispersal modes influenced biome shifts. Main Conclusions Our analysis reveals a paradox as drift fruit greatly enhanced dispersal distances at intercontinental scale but did not affect the strong biome conservatism observed.

Key to the Species 1. Mature leaves with upturned rim, rim moderate to high (8– 10% of blade length), sigmoid in cross- section; mature bud concave towards apex; carpellary appendages with a cuneate base, arising 45 ◦ from the point of attachment (see Figure 2); prickles associated with the flowers tapering abruptly to a point, covering ovary and either absent, or covering the basal 1/3 of abaxial surface of the outer tepal, prickles 0 – 100 per tepal; seeds globose, raphe faintly visible. Paraná river basin, lower course of Paraguay river V. cruziana (Figure 6) 1. Mature leaves with no upturned rim, or where present the upturned rim low to moderate (4–7% of blade length), sigmoid or vertical in cross-section; mature bud convex towards apex; carpellary appendages with an auriculate or subauriculate base, arising 45 ◦ from point of attachment or not (see Figure 3); prickles associated with the flowers tapering smoothly or abruptly to a point, covering both the ovary and entire abaxial surface of the outer tepal, or only the ovary, 0–1000 prickles per tepal; seeds ellipsoid or globose, raphe faintly visible or prominent. Amazon river basin, Pantanal 2 2. Mature leaves with no or moderate upturned rim, which, where present, is vertical in cross-section. Prickles associated with the flowers tapering smoothly to a point and covering both ovary and entire abaxial surface of the outer tepals, 55–300 prickles per tepal; stigmatic chamber deeply concave, obdeltate in longitudinal profile, carpellary appendage auriculate and hanging free from, not arising 45 ◦ from point of attachment (see Figures 2, 3); seeds ellipsoid, the raphe faintly visible. Amazon river basin excluding V. amazonica (Figure 4) 2. Mature leaves with a moderate upturned rim, which is sigmoid in cross-section. Prickles associated with the flowers tapering abruptly to a point, covering both the ovary and entire abaxial surface of the outer tepals, or only the ovary, 0–1000 prickles per tepal; stigmatic chamber shallowly concave and oblong in longitudinal profile, carpellary appendage subauriculate at point of attachment not arising 45 ◦ from point of attachment (see Figure 3) (not known for V. cruziana f. mattogrossensis); seed ovoid, the raphe prominent. Llanos de Moxos or Pantanal 3 3. Prickles associated with the flowers covering the ovary and either absent from or sparsely distributed over the abaxial surface of the outer tepals, 0–10 per tepal; apical portion of the carpellary appendage longer than the basal portion. Llanos de Moxos V. boliviana (Figure 5) 3. Prickles associated with the flowers covering the ovary and distributed densely and evenly over the abaxial surface of the outer tepals, 500–1000+ per tepal; apical portion of the carpellary appendage shorter than the basal portion. Pantanal V. cruziana f. mattogrossensis taxon incertum taxon incertae Victoria R. H. Schomb., Athenaeum (London) 1837 (No. 515): 661 (September 9 1837). Victoria Lindl., Monog. 3 (October 16 1837). Victoria J. E. Gray, Mag. Zool. Bot. 2(10): 373 (December 1 1837). Aquatic perennial herb, rhizome erect, tuberous, elongate to cylindrical, roots adventitious. Leaves floating, orbicular, peltate, perforated by stomatodes, adaxial surface of lamina glabrous, lacking prickles, green; abaxial surface of lamina with prominent radial and reticulate ribs, juvenile leaves sagittate; leaf margins flat or upturned; prickles covering petiole and ribs. Inflorescences uniflorate, bracteate. Flowers axillary, solitary, multiple buds per plant; pedicel with 4 primary air chambers, 8 minor chambers, covered in prickles; flowers opening one at a time, projecting above water surface shortly before anthesis, projecting above or resting on water surface at anthesis, each