Oiapoque, Aeroporto, campina de areia branca, 20 Mar. 1982 (fl), *B.V. Rabelo RN 1545* (UB); Mun. Macapá, km 104 da BR-156, campo cerrado, 5 May 1979 (fl), *B.V. Rabelo s.n.* (HAMAB No. 1412); Mun. Oiapoque, Aldeia do Manga, Tribo Karipuna, capoeira próximo da Aldeia, 20 Aug. 1981 (fl), *B.V. Rabelo 1303* (HAMAB); Mun. Oiapoque, aeroporto, campina de areia branca, 20 Mar. 1982 (fl), *B.V. Rabelo 1545* (HAMAB); BR-156, km 103, campina de areia branca, 29 Feb. 1984 (fl), *B.V. Rabelo et al. 2544* (HAMAB); Ferreira Gomes [ca. 0°51'N, 51°10'W], cerrado, 23 Jul. 2009 (fl, fr), *A.E.S. Rocha 1131* (HAMAB, MG); Calçoene, savanna, Estrada para Cunasin, 27 Jun. 2010 (fl, fr), *A.E.S. Rocha & S.V.C. Neto 1327* (HAMAB, MG); Mun. Amapá, perto do aeroporto, terreno úmido, campo, erva rastreira, occasional à margem da Estrada, 25 Aug. 1978 (fl), *W. Rodrigues 10059* (INPA); Mun. Amapá, quadrícula SB-22-VA, ponto 44, campos naturais de terra firme, 9 Apr. 1982 (fl), *N.A. Rosa et al. 4201* (MG, NY); Mun. Pracuúba, Vila Porto Franco, cerrado, subarbusto 35 cm altura, 1 Jul. 2002 (fl), *M.A.C. Santo & M.H.A. Martins 283* (HAMAB); Porto Grande, savana, 10 Oct. 103 (fl), *M.V.B. Soares & W.L.S. Silva 103* (MG); Mun. Pracuúba, ruderal, 9 May 1987 (fl), *C. Souza & B.V. Rabelo C-50* (HAMAB). **Pará:** Mun. Faro, Fazenda Santa Olimpia, Campo Umirí, 6 Dec. 1950 (fl), *G.A. Black & P. Ledoux 50-10577* (IAN, U); Óbidos, em torno da Serra do Escama, perto do Lago Pauxí, 22 May 1957 (fl), *G.A. Black & W.A. Egler 57-19454* (IAN), *P. Cavalcante 224* (MG); Prainha [ca. 1°48'S, 53°28'W], capoeira, 10 May 1903 (fl), *A. Ducke s.n.* (*MG 3601*) (MG); campos a E de Faro, 21 Aug. 1907 (fl, fr), *A. Ducke s.n.* (*MG 8436*) (MG); Mun. Almeirim [ca. 1°31'S, 52°34'W], Estrada de accesso a Monte Dourado, saindo da cidade, 12 May 2005 (fl), *L.C.B. Lobato 3246* (MG); arredor do Campo de Aviação, margem do Rio Parú, 2 Jun. 1974 (fl), *N.A. Rosa 6* (IAN); Região do Rio Jari, Água Branca, km 3, campo cerrado, 28 Jul. 1969 (fl, fr), *N.T. Silva 2491* (IAN, NY, UB); Mun. Óbidos, Estação Ecológica do Grão Pará, trilha 3, 0°38'33''N, 55°41'47''W, 22 Jan. 2009 (fl), *M.G.C. Souza & G. Teixeira 456* (MG), 24 Jan. 2009 (fl), 533 (MG); Macau airstrip, 1.5 h upstream from Lageira airstrip, on Rio Maicuru, 0°55'S, 54°26'W, 270 m, 24 Jul. 1981 (fl, fr), *J.J. Strudwick et al. 3481* (INPA, MG, MO).

8-5b. *Sipanea pratensis* var. *dichotoma* (Kunth) Steyerm. in Mem. New York Bot. Gard. 17: 273. 1967. (Figures 49-50).

(=) *Sipanea dichotoma* Kunth in Humb. & Bonpl., Nov. Gen. Sp. 3 (quarto): 397. 21 Nov. 1819 ["1820"]).

Type: VENEZUELA. "Atures, cataractam Orinocensium," [Atures, Orinoco waterfalls], s.d. [15-17 Apr. 1800], *A.J.A. Bonpland & F.W.H.A. Humboldt 855* (*lectotype* P [barcode P00729459], **here designated**; *isolectotype* B [barcode B-W 04124-1 0]).

(=) "*Ptychodea pedunculata*" in Cham. & Schltdl. in Linnaea 4: 168. 1829, *nom. illeg. superfl. pro syn.*

Type: VENEZUELA. "Atures, cataractam Orinocensium," s.d. [15-17 Apr. 1800], *A.J.A. Bonpland & F.W.H.A. Humboldt 855* (*lectotype* B [barcode B-W 04124-1 0]) **here designated**; *isolectotype* P [barcode P00729459]). This name was cited by Chamisso & Schlectendal (1829: 168) as a synonym of *Sipanea dichotoma* Kunth.

(=) *Sipanea trichantha* Miq., Linnaea 18(3): 293. 1845 ["1844"]).

Type: SURINAME: Onoribo, s.d. (fl, fr), *H.C. Focke 60* (*lectotype* U [barcode U00116946], **here designated**).

(=) *Sipanea colombiana* Wernham, J. Bot. 55: 174. 1917

Type: COLOMBIA. [Norte de Santander]: Ocaña, Crecenoche, 300 m, May [1846-1852] (fl), *L. Schlim 548* (holotype BM [barcode 000614320]; isotypes BR [barcode BR824758], K [without barcode], P [2 sheets, barcodes P00729397 and P00748109]; photo-K NYBG Neg. No. 3247]; photo at NY).

(=) *Sipanea pratensis* var. *dichotoma f. brachycarpa* Steyerm., Mem. New York Bot. Gard. 17: 276. 1967

Type: VENEZUELA. Amazonas: Danta (Tapir) Falls, Rio Cuao, Río Orinoco, 125 m, 19 Nov. 1948 (fl, fr), *B. Maguire & L. Politi 27324* (holotype VEN not found; isotypes NY [2 sheets, barcodes NY00133317, and NY 00133318], US [barcode 00137727]).

(=) *Sipanea pratensis* var. *dichotoma f. glabriloba* Steyerm., Mem. New York Bot. Gard. 17: 276. 1967

Type: VENEZUELA. Amazonas: Yavita-Pimichín trail, near Yavita, Río Atabapo, 125-140 m, 10 Jun. 1959 (fl, fr), *J.J. Wurdack & L. Adderley 42916* (holotype NY [barcode 0013323]; isotypes S [Acc. No. 10-27872], US [Acc. No. 2320890, barcode 00137728]).
<figure><img src="image_54.png" /><figcaption>**Figure 50.** *Sipanea pratensis* var. *dichotoma*. A. branch with terminal inflorescence. B. Node with stipule and leaf bases. C. Longitudinal section of hypanthium, ovary and calyx (note colleter at each calyx lobe sinus). D. Inflorescence branchlet. E. Seeds. F. Dissected corolla with detail of hairs at mouth and corolla throat. Illustration by Bruno Manara (Reproduced with permission from Flora de Venezuela vol. 9(1), fig. 59. 1974).</figcaption></figure>
(=) *Sipanea pratensis* var. *dichotoma* f. glabrior Steyerm., Mem. New York Bot. Gard. 17: 277. 1967

Type: VENEZUELA. Bolivar: Cerro San Borjas, 100-300 m, 12 Dec. 1955 (fl), J.J. Wurdack & J.B. Monachino 39823 (holotype NY [barcode 00133324]; isotype VEN [Acc. No. 65160]).

(=) *Sipanea pratensis* var. *dichotoma* f. breviflora
Steyerm., Mem. New York Bot. Gard. 17: 277. 1967

Type: VENEZUELA. Bolivar: Cerro between Las Nieves and base of slopes SE of Cerro Picacho, N of Las Nieves, 45 km N of Tumeremo, Altiplanicie de Nuria, 100-300 m, 1-11 Feb. 1961 (fl, fr), J.A. Steyermark 89276 (holotype VEN [Acc. No. 64017]; isotypes NY [4 sheets, barcodes NY00133319, NY00133320, NY00133321, NY00133322]). Steyermark (1967: 277) cited the type of forma breviflora as "*Steyermark* 89276 (holotype VEN)." Therefore, although the VEN specimen Acc. No. 64017 was annotated by Steyermark as isotype, it is the holotype of this taxon.

(=) *Sipanea pubinoda* Steyerm., Mem. New York Bot. Gard. 17: 278. 1967

Type: VENEZUELA. Amazonas: Capibuara, Alto Casiquiare, 120 m, 29 May 1942 (fl, fr), Ll. Williams 15656 (holotype VEN [Acc. No. 16075]; isotypes NY [without barcode], RB [Acc. No. 108624, barcode 00364140], US [2 sheets, barcodes 00604080 and 00137729]; photo at NY).

*Perennial herb or subshrub* (basal internodes woody), 0.3-1.5 m tall, basal portion sometimes woody, terrestrial or epipetric (on inselbergs), erect or ascending, rarely decumbent, often forming large populations; stems to 1.8 m long, rarely geniculate, sometimes rooting at nodes; young branches thin to stout, 1-3.3 mm thick, terete, sparsely to densely, appressed or spreading, pubescent or sericeous (hairs whitish or beige, 0.8-2 mm long). *Leaves* with blades broadly ovate, ovate, ovate-elliptic, elliptic, oblong-elliptic, oblong-lanceolate, lanceolate, narrowly lanceolate to narrowly oblong-elliptic, 1.8-7.5 x 0.5-2.5 cm, round, obtuse to acute, rarely decurrent, at base, acute to acuminate at apex; secondary veins 4-8 on each side of midrib, obsolete or slightly impressed above, prominent below. *Inflorescences congested-cymose during anthesis*, 4-8-flowered, heads 0.5-1.8 x 0.3-1.5 cm (excluding corollas) during anthesis, moderately or much expanding and obvi-

ously branched during and/or after anthesis; peduncle (when present) to 7.5 cm long; bracts subtending secondary branches lanceolate, oblong-lanceolate to narrowly lanceolate, 3-5.5 x 0.5-1.1 mm, acute to acuminate at apex, sparsely to densely antrorse-pubescent outside; bracteoles subtending flowers (when present) lanceolate to narrowly lanceolate, 2-3.5 x 0.4-0.5 mm, acute to acuminate at apex, sparsely to densely antrorse-pubescent outside; *inflorescences evidently branched during anthesis*, 4-37-flowered, 1-10 x 0.8-7 cm (excluding corollas) during anthesis, with lateral axes 0.5-7 cm long, more or less elongating during and/or after anthesis, 1-10 cm long (excluding corollas), scorpiod, producing a subsecund monochasial of dichasial appearance; peduncle (when present) to 7.5 cm long, sparsely to densely, appressed or spreading, pubescent or sericeous (hairs whitish or beige, 1.5-2 mm long); bracts subtending secondary branches (when present) lanceolate, oblong-lanceolate, narrowly lanceolate to linear, 2.5-7 x 0.5-1.2 mm, acuminate at apex, sparsely to densely antrorse-pubescent outside, sometimes absent; bracteoles subtending flowers (when present) lanceolate, narrowly lanceolate to linear, 2-7 x 0.4-0.7 mm, appressed pubescent to sericeous outside (hairs whitish or beige, 1.5-2 mm long). *Hypanthium* 0.7-1.5 x 0.5-0.9 mm. *Calyx* lobes lanceolate, narrowly oblong-lanceolate to linear, 2.3-6 x 0.2-0.5 mm. *Corolla* 14-28 mm long; tube (8-)10-17 mm long, 0.4-0.7 mm wide at base, 1.5-3 mm wide at mouth; lobes broadly elliptic, elliptic, obovate, oblong-elliptic to oblanceolate, (sometimes varying in shape and size within the same population), sometimes unequal or of slightly different shape within the same corolla, (2.5-)4-11 x (2-)3-7 mm, obtuse to acute, rarely apiculate when subobtuse or acute, at apex. *Long-styled flowers* [Delprete & Marjanom 12441 (corolla tube 14 mm long), Delprete & Benjamin 12861 (corolla tube 14 mm long), Jansen-Jacobs et al. 1114 (corolla tube 8 mm long)]: stamens inserted at 6-7 mm from base or about the middle of corolla tube; filaments 0.5-0.7 mm long; anthers narrowly oblong-elliptic, 1.8-2.5 mm long, acute at both ends; style exerted 1.5-2.5 mm beyond corolla mouth, (9.5-)15-16.5 mm long; style branches narrowly oblong to oblong-ovate, (0.6-)1.3-1.5 mm long, round at apex. *Short-styled flowers* [Delprete & Marjanom 12440 (corolla tube 12.5 mm long), Delprete & Marjanom 12596 (corolla tube 14 mm long)]: stamens inserted at 4-5.5 mm below corolla mouth; filaments 0.5-0.7 mm long; anthers linear to narrowly oblong-elliptic, 2.5-3 x 0.2-0.3 mm, acute at both ends; style included, 6-8 mm long, glabrous; branches linear, 2-3 mm long, acuminate at apex. *Infructescences* with mature fruits dichotomously branched, more or
less expanded, with scorpioid secondary branches (0.5–)1–10 cm long. *Capsules* longer than broad, cylindrical-oblong to oblong-narrowly ovoid, (3–)4.5–8 × 1.5–3 mm, round to acute at base, thinly woody, commonly smooth or faintly costate when dry, sparsely to densely pubescent to sericeous (hairs 0.5–1.5 mm long) to almost glabrate, pale green when young, turning beige to pale brown when dry.

**Taxonomic notes:** The fate of the botanical specimens gathered by Bonpland and Humboldt has been discussed by several authors (Delprete 2001; Hiepko 2006; Kirkbride & Wiersema 2020; Lack 2004a, 2004b, 2009, 2018; Stauffer et al. 2012). Humboldt divided these collections in several sets. The main set, although incomplete, is currently kept in the Bonpland Herbarium (P-Bonpl.) at the Museum of Natural History in Paris. Bonpland was initially in charge of writing the descriptions for the *Nova genera et species plantarum*, but after several months he renounced to this task. Then, Humboldt invited Carl Ludwig Willdenow (1765–1812) to take up this huge task, to whom he sent a set in Berlin, but he died shortly after. Subsequently, Humboldt invited Carl Sigismund Kunth (1788–1850), then in Berlin, to take up this enormous work. Kunth arrived in Paris in 1813, and started immediately to write the descriptions for the *Nova genera*. Three years later, in 1816, Bonpland emigrated from France to Argentina, with his personal herbarium, which was a third, incomplete set of the collections that he made with Humboldt. Therefore, Kunth and Bonpland where both in Paris from 1813 to 1816, and, as far as I know, it is not possible to exclude the possibility the Kunth was able to study the specimens in Bonpland's personal herbarium during that period. Kunth went to La Rochelle, just before Bonpland's departure, to convince him to leave his personal set and his field book in France, but he was only able to obtain Bonpland's field book ("Journal Botanique"). Bonpland returned his personal herbarium from Argentina in 1832, which arrived in Paris in 1833, and was then integrated in the P general herbarium in 1837. These specimens have the label "Herbier de l'Amérique équatoriale, donné par M. A. Bonpland."

No original specimen of *Sipanea dichotoma* was found in P-Bonpl. and P-JU, but two specimens were found at P and B-W. This is not surprising, as the absence of original specimens in P-Bonpl. has puzzled many botanists (Stauffer et al. 2012). For example, out of the about 350 monocotyledon names described by Kunth, Stauffer et al. (2012) reported that 235 original specimens are not present in P-Bonpl. The P specimen of *S. dichotoma*, with barcode P00729459, has the

label "germ. an 8. Orinoco, no. 855. Pent.¹ª, in Guainia. Atures" handwritten by Bonpland, where "germ. an 8" means that it was collected in the month of Germinal (March–April) and the 8th year of the French Republican calendar, which corresponds to 1800. This specimen has a second label with the printed heading "Herb. Mus. Paris" and the print "Herbier de l'Amérique équatoriale, donné par M. A. Bonpland" at the bottom of the label. On the second label it is handwritten (author unknown) "Hedyotis, Sipanea pratensis Aubl., K. Schum. Fl. Bras. VI, pars VI, p. 249." Therefore, this specimen was part of the set that Bonpland returned to Paris from Argentina. It has a simple branch with a few nodes with leaves and axillary brachyblasts, and a terminal, dichomously branched inflorescence with a few open capsules. It is here designated the lectotype of *Sipanea dichotoma*. The image of this specimen is available at: https://science.mnhn.fr/institution/mnhn/collection/p/item/p00729459

The original specimen of *S. dichotoma* at B-W, with barcode B-W 04124 -01 0, is inside a folder with the label "Pentandria Monogynia, Ptychodea pedunculata, floribus terminalibus pedunculata, Habitat ad Orinocum" handwritten by Willdenow. On the sheet there are three labels. The first two labels are handwritten by two unknown authors "Sipanea dichotoma H. B. Kth. 3. p 110" and "855 Pent.¹ª, Orinoco (Humboldt)". The third label has "Sipanea pratensis Aubl." handwritten by K. Schumann and the the print "det. Schumann in Fl. Bras." At the lower right corner is handwritten directly on the sheet "Humboldt. W." meaning that this is a specimen that Humboldt sent to Willdenow. This specimen has the same collection number of P00729459, to which is similar by bearing a small branch with a few leafy nodes, and two branched inflorescences with open capsules. This specimen is the isotype of *S. dichotoma*. This specimen is also original material of "Ptychodea pedunculata", an illegitimate superfluous name that Chamisso & Schlechtendal (1829: 168) cited as a synonym of *Sipanea dichotoma*. The image of this specimen is available at: https://ww2.bgbm.org/Herbarium/specimen.cfm?Barcode=BW04124010

Kunth (in Humboldt & Bonpland (1821 ["1820"): 397 (quarto)) described *Sipanea dichotoma* as having hypocrateriform, pink corollas with a tube longer than the calyx, 5-lobed, yellow-pubescent at mouth, with 5 included stamens, and a bilobed style. As both specimens of *Bonpland & Humboldt 855* only have open capsules, Kunth might have used another specimen to describe the flowers, or he might have used Bonpland's field book to complete the description of this species.

Miquel (1844: 49–50), in the introduction of his *Symbolae ad Floram Surinamensem* wrote that Hendrik
Focke (1802–1858) had a profound botanical knowledge, and made excellent herbarium specimens, pickled flowers, drawings made from living plants, and detailed species descriptions. Miquel wrote that he used these specimens and accompanying information for the description of taxa in the *Symbolae ad Floram Surinamensem*. Miquel (1845: 293–294) along with the description of *Sipanea trichantha* Miq. stated that this species is frequent in Suriname, without citing the collection locality or the herbarium of deposit. He started the species description with “Annua? ex cl. Focke perennae, [...]”, which means that it is an annual species that, according to Focke, might also be perennial. Therefore, as explained above, original collection to be considered for type designation of new taxa described by Miquel should be specimens collected by Focke in Suriname.

Steyermark (1967: 274) for *S. trichantha* cited “Suriname: [...] *Hostmann 663* (photo of type of *Sipanea trichantha*),” and in the following page (1967: 275) he wrote “*Sipanea trichantha* Miq., described from Suriname, is conspecific *S. dichotoma* H.B.K. The photo of the type of *S. trichantha* (*Hostmann 663* from the Delessert Herb.) shows loosely moderately villous stems with elongated leaves with a short, appressed pilosity on the lower nerves and midrib. The lower surface appears glabrate, while the upper surface appears appressed-pilose with short hairs. The photo shows elongate, setaceous calyx-lobes which appear to measure 3.5–4.5 mm long. The corolla lobes in bud appear to show slight pilosity along the outer margins. The characters shown by the photo of the *S. trichantha* type fall within the range of variation matched by various other collections from Suriname with similarly elongated leaves, similar type of inflorescence, calyx-lobes, and corolla-tube, which are to be identified with *S. dichotoma* H.B.K. (= *S. pratensis* var. *dichotoma*), tending toward the variation with longer, slender, calyx-lobes.” However, as discussed above, Miquel stated that all his species descriptions were based on specimens collected by Focke. Therefore, Steyermark’s citation of *Hostmann 663* at G the as the type of *S. trichantha* cannot be interpreted as an inadvertent neotypification, and should be superseded by a specimen collected by Focke.

According to Stafleu and Cowan (1981), Friedrich Miquel (1811–1871) worked at Utrecht during 1859–1871, his herbarium and types are mostly at U, and additional specimens are present at E, K, L, and P. Accordingly, Ek (1991: 38) confirmed that Focke collected in Suriname and sent his collections to Miquel. At U are present two specimens collected by Focke with the labels “Sipanea trichantha sp.” handwritten by Miquel. One specimen, barcode U0016947, has a label “Sipanea trichantha [...]

a forte dichotoma HBK, Surinami, Para, poste Wran, s. April, legit Il. C. Focke 153.” The other specimen, barcode U00116946, has the label “S. trichantha Miq., Sipanea pratensis Aubl., Onoribo, Surinami, in Martu [Isic, Martius?] 1023, Focke 60.” The latter specimen, barcode U00116946, collected by Focke near the town of Onoribo, has three branches with flowers in anthesis and fruits, and is here designated the lectotype of *Sipanea trichantha*.

**Distribution and ecology:** Distributed in Colombia, Venezuela, the Guianas, and northern Brazil, in the states of Roraima, Amapá, Amazonas, Pará, and Maranhão (Figure 49). Often forming large populations, in white-sand savannas, brown-sand savannas, on granitic inselbergs, granitic outcrops, swampy margins in savannas, seasonally inundated savannas, open field savannas or savannas with sparse trees of *Curatella Americana* and *Byrsonima* spp., margins of semi-shady forest, creek margins in forests, margins of forest trails, disturbed or secondary vegetation, roadside vegetation; in sunny or semi-shady exposures; on sandy, granitic or lateritic soils; from near sea level to 1200 m altitude.

**Phenology:** Due to the short life span of this species, it is common to find individuals with flowers and fruits. Flowering and fruiting the whole year.

**Suggested conservation status:** Least Concern (LC). This variety is amply distributed in Colombia, Venezuela, the Guianas, and Amazonian Brazil (Roraima, Amapá, Amazonas, Pará, Maranhão). It is most common in various kinds of savannas (see ecology, above), and is also a ruderal species growing at margins of forest trails, in secondary vegetation, and roadside vegetation, often forming large populations. As it is quite common in savannic vegetation and is a ruderal species known by numerous collections, this variety belongs to the Least Concern (LC) category, following IUCN criteria (IUCN 2012, 2019).

**Vernacular names:** VENEZUELA: clavel del agua (Spanish, *Williams 11770*), clavel de pozo (Spanish, *Williams 11953*). SURINAME: nepoko (Karib, *Gonggrijp & Stahel 805*), witbaka (N.E., *Gonggrijp & Stahel 805*). BRAZIL: Roraima: malva do campo (*Jaccoud s.n.* (INPA 285)).

**Specimens examined:** COLOMBIA: Amazonas: Rio Igara-Parana (affl. Río Putumayo), corr. La Chorrera, territoire des indiens Witoto Jitomagaro, famille Kuiru, 17 km en aval de La Chorrera [ca. 1°27'S,
72°45'W], 25 Dec. 1973 (fr), *J. Gasche & J. Desplats* 91 (P), Parcelle 107, 3 Apr. 1974 (fr), 1268 (P), Parcelle 124, 26 Jun. 1974 (fl, fr), *C. Sastre* 3507 (P). **Casanare:** Tauramena [ca. 5°1'N, 72°45'W], sabanas o llanos al S de la población, 550 m, 30 Jun. 1961 (fl), *L. Uribe Uribe* 3757 (US). **Guainía:** Mun. Inírida, Corregimiento de Cacaual, Río Atabapo, Comunidad de Pato Corona, bosque de tierra firme, 3°21'11"N, 67°19'48"W, 10 May 2007 (fl-fr), *D. Cárdenas et al.* 20642 (COAH, NY); Mun. Inírida, Caño Chaquita, sabanas naturales de arenas blancas inundadas, 3°31'12"N, 67°39'11"W, 9 Jul. 2008 (fl-fr), *D. Cárdenas et al.* 21658 (COAH, NY). **Magdalena:** Bajo Magdalena, Sep. 1916 (fl, fr), *M.T. Dawe* 445 (US); 17 km N of El Banco [ca. 9°0'N, 73°58'W], on rd to Chimichagua, very abundant in sabana, 1 Aug. 1938 (fl, fr), *O. Haught* 2216 (NY, US); La Jagua, 100 m, 3 Aug. 1943 (fl, fr), *O. Haught* 3587 (US). **Meta:** Puerto Gaitán, Estación Carimagua, region de Core-Alegría, 4°35'N, 71°18'W, 200-300 m, sabanas arenosas, 10 Jan. 1995 (fl), *F. González & R. Serna* 3378 (NY). **Vaupés:** San José del Guaviare [ca. 2°33'N, 72°38'W], sabana, 240 m, 4 Nov. 1939 (fl-fr), *J. Cuatrecasas* 7411 (US); Río Negro, San Felipe [ca. 1°54'N, 67°4'W], 200 m, 13-25 Nov. 1952 (fl, fr), *H. Humbert* 27496 (P). **Vichada:** Raudal San Borja [ca. 6°2'N, 67°25'W], Ventanas, margen izquierda del Río Orinoco, 100 m, 17 Mar. 1971 (fl, fr), *P. Pinto & C. Sastre* 1291 (P).

**VENEZUELA:** **Amazonas:** Dpto. Río Negro, selvas pluviales a 8 km SE de San Carlos de Río Negro, por la carretera que va a Solano, 1°10'N, 67°0'W, 12 Apr. 1985 (fl), *G. Aymard et al.* 3507 (MO, MY, PORT); Depto. Atures, Puerto Ayacucho [ca. 5°39'N, 67°37'W], near MARNR fish culture station in fields and woods south of the station, area of many granite outcrops, 90 m, 21 Jan. 1985 (fl), *B.M. Boom & A.L. Weitzman* 5283 (NY); Depto. Río Negro, San Carlos de Río Negro, ca. 20 km S of confluence of Río Negro and Brazo Casiquiare, 1°56'N, 67°3'W, 119 m, 24 Jan. 1985 (fl, fr), *B.M. Boom et al.* 5340 (INPA, MO, NY, US); bords de l'Orenoque, Cerro Carichand, s.d. (fl), *J. Chaffanjon* 101 (P [2 sheets]); San Carlos de Río Negro, ca. 20 km S of confluence of Río Negro and Brazo Casiquiare, 1°56'N, 67°3'W, 119 m, near secondary radar construction site, 5.3 km NNE of San Carlos de Río Negro on Solano Rd., on bulldozed white sand, 23 Mar. 1981 (fl), *H.L. Clark et al.* 7835 (INPA, NY); Alto Orinoco, La Esmeralda [ca. 3°10'N, 65°31'W], scarse in the savanna, 14 Jul. 1951 (fl), *L. Croizat* 86 (NY); Puerto Ayacucho, 24 Mar. 1950 (fl), *H.M. Curran* 136A (NY), 140 (NY); Depto. Río Negro, km 11 NE of San Carlos de Río Negro, along rd to Solano, 1°53'N, 67°2'W, 75 m, 24 Jun. 1984 (fr), *G. Davidse & J.S. Miller* 26493 (NY), 26533 (MO, NY); Depto. Río Negro,

