Type: S. wilson-brownei Cowan

Herbs or subshrubs, erect or decumbent; corolla mouth glabrous or with sparse white hairs.

8-13. *Sipanea ayangannensis* Steyerm., Mem. New York Bot. Gard. 17: 269. 1967. (Figures 63, 65).

Type: GUYANA. Upper Mazaruni River Basin, Mount Ayanganna [ca. 5°24′N, 59°57′W], on shoulder of E flank, about Thompson Camp, 1418 m, "Small herb; leaves medium green, papery; calyx green, corolla tube red, limb pinkish-white, slightly irregular; frequent, dense scrub and low forest (8-10 m)", 12 Aug. 1960 (fl), S.S. Tillett, C.L. Tillett & R. Boyan 45114 (holotype NY [barcode 00133308]). [Section Nudae].

Herbs, decumbent, rooting at basal nodes; young branches slightly geniculate at least at basal nodes, densely antrorse-strigose (hairs yellowish-beige). *Stipules* adnate to the petioles, subulate, base triangular and medio-distal portion narrowly triangular to linear, 2.5-4 x 1.5-2 mm, densely strigillose outside, margins glabrous or sparsely antrorse-strigillose; internal side and presence of colleters not observed. *Leaves* often unequal in size on the same node, petiolate; petioles 4-6 mm long, densely antrorse-strigose (hairs yellowish-beige); blades lanceolate-elliptic, 4-5.5 x 1.7-2.2 cm, acute to decurrent at base, acute at apex, chartaceous, brown above and olive-green below when dry, lamina densely appressed-strigose above, sparsely strigose below, midrib and secondary veins strigose; secondary veins 5-7 on each side of midrib, slightly impressed above, prominent below. *Inflorescences* cymose, dichotomous, 5-7-flowered, condensed at initial stage of anthesis, 4-6 cm long (including peduncle and flowers in anthesis), expanding during and after anthesis; peduncle 1.3-2 cm long during anthesis and 2.7 cm long after anthesis, densely griseous-strigose; lateral branches 0.5-1 cm long, 3-5-flowered; bracts subtending branchlets narrowly lanceolate to linear, 4-6 x 0.3-0.5 mm, sparsely antrorse-strigose outside, margins antrorse-strigose; bracteoles opposing each flower narrowly lanceolate to linear-lanceolate, 3.5-7 x 0.5-0.7 mm, sparsely antrorse-strigose outside, margins antrorse-strigose. *Flowers* most likely distylous, 5-merous; the flower at base between the two branches with pedicels 0.7 mm long, densely antrorse-strigose; the other flowers sessile to subsessile. *Hypanthium* broadly turbinate to obovoid, 1.5-2 x 1.3-1.5 mm, acute to obtuse at base, round at apex, not tuberculate, densely

antrorse-strigose; hairs 0.7-1 mm long, not tuberculate. *Calyx* lobes subequal, oblong-lanceolate to oblanceolate, unequal, 5.5-7 x 0.7-1 mm, evidently veined, glabrous, midvein and margins strigose-ciliate; with 1 colleter in each calcine sinus, conical, obtuse at apex, 0.2 mm long. *Corolla* hypocrateriform, 24.9 mm long, corolla tube red, lobes pinkish-white, with a glabrous white ring at mouth; tube subcylindrical, gradually wider towards the mouth, 14.5 mm long, 0.6 mm wide at base, 1.5 mm wide at mouth, hirsutulous at medio-basal portion and densely griseous-strigose at distal portion outside, glabrous at basal 2.5 mm, villosulous above, puberulent at anther portion; lobes elliptic-oblong to obovate-oblong, 10.4 x 4.3 mm, round at apex, margins glabrous or sparsely ciliate. *Long-styled flowers* unknown. *Short-styled flowers* (Tillett et al. 45114): stamens inserted at 5.5-6.5 mm below corolla mouth; filaments 3 mm long, glabrous; anthers linear, 2 x 0.2-0.3 mm; style included, 6.5 mm long, glabrous; branches narrowly lanceolate, ca. 1 mm long, acute at apex. *Infructescences*, *capsules* and *seeds* unknown.

**Notes:** Steyermark (1967: 269) in the original description of *Sipanea ayangannensis* Steyerm. wrote that the corolla tube is 18 mm long and the corolla lobes are 9-9.5 x 4.5-4.7 mm. However, the sole corolla in anthesis of the holotype specimen has a tube 14.5 mm long and lobes 10.4 x 4.3 mm.

Steyermark (1967: 269-270) also wrote: "This species is distinct from *S. stahelii* and *S. wilson-brownei* in the larger corolla tube, prominently ciliate corolla-lobes, prominently strigose midrib of the calyx-lobes, externally hirsutulous spreading hairs of the corolla-tube throughout most its length, and obsolete calcine squamellae [i.e., colleters] or these not manifest." I concur with these observations.

**Distribution and ecology:** Only known from the type, collected on Mount Ayanganna, Guyana (Figure 63); growing in dense scrub and low forest, at 1418 m altitude.

**Phenology:** The sole flowering collection was made in August.

**Suggested conservation status:** *Critically Endangered (CR)*. A species only known by the type, collected in 1960 on the shoulder of the East flank of Mount Ayanganna, Guyana, at 1418 m altitude. Mount Ayanganna is part of the Pakaraima Mountain range, and is sandstone tepui reaching 2041 m altitude. One flank of the mountain is slanted, while the other flank has sev-
<figure><img src="image_69.png" /><figcaption>**Figure 65.** *Sipanea ayangannensis*. A. Habit. B. Dissected short-styled flower, with stamens inserted at distal portion of corolla tube, and included style. A–B: drawn from Tillett *et al*. 45114 (NY, holotype). Illustration by Piero Delprete.</figcaption></figure>
eral escarpments, these being an alternation of plateaux and steep slopes. Mount Ayanganna is also considered a national landmark, and each year, on 23 February, members of the Guyana Defense Force climb to its top and hoist the national flag to celebrate the anniversary of Guyana as a republic. It is also recognized as a center of biodiversity, with a high number of endemic frogs and plant species, many of which have recently been described (e.g., MacCulloch & Lathrop 2005; Cobb et al. 2007; Skog & Clark 2015; Wurdack & Michelangeли 2019). A description of the ascension, observations about the local vegetation, and endemic species, as well as botanical collections made by Bruce Hoffman and collaborators in October–November 1992 is available in Alexander et al. (2014).

The label of the type specimen reports this species as a small herb, locally frequent, growing in dense scrub and low forest (8–10 m tall). The population size was not reported. Although several botanical expeditions have been made on Mount Ayanganna, in 1955 (e.g., *Maguire et al. 40637*), 1987 (e.g. *Pipoly et al. 11177*), 1992 (e.g., *Henkel & Hoffman 154*), and 2001 (e.g., *Clarke et al. 9490*), this species remains known only from the type collection. The Pakaraima Mountain range is known to be an area of indiscriminate gold and diamond mining, and Mount Ayanganna is not included in a protected area. Therefore, because of its rarity, and the potential of gold mining exploitation in the area, this species is treated as Critically Endangered (CR) following IUCN criteria (IUCN 2012, 2019; area of occurrence < 10 km², and a single known locality).

8-14. *Sipanea prancei* Steyerm., Brittonia 33: 400. 1981.
(Figures 63, 66).

Type: BRAZIL. Roraima: [Yanomami Indigenous Territory], Indian trail from Surucucu to Uaicá, near Maitá Indian Village, 3°20'N, 63°24'W, terra firme forest, 14 Feb. 1971 (fl, fr), G.T. Prance, W.C. Steward, F.P. Harter, J.F. Ramos, W.S. Pinheiro & O.P. Monteiro 10528 (holotype NY [barcode 00133316]; isotypes INPA [Acc. No. 28860], MG [Acc. No. 42032], MO [Acc. No. 2988539], R [Acc. N. 180079], S [Acc. No. 10-28005], U [1574627], US [Acc. No. 2801068], VEN [Acc. No. 134008, 2 sheets]). [Section Nudae] (Figures 63, 66).

Herbs or *subshrubs* (basal internodes woody), to 50 cm tall, erect, sparsely branched; stems with pubescence in two lines on the internal margins of two opposite sides, antrorsely appressed-pubescent, the other

two sides glabrous. *Stipules* reduced to a line between the petioles, with 5-7 colleters intermixed with a few hairs. *Leaves* petiolate; petioles 0.3-3.7 cm long, antrorse appressed-strigose, often with curved hairs; blades ovate to elliptic, 2-7.5 x 1-4.5 cm, acute-decurrent to obtuse and shortly decurrent at base, acute or acuminate at apex, membranaceous, drying olive-green, sparsely strigose above and below, midrib and secondary veins densely to sparsely antrorse-strigose above and below (hairs 0.7-1 mm long); secondary veins 5-7 each side. *Inflorescences* laxly cymose during anthesis, dichomously branched, (3-)5-32-flowered, (1.5-)3-6 cm long when fully expanded; rachis thin, filiform, with pubescence in two lines on two opposite sides, antrorse appressed pubescent, the other two sides glabrous; peduncles 1-3 cm long; with two lateral branches dichotomously divided into two terminal branchlets; ultimate branchlets scorpiod when fully expanded, 3-8-flowered; or rarely the whole inflorescence extremely reduced, unbranched, (2-)5-8-flowered (on the same plant of the other dichomously-branched inflorescences), or with two small 1-2(-3)-flowered branches, inflorescences 3-5-flowered; bracteoles opposite to the insertion of each flower, linear to narrowly lanceolate, 1.7-4 x 0.2-0.4 mm, glabrous or sparsely appressed-pubescent. *Flowers* distyly not observed (see Morphological Observations, below), 5-merous, sessile. *Hypanthium* obovoid to subglobose, 0.7-0.9 x 0.5-0.7 mm, densely antrorse-pubescent; hairs inflate at base, not tuberculate. *Calyx* lobes subequal, linear to setaceous, 2.2-3 x 0.1-0.2 mm, margins sparsely appressed-ciliate; with 1 colleter in each calycine sinus, narrowly conical, acute at apex, 0.2 mm long. *Corolla* hypocrateriform, 8.5-11 mm long, tube pale green to greenish-white and lobes pink, with a glabrous white ring at mouth; tube narrowly cylindrical, slightly wider at median portion, at the area with anthers, 5.7-7.5 mm long, 0.5-0.7 mm wide at base, 1-1.2 mm at medio-distal portion, the anther position, and at mouth, sparsely pilose at distal portion and glabrous below outside, antrorse pubescent at medio-distal portion inside, with a thickened, glandular ring at mouth; lobes ovate, 2.7-3.5 x 1.4-1.5 mm, obtuse to acute at apex, glabrous outside, papillose inside. *Stamens* inserted 3.7-4.3 mm above the base of corolla tube [Steyermark (1981: 400) wrote "at 6 mm from base"); filaments 0.5 mm long; anthers linear, 1.2-1.5 x 0.1-0.2 mm. *Style* not completely elongated, included, filiform, 4-4.5 mm long; style branches still appressed against each other, linear, 1.5 mm long. *Infructescences* laxly cymodichomously branched, (3-)5-32-fruited, rachis filiform, similar to or slightly larger than inflorescence. *Capsules* globose, 2.5-3.5 mm diam., crustaceous, sparsely antrorse-strigose; hairs
<figure><img src="image_70.png" /><figcaption>**Figure 66.** *Sipanea prancei*. A. Habit. B. Node with stipule reduced to a line between petioles, exserted colleters, basal portion of petioles, and axillary vegetative buds. C. Terminal portion of inflorescence, with a flower in anthesis (dissected) and flower buds. D. Dehisced capsule. A, B, D: drawn from Jansen-Jacobs *et al.* 6010 (CAY). C: drawn from Daniels & Jonker 906 (U). Illustration by Piero Delprete.</figcaption></figure>
0.9–1 mm long, inflated at base, not tuberculate. *Seeds* irregularly polyhedral with smooth angles, 0.4–0.6 mm in diam.; testa foveolate.

**Morphological observations:** Several collections of *Sipanea prancei* were studied; however, most of them only have one or a few flowers in anthesis. Because of this, the flowers of only three collections were analyzed. In the specimens studied, the corolla tubes are 6.5–7.5 mm long, and the stamens are inserted at about the middle of the corolla tube (1/2 to 3/5 from base, i.e., 3.7–4 mm from the base). In the flowers analyzed, the style is slightly longer than the points of insertion of the stamens, with the style branches still appressed against each other or opened. With the material at hand it is impossible to confirm the heterostyly of this species. Additional specimens with flowers in anthesis are wanted to determine the possible distylous condition of this species. The three specimens analyzed provided the following proportions:

**Suriname:** Daniels & Jonker 906 (U): Corolla tube 7.5 mm long; stamens inserted at 4 mm from the base of the corolla tube (3/5 of corolla tube); style 4.5 mm long, branches opened.

**Guyana:** Jansen-Jacobs et al. 6010 (CAY, U; two flowers analyzed): Corolla tube 6.5 mm long; stamens inserted at 3.8 mm from the base of the corolla tube (3/5 of corolla tube); style 4.5 mm long, branches not opened.

**Guyana:** Kennedy & Ek 4759 (US): Corolla tube 7.3 mm long; stamens inserted at 3.7 mm from the base of the corolla tube (1/2 of corolla tube); style 4 mm long, branches not opened.

**Notes:** Steyermark (1981: 400) wrote that *Sipanea prancei* Steyerm. "is most closely related to S. stahelii Brem., from which it differs in the absence of conspicuously enlarged tuberculate-based hairs on the fruiting hypanthium, the longer hairs of the hypanthium, the conspicuously appressed pilose midrib and nerves on both upper and lower leaf surfaces, the elongate ciliate hairs on the leaf margins, and the shorter corollas. From S. gleasonii Steyerm. and S. ovalifolia Brem. it differs in the absence of the white villous pubescence within the orifice of the corolla tube, and the different habit, inflorescence, and leaf blades." I concur with Steyermark's statements.

*Sipanea prancei* and *S. stahelii* Bremek. are also similar to *Neobertiera gracilis*, because of the dichotomously branching inflorescence with a thin, filiform rachis, and globose fruits. However, *Neobertiera* has indehiscent crustaceous fruits falling off as dispersal units, whereas *Sipanea* has persistent loculicidal capsules with wind-dispersed seeds.

**Distribution and ecology:** Known by several collections from Guyana and Suriname, and the type specimens collected in the nearby Brazilian state of Roraima (Figure 63). Found in forests on granitic rocks, forested slopes, along trails in lowland forests, and secondary vegetation, at 75–500 m altitude.

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

**Suggested conservation status:** Vulnerable (VU). Known from several collections from remote localities in Suriname and Guyana, and one in the contiguous Brazilian state of Roraima. In Brazil it is only known by the type collection made in 1971 in the northern portion of Roraima, along the indigenous trail from Surucucu to Uaicá villages, near the Maitá Indigenous Village (ca. 3°20′N, 63°24′W), in *terra firme* forest. This locality is now within Yanomami Indigenous Territory.

In Suriname it is known from two collections made in 1959 on the Emmaketen Mountains, ca. 4°5′N, 56°12′W, at 325 and 350 m altitude.

In Guyana it is known from eight collections from five different localities. The first collection was made in 1992 in the Upper Demerara-Berbice Region, Mabura, DTL Concession, West Pibiri Compartment, Pibiri Creek and vicinity, 5°20′N, 58°30′W, 0–75 m, in secondary vegetation. Two collections were made in the Potaro-Siparuni Region, one in 1994 on the Pakaraima Mountains, Upper Ireng River, Julong Falls, and the other in 1997 near the Iwokrama Mountains, at 60 m elevation, inside the Iwokrama Forest Reserve (ca. 4–5°N, 58–59°W). Three collections were made in the Upper Takutu-Upper Essequibo Region in 1999 on the Wassarai Mountains, 1°35′N, 59°14′W, at 215, 300 and 715 m altitude. The most recent collection was made on 20 September 1999 along the Rewa River, Spider Mountain, Upper Essequibo River, 3°08′N, 58°32′W, at 400–500 m, in an open spot within forest growing on rock.