flower opening over two nights and partially closing in between, protogynous. Epigynous, ovary globose, covered in prickles externally, ovules parietal, attached by short funiculi, globose. Outer tepals 4, triangular, apex acute to rounded. Inner tepals 40–c.100, arranged in spiral series, creating a torus (attachment point of tepals forming a ring of tissue), tepals gradually reducing in size towards the center and changing shape from apically rounded to acute from outer to innermost; outer staminodia in 1 or 2 whorls, thick, rigid, apiculate; stamens> 100, borne in c. 3 series, subulate, introrse; anthers linear-elongate; inner staminodia,>50, sigmoid, subulate, partially adnate to carpellary appendages, detaching at second-night anthesis; carpellary appendages L-shaped, arising from extension of stigmatic surface, lower parts adnate to tissue extending from tepal base attachment, corresponding in position and number with stigmatic surface ridges and locules. Fruit ripening just below surface of water, 10–15 cm in diameter (excluding prickles) at maturity, fleshy, oblate, topped by a shallow cylinder-shaped mass of dark reddish to maroon ring of persistent hard tissue formed by the remnant bases of tepals; inner staminodia persistent and curved over concave stigmatic surface while ripening; outer layers of pericarp disintegrating to release seeds. Seeds smooth, surrounded by a mucilaginous aril. Three, possibly four species, tropical and temperate South America., Published as part of 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, pp. 1-31 in Frontiers in Plant Science 13 on pages 19-20, DOI: 10.3389/fpls.2022.883151, http://zenodo.org/record/7576624

Victoria cruziana Orb., Ann. Sci. Nat., Bot. , sér. 2, 13: 57 (January 1840). Type: Bolivia [Argentina], Corrientes, banks of the Paraná river, Arroyo de San José, beginning of 1827, d’Orbigny s.n. (lectotype: P (P02048598 ∗) (designated by de Lima etal., 2021); isolectotypes: P (P02048599). Vernacular names: Irupé (yrupé), yacare yrupé, naanók lapotó (poncho del otany), maíz de agua, Santa Cruz Waterlily, Victoria regia. Figures 1C, 3G–I, 6, 11. Victoria regia var. cruziana (Orb.) G. Lawson, Proc. & Trans. Roy. Soc. Canada 6(4): 109 (1889) Euryale brasiliana Steud., Nomencl. Bot. [Steudel], ed. 2. 1: 617 (November 1840). Euryale policantha Rojas Acosta, Cat. Hist. Nat. Corrientes 65 (1897). Euryale bonplandia Rojas Acosta, Cat. Hist. Nat. Corrientes 151 (1897). Victoria cruziana f. trickeri Henkel ex Malme, Acta Horti Berg. 4(5): 12. 1907 (“Trickeri”), nom. nud. Leaves up 2.4 m broad, adaxial surface of lamina green, abaxial surface of lamina green or dark blue-green, radial and reticulate ribs yellow or green; leaf margins form a high rim 8–10% of leaf length, rim ± perpendicular to or slightly recurved over adaxial surface at base, flared outwards at top (sigmoid in profile), abaxial surface of rim green or tinged maroon, hairs 1–3 mm, simple, multicellular, 10–15 segmented. Flower bud broadly ovoid, concave just before apex, up to 30 cm diameter at secondnight anthesis. Ovary 7–10 cm diameter, outer surface covered in prickles 1–22 mm (dried), prickles abruptly tapering from c. half their length to sharp apex; hairs absent or present, where present 0.1–12 mm; inner surface of ovary with moderately concave stigmatic surface, rounded to triangular in longitudinal profile, ridged with lines corresponding with 25–38 radially arranged locules, each containing 20–25 ovules 1.5–1.8 mm (fresh). Outer tepals 4, 10–13 × 4–9 cm when fresh, abaxial surface green and/or tinged maroon abaxially prickles absent or present, where present up to 100 per tepal, prickles tapering abruptly at their midpoint to a sharp point, 1–10 mm (dried), distributed up to lower one third of surface, hairs absent or present, where present 0.1–1 mm. Inner tepals 7–10 × 1.5–9 cm (fresh), innermost all white both in bud and during first-night anthesis, crinkled in appearance, turning pale to