Mamurividi, laja along Río Pasimoni, E bank, 1°50'N, 66°38'W, 75 m, 27 Jun. 1984 (fl, fr), *G. Davidse & J.S. Miller* 26710 (MO, NY, US); Depto. Río Negro, Mucuripari, small laja along the lower part of the Río Baria, 1°20'N, 66°26'W, 80 m, 28 Jun. 1984 (fl, fr), *G. Davidse & J.S. Miller* 26770 (MO, NY, US); Depto. Casiquiare, Culimacare, Indian village on the N bank of the Brazo Casiquiare, 1°58'N, 66°50'W, 75 m, 27 Jun. 1984 (fl), *G. Davidse & J.S. Miller* 26730 (MO, NY, US); Depto. Río Negro, Río Pasimoni, between its mouth and its junction with the Río Baria and the Río Yatua, 1°10'-27'N, 66°25-32'W, 80 m, inundated forest along river, 22-23 Jul. 1984 (fl), *G. Davidse* 27706 (NY); Depto. Río Negro, lower part of the Río Baria, 1°27-35'N, 66°32-35'W, 80 m, inundated forest along river margin, 23-25 Jul. 1984 (fl), *G. Davidse* 27807 (NY); Depto. Atures, Mun. Puerto Ayacucho, Setor Corocito, ca. 3 km N of Puerto Ayacucho, on the road toward El Burro, sabaneta inundable con pequeños arboles (3-7 m) esparsos, vegetation dominated by Poaceae and Cyperaceae, 5°42'N, 67°35'W, 85 m, 5 Jan. 2001 (fl), *P.G. Delprete et al.* 7438 (CAY, MO, P, TFAV, VEN), 7439 (L, NY, TFAV, VEN), 7440 (F, TFAV, VEN), 7441 (CAY [2 sheets], TFAV, VEN), 7442 (VEN); Dpto. Atures, 9 km por arriba del Raudal Remo, 4°34'N, 67°18'W, 120 m, May 1989 (fl), *E. Foldats & J. Velazco* 9528 (NY); bords de l'Orenoque, Savane d'Atures, 28 Jul. 1887 (fr), *A. Gaillard* 77 (P); bords de l'Orenoque, bois humides des bords de l'Atabapo, 31 Aug. 1887 (fl), *A. Gaillard* 191 (P); San Carlos, Río Negro, 100 m, 28-29 Jan. 1930 (fl), *E.G. Holt & W. Gehriger* 305 (US), 312 (NY, US), 329 (NY, RB, US); San Carlos, Río Negro, 100 m, 27 Jan. 1931 (fl), *E.G. Holt & E.R. Blake* 640 (NY, US); terreno situado entre la carretera hacia Samariapo y la pista de aterrizaje del Aeropuerto de Puerto Ayacucho, entre las macollas elevadas de la "cowfoothill" sabana, 5°37'N, 67°36'W, 75 m, 13 Apr. 1977 (fl), *O. Huber* 555 (NY); entre la carretera hacia Samariapo y la pista de aterrizaje del aeropuerto de Puerto Ayacucho, sabana semi-inundada de tierra negra SW de la Estación, 5°37'N, 67°36'W, 75 m, 23 May 1977 (fl, fr), *O. Huber* 779 (NY, US, VEN); terreno situado entre la carretera hacia Samariapo y la pista de aterrizaje del aeropuerto de Puerto Ayacucho, sabana húmeda a lo largo de un pequeño arroyo casi sin agua, al pié de la Laja Grande, 5°37'N, 67°36'W, 75 m, 6 Dec. 1977 (fl), *O. Huber* 1283 (NY); Depto. Atures, ca. 30 km N of Puerto Ayacucho, Sabana de Rincones de Cachorro, 5°48'N, 67°20'W, 80 m, 23 May 1979 (fl), *O. Huber* 3785 (US); Depto. Río Negro, carrettera San Carlos-Solano, entre km 4 y 20, 1°56'W, 67°3'W, 120 m, 15 Sep. 1980 (fl, fr), *O. Huber & E. Medina* 5654 (NY, US); Depto. Atures, 56 km NE of Puerto Ayacucho, 170 m, 22 Nov. 1984 (fl, fr), *R. Kral et*
al. 71760 (NY); Dpto. Casiquiare, near Chapezon, between Boca de Casiquiare and Solano, large but flat laja with many depressions with patches of grasses and sedges and patches of shrubs, 9 Nov. 1987 (fl), R. Liesner & G. Carnevali 22930 (NY); Río Orinoco, Río Cuao, near Danta (Tapir) Falls, 125 m, 19 Nov. 1948 (fl, fr), B. Maguire & L. Politi 27326 (MO, NY); mouth of Cano Arapacua, Río Pacimoni, 120 m, 8 Apr. 1953 (fl, fr), B. Maguire & J.J. Wurdack 34918 (NY, UB, US); Ríos Pacimoni, Yatua, Casiquiare, Piedra Catipan, 100-140 m, 30 Nov. 1953 (fl), B. Maguire et al. 36474 (NY [2 sheets], US, VEN); Río Negro, slopes of Piedra Nunca (just N of Piedra Cocuy), 200-250 m, 10 Apr. 1953 (fl), B. Maguire & J.J. Wurdack 34951 (NY); Depto. Atures, near Puerto Ayacucho [ca. 5°39'N, 67°37'W], 120 m, 17 Jan. 1968 (fl), L. Ruiz-Terán 4286 (NY); Depto. Río Negro, near San Carlos de Río Negro [ca. 1°50'N, 67°5'W], 120 m, 17 Jan. 1968 (fl-imm fr), L. Ruiz-Terán 4968 (BR, NY, US); San Carlos de Río Negro, ca. 4-5 km along track running eastwards out of the village from airstrip, sandy soil, 120 m, 8 Mar. 1984 (fl, fr), B.L. Stannard 67 (U); Selvas pluviales y de rebalse del Río Pasimoni, entre la boca y Piedra Arapacoa, 1°50'N, 66°45'W, 175 m, 20-30 Oct. 1986 (fl, fr), B. Stergios et al. 9537 (MO, NY, PORT); Piedra Aracapua y laguna adyacente, selvas de rebalse (Iguapó), Misión Proyecto Vavilov, 1°51'5"N, 66°35'11"W, 3 Nov. 1994 (fl, fr), B. Stergios et al. 16213 (NY, PORT, US); San Carlos de Río Negro, sabana y selva alrededores del aeropuerto, 125 m, 17-18 Apr. 1970 (fl), J.A. Steyermark & G. Bunting 102712 (NY, P); 10-30 km E of Puerto Ayacucho towards Gavilan, 5°35'N, 67°22'W, 120 m, 11 Jun. 1977 (fl), J.A. Steyermark et al. 113887 (NY); Puerto Ayacucho, 100 m, 18 May 1940 (fl), Ll. Williams 12979 (US), Apr. 1940 (fl), Ll. Williams 13450 (US); camino de Yavita, sítios abierto a lo largo del camino, 128 m, 22 Jan. 1942 (fl, fr), Ll. Williams 13876 (S, US), 2 Feb. 1942 (fl, fr), Ll. Williams 14147 (NY, S, US); sabana de Atures, 120 m, 26 Jun. 1942 (fl), Ll. Williams 15914 (US). **Bolívar:** Mun. Uriman, Campo de Uriman, Morichal, 400 m, 7 Sep. 1954 (fl), L. Bernardi 1683 (MER, NY); Río Carrao, Alto Caroní, playos del Salto Hacha, 6°15'N, 62°51'W, 350 m, Mar. 1954 (fl), F. Cardona Puig 2871 (NY, US, VEN); Via Upata, Río Caroni, 25 Aug. 1947 (fl), H.M. Curran 216 (NY); 61 km SE of Upata (20 km SE of Villa Lola), *Trachypogon* savanna on low hills with very widely spaced *Curatella americana*, 2 Dec. 1973 (fl), G. Davidse et al. 4625 (MO, NY); Dto. Piar, Kamarata, disturbed *Trachypogon* savanna around village, 5°42'N, 62°21'W, 470 m, 23 Nov. 1982 (fl), G. Davidse & O. Huber 22519 (NY); Mun. Sucre, sabana graminosa abierta dom. por *Tragopogon*, 7°30'N, 65°16'W, 50 m, Jul. 1990 (fl), L. Delgado 1246 (NY, PORT, US, VEN); Mun. Piar, sabana

graminosa arbustiva densa, 7°4'N, 62°5'W, 100 m, Apr. 1994 (fl), *L. Delgado 2201* (NY, PORT, US, VEN); Mun. ASC. Farreras, Maripa-Aripao, 7°29'N, 65°20'W, 80 m, Feb. 1990 (fl), S. Elcoro 680 (NY, PORT); Distrito Pilar, sabanas en los alrededores del Hato Terecay, ca. 16 km N de El Manteco, 7°28'N, 62°27'W, 300 m, 24 Sep. 1982 (fl), *O. Huber & C. Alarcón 6592* (NY, US, VEN); N of Guasipati [ca. 7°40'N, 61°57'W], 1 km S of Santa Rosa, 260 m, 19 Mar. 1974 (fl, fr), *A.H. Gentry et al. 10752* (MO, US); Villa de Upata [ca. 8°10'N, 62°24'W], dans la savanne, 1864 (fl), *R. Gros sourdy 43* (P); Mun. Sifontes, Tumeremo, Via Los Guaicas, roadside, 7°13'N, 61°26'W, 160 m, 19 Feb. 2000 (fl, fr), *C. Knab-Vispo 1185* (NY); La Gran Sabana, 2 km W of Kamarata, 450 m, 22 Feb. 1967 (fl), *T. Koyama & G. Agostini 7228* (NY [2 sheets]); Mun. Piar, entre El Miamo y El Palmar, 7°40'N, 61°46'W, 240 m, Apr. 1986 (fl), *E. Sanoja 1027* (NY); Reserva Forestal Imataca, Pica de la CGV que conduce al medio Río Botanamo (cerca de la confluencia con le Río Corumo) desde la carretera Casa Blanca-Anacoco, selva pluvial, 19-20 Feb. 1983 (fl), *B. Stergios et al. 5392* (MO, NY, PORT); between Ciudad Bolívar and Río Caroní, sandy openings along small stream in savana, 100 m, 31 Jul. 1944 (fl), *J.A. Steyermark 57576* (NY, US); vicinity of Urimán, large savanna, 300 m, 30 Apr. 1953 (fl, fr), *J.A. Steyermark 75281* (NY, US, VEN); Altiplanicie de Nuria, more or less level forest in vicinity of Quebrada Agua Linda, 8-11 km E of Campamento Nuria, E of Hato de Nuria, near pica 35 and 106, 19 Jul. 1960 (fl), *J.A. Steyermark 86515* (NY); Río Toro (Río Grande), between Río La Reforma and Puerto Rico, N of El Palmar, 200-250 m, 12 Dec. 1960 (fl), *J.A. Steyermark 87966* (F, NY [3 sheets], US); NW of Camp, 5 km from Hato de Nuria, E of Miamo, Altiplanicie de Nuria, 400 m, avanna bordering morichal, 400-450 m, 10 Jan. 1960 (fl), *J.A. Steyermark 88332* (NY [3 sheets]); 261 km SW of Caicara del Orinoco, small stream in savanna, 6°10'N, 67°16'W, 110 m, 5 Sep. 1985 (fl), *J.A. Steyermark et al. 131389* (MO, NY); Distrito Piar, vicinity of Guadequen (Buadequen), Río Acanán (affluent of Río Carrao), W of Cerros Los Hermanos, brushy draw at head of low place of savanna by forest border, 5°56'N, 62°17'W, 470 m, 13-3 and 20 May 1986 (fl), *J.A. Steyermark et al. 131970* (MO, NY, US, VEN); Bajo Caura, Guayapo, 100 m, 12 Apr. 1939 (fl, fr), *Ll. Williams 11770* (US); Bajo Caura, 100 m, 21 Apr. 1939 (fl, fr), *Ll. Williams 11953* (US). **Delta Amacuro:** Río Amacuro, Venezuela-Guyana frontier, Sierra Imataca, downstream from mouth of Yarakita River, 65-80 m, 9 Nov. 1960 (fl), *J.A. Steyermark 87445* (NY [3 sheets]). **Monagas:** Carretera de Barrancas a Temblador, Puente sobre el Río Uracoa, 8°49'N, 62°27'W, 50 m, vegetación de morichal, 19 Apr. 1973 (fl), *G. Agostini & T. Agostini*
1663 (NY, U, US [2 sheets], VEN); between Montaña de Aguate and Colorado, along Río Caripe, E of Caripe [ca. 10°10'N, 63°30'W], 700–800 m, 19 Apr. 1945 (fl), *J.A. Steyermark* 62237 (NY [2 sheets], US, VEN); Caripito [ca. 10°6'N, 63°6'W], praderas megatermicas, 11 Mar. 1948 (fl), *F. Tamayo* 3492 (NY, US, VEN); open swampy margins of Laguna Grande, just S of La Pica, 3 Oct. 1955 (fl), *J.J. Wurdack & J.V. Monachino* 39556 (NY). **Tachira:** W of El Piñal [San Rafel del Piñal, 7°31'N, 71°57'W], W of bridge over Río Frio, 250–300 m, 27–30 Aug. 1966 (fl), *J.A. Steyermark & M. Rabe* 96664 (NY, U); Montañas de Guafitas, just W and N of El Piñal, sandy steep slopes, ridges and quebradas, 7°32'30"N, 71°58'20"W, 250–300 m, *J.A. Steyermark et al.* 119526 (NY). **Zulia:** Perijá, 30 m, 3 Jan. 1951 (fl, fr), *Hermano Gines* 1569 (US). **State unknown:** Without locality, s.d. (fl), *L. Ruíz Terán* 4295 (NY).

**TRINIDAD:** Erin Savanna, 20 Dec. 1977 (fl, fr), *C.D. Adams* 14323 (NY); Piarco Savanna [ca. 10°35'N, 61°20'W], S of Arouca, 27 Feb. 1920 (fl, fr), *E.G. Britton et al.* 86 (NY); Piarco Savannah, 12 Dec. 1919 (fl, fr), *W.E. Broadway s.n.* (US 1047665) (US); Piarco Savannah, S of Arouca, 27 Feb. 1920 (fl, fr), *W.E. Broadway* 86 (US); Piarco Savannah, 10 Mar. 1929 (fr), *W.E. Broadway* 7142 (BM, US [2 sheets]); locality unknown, s.d. (fl), *Collector Unknown* 3125 (US); 3 miles S of Cunupia, along the Southern Main Road, 40 m, treeless savanna dominated by *Axonopus & Andropogon*, 8 Aug. 1970 (fl), *G. Davidse* 2602 (US); Piarco Savanna, 100 ft [30 m], 19 Jun. 1963 (fl), *M.B. Kalloo* B-643 (NY), 21 Feb. 1971 (fl), *B-984* (NY); Erin Savanna, 11 Aug. 1977 (fl), *D. Philcox & E. Ramcharan* 8167 (NY, P); Erin Savanna, open savanna, 65 m, 25 Aug. 1962 (fl, fr), *J.W. Purseglove* 6462 (K, NY, US); Piarco Savanna, 7 Dec. 1891 (fl), *E. Warming* 126 (C, US), s.d., 127 (C), 7 Dec. 1891 (fl), 150 (C, US); St. George, Mausica Savanna, 2–2.5 mi S of D’Abadia P.O., 25 Aug. 1959 (fl), *G.L. Webster* 9887 (S); St. Patrick, Erin Savanna, E of Buenos Ayres, 26 Aug. 1959 (fl, fr), *G.L. Webster et al.* 9921 (S); Pitch Lake, Sep. 1901 (fl), *F. Went* 1045 (U).

**GUYANA:** Waruni-Ituni Savannahs, along Berbice-Rupununi Cattle Trail, 1919 (fl, fr), *A.A. Abraham* 27 (NY); Waruni-Ituni Savannahs, along Berbice-Rupununi Cattle Trail, 24 May 1919 (fl, fr), *A.A. Abraham* 129 (NY); Kaieteur Plateau, Potaro River, 1300 ft, May 1926 (fl, fr), *A.H.G. Alston* 549 (NY); Koriabo River, savanna, 20–22 Jul. 1934 (fl-fr), *W.A. Archer* 2361 (US); Upper Mazaruni River Region, vicinity of Kako, an Akawaio Indian Village on the Kako River near its junction with the Mazaruni River, 5°45'N, 60°35'W, 500 m, 14 Apr. 1987 (fl), *B.M. Boom & D. Gopaul* 7289 (MO, NY); Immatumari, sandy soil, s.d., *A.J. Cheong* 3 (US); Upper

Demerara-Berbice Region, Mabura Hill, 60 km N on Mabura Hill-Linden Hwy, 5°50'N, 58°30'W, 60 m, 4 Jan. 1996 (fl), *H.D. Clarke* 769 (CAY, NY, U, US); Upper Takutu-Upper Essequibo Region, Sand Creek to Shea Village, 2°58'N, 59°31'W, 28 May 1996 (fl), *H.D. Clarke* 1762 (U, US), 1792 (NY); Upper Takutu-Upper Essequibo Region, Maparri River, base of waterfall, 3°20'N, 59°15'W, 2 Jun. 1996 (fl), *H.D. Clarke* 1879 (MO, US); Upper Takutu-Upper Essequibo Region, Bat Mountain, 2°10'N, 59°10'W, 25 Jul. 1996 (fl), *H.D. Clarke* 2459 (MO, US); Upper Takutu-Upper Essequibo Region, Makarapan Mountain, base of S side, 1 m S of camp, 3°56'59"N, 58°52'3"W, 100 m, 29 Sep. 1997 (fl), *H.D. Clarke et al.* 6943 (NY); Cuyuni-Mazaruni Region, Ataro River, below falls of Wayalayeng Village, 5°14'8"N, 60°30'37"W, 488 m, 29 Jun. 2004 (fl, fr), *H.D. Clarke et al.* 12260 (P); Warenero [?], Berbice, brown sand savanna, Oct. 1973 (fl, fr), *A. Cooper* 37 (U); St. Cuthbert’s Mission, Mahaica River, white sand savanna, Sep. 1966 (fl), *D.H. Dans* 28 (NY); N Rupununi, savanna, Apr. 1968 (fl), *D.H. Dans* 755 (NY), 902 (NY); Pomeroon District, Moruka River, Mora Landing, 14 Aug. 1921 (fl, fr), *J.S. De La Cruz* 975 (NY, US), 26 Aug. 1921 (fl), 1057 (NY, US); between Demerara and Berbice Rivers, 5°50'N, 15–19 Jul. 1922 (fl, fr), *J.S. De La Cruz* 1578 (NY, US), 1603 (NY, US), 1604 (NY, US); Upper Rupununi River, near Dadanawa, 24–29 Jul. 1922 (fl), *J.S. De La Cruz* 1800 (NY); Pomeroon District, Moruka River, Mora Landing, 21–23 Aug. 1922 (fl, fr), *J.S. De La Cruz* 1816 (NY, US); vicinity of Bartica, on Essequibo River, 6°25'N [58°37'W], 3–12 Sep. 1922 (fl), *J.S. De La Cruz* 1860 (NY, US); Upper Mazaruni River, ca. 60°10'W, 22 Sep.–6 Oct. 1922 (fl, fr), *J.S. De La Cruz* 2118 (NY, US), 2239 (NY, US), 2392 (NY, US); Demerara River, Malali, ca. 5°35'N, 30 Oct.–5 Nov. 1922 (fl, fr), *J.S. De La Cruz* 2630 (NY, US); Upper Mazaruni River, Kamakusa, ca. 59°50'W, 23–29 Nov. 1922 (fl, young fr), *J.S. De La Cruz* 2765 (MO, NY, US); Northwest District, Waini River, 8°20'N, 59°40'W, 3–18 Apr. 1923 (fl, fr), *J.S. De La Cruz* 3573 (NY, US); Northwest District, Wanama River, 7°45'N, 60°15'W, 10–23 May 1923 (fl, fr), *J.S. De La Cruz* 3882 (NY, US), 4047 (NY, US); Kaieteur Falls, Potaro River, 23 Oct–3 Nov. 1923 (fl, fr), *J.S. De La Cruz* 4448 (NY, US); Potaro-Siparuni Region, Upper Potaro River, Chenapou Village, Aklapanan Savanna, motorkike trail from Chenapou Creek mouth towards Paramakatoi Village, eight hills from creek mouth (ca. 10 km), then small trail to the left, crossing a small waterfall, savanna frequently burned, dominated by *Pteridium arachnoideum* and *Brocchinia micrantha*, on white-sand soil, 4°55'28"N, 59°34'33"W, 510 m, 16 Jun. 2016 (fl, imm fr), *P.G. Delprete & P. Benjamin* 12860B (BRG, CAY, MO); Potaro-
Siparuni Region, Upper Potaro River, Chenapou Village, disturbed vegetation around village, frequently cut, on lateritic-sandy soil, 4°59'0"N, 59°34'35"W, 470 m, 17 Jun. 2016 (fl, imm fr), *P.G. Delprete & P. Benjamin 12861* (B, BRG, CAY, K, L, MG, MO, NY, P, US), *12863* (BR, BRG, CAY, F, K, US); Potaro-Siparuni Region, Iwokrama Rainforest Reserve, N of Surama Lake, trail to Lake Surama, frequently burned savanna, brown sand, 4°10'N, 59°3'W, 200 m, 15 Mar. 1995 (fl), *C. Ehringhaus et al. 60* (CAY, NY, U, US); Rupununi Distr., S Kanuku Mountains, Sand Creek Village, 6 Feb. 1985 (fl), *C. Feuillet 1628* (CAY); Rupununi River, Sand Creek, Sep. 1948 (fl), *Forest Department of British Guiana BW 156* (NY); Cock of the Rock, Dec. 1948 (fl, imm fr), *Forest Department of British Guiana BW 649* (NY); Rocky hillside, 0.5 mi N of Camp 1, 6.1952 (fl, imm fr), *Forest Department of British Guiana Field No. G318* (*Record No. 7397*) (NY); Parakara Savanna, 8 Dec. 1952 (fl), *Forest Department of British Guiana Field No. G633* (*Record No. 7648*) (NY); Upper Takutu-Upper Essequibo Region, Baboon Hill, 1.5 km S of Sand Creek Village, 3°0'N, 59°13'W, 120-150 m, 21 Jun. 1989 (fl, fr), *L.J. Gillespie et al. 1734* (NY, U, US); Upper Takutu-Upper Essequibo Region, SE Kanuku Mountains, Mt. Ishtaban, upper slopes, 3°15'N, 59°25'W, 500-700 m, 26 Jun. 1989 (fl), *L.J. Gillespie et al. 1895* (NY); Upper Demerara-Berbice Region, ca. 27 km from Ituni, along Ituni-Kwakwa rd., 5°22'N, 59°7'W, 30-60 m, 16 Jan. 1990 (fl, fr), *L.J. Gillespie et al. 2965* (CAY, NY, U, US); second-growth thickets along Potaro River, Tumutumari [ca. 5°22'N, 59°0'W], 18 Jun-8 Jul. 1921 (fl), *H.A. Gleason 44* (NY, US), *45* (NY, US); Rockstone, in and about the village, 13-30 Jul. 1921 (fl), *H.A. Gleason 703* (NY, US); 2 miles S of St. Ignatius, near Takutu, grassland with scattered trees, dominant: *Curatella, Byrsonima, Trachypogon, Fimbristylis*, 115 m, 1 Aug. 1963 (fl-fr), *R. Goodland 265* (NY [2 sheets], US); Stand 7, SE of Kumu Creek, high lateritic outcrops, sand and gravel, grassland with scattered trees, dominant: *Curatella, Byrsonima, Trachypogon, Fimbristylis*, 350 ft [ca. 106 m], 14-15 Aug 1963 (fl-fr), *R. Goodland & P.F. Maycock 422* (NY); Stand 21, Nappi Flats, grassland with scattered trees, dominant: *Curatella, Byrsonima, Trachypogon, Fimbristylis*, 350 ft [ca. 100 m], 11 Sep. 1963 (fl), *R. Goodland & R. Persaud 653* (NY); Inasa Creek, grassland with scattered trees, dominant: *Curatella, Byrsonima, Trachypogon, Fimbristylis*, 115 m, 20 Sep. 1963 (fl-fr), *R. Goodland 846* (US); Rupununi Savanna, natural pond near Karanambo Ranch, 3°45'N, 59°19'W, 100-250 m, 21 Nov. 1992 (fl, fr), *A.R.A. Gorts-van Rijn et al. 361* (CAY, NY, U [2 sheets]); Savanna beyond Aratak Mission across Kamuni Creek from Santa Mission, 6°30'N, 58°23'W, 0-200 m, 28 Nov. 1992 (fl, fr), *A.R.A. Gorts-van Rijn et al. 453* (CAY, P, U); Junction of Mazaruni and Cuyuni Rivers, at Kalacoon, Jun. 1927 (fl), *E.H. Graham 101* (NY); Ituni trail by goat farm, 1 Jul. 1976 (fl, fr), *M.S. Grewal 212* (U); Potaro-Siparuni Region, Kaieteur Falls National Park, around guest house, 5°10'N, 59°29'W, 500 m, 18 Apr. 1988 (fl), *W. Hahn et al. 4744* (NY, U, US); St. Ignatius, Rupununi, most marginal land between savanna and marsh, 14 Jul. 1958 (fl, imm fr), *S.G. Harrison 1283* (NY); Upper Demerara-Berbice Region, lower 3 km of Macouria River, N and S banks, 6°25'N, 58°34'W, 0-15 m, 7 Dec. 1992 (fl), *T.W. Henkel & M. Chin 335* (NY, U, US); Essequibo Island-West Demerara Region, lower 7 km of Tiger Creek, 6°30'N, 58°39'W, 0-15 m, 12 Dec. 1992 (fl, fr), *T.W. Henkel & M. Chin 472a* (INPA, NY); Potaro-Siparuni Region, Kaieteur Pateau, 3 km WNW from falls along Kaieteur-Kuribong trail, 5°12'N, 59°30'W, 540 m, 22 Jul. 1993 (fl), *T.W. Henkel & R. Williams 2390* (NY, U, US); Est Berbice Region, Berbice Savanna near Takama Army Base, 5°43'N, 57°57'W, 100 m, 14 Aug. 1993 (fl, fr), *T.W. Henkel et al. 2515* (CAY, MO, NY, US); Upper Takutu-Upper Essequibo Region, S Rupununi savanna, 5 km S of Aishton along rd to Marudi Mountains, 2°23'N, 59°18'W, 200 m, 9 Sep. 1993 (fl, fr), *T.W. Henkel et al. 2799* (CAY, MO, NY, U, US); Upper Takutu-Upper Essequibo Region, South Rupununi Savanna, Aishalton village, 2°31'N, 59°20'W, 200 m, 3 Aug. 1993 (fl, fr), *T.W. Henkel & R. James 3970* (NY, U, US); Potaro-Siparuni Region, Pakaraima Mts., Upper Ireng River watershed, Kaatnang River, near base of Malakwalai-Tipu, 4°48'N, 60°12'W, 700 m, 9 Jul. 1994 (fl), *T.W. Henkel & M. Chin 5489* (CAY [2 sheets], NY, U, US); Upper Takutu-Upper Essequibo Region, northern Rupununi Savanna, 2.5 km NW of Karanambo Ranch, 3°45'N, 59°19'W, 100-120 m, 11 Mar. 1992 (fl, fr), *B. Hoffman 1248* (CAY, NY, U, US); Cuyuni-Mazaruni Region, Pakaraima Mountains, Kurupung River, landing at base of Makrega Falls, 6°7'N, 60°20'W, 85 m, 18 Jul. 1992 (fl), *B. Hoffman 2075* (MO, NY, US); Upper Takutu-Upper Essequibo Region, NW Kanuku Mountains, 12 km ESE of Nappi Village in foothills, 3°23'N, 59°29'W, 170 m, 10 Feb. 1993 (fl, fr), *B. Hoffman 3664* (MO, US); Kaieteur Savanna, 1936 (fl), *G. Hollister s.n.* (NY); Rupununi District, Lethem, stream bank 1 mile N of city, 14 Apr. 1956 (fl), *H.S. Irwin 570* (US); Rupununi District, Lethem, Swail S of Lethem, near Takutu River, 15 Apr. 1956 (fl, fr), *H.S. Irwin 620* (US), 622 (US); Rupununi Savanna, Lethem, 3°23'N, 59°47'W, 80 m, 27 Oct. 1987 (fl, fr), *M.J. Jansen-Jacobs et al. 552* (CAY, NY, P [2 sheets], U, US); Rupununi Savanna, Mora Savanna, near Toroebaroe Creek, 3°23'N, 59°29'W, 145 m, 21 Nov. 1987 (fl, fr), *M.J. Jansen-Jacobs et al. 1114* (CAY, U, US); Rupununi Savanna, Nappi Village, 3°25'N, 59°35'W, 110 m, 1
Dec. 1987 (fl, fr), *M.J. Jansen-Jacobs et al. 1379* (CAY, NY, P, U, US); Essequibo River, Gunn's Savanna, 1°39'N, 58°38'W, 240-260 m, 3 Sep. 1989 (fl, fr), *M.J. Jansen-Jacobs et al. 1434* (CAY, MO, NY, U), 26 Sep. 1989 (fl, fr), 1836 (CAY, NY, U); Rupununi District, Kuyuwini Landing, Kuyuwini River, savanna, 10 Feb. 1991 (fl, fr), *M.J. Jansen-Jacobs et al. 2512* (CAY, NY, P, U); Rupununi District, Kuyuwini Landing, Kuyuwini River, savanna, 29 Oct. 1992 (fl), *M.J. Jansen-Jacobs et al. 3156* (CAY, NY, P, U, US); Rupununi District, Manari, savanna, 3°26'N, 59°46'W, 80 m, 12 Jul. 1995 (fl), *M.J. Jansen-Jacobs et al. 4419* (CAY, NY, P, U); Kanuku Mountains, Moco Moco River, 3°18'N, 59°39'W, 140-180 m, 14 Jul. 1995 (fl), *M.J. Jansen-Jacobs et al. 4463* (MO, U, US); South Rupununi District, Wakadanawa Savanna, 1°60'N, 59°34'W, 290 m, 8 Sep. 1997 (fl, fr), *M.J. Jansen-Jacobs et al. 5424* (MO, NY, P, U); Corentyne River, Nov. 1879 (fl), *G.S. Jenman 294* (P); Bartica, Nov. 1888 (fl, fr), *G.S. Jenman 4723* (BM, NY [mounted on the same sheet with Jenman 5453], US); Potaro River, Kaieteur Savannah, Sep.-Oct. 1881 (fl), *G.S. Jenman 855* (P); Mazaruni River, Aug. 1889 (fl, fr), *Jenman, G.S. 5453* (BM, NY, US); Rupununi River, Ruimatter, Oct. 1889 (fl, fr), *G.S. Jenman 5520* (US); Japacourna [?], Jan-Feb. 1901 (fl, fr), *G.S. Jenman 7829* (NY); creek 2 miles S of Araquoi, 2 Nov. 1957 (fl, fr), *C.D.K. Kooh 88* (NY, U); Bartica, Waipiole Luce, 7 Oct. 1922 (fl), *H. Lang 133* (US); Rupununi Savanna, in directionem borealem de montibus Kanuku, maurisielaagte a.d. voet v.d. Manakaparuheuvels ten z.o. van Jawarie, 14 Feb. 1959 (fl fr), *J. Lanjouw 760* (U); Cuyuni River, Camaria Road, 15-30 m, 12 Feb. 1951 (fl, fr), *Lorkie, J.R. 214* (S); Rupununi District, Manari, 3°28'N, 59°41'W, 21 Oct. 1979 (fl, fr), *P.J.M. Maas & L.Y.T. Westra 3702* (U), 22 Oct. 1979 (fl), 3754 (S, MO, NY, U); Sand Hills, W of Abary River, 5°55'N, 57°54'W, 4 Oct. 1981 (fl), *P.J.M. Maas et al. 5512* (MO, NY, P, S, U); Rupununi River, Monkey Pond Landing, SW of Mt. Makarapan, 3°53'N, 58°55'W, 10 Sep. 1988 (fl), *P.J.M. Maas et al. 7346* (MG, MO, U, US), 11 Sep. 1988 (fl), 7380 (CAY, U, US); Kaieteur Plateau, savannas, 14 May 1944 (fl), *B. Maguire & D.B. Fanshawe 23472* (NY); Mount Ayanganna, Pakaraima Mountains, between Ayanganna and Chinowieng, 1000-1200 m, 7-8 Feb. 1955 (fl), *B. Maguire et al. 40639* (NY); Southern Pakaraima Mountains, Kamana Savanna, en route to Kopinang Falls, pebbly red laterite, 28 Aug. 1961 (fl), *B. Maguire et al. 45974A* (NY); Ando Savanna, Sukabi River (affl. Ireng River), 800 m, 16 Sep. 1961 (fl, fr), *B. Maguire et al. 46184A* (MO, NY); Waranama Ranch, Wiruni Ituni Savannah, Sep. 1929 (fl, fr), *E.B. Martyn 86* (NY); Upper Takutu-Upper Essequibo Region, woods between Lake Surama and Surama Village, 4°9'N,