The only collection of this species from a preserved area was made in 1997 at the base of the Iwokrama Mountains, inside the Iwokrama Forest Reserve, Guyana. No recent field observation has been made to assess the conservation status of the localities where it has been collected in Brazil, Suriname and Guyana. Using all the collection localities, the extent of occurrence (EOO) of this species is calculated to be of ca. 151,000 km². Several collections sites in Guyana and Suriname within the EOO are influenced by indiscriminate, often illegal, gold hunting activities. Because the localities where it has been collected are threatened by these human activities,
it is here treated as Vulnerable (VU), following IUCN criteria (IUCN 2012, 2019).

**Specimens examined: SURINAME:** Emmaketen Mountains [Emma Range, ca. 4°5′–13′N, 56°11′–12′W], on bank of waterfall 1 km E of Main Camp, 350 m, 1 Sep. 1959 (fl, fr), A.G.H. Daniëls & F.P. Jonker 906 (U); Emmaketen Mountains, on top of a large granite boulder, trail upstream of Main Camp, 325 m, 7 Sep. 1959 (fl, fr), A.G.H. Daniëls & F.P. Jonker 946 (U).

**GUYANA:** Potaro-Siparuni Region, Iwokrama Rainforest Reserve, Iwokrama Mountains, 4.5 km transect from Karupurani-Annai Rd., to camp, dense forest on brown sand, with *Mora, Carapa* and *Socratea*, 4°20'15"N, 58°48'37"W, 60 m, 27 Mar. 1997 (fl, fr), *D. Clarke et al.* 4350 (MO, NY, US); Upper Takutu-Upper Essequibo Region, Wassarai Mountains, 0.2–0.5 km N of camp at base of mountains, semi-disturbed forest at base of escarpment on brown sandy clay and boulders, with *Euterpe, Jacaratia* and *Cecropia*, 1°35'57"N, 59°14'17"W, 300 m, 27 Aug. 1999 (fl, fr), *D. Clarke et al.* 7992 (MO, NY, US); Upper Takutu-Upper Essequibo Region, Wassarai Mountains, summit of unnamed peak, 8 km S of S. Kassikaityu River, dense forest on reddish sandy clay, with *Couratari, Pouteria* and *Inga*, 1°34'57"N, 59°14'17"W, 715 m, 31 Aug. 1999 (fl, fr), *D. Clarke et al.* 8092 (MO, NY, US); Upper Takutu-Upper Essequibo Region, Wassarai Mountains, near camp, along creek at base of highest peack in Wassarai Mountains (1230 m), dense forest on brown sandy clay, with *Inga, Socratea* and *Cecropia*, 1°34'8"N, 59°14'18"W, 275 m, 17 Sep. 1999 (fl, fr), *D. Clarke, R. Williams & C. Perry* 8744 (MO, NY, US); Upper Essequibo River, Rewa River, Spider Mountain, forest on rock, open spot, 3°08"N, 58°32'W, 400–500 m, 20 Sep. 1999 (fl, fr), *M.J. Jansen-Jacobs et al.* 6010 (CAY, NY, P, U [2 sheets]); Upper Demerara-Berbice Region, Mabura, DTL Concession, West Pibiri Compartment, Pibiri Creek and vicinity, secondary vegetation, 5°20"N, 58°30'W, 0–75 m, 3 Nov. 1992 (fl, fr), *H. Kennedy & R. Ek* 4759 (U, US); Potaro-Siparuni Region, Pakaraima Mountains, Upper Ireng River, 15–20 km upriver from Cipo settlement, Julong Falls, creekside and falls spray zone, waterfall meadow, 5°0"N, 59°57'W, 60 m, 2 Nov. 1994 (fl, fr), *P. Mutchnick et al.* 361 (MO, US).

8-15. *Sipanea saxicola* J.H. Kirkbr., Brittonia 49: 362, fig. 3. 1997. (Figures 67–68).

Type: BRAZIL. Pará: Mun. Oriximiná, 15 km NW of Cachoeira Porteira, Rio Cachorro, Serra do Cachorro [ca. 0°58'S, 57°10'W], terra firme, afloramento rocho-

so, "saxicola, ciófila, crescendo nas grutas formadas por rochas de salitre, local seco; pecíolos avermelhados, folhas levemente discolores, inflorescencia com brácteas verdes, calice verde, corola rósea, estames alvos, frutos imaturos verdes", 150 m, 8 Jun. 1980 (fl, fr), G. Martinelli, C. Davidson & C.D.A. Mota 7074 (RB203259) (holotype RB [Acc. No. 203259, barcode 00543717]; isotypes BRIT [barcode 24069], INPA [Acc. No. 101666], MG [Acc. No. 101066], MO [barcode 2292872], NY [2 sheets, barcodes 00658722 and 1747355], RB [2 sheets; barcodes 00543717 and 00560065], SPF [SPF 00163000], UB [Acc. No. 61930, barcode UB0040404], US [barcode 00512891]). [Section Nudae].

*Herbs* 15–20 cm tall, terrestrial or epipetric, erect, with a central taproot; young branches terete, green, spreading erect-pubescent (hairs thin, 0.9–1.2 mm long). *Stipules* reduced to a line between the petioles or very shallowly transversally ovate, 0.5–3 mm tall, sometimes with a central seta 0.5–0.7 mm long, glabrous outside, glabrous inside, with 5–8 colleters inserted in a row just below the margin inside, 1–2.2 mm long; margins with long-pubescent hairs (ca. 1.5 mm long). *Leaves* petiolate; petioles 0.2–1.5 cm long, reddish when fresh, winged on one or both sides (extensions of decurrent blade), sparsely spreading erect-pubescent; blades narrowly obovate, 9–12 × 2.5–4.3 cm, acute to decurrent at base, acute to obtuse at apex, membranaceous, grass-green above and dull pale green below when fresh, olive-green above and below when dry, lamina sparsely spreading erect-pubescent throughout, margin erect-pubescent, hairs thin, erect-sericeous; secondary veins 8–9 on each side of midrib, weakly evident. *Inflorescences* terminal, modified dichasia, 8–20-flowered, 5–11 cm long, expanding after anthesis, rachis thin, almost capilliform during anthesis, slightly thicker at fruiting stage, peduncle terete, 3–7 cm long, sparsely pubescent, with 1–3 pairs of lateral branches; lateral branches 0.5–4 cm long during anthesis, increasing in length when fully expanded (fruiting stage), scorpiod, more evident when fully expanded, 3–9-flowered; bracts subtending branches narrowly elliptic, 3–9 × 0.6–1.1 mm, acute at apex, sparsely erect-pubescent (hairs thin, erect-sericeous, 0.9–1.2 mm long), ciliate-pubescent at margins; rarely inflorescences simple, unbranched, 3–4-flowered (e.g., *Ferreira* 9718); bracteoles opposing each flower linear-lanceolate, 3–6 × 0.5–0.7 mm, sparsely pubescent, midvein and margins ciliate-pubescent, (hairs thin, erect-sericeous). *Flowers* distylous, 4- or 5-merous, pedicellate; pedicels 1.5–3 mm long, terete, sparsely erect-pubescent (hairs thin, erect-sericeous). *Hypanthium* globose, 1–1.5
<figure><img src="image_71.png" /><figcaption>**Figure 67. Sipanea saxicola.** A. Habit. B. Node with stipule reduced to a line between petioles, exserted colleters, and basal portion of petioles (note long, erect trichomes). C. Dissected short-styled flower, with stamens inserted at median portion of corolla tube, and included short style. D. Dehisced capsule with permanent calyx lobes. A–D: drawn from Martinelli *et al.* 7074 (MG). Illustration by Piero Delprete.</figcaption></figure>
× 1.5–2 mm, round at both ends, densely spreading erect-pubescent (hairs thin, 0.9–1.2 mm long). *Calyx* lobes subequal, green when fresh, very narrowly elliptic to almost linear, 7–10 × 0.6–0.8 mm, acute at apex, sparsely pubescent, margins spreading pubescent; with 1 colleter in each calcine sinus, linear, 0.1 mm long. *Corolla* hypocrateriform, 14–16.5 mm long, pink, with a glabrous white ring at mouth; tube narrowly cylindric, gradually expanding towards the mouth, 9–10.5 mm long, 1–1.5 mm wide at base, 2.5–4 mm wide at mouth, lower half glabrous and upper half sparsely pubescent outside, glabrous inside; lobes obovate, 5–6 × 3–4 mm, obtuse at apex, glabrous throughout. *Long-styled flowers* unknown. *Short-styled flowers* (Martinelli et al. 7074; Coelho et al. 213): stamens inserted at 4–5 mm below corolla mouth, about the middle of the tube; filaments

0.3–0.5 mm long; anthers linear, 2.5–3 × 0.2 mm, round at base, acute at apex; style included, 3–5 mm long, glabrous; branches narrowly oblong-ovate, 2.5–3.5 mm long, acute at apex. *Infructescences* terminal, modified dichasia, 8–20-fruited, with 1–3 pairs of lateral branches, more expanded than the inflorescences, rachis capilliform or slightly thicker; lateral branches scorpiod, 3–9-fruited. *Capsules* globose to very broadly ellipsoid, 5.5–6 × 4–5.5 mm, round at base and at apex, crustaceous, erect-pubescent; hairs thin, sericeous, 0.9–1.2 mm long, pale green when young, turning pale brown-beige when dry. *Seeds* irregularly globose to subglobose, 0.35–0.4 mm in diam., testa reticulate-foveolate.

**Distribution and ecology:** Endemic to Serra do Cachorro, Municipality of Oriximiná, state of Pará, Brazil.

<figure><img src="image_72.png" /><figcaption>**Figure 68.** Distribution of *Sipanea saxicola* (star) and S. wilson-brownei (circles).</figcaption></figure>
zilian Amazon (Figure 68). Growing on sandy soils, or sandy soil with saltpeter accumulation, at ca. 150 m altitude.

**Phenology:** It is common to find flowers and fruits on the same individual or even in the same inflorescence. Specimens were collected in May, June and November.

**Suggested conservation status:** Critically Endangered (CR). A species known by three collections near and at the base of the Serra do Cachorro, Municipality of Oriximiná, state of Pará, Brazil. The Serra do Cachorro is a tabular mountain at ca. 0°58'S, 57°10'W, and ca. 3 km from Rio Cachorro, a tributary of Rio Trombetas. This mountain has been visited by at least three botanical expeditions in 1980, 1985, and 1987. The Serra do Cachorro is now included inside the Faro State Forest, which is in the municipalities of Faro and Oriximiná. The label of the 1985 collection describes this species as a creeping herb growing on the sandy margins of the Rio Cachorro, which flows near the Serra do Cachorro. The label of the type specimen and that of the 1987 collection report it as an herb growing on sandy soil with saltpeter accumulation, at the base of Serra do Cachorro. The size of the populations was not reported on the specimen labels. Because of its rarity, the probably small populations, and its extent of occurrence (EOO) less than 100 km², this species is treated as Critically Endangered (CR) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** BRAZIL. Pará: Mun. Oriximiná, Serra do Cachorro [ca. 0°58'S, 57°10'W], Rio Cachorro, afluente da margem direita do Rio Trombetas, planta rasteira com flor lilás e botôes florais esverdeados, 20 Nov. 1985 (fl, fr), L.S. Coelho et al. 213 (INPA, NY); Mun. Oriximiná, Serra do Cachorro, 3 km do Rio Cachorro, afluente do Rio Trombetas, encosta da serra, crescendo sobre arenito e salitre, 0°58'S, 57°10'W, 28 Nov. 1987 (fl, fr), C.A.C. Ferreira (as "C.A. Cid") 9718 (INPA, MG, MO, RB).

8-16. *Sipanea stahelii* Bremek., Recueil Trav. Bot. Néerl. 31: 258. 1934. (Figures 69-71).

Emend. Steyerm., Mem. New York Bot. Gard. 17: 268-269. 1967.

Type: SURINAME. Wilhelmina Range ["Wilhelminageberte"], 20 Jun. 1926, G. Stahel 7168 (lectotype

U [barcode 0006313], here designated; isolectotypes U [3 sheets, barcodes U 006314, U 006315, U 006316]). [Section Nudae].

Herbs or *subshrubs* (basal internodes woody), 0.25-4 m tall, unbranched or sparsely branched, erect or decumbent, sometimes basal internodes prostrate, rooting at basal nodes; stems terete, reddish, terminal internodes slightly quadrangular, sparsely antrorsely appressed sericeous-pubescent. *Stipules* adnate to the petioles, broadly to shallowly triangular at base, narrowly triangular to almost linear distally, 3.5-4 × 1.5-2 mm, acuminate, sparsely appressed sericeous-pubescent outside, sparsely pubescent, with 6-9 colleters in a row just below the margin or concentrated just below the base of the central lobe inside; margins pubescent. *Leaves* of the same node often unequal, petiolate; petioles 0.7-3 cm long, sparsely antrorsely-appressed sericeous-pubescent; blades ovate, elliptic to oblong-elliptic, 3.5-9 × 1.2-4.7 cm, obtuse and shortly decurrent at base, acute to acuminate at apex, chartaceous or rarely membranaceous, dark green above and pale green below when fresh, drying reddish-brown above and pale brown to olive green below, lamina glabrous or very sparsely pubescent above, sparsely pubescent below, midrib and secondary veins glabrous or sparsely to densely antrorse-pubescent above, sparsely to densely antrorse-pubescent, red to reddish, slightly prominent below; secondary veins 6-10 each side. *Inflorescences* laxly cymose during anthesis, dichomously branched, (13-)16-48-flowered, 4.5-13 cm long when fully expanded, with a pair of lateral branches dichotomously divided into two terminal branchlets (for a total of four terminal branchlets), rarely with two pairs of lateral branches dichomously divided for a total of 8 terminal branchlets; rachis filiform; ultimate branchlets scorpiod when fully expanded, 4-8-flowered; peduncles 1.8-6.5 cm long; basal branches subtended by bracts narrowly lanceolate, 2.5-6 × 0.3-2.5 mm, or subtended by two leaf-like bracts (i.e., pherophylls), elliptic, up to 4 × 2 cm; rachis filiform, uniformly sparsely antrorse appressed pubescent; bracteoles opposite to insertion of each flower, linear to acicular, 0.8-2.5 × 0.1-0.3 mm, sparsely appressed-pubescent. *Flowers* distylous, 5-merous, subsessile to shortly pedicellate; pedicels 0.3-1.2 mm long, sparsely to densely pubescent. *Hypanthium* subglobose, 0.7-1.1 × 0.8-0.9 mm, densely antrorse-pubescent; hairs basally tuberculate. *Calyx* lobes subequal or unequal, linear to acicular, 1.5-3 × 0.2-0.5 mm, acuminate, glabrous, margins sparsely appressed-ciliate; with 1 colleter in each calyxine sinus, conical or linear, obtuse at apex, 0.2-0.3 mm long. *Corolla* hypocrateriform, 15-20 mm long, tube pale
green to greenish-white, lobes pink to lavender, with a thickened glabrous white ring at mouth; tube narrowly cylindrical, gradually wider towards the mouth, 10.5-12 mm long, 0.6-0.8 mm wide at base, 1.2-1.5 mm wide at mouth, sparsely pubescent at medio-distal portion and glabrous below outside, basal portion glabrous and puberulent at anthers position inside; lobes elliptic, oblong-elliptic to oblong-obovate, 4.5-8 × 1.8-3 mm, obtuse to subacute at apex, glabrous outside, papillose inside. *Long-styled flowers* (Schultz & Wessels Boer LBB 10223 and *Florschütz & Maas* 3116): stamens inserted at 2-3 mm from base of corolla tube; filaments 0.5-1 mm long; anthers linear, 1.2-2 × 0.1-0.2 mm; style about the same length or slightly longer than the corolla tube, 11-12.5 long, glabrous; style branches linear to filiform, 1.5-2 mm long, acuminate at apex, densely papillose when receptive. *Short-styled flowers* (Maguire 24314): stamens inserted at distal portion of corolla, at 4-4.5 mm below corolla mouth; filaments ca. 1 mm long; anthers linear, 1.8-2 × 0.2 mm, obtuse at both ends; style much shorter than corolla tube, 5-5.5 mm long, glabrous; style branches linear, 1.4 mm long, densely papillose when receptive. *Infructescences* laxly cymose, dichomously branched, 16-48-fruited, with a pair of lateral branches dichotomously divided into 2 terminal branchlets for a total of 4 terminal branchlets, or with 2 pairs of lateral branches dichomously divided for a total of 8 terminal branchlets; ultimate branchlets scorpiod when fully expanded, 4-8-fruited. *Capsules* apically loculicidally dehiscent at full maturity, globose, 3.7-4.5 × 3.5-4 mm, crustaceous, antrorse-pubescent; hairs basally tuberculate, more evident in fresh or rehydrated specimens, 0.4-0.6 mm long, basal tubercules conical, 0.2-0.3 mm long. *Seeds* irregularly polyhedral, subcompressed, with smooth angles, 0.6-0.9 mm in diam.; testa foveolate-reticulate.