dark pink on second-night; outer staminodia, 6– 7 × 1–1.5 cm, thick, rigid, apiculate; stamens 4–6 × 0.5–1 cm inner staminodia> 50, 4–6 × 0.5 cm; base of lower parts of carpellary appendage flat, arising from stigmatic surface at 45 degree angle, cuneate, length of upper parts not exceeding that of lower parts. Flower at first night of anthesis: all inner tepals white, outer staminodia tipped pink; at second night anthesis, outer tepal adaxial surface pink, inner tepals pale or dark pink at base, white or pink towards apex, outer staminodia dark pink for basal two-thirds of their length, white then pink towards apex, inner staminodia pink at base. Seeds, c. up to 1000 per fruit, 7–9 × 8–10 mm, globose, raphe faintly visible, brown to black, surrounded by a mucilaginous aril. Distribution and Conservation Status Victoria cruziana f. mattogrossensis taxon incertum has hitherto been included within Victoria cruziana until this study. Because our genomic analyses are limited with respect to the rank of this taxon and of its relationship to the other species, we have not considered forma mattogrossensis taxon incertum as conspecific with V. cruziana for the purposes of an extinction risk assessment. Victoria cruziana is restricted to the Paraná river basin and tributaries, from Paraguay to Argentina, and possibly Bolivia. Based on the maximum potential habitat of wetlands (including Esteros de Ibera National Park Wetlands) and a combination of verified herbarium collections and iNaturalist images we estimate the EOO of V. cruziana to be between 46,563 and 132,945 km 2. Thisexceeds the threshold fora threatened category under criterion B (IUCN Standards and Petitions Committee, 2019). We calculate the AOO to be 120 km 2, although an upper estimate based on the extent of the river may exceed 2,000 km 2 (but not more than 3,000 km 2). This would assess V. cruziana as between Endangered and Vulnerable categories. There are more than 10 locations but no information about population fragmentation. We infer a continuing decline in habitat quality due to the increasing frequency of droughts, the abstraction of water, deforestation (Caivano and Calatrava, 2021; Comisión Nacional de Actividades Espaciales, 2021) and big hydroelectric dams. For example, Itaipu is one of the largest dams in the world (Stevaux et al., 2009) and lies within the V. cruziana range. Whilst the AOO upper estimate is close to the threshold for VU, there are more than sufficient criteria for a threatened category under criterion B, and we therefore assess V. cruziana as LC. We recommend further documentation of the distribution and size of V cruziana populations and investigations into their fluctuation through time as we believe that it may be vulnerable to an increase in the frequency and severity of droughts associated with climate change and increased sedimentation caused by the construction of large dams within its habitat (Stevaux et al., 2009). Notes. Victoria cruziana forms the proportionately highest leaf rims of all the species, and these are always slightly recurved over the flat part of the lamina, flaring out at the top. The concavity of the outer tepals before their apex gives the bud a pinched-in appearance. Prickles are absent or occur on the outer tepal abaxial surface, but only up to one-third of their length from the base. In this species, hairs which are sometimes present on the lower outer tepal abaxial surface and ovary are the only ones large enough to see without magnification. At first night anthesis, inner tepals have a crinkled appearance. Victoria cruziana f. mattogrossensis taxon incertum was described by Malme based on material present in the spirit collection of the Swedish Museum of Natural History comprising two jars of the same gathering. S07-84 comprises a longitudinal section through the ovary and perianth showing clearly the distribution of prickles and their form on both the ovary and outer tepals, also the morphology of the