59°3'W, 65-75 m, 22 Feb. 1990 (fl), *T. McDowell et al. 1984* (CAY, NY); East Berbice-Corentyne Region, Baba Grant Sawmill landing, along banks of Corentine River, above Cow Falls, 5°0'N, 57°42'W, 10-25 m, 21 Apr. 1990 (fl, fr), *T. McDowell & D. Gopaul 2403* (CAY, NY, U, US); Cuyuni-Mazaruni Region, Waramaden, around village and between airstrip and creek, 5°48'N, 60°44'W, 490 m, 12 Jun. 1990 (fl, fr), *T. McDowell & D. Gopaul 3209* (CAY, MO, NY, US); Potaro-Siparuni Region, in woods along creek at SW of Eagle Mt., along Potaro-Konawaruk Rd., 5°12'N, 59°7'W, 107-152 m, 12 Oct. 1990 (fl, fr), *T. McDowell & D. Gopaul 3506* (CAY, MO, NY, U, US); Cuyuni-Mazaruni Region, along road from Arching airstrip to Mazaruni River 6°10'N, 60°7'W, 137 m, 22 Feb. 1991 (fl, fr), *T. McDowell 4055* (MO, NY, U, US); Cuyuni-Mazaruni Region, route to Crommer's Farm, 2 km SE of Haiamatiupu, 5°27'N, 60°31'W, 550 m, 21 Jun. 1991 (fl-imm fr), *T. McDowell & B. Hoffman 4761* (NY); Mazaruni River, settlement, 8 Aug. 1933 (fl, fr), *C.D. Mell & R.C. Mell 226* (NY, US); on rd. to St. Cuthbert's Mission, ca. 10 km in from the Linden-Soesdyke Hwy, ca. 50 km S of Georgetown, open savannah, 8 Aug. 1976 (fl), *S.A. Mori et al. 8039* (NY, US); Mazaruni-Potaro District, Bartica-Potaro Road, 24 miles S of Bartica at Forest Service "24 mile Camp", 50-125 m, 20 Aug. 1976 (fl), *S.A. Mori et al. 8183* (NY, US); East Berbice-Corentyne Region, Baranka Landing, Piruru Savanna, 5°11'21"N, 57°17'7"W, 20 m, 1 Apr. 1995 (fl, fr), *P. Mutchnick 1143* (NY, U, US); Upper Mazaruni District, Kamarang River, Waramadong, 2800 m, 6 Oct. 1938 (fl, fr), *A.S. Pinkus 21* (NY, US); Mazaruni-Potaro Region, Pakaraima Mts., N of Imbaimadai, 5°43'N, 60°18'W, 518 m, 20 Jun. 1986 (fl), *J.J. Pipoly & K. Alfred 7880a* (US); Mazaruni-Potaro Region, Pakaraima Mts., Imbaimadai Creek, W of Imbaimadai, 5°42'N, 60°18'W, 500 m, 22 Jun. 1986 (fl), *J.J. Pipoly & K. Alfred 7969* (CAY, NY, P, U, US); East Berbice-Corentyne Region, S Awara Savanna, ca. 35 mi S of Torani Canal, on Canje River, 3 mi SE of Digitima Creek 5°43'N, 57°32'W, 0-25 m, 18 Dec. 1986 (fl), *J.J. Pipoly et al. 9385* (NY); East Berbice-Corentyne Region, Left (S) bank of Upper Canje River, W of Digitima Creek, 5°36'N, 57°35'W, 1-25 m, 19 Dec. 1986 (fl), *J.J. Pipoly et al. 9457* (MO, NY, US); Upper Demerara-Berbice Region, ca. 15 km E of Rockstone, on Linden-Rockstone rd., secondary *Eperua* forest, *Dimorpandra* thickets & Savannas, deep white sand, sandy loam, loamy sand, 5°58'N, 58°30'W, 1-50 m, 19 Dec. 1986 (fl), *J.J. Pipoly & G. Gharbarran 9575* (NY); East Berbice-Corentyne Region, Canje River, 0.75 km N of Ekwarun River, disturbed tall evergreen forest on brown loamy sands, 5°20'N, 57°38'W, 1-25 m, 9 Apr. 1987 (fl, fr), *J.J. Pipoly et al. 11332* (CAY, MO, NY, US); Bartika [now
Bartica], along road in the jungle 12–15 miles from the town, 28 Aug. 1935 (fl), *D. Potter* 5341 (BR); Upper Mazaruni District, Kamarang, footpath in the village and along airstrip, man-made savanna, 5°52′N, 60°37′W, 525 m, 18 Oct. 1985 (fl), *H.E. Robinson* 85-0054 (NY, US); Cuyuny River, sides of portage road at lower Camaria Landing, 23 Nov. 1929 (fl), *N.Y. Sandwith* 655 (NY, RB); without locality, s.d. (fl), *R.H. Schomburgk s.n.* (FI-Webb); without locality, s.d. (fl), *R.H. Schomburgk s.n.* (P P00748113); Berbice, moist savannahs, 1837 (fl), *R.H. Schomburgk Ser. I, 15* (BM, BR, P, U [2 sheets], US [2 sheets]); without locality, moist savannhas, 1837 (fl, fr), *R.H. Schomburgk Ser. I, 95* (BM, BR, FI-Webb, NY, P, US); Pirara, 1838 (fl, fr), *R.H. Schomburgk Ser. I, 723* (FI-Webb, mounted on the same sheet with two branches of *Declieuxia chiococcoides* Kunth); Basin of Rupununi River, near mouth of Charwair Creek, ca. 2°35′N, 1–4 Nov. 1937 (fl, fr), *A.C. Smith* 2390 (NY, P, S, U, US); without locality, s.d., *F.L. Splitgerber s.n.* (P P00729339); West-Demerara, Mabura Hill, 180 km SSE of Georgetown, Demerara Landing, 7 km up the Great Falls, 5°20′N, 54°40′W, 0–50 m, 14 Jun. 1988 (fl), *H. ter Steege & G.J. ter Steege* 407 (CAY, U, US).

**SURINAME:** Potaro River, Kangaruna Landing, Oct. 1922 (fl, fr), *A.A. Abraham* 349 (U); Tumuc-Humac Mountains, Talouakem, 2°31′N, 54°45′W, 430–530 m, open vegetation at base of granite, 9 Aug. 1993 (fl, fr), *P. Acevedo-Rodriguez et al.* 5958 (CAY [2 sheets], MO, NY, U, US); without locality, 1841 (fl, fr), *M. Berthoud-Coulon* 176 (BM, NY [2 sheets]); Coppename River, Aug. 1901 (fl), *H. Boon* 1077 (U), Sep. 1901 (fl), 1147 (U); Suriname River, 1908 (fl, fr), *Coll. Indigen. s.n.* (U 001806) (U), s.n. (U 0016988) (U); without locality, s.d. (fl), *Collector Unknown [probably Hostmann] s.n.* (BR 825740) (BR, probable type material of *S. trichantha*); Brokopondo, Brownsberg Nature Park, trail to Mazaroni Val., primary forest, light gaps, 4°56′N, 55°11′W, 400–450 m, 23 Jan. 1999 (fl, fr), *P.G. Delprete & Flora of the Guianas Symposium Tour Participants* 7082 (BBS, CAY, MO, NY, P, U, US); Sipaliwini Distr., Sipaliwini Village, savanna N of the airstrip, with sparse trees of Curatella americana (3–5 m tall), herbaceous layer dominated by grasses and sedges, with many species of flowering plants, 2°1′39″N, 56°7′23″W, 240 m, 16 Apr. 2014 (fl, imm fr), *P.G. Delprete & G.P. Marjanom* 12349 (BBS, CAY, GB, K, L); Sipaliwini Distr., Sipaliwini Village, trail from the Sipaliwini Village to the Sipaliwini River, secondary forest with sparse trees 25–30 m tall and canopy 15–20 m tall, herb 50–130 cm tall, found as sparse individuals in large standing of more than 100 individuals, 2°1′14″N, 56°7′23″W, 250 m, 16 Apr. 2014 (fl, imm fr), *P.G. Delprete & G.P. Marjanom* 12386 (B, BBS, CAY, F, GB, K, L,

MO, US); Sipaliwini Distr., Sipaliwini Village, trail from the Sipaliwini airstrip towards the Sipaliwini Nature Reserve, ca. 2 km of the airstrip, savanna with sparse *Curatella Americana* trees (3–5 m tall) and herbaceous layer dominated by grasses and sedges, white-sand soil, herb 70–100 cm tall, 2°1′53″N, 56°6′53″W, 230 m, 18 Apr. 2014 (fl, imm fr), *P.G. Delprete & G.P. Marjanom* 12440 (BBS, CAY, F, G, L, K, NY, P), 12441 (B, BBS, BR, C, CAY, F, GB, L, K, MO, NY, P, RB, S, US); Sipaliwini Distr., Sipaliwini Village, trail from the Sipaliwini Village to the Sipaliwini River, secondary forest with sparse trees 25–30 m tall and canopy 15–20 m tall, herb 50–130 cm tall, vegetation at the edge of the trail, fine gray sandy soil, 2°1′14″N, 56°7′23″W, 250 m, 19 Apr. 2014 (fl, imm fr), *P.G. Delprete & G.P. Marjanom* 12449 (B, BBS, CAY, K, L, MG, MO, NY, P, US), 12450 (BBS, BR, CAY, F, G, GB, RB, S, US); Sipaliwini Distr., Sipaliwini Village, small savanna on the other side of the Sipaliwini River with respect to the Sipaliwini Village (S of the village), savanna with rolling hills, with sparse with sparse trees of *Curatella Americana* and a few other species, grass layer dominated by grasses and sedges, 2°1′6″N, 56°7′23″W, 220–230 m, 20 Apr. 2014 (fl, imm fr), *P.G. Delprete & G.P. Marjanom* 12466 (BBS, C, CAY, FT, L, P); Sipaliwini Distr., Sipaliwini Village, disturbed area around the village, adventitious plants, 2°1′28″N, 56°7′29″W, 240 m, 26 Apr. 2014 (fl, imm fr), *P.G. Delprete & G.P. Marjanom* 12596 (BBS, CAY, L, P); Zanderij-complex, kleihoudende zand-savanne, 31 Jul. 1952 (fl, fr), *I.J.G. Dirven* LP297 (U); Lobin-savanna inter Zanderiji I et Hannover, Aug. 1958 (fl), *J. van Donselaar & W.A.E. van Donselaar* 319 (NY); Paramaribo, in pratis et cortis cultis, 2 Jul. 1902 (fl), *E. Essed* 93 (U); Saramacca District, Experimental Farm Coebiti, 5°20′N, 55°30′W, 10 Nov. 1981 (fl), *A.P. Everarts* 524 (CAY, U, US); Dam, langs de spoorbaan, 9 Nov. 1950 (fl, fr), *J. Florschütz & P.A. Florschütz* 156 (U); Brokoboto-val in de Boven-Saramacca, 7 Feb. 1951 (fl, fr), *J. Florschütz & P.A. Florschütz* 1214 (NY, U); Paka Paka (Saramacca), naar Ebbatop (v. Asch v. Wijks-Geb.), bij km 8, granitic plateau, 10 Feb. 1951 (fl), *J. Florschütz & P.A. Florschütz* 1309 (BR, C, NY, U); Warrah-savanna on W bank of Maratakka River, 7 Apr. 1951 (fl, fr), *J. Florschütz & P.A. Florschütz* 1943 (C, NY, U); without locality, s.d. (fl, fr), *H. Focke s.n.* (U 0016948) (U); Para, poste Wran, Apr. [without year] (fl, fr), *H. Focke* 153 (U); South Central Suriname, Kaysergeberte Airstrip [ca. 3°5′N, 56°28′W], sandy soil along airstrip, 2 Dec. 1961 (fl, fr), *R. Freund & S. Freund R-28-B* (US); Powakka Savanne, 1936 (fl, fr), *J. Frickers & H.J. Müller* 7 (U), 22 (U); Palaime Savanne, Boven Tapanahoni, 2 May 1952 (fl, fr), *D.C. Geijskes s.n.* (U 0016949) (U); Coppename River, Cabalebas Creek, 26
Jan. 1915 (fl), J.W. Gonggrijp & G. Stahel 805 (U); Massif de Tumuc-Humac, Inselberg Talouakem, N slope, 2°29′N, 54°45′W, 550 m, herbaceous vegetation de savane roche (inselberg), 9 Aug. 1993 (fl), J.J. de Granville et al. 12160 (BBS, CAY, MO, P, U, US); Voltzberg, base of inselberg, 4°41′N, 56°8′W, 200 m, 28 Jan. 1999 (fl), J.J. de Granville et al. 13739 (BBS, CAY, NY, P); Central Suriname Nature Reserve, Voltzberg, Top I, 4°41′N, 56°11′W, 245 m, 13 Apr. 2002 (fl), A. Gröger et al. 1306 (U); Jodensavanne (fluv. Suriname), 14 Jun. 1957 (fl, fr), Heyligers, P.C. 342 (U); without locality, s.d. (fl, fr), F.W.R. Hostmann & A. Kappler 507 (BM, FI-Webb, NY, P, S); "In Surinam", s.d. (fl), F.W.R. Hostmann 663 (P [2 sheets, P00729342 and P05030245]); without locality, s.d., F.W.R. Hostmann 663a (MO [Acc. No. 5448229], FI-Webb [Acc. No. 080089], G [without barcode], S [Acc. No. S05-478], U [barcode 0016778]); Zuid Rivier, 3°10'–20'N, 56°29'–49'W, Kayser Airstrip, 25 km above confluence with Lucie River, 270 m, 1 Jul. 1963 (fl), H.S. Irwin et al. 53958 (NY, P, S); Zuid Rivier, 3°10'–20'N, 56°29'–49'W, 2 km above confluence with Lucie River, 220 m, 3 Jul. 1963 (fl, fr), H.S. Irwin et al. 54056 (NY, RB, UB, US); Wilhelmina Geberte, savanna ca. 2 km N of Lucie River, 3°20'–32'N, 56°26'–49'W, 225 m, 16 Jul. 1963 (fl, fr), H.S. Irwin et al. 54214 (NY); Wilhelmina Geberte, forested hill 9 km N of Lucie and 12 km W of Oost River, 3°36'–41'N, 56°30'–34'W, 275 m, 16 Jul. 1963 (fl, fr), H.S. Irwin et al. 54223 (F, NY, RB); Zuid Rivier, 3°10'–20'N, 56°29'–49'W, Kayser Airstrip, 45 km above confluence with Lucie River, 270 m, 22 Sep. 1963 (fl), H.S. Irwin et al. 55946 (MO, NY [2 sheets], RB, UB, US), 55963 (MO, NY, S); along railway near LBB, Zanderij, white-sand savanna, 2 Mar. 1976 (fl, fr), J. Jansma s.n. (LBB 15525) (U); nieuwe weg van Zanderij, naar Matta Zand, 9 Dec. 1955 (fl), A.M.E. Jonker-Verhoef & F.P. Jonker 151 (U); Zanderij, vliegveld, 30 Dec. 1955 (fl), A.M.E. Jonker-Verhoef & F.P. Jonker 323 (U); ad fl. Tap-anahoni, Oct-Nov. [without year] (fl), A. Kappler 2084 (P); Joden Savanne - Mapane Creek area, Suhoza (Suriname River), 19 Dec. 1960 (fl, fr), K.U. Kramer & W.H.A. Hekking 2380 (U); Zanderij [ca. 50 km S of Paramaribo], savanna, 26 Dec. 1960 (fl), K.U. Kramer & W.H.A. Hekking 2503 (U); Saramacca Distr., Tafelberg [ca. 3°54′N, 56°10′W], Lareco-savanne, 300 m, 25 Feb. 1961 (fl), K.U. Kramer & W.H.A. Hekking 3032 (U); Paramaribo, Oct. 1972 (fl), C. Kock s.n. (U); Potaro-Siparuni Region, Kaie-teur Falls, close to the falls and the government house, disturbed scattered shrubby vegetation on white sand and exposed bedrock, terrestrial and epipetric, 5°11′N, 59°29′W, 425 m, 6 Oct. 1987 (fl, fr), L.P. Kvist et al. 3 (CAY, NY, P, U, US); Kleisavanne I, bij Vierkinderen, 5 Sep. 1948 (fl), J. Lanjouw & J.C. Lindeman 183a (U), 198

(U); Coppename River, near Raleighfalls, 10 Sep. 1933 (fl), J. Lanjouw 755 (U); via secta ab Moengo tapoe ad Grote Zwiebelzwamp, km 11.5 in savanna, 9 Oct. 1948 (fl), J. Lanjouw & J.C. Lindeman 757 (U); Indigenous Village, Kalebas Creek, 1 Oct. 1933 (fl), J. Lanjouw 1006 (INPA, U); Savannah near Brownsweg, Nov. 1933 (fl, fr), J. Lanjouw 1251 (U); Tibiti Savanne, 5 Jan. 1949 (fl), J. Lanjouw & J.C. Lindeman 1601 (IAN, NY, U, US), 6 Jan. 1949 (fl), 1618 (IAN, NY, U), 1619 (U), 1621 (U); Jodensavanne, bij ruine, 12 Apr. 1949 (fl), J. Lanjouw & J.C. Lindeman 2999 (U), 3000 (U); Zanderij, zandsavanne B, 10 Mar. 1949 (fl), J. Lanjouw & J.C. Lindeman 3270 (NY, U); savanne langs spoorbaan bij km 100, 16 Mar. 1949 (fl), J. Lanjouw & J.C. Lindeman 3315 (U), 3316 (U); Blanche Marie Vallen, Upper Nickerie River, 100 m, Apr. 1984 (fl, fr), H. Lardinois 25 (BR); Kasiwinika, yearly burned clay savanna near Kopie, 15 Jul. 1953 (fl, fr), J.C. Lindeman 4263 (U); Maratablow bij Esparracreek, bogere savanne, 8 May 1965 (fl, fr), P.J.M. Maas & J.A. Tawjoeran s.n. (LBB 5215) (U); 3 km ZO van basiskamp, 2 km beneden Blanche Marievallen, Nickerierivier, 19 Jun. 1965 (fl, fr), P.J.M. Maas & J.A. Tawjoeran s.n. (LBB 10910) (U); W-oever van Nickerie River, 3 km N van Kamisavallen, struiksavanne, 2 Jul. 1965 (fl, fr), P.J.M. Maas & J.A. Tawjoeran s.n. (LBB 11037) (U); Sanderij, Savanna II, 3 Jun. 1944 (fl), B. Maguire & G. Stahel 23663 (NY); Wilhelmina Geberte, 3°34'–36′N, 56°30'–34'W, 9 km N of Lucie and 12 km W of Oost River, 275 m, 16 Jul. 1963 (fl, fr), B. Maguire et al. 54223 (NY, UB, US); Berseba, small savanna, 12 Sep. 1954 (fl, fr), A.M.W. Mennega 13 (U); Boven Tanjimama, 20 Nov. 1954 (fr), A.M.W. Mennega 491 (U); clay savanna Berseba, near Republiek, 14 Nov. 1974 (fl, fr), A.M.W. Mennega & J. Koek-Noorman 894 (U); vicinity of Blanche Marie Waterfall, on Nickerie River, along the entrance road to the Blanche Marie Guest House, 4°45'20"N, 56°54'19"W, 60 m, 3 Feb. 1998 (fl, fr), J.S. Miller et al. 9300 (MO, U), 9306 (MO, U); western part of Great Sipaliwini Savanna, near bank of the 4-Gebroeders Creek, transect D near base camp, 283 m, 4 Jan. 1968 (fl), F.H.F. Oldenburger et al. 674 (U); Coesterie Creek (Coesewijne), creek bank, Jan. 1978 (fl, fr), J. Procter 4700 (U, UB); Upper Coppename River, in Monte Volzberg, 50 m, 21 Aug. 1920 (fl, fr), A. Pulle 226 (U); Upper Coppename River, in summit Monte Volzberg, 250 m, 22 Aug. 1920 (fr), A. Pulle 250 (U); Skull Point, near Bartica, 9 Apr. 1956 (fl), J.R. Ramsammy 13 (US); Awarra Savanne aan de Maratakka, 2 Mar. 1971 (fl, fr), D. Reeder s.n. (LBB 13464) (U); Corantyne River, Wonotobo Falls, 6 Aug. 1935 (fl, fr), H.E. Rombouts 101 (U); Upper Sipaliwini River, savanna, 23 Oct. 1935 (fl, fr), H.E. Rombouts 223 (U); Upper Sipaliwini River, savanna near Kamp II, 6 Dec. 1935 (fl),
H.E. Rombouts 332 (U); Upper Tapanahoni River, Iden-
burg Rapids, granitic islet, 13 Sep. 1936 (fl, fr), H.E.
Rombouts 603 (C, U); Litani River, Mt. Knopaiamoi [ca.
2°28′N, 54°30′W], 12 Aug. 1937 (fl), H.E. Rombouts 807
(BR, U); Upper Litani River, bij oorsprong granite, 8
Nov. 1937 (fl), H.E. Rombouts 885 (IAN, U); without
locality, s.d. (fl, fr), J.P. Schultz s.n. (U 0016944) (U);
along Zuidrivier, near confluence with Lucie River, 3 Jul.
1963 (fl, fr), J.P. Schultz s.n. (LBB 10023) (U); Zuidrivier,
savanna near Kayser Landing, 13 Aug. 1963 (fl, fr), J.P.
Schultz s.n. (LBB 10432) (U); Voltzberg (Boven-Wayom-
bo), op dun verwerings-laagje, 22 Apr. 1956 (fl), J.P.
Schultz 7610a (U); Upper Copename River, near
Raleigh Falls, 23 Sep. 1956 (fl, fr), J.P. Schultz 7819 (U);
without locality, s.d. [1837–1838] (fl), F.L. Splitgerber s.n.
(BM 000614321) (BM); Wilhemina Geberte, 11 May 1926
(fr), G. Stahel 7049 (U [2 sheets]); Corantyne River,
Wonotobo Falls, 14 Oct. 1916 (fl, fr), G. Stahel & J.W.
Gonggrijp 3157 (U); Makatui Savanna, ca. 3.5 km SW of
Aishalton, 2°30′N, 59°15′W, white sand savanna, 18 Nov.
1982 (fl), A.L. Stoffers et al. 360 (MO, U); near Aishalton
Hospital, 2°30′N, 59°15′W, along savanna pond on clay,
18 Nov. 1982 (fl), A.L. Stoffers et al. 388 (U); near Aishal-
ton Hospital, 2°30′N, 59°15′W, white sand savanna, 19
Nov. 1982 (fl, fr), A.L. Stoffers et al. 435 (NY, S), 440 (C,
CAY, MG, MO, RB, U); 5 km SE of Aishalton Hospital,
near Lake Awakawau, 2°30′N, 59°15′W, white sand
savanna, on clay, 20 Nov. 1982 (fl), A.L. Stoffers et al. 508
(CAY, MO, U); Suriname River, near Misikai, 13 Jul.
1908 (fr), J.H.A.T. Tresling 128 (MG, U); Suriname River,
“jp. pr. coth. Kwemba” 25 Jul. 1908 (fl), J.H.A.T. Tresling
225 (U); Suriname River, near Kapiai, 5 Aug. 1908 (fr),
J.H.A.T. Tresling 287 (U); Upper Suriname River, 19 Sep.
1908 (fr), J.H.A.T. Tresling 449 (U); Suriname River,
Gran Rio, 20 Sep. 1908 (fr), J.H.A.T. Tresling 469 (U), 21
Sep. 1908 (fr), J.H.A.T. Tresling 471 (U); without locality,
s.d. (fl), J.E. Tulleken 152 (MO); Marowini River, Jul.
1904 (fl, fr), G.M. Versteeg 614 (U); Brokopondo Distr.,
Brownsberg National Park, 5 Nov. 1973 (fl), C. Vreden
s.n. (LBB 14723) (U); Brokopondo, Brownsberg Nature
Reserve Plateau weg, 4°55–57′N, 55°11–12′W, 475–500
m, 23 Jul. 1979 (fl, fr), G.L. Webster 24103 (U); Para,
between Hannover and the Onverwacht-Zanderij road,
5°28′N, 55°10′W, 20 m, 10 Jul. 1979 (fl, fr), G.L. Webster
24034 (U); Para District, prope Berlijn, Sep. 1901 (fr), F.
Went 386 (U); along Affobaka road near Berg en Dal, S
of Blauwe Berg, 20 Dec. 1962 (fl, fr), J.G. Wessel Boer 400
(U); along Kabel railroad, at km 116 near Brownsberg,
on white sand, 14 Feb. 1963 (fl, fr), J.G. Wessel Boer 684
(U); inter Flum. Copename Dextrum et montes Emma
dictos, 4°10′N, 56°20′W, 20 May 1963 (fl, fr), J.G. Wessel
Boer 14010 (U); Nat. Res. Brinckheuvel, Saban pasi

savanne, 16 Jan. 1968 (fl), J.T. Wildschut & P.A. Teunissen s.n. (LBB 12175) (U); “Surinam: Hmenge [? illegible], 1864 (fl, fr), H.R. Wullschlaegel s.n. (BR 824613) (BR); Paramaribo, 1851 (fl, fr), H.R. Wullschlaegel 257 (BR).

FRENCH GUIANA: Upper Tampoc River, Saut
Pier Kourou, on islets just below the waterfall, 6 Apr.
1977 (fl), G. Cremers 4636 (CAY [2 sheets], P, U); Tumuc
Humac, layon Orstom Koulimapopann-Mitaraka (bor-
der between French Guiana and Brazil), p.k. 7.5, great
inselberg, 550 m, 6 Aug. 1972 (fl, fr), J.J. de Granville
1129 (CAY [2 sheets], P); Tumuc Humac, base of Touk-
ouchipann, N slope, 480 m, 22 Aug. 1972 (fl, fr), J.J. de
Granville 1339 (CAY, P); Haut Oyapock, Mont St. Mar-
cel [ca. 2°36′N, 53°1′W], inselberg on S slope, 300–350
m, 30 Jul. 1975 (fl), J.J. de Granville 2603 (CAY); Upper
Mana River, Saut Ananas, 12 Aug. 1981 (fl), J.J. de Gran-
ville 4820 (BR, CAY, P, UB); Oyapock River Basin, Roche
Touatou (Inselberg), vegetation herbacée de savane-roche
[herbaceous vegetation on inselberg], 2°57′N, 52°32′W,
160 m, 16 May 1995 (fl, fr), J.J. de Granville & G. Cre-
mers 12935 (B, CAY [2 sheets], K, MO, P, U, US [some
duplicates of this collection have the label “Montagne
Emérillons, alt. 160 m, 3°15′N, 53°5′W” but this label
is wrong]); Upper Mana River, Saut Ananas, bord d'un
sentier en sous-bois clair, 31 Aug. 1962 (fl), F. Hallé 741
(CAY, NY, P, U, US); Maripasoula, Massif du Mitaraka,
Crique Alama, Inselberg Sommet Cloche, éperon de roche
affleurante au nord du dome, 2°13′40″N,
54°28′10″W, 500 m, 5 Mar. 2015 (fl), O. Poncy 2869 (B,
CAY, MO, P); Mitaraka Sud, sommet inselberg, border
of inselberg, 2°16′N, 54°31′W, 660 m, 4 Mar. 2001 (fl), C.
Sartou 807 (CAY, MO, P); Tumuc-Humac (border Brazil-
French Guiana-Suriname), Montagne Paloulouiméem-
peu, E slope, 450–500 m, granitic inselberg, 2 Aug. 1972
(fl, fr), C. Sastre 1538 (CAY [2 sheets], IAN, NY, P); Mas-
sif Mitaraka, Mitaraka S, versant S, granitic slopes, 550
m, 17 Aug. 1972 (fl, fr), C. Sastre 1713 (CAY [2 sheets],
P); Upper Maroni River (Litany), Antecum-Pata (Mala-
vate) [ca. 3°19′N, 54°4′W], 21 Apr. 1975 (fl), C. Sastre et
al. 3855 (CAY, MO, P); gros saut riviére Portal (Mana),
29 Mar. 1949 (fl), Service Forestier s.n. (P00729315) (P);
Mana River, Station Rivière Forestal, 29 May 1949 (fl),
Service Forestier 4544 (CAY [2 sheets], P).