**Notes:** Bremekamp (1934b: 258) along with the protologue of *Sipanea stahelii* Bremek. cited the gathering "Wilhelmina Range, Stahel 7168" without citing the herbarium of deposit. At U, where he worked, there is a type folder with four sheets of *Stahel 7168*, each sheet has a different label, and these specimens are not consecutively numbered. According Art. 8.3 of the *Code* (Turland et al. 2018) "A specimen may be mounted as more than one preparation, as long as the parts are clearly

labelled as being part of that same specimen, or bear a single, original label in common. Multiple preparations from a single gathering that are not clearly labelled as being part of a single specimen are duplicates, irrespective of whether the source was one individual or more than one." Therefore, these four sheets are duplicate specimens, and one of them should be designated lectotype. The specimen with barcode 0006313 has Stahel's original field label "510, 7168, 20 VI 26" and was annotated as holotype by Steyermark in 1965. However, Steyermark (1967: 268) cited the gathering without specifying that there are four sheets, and without designating a lectotype. Therefore, specimen 0006313 is here designated the lectotype of this name.

*Sipanea stahelii* is similar to *S. wilson-brownei*, and the two species have sometimes been confused in herbaria. Both of them can be found as herb, subshrub or shrub (basal internodes woody), 0.25-2(-4) m tall, with stipules adnate to the petioles, triangular at base and narrowly triangular to almost linear distally, leaves with 6-10 secondary veins on each side, inflorescences dichomously branched with scorpioid terminal branchlets, and hypanthia and fruits with hairs basally tuberculate. *Sipanea wilson-brownei* differs from *S. stahelii* by the inflorescence rachis thin but not filiform (vs. filiform in *S. stahelii*), bracteoles opposing each flower elliptic, narrowly elliptic, lanceolate, narrowly lanceolate to narrowly oblong-lanceolate, 3.5-7 × 0.3-1.7 mm (vs. linear to acicular, 0.8-2.5 × 0.1-0.3 mm), hypanthium narrowly obconical, narrowly ovoid to obovoid, 1-1.5 × 0.7-1.5 mm (vs. subglobose, 0.7-1.1 × 0.8-0.9 mm), calyx lobes oblong-lanceolate, lanceolate to narrowly lanceolate, 3.5-8 × 0.5-1.7 mm (vs. linear to acicular, 1.5-3 × 0.2-0.5 mm), corolla tube pink or red, sometimes red-dish-green at base, and lobes pink, often with a median linear white area (microscopically glandular) at base, the 5 lobes sometimes forming a white star-like pattern (vs. tube pale green to greenish-white and lobes pink to lavender), and capsules subglobose, broadly ovoid, ovoid to narrowly ovoid, 3.5-7 × 3-5 mm (vs. globose, 3.7-4.5 × 3.5-4 mm).

*Sipanea stahelii* is also similar to *S. prancei*, from which it differs by the hypanthia and fruits with hairs conspicuously tuberculate at base (vs. with hairs slightly tuberculate at base in *S. prancei*), and the corollas 15-20 mm long, with tubes 10.5-12 mm long and lobes 4.5-8

**Figure 69.** *Sipanea stahelii.* *A.* Habit. *B.* Node with stipule and basal portion of petioles. *C.* Dissected long-styled flower, with stamens inserted at median portion of corolla tube, and exserted style. *D.* Dissected short-styled flower, with stamens inserted at distal portion of corolla tube, and included short style. *E.* Dehisced capsule with permanent calyx lobes. *F.* detail of basally tuberculate hair present on the capsule. *A, B, D:* drawn from *Maguire 24314* (US). *C:* drawn from *Schultz & Wessel Boer LBB 10223* (US). *E-F:* drawn from *Irwin et al. 54918* (US). Illustration by Piero Delprete.
<figure><img src="image_73.png" /><figcaption>Figure 69.</figcaption></figure>
<figure><img src="image_74.png" /><figcaption>**Figure 70.** *Sipanea stahelii*. A. Habit. B. Inflorescence, side view. C. Flower in anthesis (note glabrous mouth). D. Inflorescence with flowers in anthesis, top view. E. Inflorescence with flower buds, flower in anthesis, and young fruits. Photos taken by Hervé Galliffet on 28 April 2019, Sentier Roche Bateau, Saül, French Guiana (reproduced with permission from the author, plants not collected).</figcaption></figure>
× 1.8–3 mm (vs. corollas 8.5–11 mm long, with tubes 5.7–7.5 mm long and lobes 2.7–3.5 × 1.4–1.5 mm).

*Sipanea prancei* and *S. stahelii* are also similar to *Neobertiera gracilis*, because of the erect habit, inflorescences dichotomously branched with a thin, filiform rachis, and globose fruits. *Neobertiera* differs from *Sipanea* by the indehiscent fruits falling off as dispersal units, while in *Sipanea* the fruits are persistent, loculicidal capsules, and the wind-dispersed seeds are released at capsule dehiscence. Due to the above-mentioned similarities, Delprete (2015) erroneously cited the Suriname collections Irwin et al. 54918, Schultz & Wessels Boer s.n. (LBB 10223) and Schultz J.P. s.n. (LBB 10471) under *Neobertiera gracilis* Wernham. However, after detailed observations of additional duplicate specimens, these three collections have apically dehiscent capsules, and hypanthium and capsules with basally tuberculate hairs (vs. with hairs basally inflated but not basally tuberculate in *N. gracilis*). The dry fruits of *Sipanea stahelii* at

the initial stage of maturity seem to be indehiscent, which lead to the misidentification of Delprete (2015). The dehiscence of these capsules takes place only at the final stage maturity, which was observed in recently analyzed duplicates, confirming that these three Surinamese collections belong to *S. stahelii*.

**Common name:** FRENCH GUIANA: alakua?a (Wayampi).

**Distribution and ecology:** Known only from several collections from Suriname and two collections from French Guiana (Figure 71). Growing in forests undercanopy on forested slopes, and top of table mountains, at (200–)390–1027 m altitude.

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

<figure><img src="image_75.png" /><figcaption>**Figure 71.** Distribution of *Sipanea stahelii*.</figcaption></figure>
**Suggested conservation status: Vulnerable (VU).**
This species is known from several collections from Suriname and two from French Guiana. In Suriname it is known from six collections from three localities. The first collection was made in 1944 on the flank of Tafelberg Mountain ("Table Mountain"), ca. 3°54'N, 56°10'W, at 390 m altitude. Four collections were made in 1963 on the Wilhelmina Mountains ("Wilhelminageberte"): at 1 km SE of Juliana Top at 800 m; at 3.5 km SSE of Juliana Top, 3°30'–36'N, 56°30'–34'W, at 500 m; on a nameless top SE of camp km 14 in line to Lucie River, at 700 m; and on Top 1059, SE of Julianatop, at 900 m. The most recent collection in Suriname was made in 1965 on the Bakhuis Mountains, ca. 4°21'N, 56°45'W, at 1027 m. Tafelberg Mountain is one of the highest mountains in Suriname, and is included in the Central Suriname Nature Reserve. The Wilhelmina Mountains, which include Julianatop, ca. 3°41'N, 56°32'W, 1280 m, the highest mountain in Suriname, are also part of the Central Suriname Nature Reserve. This reserve is a UNESCO World Heritage Site. The Bakhuis Mountains, a continuation of the Wilhelmina Mountains, are outside of the Nature Reserve, are rich in bauxite, nickel and copper, and have been the focus of several potential mining projects, which have not been realized due to local protests.

In French Guiana, this species is known from two collections. The first one was made in 1975 on Mont Saint Marcel, at 500 m altitude. Mont Saint Marcel, ca. 2°23'N, 53°1'W, with its 645 m altitude, is one of the highest mountains in French Guiana and is included within the French Guiana Amazonian Park (Parc Amazonienne de Guyane). Numerous sites of illegal gold mining sites are present within the park, although most of them are at low altitudes and in the central zone of French Guiana, while Mont Saint Marcel is located at the southern limit of the French Guiana.

The second collection of *Sipanea stahelii* in French Guiana was made in 1994 on Monts Belvédère, 3°38'N, 53°10'W, at 200–400 m altitude, located at 1.8 km from the town of Saül. This small mountain is inside the free-adhesion area of the French Guiana Amazonian Park, which means that it is under limited human pressure; for example, locals are allowed to fish and hunt in this area. On the other hand, no gold mining is allowed in the immediate vicinity of the town of Saül.

The number of individuals of populations present in Suriname and French Guianas is not reported in the specimen labels, and no recent field work has been made to access the status of conservation on the above described localities. The extent of occurrence (EOO) of this species is of ca. 40,000 km². It is tempting to say

that the localities within the Central Suriname Nature Reserve are without any immediate threat because the collection localities of this species are at 390–1027 m altitudes; while illegal gold mining is usually present at lower altitudes. This species is known from five distant localities and has a large EOO. Because the last collection of this species was made in 1994, and some localities might be impacted by illegal gold mining, it is estimated to be Vulnerable (VU) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined: SURINAME:** In montibus Bakhuis inter flum. Kabalebo et Coppename Sinistrum, Bakhuis Mountains [ca. 4°21'N, 56°45'W], top of 1027 m, 3 Mar. 1965 (fl), *J. Florschütz & P.J.M. Maas* 3116 (NY, U); Wilhelmina Geberte, forested lower slopes of Frederick Top, 3.5 km SSE of Juliana Top, 3°30'–36'N, 56°30'–34'W, 500 m, 20 Aug. 1963 (fl, fr), *H.S. Irwin et al.* 54918 (MO, NY, P, S, U, US); Wilhelmina Mountains, 1 km SE of Juliana Top [ca. 3°41'N, 56°32'W], headwaters of West River, frequent on forested slopes and granitic peak, 800 m, 27 Aug. 1963 (fl-fr), *H.S. Irwin et al.* 55129 (F, NY [2 sheets], RB, UB, US); Tafelberg ("Table Mountain"), [ca. 3°54'N, 56°10'W], base of N escarpment, 390 m, 11 Aug. 1944 (fl, fr), *B. Maguire* 24314 (NY, U, US), 15 Sep. 1944 (fr), 24774 (F, NY); Wilhelmina Mountains, top 1059, SE of Julianatop, 900 m, 24 Jul. 1963 (fl), *J.P. Schultz & J.G. Wessels Boer s.n.* (*LBB* 10223) (BBS [2 sheets], MO, U); Wilhelmina Mountains, nameless top SE of camp km 14 in line to Lucie River, 700 m, 24 Jul. 1963 (fl), *J.P. Schultz s.n.* (*LBB* 10471) (BBS, K, NY, U).

**FRENCH GUIANA:** Upper Oyapock River, Mont Staint Marcel [ca. 2°23'N, 53°1'W], NE Summit, foret basse broussailleuse, 500 m, 31 Jul. 1975 (fr), *J.J. de Granville* 2624 (CAY [3 sheets]); vicinity of Saül, Monts Belvédère, 3°38'N, 53°10'W, 200–400 m, 27 Mar. 1994 (fl), *J.H.E. Rova et al.* 2068 (S).

8-17. *Sipanea wilson-brownei* Cowan, Cowan et al. in Brittonia 7: 412. 1952. (Figures 5H, 68, 72–73).

Type: GUYANA. Komo Creek, Dec. 1948 (fl, fr), *G. Wilson-Browne* 637 (holotype NY [barcode 00133328]; isotypes U [barcode 000317], US [barcode 00137731]). [Section *Nudae*].

*Herbs, subshrubs or shrubs* (basal internodes woody), 0.25–2 m tall, terrestrial or epipetric, dichotomously branched, erect or decumbent, rooting at basal nodes; young branches geniculate or slightly geniculate at least at basal nodes, reddish, sparsely to densely antrorse-
pubescent or antrorse-sericeous. *Stipules* adnate to the petioles, deltoid, narrowly triangular or broadly to shal- lowly triangular to truncate at base and narrowly tri- angular to narrowly oblong-triangular distally, 2–6.5 × 1–3.5 mm, acute or acuminate at apex, sometimes becoming bifid at apex at later stages, apical teeth (when present) to 2 mm long, equal or sometimes unequal, sparsely to densely appressed-pubescent to appressed strigose-pubescent, sparsely antrorse-sericeous or rare- ly glabrous outside, glabrous and with 3–7 colleters at base or a 3–4 colleters on each side at base inside; mar- gins sparsely to densely pubescent or sericeous. *Leaves* petiolate; petioles 3–28 mm long, sparsely antrorse- pubescent or antrorse-sericeous; blades elliptic to ellip- tic-lanceolate, 2.5–10.5(–14.5) × 1–4(–5.5) cm, acute to decurrent at base, acuminate at apex, chartaceous, dark green above and dull pale green or reddish below when fresh, dark reddish-brown or olive-green above and pale brown below when dry; lamina glabrous or sparsely antrorse-puberulent above, glabrous, sparsely to densely appressed-puberulent or appressely hispidulous on secondary and tertiary veins below, midrib and sec- ondary veins sparsely to densely appressed-puberulent, often reddish below; secondary veins 6–10 on each side of midrib, complanar above, prominent below. *Inflores- cences* condensed-cymose (capitate) to shortly cymose at initial stage of anthesis, pauci- or multiflorous, 2.5–5.5 cm long long (excluding corollas), expanding during and after anthesis to 8 cm long; peduncle 2–3.3 cm long, sparsely to densely antrorse-puberulent or antrorse-seri- ceous, with 1–3 pairs of lateral branches; rachis thin (i.e., not filiform), slightly thicker in infructescences, second- ary branches cymose, 0.8–4 cm long during anthesis, dichotomously divided, ultimate branchlets scorpiod, more evident when fully expanded, 3–9-flowered; bracts subtending the flowering portion of secondary branches broadly to narrowly obovate, lanceolate, lanceolate, nar- rowly lanceolate to linear, 4.5–9 × 0.5–2 mm, glabrous, or with very few hairs, or sparsely antrorse-sericeous outside, margins antrorse-pubescent or antrorse-seri- ceous; bracteoles opposing each flower elliptic, narrow- ly elliptic, lanceolate, narrowly lanceolate to narrowly oblong-lanceolate, 3.5–7 × 0.3–1.7 mm, glabrous, or with very few hairs, or sparsely antrorse-pubescent outside, margins ciliate-pubescent or antrorse-sericeous. *Flowers* distylous, 5-merous, subsessile to pedicellate; pedicels 0.5–1 mm long, sparsely to densely antrorse-puberu- lent to densely antrorse-pubescent (strigose) or densely antrorse-sericeous. *Hypanthium* narrowly obconical, narrowly ovoid to obovoid, 1–1.5 × 0.7–1.5 mm, acute to obtuse at base, round to obtuse at apex, densely antror- se-strigose or densely antrorse-pubescent; hairs basally

tuberculate, 0.7–1.4 mm long. *Calyx* lobes subequal or unequal, green to reddish when fresh, oblong-lanceolate lanceolate to narrowly lanceolate, (2.5–)3.5–8 × 0.5–1.7 mm, glabrous, margins sericeous or long-hispidulous; with 1 colleter in each calcine sinus, narrowly coni- cal, acute at apex, 0.2–0.3 mm long. *Corolla* hypocra- teriform, 12.5–28 mm long, tube pink or red, sometimes reddish-green at base, lobes pink, often with a median linear white area (microscopically glandular) at base, sometimes forming a white star-like pattern with the 5 lobes, with a glabrous white ring at mouth; tube narrow- ly cylindrical, slightly wider at mouth, (7–)9.5–15 mm long, 0.5–1 mm wide for most length slightly wider at the distal portion below the mouth (at anther position in short-styled flowers), 0.8–2 mm wide at mouth, sparsely spreading or appressed antrorse-pubescent or antrorse- sericeous outside, densely puberulent from middle to distal portion of corolla tube, glabrous below and above inside; lobes elliptic, narrowly elliptic, narrowly obovate to obovate, 4–9.5(–14) × 2–6(–8) mm, round, obtuse or acute, rarely mucronate at apex, glabrous through- out (with a median linear white area (microscopically glandular) at base above, or microscopically glandular above and glabrous below. *Long-styled flowers* (*Crem- ers* 12781): stamens inserted at 3.5–4.5 mm from base corolla tube; filaments 0.5–1 mm long; anthers linear, 2–2.5 × 0.2–0.3 mm, round at base, acute at apex; style about the same length as corolla tube (tip of branches exserted) or exserted, 8.5–15 mm long, glabrous; style branches linear, 1–1.5 mm long, acute at apex. *Short- styled flowers* (Granville 14801, Granville 13588, Ferreira et al. 7630 [Oriximiná]): stamens inserted at 4–7 mm below corolla mouth; filaments 0.3–0.5(–0.7) mm long; anthers linear, 1.7–2.5 × 0.2 mm, round or acute at base, acute, often apiculate at apex; style included, 3.5–5.5 mm long, glabrous; branches narrowly lanceolate to linear, 1–1.3 mm long, acute at apex. *Infructescences* slightly or much expanded with respect to the inflorescence size, with lateral branches scorpioid, (2–)3.5–6.5 cm long; bracteoles persistent, even after the corresponding cap- sules have fallen off. *Capsules* broadly ovoid, ovoid to narrowly ovoid, 3.5–7 × 3–5 mm, round at base, obtuse to subacute at apex, crustaceous, sparsely ascending or spreading pubescent; hairs basally tuberculate, 0.9–1.4 mm long; pale green when young, turning pale brown to reddish-brown when dry; pale green when young, turn- ing pale brown when dry. *Seeds* irregularly tetrahedral- globose, 0.6–0.7 mm in diam., testa foveolate.