carpellary appendages; S07-85 comprises sections of the petiole apex and either the petiole or pedicel. Both are preserved in excellent condition. S07-84 was selected as lectotype as it displayed a greater number of diagnostic morphological features. Victoria cruziana f. mattogrossensis taxon incertum was described from, and material corresponding to it has only been observed from the Pantanal, in Bolivia, Brazil and Paraguay in the Uruguay river basin. Phylogenomic data were unable to confirm whether the material sampled represents a distinct evolutionary lineage (see section “Discussion”). It may be that further sampling and research supports the recognition of this name as a distinct taxon. Material Examined. ARGENTINA. Chaco: 1st De Mayo, Laguna en Chacra, al lado del arroyo Ine, –58.849444, – 27.416389, 02/03/2006, Mulgura de Romero, M. E., Anisko, T., Harbage, J., Illarrage, H. 4249 (SI); San Fernando, Entre Barranqueras e Isla Antequera, –58.87944, –27.416389, 18/03/1967, Krapovickas, A. and Cristobal, C. L. 12752 (MO). Corrientes: Capital, Riachuelo, off Ruta 12 ca 17 km Sof Corrientes, –58.749444, –27.554444, 06/04/1982, Schinini, A. Wiersema, J. H. 2243 (MO); Esquina, Isla Correntina frente a curuzu – Chali, en el Paraná medio, –59.630833, –30.341944, 10/04/1968, Burkart, A., Troncoso, N. S., Guaglianone, E. R. and Palacios, R. A. 26963 (SI); San Roque, R. Santa Lucia, – 58.738056, –28.576944, 27/02/1957, Pedersen, T. M. 4486 (K, MO); Bella Vista, Cruce Ruta Nacional 12 Ey Puente Sobre Lel Río Santa Lucia, –58.72, –28.57, 12/04/2008, Mulgura de Romero, M. E., Aniśko, T., Belgrana, M. J. and Harbage, J. 4474 (SI); Dep. Esquina, Río Guayguiraro, –59.56, –30.374444, 26/02/1974, Quarin, C., Schinini, A. Gonzales, J. M., Ishikawa, A. 2196 (K). Santa Fe: La Capital, Ruta Nacional 168 y Puente no. 9, Sobre Arroyo mini, al E-SE de la Ruta Prov. 1. W, – 60.575833, –31.672778, 12/04/2008, Mulgura de Romero, M. E., Anisko, T., Belgrana, M. J. and Harbage, J. 4477 (SI). BOLIVIA. Santa Cruz: Angel Sandoval, channel at southern end of Laguna Mandiore, –57.483333, –18.216667, 16/07/1998, Ritter, N., Crow, G. E., Garvizu, M. and Crow, C. 4562 (LPB, MO) [f. mattogrossensis], Ritter, N., Crow, G. E., Garvizu, M. and Crow. C. 4560 (LPB, MO). BRAZIL. Mato Grosso du Sul: Corumba, Cacimba da Saude, – 57.664167, –18.99833, 13/12/2002, Avellar, A. L. F. 13 (COR) [f. Mattogrossensis]; Ladario, terminal da Branave, no porto de Ladario, –57.584722, –19.0225, 12/08/1994, Sanches, A. L., Bortolotto, J. M., Damascenos Jr., G. 44 (COR) [f. mattogrossensis]; Corumba, Ladario, CODRASA Brejo (swamp), – 57.516389, –19.021111, 27/11/2004, Souza Jr., A. F., and Siqueira, C. S. 39 (COR) [f. Mattogrossensis]; Corumba, Ladario, –57.580556, –19.001111, –/07/1894, Anon s.n. [G. O. A. Malme] S07-785, S07-784 (S). PARAGUAY. Distrito Capital: L’Assumption [in the swamps], –57.604167, –25.256667, 17/03/1875, Balansa, R. 523 (K, P). Central: Piquete Cue, –57.666667, –25.166667, 29/11/2000, Zardini, E. M. and Guerrero, L. 55181 (MO); Bay of Ascuncion, wetlands, –57.61922, –25.273333, 15/03/1988, Ericsson, K. 577 (MO). Neembucu: Yataity, –58.040556, –26.788611, –/03/1975, Walter, M. A. 86 (K); Pilar Garden Club S, –58.276389, –26.867778, 28/01/2005, De Egea Juvinel, J., Pena-Chocarro, M.., Vera, M., Torres, M.. and Elsam, R. 738 (MO). Presidente Hayes: Riacho Pucu., – 57.083333, –24.666667, 20/08/2000, Zardini, E. M and Guerrero, L. 54765 (MO); 12/02/1987, Sparre 2363/51 (P)., Published as part of 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, pp. 1-31 in Frontiers in Plant Science 13 on pages 25-26, DOI: 10.3389/fpls.2022.883151, http://zenodo.org/record/7576624, {"references":["de Lima, C. T., Machado, I. C., and Giulietti, A. M. (2021). Nymphaeaceae of Brasil. Sitientibus serie Ciencias Biologicas 21. doi: 10.13102 / scb 4986","Comision Nacional de Actividades Espaciales (2021). Los Satelites Argentinos SAOCOM Monitorean la Bajante del Rio Parana. Ministerio de Ciencia, Tecnologia e Innovacion. Buenos Aires, AC: Comision Nacional de Actividades Espaciales.","Stevaux, J. C., Martins, D. P., and Meurer, M. (2009). Changes in a large regulated tropical river: the Parana River downstream from the Porto Primavera Dam, Brazil. Geomorphology 113, 230 - 238. doi: 10.1016 / j. geomorph. 2009.03.015"]}