BRAZIL: Amapá: Massif Tumuc-Humac, Inselberg Talouakem, 2°29′N, 54°45′W, 650 m, 11 Aug. 1993 (fr), J.J. de Granville et al. 12244 (BBS, CAY, MO, P, US); Mun. Oiapoque, Aldeia do Manga [ca. 3°43′N, 51°43′W], tribo Karipuna, 20 Aug. 1981 (fl, imm fr), B. Rabelo 1303 (NY). Amazonas: Rio Cuieras, at junction with Rio Negro [ca. 2°47′S, 60°27′W], campina on white sand, 5 Mar. 1976 (fl), A.B. Anderson 237 (INPA), 238 (INPA); Manaus, Taracuá Campo, 10 Dec. 1962 (fl), A.G.
Andrade & M. Emmerich 1455 (R [3 sheets]), 1456 (R); without locality, 2°55'46"S, 59°58'24"W, 138 m, 28 Aug. 2010 (fl), A. Antonelli et al. 431 (INPA); Manaus, rod. AM-010, Manaus-Itacoatiara, km 26, Reserva Florestal Adolfo Ducke [ca. 3°0'S, 59°56'W], prox. ao alojamento, 11 Feb. 2000 (fl-fr), E.A. Anunciação & E.C. Pereira 810 (INPA, UB); Itacoatiara, rod. AM-010, Manaus-Itacoatiara, km 227, MIL-Madeira, beira da estrada em direção ao Mun. de Silves, km 22, 7 Mar. 2000 (fl), E.A. Anunciação et al. 833 (INPA); Ilha do Piranadé, Rio Negro above Barcelos, 9 Feb. 1944 (fl), J.T. Baldwin Jr. 3307 (F, IAN, NY, US); Rio Negro, Barcelos [ca. 0°58'S, 62°55'W], capoeira, 23 Apr. 1948 (fl, fr), G.A. Black 48-2430 (IAN, U [2 sheets]); Rio Içana, Tunuí, floresta de caatinga, 10 May 1948 (fl, fr), G.A. Black 48-2595 (IAN, P), Rio Içana, Camrão, capoeira, 7 May 1948 (fl, fr), G.A. Black 48-2627 (IAN); 72 km NNE of Manaus, Distrito Agropecuário, Fazenda Dimona of the WWF/INPA MCS Project, 2°19'S, 60°5'W, 50-125 m, mata de terra firme, 26 Aug. 1991 (fl, fr), B.M. Boom & M. Pacheco 8486 (INPA, NY); Estrada Manaus-Itacoatiara, km 64, picada do INPA, 800 da rodovia, lagoa ao lado da picada, solo úmidoadrenoso, 2 Feb. 1969 (fl, fr), *Byron & D. Coelho* 69-126 (INPA, US); 2 km S of Maués [ca. 3°23'S, 57°43'W], near Maués River, deserted farm and nearby forest, disturbed open area on terra firme, 26 Apr. 1974 (fl), D.G. Campbell et al. P22132 [Prance collection number] (INPA, MO, NY, R, S, U, US); Maués airport road, between Maués and airport, 27 Apr. 1974 (fl), D.G. Campbell et al. P22173 [Prance collection number] (INPA, NY, US); Reserva Florestal Adolpho Ducke, Manaus-Itacoatiara, km 26, 2°53'S, 59°58'W, 9 Dec. 1994 (fl), M.T.V. Campos et al. 580 (INPA, NY); Estrada do Aleixo, Lajes, solo de terra firme, 19 May 1975 (fl), L. Carreira 64 (INPA); Mun. Presidente Figueiredo, Balbina, Estrada de acesso ao antigo porto, vegetação aberta, degradada, 1°0'N, 59°0'W, 1 Aug. 2006 (fl), J.G. de Carvalho-Sobrinho & K.M. Silva 824 (INPA), 826 (INPA); Manaus, antiga estrada de de São Raimundo, terra úmida, 7 Dec. 1954 (fl, imm fr), J. Chagas s.n. (INPA 330) (INPA), (MG No. 254545) (MG), s.n. (MG 21681) (MG); Manaus, margem do Igarapé do Parque 10, 6 Dec. 1955 (fl), J. Chagas s.n. (INPA 3049) (IAN, INPA, MG), 23 Jul. 1956 (fl), J. Chagas s.n. (INPA 3971) (INPA, MG); Manaus, Cachoeira Grande dos Bilhares, 16 Mar. 1959 (fl), J. Chagas s.n. (INPA 7220) (INPA), (MG 25061) (MG), s.n. (MG 25134) (MG), s.n. (MG 25068) (MG), s.n. (INPA 330) (IAN 109888) (IAN), s.n. (INPA 7220) (RB), 11 Apr. 1958 (fl), s.n. (INPA 6342) (IAN, INPA, RB); Reserva Florestal Adolpho Ducke, Manaus-Itacoatiara, km 26, 2°53'S, 59°58'W, 9 Dec. 1994 (fl), M.A. Costa & J.R. Nascimento 43 (INPA, MG, NY); Manaus, RDS do Tupé, campinara-

na, prox à comunidade Agrovila, trilha que segue a 4 km a dentro da mata de terra firme, 2°98'22"S, 60°21'8"W, 35 m, 31 May 2015 (fl), L.O. Demarchi, J. Alves & M.C. Melo 409 (INPA); Parintins, Tribo Sateré Maué, mata secundária, Araticu, 8 May 2005 (fl), P.A. Ferreira 26 (INPA); Região de Parintins, Lago do Jurutí, 18 Jan. 1957 (fl, fr), R.L. Frões 33079 (IAN, US); Parintins, pastagem de terra firme, 2°53'14"S, 56°40'40"W, 26 Jan. 2009 (fl), A.K.L. Galvão et al. 35 (INPA); Acampamento da D.B.F.F./W.W.F., km 41, Res. 1501, 25 Jul. 1985 (fl, fr), M.V.B. Garcia 008 (R); Igarapé da Bolivia, woods along stream near turnoff of Caracaraí Road from Itacoatiara Road, ca. 8 km from Manaus, 14 Dec. 1974 (fl), A. Gentry & J. Ramos 13298 (INPA, MO); Santarém, Dto. Alter do Chão, Lago Verde dos Muiraquitãs, Igarapé do Caranazal, margem do igapó, 2°31'17"S, 54°55'54"W, 2 Oct. 2015 (fl), L.L. Giacomin et al. 2680 (RB); Manaus, Reserva Florestal A. Ducke, Estrada Manaus-Itacoatiara, km 26, 2°53'S, 59°58'W, 4 Dec. 2001 (fl), M. Groppo Jr. et al. 916 (INPA); Manaus, margem do igarapé da Normandia, 28 Oct. 1954 (fl), [initials unknown] Jaccoud s.n. (INPA No. 285) (IAN, MG); Cachoeira do Tarumã Mirim, 7 Dec. 1974 (fl), W. Junk 175 (INPA); CacauPirêra, margem direita do Rio Negro, campo arenoso na terra firme, com argila branca, explorada para tijolos, 28 Jun. 1986 (fl), M.L. Kawasaki et al. 2 (INPA); Igarapé Tarumã, N of junction of Rio Negro and Igarapé Tarumã, white sand, black water igapó, 3°2'S, 60°8'W, 23 Nov. 1977 (fl), S. Keel & J. Guedes 300 (MG, NY, RB, S, U); Manaus, road Manaus-Itacoatiara, km 13.5-14, along roadside, 22 Feb. 1969 (fl), H. Kennedy 126 (INPA); "Manáos, civ. Amazonas" [Manaus], Oct. 1912 (fl), J.G. Kuhlmann s.n. (RB2842) (S 10-27921); Uucajatuba, Rio Paurary, affluente do Rio Maués, 17 Mar. 1924 (fl, fr), J.G. Kuhlmann 1648 (RB 110698) (RB [3 sheets]); estrada Manaus-Caracaraí, km 585, campina, 29 Jun. 1979 (fl), J. Lima 464 (INPA); Road to Praia Dourada, ca. 10 km NW of Manaus, 30 Aug. 1980 (fl), J. Lowe 3953 (INPA); near the ferry, other side of the Rio Negro from Manaus, roadside herb, 28 Sep. 1980 (fl), J. Lowe 4028 (INPA); Boa Vista road, 55 km N of Manaus, herb in road cutting, 17 Apr. 1981 (fl, fr), J. Lowe 4080 (INPA); Road AM-010, km 246 (25 km WNW of Itacoatiara), 10 Jun. 1981 (fl), J. Lowe 4271 (INPA); Manaus, Fazenda Aurora, 22 Aug. 1922 (fl, fr), P. Luetzelburg 22041 (R), 22073 (R); Tunuy, Rio Içana, 21 Oct. 1927 (fl, fr), P. Luetzelburg 22412 (R); Manaus, Jardim Zoologico, secondary vegetation along creekside, 18 Apr. 1971 (fl), P.J.M. Maas & H. Maas 278 (INPA); Manaus, Arquipélago das Anavilhanas, Foz do Rio Cuieras, solo argiloso-arenoso, 4 Mar. 1976 (fl, fr), M.E. Medri 51 (INPA); Manaus, igarapé da Cachoeira Grande dos Bilhares, 26 Sep. 1955 (fl), F.C.
Mello s.n. (MG 25352) (MG), (INPA 2040) (IAN, INPA); Manaus, campina nos fundos do Conjunto dos Jornalistas, solo arenoso, 5 Jun. 1984 (fl), F.E. Miranda & M.C.C. Miranda 817 (INPA); estrada Manaus-Itacoatiara, Reserva Florestal Ducke, próximo ao campo de futebol, 12 Mar. 1977 (fl), O.P. Monteiro & R. Lisboa 1339 (INPA); Mun. Prainha, sub-base do Projeto RADAM/BRASIL, capoeira baixa de terra firme, 29 Sep. 1976 (fl, fr), C.D. Mota s.n. (INPA); Cucuí, 26 Apr. 1975 (fl, fr), O.C. Nascimento et al. 198 (IAN, RB, US); Rio Negro, L margin, 50 km acima de Manaus, Campo Amélia, Faz. Belo Horizonte, campo natural de terra firme, dominado por Antonia ovate, *Bulbostylis* spp. e *Curatella americana*, solo argiloso, 18-19 Aug. 1985 (fl), B. Nelson et al. 1414 (NY, UFMT, US); Manaus, área da Fenologia de EEST (Estação Experimental de Silvicultura Tropical), BR-174, 45 km de Manaus, 10 Dec. 1996 (fl, fr), A. Nitta & A. Yoshida 17565 (INPA); 90 km NNE of Manaus, Distrito Agropecuário, Reserva 1501 (km 41), 2°24'26"-2°25'31"S, 59°43'40"-59°45'50"W, 50-125 m, mata de terra firme, 26 Aug. 1991 (fl, fr), A.A. Oliveira et al. 176 (INPA, NY); Manaus, campina atrás do Conjunto dos Jornalistas, solo arenoso, 5 Jun. 1984 (fl), N. de Paula et al. 16 (INPA); Manaus, Reserva Florestal Adolfo Ducke, floresta de terra firme, 2°33'35"S, 59°35'7"W, 17 Sep. 2013 (fl), M.C. Pessoa et al. 855 (INPA); Manaus, Reserva Florestal Adolpho Ducke, caminho para Harpia, terra firme, clareira, solo areno-aregiloso, 21 Oct. 2015 (fl), R.B. Pinto et al. 528 (INPA); Rio Içana, Tunuí, pé da serra, terreno pedregoso, 24 Oct. 1947 (fl, fr), J.M. Pires 756 (IAN); Parintins, 11 Mar. 1946 (fl, fr), J.M. Pires & G.A. Black 1143 (IAN); Manaus-Itacoatiara Highway, km 64, Reserva Florestal Walter Egler, beside small lake, trailing herb, corolla pale purple, 13 Dec. 1966 (fl), G.T. Prance et al. 3623 (INPA, NY, S); vicinity of Manaus, Tarumã Grande, 1 km N from the junction of Rio Negro and Manaus, Rio Negro, Baia do Bueussu, 15 km above Manaus, 18 Mar. 1969 (fl), G.T. Prance et al. 10436 (F, INPA, MG, NY, S, US); Campo Amélia, Faz. Belo Horizonte, entre Igapó Acajatuba e margem direita do Rio Negro, extenso campo graminoso de terra firme, solo argiloso, 21 Apr. 1986 (fl), G.T. Prance et al. 30048 (NY, US); Reserva Florestal Adolpho Ducke, Manaus-Itacoatiara, km 26, ramal para a torre, terra firme, 28 Apr. 1988 (fl, fr), J.F. Ramos 1890 (IAN, INPA, NY); Manaus, Bilhares-Chapada, campo aberto, solo arenoso, úmido, 30 Dec. 1958 (fl, fr), W. Rodrigues 770 (IAN, INPA, MG); Manaus, Igarapé Passarinho, terra firme, solo arenoso, descampado, 21 Nov. 1961 (fl, fr), W. Rodrigues & J. Chagas 3607 (INPA, US); Manaus, Cachoeira Alta do Tarumã, Igarapé do Passarinho, terra firme, solo arenoso, descampado, 30 Nov. 1961 (fl), W. Rodrigues & J.

Chagas 3640 (INPA); Rio Tiririca, afluente do Rio Preto, terreno úmido, sujeito a inundação temporária, 1 Jun. 1964 (fl), W. Rodrigues & D. Coelho 5866 (INPA); Parintins, Comunidade Vinte Quilo da Tribo Mawé, 24 Nov. 2009 (fl, fr), O.A. Santos 221 (INPA); Région de Manaus, clariére herbeuse sur sable claire, Oct. 1958 (fl), R. Schnell 9223 (P); Manaus, Cachoeira Grande, 30 Mar. 1982 (fl, fr), C.A.W. Schwacke 181 (coll. III) (R); Mun. Presidente Figueiredo, km 189 da BR-174, 4 Oct. 1998 (fl), J.A. Silva et al. 857 (INPA); Rio Xié, prox. a Cachoeira de Cumati [ca. 1°18'N, 67°15'W], capoeira de solo argiloso, 9 May 1973 (fl, fr), M.F. Silva et al. 1404 (INPA, US); Estrada Manaus-Caracarai, trecho perdido, 11 Mar. 1978 (fl), N.T. Silva 4583 (MG, NY); Manaus, atrás do Conjunto dos Jornalistas, campina perturbada, solo arenoso, 5 Jun. 1984 (fl), H.C. de Souza 19 (INPA); "Prov. Rio Negro, prope Barra [now city of Manaus]", Dec. 1850-Mar. 1851 (fl), R. Spruce 994 (NY), s.n. (NY 02693444, BM 000614335, FI-Webb 080079, P [2 sheets, P00748125, P00748124], RB 17399 [2 sheets], C); "Barra" [Manaus], Dec. 1850 (fl), R. Spruce 1132 (P); prope Panuré ad Rio Uaupés, Oct. 1852-Jan. 1853 (fl), R. Spruce 2908 (P); Mun. Nova Olinda [Nova Olinda do Norte, ca. 3°53'S, 59°5'W], Lower Rio Mari Mari, below Rio Pará, Laranjal, along river and igarapé, sandy soil, 1 Jul. 1983 (fl), C. Todzia et al. 2286 (INPA, MG, NY, RB, US); SWW of Manaus, en route from Cacau Pirera to Manacapuru, along rd. AM-070, sunny place, roadside, 50-60 m, 6 Aug. 1987 (fl), S. Tsugaru B-757 (NY); Manaus, Jan. 1901 (fl), E. Ule 5374 (MG). Maranhão: Turiaçu [ca. 1°39'S, 45°22'W], km 6 da BR-106 Maracaçumé-Sta. Helena, Fazenda Maracaçumé Agro Industrial Grupo Mesbla, solo arenoso, 1 Dec. 1978 (fl, fr), N.A. Rosa & H. Vilar 2780 (MG, NY). Pará: Rio Parú, Parú do Oeste, Missão Tiriyo, arredores da Missão, 2°20'N, 55°45'W, 23 Feb. 1970 (fl), P. Cavalcante 2491 (MG, NY, US); Rio Tiriós, parte norte occidental do E Pará, 12 May 1962 (fl, fr), E. Oliveira 1843 (IAN, UB); Rio Arapuns, Campo da Raposa, 8 Jun. 1952 (fl), J.M. Pires & N.T. Silva 4226 (IAN); Mun. Juruti [ca. 2°9'S, 56°05'W], erva comum a margem do rio em locais úmidos, 3 Jun. 2014 (fl, fr), R.P. Salomão et al. 1384 (MG); Sete Varas airstrip on Rio Curua [ca. 5°20'S, 54°30'W], open savanna bordered by medium-sized forest and low scrub, 8 Aug. 1981 (fl, fr), J.J. Strudwick et al. 4348 (MG). Roraima: Boa Vista [ca. 2°49'N, 60°40'W], 8 Jul. 1955 (fl), M. Alvarenga s.n. (RB 90577) (RB); Caracaraí, Parque Nacional do Viruá, na margem da Estrada Perdida, 1°25'35"N, 60°59'0"W, 62 m, área alagada, solo argiloso, 17 Jul. 2010 (fl), T.D.M. Barbosa & S.M. Costa 1138 (INPA); Caracaraí, Parque Nacional do Viruá, na margem da Estrada Perdida, 1°26'4"N, 60°58'51"W, 63 m, área alagada, solo
argiloso, 19 Jul. 2010 (fl), *T.D.M. Barbosa & S.M. Costa 1178* (INPA); Mun. Boa Vista, Igarapé Aguas Boas, 20 Aug. 1951 (fl, fr), *G.A. Black 51-12816* (IAN, U); entre Igarapé Azul e Branco e a Estrada Bõa Vista-Caracaraí, 17 Sep. 1951 (fl, fr), *G.A. Black 51-13562* (IAN); Ilha de Maracá, erva lenhosa, flores róseas, 19-26 Jun. 1997 (fl, fr), *L. Carreira et al. 1224* (MG); NE Roraima, ca. 110 km NE of Boa Vista, 8 km SW of Bonfim, Fazenda Valparaiso, 133 m, savannas (campo) with moist sandy soil, 20 Oct. 1977 (fl), *L. Coradin & M.R. dos R. Cordeiro 820* (IAN, INPA, NY); along Boa Vista-Mucajaí road (BR-174), km 2, 3 Nov. 1977 (fl), *Coradin, L. & M. dos R. Cordeiro 965* (INPA); Along Mucajaí-Caracaraí road (BR-174), km 17, dry, sandy soil, roadside, 8 Nov. 1977 (fr), *L. Coradin & M.R. dos R. Cordeiro 1012* (INPA); Caracaraí, Parque Nacional do Viruá, Estrada Perdida, 1°48'67"N, 61°1'57"W, Jun. 2006 (fl), *F.R.C. Costa 1255* (INPA); Caracaraí, Parque Nacional do Viruá, campina, às margens de rio de água preta, 1°3'0"N, 61°14'59"W, 23 Mar. 2011 (fl), *S.M. Costa 941* (INPA); Caracaraí, Parque Nacional Viruá, campinarana (emergentes até 15 m), solo arenoso, inundado periodicamente, Lat. 1.48219, Long. -61.0207, 64 m, 30 May 2010 (fl, fr), *N.C. Dávila & F.N. Cabral 6178* (INPA), 6179 (INPA); Caracaraí, Parque Nacional Viruá, vegetação secundária, Lat. 1.816, Long. -61.128, 2 Jun. 2010 (fl), *N.C. Dávila et al. 6211* (INPA); Caracaraí, Parque Nacional do Viruá, Estrada Perdida, área aberta, alagada, com solo arenoso, 22 Jul. 2010 (fl), *G.C. Delgado Jr. et al. 106* (INPA); Caracaraí, Parque Nacional do Viruá, Rio Anauá, localidade Campinho, campina aberta, solo hidromorfico, 12 Mar. 2007 (fl), *C.A.C. Ferreira et al. 12979* (INPA); Caracaraí, Estação Ecológica de Niquiá, floresta de terra firme, 0°48'5"N, 61°39'37"W, 50 m, 25 Mar. 2012 (fl), *R.C. Forzza et al. 7007* (RB); SEMA Estação, open grassy savanna close to estação, savanna with Curatella americana, 3°22'N, 61°20'W, 7 Jul. 1986 (fl), *M.J.G. Hopkins 545* (MG); Rorainopolis [ca. 0°56'N, 60°25'W], Rio Xipariná, área alterada no igapó, sazonalmente inundada, 6 Mar. 2010 (fl), *M.J.G. Hopkins & E.L. da Encarnação 1978* (INPA); Ilha de Maracá, Mun. Alto Alegre, near Estação Ecológica de Maracá, 3°22'N, 61°20'W, 12 Jul. 1986 (fl), *A. Henderson & R. Pereira de Lima 545* (NY, US); Caracaraí, Parque Nacional Viruá, Estrada perdida, near BR-174, along stagnant water channel dominated by Maurititia flexuosa, 5 Apr. 2013 (fl), *J.E. Householder 2576* (INPA); Margem do Igarapé da Normandia, 28 Oct. 1954 (fl), [initials unknown] *Jaccoud s.n.* (INPA No. 285) (INPA, MG, RB); Amajari, Serra do Tepequém, submontana, savanna, 3°45'45"N, 61°42'57"W, 723 m, 3 Dec. 2013 (fl, fr), *D.L. Komura et al. 1769* (INPA), 6 Dec. 2013 (fl, fr), 1778 (INPA); Manaus, Oct. 1912 (fl), *J.G. Kuhlmann 37*

(RB); Boa Vista, Jul. 1913 (fl, fr), *J.G. Kuhlmann 814* (RB); Estrada Caracaraí, Boa Vista, km 59, beira da Estrada, solo argiloso-arenoso, 28 Jun. 1979 (fl, fr), *J. Lima 450* (INPA); Mun. Caracaraí, Jualpiri, várzea, 30 Jun. 1979 (fl, fr), *J. Lima 467* (INPA); Mun. Boa Vista, 5 km N do alojamento, campo de terra firme, graminoso, com *Curatella e Byrsonima*, solo areia quartzosa, 16 May 1987 (fl), *J. Lima & B. Nelson 734* (INPA, NY, U); Serra do Xiriry, campos serranos, Aug. 1927 (fl), *P. Luetzelburg 20708* (R); Serra da Moça, campos humidos, Aug. 1927 (fl), *P. Luetzelburg 20811* (R), 20816 (R); Serra do Murupú, campos humidos, Aug. 1927 (fl), *P. Luetzelburg 20821* (R); Serra do Camelo, campos humidos, Aug. 1927 (fl, fr), *P. Luetzelburg 20868* (R); Valle do Rio Quinó, Igarapé do Ignacio, campos humidos, Sep. 1927 (fl), *P. Luetzelburg 21502* (R); Igarapé Ouato, campos humidos, Sep. 1927 (fl), *P. Luetzelburg 21588* (R); Bõa Vista, Apr. 1952 (fl, fr), *D. Magalhães 1* [= G.A. *Black 51-12816*] (IAN); Serra Tepequem, occasional in savanna below airstrip, 700-800 m, 2 Dec. 1954 (fl), *B. Maguire & C.K. Maguire 40147* (NY); Caracaraí, Rio Catrimani, 4 km acima da boca do Rio Branco, 0°28'39"N, 61°45'0"W, 45 m, 26 Mar. 2012 (fl, fr), *G. Martinelli et al. 17518* (RB); Caracaraí, Parque Nacional do Viruá, Estrada Perdida, após a primeira bueira, campinarana alagada, 22 Jul. 2010 (fl), *A. Melo et al. 399* (INPA); Caracaraí, Parque Nacional do Viruá, margem do Rio Iruá, 23 Aug. 2012 (fl), *A. Melo et al. 1016* (INPA); Rod. RR-205, km 67, savan arbórea aberta, 23 Jun. 1994 (fl), *I.S. Miranda 187* (INPA); Boa Vista, Lago Redondo, campo aberto, mal drenado, solo arenoso, 29 Jun. 1994 (fl), *I.S. Miranda 242* (INPA), 257 (INPA [2 sheets]); Estrada Velha para a Vila do Cantá, savanna arbustiva, solo arenoso, 27 May 1995 (fl), *I.S. Miranda 715* (INPA); Lago Redondo, à esquerda do km 1 da estrada para Vila São Francisco, campo úmido, 6 Oct. 1995 (fl), *I.S. Miranda 881* (INPA); Mutum, Igarapé do Rebenque, subindo a serra, savanna, 13 Oct. 1995 (fl, imm fr), *I.S. Miranda 1122* (INPA); Alto Alegre, Estação Ecológica de Maracá, Grade do PPBio, trilha No. 1, ilha de savanna arborizada, 3°22'45"N, 61°26'32"W, 128 m, 18 May 2012 (fl), *P.O. Perdiz et al. 1354* (INPA, RB); Caracaraí, Parque Nacional do Viruá, Estrada de acesso à sede do Parque, sentido sede-guarita, floresta ombrófila densa, 1°29'24"N, 61°0'10"W, 10 Aug. 2013 (fl, imm fr), *P.O. Perdiz et al. 1931* (INPA); Rorainopolis, Rio Xipariná, Comunidade Xixuaú, 0°38'56"S, 61°27'10"W, 7 Sep. 2013 (fl), *M.C. Pessoa & E.A. Santos 826* (INPA); Caracaraí, Parque Nacional do Viruá, floresta de terra firme, 20 Sep. 2013 (fl), *M.C. Pessoa & J. Maciel 858* (INPA); Caracaraí, campo, *J.M. Pires et al. 14330* (IAN, M); rd. Rio Uraricoera-Tepequem, grassy campo, 9 Feb. 1967 (fl, fr), *G.T. Prance et al. 4245* (F,
INPA, MG, NY, P, R, S); foothills of Serra da Lua, 2°25'–29'N, 60°11'–14'W, 10 Jan. 1969 (fl), *G.T. Prance et al.* 9183 (F, INPA, MG, NY, S); Reserva Florestal Ducke, Manaus-Itacotiara, km 26, 2°53'S, 59°58'W, ramal para a torre, 28 Apr. 1988 (fl, fr), *J.F. Ramos* 1890 (RB); Ilha de Maracá, SEMA Ecological Reserve, Santa Rosa Area, 3°22'N, 61°25'W, damp campo below the station, 23 Feb. 1987 (fl, fr), *J.A. Ratter et al.* R-5393 (INPA, NY, U); Ilha de Maracá, SEMA Ecological Reserve, Santa Rosa Area, ca. 4 km NE of Station buildings, 3°22'N, 61°25'W, scrubby transition zone between campina forest ans seasonally inundated campo, 1 Feb. 1988 (fr), *J.A. Ratter et al.* R-6189 (NY); Estrada Boa Vista-Venezuela, perto do Igarapé Jacitara, campo alagado com buritizal, 1 May 1979 (fl), *I.A. Rodrigues et al.* 803 (IAN); Normandia, 6 Nov. 1954 (fl), *W. Rodrigues* 11 (IAN, INPA, MG, RB); margem do Igarapé Canan, 22 Nov. 1954 (fl), *W. Rodrigues* 145 (INPA); Mun. Boa Vista, Estação Ecológica de Maracá, campo, 19 Nov. 1978 (fl), *J.L. dos Santos* 260 (INPA [2 sheets]); Boa Vista, Estrada do Cantá, 2°50'N, 60°40'W, beira da Estrada, solo argiloso, 31 Jul. 1986 (fr), *J.A. Silva et al.* 550 (INPA, MG, UB); Boa Vista, Estrada da Serra Grande, 2°50'N, 60°40'W, beira da Estrada, solo argiloso, 2 Aug. 1986 (fr), *J.A. Silva et al.* 585 (INPA, MG, UB); Aldeia da Raposa, 1 Apr. 1964 (fl, fr), *M. Silva* 262 (MG); **State Unknown**: Without locality, s.d. (fl, fr), *Collector Unknown* s.n. (BR 824757) (BR).

**Specimens with erroneous locality:** **BRAZIL:** "Rio de Janeiro: São João da Barra, dans les bois", 10 February 1876 (fl, young fr), *A. Glaziou* 9887 (C [2 sheets, without barcode], P [P00748138], R [Acc. No. 10469]).

The specimens *Glaziou* 9887 and *Glaziou* 9887a, belonging to two different species, have labels reporting a collection locality in the state of Rio de Janeiro, with different collection dates and different ecologies. The labels of both gatherings indicate that the specimens were collected in São João da Barra [ca. 21°38'24'S, 41°3'3"W], which is a coastal town in the northern portion of the State of Rio de Janeiro, mostly with restinga vegetation and patches of "floresta de tabuleiro". The gathering *Glaziou* 9887 is *Sipanea pratensis* var. *dichotoma*. The label of the specimen *Glaziou* 9887 at R [Acc. No. 10469, barcode R010053882] has the heading "PLANTAS DO BRASIL CENTRAL, GLAZIOU N°" and the handwritten text "9987. Sipanea pratensis var. trichantha, K. Sch. Flora Bras. VI. VI, 255. São João da Barra (Rio de Jan. [Rio de Janeiro]), 10 de Fevereiro de 1876. Herbacea flores rosas." The specimen of *Glaziou* 9887 at P, barcode P00748138, has two labels. At the lower left corner is a label handwritten by Glaziou in pencil reporting "São João da Barra [...] 10 fevrier 1876. Herb. fl. rose."

The other label, at the lower right corner, has the printed heading "HERB. MUS. PARIS" and the text, handwritten in black ink by Glaziou "Sipanea pratensis Aubl. var. tricantha. BRÉSIL (Rio Janeiro) São João da Barra, dans les bois, 10 fevrier 1876, Herbacé - fl. rose, n. 9887" and the lower heading "Herbier de A. Glaziou, donné par Mme SIMART, sa fille, en 1907." There are two specimens of *Glaziou* 9887 at C, without barcode, and without indication of locality or collection date. Each specimen has a label with the printed heading "Herbarium Eug. Warming" and the handwritten text "Sipanea pratensis Aubl. [Legit] Glaziou 9887, [determ.] Schumann." The specimens of *Glaziou* 9987 at C, P, and R, correspond with *S. pratensis* var. *dichotoma*; however, Schumann (1889) in *Flora Brasiliensis* did not cite the gathering *Glaziou* 9887 under *S. pratensis* or any other species of *Sipanea*. This variety ranges throughout the Guiana Shield and the Amazon Basin, in Colombia, Venezuela, the Guianas, and northern Brazil (Roraima, Amapá, Pará, and Maranhão). The southernmost collection of this variety is reported from Nova Olinda do Norte [ca. 3°53'S, 59°5'W, Todzia et al. 2286], in the Brazilian state of Amazonas, which is more than 2000 km air distance from São João da Barra [ca. 21°38'S, 41°3'W], state of Rio de Janeiro, which is also a completely different habitat, with remnants of "floresta de tabuleiro" and "restinga" vegetation. Glaziou collected in the states of Rio de Janeiro, São Paulo, Espirito Santo, Minas Gerais and Goiás (Lanjouw & Stafleu 1957; Urban, 1906), and never set foot in the Amazon Basin. It is well-known among botanists (Santos, 2016; Sleumer, 1954; Smith, 1966; Wurdack, 1970) that for numerous specimens Glaziou reported localities where he never was, and included Amazonian collections in his herbarium citing them as collected by him in Rio de Janeiro. Therefore, the specimen labels of *Glaziou* 9987 are treated as reporting an erroneous collection locality.

8-6. *Sipanea veris* S. Moore, Trans. Linn. Soc. 2, 4: 368. 1895. (Figures 51-53).

Type: BRAZIL. Mato Grosso: Rio dos Bugres, near the confluence with the Paraguay River, "in apertis arenosis ad ripas fl. dos Bugres, itaque ad confluentes fl. Paraguay et dos Bugres prope Santa Cruz," 1891-1892 (fl), S. Moore 435 (Lectotype BM [barcode 000614332], here designated; isolectotypes K [without barcode], NY [barcode 01085914]; photo-BM at NY). [Section *Sipanea*].

(=) *Sipanea acinifolia* R. Spruce ex Sprague, Trans. Proc. Bot. Soc. Edinb. 22: 433. 1905
Type: COLOMBIA. Vichada: Maipures, Río Orinoco [border between Colombia and Venezuela], "Prope Maypures, ad flumen Orenoco", Jun. 1854 (fl), R. *Spruce 3652* (**lectotype** LD [Acc. No. 1219703], **here designated**; isolectotypes K [without barcode], MPU [barcode MPU021332], P [barcode P007294432, as "3652 (299)"] RB [Acc. No. 15356, barcode 00364227]).

(=) *Sipanea spraguei* Wernham, J. Bot. 55: 172. 1917.

Type: VENEZUELA. Bolívar: Río Orinoco, Caicara, near a clump of moriche palms, Nov. 1898 (fl), C.J. *Sprague* 7 (holotype BM [barcode 000614324]; isotype K [without barcode]).