**Morphological notes:** *Sipanea wilson-brownei* Cow- an is highly variable in terms of corolla size, like several other species of the genus. Sometimes the corolla size
varies even within the same individual; for example, in *Jansen-Jacobs et al. 1611* (U), the corollas vary from 12.5 mm long (tube 9 mm long and lobes 3.5–3.7 × 1.1–1.3 mm) to 18 mm long (tube 11 mm long and lobes 6–7 × 2.9–3.2 mm). This species is also variable in vestiture of vegetative and reproductive parts. Young branches, stipules and petioles can be sparsely to densely antrorse-pubescent or antrorse-sericeous. The leaf lamina can be glabrous or sparsely antrorse-puberulent above, glabrous or sparsely to densely appressed-puberulent or appresse-ly hispidulous, with midrib, secondary and tertiary veins sparsely to densely antrorse or spreading puberulent.

*Sipanea wilson-brownei* is similar to *S. stahelii* and has sometimes been confused in herbarium specimens. For comparison and differentiation of this two species, see discussion under *S. stahelii*.

Two collections from a single locality in the state of Pará, Brazil, differ from the specimens from all other collection localities. The locality is within the municipality of Oriximiná, at ca. 0°58'48"S, 56°59'23"W, at 14 km north of the waterfall called Cachoeira Porteira, at the edge of Rio Trombetas. The two collections C.A.C. Ferreira & J. Ramos 1074 and C.A.C. Ferreira 7630 differ from all the other specimens by having inflorescences with both the bracts subtending the branched portion of secondary branches and the bracteoles opposing each flower sparsely antrorse-sericeous outside (vs. glabrous or with very few hairs outside in all other specimens), the corolla tubes 9.5–10 mm long (vs. 10–15 mm long), the corolla lobes 4–4.5 × 3 mm, round to obtuse at apex (vs. 4.5–14 × 2–8 mm, obtuse to acute, sometimes mucronate at apex), and the capsules 3.5 × 3 mm (vs. 5–7 × 2.8–5.5 mm). The features peculiar to these two collections from the municipality of Oriximiná are here interpreted as morphological variation of a variable, wide-ranging species.

**Distribution and ecology:** A species distributed throughout the Guiana Shield, in Amazonian Venezuela (Amazonas, Bolívar), Guyana, Suriname, French Guiana and Amazonian Brazil (Amapá and Pará) (Figure 68). A species adapted to a wide range of habitats, commonly found in the undercanopy of lowland forests, lower to higher montane, common among igneous outcrops and over sandstones, in forests of mountain summits, or high altitude cloud forest with *Clusia*, *Bonnettia*, *Helico-

nia, *Cecropia*, Rubiaceae and Myrtaceae, or in dense forest on brown sand and laterite, with *Caryocar*, *Sloanea* and *Aparisthmium*, or in semi-shady places of forested mountain slopes and in forests at the edges of inselbergs, often found at edges of forest creeks, and sometimes also found in secondary vegetation; on granitic, sandy, lateritic or clay soils, brown sand, or soils with a superficial layer of forest litter, at 50–1500 m altitude.

**Vernacular name:** FRENCH GUIANA: Pililaåga (Wayampi; *Sastre 4480*).

**Phenology:**—It is quite common to find fruits at the basal portion and flowers at the distal portion of the same inflorescence. However, specimens at the early stages of anthesis might not have developing fruits present. Flowering and fruiting specimens were collected the whole year around.

**Conservation status:** Least Concern (LC). A species widely distributed in southern Venezuela (Amazonas, Bolívar), Guyana, Suriname, French Guiana and eastern Amazonian Brazil (Amapá and Pará), adapted to a wide range of habitats and soil types (see Distribution and ecology, above), and ranging from 50 to 1500 m altitude. Because of its vast extent of occurrence (EOO > 500,000 km²) and its numerous collection localities, some of them included within protected areas, this species belongs to the Least Concern (LC) category, following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** VENEZUELA: Bolívar: Carretera El Dorado, ca. km 88, Nov. 1958 (fl, fr), *L. Aris-teguieta 3722* (NY, US, VEN); entre el km 88 y el 126, 8 Apr. 1970 (fl, fr), *A. Fernández 1100* (F, MY); 105 km de El Dorado hacia Santa Helena, 29 Dec. 1956 (fl), *E. Foldats 2679* (F); La Escalera, 107–118 kms S of El Dorado, bosque montano, 5°50'N, 61°30'W, 450 m, 2 Apr. 1985 (fl, fr), *B.K. Holst et al. 2052* (MO, VEN); N portion of Cerro Uroi, Río Uroi [ca. 6°18'5"N, 62°3'47"W], Río Chicanán [ca. 6°44'N, 61°37'W], 250–400 m, 12 Sep. 1962 (fl, fr), *B. Maguire et al. 53733* (F, NY [2 sheets], RB, UB, US); along forested dirt road between km 109 and 125, between El Dorado and Cerro Uei, 800–1000 m, 30 Apr. 1960 (fl, fr), *J.A. Steyermark & S. Nilsson 837* (NY, US, VEN).

**Figure 72.** *Sipanea wilson-brownei.* *A.* Habit. *B.* Dissected long-styled flower, with stamens inserted at median portion of corolla tube, and exserted style. *C.* Dissected short-styled flower, with stamens inserted at distal portion of corolla tube, and included short style. *D.* Dehisced capsule with permanent calyx lobes. *E.* detail of basally tuberculate hair present on the capsule. *A:* drawn from *Granville & Crozier 13588* (CAY). *B:* drawn from *Clarke et al. 4158* (CAY). *C:* drawn from *Clarke 2554* (CAY). *D-E:* drawn from *Mori & Smith 25056* (CAY). Illustration by Piero Delprete.
<figure><img src="image_76.png" /><figcaption>Figure 72.</figcaption></figure>
**GUYANA:** Potaro-Siparuni Region, Iwokrama International Rainforest Reserve, Iwokrama Mountains, summit of highest peak, 4°20′N, 58°46′W, 1000 m, 23 Sep. 1996 (fl), *H.D. Clarke* 2554 (CAY, MO, U, US); Upper Tukutu-Upper Essequibo Region, Kamoa Mountains, 2 km N of camp on Kamoa River, scrub forest on granitic outcrop, 1°32′50″N, 58°50′4″W, 520 m, 9 Nov. 1996 (fl, fr), *H.D. Clarke* 2992 (MO, US); Potaro-Siparuni Region, Iwokrama Rainforest Reserve, Karupukari-Annai Rd, below summit of unnamed peack, dense forest on brown sand and laterite, 4°28′N, 58°47′W, 500–600 m, 20 Mar. 1997 (fl), *H.D. Clarke* et al. 4158 (CAY, MO, NY, U, US); Cuyuni-Mazaruni Region, Pakaraima Mountains, transect 4 km along NE Plateau of Mout Ayanganna, 5°24′N, 59°57′W, 1100–1500 m, 6 Nov. 1992 (fl, fr), *T.W. Henkel & B. Hoffman* 154 (MO, US); Upper Tukutu-Upper Essequibo Region, NW of Kanuku Mountains, top to Nappi Mountain, 12 km S of Nappi village, 3°18′N, 59°33′W, 750–950 m, 8 Feb. 1993 (fl, fr), *B. Hoffman* & *R. Foster* 3549 (BRG, F, MO, US); Kanuku Mountains, Nappy Mt., 3°19′N, 59°34′W, 700–960 m,

11 Nov. 1987 (fl, fr), *M.J. Jansen-Jacobs et al.* 884 (CAY, MO, NY, P, U, US); Kamoa River, Toucan Mountain, forest on mountain slope, 1°33′N, 58°50′W, 260–360 m, 16 Sep. 1989 (fl), *M.J. Jansen-Jacobs et al.* 1611 (CAY, MO, NY, P, U, US); Potaro-Siparuni Region, ridge of Eagle Mountain, near trail of Golden Star, upper plateau of ridge, 5°12′N, 59°7′W, 300–685 m, 10 Oct. 1990 (fl, fr), *T. McDowell & D. Gopaul* 3478 (MO, US); Cuyuni-Mazaruni Region, Mount Ayanganna, E side on steep slopes, cloud forest dominated by *Clusia*, *Bonnettia*, *Myrtaceae* and *Rubiaceae*, 5°27′N, 59°57′W, 1250–1300 m, 12 Mar. 1987 (fr), *J.J. Pipoly et al.* 11177 (CAY, INPA, MO, NY, US).

**SURINAME:** Sipaliwini District, Sipaliwini Village, trail from village towards the Sipaliwini Nature Reserve, ca. 2 km from village, primary forest with canopy 15–25 m tall, lateritic soil and white sand, 2°2′1″N, 56°6′45″W, 220 m, 17 Apr. 2014 (fl), *P.G. Delprete & G.P. Marjanom* 12408 (CAY); Massif de Tumuc-Humac, Inselberg Talouakem, 2°29′N, 54°45′W, 10 Aug. 1993 (fl, fr), *J.J. de Granville et al.* 12225 (CAY, US).

<figure><img src="image_77.png" /><figcaption>**Figure 73.** *Sipanea wilson-brownei*. Inflorescence with flowers in anthesis, flowers buds, and immature fruits. Photo taken by Scott Mori at Pic Matécho, central French Guiana, on 8 September 2000 (reproduced with permission from the author), gathering Mori & Smith 25056.</figcaption></figure>
**FRENCH GUIANA:** Bassin de l'Approuague, Savane Roche (Inselberg) Virginie, lisière de savane roche, 4°11'N, 52°9'W, 50 m, 12 Feb. 1991 (fl), G. Cremers & P. Petronelli 11857 (CAY, P, U, US); Montagnes de la Trinité, Bassin de la Mana, 4°35'N, 53°21'W, 440 m, 17 Mar. 1992 (fl, fr), G. Cremers 12781 (CAY, P, U, US); Région des Emerillons, Mont Bakra, foret primaire, 3°18'N, 52°57'W, 550 m, 14 Apr. 1993 (fl, fr), G. Cremers 13075 (CAY, B, MO, NY, P, U, US); Crique Jean-Pierre, Bassin de l'Approuague, 4°17'N, 52°12'W, 75 m, 5 Dec. 1994 (fl, fr), G. Cremers et al. 13537 (BR, CAY, MO, NY, P, US); along rd between Cayenne and Regina, pk 93, 4°18'N, 52°10'W, 21 Feb. 1993 (fl, fr), T.B. Croat 74314 (CAY, MO); Réserve Naturelle des Nouragues, foret basse sur inselberg, 4°5'30"N, 52°42'W, 300 m, 17 Oct. 2007 (fl, fr), C. Delnatte et al. 1507 (CAY, MO, P); Mun. Saint George de l'Oyapock, site of future road to the bridge French Guiana-Brazil, ecological corridor, ca. 2 km from the Oyapock River, primary disturbed forest, at forest disturbed edge, 3°52'33"N, 51°49'35"W, 50 m, 2 Jul. 2009 (fl), P.G. Delprete 10575 (CAY); Mun. Saint George de l'Oyapock, on the road to future bridge to Brazil, primary forest with canopy 30-35 m tall, on rich lateritic soil, 3°52'33"N, 51°49'35"W, 50 m, 8 Jul. 2009 (fl), P.G. Delprete 10629 (CAY, F, G, L, MO, NY, P); Route National 2, pk 91, near Regina, 5 Apr. 1992 (fl, fr), Deroin, T. 133 (CAY, P); Grégoire, abbatis N. 11, 3 Jun. 1972 (fl, fr), G. Deward 237 (CAY [2 sheets], P); Montagne des Nouragues, Bassin de l'Arataye, foret basse sur inselberg, 4°3"N, 52°42'W, 23 Aug. 1987 (fl, fr), C. Feuillet 4264 (CAY), 25 Aug. 1987 (fl, fr), 4301 (MO, P, US); Station Hydrologique de la Crique Grégoire, vers le saut, Haute Sinnamary, 21 Jan. 1974 (fl), J.J. de Granville B-5104 (CAY, US); Tumuc Humac Range, layon Koulimapopann-Mitaraka (frontier), pk 7.5, grande savane roche (inselberg) sur crête granitique, 500 m, 6 Aug. 1972 (fl, fr), J.J. de Granville 1135 (CAY, P, VEN); trail from Saül to Carbet Maïs, granitic hill 2 km N of Carbet Mitan, 7 Jul. 1979 (fl), J.J. de Granville & G. Cremers 3030 (CAY, NY, P, UB); Pic Matécho, S summit, 30 km NE of Saül, 520 m, 22 Jan. 1980 (fl, fr), J.J. de Granville 3368 (CAY [2 sheets], NY, P, U, UB); Massif des Emerillons, zone centrale, affleurement granitique, 10 Sep. 1980 (fl, fr), J.J. de Granville 3808 (CAY, P, U, UB); Monts Bakra, versant Sud, at 3.5 km W of Pic Coudreau [ca. 3°17'N, 52°55'W], 450 m, 1 Oct. 1980 (fl, fr), J.J. de Granville 4047 (CAY [2 sheets], P, U [2 sheets], UB); granitic inselberg at NW limit of Monts de la Trinité, 4 Aug. 1981 (fr), J.J. de Granville 4746 (CAY, P, UB); Montagnes Trinité, N summit, 350 m, 13 Jan. 1987 (fl, fr), J.J. de Granville et al. 5953 (BR, CAY, G, MG, MO, NY, P, S, U, US); ibid., 400 m, J.J. de Granville et al. 5974 (CAY);