Victoria boliviana Magdalena and L. T. Sm., sp. nov. Type: Bolivia, Beni Department, Provincia Ballivían, subiendo el Río Yacuma desde Puerto Espíritu, laguna en conexíon al Río Yacuma, unos 20 m al N, 29 Mar. 1988, S. G. Beck 15173 (holotype: LPB; isotype: K (K000798309). Vernacular names: Reina Victoria, Victoria regia. Figures 1B, 3D–F, 5, 10. Most similar to V. cruziana Orb., from which it can be distinguished by the lower rim of the floating leaf, convex apex of the flower bud, length of the upper part of the carpellary appendages exceeding that of the lower part and the larger seeds. The V. boliviana plastid genome differs from that of other Victoria species by a 14 bp insertion between plastid genes ndhC and trnV in the large single copy region (LSC), a 5 bp deletion between trnK and rps16, a 7 bp deletion adjacent to trnC in the LSC and a 42b p deletion in the CDS of gene ycf1, within the SSC. Finally, a 4 bp transversion unique to V. boliviana sp. nov. was found inthe LSC. Leaves up to 3.2 m broad, adaxial surface of lamina green; abaxial surface of lamina dark green, maroon or dark-blue, radial and reticulate ribs yellow or green; leaf margins upturned to form a moderate rim c. 4–7% of leaf length, rim recurving strongly over blade surface at base and curving inwards or flared outwards at top, abaxial surface of rim deep maroon or very pale green/white in color, glabrous or with hairs, where present 1.2–3 mm, simple, multicellular, 6–15 segmented. Flower bud broadly ovoid, convex at apex, up to 36 cm in diameter at second-night anthesis. Ovary 8–10 cm diameter, outer surface covered in prickles, 1–10 mm (dried) glabrous; prickles abruptly tapering from c. half of length to sharp apex; inner surface of ovary with shallowly concave stigmatic surface, oblong in longitudinal profile, ridged with lines corresponding with 25–36 radially arranged locules, each containing 8–14 ovules, 2–2.5 mm diameter (fresh). Outer tepals 4, 10–15 × 8–10 cm when fresh, abaxial surface predominantly green, or tinged maroon, prickles absent or present, where present up to 10 per tepal, prickles tapering abruptly at their midpoint to a sharp point, 1–10 (dried), distributed irregularly over entire surface, glabrous. Inner tepals 6–15 × 1.5–9 mm (fresh), innermost remaining white or turning pale pink at their base at the second-night anthesis; outer staminodia> 50,3– 4 × 0.5– cm, thick, rigid, apiculate; stamens, 4–5 × 0.5–1 cm; inner staminodia 4– 5 × 0.5–0.7 base of lower parts of carpellary appendage angular in shape and arising at 45 degree angle from stigmatic surface, length of upper parts exceeding that of lower parts. Flower at first night of anthesis, inner tepals white, outer staminodia tipped blue-violet; second night anthesis, inner tepal adaxial surface pink, inner tepals pale pink at base, white or pink towards the apex, outer staminodia dark pink for basal twothirds of their length, white then violet towards the apex, inner staminodia pink at base. Seeds c. 300 per fruit, 12–13 × 16– 17 mm, globose with a prominent raphe (especially when dry), dark brown to black, surrounded by a mucilaginous aril. Distribution and Conservation status Victoria boliviana Magdalena and L. T. Sm. is restricted to Bolivia and the flood plains of the Llanos de Moxos, Mamoré watershed, identified as a Centre of Plant Diversity and Endemism (Site SA24) (Beck and Moraes, 1997). These area is considered by Langstroth Plotkin (2012). Moxos is surrounded by the forests of the Upper Madeira basin and is an area of largely open vegetation – herbaceous wetlands, grasslands, savannas, and woodlands (Beck, 1983, 1984; Langstroth Plotkin, 2012). Images (not included in the minimum calculation of the EOO or AOO) suggest that V. boliviana ’s range extends further west (natural or cultivated) to Rurrenabaque. We estimated a minimum and maximum EOO and AOO. The maximum range was based on the potential habitat across the Llanos de Moxos region (including geographical information from public unverified images) and verified herbarium collections and iNaturalist images. The minimum range was based on the coordinates of herbarium collections and iNaturalist images alone. We estimated that the EOO of V. boliviana ranges between 8,006 km 2 (minimum) and 33,151 km 2 (maximum), falling close to the thresholds of the Vulnerable category under criterion B. We estimated that the AOO of V. boliviana ranges from 32 km 2 (minimum) to 2,000 km 2 (maximum), falling between the Endangered and Vulnerable categories. There are less than five known locations for V. boliviana. We could find no information about population fragmentation but believe that V. boliviana may be vulnerable to fluctuations in flooding and drought throughout the year. For example, Beni Department has been recently affected by seasonal floods, fires and droughts due to El Niño and La Niña climate events (Vásquez, 2015). Arecent increase in agriculture-lead deforestation has been documented along the Trinidad-Santa Cruz highway (Langstroth Plotkin, 2012), to the south of known V. boliviana populations and satellite images from Google Earth Pro and i-Terra suggest extensive deforestation along the edges of roads (i-Terra, 2021; NASA, 2021; GoogleEarth, 2022) which we use to infer an active decline in habitat quality. Taking a precautionary approach and based on the small EOO and AOO, small number of locations (5), and continuing decline in habitat, we assess V. boliviana as Vulnerable (VU), according to criteria B1ab(iii)+B2ab(iii). Notes. Victoria boliviana sp. nov. has the largest observed leaves of the three species, with laminae> 3 m in length having been observed. The abaxial surface of the upturned rim surface varies between individual plants in the same locality from dark maroon to very pale, almost “white” green, a characteristic not found in other species. Prickles on abaxial outer tepal surfaces are absent or very few, and if present are not confined to the lower portion of the outer tepals unlike V. cruziana where the smaller number of prickles are confined to the lower one-third of the abaxial tepal surface. Victoria boliviana is the only species of Victoria whose carpellary appendages have upper portions that are longer than the lower portions. In addition, V. boliviana ’s stigmatic chambers are the shallowest of the three Victoria species. Haenke saw Victoria plants in Yacuma in 1801, during the Malaspina expedition (Gickelhorn, 1966, p. 105; Ibañez Montoya, 1984), but no identifiable description is available and no voucher has been found. d’Orbigny (1840, p. 57) reported seeing this species on the banks of the Mamoré river in 1832 and mistakenly assigned this species to V. amazonica when publishing his description of V. cruziana. Further investigations and surveys are required to better understand the species’ current range, population fluctuations and habitat and thus better predict the impacts of the threats identified. Field observations of the flowers from a single population in the Llanos de Moxos suggest that whilst pollinated by beetles, V. boliviana sp. nov. flowers may host fewer individuals of pollinators than V. amazonica, only 4–10 individuals being observedintheflowersof theformer, comparedto> 20 inthe flowers of the latter. This could be due to a lower density of pollinators in their area of occupation. Additional Material – BOLIVIA. Beni: Santa Ana del Yacuma, –65.4236, –13.74, –/6–7/1845, Bridges, T. s. n. (K); Cercado: Laguna Suarez 5 km sur de la ciudad de Trinidad, –64.864167, – 14.872222, 