Herbs, prostrate or sometimes decumbent, root-
ing at nodes, commonly forming thick patches; stems
thin, 0.5-0.8 mm thick, terete or slightly quadrangular,
green, sparsely strigulose. *Stipules* adnate to the peti-
oles, sheathing, free portion truncate, shallowly triangu-
lar or shortly suborbicular, 0.5 × 0.5-0.8 mm, truncate
to shallowly obtuse at apex, membranaceous, strigulose
or glabrate outside, glabrous, slightly thickened at mar-
gin inside; with 5-6 colleters inserted along the mar-
gin. *Leaves* very variable in size, often varying in size
on the same individual, petiolate; petioles 0.5-6 mm
long, olive-green when dry, strigulose below, glabrous
or glabrate above, commonly canaliculate, not winged,
sparsely appressed strigose; blades ovate, ovate-lanceo-
late to rhombic-ovate, 0.5-2.6 × 0.2-1.7 cm, subacute,
cuneate to obtuse at base, obtuse to acute at apex, mem-
branaceous to papyraceous, grass-green above and dull
pale green below when fresh, olive-green above and
below when dry, lamina glabrous or sparsely strigulose
above (hairs 0.5 mm long) with midrib and secondary
veins sparsely short strigose above, with midrib and sec-
ondary veins sparsely strigose below, margin glabrous
or appressed-strigulose ciliate; secondary veins 2-3
on each side of midrib, obsolete above, slightly promi-
nulous below. *Inflorescences* terminal or subaxillary,
originally terminal, uniflorous or 2-flowered, sessile to
short-pedunculate, peduncle (when present) to 4.5 mm
long, appressed strigose; bracts and bracteoles lacking.
*Flowers* distylous, 5(6)-merous, sessile to short-pedun-
culate; pedicels (when present) to 1 mm long, sparsely
strigulose. *Hypanthium* obovoid, 1.5-3.5 × 2-2.2 mm,

acute to obtuse at base, round to obtuse at apex, densely
appressed-strigillose. *Calyx* lobes subequal or unequal,
green to reddish-green when fresh, linear-lanceolate,
(2-)3-7.5 × 0.1-0.6 mm, acute to acuminate at apex,
sparsely strigulose outside, especially along the midvein,
margins sparsely appressed ciliolate; with 1-2 colleters
in each calycine sinus, narrowly conical, acute at apex,
0.2-0.4 mm long. *Corolla* hypocrateriform, (14-)16-23.5
mm long, variable in size throughout the geographic
range, sometimes varying in size within the same pop-
ulation or on the same individual, pale to intense pink,
turning paler after anthesis, lobes basally white, non-
glandular, forming a white ring at mouth, with a dense
ring of yellow hairs exserted beyond the mouth; tube
(8-)10-13.5 mm long, cylindrical and gradually expand-
ing towards the mouth, 0.9-1.1 mm at base, 2-3 mm
wide at mouth, glabrous and shortly strigillose in lines
outside, lower half glabrous and puberulous above, and
yellow-villous where anthers are inserted, densely yel-
low-villous at corolla mouth; lobes 5(6), obovate, broadly
obovate to round, 6-10 × 4.5-8 mm, obtuse to suba-
cute at apex, glabrous throughout. *Long-styled flowers*
[Molina & Barkley 161 (corolla tube 10 mm long)]: sta-
mens inserted at 3.7 mm below corolla mouth; filaments
0.3 mm long; anthers linear, 1.7 × 0.2 mm, acute at both
ends; style exserted to 1.5 mm beyond corolla mouth,
glabrous; style branches obovate, 1 mm long, round at
apex, papillose. *Short-styled flowers* [Beck et al. 9898
 corolla tube 13.5 mm long), *Haugh 2622* (corolla tube
  12.5 mm long), *Velez 2713* (young fruits, US); stamens
  inserted at 3-3.5 mm below corolla mouth; filaments
  0.3 mm long; anthers linear, 1.8 × 0.2 mm, acute at both
  ends; style included, 7-7.5 mm long, glabrous; branches
  linear to very narrowly lanceolate, ca. 2 mm long, acute
  at apex. *Infructescences* terminal or subaxillary, original-
  ly terminal, uniflorous or 2-flowered, similar to inflores-
  cence. *Capsules* oblong-elliptic, 7 × 3 mm, round at both
  ends, crustaceous, densely antrorse hispidulous-strigil-
  lose, pale green when young, turning pale brown when
  dry. *Seeds* irregularly polyhedral, ca. 0.5 mm in diam.,
  testa foveolate.

Notes: Moore (1895: 368–369) described *Sipanea veris* and cited his own gathering *Moore 435*, without mentioning the herbarium of deposit. Steyermark (1967: 266–267) cited *Moore 435* as type, without indicating any herbarium. Delprete (2010c: 1150) cited a speci-

**Figure 51.** *Sipanea veris.* A. Habit. B. Node with sheathing stipule and partially exserted colleters, and leaf bases. C. Dissected short-styled flower, with stamens inserted at distal portion of corolla tube, and included style. D. Dissected long-styled flower, with stamens inserted at distal portion of corolla tube, and exserted style. A–B, D: drawn from *Molina & Barkley 18-V-161* (US); C: drawn from *Haught 2622* (US). Illustration by Piero Delprete.
<figure><img src="image_55.png" /><figcaption>Figure 51.</figcaption></figure>
<figure><img src="image_56.png" /><figcaption>**Figure 52.** *Sipanea veris* – Color plate. A. Habitat; vegetation frequently cut down, in front of a house; ground about 5 m above river level at the time of collection; during the rainy season (June–September) the water level is 0.5–1 m above the ground. B. Habit. C. Detail of habit, with plants in anthesis. D–F. Corolla variation in lobe shapes and merosity. D. Corolla with four equal lobes. E. Corolla with five equal lobes. F. Corolla with five unequal lobes (two wider than the others). Photos taken by Piero Delprete on 7 January 2001, Mun. San Fernando de Atabapo, locality Súpiro, at the margin of the Orinoco River, Venezuela, where *Delprete et al.* 7457–7459 were collected.</figcaption></figure>

men at BM as holotype; however, this citation cannot
be treated as an inadvertent lectotypification because, as
being published after 2001, it should have been accom-

panied by “here designated” or a similar expression.
Therefore, the BM specimen, barcode 000614332, is here
designated the lectotype of this name.
Sprague (1905: 433–434) described *S. acinifolia* Spruce ex Sprague as having corolla tubes sparsely pubescent outside and cited two gatherings “Orinoco: Caicara, *Sprague*, No. 383; Maypures, *Spruce*, No. 3562” without mentioning any herbarium. Steyermark noted that both gatherings cited by Sprague have corolla tubes externally glabrous; therefore, to be in agreement with Sprague’s protologue, he designated as type of *S. acinifolia* “Vicinibus Santarem, Prov. Para, Brazil, Nov–Mar. 1849–1850, *R. Spruce* 329 (lectotype)”, which has corolla tube sparsely pubescent outside. As the corolla outer vestiture is here considered a variable character within the species, specimen *Spruce* 3652 at LD [Acc. No. 1219703] is here designated as the lectotype of *S. acinifolia*.

Wernham (1917), in the introduction of the same article where he published the revision of *Sipanea* wrote “The types of all the novelties described are in the National Herbarium [BM].” Therefore the specimen *Sprague* 7 at BM, barcode 000614324, is the holotype of *Sipanea spraguei* Wernham. In the same article he also treated *Sipanea acinifolia* as a synonym of *S. veris*.

Seyermark (1967: 266–267) explained that the difference in size and vestiture of leaves and corollas of *Sipanea veris* and *S. spraguei* are overlapping in several specimens, therefore he treated them as synonymous. A conclusion with which I concur.

Seyermark (1967: 263; 1974: 361) distinguished *Sipanea acinifolia* from *S. veris* by having corolla tube externally sparsely strigillose (vs. externally glabrous throughout in *S. veris*), and leaf blades more or less densely strigose beneath (vs. glabrous or nearly so beneath, only the midrib and lateral nerves strigose in *S. veris*), although admitting that “there is little or no demarcation within these intergradations, and it is believed at this time, until further collections are available, justifiable to encompass the variability of leaf pubescence in *S. acinifolia* as one taxon.”

Delprete and Steyermark (2004e: 828–832) treated *S. veris* and *S. acinifolia* as synonymous, without any comment. After an analysis of numerous specimens from throughout the geographic range, I concluded that the variation in external corolla vestiture is a trivial character that can vary even within the same population, and it is here confirmed that these two names are conspecific. Therefore, Steyermark’s typification of *S. acinifolia* is superseded, and the lectotype of this name is here selected from the two gatherings cited by Sprague. Specimens of *Spruce* 3562 are at K, LD and MPU. The specimen at LD, Acc. No. 1219703, has Spruce’s handwritten label “3652 (299) Sipanea acinifolia sp. n., Prope Maypures, ad flumen Orenoco, coll. R. Spruce, Junio 1854” and is here designated the lectotype of this name.

**Distribution and ecology:** Distributed in the Amazon Basin, in the “Llanos del Orinoco” or Orinoquía” of Colombia and Venezuela, as well as Brazil and Bolivia (Figure 53). Spreading herbs, sometimes forming small patches, growing in seasonally flooded areas, in open savannas, at river margins, or at edges of forests and *Mauritia* palm swamps, in sunny or semi-shady exposures, in sandy soils, or soils with sand and clay intermixed with organic material; at 70–450 m altitude.

**Phenology:** Flowering specimens were collected from January through June, and from October through December. Specimens with flowers and fruits were collected in March, April, July and October.

**Suggested conservation status:** Least Concern (LC). This species is amply distributed in Colombia, Venezuela, the Guianas, and Amazonian Brazil (Roraima, Amapá, Amazonas, Pará, Maranhão). It is most frequent in the various kinds of savannas and forest edges (see Distribution and Ecology, above), and is also a common ruderal species growing at margins of forest trails, in secondary vegetation, and roadside vegetation, often forming large patches. Because of its wide geographic distribution, common occurrence in savannic vegetation and as a ruderal species, this species belongs the Least Concern (LC) category, following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** COLOMBIA: Boyacá: Near mouth of Río Güira (Río Meta), 150 m, wet soil, partly shaded, 22 Feb. 1939 (fl), *O. Haught* 2622 (NY, US); Cañada Castero, 20 km de Santa Elena, piste Mani-Sta. Elena, 25 Feb. 1971 (fl), *C. Sastre* 863 (P); Caño Casimena, 16 km de Santa Elena, 26 Feb. 1971 (fl), *C. Sastre* 883 (P). **Casanare:** Río Meta, Orocué [ca. 4°47'N, 71°20'W], 140 m, 3 Nov. 1938 (fl, fr), *J. Cuatrecasas* 4378 (US). **Guainía:** Ribeiras del Río Inírida, sítio Sabanita, a medio camino entre “Raudal Alto” y “Morichal,” 180–200 m, 7 Feb. 1953 (fl), *A. Fernández* 2184 (US). **Guaviare:** San José del Guaviare [ca. 2°33'N, 72°38'W], Río Guaviare, sabana, 240 m, 13 Nov. 1939 (fl, fr), *J. Cuatrecasas* 7722 (US). **Meta:** Mun. Puerto Gaitán, cerca al Río Manacacias, 4°18'N, 72°5'W, 220 m, en sabana, 9 Dec. 1993 (fl), *R. Cortés et al.* 1103 (P); 2 km E of Puerto Gaitán, in the flood-plain of Río Manacacias, low lying savanna and scattered secondary forest along river bank, 150 m, 31 Dec. 1973 (fl), *G. Davidse & F. Llanos* 5408 (F, MO, NY); margem derecho del Río Manacacias, frente a Puerto Gaitán, fomación Serrania, ca. 200 m, 26 Mar. 1971 (fl), *E.P. Pinto & O.P. Bernal* 1575 (P). **Vaupés:** Rio Vaupés, a Mitu [1°11'N, 70°10'W], 320 m, 7–8 Nov. 1952
(st), *H. Humbert & A. Fernandez s.n.* (P P06772733), (fl),
*s.n.* (P P06772734). **Vichada:** Hacienda de Hector Pérez,
Sacuraré, on the Río Vichada, ca. 40 km W of San José
de Ocuné, 18 Jan. 1944 (fl), *F.J. Hermann* 10964 (F, NY,
US); Parque Nacional El Tuparro, La Linea Roja, just S
of Río Toro, open savanna and wooded edge of small
stream, 5°32'N, 68°32'W, 100 m, 12 Mar. 1985 (fl, fr), *J.L.
Zarucchi & C.E. Barbosa* 3684 (COL, FMB, MO, NY).

**VENEZUELA. Amazonas:** Orinoco River, south-
ern part of Isla del Raton, savannah, grass vegetation,
light shade, 5°2'N, 67°46'W, 90 m, 22 Nov. 1965 (fl), *F.J.
Breteler* 4786 (MER, NY, U, US, WAG); carretera Puer-
to Ayacucho-Sanariapo, 100 m, s.d., *G. Bunting et al.*

3543 (U, VEN); San Fernando de Atabapo [ca. 4°3′N, 67°38′W], 29 Mar. 1950 (fl), *H.M. Curran* 109 (NY); Isla Ratón, area seasonally inundated, dominated by Poaceae, soil of sand intermixed with rich organic material and clay, at moment of collection soil moist but not inundated, 5°4′N, 67°48′W, 90 m, 6 Jan. 2001 (fl), *P.G. Delprete et al.* 7446 (CAY, MO, NY, P, TFAV, US, VEN), 7447 (CAY, F, L, NY, TFAV, VEN), 7448 (B, CAY, K, TFAV, VEN), 7449 (CAY, P, TFAV, US, VEN), 7450 (CAY, VEN); Mun. San Fernando de Atabapo, Community of Minicia Nuevo, casério near river margin, open, sunny exposed area, soil with sand and clay intermixed, 3°56′N, 67°27′W, 130 m, 7 Jan. 2001 (fl), *P.G. Delprete et al.*

<figure><img src="image_57.png" /><figcaption>**Figure 53.** Distribution of *Sipanea veris*.</figcaption></figure>
al. 7454 (CAY, P, TFAV, US, VEN), 7455 (CAY, MO, NY, TFAV, VEN), 7456 (B, K, L, TFAV, VEN); Mun. San Fernando de Atabapo, locality Súpiro, at the margin of the Orinoco River, vegetation frequently cut down on front of a house, soil about 5 m above river level at the time of collection, during the rainy season (June-September) the water level increases to 0.5-1 m above soil level (and the plants are submersed), soil of clay and sand intermixed, 3°56'N, 67°27'W, 120 m, 7 Jan. 2001 (fl), P.G. Delprete et al. 7457 (CAY, MO, TFAV, US, VEN), 7458 (L, NY, P, TFAV, VEN), 7459 (BR, K, NY, TFAV, VEN); [Parque Nacional Cerro Yapacana, ca. 4°0'N, 66°35'W], Cerro Yapacana, upper Río Orinoco, 100 m, Apr. 1931 (fl), E.G. Holt & E.R. Blake 774 (BR, NY); Depto. Atabapo, near Canaripó, L margin (S) of Río Ventuari, ca. 20 km from junction with Río Orinoco, sabana inundada con *Caraipa* y montículos de termiteros hasta 2 m de alto, 4°3'N, 66°49'W, 98 m, 31 May 1978 (fl), O. Huber 1974 (NY, VEN); clearing at Santa Barbara, at junction of Ríos Orinoco and Ventuari, 100 m, 8 Jan. 1951 (fl), B. Maguire et al. 30811 (NY [2 sheets], VEN); Alto Río Orinoco, Santa Barbara, at junction of Ríos Orinoco and Ventuari, 120 m, 16 Mar. 1953 (fl), B. Maguire & J.J. Wurdack 34479 (NY [2 sheets], UB, VEN); orilla del Río Orinoco en los alrededores de San Fernando, 250 m, terrenos arenosos y húmedos, 24 Nov. 1948 (fl), J.A. Molina & F.A. Barkley 18-V-161 (US); gallery forest along Caño Cataniapo, 14 km S of Puerto Ayacucho, 75 m, 4 May 177 (fl, fr), J.A Steyermark & O. Huber 113871 (F, VEN); silvestres en formación, graminiforme e chaparral sobre pequeñas planicies y/o lajas graníticas, en el Tobogancito, alrededores de Puerto Ayacucho, al sur, 16 Apr. 1986 (fl), B. Truillo et al. [additional collectors unknown] 20441 (MG, VEN); along trail between San Fernando de Atabapo and Río Orinoco, 125 m, 29 May 1959 (fl), J.J. Wurdack & L.S. Adderley 42664 (NY [2 sheets], US, VEN). **Apure:** Dtto. Muñoz, Hato El Polvero, 79 km S de Elorza, sabanas de banco adyacentes al Caño Cicature (afluente del Alto Río Cinaruco), 6°38'N, 69°37'W, 80 m, 28 Apr. 1987 (fl), G. Aymard et al. 5666 (MO, NY); Dtto. Pedro Camejo, near Río Meta, at Fundo El Algarrobo, 25 airline km WNW of Buena Vista, Morichal de La Madera, 6°13'N, 68°49'W, 70 m, 16-18 Feb. 1978 (fl), G. Davidse & A. González 14188 (MO, NY, P, VEN); Puerto Paez [ca. 6°12'N, 67°27'W], 8 May 1946 (fl), I. Velez 2713 (US). **Bolívar:** Dtto. Cedeño, km 12-120 de la carretera Caicara del Orinoco-Puerto Ayacucho, s de Caicara del Orinoco, 6°50'N, 66°30'W, 100 m, 18 Nov. 1984 (fl), G. Aymard & B. Stergios 3286 (MO, NY, PORT); Caicara [ca. 7°50'N, 66°10'W], Estero Santa Rita, 60 m, 4 Jun. 1984 (fl), S. Lopez-Palacios et al. 4334 (MERF, NY, VEN); Distrito Cedeño, 14 km SW of Caicara del Ori-

noco, wet savanna, 2 Sep. 1985 (fl), J.A. Steyermark et al. 131116 (MO, VEN); Guayapo [ca. 7°20'N, 65°11'W], Bajo Caura, 100 m, 20 Apr. 1939 (fl, fr), Ll. Williams 11917 (F, IAN, US); sítios bajos y periódicamente anegados, en la sabana de El Tigre, 90 m, 20 Jun. 1940 (fl, fr), Ll. Williams 13411 (US). **Guárico:** Hato Flores Moradas, 60 km S of Calabozo, s.d., L. Aristeguieta & Hernández 2324 (VEN); Dtto. Infante, Parque Nacional Aguaro-Guariquito, Aguas Muertas, orillas del Río San Bartolo, 8°92-96'N, 67°52-60'W, 50 m, Dec. 1981 (fl), F. Delascio et al. 11282 (NY); Carretera Santa Rita-Cabruta, s.d., B. Trujillo 5507 (VEN). **State Unknown:** without locality, 7 Apr. 1921 (fl), M. Grisol s.n. (P P00729413).

**BRAZIL. Amazonas:** Mun. Nova Olinda [Nova Olinda do Norte], lower Rio Mari, below Rio Pará, Laranjal, along river and igarapé, sandy soil, 1 Jul. 1983 (fl, fr), C. Todzia et al. 2288 (INPA, MG, NY). **Mato Grosso:** Parque Nacional do Xingú [now Parque Indígena do Xingú], Rio Tutuarí, margem alagável do rio, Sep. 1965 (fl), D. Coelho s.n. (INPA 15866) (INPA); Cidade de Barra dos Bugres, beira do Rio Paraguai, mata de beira do rio, solo arenoso, 15°5'S, 57°10'W, 160 m, 27 Aug. 1984 (fl), L. Coradin et al. 6996 (CEN); Serra do Roncador, Mun. Barra do Garças, Córrego do Gato, 210 km along new road NNE of village of Xavantina, 50 km due S of Royal Society-Royal Geographic Society Base Camp, base camp at 12°51'S, 51°45'W, 450 m, 9 Sep. 1968 (fl, fr), G. Eiten & L.T. Eiten 8649 (RB, UB), 4 Oct. 1968 (fl, fr), 9074 (NY, SP, US); without locality, 11°27'S, 50°19'W, 25 Nov. 1977 (fl), C.T. Falcão 5096 (RB [2 sheets]); Nova Ubiratã, Esctação Ecológica (ESEC) Rio Ronuro, Cruz de Malta I, margem direita do Rio Ronuro, floresta estacional semicidical, transição Cerrado-Amazônia, mata ciliar, 13°5'32"S, 54°25'19"W, 323 m, Oct. 2018 (fl), D.R. Giacoppini et al. 486 (CNMT); Córrego do Gato, ca. 51 km S of Base Camp (12°49'S, 51°46'W), drying up margin of small lake, 3 Oct. 1968 (fl), R.M. Harley, R. Souza & A. Ferreira 10443 (NY, P, RB, U, UB); 1º de Fevereiro, margem do Rio Cautario, Jan. 1919 (fl), J.G. Kuhlmann 2362 (RB [2 sheets]); Rio Sangrador, in nemore uliginoso, 9 Dec. 1893 (fl), C.A.M. Lindman A2443 (S [2 sheets]); Fazenda Espírito Santo, Sesc Pantanal [ca. 16°30'S, 56°30'W], borda Cerrado, 15 Nov. 2003 (fl), G.A. Lima Jr. 31 (UFMT); 0.5 km E of km 235.5 of Xavantina-Cachimbo road, 18 Dec. 1967 (fl), D. Philcox et al. 3656 (NY, P, UB); Poconé, Fazenda Ipiranga, seco, 25 Jul. 1991 (fl), A.L. Prado 959 (UEC); Poconé, Fazenda Ipiranga, ca. 4 km a esquerda no km 10 da rodovia transpantaneira, borda de campo, sazonalmente alagável, 21 Jul. 1991 (fl), A.L. Prado 5103 (UFMT); Pantanal de Poconé, Fazenda Nossa Senhora Aparecida, campo inundável, solo argilo-arenoso, 15 Nov. 2003 (fl), L. Rebellato & L.
<figure><img src="image_58.png" /><figcaption>**Figure 54.** *Sipanea biflora*. A. Habit. B. Portion of inflorescence. C. Dissected corolla with detail of hairs at mouth and corolla throat. D. Dehisced capsule. E. Seeds. F. Node with bilobed stipule and leaf bases. Illustration by Bruno Manara (Reproduced with permission from Flora de Venezuela vol. 9(1), fig. 60. 1974).</figcaption></figure>
Ferraz 212 (UFMT); Rio Xingú, Oct. 1947 (fl), H. Sick B-416 (RB); ca. 66 km S of Base Camp, along rd, 9 Sep. 1968 (fl), R. Souza in Richards 6878 (NY, P, RB, U); Mun. Luciara, lake 2 km NW of Luciara, cerradão forest and marsh, 11°11'S, 50°44'W, 8 Oct. 1985 (fl), W.W. Thomas et al. 4303 (INPA, MG [2 sheets], NY [2 sheets]), SPF; Jacaré (acampamento), Rio Xingu, 15 Jan. 1948 (fl), M. Victorino 1 (R 3 sheets)); Cotriguaçu [ca. 9°51'S, 58°24'W], Fazenda São Nicolau, 3 Mar. 2011 (fl), M.C. Vilela-Santos & L.F. Barbosa 4832 (IAN); Pantanal de Poconé, Pirizal, Fazenda Aparecida, campo de pastagem, 18 Nov. 2000 (fl), D.A.A. Vilhalva & E.C. Arruda 95 (UFMT). **Mato Grosso do Sul:** Mun. Corumbá, Fazenda Marilândia [ca. 19°S, 57°W], campo arenoso seco, 7 Oct. 1953 (fl), E. Pereira et al. 245 (NY, R, RB); Mun. Corumbá, Fazenda Campo Dora, Vazante, 10 Jun. 1994 (fl), G. Hatschbach et al. 60916 (C, MBM, US). **Pará:** Mun. Santarém, Praia do Lago Verde de Alter do Chão, 2°29'2"S, 54°56'56"W, 3 Apr. 2016 (fl), T. André et al. 82 (RB); Belterra, Praia de Porto Novo, 29 Oct. 1947 (fl), G.A. Black 47-1841 (IAC, IAN, U); Santarém, embodura do Rio Tapajós, lado direito, campo arenoso com vegetação arbustiva, 12 Dec. 1966 (fl), P. Cavalcante & M. Silva 1727 (IAN, MG, RB); Bõa Esperança, abaixo da Cachoeira, capoeira, margem esquerda, 29 Dec. 1951 (fl, fr), J.M. Pires 3644 (IAN); Parque Nacional Tapajós, km 60 da estrada Itaituba-Jacarecanga, margem esquerda do rio, capoeira de várzea, 23 Nov. 1978 (fl, fr), M.G. Silva & C. Rosário 3904 (MG, NY, UB); Santarem, campos de Maracá, soil peaty, Nov. 1849 (fl), R. Spruce 299 (C, K, P, S, MPU); in vicinibus Santarem, Nov. 1849-Mar. 1850 (fl), R. Spruce s.n. (FI-Webb, P); Vitória do Xingu, 3°12'54"S, 52°6'46"W, 20 Feb. 2015 (fl), C.R.D. Vieira, PSACF_EX05366 (RB). **Rondônia:** São Francisco, Porto Murtinho, Reserva Biológica Guaporé [ca. 12°30'S, 62°53'W], Rio São Miguel, permanenetly flooded wetland dominated by Mauritia flexuosa, 150 m, 1 Oct. 2013 (fl), J.E. Householder 3025 (INPA). **Tocantins:** Mun. Santa Isabel, Ilha do Bananal, moist edges of campo, growing in large masses, 25 Jul. 1968 (fl), P.W. Richards 6515 (NY, P, RB, U, UB). **State Unknown:** Without locality, s.d. (fl, fr), L. Riedel 141 (BR).

**BOLIVIA:** **Beni:** Prov. Ballivian, en la zona de influencia del Río Yacuma, Isla de Espiritu, [14°9'S, 64°23'W], 200 m, sabana húmeda, 16 Oct. 1980 (fl), G. Beck 5102 (UB); Prov. Itenez, S side of Río Guaporé, "campo verde" ca. 15 km ESE of Costa Marques [Estado de Rondônia, Brazil], flat grassland with scattered tortuous trees, 12°30'S, 64°8'W, 150 m, 31 Mar. 1987 (fl), M. Nee 34644 (NY, P, SP). **La Paz:** Prov. Iturralde, Luisita, W del Río Benio, pastizal al lado del Río Muqui, sabana húmeda, 13°5'S, 67°15'W, 180 m, 22 Feb. 1984 (fl), G. Beck & R.

Haase 9898 (US). **Santa Cruz:** Prov. Velasco, Reserva Ecológica El Refugio, Campamento La Toledo, a 5 km de la casa, pampa inundada estacionalmente, 14°45'42"S, 61°8'26"W, 220 m, 29 Oct. 1996 (fl), M. Garvizu & A.M. Carrión 234 (MO); Prov. Velasco, Parque Nacional Noel Kempff Mercado, pampa humeda Los Fierros, con termineros atualmente seca sin incendio recente, atualmente inundada, 14°48'41"S, 60°23'45"W, 225 m, 15 Oct. 1993 (fl), T. Killeen et al. 5562 (MO, NY); Prov. Velasco, Reserva Ecológica El Refugio, pampa inundada estacionalmente, 14°45'42"S, 61°8'26"W, 200 m, 19 Oct. 1994 (fl), T. Killeen et al. 7012 (NY).

**8B. Sipanea section Virecta** (Rottb.) Steyerm., Mem. New York Bot. Gard. 17: 263. 1967.
Virecta L.f., Suppl. Pl. 17, 134. 1782 ["1781"]).

Type: *Virecta biflora* (Rottb.) L.f [= *Sipanea biflora* (Rottb.) Cham. & Schltdl.].

*Sipanea* sect. *Panisea* K.Schum., Martius et al., Fl. Bras. 6(6): 249. 1889.

Type: *Sipanea biflora* (Rottb.) Cham. & Schltdl.

*Sipanea* sect. *Sipanea* subsect. *Cryptotricha* Steyerm.,
Mem. New York Bot. Gard. 17: 263, 264. 1967.
Type: *Sipanea biflora* (Rottb.) Cham. & Schltdl.

Herbs, prostrate, rarely decumbent; corolla mouth with
a ring of white hairs.

8-7. **Sipanea biflora** (Rottb.) Cham. & Schltdl., Linnaea 4: 168. 1839. (Figures 5G, 54, 55).

(=) *Rondeletia biflora* Rottb., Descr. Rar. Pl. Surin. 7-8. 1776 [Rottb., Acta Lit. Univ. Hafn. 1: 276, tab. 2, fig. 2. 1778; Ed. 2: 8. 1798]. - *Virecta biflora* (Rottb.) L. f., Suppl. Pl. 134. 1782 ["1781"). - *Rondeletia biflora* A. Rich., Mém. Fam. Rubiacées 191. Dec. 1830 [reimpr. Mém. Soc. Hist. Nat. Paris, ser. 3, 5: 271. 1834], isonym *Lectotype (here designated)*: Drawing commissioned by Christen Friis Rottbøll in the library archives of the Natural History Museum, Copenhagen, the plant and analysis on the right side of the drawing [illustrated in Moraes, Dobreff & Reinhammar, Phytotaxa 165(1): Fig 21. 16 Apr 2014].

(=) *Sipanea radicans* Endl., Atakt. Bot. 7, fig. 7. 1833
Type: BRAZIL. Bahia: Without locality, s.d., *J.S. Blanchet s.n.* (lectotype W [Acc. No. W0072693], here designated).

(=) *Sipanea cowanii* Steyerm., Mem. New York Bot. Gard. 17: 265. 1967.

Type: GUYANA. Kaieteur Plateau, near Johnson's View, vicinity of Kaieteur Falls, and along western rim of Potaro Gorge, 425 m, 18 Feb. 1962, *R.S. Cowan & T.R. Soderstrom 1863* (holotype US [Acc. No. 2438240, barcode 00137724]; isotypes NY [barcode 00133310], VEN [Acc. No. 309636]; photo-US at NY).

(=) *Manettia hydrophila* Dwyer, Ann. Missouri Bot. Gard. 67: 280. 1980.

Type: PANAMA. Veraguas: Mouth of Río Concepción, in swamp, 4 Dec. 1967 (fl, fr), *W.H. Lewis, T.B. Croat & J.L. Hawcker 2822* (holotype MO [Acc. No. 312080]).