Région des Emerillons, Mont Bakra, Pic Coudreau, foret basse sur inselberg, 3°18'N, 52°57'W, 650 m, 19 Apr. 1993 (fl, fr), J.J. de Granville & G. Cremers 11825 (CAY, MO); Montagnes de la Trinité, S zone, Bassin de la Mana, foret basse sur inselberg, 4°34'N, 53°21'W, 350 m, 14 Jan. 1998 (fl, imm fr), J.J. de Granville & F. Crozier 13588 (B, CAY, K, MO, NY, P, U, US); Pic Matécho, zone sommitale 2, 3°45'N, 53°2'W, 400 m, foret basse sur inselberg, 15 Sep. 2000 (fl, fr), J.J. de Granville et al. 14079 (CAY, MO, NY); Monts Bakra, 1.5 km W of Pic Coudreau, foret basse sur inselberg, 3°18'N, 52°57'W, 520 m, 16 Jun. 2002 (fl, fr), J.J. de Granville et al. 14801 (CAY, MO, P, US); Chemin de fer Gare-Tigre-Saint Elie, de pk 11 à pk 17, foret primaire, 4°54'N, 53°11'W, 20 m, 7 Apr. 1990 (fl, fr), M. Hoff et al. 6536 (CAY, MO, NY, P, U); Montagnes des Nouragues, Bassin de l'Approuague-Arataye, 4°3'N, 52°42'W, 260 m, Oct. 1989 (fl), D. Larpin 697 (CAY); Pic Matécho, ca. 22.5 km NE of Les Eaux Claires, 3°45'N, 53°2'W, 515 m, on rock outcrop, 8 Sep. 2000 (fl, fr), S.A. Mori & N.P. Smith 25056 (CAY, NY); Saül, Roche Bateau trail, ca. 2 km SE of the village, 3°37'N, 53°12'W, 200 m, 15 May 2006 (fl), S.A. Mori et al. 25857 (CAY); Route de l'Est (RN2), près de Regina, vegetation secondaire, 4°18'N, 52°12'W, 17 Sep. 1999 (fl, fr), M.-F. Prévost & D. Barthélémy 3699 (CAY, MO, NY, P, U); Pic Coudreau du Sud, Bassin de la Malani, 2°14'53"N, 54°19'40"W, 320 m, 7 Feb. 2013 (fl), H. Richard & F. Bagadi 703 (CAY, P); Haut Oyapock, Mont Saint Marcel [ca. 2°23'N, 53°1'W], près du sommet, 300-450 m, 21 Mar. 1976 (fl), C. Sastre 4480 (CAY, L, P); Haut Oyapock, Mont Saint Marcel, ilet sablonneux de la Crique Camp Poivre, 300-450 m, 24 Mar. 1976 (fl), C. Sastre 4526 (CAY, NY, P); Fleuve Approuague, Rivière Arataye, Saut Pararé, sommet d'un inselberg, 375 m, 22 Oct. 1978 (fl, fr), C. Sastre 6225 (CAY, NY, P, U, US); Les Nouragues Reserve, Montagnes des Nouragues, Bassin de l'Approugue-Arataye, 4°3'N, 52°42'W, 120 m, 15 Mar. 2002 (fl, fr), E.J. Tepe et al. 561 (CAY).

**BRAZIL. Amapá:** Territoire contesté franco-brésilien, place de Carsevenne [Calçoene? ca. 2°29'N, 50°57'W], terrain aurifére, 1898 (fl), F. Geay s.n. (P P00748134); Upper Jari River, margin of an inserberg, "2°28'N, 54°46'W" [these coordinates are incorrect, because they fall in southern Suriname], 500 m, 24 Aug. 1993 (fl), J.J. de Granville et al. 12437 (CAY, MO, US); margem do Rio Vila Nova, solo umido, 8 Dec. 1976 (fl buds), B.G.S. Ribeiro 1654 (UB); Tumuc-Humac Region, Paloulouiméempeu Mountain [ca. 2°20'N, 54°38'W], E slope, 550 m, 2 Aug. 1972 (fl, fr), C. Sastre 1539 (CAY, IAN, NY, P, U). **Pará:** Mun. Oriximiná, a 200 m da margem esquerda da estrada da Cachoeira Porteira, paralela ao Rio Trombetas, km 14 [ca.
0°58'48"S, 56°59'23"W], Igarapé das Pedras, mata ciliar,
solo humoso, 19 Jun. 1980 (fl), C.A.C. *Ferreira* [as "C.A.
*Cid"] & J. *Ramos* 1074 (INPA, MG, NY [2 sheets], RB,
UB, US); Mun. Oriximiná, BR-163, estrada que dá acesso
à Perimetal Norte, a 14 km de Cachoeira Porteira [ca.
0°58'48"S, 56°59'23"W], mata de terra firme, solo argilo-
so, 9 Aug. 1986 (fl), C.A.C. *Ferreira* [as "C.A. *Cid*"] 7630
(F, INPA, MG, MO, NY, US).

Excluded species

*Sipanea angustifolia* A.Rich. ex DC., Prodr. 4: 414. Sep. 1830, *nom. illeg. pro syn.* under *Virecta multiflora* Sm. (in A.Rees, The Cyclopaedia, vol. 37, Part I. 1817).

(-) *Sipanea angustifolia* A.Rich., Mém. Fam. Rubi-
acées 196. Dec. 1830; reimpr. A.Rich., Mem. Soc.
Hist. Nat. Paris 5: 276. 1834].

Type: ANGOLA. Without locality, s.d., *Collector Unknown s.n.* (lectotype P [P03947412] designated by Verdcourt (1953b: 314).

(≡) *Pentas angustifolia* Verdc., Bull. Jard. Bot. État Bruxelles 23: 312. 1953.

**Notes:** Verdcourt (1953b: 312) published the new combination *Pentas angustifolia* (A. Richard) Verdcourt" citing *Sipanea angustifolia* A. Rich. ex DC., Prodr. IV, p. 414 (1830); Mém. Soc. Hist. Nat., Paris, p. 276 (1834)." However, *Sipanea angustifolia* A. Rich. was published in Candolle's *Prodromus* (Sep. 1830, p. 414) in synonymy under *Virecta multiflora* Sm. Therefore, the combination *Pentas angustifolia* (A. Rich. ex DC.) Verdcourt" is based on an illegitimate name, and the correct authority for this taxon is *P. angustifolia* Verdc. In addition, Verdcourt indicated the type of *Pentas angustifolia* as "Angola: sine loco, no collector's name (T.) (P.)". The citation "(T.) (P.)" means that the type is in the Paris general herbarium. Below the description, he observed that *"Sipanea angustifolia"* has always been misinterpreted and referred to *Virecta* by all authors; yet the excellent type is available at Paris. No one could have looked at it since there is no possible doubt as to its identity." With this statement, Verdcourt designated the lectotype of this name, which is specimen with barcode P03947412 at P.

*Virecta elatior* DC., Prodr. 4: 415. Sep. 1830.

(-) *Sipanea elatior* A. Rich. ex DC., Prodr. 4: 415.
Sep. 1830, *nom illeg. pro syn.* [*Sipanea elatior* A.Rich.

Mém. Fam. Rubiacées 196. Dec. 1830, *nom. superfl.*;
reimpr. Mém. Soc. Hist. Nat. Paris 5: 276. 1834].

Type: ANGOLA. Without locality, s.d., *J.J. da Silva s.n.* (holotype P [barcode 00539269]).

(≡) *Otomeria elatior* (DC.) Verdc. in Bull. Jard. Bot. État Bruxelles 23: 18. 1953.

**Notes:** Verdcourt (1953a) published the new com-
bination *"Otomeria elatior* (A.Rich. ex DC.) Verdcourt"
(Verdcourt 1953a: 18) citing *"Sipanea elatior"* A. Rich.
ex DC., Prodr. IV, p. 415 (1830), *"Virecta elatior"* A. Rich.
ex DC., loc. cit., *"Pentas elatior"* Benth. ex Walp., Repert.,
IV, p. 57 (1846)*" along with two heterotypic synonyms.
However, *"Sipanea elatior"* A. Rich.*" was published in
Candolle's *Prodromus* (Sep. 1830) in synonymy under
*"Virecta elatior"* DC. (Sep. 1830: 415), and is an ille-
gitimate superfluous name. Therefore, the authority of
*"Otomeria elatior"* (A.Rich. ex DC.) Verdcourt (1953a:
18)*" is erroneous, and the correct authority for this tax-
on is *O. elatior* (DC.) Verdc. Verdcourt (1953a: 22) cited
the type as "Angola: s.l., unknown collector (T.) (P.)."
Below the description, he explained "This extremely var-
iable plant has long been known by Hiern's name, but is
so certainly the long misunderstood *Sipanea elatior* of
Richard that a name change is unavoidable. An excel-
lent specimen which I have accepted as the type exists in
the herbarium at Paris. It was collected by an unknown
traveler in Angola, and formed part of a collection that
was seized from Lisbon by Napoleon during the inva-
sion of Portugal. The collection was carried off to Paris
without notes of any kind." Later, Verdcourt (1976: 214)
cited the type of *Virecta elatior* as "Angola, da Silva [s.n.]
(P, holo!)." Most likely, the collector cited as "da Silva"
was Joaquim José da Silva, who collected in Angola
before 1804 (Hettie Vegter 1986). The P sheet, barcode
P 00539269, has a label with the handwriting (author
unknown) "Herb. Mus. Par. - Sipanea elatior A. Rich. -
Potius Virecta elatior DC., Prodr. IV, p. 415. - Angola."
Just above this label, there is a label reporting "Holo-
typus vel Lectotypus, *Otomeria elatior* (A. Rich. ex DC.)
Verdcourt, Determinavit N. Hallé, 30 mars 1965." This
specimen is the holotype *Virecta elatior* DC.

*Sipanea carnea* Jacques, Rev. Hort., sér. 2, vol. 2 ["vol. 5 (26)"] : 302. May 1843.

*Sipanea carnea* Neumann, *nom. illeg. superfl.*, in Rev. Hort., sér. 2, vol. 2 ["vol. 5 (32)"] : 445, fig. 15. Nov. 1843
(=) *Pentas carnea* (Jacques) Benth., Curtis's Bot. Mag. 17: tab. 4086. 1844. – *Virecta carnea* (Jacques) Baill., Hist. Pl. 7: 331. 1880. Type: Locality of original collection unknown, "Herbier de Jacques," cultivated, s.d., Jacques s.n. (*Lectotype*, P [barcode P04959285], here designated).

(=) *Pentas lanceolata* (Forssk.) Deflers, Voy. Yemen 142. 1889.

**Notes:** Jacques (May 1843: 302-303) in the introduction "*Plantes figurées dans ce numéro*" [Plants illustrated in this issue] of *Revue Horticole*, sér. 2, vol. 2, fascicle 26, described *Sipanea carnea* Jacques as a plant originally either from Madagascar or South America, sent to the *Jardin des Plantes* (Paris) in 1841 by Mr. Pervillé. Jacques was referring to the same plant described and illustrated in the same volume by Neumann, in a fascicle published six months later. Jacques described the species as an herb 30 cm tall, with ovate to lanceolate leaves, terminal multiflorous corymbs, and corolla pale pinkish-violet, with narrowly cylindrical tube. He did not mention any collection or herbarium of deposit. According to Stafleu & Cowan (1979: 404), Jacques might have had "Some material (cult. Plants from Neuilly garden) in Webb herbarium at FI." However, after exhaustive searches at FI and FI-Webb, no original material of *S. carnea* was found. In P there are four specimens labeled *S. carnea*:

1) Specimen with barcode P04959285 has a label with the heading "MUSEUM NATIONAL D'HISTOIRE NATURELLE - CULTURE" with the handwritten text "Sipanea carnea Nem. - Herbier de Jacques." At the lower left corner is a label (author unknown) with the handwritten annotation "Sipanea carnea." The specimen is a branch with numerous leaves and inflorescences with flowers in anthesis. This specimen is original material and is here designated the lectotype of *Sipanea carnea*.

2) Specimen with barcode P04959286 has a label with the heading "HERB. MUS. PARIS" and the printed text "herbier d'Édouard Spach, donné au Muséum par sa famille, en jullet 1879." At the lower left corner is a label (author unknown) with the handwritten annotation "Sipanea carnea det. Brongn. (Pentas carnea Lindl) S. Par 1841 seminibus borbonides." The specimen is composed of two branches with numerous leaves and terminal inflorescences. This specimen is not original material of *Sipanea carnea*.

3) Specimen with barcode P04959287 has a label with the heading "HERB. MUS. PARIS" and the printed text "Herbier donné par M. Houllet, Jardinier en chef des Serres. (1869. N° 1.)" At the lower left corner is a label (author unknown) with the handwritten annotation "Sipanea carnea Ad. Brong. - Pentas carnea Bot. veg. 1844., S. ch., en herb. Houllun, Ile Bourbon. Juillet 1844, hort. Paris, Pervillé." The specimen is composed of two branches with numerous leaves and terminal inflorescences. This specimen is not original material of *Sipanea carnea*.

4) Specimen with barcode P04959288 has a label with the heading "HERB. MUS. PARIS" and the printed text "Herbier d'Adrien de Jussieu, donné au Muséum par ses enfants, en 1857." At the lower left corner is a label (author unknown) with the handwritten annotation "Sipanea carnea. ad. Br. - Pentas carnea Lindl. H.P. (Serr. Ch.) Mars 1846." The specimen is composed of two branches with numerous leaves and terminal inflorescences. This specimen is not original material of *Sipanea carnea*.

In the fascicle 32 of the same volume where Jacques (May 1843: 302-303) first published *Sipanea carnea*, J.H.F. Neumann (Nov. 1843: 445), six month later, described and illustrated *S. carnea*. He described the species as a herbaceous plant from South America, with corymbose inflorescence, corolla tube four times longer than the calyx, with (4)5 lobes, filiform style longer than the corolla, and pale violet corollas. The beautiful illustration, drawn by the same author, corresponds entirely with the species description. Neumann did not indicate a collection or herbarium, and, according to Stafleu & Cowan (1981: 731), he did not keep a personal herbarium.

The following year, Bentham (1844: tab. 4086) published the new genus *Pentas* Benth., with the species *P. carnea* Benth. and *P. parviflora* Benth. Along with the description of *P. carnea*, he wrote "Our stoves have lately exhibited a fine flowering plant, under the name of *Sipanea carnea*, well worthy of general cultivation. Mr. Makoy, of Liege, I believe, introduced it to England: at least, the Royal Botanic Garden of Kew is indebted for the possession of it to that liberal cultivator. A slight examination of the flowers, however, show that it has nothing to do with *Sipanea*, and that it is one of the Hedyotideae;-a new genus, to which Mr. Bentham has given the name of *Pentas* [...] although not so stated, there is every reason to believe the plant is from tropical Africa". Bentham (1844: tab. 4086) cited *S. carnea* in synonymy under *P. carnea*. Bentham's citation is an indirect reference to *S. carnea* Jacques, and it should be treated as the publication of the new combination *Pentas carnea* (Jacques) Benth., which has the same type of *S. carnea*.
## 9. SIPANEOPSIS

*Sipaneopsis* Steyerm., Mem. New York Bot. Gard. 17: 284-289. 1967

Steyerm. in Lasser & Steyermark, Fl. Venezuela 9: 272-282. 1974; J.H. Kirkbr., Acta Amazonica 10: 113-115. 1980; Delprete & Steyerm. in Steyermark et al., Fl. Venezuela. Guayana 8: 833-835, figs. 648-650. 2004.

Type: *Sipaneopsis rupicola* (K.Schum.) Steyerm.