08/05/2019, Magdalena, C. Melgar, D. G., Salazar, C. D., Alvarez, C., Gutierrez, G., Arias, J. 154 (German Coimbra Sanz Herbarium, Jardin Botanico Municipal); Moxos, pasando el Río Mamoré, cerca al puente del Río Tijamuchi a lado del camino, –65.145278, –14.851111, 09/05/2019, Magdalena, C. Melgar, D. G., Salazar, C. D., Alvarez, C., Gutierrez, G., Arias, J.155 (Herbario German Coimbra Sanz Jardín Botanico Municipal)., Published as part of 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, pp. 1-31 in Frontiers in Plant Science 13 on pages 22-25, DOI: 10.3389/fpls.2022.883151, http://zenodo.org/record/7576624, {"references":["Langstroth Plotkin, R. (2012). Biogeography of the Llanos de Moxos: natural and anthropogenic determinants. Geographica Helvetica 66, 183 - 192. doi: 10.5194 / gh- 66 - 183 - 2011","Beck, S. G. (1983). Vegetationsoekologische Grundlagen der Viehwirtschaft in den Ueberschwemmungs-Savannen des Rio Yacuma (departamento Beni, Bolivien) Dissertationes Botanicae Bd. Vaduz: J. Cramer, 186.","Beck, S. G. (1984). Comunidades vegetales de las sabanas inundadizas en el NE de Bolivia. Phytocoenologia 12, 321 - 350. doi: 10.1127 / phyto / 12 / 1984 / 321","Vasquez, G. C. (2015). \" Indigenous people and climate change: causes of flooding in the Bolivian Amazon and consequences for the indigenous populaton, \" in Inequality and Climate Change: Perspectves from the South, ed. G. C. D. Ramos (Dakar: Council for the Development of Social Science Research in Africa (CODESRIA )), 121 - 136. doi: 10.2307 / j. ctvh 8 r 0 w 3.12","i-Terra (2021). CIAT-Terra-I. Terra-i Peru. Available online at: http: // terra-i. org / terra-i / data / data-terra-i _ peru. html (accessed October 20, 2021)","NASA (2021). Earth Observatory. Deforestation. Washington, DC: NASA.","GoogleEarth (2022). 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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., Published as part of 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, pp. 1-31 in Frontiers in Plant Science 13 on pages 20-22, DOI: 10.3389/fpls.2022.883151, http://zenodo.org/record/7576624, {"references":["de Lima, C. T., Machado, I. C., and Giulietti, A. M. (2021). Nymphaeaceae of Brasil. Sitientibus serie Ciencias Biologicas 21. doi: 10.13102 / scb 4986","i-Terra (2021). CIAT-Terra-I. Terra-i Peru. Available online at: http: // terra-i. org / terra-i / data / data-terra-i _ peru. html (accessed October 20, 2021)","USGS Environmental Health Program (2019). Mercury Isotope Ratios used to Determine Sources of Mercury to Fish in Northeast U. S. Streams. Reston, VA: USGS.","Swenson, J. J., Carter, C. E., Domec, J. C., and Delgado, C. I. (2011). Gold mining in the Peruvian Amazon: global prices, deforestation, and mercury imports. PLoS One 6: e 18875. doi: 10.1371 / journal. pone. 0018875","Hutukara Associacao Yanomami, and Associacao Wanasseduume Ye'kwana (2020). Scars in the Forest - The Growth of Illegal Mining in the Yanomami Indigenous Territory (YIL) in 2020. Report. Available Online at: https: // www. amazoniasocioambiental. org / en / radar / em- 2020 - garimpoavancou- 30 - na-terra-indigena-yanomami-aponta-relatorio / (accessed October 20, 2021)","MapBiomas (2021). MapBiomas Project- Collection Cobertura of the Annual Series of Land Use and Land Cover Maps of Brazil. Available online at: https: // plataforma. brasil. mapbiomas. org (accessed October 20, 2021).","Mercado, J. (2021). Tras el dorado. Cronicas de la explotacion del oro en la Amazonia. Cochabamba: La Libre."]}

La période entre 2000 et 2012 a vu une progression significative du nombre de nouveaux docteurs en Europe (+56 %). La formation par la recherche reste une expérience unique et formatrice sur les plans personnels et professionnels. Chaque thèse est en effet unique, non seulement en raison du sujet et de son domaine de spécialisation, mais également par le parcours personnel et académique du candidat. Une thèse de doctorat est par définition jalonnée de défis, de joies, de doutes, de remises e...