*Herbs*, terrestrial on forest floor, prostrate or sometimes decumbent on terrestrial habitats, or floating or semi-submersed in flooded areas, rooting at nodes, commonly forming thick patches, very variable in leaf and corolla size and shape; stems thin, 1.5-2 mm in diam., terete or slightly quadrangular, green, sparsely antrorse-pubescent or sparsely strigulose. *Stipules* adnate to the petioles, broadly triangular, 0.3-1.2(-2) × 0.7-0.9(-1.2) mm, shallowly obtuse to subacute or very rarely bifid at apex, membranaceous, glabrous or sparsely strigillose outside, glabrous, with a line near the base with a row of 2-6 colleters inside; margins hispid. *Leaves* very variable in size, often varying in size on the same individual, petiolate; petioles 4-18 mm long, green when fresh, commonly canaliculate, not winged or sometimes narrowly winged on both sides, sparsely short-strigose to shortly villous; blades ovate, rhombic-ovate or rarely oblong, (0.4-)1.2-5 × (0.35-)0.8-2.8 cm, truncate, round to cuneate, sometimes decurrent, rarely asymmetric, at base, obtuse to acute at apex, membranaceous to papyraceous, grass-green above and dull pale green below when fresh, olive-green above and below when dry, lamina glabrous, sparsely strigulose-puberulous or strigillose above with midrib and secondary veins sparsely short puberulous-strigulose above, glabrous, with midrib and secondary veins sparsely short puberulous-strigulose or strigillose below, margin sparsely appressed pubescent to appressed-strigulose ciliate; secondary veins (3-)4-6 on each side of midrib, ascending, weakly evident above,

slightly prominulous below. *Inflorescences* terminal or subaxillary, originally terminal, cymose, monochasial or rarely dichotomous, 1-3(-7)-flowered, short to long-pedunculate, rachis thin to almost capilliform, peduncle 5-15 mm long, glabrous; bracts subtending terminal cyme lanceolate, 1.5-3 × 0.2-0.4 mm, acute or acuminate at apex, glabrous, sparsely ciliate at margins; bracteoles subtending central flowers subulate, 0.5-1.5 mm long. *Flowers* distylous, (4)5-merous, pedicellate or sesile; pedicels of lateral flowers 5-15 mm long, those of central flower absent (flower sessile) or 1-1.5 mm long, appressed-pubescent and ciliolate at apex. *Hypanthium* obconical, obovoid or subglobose, 1.5-2 × 0.7-1.2 mm, acute obtuse at base, round to obtuse at apex, sparsely appressed-strigillose at distal half (glabrous below), or appressed-strigillose throughout. *Calyx* lobes subequal or unequal, green when fresh, linear-lanceolate to setaceous, 1-5 × 0.2-0.6 mm, acuminate at apex, glabrous or puberulent, with a few sparse hairs or appressed-pubescent, margins with a few antrorse-ascendent or appressed hairs; with 1 colleter in each calycine sinus, conical, obtuse at apex, 0.25-0.3 mm long. *Corolla* hypocrateriform, (8-)15-21(-29) mm long, very variable in size throughout the geographic range, sometimes varying in size within the same population or on the same individual, pink deep pink to pale purple turning paler after anthesis, lobes with a basal narrowly triangular area with white, glandular hairs, forming a white star-shaped area at mouth, with a ring of white hairs at mouth; tube cylindrical, 5-13(-15) mm long, 1-1.1 mm wide throughout, or slightly widening distally, 1-1.2 mm at base and to 1.4-2 mm wide at mouth, glabrous or glabrate at lower half and sparsely short-pubescent outside, lower half glabrous and shortly puberulous above, and white-villous near the anthers, with papillose hairs inside, densely white-villous at corolla mouth; lobes oblong-obovate, obovate to round, 3-8.5(-14) × 2-5.5(-7) mm, obtuse to round at apex, glabrous throughout. *Long-styled flowers* (Henkel et al. 4607, Wessel-Boer 636, Sastre et al. 8150; corolla tubes 7.5-13.5 mm long): stamens inserted at 3.7-5.5(-9) mm below corolla mouth; filaments 0.5-0.7 mm long; anthers narrowly oblong to linear, 1.5-2.5 × 0.15-0.2 mm, acute at both ends; style about the same length as corolla tube (tips of style branches exserted) or exserted to 1.5 mm beyond corolla mouth, glabrous; style branches linear, acute or very narrowly lanceolate and acuminate at apex, 0.8-2 mm long. *Short-styled flowers* [Pipoly & Gharbarran 10147; corolla tubes 12-12.5 mm long]: stamens inserted at 4-4.5 mm below corolla mouth; filaments 0.5-0.7 mm long; anthers linear, 2.2 × 0.2 mm, acute at both ends; style included, 5 mm long, glabrous; branches linear, 1.2
nally terminal and becoming subaxillary by the reitera-
ion of an axillary bud), cymose, commonly with 1-2
pedicellate flowers (or exceptionally with 3(-7) flowers).
In fact, in figure 2 of table 2 of *Rondeletia biflora* pub-
lished by Rottbøll (1778) the plant is depicted with three
2-flowered inflorescences. The identification of speci-
men SBT 4.2.6.60 as *Coccocypselum guianense* was fur-
ther confirmed by Cristina Bestetti Costa, a specialist
that wrote a monograph on *Coccocypselum* (Costa 2004).
Therefore the lectotypification of *Rondeletia biflora* by
Moraes et al. (2010) is erroneous and is here superseded.

According to Stafleu & Cowan (1983: 930–931), Cris-
ten Friis Rottbøll studied botany with Linnaeus in 1756–
1757, was director of the Botanical Garden in Copenha-
gen in 1770–1797, and his “herbarium and types are at
C, with some further material in LINN.” Rottbøll (1776)
first published the description of *Rondeletia biflora* in his
*Descriptionis Rariorum Plantarum*. Moraes et al. (2010:
183) cited that Table 2 Figure 2 was published in Rott-
bøll’s 1776 publication; however, no plate is included in
that publication. The illustration of *Rondeletia biflora*
was first published two years later by Rottbøll (1778) in
Table 2, Figure 2 of *Descriptiones Plantarum quarundam
Surinamensium*, in *Acta Literaria Universitatis Hafnien-
sis*. As figure 2 of table 2 of Rottbøll (1778) was pub-
lished after the original description, it cannot be treated
as original material. Therefore, original material of *Ron-
deletia biflora* is either a specimen collected by Rolander
and examined by Rottbøll, or an illustration that was
available to him by 1776.

Two specimens of *Rondeletia biflora* were located at
C. Both of them were without barcode at the time they
were studied. Both of them have the stamp “HB VAHL-
II” on the upper right-hand corner on the back of the
sheet, confirming that they were in Martin Vahl’s Her-
barium. In Vahl’s vast card catalogue, intended for his
*Enumeratio Plantarum*, he positioned the species in the
genus *Virecta*. On the card dedicated to *Virecta biflora*
he wrote “biflora V. [Virecta] foliis ovatis pedunculis
bifloris – Rondeletia biflora, caulis filiformis repente;
pedunculis bifloris – Rottb. descr. pl. Surinamensium
pag. 7 tab. 2 fig. 2. * Virecta biflora Lin. Suppl. pag. 134
S.V. – Habitat Surinami humidis Ø - [followed by the
plant description].” One C specimen is composed by
two branches with numerous adventitious roots at each
node, and numerous leaves; one branch has at least one
fruit, and the other branch has a flower in anthesis. On
the back of the sheet, on the lower portion, is Vahl’s
handwritten note “Virecta biflora Rottb. Des. pag. 7 no.
6 tab. 2 fig. 2.” This specimen much resembles the plant
depicted in figure 2 of table 2 of Rottbøll’s 1778 publi-
cation. However, on this specimen the collection local-

ity or the collector's name is not indicated, and there is
no evidence that it was studied by Rottbøll. The other C
specimen of *R. biflora* is a very small branch with a few
leaves, one flower, and one young fruit. On the back of
the sheet, on the lower portion, is Vahl's handwritten
note "Virecta biflora Linn. Suppl. - Rondeletia biflora
Rott. Descr. Sur. 7 ns. 6 tab. 2 fig 2." On this specimen
the locality and collector's name is also not supplied,
and there is no evidence indicating that it was studied by
Rottbøll. In conclusion, because it cannot be proven with
certainty that the two C specimens were collected by
Rolander nor that they were examined by Rottbøll, they
cannot be treated with certainty as original material.

In Table 2 Figure 2 of Rottbøll’s 1778 publication is
depicted a ramified branch with numerous adventitious
roots, leaves acute at base, and three terminal 2-flowered
inflorescences, each of them with one terminal flower in
anthesis, and a fruit just below it. In the lower portion of
the figure are depicted flowers in anthesis (side view and
dissected), one unopened fruit, and one open capsule. As
this figure was published two years after Rottbøll’s 1776
publication where he first described *Rondeletia biflora*, it
cannot be treated as original material.

The only entities that can be treated as original
material of *Rondeletia biflora*, available to Rottbøll by
1776 are the right portion of the drawing commissioned
by Rottbøll, and the right portion of the copper plate
engraving corresponding to Table 2 Figure 2 of Rott-
bøll’s 1778 publication. The original copper plate engrav-
ing has been lost (Henning Knudsen, pers. comm.).
Therefore the only original material left is the drawing
commissioned by Rottbøll. The drawing commissioned
by Rottbøll was reproduced in Figure 21 of Moraes et
al. (2014) and is an ink drawing of two different spe-
cies, and is similar to Table 2 Figure 2 of Rottbøll’s 1778
publication. On the left side of the drawing is *Eriocau-
lon fasciculatum* Rottb. and on the upper margin is
depicted a series flower and fruits of that species. On the
right side of the drawing is depicted a plant of *Rondele-
tia biflora* with several branches and adventitious roots.
On the lower right portion of the drawing is glued a
rectangular piece of paper with the same series of parts
depicted in Table 2 Figure 2 of Rottbøll’s 1778, which
is to say flowers in anthesis (side view and dissected),
one unopened fruit, and one open capsule. Aside from
the small piece of paper glued on the sheet, it is possible
to see that Figure 21 of Moraes et al. (2014) is a photo-
graph of the original drawing by the calligraphy of the
letters present on the drawing, which differs from that
of the letters on Table 2 Figure 2 of Rottbøll’s 1778 pub-
lication. Rottbøll’s commissioned drawing (reproduced
in Figure 21 of Moraes et al. (2014)) was present in the
<figure><img src="image_59.png" /><figcaption>**Figure 55.** *Sipanea biflora*. A. Habit. B. Detail of habit with flowers in anthesis. C. Detail of habit with flower buds. D. Habit, showing variation in leaf size within the same population. Photos by Piero Delprete in the Kaieteur National Park on 22 June 2016, Guyana, plants not collected.</figcaption></figure>
archives of the library of the Natural History Museum of Copenhagen. That Museum is currently under restruc-
turation and reorganization. The librarian responsible for the archives at the time that the drawing commis-
sioned by Rottbøll was photographed has retired, and a new librarian has not yet been appointed. In addition, as the museum is being renovated, the library and archives are currently closed to the public. Henning Knudsen, the person temporarily responsible of the library and the archives of that institution, informed me that Rottbøll's original copper plates were historically destroyed and melted to produce new copper plates. Therefore, the por-
tion of the drawing commissioned by Rottbøll (repro-
duced in Figure 21 of Moraes et al. (2014)), representing the plant on the right side of the drawing and the small piece of paper with the flower and fruit series is here designated as the lectotype of *Rondeletia biflora*.

Endlicher (1833: 7, fig. 7) described *Sipanea radicans*
Endl. and cited a gathering collected by Blanchet in the
state of Bahia, Brazil, without indicating the herbar-
ium of deposit. A specimen at W, Acc. No. W0072693,
has two handwritten labels “Hb. Endl., Sipanea radi-
cans Endl., Bahia, Blanchet” and “Sipanea biflora L. fil.,
determ. C. Schumann.” This specimen is here designated
the lectotype of this name.

Steyermark (1967) separated *S. cowanii* from *S. biflora* by the hypanthium densely strigose (vs. glabrous except for sparse hairs on upper half in *S. biflora*), upper leaf moderately appressed-pubescent (vs. upper surface glabrous), and capsules moderately to inconspicuously strigillose (vs. glabrous except for a few inconspicuous short hairs). A detailed study of specimens of *S. biflora* from throughout the geographic range, and personal field observations in Guyana, French Guiana, and Brazil,

<figure><img src="image_60.png" /><figcaption>**Figure 56.** Distribution of *Sipanea biflora*.</figcaption></figure>
revealed that the vestiture of this polymorphic species is highly variable (see below), and the characters used by Steyermark to separate the two taxa are broadly intergrading. Therefore, the two names are here treated as synonymous.

**Notes on variation in vegetative parts and corolla dimensions:** After detailed study of herbarium specimens of more than 300 collections from throughout the geographic range, I concluded that the variations in leaf shape and size, hypanthium vestiture, corolla tube and lobes shape and dimensions of this species is not correlated with any geographical pattern. A certain clinal variation is observed in leaf dimensions, which are generally smaller at higher elevations. Leaves and corollas of *Sipanea biflora* are widely variable in the same patch and often on the same plant. To convey the degree of variation of this species, a detailed field study of three populations occurring in Suriname and Guyana is presented below:

**Suriname:** Sipaliwini District. Sipaliwini trail from Sipaliwini Village towards Sipaliwini Nature Reserve. Primary forest with canopy 15-25 m tall, with some large emergent trees, lateritic and white sand soil, 2°2'1"N, 56°6'45"W, 220 m, 19 Apr. 2014 (fl, fr), P.G. Delprete & G. Marjanom 12446 (B, BBS, BR, CAY, K, L, MO, P, RB, US) and 12447 (BBS, C, CAY, F, G, GB, K, NY, MG, ). **Plant description:** Decumbent herbs, rooting at basal nodes, forming patches several meters wide, area seasonally inundated. **Leaves:** blades ovate narrowly ovate to rhombic, 0.6-3.4 × 0.4-2.1 cm. **Corollas:** tubes 7-8.5 mm long; lobes oblong, obovate to broadly ovate, 3.5-5 × 1-2.5 mm.

**Guyana:** Potaro-Siparuni Region, Kaieteur National Park, trail between Guest House and Johnson's view, evergreen wet forests, thin sandy soil, mixed with bare rocks, 5°11'N, 59°29'W, 420-450 m, 23 Jun. 2016 (fl, fr). Populations studied and photographed, but no specimens collected (Figure 55). **Plant description:** prostate to somewhat decumbent herbs, rooting at basal nodes, forming dense patches 0.5-2 meters wide, growing on leaf litter accumulated on white-sand soil. **Leaves:** blades ovate, narrowly ovate to rhombic, 0.4-3.5 × 0.3-3 cm. **Corollas:** tubes 8.3-13.5 mm long; lobes ovate to oblong-ovate, 5-10 × 3-5 mm.

**Guyana:** Potaro-Siparuni Region. Upper Potaro River, Chenapou Village, trail between Kamo Creek and Urakak Creek, Mora forest on white sand, soil moist but not inundated, 4°59'43"N, 59°34'6"W, 220 m, 18 Jun. 2016 (fl, fr), P.G. Delprete & P. Benjamin 12869 (BRG, CAY, MO, NY). **Plant description:** prostate to somewhat decumbent herbs, rooting at basal nodes, forming patches

0.5-1.5 meters wide, growing on leaf litter accumulated on white-sand soil. **Leaves:** blades ovate to narrowly ovate, 0.6-3.2 × 0.4-2.8 cm. **Corollas:** tubes 8-12.5 mm long; lobes narrowly ovate to oblong-ovate, 6-11 × 3-4.5 mm.

In addition, a few gatherings have exceptionally large flowers. For example, Hoffman & Marco 2324 (CAY, U), from Pakaraima Mountains, Guyana, has corolla tubes 13.5-15 mm long and lobes 11.5-14 × 5-7 mm; however, the rest of the vegetative and reproductive characters are well within the species morphological range. At the other extreme, several specimens of *Sipanea biflora* have exceptionally small flowers. For example, Versteeg 863 (U), from Tapanahoni River, Suriname, has corolla tubes 5.5-6.5 mm long.

**Notes on variation in stamen insertion and style presentation:** As a result of a detailed analysis of numerous natural populations and herbarium specimens, *Sipanea biflora* is here confirmed to be distylous. In addition, there is a certain degree of variation in stamen insertion and style length within the two morphs. For example, in long-styled flowers the style could be either about the same length as the corolla tube or exserted just beyond the corolla mouth, and the stamens could be inserted at variable height of the corolla tube. To demonstrate this variation, a few examples are presented below.

*Long-styled flowers with stamens inserted at distal portion of corolla tube:* **Guyana:** Akarai Mts., Henkel et al. 4607: corolla tube 7.5 mm long; stamens inserted 3.7 mm below corolla mouth; filaments 0.5 mm long; anthers narrowly oblong, 1.5 × 0.15 mm, acute at both ends; style about the same length as corolla tube (tip of branches exserted), 7.5 mm long, glabrous; style branches linear, 0.8 mm long, acute at apex. **Suriname:**-Brownsberg, Wessel-Boer 636: corolla tube 10.5 mm long; stamens inserted 4.5 mm below corolla mouth, filaments 0.7 mm long; anthers 2.5 mm long; style about the same length as corolla tube (tip of branches exserted) style branches very narrowly lanceolate, 2 mm long, acuminate at apex. **French Guiana:** Lawa River, Papachon, Sastre et al. 8150: corolla tube 13.5 mm long; stamens inserted 5.5 mm below corolla mouth; filaments 0.7 mm long; anthers narrowly oblong, 2 × 0.2 mm, acute at both ends; style exserted 3 mm beyond corolla mouth, 17 mm long, glabrous; style branches linear, 1.7 mm long, acute at apex.

*Long-styled flowers with stamens inserted at about the middle of corolla tube:* **Suriname:** Upper Demerara River, Jenman s.n. (US Acc. No. 223383): corolla tube 15 mm long; stamens inserted 9 mm below corolla mouth; filaments 0.5 mm long; anthers linear, 2 × 0.2 mm,
acute at both ends; style exserted 1.5 mm beyond corolla mouth, 16.5 mm long, glabrous; style branches linear, 1.5 mm long, acute at apex.

*Short-styled flowers with stamens inserted at distal portion of corolla tube: Guyana:* Kaieteur Falls National Park, *Pipoly & Gharbarran 10147:* corolla tube 12.5 mm long; stamens inserted at 4–4.5 mm below corolla mouth; filaments 0.5–0.7 mm long; anthers linear, 2.2 × 0.2 mm, acute at both ends; style included, 5 mm long, glabrous; branches linear, 1.2 mm long, acute at apex.

**Distribution and ecology:** A species broadly distributed, ranging from Nicaragua, Costa Rica, Panama to Colombia, Venezuela, the Guianas, and Amazonian Basin and Atlantic forest of Brazil (Figure 56). Commonly found in riparian forests, semideciduous forests, evergreen lowland forests to lower mountain forests; growing on the forest floor, usually in areas rich in organic material and leaf litter, in lateritic soils or white-sand; sometimes found as aquatic/amphibian plant floating or semi-submersed in seasonally flooded area or in muddy soils of wet forests; at 5–750 m altitude.

**Phenology:** Due to the short life span of this species, it is common to find flowers and fruits on the same gathering. In Central America, specimens with flowers and fruits were collected in March, June, July, September, November and December. In South America, specimens with flowers and fruits were collected the whole year around.

**Suggested conservation status: Least Concern (LC).** A species distributed from Nicaragua, Costa Rica, Panama, Colombia, Venezuela, the Guianas, to the Amazonian Basin and Atlantic forest of Brazil. It is rare in Panama (collected in 1967) and Colombia (collected in 1854), where it is known only by a single collection in each country, and where it is locally endangered (or extinct?). This shade-loving species is most common on forest floor, and less frequent as an aquatic/amphibian plant in seasonally flooded, shaded areas, and sometimes often forming large patches. Because of its wide geographic distribution and its numerous collections, it is placed in the Least Concern (LC) category, following IUCN criteria (IUCN 2012, 2019).

**Specimens examined: NICARAGUA: Río San Juan:** San Juan del Norte, Río Indio, Caño La Paloma, 11°3'N, 83°53'W, 5 m, 1 Dec. 1982 (fl), *M. Araquistan* 3357 (MO, P); entre el pueblo de San Juan del Norte Nuevo y la casa de Ramón Castillo, viajando por el Caño

San Juanillo, 10°55'N, 83°49'W, 5 m, 7 Jul. 1994 (fl), *R.M. Rueda et al.* 1847 (MO).

**COSTA RICA: Alajuela:** Llanura de San Carlos, Lagunas Lagarto Lodge, 7 km N of Boca Tapada, 37 km N of Pital, 10°41'N, 84°10'W, 90 m, 6 Jun. 2000 (fl), *G.E. Crow* 9835 (CR, MO, NHA n.v.); San Carlos, Boca Tapada, Lagunas Lagarto Lodge, 7 km N of Boca Tapada, pantano en bosque, agua no muy profunda, 10°40'48"N, 84°10'48"W, 90 m, 27 Jul. 1996 (fl), *B.E. Hammell* 20356 (INB n.v., MO). **Limón:** Inundated forest at northern base of SW-most extension of Cerro Coronel, inundated lowland dominated by *Symphonia*, *Campnosperma*, *Manicaria* and *Cyclathaceae* understory, small clearing in forest for rice field, 10°40'30"N, 83°39'30"W, 5 m, 19 Sep. 1986 (fl, fr), *G. Davidse & G. Herrera* 31444 (MG, MO); Refugio Barra de Colorado, forest and pastures between Río Cirripocito and Río Sardina, marshy sites along carril in primary forest, 10°38'N, 83°45'W, 12 Nov. 1988 (fl), *M.H. Grayum et al.* 9018 (MO); Refugio Barra de Colorado, forest and pastures between Río Cirripocito and Río Sardina, muddy sites along trail in low, disturbed forest, 10°38'N, 83°45'W, 12 Nov. 1988 (fl), *M.H. Grayum* 9755 (F, INBIO n.v., MO); E of Cerro Coronel, opposite Laguna Danto, Raphia swamp, in standing water, 10°41'N, 83°38'W, 5 m, 20 Sep. 1986 (fl), *W.D. Stevens & O.M. Montiel* 24668 (MO); ca. 2 km S of Río Colorado, along new rd and ca. 1 km E; inundated forest with *Manicaria* and clumped *Sphaeradenia* dominant, margin of swamp forest, 10°40'N, 83°40'W, 5 m, 19 Mar. 1987 (fl, fr), *W.D. Stevens*, *G. Herrera & O.M. Montiel* 25080 (CAY, MG, MO).

**PANAMA: Veraguas:** Mouth of Río Concepción, swamp, 4 Dec. 1967 (fl, fr), *W.H. Lewis et al.* 2822 (MO, type of *Manettia hydrophila* Dwyer).

**COLOMBIA: Vichada:** “prope Maypures, flumen Orenoco” [Now Maipures, ca. 5°12'N, 67°50'W, on Río Orinoco], Jun. 1854 (fl), *R. Spruce* 3620 (BM, BR, NY, P).

**VENEZUELA: Amazonas:** Río Metacuni, Frente No. 5, selvas ribereñas y del banco, bajo del Salto “Pura Wániti’ma”, 22 Jan. 1990 (fl, fr), *B. Stergios & J. Velazco* 14114 (MO, NY, PORT, VEN); Mun. Río Negro, 1°49'N, 64°16'W, 427 m, Nov. 1992 (fr), *A. Valera* 152 (MO, NY, PORT). **Bolívar:** Región de Urimán, selva pluvial de Alto Río Chirca, 750 m, Aug. 1953 (fl, fr), *L. Bernardi* 950 (MER, NY); 32 km E of El Palmar, moist virgin forest, 4 Jun. 1960 (fl, fr), *J.A. Steyermark* 86259 (NY, VEN); Altiplanicie de Nuria, forest trail between El Cruzero and slightly beyond pica 101, ESE of Villa Lola, 315 m, 15 Jul. 1960 (fl), *J.A. Steyermark* 86390 (NY, P, US, VEN); low wet woods along trail at base of Cerro Pauji [ca. 8°8'N, 62°32'W], along Quebrada 94 at km 94 S of El Dorado, 250 m, 27 Jul. 1960 (fl), *J.A. Steyermark* 86713 (NY, VEN), 86731 (NY, VEN); near El Palmar, 8°5'N,
61°50′W, 75 m, 23 Nov. 1967 (fl, fr), J.G. Wessel Boer 2078 (NY, U). **Delta Amacuro:** Depto. A. Diaz, bosques húmedos primarios del medio Río Grande, 60 km al NE de El Palmar, 15 Mar. 1987 (fl, imm fr), G. Aymard 5440 (MO, PORT, VEN); E of Río Grande, directly E of El Palmar, swampy mature forest, 6 Jul. 1975 (fl, fr), A. Gentry & P. Berry 14979 (MO, NY, U); Río Amacuro, Venezuela-Guyana frontier, Río Amacuro, Sierra Imataca, in low places along water at Salto de Quebradero, upstream from San Victor, 65–80 m, 2 Nov. 1960 (fl), J.A. Steyermark 87252 (F, NY [3 sheets], VEN).

**GUYANA:** Kaieteur, 29 Dec. 1948 (fl), D.J. Atkinson 39 (BM); Upper Takutu-Upper Essequibo Region, Kuyuwini River, area near camp, disturbed vegetation, 2°5′N, 59°17′W, 240 m, 19 Jul. 1996 (fl), D. Clarke 2309 (MO, US); Upper Takutu-Upper Essequibo Region, Kassikaityu River, 0–1 km N of landing at terminus of trail from Kuyuwini River, 1°50′N, 59°5′W, 240 m, 18 May 1997 (fl), D. Clarke 4658 (MO, US); Potaro-Siparuni Region, Upper Potaro River, Chenapou Village, between Kamo Creek and Urakak Creek, mora forest on white sand, not inundated (terra firme), 4°59'43"N, 59°34'6"W, 450 m, 18 Jun. 2914 (fl), P.G. Delprete & P. Benjamin 12869 (BRG, CAY, MO, NY); Loc. Mapuera Camp, Riverside, Acarai Mountains [ca. 1°50′N, 57°30′W], 25 Oct. 1952 (fl), Forest Department of British Guiana Field No. G-495 (Record No. 7510) (NY); Potaro-Siparuni Region, Kaieteur Falls National Park, mostly evergreen forest to 20 m, 5°10′N, 59°29′W, 500 m, thin brown-white sand-gravel substrate, 29 Mar. 1988 (fl, fr), L.J. Gillespie & H. Persaud 973 (MO, US); Cuyuni-Mazaruni Region, Aurora, creek ca. 1 km WSW of camp, 1 km W of camp along 10,400 line, 6°47′N, 59°45′W, 70 m, 16 Oct. 1989 (fl, fr), L.J. Gillespie & S. Tiwari 2387 (MO, NY, US); Potaro-Siparuni Region, Kaieteur Falls National Park, mostly evergreen forest to 20 m, 5°10′N, 59°29′W, 500 m, thin brown-white sand-gravel substrate, 29 Mar. 1988 (fl, fr), W. Hahn et al. 4023 (MO, NY, US); Potaro-Siparuni Region, Kaieteur National Park, Kaieteur Gorge, W bank of Potaro River, 0.75 km from falls, 5°11′N, 59°28′W, 200–250 m, sandstone and conglomerate with medium stature forest, 50–100 m upslope from river, 13 Jul. 1993 (fl, fr), T.W. Henkel & R. Williams 2187 (MO, NY, US); Upper Takutu-Upper Essequibo Region, Acarai Mountains, Watuwau Creek, 8–10 km upstream of junction with Chodikar River, 1°22′N, 58°42′W, 250 m, 22 Feb. 1994 (fl, fr), T.W. Henkel et al. 4607 (CAY, MO, NY, U, US); Cuyuni-Mazaruni Region, Pakaraima Mountains, Meanu River, at foot of Marali Falls, on rocks, growing in mixed herb, bryophyte, pterydophyte community, 6°15′N, 60°26′W, 150–200 m, 6 Aug. 1992 (fl), B. Hoffman & G. Marco 2324 (CAY, MO, NY, U, US); near

Kaieteur Falls, Potaro River, Essequibo County, 17 Jan. 1954 (fl), H.S. Irwin BG-94 (US); Upper Demerara River, s.d. (fl), G.S. Jenman s.n. (US 00634732) (US); Demerara, s.d. (fl, fr), G.S. Jenman 4096 (US); Demerara River, Mar. 1898 (fl), G.S. Jenman 7306 (K); Potaro-Siparuni Region, Kaieuteur Falls, trail between Governement Guesthouse and Johnson's view, open, shrubby vegetation mixed with patches of forests, thin sandy soil, mixed with bare rocks, 5°11′N, 59°29′W, 420 m, 12 Oct. 1987 (fl, fr), L.P. Kvist et al. 227 (NY, U, US); Potaro-Siparuni Region, line from Kaieteur to Tukeit, 5°13′N, 59°27′W, 90–450 m, mixed forest, alluvial sand and gravel, growing on rocks, 15 Jul. 1991 (fl), K. Lance & C. Bird 18 (MO, US); Upper Mazaruni River, Kurupung [ca. 6°28′N, 59°10′W], 19 Nov. 1922 (fl, fr), H. Leng 159 (NY, paratype of S. cowanii); Potaro River Gorge, damp rock ledges, mixed forest, trail Tukeit to Kaiatuk Plateau, 29 Apr. 1944 (fl, fr), B. Maguire & D.B. Fanshawe 23086 (NY); N of Tinamu Falls [ca. 6°41′N, 58°59′W], Cuyuni River, Essequibo, on mossy dead log, 7 Mar. 1931 (fl, fr), E.B. Martyn 292 (K); Upper Takutu-Upper Essequibo Region, along track leading to airstrip village between Buro-Buro River and savanna, 4°8′N, 59°4′W, 70–80 m, 20 Feb. 1990 (fl, fr), T. McDowell et al. 1934 (CAY, MO, U, US); Barima-Waini Region, along trail between Baramita airstrip toward Millionaire, 7°22′N, 60°28′W, 91 m, mixed forest, lowland brown mud, red clay soil, 3 Apr. 1991 (fl, fr), T. McDowell et al. 4189 (CAY, MO, NY, U); Potaro-Siparuni Region, Kaieuteur Falls Nation Park, trail to Johnson's View, cliff area proceeding S, on W side of Potaro River, cloud and elfin forest, on sandstone, on rock wall, 5°11′N, 59°28′W, 360–400 m, 28 Jan. 1987 (fl, fr), J.J. Pipoly & G. Gharbarran 10147 (NY, P, U, US); without locality, s.d., J.P.B. Rohr s.n. ("dedit Vahl") (C [2 sheets]); gorge below Kaieteur Falls, ravine above Tukeit, 167 m, 1 Sep. 1937 (fl, fr), N.Y. Sandwith 1274 (K); Kaieteur Falls, top of precipices at edge of savannah, 400 m, 8 Sep. 1937 (fl, fr), N.Y. Sandwith 1437 (K, U); Cuyuni River, Arawak Matope, damp sand at edge of creek, 29 Jul. 1933 (fl), T.G. Tutin 399 (BM, U, US).