Subshrubs or shrubs, rarely herbs, erect, ascending or rarely decumbent. *Stipules* persistent, free or adnate to petioles, sometimes basally sheathing, free portion broadly to narrowly triangular, or base broadly triangular and distal portion linear, acuminate, bilobed or bifid at apex, or topped by 2-7 linear lobes (*S. morichensis*, *S. rupicola*), with (1)2-11 colleters at medio-basal portion or just below the margin inside, tips exserted. *Leaves* opposite, short-petiolate; blades ovate, elliptic, lanceolate or linear, chartaceous or subcoriaceous; domatia absent. *Inflorescence* terminal, cymose, few- to many-flowered. *Flowers* homostylous, 5-merous or very rare 6-merous. *Hypanthium* subglobose, globose, broadly turbinate, turbinate, subcampanulate-urceolate, broadly-obovoid, obovoid-turbinate, obconical or oblong-obovoid, scabrous, densely hispid, densely hispidulous strigillose, densely to sparsely strigose, sericeous-strigose or sericeous. *Calyx* persistent; lobes free or shallowly connate at base, small, ovate, narrowly ovate, narrowly lanceolate to linear-lanceolate. *Calycophylls* absent. *Corolla* campanulate at initial and intermediate stages of flowering with tube short and lobes already opened and erect, at ultimate stage of anthesis the corolla becoming hypocrateriform with tube completely elongate and lobes reflexed and perpendicular to the tube; corolla pink during anthesis and turning pinkish-white to white at final stage of anthesis; tube externally hirsute, internally pubescent, without a pubescent ring, with 5 triangular appendages at the base of the corolla lobes, the appendages fleshy, covered by dense tufts of strigose hairs 0.3-0.4 mm long; lobes left-contorted or rarely imbricate, obovate, ovate or round, margin entire, obtuse or round at apex. *Stamens* included, filaments extremely short, attached at the middle of the corolla tube; anthers subsessile, linear, acute at both ends, dorsifixed near the base. *Pollen* 4(5)-colporate; exine shallowly foveolate, perforate. *Ovary* with a stalked placenta; stalk inserted at the middle of the septum, terminating with a globose extension; ovules inserted on the globose extension. *Style* included or with branch tips exserted, filiform, glabrous; branches lanceolate to narily lanceolate.

*Fruit* indehiscent, globose, subglobose or urceolate, thinly woody to crustaceous, crowned by the permanent calyx lobes, falling off as a dispersal unit, leaving a round scar on the infructescence rachis. *Seeds* minute, horizontal, spherical to ellipsoidal.

**Flowering modality:** The flowering process of *Sipaneopsis* is unique within the Rubiaceae, and it can be appreciated in Figures 80 and 82. It was described in detail by Delprete (2015a, 2017). An inflorescence with flowers in different developmental stages of *Sipaneopsis maguirei* is shown in Figure 80B-C. During the initial stage of anthesis the flowers are still very small; however, their corolla lobes are already open and erect, although not fully expanded, before tube elongation (Figure 82A-B); at this stage the triangular appendages present at the base of the corolla lobes are appressed against each other, forming a convex structure at the corolla mouth (Figure 82A-B), and obstructing the entrance of any visitors or pollinators. During successive stages of anthesis, the corolla tube and lobes elongate; the corolla lobes remain erect and the corolla lobe appendages are still appressed against each other (Figure 82C-F), continuing to obstruct the entrance of any visitors or pollinators. At the final stage of anthesis, the corolla lobes reflex outwards and become perpendicular to the tube (corolla hypocrateriform), the corolla lobe basal appendages become erect, not touching each other anymore (Fig. 82G-H), allowing visitors and pollinators to access the corolla tube and nectar disk. In addition, flowers of *Sipaneopsis* are protandrous, which means that after the lobes have become perpendicular to the corolla tube, on the first day the anthers reach full maturity and dehisce, releasing pollen, while the stigmatic branches are still appressed to each other and not receptive (Figure 82G). The following day the anthers are no longer releasing pollen, the stigmatic branches spread apart and are reflexed, and the stigmatic surface become receptive. For the above reasons, for species descriptions the corolla tube and lobes of *Sipaneopsis* species should be measured only when they are fully expanded and the lobes are perpendicular to the tube.

**Distribution and ecology:** A genus with eight species endemic to white-sand savannas, occurring mostly in the lowlands, although with a few collections from higher altitudes, of the Guayana Shield, ranging from Colombia to southern Venezuela and the Amazon basin of Brazil.

**Stipules variation within the same species:** Stipules are polymorphic in several species of *Sipaneopsis*, as they
can be entire, bifid or multifid in the same species. Also, in some species the stipules are entire at distal nodes and sometimes apically splitting with age, remaining entire or becoming bifid or multifid in older nodes of the same branch. In *S. cururuensis* they are consistently bifid with narrowly triangular lobes. In *S. duckei*, *S. maguirei* and *S. pacimoniensis* they are consistently entire and broadly to narrowly triangular. In *S. foldatsii* and *S. huberi* they are broadly to narrowly triangular, and rarely (*S. foldatsii*) or consistently (*S. huberi*) bifid at apex. In *S. rupicola* they are triangular, entire and caudate, or triangular at base with 2-3 linear lobes, or basal portion extremely reduced and topped by 6-7 linear lobes or by 2 longer lobes intercalated with shorter ones.

Key to *Sipaneopsis* species

Stipule shape and vestiture of vegetative parts are important characters for species identification in *Sipaneopsis*. As in several species stipules are polymorphic, a few species are present in multiple entries in the key below. Corolla measurements were made only when fully expanded, with the corolla lobes perpendicular to the tube, and the corolla lobe basal appendages erect, no longer touching each other.

1. Herbs, 10-12 cm tall, single-stemmed (Venezuela: Amazonas, Río Pacimoni)................................................ 7. *S. pacimoniensis*

1. Subshrubs or shrubs, (8-)18-200 cm tall, many-stemmed or branched, or ranging from herb to subshrub in *S. cururuensis*..................................................................................2

2. Stipules with 2 narrowly triangular lobes (i.e., stipules bifid) or with (1-)2-7 linear lobes..................................................3

2. Stipules entire ..................................................................................................................9

3. Stipules with (1-)2-7 linear lobes..................................................................................4

3. Stipules with 2 narrowly triangular lobes (i.e., stipules bifid ..................................................................................5

4. Stipules ochroleucous-pubescent outside, with 1-4 linear lobes (Venezuela: Amazonas: Cerro Moriche..................................................6. *S. morichensis*

4. Stipules densely white-strigulose outside, with 2-3 linear lobes, or base reduced to a transversal line with 6-7 linear lobes (Colombia: Vaupés: Cerro Chiribiquete; Caquetá: Araracuara; Venezuela: Amazonas: Casiquiare53 ..................................................8. *S. rupicola*

5. Leaf blade sericeous below; corollas 13-14 mm long, with tubes 9.5-10 mm long and lobes 4-4.5 mm long when fully expanded (Venezuela: Amazonas, Dpto. Río Negro)..................................................4. *S. huberi*

5. Leaf blade glabrous, scabrous or strigose below; corollas 14.5-22 mm long, with tubes 11-16 mm long and lobes 3.5-7 mm long when fully expanded ..................................................6

6. Young stems strigulose; leaf blade glabrous or sparsely to densely strigose, or densely sericeous-strigose with midvein and secondary veins white- or ochroleucous-strigose below; corollas 14.5-16.2 mm long, with tubes 11-11.5 mm long and lobes 3.5-4.7 mm long when fully expanded (Colombia: Vaupés: Cerro Chiribiquete; Caquetá: Araracuara; Venezuela: Amazonas: Casiquiare) ..................................................8. *S. rupicola*

6. Young stems glabrous, sparsely to densely strigose, or sparsely to densely scabrous; leaf blade glabrous, sparsely to densely strigose, or sparsely to densely scabrous below; corollas 17.5-22 mm long, with tubes 12-16 mm long and lobes 4.2-7 mm long when fully expanded ..................................................7

7. Young stems and leaf lamina sparsely to densely scabrous below; stipules always bifid; stamens inserted at 9-9.5 mm below corolla mouth, filaments 3.5-4 mm long (Brazil: Pará, Tapajos River Basin) ..................................................1. *S. cururuensis*

7. Young stems and leaf lamina glabrous or sparsely to densely strigose or densely hispid-strigose below; stipules entire or rarely bifid at older nodes; stamens inserted 2-3.5 mm below corolla mouth, filaments 0.3-0.4 mm long (anthers subsessile)..................................................................................8

8. Leaf lamina glabrous and midrib appressed-strigose above, glabrous or sparsely strigose and midrib and secondary veins appressed strigose below; corollas 17.5-22 mm long, with tubes 12-16 mm long and lobes 4.5-7 mm long when fully expanded; fruits globose, 4-5 x 4-5 mm, appressed strigose (Brazil: Pará, Campos do Ariramba, and Amazonas, Serra do Aracá) ..................................................................................2. *S. duckei*

8. Leaf lamina densely hispid-strigose above and below; corollas 20.5-24 mm long, with tubes 14-16 mm long and lobes 6.5-8 mm long when fully expanded; fruits urceolate, 3.5 x 3 mm, densely sericeous (Venezuela: Amazonas, Río Atabapo)..................................................................................3. *S. foldatsii*

9. Corollas 14.5-16.2 mm long, with tubes 11-11.5 mm long and lobes 3.5-4.7 mm long when fully expanded (Colombia: Vaupés: Cerro Chiribiquete; Caquetá: Araracuara; Venezuela: Amazonas: Casiquiare) ..................................................8. *S. rupicola*

9. Corollas 17.5-30 mm long, with tubes 12-22 mm long and lobes 4.5-9 mm long when fully expanded ..................................................10

10. Stems thin; inflorescences 2-8(-13)-flowered (Venezuela: Amazonas, Dptos. Atabapo, Atures, Casiquiare; Brazil: Amazonas, Rondônia) ..................................................................................5. *S. maguirei*

10. Stems stout; inflorescences (12-)19-58-flowered ..................................................11

11. Leaf lamina glabrous and midrib appressed-strigose above, glabrous or sparsely strigose and midrib and secondary veins appressed strigose below; corollas 17.5-22 mm long, with lobes 4.5-7 mm long when fully expanded; fruits globose, 4-5 x 4-5 mm, appressed strigose (Brazil: Pará,
Campos do Ariramba, and Amazonas, Serra do Aracá)......
.................................................................................. 2. *S. duckei*

11. Leaf lamina densely hispid-strigose above and below; There is absolutely no contrast in this character.; corollas 20.5–24 mm long, with lobes 6.5–8 mm long when fully expanded; fruits urceolate, 3.5 x 3 mm, densely sericeous (Venezuela: Amazonas, Rio Atabapo).................................... 3. *S. foldatsii*

9-1. *Sipaneopsis cururuensis* J.H. Kirkbr., Acta Amazon. 10: 113, figs. 19, 24. 1980. (Figures 74-75).

Type: BRAZIL. Pará: Upper Tapajós River Basin, Cururu River, Erereri [River, ca. 7°37'S, 57°37'W], 8 Feb. 1960 (fl, fr), W.A. Egler & Raimundo [R.R. Silva] 1265 (MG 24312) (holotype IAN [barcode and Acc. No. 123604]; isotypes MG [barcode unknown, Acc. No. 24312]; isotype fragment US[barcode 00074187]; photo-IAN at NY, UB, S).

Herbs or subshrubs 25-50 cm tall, basal portion sometimes prostrate, rooting at basal nodes; stems scabrous to densely scabrous; hairs antrorse, white or pale brown. *Stipules* free at base, bifid, the lobes narrowly triangular, acute at apex, 2-5 × 0.5-1 mm, scabrous to densely scabrous outside, sparsely scabrous and with a few colleters at base inside. *Leaves* petiolate; petioles 2-3 mm long, scabrous to densely scabrous; blades elliptic to narrowly elliptic, 4-7 × 1.5-2.5 cm, attenuate at base, acute at apex, papyraceous, dark green above, pale green below, drying brown above and pale brown below, sparsely scabrous above, scabrous to densely scabrous below; with 7-10 secondary veins each side. *Inflorescences* cymose, a compound dichasium with scorpiod terminal branches, 23-52-flowered, 2.5-5-(13) cm tall, (including peduncle and flowers), 3-4.5 cm broad, rachis scabrous to densely scabrous; peduncle 0.5-3(-6.5) cm long; with 9-14 terminal branchlets; branchlets secundiflorous, scorpioid, 7-14-flowered. *Flowers* with pedicels 0.4-0.6 mm long, scabrous to densely scabrous, subtended by one bracteole narrowly triangular to oblong-lanceolate, 2.2-5.8 × 1-2.4 mm, acute at apex, scabrous. *Hypanthium* globose, ca. 1 mm in diam., scabrous. *Calyx* lobes subequal, narrowly ovate, 2-2.3 × 0.5-0.7 mm, sparsely strigose outside, glabrous inside, acute at apex; with 1-2 colleters in each calycine sinus, narrowly oblong, 0.2 mm long. *Corolla* hypocrateriform, 18.2-20 mm long when fully expanded, tube cylindrical and slightly wider at mouth, 14-15.4 mm long when fully elongated, 0.8 mm wide at base, 1.2 mm wide at mouth, strigose outside, glabrous at base and sericeous at upper 3/4 inside; lobes irregularly imbricate in bud,

1-3 lobes interior, elliptic to oblong-ovate, 4.2-4.6 × 1.8-2 mm when fully expanded (perpendicular to the tube), broadly acute at apex, antrorse-strigose outside, glabrous inside. *Stamens* included, inserted at 9-9.5 below corolla mouth; filaments 3.5-4 mm long; anthers narrowly oblong, 1.6-1.7 mm long, acute at both ends. *Style* included, ca. 13 mm long, glabrous; style branches lanceolate, ca. 1 mm long. *Infructescences* a compound dichasium, 5-14.5 cm long, terminal branches scorpiod, 3-7 cm long; rachis scabrous to densely scabrous; peduncle 0.5-3(-6.5) cm long; fruits subtended by the persistent bracteoles. *Fruit* globose, 3-4 mm in diam., strigose to sparsely strigose. *Seeds* globose, 0.5-0.8 mm in diam.; testa reticulate-foveolate, cells polygonal.

**Notes:** In agreement with Kirkbride's (1980) discussion, *Sipaneopsis cururuensis* J.H.Kirkbr. is similar to *S. rupicola* (K. Schum.) Steyerm. and *S. morichensis* Steyerm. because of the lobed stipules, shorter corolla tubes and taller habit with respect to the other species of the genus, from which it could be distinguished by the scabrous leaves and stems (vs. glabrous in *S. rupicola* or villosulous in *S. morichensis*), and corolla lobes 4.2-4.6 mm long (vs. 3-3.5 mm long in *S. rupicola* or 4.5-6 mm long in *S. morichensis*).

*Sipaneopsis cururuensis* is also similar to *S. duckei* Delprete in being herbs or subshrubs with stout stems, leaves with 6-10 secondary veins on each side of midrib, inflorescences compound dichasia, corolla tube length (14-15.4 mm long in *S. cururuensis* and 12-16 mm long in *S. duckei*), and globose fruits. *Sipaneopsis cururuensis* differs from *S. duckei* in the young stems scabrous to densely scabrous (vs. hispid-strigose or appressed to spreading strigose in *S. duckei*), stipules consistently bifid (vs. entire or bifid), leaf blades sparsely scabrous above (vs. completely glabrous, or lamina glabrous and midrib appressed strigose, or lamina and midrib sparsely strigose above in *S. duckei*) and scabrous to densely scabrous below (vs. completely glabrous, or sparsely strigose and midrib and secondary veins sparsely to densely appressed strigose below), hypanthium globose, ca. 1 × 1 mm, scabrous (vs. globose, broadly obovoid to obovoid-turbinate, 1-5 × 0.8-1.5 mm, densely to sparsely strigose in *S. duckei*), corolla lobes elliptic to oblong-ovate, 4.2-4.6 × 1.8-2 mm when fully expanded (vs. roundish, obovate to elliptic, 4.5-7 × 2-4 mm when fully expanded in *S. duckei*), stamens inserted 9-9.5 mm below corolla mouth, filaments 3.5-4 mm long (vs. inserted 2-3 mm below corolla mouth, anthers subsessile in *S. duckei*), fruits 3-4 × 3-4 mm (vs. 4-5 × 4-5 mm in *S. duckei*), and seeds globose, 0.5-0.8 × 0.5-0.8 mm (vs. irregularly ellipsoid to subglobose, 0.9-1.1 × 0.8-1 mm).
<figure><img src="image_78.png" /><figcaption>**Figure 74.** *Sipaneopsis cururuensis*. A. Branch with several inflorescences and infructescences (fruits fallen off). B. Node with stipule and basal portion of petioles. C. Flower at intermediate stage of anthesis. D. Flower at final stage of anthesis, with corolla lobes reflexed, and receptive stigmas. A–D: drawn from *Egler & Raimundo* 1265 (MG, isotype). Illustration by Piero Delprete.</figcaption></figure>
A few specimens from Guyana, i.e., Henkel *et al.* 1449, McDowell & Stobey 3853, and Mutchnick *et al.* 156, have been incorrectly identified as *Sipaneopsis cururuensis*; however, *S. cururuensis* is known only from the type collection from the Tapajos River Basin, Brazilian state of Pará, south of the Amazon River, on the Brazilian Shield. Those specimens are *Neobertiera pakaraimensis* Delprete (2015: 126) instead. *Sipaneopsis* and *Neobertiera* are similar by having indehiscent fruits falling off as dispersal units; the former differ from the latter by the homostylous flowers, and the corolla lobes that open before tube elongation and possess basal appendage covered with a tuft of hairs; while in *Neobertiera* the flowers are distylous, and the corolla lobes open only after complete tube elongation and lack basal appendages.