**SURINAME:** Sipaliwini District, trail from Sipaliwini Village towards Sipaliwini Nature Reserve, primary forest with canopy 15–25 m tall, with some large emergent trees, lateritic and white sand soil, 2°2′1′N, 56°6'45"W, 220 m, 19 Apr. 2014 (fl, fr), P.G. Delprete & G. Marjanom 12446 (B, BBS, BR, CAY, L, K, MO, P, RB, US), 12447 (BBS, C, CAY, F, G, GB, K, MG, NY); Berggakkij lands, 1835 (fl, fr), H.C. Focke 61 (U); without locality, s.d. (fl, fr), F.W.R. Hostmann & A. Kappler 150 (S); without locality, s.d. (fl), F.W.R. Hostmann 1121 (BM, P, U); Spoorlijn bij km 129.5 bij Kabel in bos, 25 Feb. 1956 (fl, fr), A.M.E. Jonker-Verhoef & F.P. Jonker 621 (U);
in sylvis humidis ad fl. Marowyne, Aug. 1846 or 1847 (fl), *A. Kappler 1890* (C, FI-Webb, P [2 sheets], S, U); Joden Savanne [ca. 5°25'N, 54°59'W], Mapanecreek area, Kamp 8, 21 Dec. 1960 (fl, fr), *K.U. Kramer & W.H.A. Hekking 2437* (U); Brokopondo Distr., Brownsweg [ca. 5°0'N, 55°9'W], ad viam ferream prope km 115-116, 5 Apr. 1961 (fl, fr), *K.U. Kramer & W.H.A. Hekking 3200* (C, F, U); road Afobaka-Brownsweg, N of Brokopondo Lake, 10 Nov. 1974 (fl, fr), *P.J.M. Maas et al. 2332* (NY, U); Paramaribo, "in umbrosis Surinami prope Paramaribo", May 1838 (fl, fr), *F.L. Splitgerber 937* (L); Brownsberg, Wallibase creek, 1 Jul. 1924 (fl, fr), *G. Stahel & J.W. Gonggrijp BW 6552* (U); without locality, s.d. (fl, fr), *Sulitterber s.n.* (P P00729417); Tapanahoni River, Sep. 1904 (fl), *G.M. Versteeg 863* (U); from Brownsberg at Kabel railroad to Mazaroni Peak, in desne forest, frequent in very wet, muddy places, 11 Feb. 1963 (fl, fr), *J.G. Wessel Boer 636* (NY, U).

**FRENCH GUIANA:** Saint Jean du Maroni [ca. 5°24'N, 54°4'W], lieux humides sous bois, 8 Jul. 1914 (fl), *R. Benoist 862* (P); Piste de Saint Laurent vers Paul Isnard, km 15, forêt primaire, 10 Nov. 1983 (fl, fr), *F. Billet & B. Jadin 1996* (BR, CAY); Beira do Rio Cassa, 9 Nov. 1954 (fl, fr), *G.A. Black et al. 54-17651* (IAN); Bassin de l'Approuague, Rivière Kourouaï, junction de la Kourouaï et la R.N. 2, 4°25'N, 51°56'W, 16 Oct. 1996 (fl), *M. Blanc 141* (CAY, MO, NY, P); Projet de route d'Apatou, entre 2 et 3 km E de la Crique Sparouine, Plateau Bastien, 5°15'50"N, 54°13'40"W, 130 m, forêt dense a sol bien drainé, 10 May 2004 (fl), *B. Bordenave & L. Betian 7363* (CAY, MO); Bassin de l'Approuague, R.N. 2, Crique Kapiri, 4°7'N, 52°5'W, 5 m, forêt inondable primaire, 12 Jan. 1991 (fl, fr), *G. Cremers 11505* (CAY); D.Z. 5, Route Régina-Saint-George, Bassin de l'Approuague, petite crique dans une forêt de basse altitude, 1 Dec. 1995 (fl), *G. Cremers & J.J. de Granville 14290* (CAY); piste forestière de St. Laurent à Paul Isnard, entre PK 10 et 40, 14 Feb. 1983 (fl, fr), *C. Feuillet 735* (CAY, P, UB); Upper Camopi River, ca. 1.5 km NE of Mont Belvédère, 4 Dec. 1984 (fl), *J.J. de Granville 7119* (CAY [2 sheets], MO, P); route Regina-Saint George, Bassin de l'Approuague, foret ripicole sur sable, 4°2'N, 52°1'W, 100 m, 26 Nov. 1995 (fl, fr), *J.J. de Granville & G. Cremers 13154* (CAY, K, MO, P, U); Projet routier entre Saint-Georges et le pont transfrontalier sur l'Oyapock, zone Crique Mazy, forêt sur flat, 3°52'20"N, 51°49'30"W, 5 m, 8 Dec. 2005 (fl, fr), *J.J. de Granville et al. 17184* (CAY, MO); Maroni River, s.d. (fl, fr), [initials unknown] *Hachenheim 150* (P); Région Organabo, Mana [ca. 5°40'N, 53°46'W], 8 Aug. 1962 (fl, fr), *F. Hallé 504* (P); Saül, Piste Sophie, dans un ruisseau sous bois, 7 Sep. 1962 (fl, fr), *F. Hallé 805* (US), *805a* (P); Haut Oyapock,

Trois Sauts [ca. 2°14'N, 52°52'W], vegetation secondaire, 11 Jul. 2006 (fl, fr), *C. Haxaire 1028* (CAY, MO); Camp Arataï, right bank of the Arataï River, a short distance from the mouth of the Approuague River, ca. 3°59'N, 52°34'W, 20 Feb. 2003 (fl), *S.A. Mori 25584* (CAY, NY); Basse Crique Kourouaïe (affluent basse Approuague), ca. 700 en mont du deuxieme camp, 13 Jan. 1970 (fl, fr), *R.A.A. Oldeman B-2724* (CAY [2 sheets]); Basse Crique Kourouaïe (affluent basse Approuague), ca. 900 en mont des Deux Fourches (dé fouca), rive droite, 15 Jan. 1970 (fl, fr), *R.A.A. Oldeman B-2750A* (CAY [2 sheets]); Route de Paul Isnard, km 40, selectively logged forest, 5°44'N, 53°58'W, 100 m, 18 Mar. 1994 (fl, fr), *J.H.E. Rova et al. 2005* (CAY, MO, NY, S); Fleuve Approuague[Basin], Rivière Arataye, Saut Pararé, layon X, 25 Aug. 1977 (fl), *C. Sastre 5765* (CAY, P); Fleuve Lawa, Papaichon, layon de la source, 3°49'N, 54°10'W, 80-120 m, 30 Aug. 1986 (fl, fr), *C. Sastre et al. 8150* (CAY, P, US); Prés de Saint Laurent [du Maroni], bas-fonds humides, 21 Aug. 1961 (fl), *R. Schnell 11375* (P); Znieff Abattis Kotica, 4°1'23"N, 54°18'50"W, 82 m, 15 Jan. 2012 (fl, fr), *P. Silland & R. Girault 372* (CAY); Znieff du flat à palmiers bâches de la Waki [Waki River], 3°10'50"N, 53°28'39"W, 129 m, 16 Aug. 2012 (fl, fr), *P. Silland et al. 835* (CAY).

**BRAZIL: Alagoas:** Mun. Matriz do Camaragibe [ca. 9°9'S, 35°31'W], Usina Serra d'Água, mata atlantica, 31 Oct. 2009 (fr), *Chagas-Mota & J.W.A. Silva 6357* (MAC); Ibateguara, Usina Serra Grande, Mata Coimbra, mata atlantica, 8°38'32"S, 35°55'46"W, 13 Oct. 2010 (fl), *R.P. Lyra-Lemos et al. 12082* (MAC). **Amapá:** ca. 1-3 km N of Cachoeira Tres Saltos, 2°12-13'N, 52°52-53'W, 12 Sep. 1960 (fr), *H.S. Irwin et al. 48214* (IAN, NY, US). **Bahia:** Mun. Una, Reserva Biológica do Mico-leão (IBAMA), entrada no km 46 da Rod. BA-001 Ilhéus-Una, 15°9'S, 39°6'W, Picada paralela ao Rio Maruim, 8-12 Mar. 1993 (fl), *A.M. Amorim et al. 1156* (CEPEC, NY, US); without locality, s.d. (fl, fr), *J.S. Blanchet s.n.* (P P00729425); without locality, 1835 (fl, fr), *J.S. Blanchet 1439* (FI-Webb); without locality, s.d. (fl), *J.S. Blanchet 1639* (P); Mun. Porto Seguro, Estação Biológica Pau-Brasil [ca. 16°22'S, 39°10'W], CEPLAC, erva crescendo na sombria da mata, 19 Apr. 1982 (fl), *A.M. Carvalho et al. 1189* (CEPEC, UB); Mun. Ilhéus, Mata da Esperança, acesso pelo Banco da Vitória, trilha que da acesso à represa, 14°46'60"S, 39°50'0"W, 22 Jan. 2010 (fl, fr), *L. Daneu et al. CEPEC 228* (HUEFS, RB); Porto Seguro, BR-5, km 21, Fazenda Iraipe, 6 Sep. 1961 (fl), *A.P. Duarte 6168* (RB); Una, Posse, 17 Dec. 1966 (fl, fr), *L. Emydio [de Mello] et al. 2474* (R [3 sheets]); Faz. Olhos d'Água do Laranjeiras, 11 Jan. 1976 (fr), *Gusmão, E.F. de 269* (RB, UB); Mun. Cravolândia, rd. Ilha Formosa de Cravolândia-Ubaira, ca. 8.5 km, 13°28'5"S, 39°39'25"W, 5 May 2007
(fl, fr), *J. Jardim et al. 5097* (CEPEC, NY); near the Ilheus River [“in viis umbrosis silvarum primaeverum prope fluvium Ilheos”], s.d. (fl, fr), [initials unknown] *Klaenze 114* (BR 824571) (BR); Cruz de Cosme, s.d. (fl, fr), *B. Luschnath 87* (C.F.P. Martius Herb. Fl. Bras. N. 594); Cruz de Cosme, Feb. 1835 (fl, fr), *B. Luschnath 87* (C.F.P. Martius Herb. Fl. Bras. N. 1315) (BR, FI, NY); without locality, s.d. (fl, fr), *B. Luschnath 100* (BR 824559) (BR); Cruz de Cosme, s.d. (fl, fr), *Collector Unknown* (probably Luschnath; C.F.P. Martius Herb. Fl. Bras. N. 594, p. 309) (NY 02690175); without locality, 1851 (fl, fr), *C.F.P. Martius s.n.* (*Herbar. Florae Brasil. No. 594*) (P); Mun. Una, km 27 da Rod. São José-Una (Jct. with BR-101), prox. Fazenda Piedade, 20 Jul. 1981 (fr), *L.A. Mattos et al. 1291* (IPA, NY); Mun. Ilheus, rd. between Sururú and Vila Brasil, 6-14 km from Sururú, 12 km SE of Buerarema, ca. 100 m alt., mata em destruição, 27 Oct. 1979 (fl, fr), *S.A. Mori & F. Benton 12892* (CEPEC, UB); Ilheus, Faz. Pirataquisse, 12 Mar. 1944 (fl, fr), *H. Pimenta Velloso 837* (R [2 sheets]); “in humidis pr. Ilhéos” [near Ilheus], May 1821 (fl, fr), *L. Riedel 142* (*Herb. Hort. Petropol. 139*) (BR, P); Igrapiúna, Faz. Michelin, 28 Feb. 2001 (fl, fr), *S. Sampaio 24* (MBM); Mun. Una, Reserva Biológica do Mico-leão (IBAMA), entrada no km 46 da Rod. BA-001 Ilhéus-Una, 15°9’S, 39°6’W, Picada da Bandeira, 25 Jul. 1996 (fl), *S.C. de Sant’Ana et al. 615* (MO, NY); Mun. Teixeira de Freitas, Vale do Rio Alcobaça, 15 Aug. 1972 (fl), *T.S. dos Santos 2374* (CEPEC, NY); Mun. Porto Seguro, Parque Nacional Monte Pascoal, 3 km S of entrance along rd on N side of Park, 16°51’S, 39°16’W, 6 Feb. 1999 (fl, fr), *W.W. Thomas et al. 12065* (CEPEC, MO, NY). **Pará:** Mun. Vitória do Xingu, 3°11'44"S, 51°46'52"W, 15 Jun. 2014 (fl, fr), *B.S.P. da Costa PSACF_EX03392* (RB); Mun. Vitória do Xingu, 3°11'59"S, 51°47'3"W, 30 Jun. 2014 (fl, fr), *B.S.P. da Costa PSACF_EX03439* (RB); Obidos, Igapó do Curuçambá, 8 Aug. 1912 (fr), *A. Ducke s.n.* (*Herb. Amaz. 12166*) (MG, RB); S. Manuel, Tapajóz, paludicola, fl roseas, Mar. 1912 (fl, fr), *F.C. Hoene 5202* (R); Rio Maracaná, Igarapé Açu, beira do Igarapé do Prata, 24 Sep. 1903 (fl, fr), *J. Huber s.n.* (*Herb. Amaz. 3809*) (MG, RB); Mun. Óbidos, Flota de Trombetas, floresta de transição de cerrado para floresta ombrófila submontana, 0°58'10"S, 55°31'3"W, Apr. 2008 (fl, young fr), *L.C.B. Lobato 3452* (MG); rodovia Belém-Brasília, km 54-62, 29 Nov. 1959 (fl, fr), *E. Oliveira 184* (IAN); Mun. Peixe Boi, estrada de ferro de Bragança, campo alagado, 22 Jun. 1951 (fl), *J.M. Pires 3381* (IAN, U); vicinity of Cachoeira, km 96, varzea forest, 30 Oct. 1965 (fl, fr), *G.T. Prance & T.D. Pennington 1867* (IAN, MO, NY, S, US). **Without State:** without locality [probably state of Bahia], s.d. (fl, fr), *P. Clausen 6* (BR); without locality, s.d. (fl, fr), *Collector Unknown s.n.* (BR [barcode 824568]), s.n. (BR

[barcode 824574]); without locality [probably state of Bahia], s.d. (fl, fr), *J.S. Blanchet s.n.* (C), s.n. (BM [barcode 000614331]); without locality, s.d. (fl, fr), *C.F.P. Martius s.n.* (NY [barcode 02690174]); without locality, s.d. (fl), *C.F.P. Martius Herb. Florae Brasil. 594* (L [barcode L0275963]); without locality, s.d. (fl, fr), *F. Sellow s.n.* (BM [barcode 000614345]), s.n. (S [Acc. No. 10-27501]), s.n. (S [Acc. No. 10-27503]).

## 8-8. *Sipanea carrenoi* Steyerm., Ann. Missouri Bot. Gard. 71: 333. 1984. (Figures 57-58).

Type: VENEZUELA. Bolívar: Gran Savana [Canaima National Park], at base of Salto del Aponguao [5°34'59"N, 61°19'12"; also known as Salto Aponwao or Salto Chinak-Merú], 42.5 km NE of Misión de Santa Teresita de Kavanayén, open densely covered slopes at base of salto, 1130 m, 22 Feb. 1978 (fl, fr), *J.A. Steyermark, V. Carreño E., G.C.K. Dunsterville & E. Dunsterville 115598* (holotype, VEN [Acc. No. 195063]; isotypes, MO [barcode 3223163], NY [barcode 00133309]). [Section *Virecta*].

*Herbs*, prostrate, trailing or sprawling, rooting at nodes; stems trailing, to 47 cm long, thin, 1.5–2 mm thick, terete, densely or sparsely antrorse- or subsreading-pubescent. *Stipules* adnate to the petioles, deltoid to triangular-lanceolate, 2–3 × 0.5–2 mm, acute at apex, membranaceous, densely strigose outside, glabrous, with 3–5 colleters at median portion inside; margins strigose. *Leaves* subsessile to short-petiolate; petioles 0.5–1.5 mm long, olive-green when dry, not winged, margins-pilose-ciliate; blades ovate-lanceolate, elliptic to elliptic-oblong, 0.7–2.3 × 0.4–0.9 cm [Steyermark (1984: 333) wrote “1–2 mm by 0.6–0.9 mm”], acute-decurrent at base, obtuse to subacute at apex, thinly subcoriaceous, dark brown above and brownish-olive green below when dry, lamina glabrous on both sides, midrib and secondary veins glabrous, margins narrowly revolute; secondary veins 2–4 on each side of midrib, obsolete to impressed above, prominulous below. *Inflorescences* terminal or appearing axillary (due to the reiteration of an axillary branch originating from a lateral bud at the base of the terminal inflorescence), a condensed dichotomous cyme, 3–7-flowered, rarely uniflorous, sessile to pedunculate, subtended by 2 leaf-like bracts (pherophylls), these ovate, elliptic to narrowly ovate, 0.4–1.5 × 0.3–0.6 cm; peduncle (when present) to 1.1 cm long, sparsely antrorse- or sub spreading-pubescent; bracteoles subtending flowers linear-lanceolate, 3–4 × 1–1.5 mm, glabrous. *Flowers* probably distylous, 5-merous, sessile to
<figure><img src="image_61.png" /><figcaption>**Figure 57.** *Sipanea carrenoi*. A. Habit. B. Node with stipule and petioles. C. Flower in anthesis. D. Dissected long-styled flower, with stamens inserted at distal portion of corolla tube, and exserted style. A–B: drawn from Steyermark et al. 115598 (MO); C–D: drawn from Steyermark et al. 115598 (NY). Illustration by Piero Delprete.</figcaption></figure>
short-pedunculate; pedicels (when present) to 1.5 mm long, sparsely antrorse-pubescent to antrorse-strigillose. *Hypanthium* narrowly obovoid, 2–2.5 × 1–1.5 mm, acute at base, round to obtuse at apex, densely antrorse-strigillose. *Calyx* lobes subequal, linear-lanceolate, 4.5–5.5 × 0.5–0.7 mm, acuminate at apex, glabrous, margins sparsely antrorse-pubescent; with 1 colleter in each calyxine sinus, conical, obtuse at apex, 0.3 mm long. *Corolla* hypocrateriform, 16.5–19.5 mm long, pink, turning pinkish-white at end of anthesis, with a dense ring of white hairs included below corolla mouth; tube cylindrical and slightly expanding towards the mouth, 10–13 mm long, 0.8–1 mm at base, 1.2–1.3 mm wide at mouth, sparsely antrorse-pubescent at distal portion and glabrous below outside, glabrous except densely white-vilous just below corolla mouth inside; lobes round, suborbicular to obovate, 5.5–6.5 × 4–5 mm [Steyermark (1984:

333) wrote “6–6.5 mm by 5–7 mm”], round to obtuse at apex, glabrous throughout. *Long-styled flowers* (type): stamens inserted at ca. 4 mm below corolla mouth; filaments 0.2 mm long; anthers linear, ca. 3 × 0.2 mm, acute at both ends; style as long as corolla tube, 11–13 mm long, glabrous; style branches obovate, 1 mm long, round at apex. *Short-styled flowers* unknown. *Infructescence* terminal or subaxillary (originally terminal), a condensed dichotomous cyme with 3–7-fruits, rarely a single fruit, similar to inflorescence. *Capsules* narrowly oblong-elliptic, 8–9 × 2–3 mm, acute at base, round at apex, crustaceous, densely antrorse hispidulous-strigillose, pale brown when dry. *Seeds* unknown.

**Notes:** This species is most likely distylous, as are the rest of the *Sipanea* species. In the only collection known, the style is as long as the corolla tube (style

<figure><img src="image_62.png" /><figcaption>**Figure 58.** Distribution of *Sipanea carrenoi* (star), *S. gleasonii* (circles), and *S. micrantha* (triangles).</figcaption></figure>
branches barely exserted), and stamens inserted ca. 4 mm below corolla mouth, which led me to conclude that these are long-styled flowers, pending future collection to confirm the heterostylous condition of this species.

Steyermark (1984) compared *S. carrenoi* Steyerm. with *S. pratensis*, and stated that the former differs in the creeping, rooting habit of the stems and the densely crowded, smaller, glabrous, obtuse to subacute leaves with fewer secondary veins. However, because of the dense ring of white hairs included below the corolla mouth, he should have positioned this species in his section *Cryptotricha*. Within this section, although Steyermark's sections are not recognized in the present treatment, it is similar to *S. biflora* as here delimited (including *Sipanea cowanii* Steyerm.), because of the creeping habit, thin stem, and small, membranous leaves, from which it differs by the stipules deltoid to triangular-lanceolate, 2-3 x 0.5-2 mm, densely strigose outside (vs. broadly triangular, 0.3-2 x 0.7-1.2 mm, glabrous or sparsely strigillose outside in *S. biflora*), leaf blades ovate-lanceolate elliptic to elliptic-oblong, 0.7-2.3 x 0.4-0.9 cm (vs. ovate, rhombic-ovate or rarely oblong, (0.4-)1.2-5 x (0.35-)0.8-2.8 cm), calyx lobes linear-lanceolate, 4.5-5.5 x 0.5-0.7 mm (vs. linear-lanceolate to setaceous, 1-5 x 0.2-0.6 mm), and capsules narrowly oblong-elliptic, 8-9 x 2-3 mm (vs. subglobose, 3-5 mm in diam.).

Because of the creeping habit, the ovate-lanceolate, elliptic, elliptic-oblong to rhombic, 0.5-2.6 cm long, leaf blades, the corolla tubes (8-)10-13.5 mm long, and the capsules oblong-elliptic, *S. carrenoi* is most similar to *S. veris* (including *S. acinifolia*), of Section *Sipanea*, from which it differs by the ring of white hairs included below corolla mouth (vs. ring of yellow hairs exserted beyond corolla mouth in *S. veris*), stipules deltoid to triangular-lanceolate, 2-3 x 0.5-2 mm (vs. broadly triangular or shortly suborbicular, 0.5 x 0.5-0.8 mm), inflorescences 3-7-flowered (vs. 1-2-flowered), and corolla lobes 5.5-6.5 x 4-5 mm (vs. 6-10 x 4.5-8 mm).

**Distribution and ecology:** Only known from the type, collected in the Gran Sabana, Venezuela, near Salto de Aponguao, at 42.5 km NE of the Mission of Santa Teresita de Kavanayen (Figure 58). Growing in open densely covered slopes, on moist sandstone slopes below waterfall, at 1130 m altitude.

**Phenology:** The sole gathering, with flowers and mature capsules, was collected in February.

**Suggested conservation status:** Critically Endangered (CE). An extremely rare species only known from

the type collection, gathered near Salto de Aponguao (also known as Salto Aponwao or Salto Chinak-Merú). This beautiful waterfall is formed by the homonymous river, and is included within the Canaima National Park, southern Venezuela. The size of its population(s) is unknown, and is suspected to be quite small, because although the area of the park has been intensely collected for decades, it was found only once. Also, this waterfall, located within the national park, is frequently visited by tourists groups, causing a considerable impact on the vegetation of this locality. Because of its rarity, the most likely small population, and the considerable human pressure on the locality of occurrence, this species is treated as Critically Endangered (CE) following IUCN criteria (IUCN 2012, 2019).

8-9. *Sipanea gleasonii* Steyerm., Mem. New York Bot. Gard. 17: 265. 1967. (Figures 58-59).

Type: GUYANA: [Potaro-Siparuni Region] Tumatumari [ca. 5°22'N, 59°0'W], dense upland forest, 18 Jun.-8 Jul. 1921 (fl, fr), H.A. Gleason 431 (holotype NY [barcode 00133311]; isotype US [Acc. No. 1122933, barcode 00137726]). [Section *Virecta*].

*Herbs*, prostrate, rooting at basal nodes; stems 3-10 cm long, thin, terete or slightly quadrangular, green, densely antrorse-strigose. *Stipules* adnate to the petioles, broadly triangular to broadly ovate, 0.5-2 x 0.8-1.2 mm, round to obtuse at apex, membranaceous, glabrous or sparsely strigose outside, glabrous, with 3-5 colleters at median portion or 7-10 irregularly distributed colleters inside; margins irregularly fringed, long-pubescent. *Leaves* petiolate; petioles 3.5-15 mm long, green when fresh, winged on both sides (extension of decurrent blade), sparsely antrorse-strigose pubescent; blades ovate to elliptic-oblong, 1.6-5 x 1-3 cm, obtuse to acute and decurrent at base, obtuse at apex, membranaceous, grass-green above and dull pale green below when fresh, olive-green above and below when dry, lamina sparsely appressed pubescent above (hairs 0.7-1 mm long), midrib and secondary veins sparsely sericeous-pubescent, margin sparsely setaceous-pubescent ciliate; secondary veins 3-4 on each side of midrib, ascending, weakly evident above, slightly prominulous below. *Inflorescences* terminal or simingly axillary (due to the reiteration of an axillary branch originating from an an axillary bud at the base of the inflorescence), long-pedunculate, with a terminal 2-3-flowered cyme; rachis filiform, almost capilliform, peduncle 0.9-3.5 cm long, densely strigillose; bracts subtending terminal cyme linear-lanceolate
<figure><img src="image_63.png" /><figcaption>**Figure 59.** *Sipanea gleasonii.* A–B. Habit (both plants drawn to same scale). C. Dissected short-styled flower, with stamens inserted at distal portion of corolla tube (partially exserted), and included style. D. Longitudinal section of fruit. E. Detail of one hair, basally tuberculate. A–B, D–E: drawn from *Gleason* 112 (US, paratype); C: drawn from *McDowell et al.* 4824 (US). Illustration by Piero Delprete.</figcaption></figure>
to setaceous, 2–3.5 × 0.2–0.3 mm, acuminate at apex, sparsely antrorse-hispidulous, ciliate at margins; bracteoles subtending flowers (when present) linear, 2.5–2.7 × 0.2 mm, acuminate, sparsely antrorse-hispidulous. *Flowers* probably distylous, 5-merous, pedicellate; pedicels of lateral flowers 2–11 mm long, those of central flower 0.7–3 mm long, sparsely antrorse-hispidulous. *Hypanthium* subglobose to broadly obovoid, 1.3–1.7 × 1–1.5 mm, obtuse at base, round at apex, densely antrorse hispid-strigose; hairs basally tuberculate, thin, 0.7–1.1 mm long. *Calyx* lobes subequal, green when fresh, setaceous, 4–4.5 × 0.1–0.2 mm, acute at apex, glabrous, margins hispidulous-ciliate, hairs 0.5–0.7 mm long, antrorse-ascendent to appressed; with 1 colleter in each calycine sinus, linear, 0.3 mm long. *Corolla* narrowly infundibular form, 12–14 mm long, pale pink to white, with a ring of white sericeous hairs at mouth; tube narrowly obconical, gradually expanding towards the mouth, 7–9.5 mm long, 1 mm wide at base, 2–2.5 mm wide at mouth, lower half glabrous and upper half sparsely antrorse-pubescent outside, lower 4.5 mm glabrous and densely white-villous above inside; lobes oblong to oblong-obovate to broadly ovate, 4–8 × 2.5–3 mm, obtuse to round at apex, glabrous throughout. *Long-styled flowers* unknown. *Short-styled flowers* (Gleason 112 – corolla tube 9 mm long; McDowell et al. 4824 – corolla tube 9.5 mm long; Tutin 541 – corolla tube 7–8.5 mm long): stamens inserted at 2.5 mm below corolla mouth; filaments 0.6–0.9 mm long; anthers partially exserted (only tip exserted), narrowly oblong to linear, 1.4–1.8 × 0.2–0.3 mm, acute at both ends; style included, 5.5–7 mm long, glabrous; branches linear, 1–1.6 mm long. *Infructescences* terminal or simingly axillary, due to the reiteration of an axillary branch originating from an an axillary bud at the base of the inflorescence, long-pedunculate, with a terminal 2–3-fruited cyme, rachis filiform, almost capilliform, similar to inflorescences. *Capsules* globose, 2.8–3 mm in diam., round at both ends, crustaceous, pale green when young, turning pale brown when dry, densely erect-hispid; hairs basally tuberculate, 1–1.5 mm long. *Seeds* irregularly tetrahedral-globose, 0.5–0.7 mm in diam., testa foveolate.

**Notes:** This species is only known from a few flowering specimens, and each of them has only one or a few flowers. In all the flowers of the specimens examined the style is included, and the anthers are inserted near the corolla mouth, indicating that most likely they are short-styled forms. Additional collections with more flowers are needed to confirm its distylous condition.

*Sipanea gleasonii* is similar to *S. ovalifolia* because of the creeping habit, thin stems rooting at nodes and internodes, broadly ovate to broadly triangular stipules, leaf blades ovate to elliptic oblong, 1.6–5 × 1–3 cm long, few-flowered inflorescences, hypanthium and capsules with basally tuberculate hairs, and corollas with a ring of white hairs at mouth; the former differs from the latter by the leaves with 3–4 secondary veins on each side of midrib (vs. 3–6 secondary veins on each side of midrib in *S. ovalifolia*), inflorescences 2–3 flowered (vs. 3–5-flowered), hypanthium 1.3–1.7 × 1–1.5 mm (vs. 0.7–0.8 × 0.7–0.8 mm), calyx lobes setaceous, 4–4.5 × 0.1–0.2 mm (vs. linear to linear-lanceolate, 2.5–4 × (0.1–)0.2–0.4 mm), and corollas narrowly infundibuliform 12–14 mm long with tubes 7.5–9.5 mm long and lobes (4–)5–8 × 2.5 × 3 mm (vs. corollas narrowly campanulate, 4.3–5.3 mm long with tube 2.8–3.6 mm long and lobes 1.5–1.8 × 1.2–1.3 mm).

**Distribution and ecology:** Endemic to lower Potaro-Siparuni Region, central Guyana, from where it is known from a few collections (Figure 58). Growing on forest floor, on brown, sandy soils, at 137–200 m altitude.

**Phenology:** Due to its short life span, it is common to find flower and fruits on the same individual. Such specimens were collected in February, May, June–July and November.

**Suggested conservation status:** Critically Endangered (CR). This species is endemic to the lower Potaro-Siparuni Region, central Guyana. It is known from three collections, including the type, made in 1921 and 1933 near the town of Tumatumari [ca. 5°22′N, 59°0′W], and a few other localities in the lower Potaro-Siparuni Region. For all these localities, the population sizes of this species are unknown. The region surrounding the town of Tumatumari has fertile soils, which have been devoted to agricultural exploitation; the area has also been profoundly impacted by timber extraction used by several local factories producing matches, pencils and tooth-picks. Currently the area is also greatly impacted by mining activities, and the natural vegetation surrounding Tumatumari has mostly been destroyed, and only a few patches of secondary vegetation are present.

The two most recent collections of this species were made in 1991, one along the North Fork River, near Porkknoker Works, at 5°9′N, 59°8′W (McDowell 4824), and the other at the nearby village of Micobe, at 5°20′N, 59°30′W (Tiwari & Mengharini 515), at 137 and 200 m altitudes, respectively. Both of these localities are heavily impacted by gold hunting activities, either by mining, panning, or dragging the bottom of rivers with dredgers; and the forest surrounding the village of Micobe is partially destroyed.
In conclusion, all the localities where this species
has been collected are profoundly impacted by human
activities, where the natural forests have been destroyed
or reduced to meager secondary remnants. Considering
all the above, this species is Critically Endangered (CR),
following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** GUYANA: Berbice or Demerara County, along the Berbice-Rupununi Cattle Trail, 1919 (fl), A.A. Abraham 306 (NY); [Potaro-Siparuni Region], Tumatumari [ca. 5°22'N, 59°0'W], dense upland forest, 18 Jun.–8 Jul. 1921, H.A. Gleason 112 (NY, US, paratype); Potaro-Siparuni Region, along North Fork River, 0.5–1.5 km N of Konawark Rd., near Porkknoker works, forest floor, 5°9'N, 59°8'W, 137 m, 28 May 1991 (fl), T. McDowell 4824 (MO, NY, U); Potaro-Siparuni Region, Micobe, mixed forest on brown sand, 5°20'N, 59°30'W, 200 m, 21 Oct. 1991 (fl), S. Tiwari & A. Mengharini 515 (NY, US); Tiger Creek, Tumatumari [ca. 5°22'N, 59°0'W], wet clay in Mora forest near the creek, 300 ft. [ca. 90 m], 24 Aug. 1933 (fl, fr), T.G. Tutin 541 (BM, U, US);

8-10. *Sipanea micrantha* Sandwith, Bull. Misc. Inform.
Kew 1928(9): 378. 1928. (Figures 58, 60).