**Distribution and ecology:** A species only known by the type locality, in the Brazilian state of Pará, South of the Amazon River, along the Cururu River, in the upper Tapajos River basin (Figure 75).

**Phenology:** The type gathering, with flowers and fruits, was collected in February.

**Suggested conservation status:** Critically Endangered (CR). An extremely rare species only known from the type gathering, collected in 1960 near the Erereti River, ca. 7°37'S, 57°37'W, a small affluent of the Cururu River, Upper Tapajós River Basin, state of Pará, Brazil. This remote locality was botanically explored only once and remains poorly collected. The size of the population of this species is unknown and suspected to be quite small. Because it is known from a single locality, and most likely a small population, and the high possibility of local deforestation, it is treated as Critically Endangered (CR) following IUCN criteria (IUCN 2012, 2019).

9-2. *Sipaneopsis duckei* Delprete, Phytotaxa 282(1): 130, fig. 2. 2018. (Figures 5I-J, 75-76).

Type: BRAZIL. Pará: Mun. Oriximiná, Campos do Ariramba, campina inundavel do Rio Jaramacarú, solo arenito, 90 m, 8 Jun. 1980 (fl, imm fr), G. Martinelli 6869 (RB 202995) (holotype MG [Acc. No. 92530]; isotypes INPA [Acc. No. 101497], NY [2 sheets, barcodes 02693457 and 01552866], RB

<figure><img src="image_79.png" /><figcaption>**Figure 75.** Distribution of *Sipaneopsis cururuensis* (star), *S. duckei* (circles), *S. foldatsii* (triangle), *S. huberi* (asterisks), *S. morichensis* (square), and *S. pacimoniensis* (inverted triangle).</figcaption></figure>
[Acc. No. 202995, barcode 00364126], UB [Acc. No. unknown]).

Subshrubs to shrubs 0.3-1.6 m tall, erect, ascending or rarely decumbent, few- to many-branched; stems stout, woody; young branches sparsely hispid-strigose or appressed to spreading strigose; hairs 0.3-0.6 mm long. Stipules free at base or adnate to petioles and basally sheathing, free portion narrowly triangular to narrowly oblong-triangular, or base broadly triangular and distal portion linear to acicular, (2.5-)3-6.5 × 1.5-3.5 mm, or very rarely bifid at apex in older stipules, probably originating from a young entire stipule, the lobes (when present) narrowly triangular to linear, sometimes with a terminal colletter, hispid-strigose to spreading-strigose outside, spreading pubescent at base and glabrous distally or sericeous, with 4-7 colleters inserted just below the margin inside, tips exerted (populations in Serra Aracá), or sparsely pubescent at base and glabrous above or completely sericeous, with (1)4-7 colleters at medio-basal portion inside (populations of Campos do Ariramba). Leaves subsessile to petiolate; petioles to 5.5 mm long, hispid-strigose or appressed to spreading-strigose (hairs 0.3-0.5 mm long); blades ovate, narrowly ovate, elliptic-lanceolate, narrowly elliptic to narrowly oblong-elliptic, (2.5-)3.5-8.5 × (0.6-)1-3.5 cm, round, obtuse to attenuate at base, acute at apex, subcoriaceous, margins planar or narrowly revolute, dark green above and pale green below when fresh, drying dark brown above and pale brown below, lamina glabrous and midrib appressed strigose above or lamina and midrib sparsely strigose/scabrous above, lamina glabrous or sparsely strigose and primary and secondary veins sparsely to densely appressed strigose below; secondary veins 6-10 each side of midrib, strongly depressed above, prominent below. Inflorescences congested-cymose, branched portion reduced, dichotomously branched, (12-)19-58-flowered, 2.2-10 cm long (including peduncle and flowers), 1-3 cm broad (expanding after anthesis), sometimes frondose, i.e. subtended by leaf-like bracts (i.e., pherophylls), these (when present) narrowly ovate to lanceolate, 1-3.5 × 0.2-1 cm, acute at apex, lamina sparsely strigose and primary and secondary veins densely appressed-strigose below, margins strigose-ciliate; peduncle 0.7-7.5 cm long, densely strigose; bracts subtending secondary branches lanceolate, narrowly elliptic to narrowly oblong-elliptic, (1-)2-4.5 × (0.3-)0.5-1.5 mm, attenuate at base, acute at apex; terminal branchlets scorpiod, 0.5-15 mm long, elongating after anthesis, densely strigose. Flowers sessile to short-pedicellate, secundiflorous, central flower sessile to sub-pedicellate, with pedicels (when present) to 1.3 mm long, lateral flowers sessile; bracteoles 2 per flower, 1

opposing and the other subtending each flower, the one subtending the flower smaller than the one opposing the same flower, sessile, lanceolate to narrowly lanceolate, decreasing in size towards branch tip, 1.5-4.5 × 0.5-1.1, acute at apex, sparsely strigose, margins ciliate-strigose. Hypanthium globose, broadly-obovoid to obovoid-turbinate, 1-1.5 × 0.8-1.5 mm, densely to sparsely strigose. Calyx lobes equal or subequal, ovate, lanceolate to narrowly lanceolate, 1.5-3 × 0.3-0.7 mm, acute at apex, strigose outside, margins strigose-ciliate, glabrous inside; with 2 colleters in each calycine sinus, ellipsoid, acute at apex, 0.25-0.3 mm long. Corolla hypocrateriform, pink, 17.5-22 mm long when fully elongated; tube cylindrical, slightly wider at mouth, 12-16 mm long when fully elongated, 0.5-0.7 mm wide at base, 1.6-2 mm wide at mouth, sparsely appressed-strigose and distally more densely so outside, glabrous at basal 3-3.5 mm and sparsely pubescent at upper portion inside; lobes left-contorted, roundish, obovate to elliptic, 4.5-7 × 2-4 mm when fully expanded (perpendicular to the tube), round, obtuse to acute at apex, sparsely appressed strigose outside, glabrous and papillose inside. Stamens included, with anthers tips ca. 1-1.3 mm below corolla mouth or about the same length as the tube, inserted 2-3 mm below corolla mouth; filaments 0.3-0.4 mm long; anthers subsessile, oblong-elliptic, 1.5-2.1 mm long, acute at both ends. Style included, slightly shorter than or about the same length as the tube, 12-14 mm long when fully elongated, glabrous; style branches narrowly lanceolate, ca. 1 mm long. Infructescences a compound dichasium, 6.5-8.5 cm long, terminal branches scorpiod, 1.5-3 cm long. Fruit globose, 4-5 mm in diam., appressed strigose, larger than in flowers, 3-4.5 mm long. Seeds 6-8 per locule, irregularly ellipsoid to subglobose, 0.9-1.1 × 0.8-1 mm; testa rugose-foveolate, pale brown.

**Notes:** *Sipaneopsis duckei* is similar to *S. maguirei* in several aspects, e.g., stipules entire (not lobed), inflorescences congested-cymose, style included (slightly shorter than or about the same length as the tube), and seeds 6-8 per locule. The former differs from the latter in having stems stout, erect (vs. thin, reclining, decumbent or ascending in *S. maguirei*); stipules (2.5-)3-6.5 × 1.5-3.5 mm (vs. 1.5-3.5 × 1-2 mm); inflorescences 19-58-flowered (vs. 2-8(-13)-flowered); corolla 17.5-20 mm long and tube 12-14 mm long when fully elongated (vs. corolla 21-30 mm and tube 15-22 mm long when fully elongated); and fruits 4-5 × 4-5 mm (vs. 3.5-4 × 3.5-4 mm).

*Sipaneopsis duckei* is also similar to *S. rupicola* (K. Schum.) Steyerm. by being a subshrub to shrub with stout, strigose to strigulose branches, subcoriaceous leaves, and inflorescences 19-58-flowered. The former
differs from the later by the stipules entire or rarely bilobed at older nodes (lobes narrowly triangular to linear, sometimes with a terminal colleter), base broadly triangular and distal portion linear, or narrowly triangular to narrowly oblong-triangular, (2.5–)3–6.5 mm long, hispid-strigose to spreading-strigose outside; whereas, in *S. rupicola* the stipules are triangular, 4–5 mm long, entire and caudate, or triangular at base, with 2 or 3 acute lobes, or the base reduced to a line with 6 or 7 linear lobes, densely strigulose outside. In *Sipaneopsis duckei* the corollas are 17.5–22 mm long when fully expanded, tubes are 12–16 mm long when fully expanded, sparsely appressed-strigose and distally more densely so outside, glabrous at basal 3–3.5 mm and sparsely puberulous at upper portion inside; lobes are broadly obovate, (almost round), obovate to elliptic, 4.5–7 × 2–4 mm when fully expanded (perpendicular to the tube), round, obtuse to acute at apex, appressed strigose outside, papillose inside. In *S. rupicola* the corollas are 16–16.2 mm long when fully expanded; tubes are 11.5 mm long when fully expanded, sericeous-strigose (densely so near the mouth) outside, glabrous at basal 3.5–4 mm and puberulent above and papillose near the mouth inside; the lobes are left-contorted or irregularly imbricate (1 or 2 external), subequal, oblong-obovate, 4.5–4.7 × 2.2–2.5 mm when fully expanded, obtuse to round at apex, sparsely strigulose outside, glabrous inside.

**Eponimy:** The specific epithet of this taxon is dedicated to Adopho Ducke (1876–1959), a major botanist and entomologist who collected for about 50 years in the Amazon basin of Brazil, often in remote places. This epithet is dedicated to him because he was the first botanist to make botanical collections in Campos do Ariramba, northern portion of the state of Pará, Brazil, where he made the first collection of this species. He described his difficult expedition to reach this locality that took place during 7–27 December 1906, going upstream on the Rio Trombetas, and then its affluent Rio Jaramacarú, followed by several days on foot on difficult terrain, in an article published the following year (Ducke, 1907). He collected *Sipaneopsis duckei* Delprete (2018: 130) two days before reaching the open fields of Campos do Ariramba, in an area of *campinarana*, on 20 December 1906 (A. Ducke s.n. (MG 8020), MG, R, RB). The gathering

*Martinelli 6869* (RB 202995), collected in the same locality, is designated the type of this taxon because it has duplicates deposited in several herbaria (MG, NY [2 sheets], RB, UB).

**Distribution and ecology:** A species occurring in two populations in Campos do Ariramba, state of Pará, and on Serra do Aracá (tepui formation), state of Amazonas, in the northernmost region of Amazonian Brazil (Figure 75). Endemic to white-sand savannas with sparse trees and shrubs, locally called *campinarana*, sometimes seasonally inundated; most collections were made at 80–100 m altitude, and one collection was made at 1200 m altitude on the plateau of the northern massif of Serra Aracá.

**Phenology:** Specimens with flowers in anthesis were collected in January, February, May, July, October and December. Three gatherings with flowers and fruits were collected in January, June and July.

**Suggested conservation status: Not Threatened (NT).** *Sipaneopsis duckei* is known from numerous collections from two distant areas of Amazonian white-sand savannas, Brazil. The two main areas are Campos do Ariramba, state of Pará, and Serra do Aracá (tepui formation), state of Amazonas, in the northern region of Amazonian Brazil. The extent of occurrence (EOO) is therefore not calculable, and the number of individuals per population is unknown. This species occurs in remote areas, without any imminent threat, as white-sand areas are not exploitable for agriculture or cow pastures because of their soils with very low nutrient content. White-sand areas are commonly subjected by seasonal fires during the dry season, and their vegetation is adapted to a fast regeneration. This species is therefore considered Not Threatened (NT) according to IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** **BRAZIL. Amazonas:** Mun. Barcelos, Parque Estadual da Serra do Aracá, vegetação arbustiva com afloramentos e alagados, com pequena mata na margem de igarapé, 0°55'24"N, 63°25'58"W, 1220 m, 1 Sep. 2011 (fl), R.C. Forzza et al. 6632 (INPA, MG, RB); Mun. Barcelos, Parque Estadual da Serra do Aracá, Serra do Aracá, acampamento do fosso,

**Figure 76.** *Sipaneopsis duckei*. A. Branch with two frondose inflorescences and one infructescence (note last fruit on the infructescence). B. Stipule. C. Detail of an inflorescence. D. Flower at early stage of anthesis. E. Flower at intermediate stage of anthesis. F. Flower at final stage of anthesis, with corolla lobes reflexed, and receptive stigmas. G. Indehiscent fruit, fallen off as dispersal unit. A–B: drawn from *Egler 273* (MG); C, G: drawn from *Rosa & Lira 2268* (MG); D–F: drawn from *Prance et al. 28853* (US). Illustration by Piero Delprete (Reproduced from fig. 2 of Delprete (2018) with permission of copyright holder Magnolia Press).
<figure><img src="image_80.png" /><figcaption>Figure 76.</figcaption></figure>
0°51'47"N, 63°19'59"W, 1020 m, 19 Apr. 2014 (fl, fr), R.C. Forzza et al. 7999 (RB); Rio Aracá, afluente do Rio Negro, campinarana, solo arenoso, 24 Oct. 1952 (fl), R.L. Frões & G. Addison 29039 (IAN); Mun. Barcelos, campina, palmar de Barcella odora y sabana sobre arena blanca, en la ribera izquierda del Río Aracá medio, ca. 16 km al SE (en línea recta) de la boca del Río Januarí, ca. 0°25'N, 63°23'W, 100 m, Jul. 1985 (fl), O. Huber et al. 10782 (INPA, NY); Mun. Barcelos, Parque Estadual da Serra do Aracá, campos de altitude sobre tepui, 0°52'15"N, 63°25'1"W, 1100 m, 1 Nov. 2011 (fl), G. Martinelli et al. 17267 (RB); Mun. Barcelos, Parque Estadual da Serra do Aracá, margem direita do Igarapé Preto, acima da Cachoeira do El Dorado, 1900 m, 17 Aug. 2011 (fl), M.A. Moraes & E. Fernandez 221 (RB); Mun. Barcelos, Serra do Aracá, Platô Sul, campo rochoso, próximo a curso d'água, 0°51'59"N, 63°19'56"W, 1000 m, 4 May 2016 (fl, fr), M.G.C. Nogueira et al. 498 (MG); base of Serra do Aracá, 0-3 km S of Central Massif, 3 km E of Rio Jauari, white-sand savanna, 0°49'N, 63°19'W, 6 Feb. 1984 (fl), G.T. Prance et al. 28853 (INPA, MG, MO, NY, UB, US); plateau of northern massif of Serra Aracá, 0°51-57'N, 63°21-22'W, 1200 m [confirmed by Prance, pers. comm.], 11 Feb. 1984 (fl), G.T. Prance et al. 28939 (F, INPA, MG, MO, NY, RB, UB, US); 3 km S of central massif of Serra Aracá, 0°49'N, 63°17'W [specimen label reports "65°17'W" which is a typographical error], 100 m, savanna on sand, dominated by *Lagenocarpus*, 18 Jul. 1985 (fl), G.T. Prance et al. 29680 (INPA, MG, NY, UFMT, US); N margin of Rio Aracá, near Serrinha, open savanna on sand, sedge dominated, subshrub with prostrate branches, corolla pink, 0°25'N, 63°23'W, 80 m, 25 Jul. 1985 (fl, fr), G.T. Prance et al. 29798 (HFSL, INPA, MG, NY, US); N margin of Rio Aracá, just above Igarapé Sauadaua, savanna on white sand, 0°13'N, 63°8'W, 80 m, 26 Jul. 1985 (fl), G.T. Prance et al. 29798 (NY; this specimen has different locality and coordinates than the other duplicates of Prance et al. 29798); 3 km S da parte central da Serra Aracá, no caminho entre a pista de baixo e a encostas, e a 8 km E do Rio Jauari, solo silicoso ao redor do campamento, campina lenhoso-graminóide [campina-rana], 0°49'N, 63°19'W, 13 Mar. 1984 (fl, imm fr), W.A. Rodrigues et al. 10653 [Aracá Inventory IL2-6] (INPA, NY); 3 km S da parte central da Serra Aracá, 8 km E do Rio Jauari, solo silicoso ao redor do campamento, campina lenhoso-graminóide [campina-rana], escandente até 0.3 m, 0°49'N, 63°19'W, 13 Mar. 1984 (fl, imm fr), W.A. Rodrigues et al. 10662 [Aracá Inventory IL7-18] (INPA, NY, RB); parte baixa ou pé da Serra Aracá, terreno arenoso ou catinga, planta de 60 cm, flores roseas, 21 Feb. 1977 (fl), N.A. Rosa & M.R. Cordeiro 1562 (MG, NY [2 sheets]); arredores do rio da Serra