Type: GUYANA. Mazaruni River, Kamakusa [ca. 5°57'N, 59°54'W, ca. 80 m], Sep. 1925 (fl), A.A. Abraham 404 (holotype K [without barcode]; isotype NY [barcode 00133314]). [Section *Virecta*].

Herbs, prostrate, rooting at nodes; stems to 35 cm long, thin, 0.3-0.5 mm thick, terete, green, appressed antrorse-pubescent. *Stipules* adnate to the petioles, broadly transversely elliptic to truncate, 0.3-0.4 x 0.4-0.6 mm, membranaceous, appressed antrorse-pubescent outside, glabrous, with 3-5 colleters inserted just below the margin inside, tips exserted beyond margin; margins longispid. *Leaves* petiolate; petioles 2-6 mm long, narrowly winged, sparsely antrorse-pubescent; blades ovate to rhombic-ovate, 5-17 x 3-7 mm, acute to obtuse at base, acute at apex, chartaceous, brownish olive-green when dry, lamina glabrous above, glabrous and with midrib and secondary veins with secondary veins sparsely strigose below, margin glabrous; secondary veins 2-3 on each side of midrib, ascending, weakly evident above, slightly prominulous below. *Inflorescences* terminal or subaxillary (originally terminal), pedunculate, rachis capilliform, with a terminal (1)2-flowered cyme, peduncle 1.2-1.8 cm long, appressed antrorse-pubescent; bracteoles subtending flowers linear-lanceolate, 0.9-1 x 0.2 mm, acute at apex, sparsely appressed-puberulent. *Flowers* most likey disty-

lous (only one flower dissected), (4)5-merous, pedicellate; pedicels of terminal flower 3-6 mm long, those of lateral flowers 1.5-2 mm long, appressed antrorse-pubescent. *Hypanthium* obovoid, 0.7-0.8 x 0.7 mm, acute at base, round at apex, glabrous or with a few appressed hairs. *Calyx* lobes subequal, linear-lanceolate, 1.3-1.7 x 0.2-0.3 mm, acute at apex, with a few appressed-pubescent hairs; with 1 colleter in each calycine sinus, conical, obtuse at apex, 0.1 mm long. *Corolla* campanulate, 4-6.5 mm long, white, with a ring of white hairs at mouth; tube obconical, 2.5-3.5 mm long, 0.5-0.6 mm wide at base, 1.5-2 mm wide at mouth, glabrous outside, lower half glabrous and shortly puberulous above, and white-villous in the areas where anthers are located inside; lobes oblong-ovate to oblong-obovate, 1.5-3 x 1-1.5 mm, obtuse to round at apex, puberulent inside, glabrous outside. *Long-styled flowers*: Stamens included, inserted 0.7-1 mm above base of corolla tube, filaments 0.4-0.6 mm long, anthers narrowly oblong, 0.75-1.2 x 0.15 mm; style about the same length as corolla tube (tips of style branches exserted), 2.5-3.3 mm long, style branches 0.6-0.7 mm long. *Short-styled flowers* unknown. *Infructescences* terminal or subaxillary (originally terminal), pedunculate, rachis capilliform, with a terminal (1)2-fruited cyme, similar to inflorescences. *Capsules* ovoid to subglobose, 2.5 mm in diam., round at both ends, crustaceous, glabrous or with a few appressed hairs. *Seeds* not seen.

**Notes:** Sandwith (1928) described *Sipanea micrantha* and cited *Abraham 404* at K as the type of this taxon; therefore this specimen is the holotype. He also cited *Jenman 2416* as the paratype of this taxon, without indicating the herbarium of deposit; this specimen has not been traced.

*Sipanea micrantha* is similar to *S. veris* in the creeping habit, thin stems, shallow stipules, leaf blades 5-17(-26) mm long, and inflorescences 1-2-flowered; the former differs from the latter in the hypanthium 0.7-0.8 x 0.7 mm (vs. 1.5-3.5 x 2-2.2 mm in *S. veris*), calyx lobes 1.3-1.7 x 0.2-0.3 mm (vs. (2-)3-7.5 x 0.1-0.6 mm), corolla campanulate, with a ring of white hairs at mouth, 4-6.5 mm long (vs. hypocrateriform, with a ring of yellow hairs at mouth, (14-)16-23.5 mm long) with tube obconical, 2.5-3.5 mm long and lobes 1.5-3 x 1-1.5 mm (vs. tube cylindrical, (8-)10-13.5 mm long and lobes 6-10 x 4.5-8 mm), and capsules ovoid to subglobose, 2.5 mm in diam. (vs. oblong-elliptic, 7 x 3 mm in diam.).

**Distribution and ecology:** A very rare species occurring in the Mazaruni River Basin, Guyana (Figure 58). Growing on forest floor, at 76-122 m altitude.
<figure><img src="image_64.png" /><figcaption>**Figure 60.** *Sipanea micrantha*. A–B. Habit (both plants drawn to same scale). C. Node with sheathing stipule with partially exserted colleters, and petioles. D–E. Dissected long-styled flowers, with stamens inserted at distal portion of corolla tube, and exserted style (drawn to same scale). F. Capsule. A, C–D: drawn from *McDowell 3779* (US); B, E–F: drawn from *Abraham 404* (K). Illustration by Piero Delprete.</figcaption></figure>
**Phenology:** The three collections, with flowers and fruits, were made in September and November.

**Suggested conservation status:** Critically Endangered (CR). This is a very rare species known from three localities in the Mazaruni River Basin, Guyana. It might be undercollected due to the minute size of its leaves and flowers. The first collection (Jenman 2416) was made in 1886, near the town of Bartica, at the triple confluence of the Cuyuni, Mazaruni and Essequibo Rivers; the corresponding specimen was cited by Sandwith (1928) but it was not traced. During the 20th century, Bartica underwent considerable expansion and the surrounding forest was generally eliminated; nevertheless, some patches of secondary forest are still present in the vicinities of the town.

The second collection, the type gathering of this species (Abraham 404), was made in 1925 near the town of Kamakusa, ca. 5°57′N, 59°54′W, at the edge of the Mazaruni River. The current status of conservation of the forest surrounding this town is unknown but is probably precarious, as the general area is subjected to mining activities.

The third and most recent collection (McDowell 3779) was made in 1990, also near the Mazaruni River, at 2–6 km upstream from Eping River Camp (Golden Star Resources, 6°5′N, 60°7′W). The current state of conservation of the forest at this locality is unknown. However, as the specimen label indicates, the area is subject of gold mining activities.

Summarizing the above discussion, the three localities where this species was collected are heavily impacted by human activities, and the population sizes are unknown, as no recent field observation in these localities has been made. Therefore, this species is estimated to be Critically Endangered (CR), according to IUCN criteria (IUCN 2012, 2019).

**Specimen examined:** GUYANA: Bartica Grove [ca. 6°24′N, 58°37′W], Nov. 1886, G.S. Jenman 2416 (cited by Sandwith (1928), not found); Cuyuni-Mazaruni Region, ranging 2–6 km upstream from Eping River Camp (Golden Star Resources), 6°5′N, 60°7′W, 76–122 m, 15 Nov. 1990 (fl), T. McDowell 3779 (MO, US).

8-11. *Sipanea ovalifolia* Bremek., Rec. Trav. Bot. Neerl. 31: 259. 1934. [Section *Virecta*. (Figures 61–63).

**Note:** For types and conservation status, see under varieties.

*Herbs*, prostrate, rooting at basal nodes and internodes; stems thin, terete or slightly quadrangular, green,

pubescent, antrorsely appressed pubescent (var. *ovalifolia*) or spreading to ascendingly villous (var. *villosissima*). *Stipules* adnate to the petioles, broadly ovate to broadly triangular, 0.4–0.7 × 0.8–1.2 mm, obtuse at apex, membranaceous, sparsely to densely antrorse-pubescent outside (hairs 1.5–1.7 mm long), glabrous, with 3–4 colleters inserted just below the margin inside, tips exserted beyond margin; margins long-hispid. *Leaves* petiolate; petioles 5–20 mm long, narrowly winged or not winged, sparsely to densely antrorse-pubescent or villous; blades broadly ovate, ovate to oblong-ovate, 1.7–4.7 × 1.2–3.2 cm, truncate, broadly obtuse to round at base, obtuse, round to subobtuse at apex, membranaceous, dark green above and pale green below when fresh, olive-green above and below when dry, sparsely appressed pubescent or lamina glabrous and veins sparsely appressed pubescent (hairs appressed, 0.4–1.3 mm long; var. *ovalifolia*), or lamina and veins villous (hairs erect, 1.5–2.5 mm long; var. *villosissima*) above, lamina and veins sparsely appressed pubescent, or lamina sparsely appressed pubescent and veins densely appressed pubescent, or lamina glabrous and secondary veins sparsely appressed pubescent below (hairs 0.6–1.3 mm long; var. *ovalifolia*), or lamina and veins villous below (hairs erect, 1.3–2.5 mm long; var. *villosissima*), margin ciliate; secondary veins 3–6 on each side of midrib, ascending, weakly evident above, prominulous below. *Inflorescences* terminal or subaxillary (originally terminal), long-pedunculate, with a terminal 3–5-flowered cyme, rachis filiform, almost capilliform; peduncle 2–4 cm long, antrorsely appressed-pubescent or antrorsely appressed-sericeous (hairs 0.5–1.2 mm long; var. *ovalifolia*), or ascendingly villous (hairs 1.3–1.5 mm long; var. *villosissima*); bracts subtending terminal cyme and bracteoles subtending flowers very similar, narrowly lanceolate to linear, 1.5–2.5 × 0.2–0.3 mm, acute at apex, glabrous, hispid at margins. *Flowers* distylous, 5-merous, subsessile to pedicellate; in 3-flowered cymes, pedicel of central flower 0.7–1.5 mm long and those of lateral flowers 1.3–3 mm long, in 4–5-flowered cymes pedicels 0.5–2 mm long, sparsely to densely antrorse-hispidulous. *Hypanthium* subglobose to broadly obovoid, 0.7–0.8 × 0.7–0.8 mm, obtuse at base, round at apex, densely antrorse-strigose, hairs basally tuberculate (hairs 0.7–1.3 mm long). *Calyx* lobes subequal, green when fresh, linear to linear-lanceolate, 2.5–4 × (0.1–)0.2–0.4 mm, acute at apex, densely antrorse-hispid (hairs antrorse-ascendent to appressed, 0.6–0.7 mm long); with 1 colleter in each calycine sinus, conical, obtuse at apex, 0.1 mm long. *Corolla* narrowly campanulate, 4.3–5.3 mm long white, with a ring of sericeous white hairs at mouth; tube basally urceolate, constricted just below the mouth,
<figure><img src="image_65.png" /><figcaption>**Figure 61.** *Sipanea ovalifolia* var. *ovalifolia*. **A.** Habit. **B.** Dissected long-styled flowers, with stamens inserted at median portion of corolla tube, and exserted style. **C.** Fruit. **A:** drawn from Wessel Boer 1554 (U). **B:** drawn from Florshutz & Florshutz 1242 (U). **C:** drawn from Oldeman B-2190 (CAY). Illustration by Piero Delprete.</figcaption></figure>
2.8 mm long (var. *ovalifolia*) or narrowly obconical, con-
stricted at medial portion and slightly wider at mouth,
3–3.6 mm long (var. *villosissima*), glabrous outside, lower
half (cylindrical portion) glabrous and upper half white-
puberulent, and densely white-villous near the mouth
inside; lobes ovate to oblong-obovate, 1.5–1.8 × 1.2–1.3
mm, obtuse to round at apex, glabrous on both sides.
*Long-styled flowers* (*Cremers* 6460, *Pulle* 222; corolla
tubes 2.8–3.5 mm long): stamens inserted at medio-
basal portion of corolla tube; anthers subsessile, oblong,
0.7–0.9 × 0.2 mm, acute at both ends; style as long as
corolla tube (tip of branches exserted), 3–3.5 mm long,
glabrous; branches obovate, 0.6 mm long, round at apex.
*Short-styled flowers* unknown. *Infructescences* terminal
or subaxillary (originally terminal), long-pedunculate,
with a terminal 3–5-fruited condensed cyme, rachis fili-
form, almost capilliform, similar to inflorescence. *Cap-
sules* globose, 2.5–2.8 mm in diam., round at both ends,
crustaceous, pale green when young, turning pale brown
when dry, densely erect hispid-strigose; hairs basally
tuberculate, 0.9–1.5 mm long. *Seeds* irregularly tetrahe-
dral, 0.5–0.6 mm wide, testa foveolate.

Notes: Steyermark (1984: 159–160) distinguished *Sipanea ovalifolia* var. *villosissima* Steyerm. from the typical variety “in having dense villous pubescence which covers the stems, petioles, peduncles, and the calyx lobes with hairs 1–2.5 mm long, the var. *ovalifolia* having only appressed pubescence.” Detailed studies with more material available supported the distinction of these varieties. For *S. ovalifolia* var. *villosissima* the only specimens available are the type specimens. Variety *villosissima* was recently photographed at Mount Itoupé, the type locality, by Sebatién Sant (Parc Amazonien de Guyane). In those photographs the villous vestiture of vegetative parts is quite evident, and proved that this variety is most likely endemic to this locality. Unfortun- nately the plants photographed in Mount Itoupé were not collected. Additional characters that can be used to distinguish these two varieties are found in the shape of the corollas, which are compared in the key below. Also, var. *ovalifolia* is known from Suriname and French Gui- ana, at 10–180 m altitude, while var. *villosissima* is only known from Mount Itoupé, Central French Guiana, at 600–700 m altitude.

Key to varieties of *Sipanea ovalifolia*

1. Stems, petioles and peduncles antrorsely appressed-pubescent or antrorsely appressed-sericeous; leaf lamina glabrous or sparsely appressed-pubescent and veins sparsely appressed-pubescent above (hairs appressed, 0.4–1.3 mm long), lamina glabrous and veins sparsely to densely

appressed-pubescent below (hairs appressed, 0.6-1.3 mm
long); corolla tube urceolate, constricted just below the
mouth, 2.8 mm long; corolla lobes oblong-ovate, 1.5 x 1.3
mm (Suriname and French Guiana, at 10-180 m altitude).
S. ovalifolia var. ovalifolia

1. Stems, petioles and peduncles spreading to ascendingly villous; leaf blades villous above (hairs erect, 1.5-2.5 mm long) and below (hairs erect, 1.3-2.5 mm long); corolla tube narrowly obconical, constricted at medial portion and slightly wider at mouth, 3-3.6 mm long; corolla lobes ovate, 1.5-1.8 x 1.2 mm (Central French Guiana, Mount Itoupé, at 600-700 m altitude)
S. ovalifolia var. villosissima

8-11a. *Sipanea ovalifolia* Bremek. var. *ovalifolia*, Rec. Trav. Bot. Neerl. 31: 259. 1934. (Figures 61, 62A-C, 63).

Type: SURINAME: Saramacca River, 16 Mar. 1903, A.A. *Pulle* 222 (holotype U [2 sheets, bar-codes 0006311 and 0006312]; isotype NY [barcode 00133315]).

Notes: The specimen *Pulle* 222 at U is mounted on two sheets that are kept together; therefore, according to Art. 8.3 of the *Code* (Turland et al. 2018), it should be treated as a single specimen with multiple preparations. The specimen *Pulle* 222 at NY is annotated as holotype by Steyermark (May 1965), but the holotype is at U.

*Stems* antrorsely appressed-pubescent or antrorsely appressed-sericeous. *Leaves* with petioles 7–20 mm long, narrowly winged or not winged, antrorsely appressed-pubescent; blades broadly ovate, ovate to oblong-ovate, 1.7–4.7 × 1.2–3.2 cm, truncate, round, obtuse to obtuse-subacute at base, obtuse, round to obtuse, often micro-scopically mucronate at apex, lamina glabrous or sparsely appressed-pubescent and veins sparsely appressed-pubescent above (hairs appressed, 0.4–1.3 mm long), lamina sparsely appressed-pubescent and veins sparsely to densely appressed-pubescent below (hairs appressed, 0.6–1.3 mm long); secondary veins 3–6 on each side of midrib. *Peduncles* antrorsely appressed-pubescent or antrorsely appressed-sericeous (hairs 0.5–1.2 mm long). *Corolla* 4.3 mm long; tube urceolate, constricted just below the mouth, 2.8 mm long; lobes oblong-obovate, 1.5 × 1.3 mm, obtuse to round at apex.

Vernacular name: FRENCH GUIANA: Yûsapilã
(Wayapi, Granville 2486).

**Distribution and ecology:** A variety known from several collections in Suriname and French Guiana (Fig-
<figure><img src="image_66.png" /><figcaption>**Figure 62.** *Sipanea ovalifolia*. A–C. *S. ovalifolia* var. *ovalifolia*. A. Inflorescence with one flower in anthesis. B. Detail of inflorescence and several leaves (note sparsely short-pubescent leaves). C. Habit. D–F. *S. ovalifolia* var. *villosissima* Steyerm. D. Inflorescence with one flower in anthesis. E. Detail of inflorescence and several leaves (note villous leaves). F. Habit. Photos by Sebastien Sant. A–C: from Gros-Saut, Papaïchton, Abounami River, French Guiana, plants not collected; D–F: from Mont Itoupé, E slope, French Guiana, plants not collected.</figcaption></figure>
ure 63). Growing on florest floor, in Mora forest, fre-
quently in humid or swampy places; at 10–180 m altitude.

**Phenology:** Specimens with flowers and fruits were collected in February, June, July and August, which corresponds with the local rainy season.

**Suggested conservation status: Near Threatened (NT).** This variety is a rare, shade-loving, prostrate herb, growing on the forest floor, known from five collections in Suriname and two in French Guiana. In Suriname it has been collected in 1903 (type), 1908, 1933, 1951, and 1963. These collections were made in remote localities, none of which has recently been visited by botanists, therefore the status of preservation of these forest areas is unknown. The most recent collection was made in 1963, in a forest south of Kappel Savanna [ca. 3°48′N, 56°8′W], a white-sand savanna surrounded by forest, located in central Suriname. A floristic survey of that savanna was published by Kramer & Donselaar (1968),

but the current status of preservation of the forest south
of it is unknown.

In French Guiana, it is known from two collections made in localities far away from each other: the first one (Oldeman B-2190) was made in 1969, in the Approuage River Basin, Ipoucin Creek, Camp Salvat [ca. 4°9′N, 52°27′W], at ca. 100 m elevation; the second one (Granville 2486) was made in 1975, in the Upper Oyapock River Basin, near the Takululi Creek [or Euleupousing Creek], ca. 1 km W of Saut Cambrouze [Cambrouze falls, ca. 2°20′N, 52°58′W], at ca. 180 m elevation; The size of the populations in these three localities is unknown, and their forests are apparently well preserved.

In conclusion, all the collections of this species have
been made in remote, uninhabited localities, and it is
supposed that they do not have any immediate threats,
although this remains to be confirmed.

**Specimens examined: SURINAME:** Lijn van Paka Paka (Saramacca) [ca. 4°37′N, 55°40′W], ca. 5 km from

<figure><img src="image_67.png" /><figcaption>**Figure 63.** Distribution of *Sipanea ovalifolia* var. *ovalifolia* (circles), *S. ovalifolia* var. *villosissima* (asterisk), *S. setacea* (star), *S. ayangannensis* (star), and *S. prancei* (squares).</figcaption></figure>
Ebbatop (v. Asch v. Wijck-Geb.), 9 Feb. 1951 (fl, fr), J. Florschütz & A. Florschütz 1242 (U); Suriname River, near Ketokvorsoe, 23 Aug. 1908 (fl), J.H.A.T. Tresling 367 (U [U0016826], paratype); S of Kappel Savanna [ca. 3°48′N, 56°8′W], near Tafelberg, in swampy place near creek in dense forest, 12 Jun. 1963 (fl, fr), J.G. Wessels Boer 1554 (NY, U).

**FRENCH GUIANA:** Upper Oyapock River Basin, Takululi Creek [or Crique Euleupousing], affluent of Ipisi Creek, ca. 1 km W of Saut Cambrouze [ca. 2°20′N, 52°58′W, ca. 180 m], forêt sur flat, 19 Jul. 1975 (fl, fr), J.J. de Granville 2486 (CAY [2 sheets], US); Approuage River Basin, Ipoucin Creek, Camp Salvat [ca. 4°9′N, 52°27′W, ca. 100 m], 25 Feb. 1969 (fl), R.A.A. Oldeman B-2190 (CAY, NY, P, U).

8-11b. *Sipanea ovalifolia* var. *villosissima* Steyerm., Britainia 38: 159. 1984. (Figures 62D-F, 63).

Type: FRENCH GUIANA. [Région des Emérillons] Sommet Tabulaire [Mount Itoupé, ca. 3°1′-5′N, 53°5′W], central zone, W slope, 600-700 m, ca. 40 km SE of Saül, 26 Aug. 1980 (fl), G. Cremers 6460 (holotype P [ex CAY] [barcode P00076246]; isotypes CAY [ex P, 2 sheets, barcodes CAY024971 and CAY024972], VEN [Acc. No. 161727]).

**Notes:** Steyermark (1984: 159), in the original publication of *Sipanea ovalifolia* var. *villosissima* Steyerm. stated that the holotype specimen of this variety was at CAY. However, as explained by Cremers (2001), in 1998 the holotypes of 31 taxa were transferred from CAY to P, and the corresponding isotypes were transferred from P to CAY. Therefore, the two isotype sheets at P (not reported in the original publication) of this variety are now at CAY, and the holotype specimen of this taxon is at P [barcode P00076246]. To confirm this, both isotype sheets at CAY have the label "HERBARIUM MUSEUM PARISIENSIS" and have the paper that at that time was used at P.

*Stems* spreading to ascendingly villous. *Leaves* with petioles 5-11 mm long, narrowly winged, villous; blades ovate to oblong-ovate, 2.2-3 × 1.2-1.7 cm, round, obtuse to acute at base, obtuse to acute at apex, lamina and veins villous above (hairs erect, 1.5-2.5 mm long), lamina and veins villous below (hairs erect, 1.3-2.5 mm long); secondary veins 5-6 on each side of midrib, ascending. *Corolla* 4.5-5.3 mm long, white; tube narrowly obconical, constricted at medial portion and slightly wider at mouth, 3-3.6 mm long; lobes ovate,

1.5-1.8 × 1.2 mm, obtuse to round at apex, glabrous on both sides.

**Distribution and ecology:** A variety only known from Mount Itoupé, central French Guiana (Figure 63). Growing on florest floor, in humid places, at 600-700 m altitude.

**Phenology:** Specimens with flowers and fruits were collected in August.

**Suggested conservation status:** Near Threatened (NT). This variety only known by the type specimens (Cremers 6460) was made in 1980 on the slopes of Sommet Tabulaire [or Mount Itoupé; ca. 3°5′N, 53°5′W], at 600-700 m altitude.

Mount Itoupé (or Sommet Tabulaire) is included within the French Guiana Amazonian Park (Parc Amazonien de Guyane). Scientific expeditions to this table mountain have been made in 2010, 2014, 2016 and 2017, and the pristine condition of its forests was confirmed. This variety has been photographed by Sébatien Sant (Parc Amazonien de Guyane) on 9 March 2017 during an expedition to Mount Itoupé, confirming its presence in this locality; unfortunately the plants photographed were not collected. Although there are apparently no immediate threats for this rare variety in such remote locality, the AOO is most likely extremely small, therefore it is here positioned in the Near Threatened (NT) category, following IUCN criteria (IUCN 2012, 2019).

8-12. *Sipanea setacea* Steyerm., Ann. Missouri Bot. Gard. 74: 114. 1987. (Figures 63-64).

Type: VENEZUELA. Bolivar: Otto. Piar, Cerro Vendo, 20 km E of Canaima, Cumbre Norte, "hierba escasa sobre rocas húmedas", 6°17′N, 62°41′W, 1300 m, 31 Aug. 1983 (fl), O. Huber, G.T. Prance & C. Alarcón 8246 (holotype NY [barcode 00133327]; isotype VEN not traced). [Section *Virecta*].

*Herbs*, creeping, growing on wet rocks, rooting at basal nodes and internodes; stems thin, 0.5 mm diam., terete or slightly quadrangular, green, densely spreading or loosely antrorse-pubescent (hairs 0.2 mm long). *Stipules* adnate to the petioles, truncate, with a central seta or two unequal setae, 2-3 × 0.1 mm, densely pilosulous; internal side not observed. *Leaves* petiolate; petioles 3.5-4.5 mm long, densely spreading or loosely antrorse-pubescent; blades ovate to elliptic-ovate, 1.2-2 × 0.9-1.2 cm, obtuse to subacute at base, acute at apex,
<figure><img src="image_68.png" /><figcaption>**Figure 64.** *Sipanea setacea*. A. Habit. B. Node with setaceous stipule and petioles. C. Inflorescence with a flower in anthesis. D. Dissected long-styled flowers, with stamens inserted at distal portion of corolla tube, and exserted style. A–D: drawn from Huber *et al*. 8245 (NY, holotype). Illustration by Piero Delprete.</figcaption></figure>
membranaceous, reddish-brown when dry (specimen pressed with alcohol 30-50%), densely subappressed-pubescent on both sides (hairs 0.2-0.3 mm long); secondary veins 5-6 on each side of midrib, ascending, weakly evident above, prominulous below. *Inflorescences* terminal, pedunculate, rachis filiform, almost capilliform, with a terminal 3-flowered cyme, peduncle 1-1.7 cm long, densely spreading or loosely antrorse-pubescent; terminal cyme subtended by two bracts, these linear-acicular, 2.2-2.4 x 0.1 mm [Steyermark (1987: 114) wrote "2.2 cm long × 0.1 cm wide"], margins setose-ciliate (hairs 0.3-0.5 mm long). *Flowers* most likely distylous, 5-merous, subsessile; pedicels to 0.5 mm long. *Hypanthium* narrowly obovoid, 1.5 x 0.8 mm, acute at base, round at apex, densely villosulous with spreading white, non-tuberculate hairs 0.4-0.5 mm long. *Calyx* lobes subequal, subulate or subteaseous, 2.5-2.6 x 0.3-0.4 mm, densely antrorse-pubescent (hairs 0.3-0.4 mm long); with 1 colleter in each calycine sinus, setiform, 0.15-0.2 mm long. *Corolla* infundibuliform, 7.5 mm long, "purple" ("corola morada" fide Huber et al. 8246), with a ring of white-sericeous hairs at mouth; tube narrowly obconical, 4 mm long, 0.7 mm wide at base, 1.5 mm wide at mouth, sparsely puberulent outside, shortly pubescent near the base of the stamens inside; lobes narrowly oblong, 3.5 x 1-1.5 mm, obtuse apex, moderately pilose outside. *Long-styled flowers:* stamens inserted at medio-distal portion of tube, filaments 0.6 mm long; anthers narrowly oblong, 1 mm long; style exserted beyond corolla mouth, glabrous, style branches linear, 1.5 mm long. *Short-styled flowers* unknown. *Infructescences, capsules* and *seeds* unknown.

**Notes:** Only the holotype specimen of *Sipanea setacea* Steyerm. was studied for this species. It only has one flower in anthesis, which was not dissected. Therefore, it is impossible to confirm if this species is heterostylous. In the sole flower observed, the style is exserted beyond the corolla mouth, and the included stamens were observed through the transparent dried corolla. Therefore, it is assumed that this is the long-styled form of this species.

Steyermark (1987: 115) wrote that the bracts subtending the inflorescence of *S. setacea* are "subulate, acuminate, 2.2 cm long, 0.1 cm wide"; however, these bracts are linear-acicular, 2.2-2.4 × 0.1 mm.

Steyermark (1987: 115) about *S. setacea* stated: "In its setaceous stipules and pubescent exterior of the corolla lobes, this species differs from other members of sect. *Virecta* subsect. *Cryptothricha*, but resembles them in habit, slender stem, and small leaves. From *S. ovalifo-

lia* Bremekamp [...], it differs in the shorter peduncle and petioles, longer corolla and corolla lobes, 3-flowered inflorescence on a shorter peduncle, shorter, non-tuberculate hairs of the hypanthium, narrower leaves, and purple instead of white corollas. From *S. micrantha* Sandw. it may be distinguished by the spreading pubescence of stems and peduncles, broader ovate leaves with pubescent upper surface, loosely pubescent calyx lobes, and 3-flowered inflorescence. From *S. gleasonii* Steyerm. it is differentiated by the subsessile flowers, shorter petiole, calyx lobes, hypanthium, and corolla tube, smaller leaf blades, shorter foliar pubescence, and non-tuberculate hairs of the hypanthium. Finally, from *S. cowanii* Steyerm. it differs in the densely spreading hairs of the hypanthium, the shorter corolla and calyx lobes, and the 3-flowered inflorescence." Steyermark's *Sipanea* sect. *Sipanea* subsect. *Cryptothricha* Steyerm. is here included in *Sipanea* sect. *Virecta*, as here delimited. Although Steyermark's subsection is not maintained, I concur with his comparisons of similar species, and *S. setacea* is here confirmed to be a distinct species.

**Distribution and ecology:** Only know from the type, collected in open scrub vegetation on tepui summit of Cerro Venado, ca. 6°17'N, 62°41'W, Bolivar state, Venezuela, at 1300 m altitude (Figure 63).

**Phenology:** The sole gathering, with flowers in anthesis, and without fruits, was collected in August.

**Suggested conservation status:** Vulnerable (VU). A species only known by the type specimen, collected in 1983 on Cerro Venado, ca. 6°17'N, 62°41'W, Bolivar state, Venezuela. Cerro Venado is located at ca. 20 km east of the town of Canaima, on the other side of Carao River, and is included within the Canaima National Park. The specimen label reports that it was a rare plant growing on humid rocks of the North Peak, at 1300 m altitude. This species might be undercollected because of its diminutive size. The population size is unknown, as it was not indicated in the specimen label and no recent field work has been made in this locality. Because of its rarity, it is treated as Vulnerable (VU) following IUCN criteria (IUCN 2012, 2019), pending future field studies to confirm the size and health of its population(s).

### 8C. *Sipanea* section *Nudae* (Steyerm.) Delprete, *sect. et stat. nov.*

*Sipanea* sect. *Sipanea* subsect. *Nudae* Steyerm., Mem. New York Bot. Gard. 17: 261, 268. 1967