Aracá, erva de 50 cm, flores roseas, 28 Jan. 1978 (fl, fr), N.A. Rosa & S.B. Lira 2268 (MG, NY, UB). **Pará:** Campo do Jamaracarú, perto do barracão, Região do Ariramba, 26 May 1957 (fl), G.A. Black et al. 57-19606 (IAN); Alto Ariramba, campinarana, 20 Dec. 1906 (fl), A. Ducke s.n. (MG 8020) (MG, R, RB; the first collection of this species); Mun. Óbidos, Jaramacarú, 27 May 1957 (fl), W.A. Egler 273 (MG); Mun. Óbidos, Campos do Ariramba, campos near Igarapé Mutum and Rio Ariramba, whitesand soil, 4 Dec. 1987 (fl), C. Farney & E.F. Batista 2057 (F, NY, US); Mun. Óbidos, 91 km de Oriximiná, nos Campos de Ariramba, entre Rio Jaramacarú e Igarapé Mutum, campina aberta, solo areno-pedregoso, ca. 1°10'S, 55°35'W, 4 Dec. 1987 (fl), C.A.C. Ferreira 9769 (INPA, MO, RB); Mun. Oriximiná, Campos do Ariramba, campinas da margem do Rio Jaramacarú, afloramento arenítico, 80 m, 8 Jun. 1980 (fl, imm fr), G. Martinelli et al. 6929 (INPA, RB).

### 9-3. *Sipaneopsis foldatsii* Steyerm., Mem. New York Bot. Gard. 17: 288. 1967. (Figures 75, 77).

Steyerm. in Lasser & Steyermark, Fl. Venez. 9: 280-281, fig. 46. 1974.

Type: VENEZUELA. Amazonas: Santa Cruz [ca. 3°15'N, 67°20'W], margen del Río Atabapo, cerca de la desembocadura del Río Atacavi, arbusto fruticoso, flores rosadas, crece en lugar pantanoso durante la estación de lluvias, 4 Sep. 1960 (fl, fr), *E. Foldats* 3678 (holotype VEN [Acc. No. 63682]).

*Subshrubs* 30-50 cm tall; stems erect or ascending (basal portion sometimes prostrate), densely appressed-strigose; hairs antrorse, white or pale brown. *Stipules* free at base, triangular to broadly triangular, 3-4 × 2-3 mm, acute at apex, rarely bifid at apex in older stipules, the lobes (when present) narrowly triangular, densely strigose outside, inside not seen. *Leaves* petiolate; petioles 2-4 mm long, hispid-strigose; blades elliptic-lanceolate, 3-6 × 0.8-3 cm, acute to subobtuse at base, acute at apex, papyraceous to subcoriaceous, dark green, slightly shiny above, pale green below, drying brown to dark olive green above and pale olive green below, densely hispid-strigose above and below; with 5-10 secondary veins each side. *Inflorescences* congested-cymose, expanding after anthesis, with branched portion extremely reduced, 21-39-flowered, 2-4.5 cm long (including peduncle and flowers), 0.6-0.8 cm broad, rachis appressed-strigose; hairs antrorse, white; peduncle 1.7-2.3 cm long. *Flowers* sessile to short-pedicel.
<figure><img src="image_81.png" /><figcaption>**Figure 77.** *Sipaneopsis foldatsii*. **A.** Branch with terminal inflorescence. **B.** Flower at final stage of anthesis, side view. **C.** Dissected flower at final stage of anthesis, with stamens at distal portion of corolla tube. **D.** Longitudinal section of hypanthium, ovary, disk and calyx. **E.** Stem. **F.** Seeds. **G.** Node with stipule and basal portion of petioles. Illustration by Bruno Manara (Reproduced with permission from Flora de Venezuela vol. 9(1), fig. 46. 1974).</figcaption></figure>

cellate, subtended by one bracteole narrowly triangular to narrowly lanceolate, 1–2 × 0.2–0.4 mm, acute at

apex, appressed-strigose. *Hypanthium* broadly turbinate, ca. 2 × 1.7 mm, densely hispid, hairs erect. *Calyx*
lobes equal or subequal, narrowly lanceolate, 2.5-2.7 × 0.5-0.7 mm, densely hispid outside, margins ciliate with appressed rigid hairs, glabrous inside, acute at apex; with 2 colleters in each calycine sinus, narrowly ellipsoid, acute at apex, 0.2 mm long. *Corollas* hypocrateriform, pink, 20.5-24 mm long when fully elongated; tube cylindrical and slightly wider at mouth, 14-16 mm long when fully elongated, 0.8 mm wide at base, 2 mm wide at mouth, densely antrorse-hispidulous outside, glabrous on basal 2.5 mm and puberulous at medio-distal portion inside; lobes left-contorted, obovate, 6.5-8 × 3-4 mm when fully expanded (perpendicular to the tube), obtuse to acute at apex, strigose-hispidulous outside, hispidulous inside. *Stamens* included, inserted at 3.5 mm from corolla mouth; filaments 0.4 mm long; anthers subsessile, oblong-lanceolate, 2 mm long, round at base, acute at top. *Style* included, ca. 8.5 mm long, glabrous; style branches unknown. *Infructescences* cymose, 3.5-5 cm long, fruits subtended by persistent bracteoles; terminal branches scorpiod, 1.5-2 cm long. *Fruits* urceolate 3.5 × 3 mm, densely sericeous. *Seeds* ca. 12 per locule, ovoid to ellipsoid, 1 × 0.9 mm, foveolate, testa dark brown.

**Notes:** For differentiation of *Sipaneopsis foldatsii* from *S. rupicola*, see notes under the latter.

**Distribution and ecology:** A rare species only known by three collections from a locality near the town of Santa Cruz, at the margin of the Río Atabapo, state of Amazonas, Venezuela (Figure 75). Growing in white-sand savannas seasonally inundated, at about 100 m altitude.

**Phenology:** The three gatherings with flowers and fruits were collected in September and October.

**Suggested conservation status:** Critically Endangered (CR). A very rare species known by three collections from the same locality near the town of Santa Cruz (ca. 3°15'N, 67°20'W), which is at the margin of the Río Atabapo, at the junction of the Río Atacavi, State of Amazonas, Venezuela. The collections were made in September and October 1960, and the specimen labels report this species as a subshrub locally abundant, growing in an area that is flooded during the rainy season. This locality is not within a protected area. The size of the population was not reported on the specimen labels, and no recent field observations have been made to assess the status of the conservation of this locality. Therefore, because of its rarity and the small Extent of Occurrence (< 100 km²), this species is estimated to

be Critically Endangered (CR) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** VENEZUELA. Amazonas: Santa Cruz [ca. 3°15'N, 67°20'W], margen del Río Atabapo, cerca de la desembocadura del Río Atacavi, arbusto de 50 cm de alto, flores rosadas, crece en lugar pantano-soso durante la época de las lluvias, 4 Oct. 1960 (fl, fr), *E. Foldats* 3692 (VEN, paratype); Santa Cruz [ca. 3°15'N, 67°20'W], margen del Río Atabapo, cerca de la desembocadura del Río Atacavi, muy abundante, flores rosadas, crece en terrenos inundados durante la estación lluviosa, 9 Oct. 1960 (fl, fr), *E. Foldats* 3830 (VEN, paratype).

9-4. *Sipaneopsis huberi* Steyerm., Ernstia 23: 37, fig. 11. 1984. (Figures 75, 78).

**Type:** VENEZUELA. Amazonas: Depto. Río Negro, domo rocoso de gneiss granitoide, ubicado en la ribeira derecha del Río Siapa medio, ca. 15 km al E de la Punta N del Cerro Aracamuni, "Arbusto/frútice hasta 1 m de alto, poco frecuente, cáliz verde claro, corola rosada, haz verde muy oscuro mate, envéz verde claro seríceo, frutos globosos de ca. 0.5 cm de diam., verdes", 1°41'N, 65°41'W, 350 m, 6 Feb. 1981 (fl, fr), *O. Huber & E. Medina* 5786 (holotype VEN [Acc. No. 309632]; isotypes MO [barcode 3235600], NY [barcode 00133329], U [barcode 0006318], US [barcode 00054383]).

*Shrubs* 50-100 cm tall; stems erect or ascending, terete, 1.5-2.5 in diam., densely antrorse-strigose. *Stipules* adnate to petioles and sheathing at basal 3 mm, free portion deltoid to narrowly triangular at base, distally bifid, each lobe linear, 2.5-3 × 0.2 mm, acute at apex, densely strigose outside, sparsely scabrous and with a few colleters at base inside. *Leaves* subsessil to short-petiolate; petioles 1-4 mm long, hispid-strigose; blades lanceolate, elliptic-lanceolate to narrowly oblong-lanceolate, 2.5-7.7 × 0.9-2.3 cm, acute to narrowly cuneate at base, acuminate at apex, papyraceous, dark green above, pale green below, drying dark olive green above and pale olive green below, sericeous above and below; with 7-9 secondary veins on each side, impressed above and prominent below. *Inflorescences* congested-cymose when young and unexpanded, during anthesis a compound dichasium, 31-147-flowered, with 5-9 axes dichomously branched, 2-6.5 cm tall, 2-4 cm broad (including peduncle and flowers); terminal secondary branchlets scorpiod, 5-9-flowered, 1-2.5 cm long; rachis densely sericeous; peduncle 2-4 cm long, 1 mm thick, densely sericeous.
<figure><img src="image_82.png" /><figcaption>**Figure 78.** *Sipaneopsis huberi*. A. Branch with inflorescences in anthesis and an old infructescence with fruits fallen off. B. Node with stipule and basal portion of petioles. C. Flower at final stage of anthesis, with corolla lobes reflexed, dehisced anthers, and receptive stigmas. A-C: drawn from *Huber & Medina 5786* (U, isotype). Illustration by Piero Delprete.</figcaption></figure>
*Flowers* secundiflorous, sessile to shortly pedicellate, subtended by one bracteole; bracteoles linear-lanceolate to setaceous, 2-4 mm long, densely antrorse-strigillose; pedicels to 0.5 mm long, densely sericeous. *Hypanthium* oblong-obovoid, ca. 1 × 1 mm, strigillose. *Calyx* lobes equal, linear-lanceolate, 1.5 × 0.2-0.5 mm, strigose outside, margins ciliate with appressed rigid hairs, glabrous inside, acute at apex; with 2 colleters in each calycine sinus, narrowly ellipsoid, acute at apex, 0.2 mm long. *Corolla* hypocrateriform, 13-14 mm long when fully elongated, densely strigillose outside; tube cylindrical and slightly wider at mouth, 9.5-10 mm long when fully elongated, 0.5 mm wide at base, 1-2 mm wide at mouth, hispidulous outside, glabrous inside; lobes irregularly left contorted in bud, obovate, 4-4.5 × 2.5-3 mm when fully expanded (perpendicular to the tube), round to obtuse at apex, strigose-hispidulous outside, hispidulous inside. *Stamens* included, inserted at 6 mm from base of corolla tube; filaments ca. 1.1 mm long; anthers positioned at upper ¼ of corolla tube, narrowly oblong, 1.5 mm long, round at base, apiculate at top. *Style* (not fully elongated) included, ca. 5.5 mm long, glabrous; style branches lanceolate, ca. 1 mm long. *Infructescences* a compound dichasium 5-8 cm long, with 5-9 axes dichomously branched, with 31-147 fruits, not all of them present at any given time, because falling off as they reach maturity; terminal branchlets scorpiod, 2-3 cm long. *Fruit* globose, ca. 5 mm in diam. *Seeds* unknown.

**Notes:** This species is similar to *Sipaneopsis rupicola* and *S. moricensis* Steyerm. because of its multifid stipules. According to Steyermark (1984c: 39) it is most similar to *S. rupicola* because of the leaf shape and inflorescence type, from which it differs by the calyx lobes densely strigose outside, the longer style, and the leaf blades densely strigose-sericeous on both sides.

*Sipaneopsis huberii* Steyerm. differs from *S. morichensis* by the stems densely antrorse-strigose (vs. densely ochroleucous-villous in *S. morichensis*), stipules densely strigose outside, with two linear lobes 2.5-3 mm long (vs. ochroleucous-villous (hairs spreading and/or acropetally oriented) outside, topped by (1-)3-4 linear lobes 8-10 mm long), leaf blades sericeous above and below (vs. ochroleucous-strigose above, densely ochroleucous-villous below), corolla 13-14 mm with tubes 9.5-10 mm long when fully elongated, hispidulous outside, and lobes strigose-hispidulous outside (vs. corolla 14.5-18 mm long with tubes 11-12 mm long when fully elongated, densely grayish-strigose outside, and lobes sericeous or appressed-pilose outside).

**Distribution and ecology:** Known from four col-

lections from the southern portion of the Venezuelan state of Amazonas (Figure 75). Growing on flat rocky outcrops (“lajas”) in savannas formations, at 200-420 m altitude.

**Phenology:** Specimens with flowers in anthesis were collected in February and November, and specimens with flowers and fruits in February and June.

**Suggested conservation status:** Endangered (EN). A rare species known from four collections from four localities in the Río Negro Department, southern portion of Venezuelan state of Amazonas, Venezuela. The first collection, the type, was made in February 1981, on a rocky dome of granitic gneiss, 1°41'N, 65°41'W, 350 m, located at the margin of the Río Siapa Medio, ca. 15 km E the N pick of Cerro Aracamuni. The specimen label reports this species as an infrequent shrub.

The second collection, a paratype, was made in June 1981 in the central portion of the plateau of the Serranía del Vinilla, 2°26'N, 65°20'W, 420 m, in hilly savanna, at ca. 20 km SW of the town of Mavaca. The specimen label reports this species as a locally frequent subshrub growing in humid savanna.

The third collection was made in 1987 near the Río Siapa, near the base of Cerro Aracamuni, 1°39'N, 65°40'W, at 250 m altitude, growing on a flat rocky outcrop. The frequency of this species in this locality was not reported on the specimen label.

The fourth and most recent collection was made in February 1990 near the Río Metacuni, 3°2'N, 65°12'W, at 200-230 m altitude, also growing on a flat rocky outcrop. The frequency of this species at this locality was not reported.

All the above described localities are within preserved areas, either within the Parima Tapirapecó National Park, the Serra da Neblina National Park, or the Reserva de Biosfera. This region is inhabited by indigenous people of the Yanomami ethnic group, who are allowed to hunt and cultivate traditional crops within the region. The size of the populations was not reported on the specimen labels, and no recent field observations have been made to assess the conservation status of these localities. Due to its rarity, its extent of occurrence (EOO) of ca. 3500 km², and human pressure in the area of occurrence, this species is treated as Endangered (EN) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** VENEZUELA. Amazonas: Depto. Río Negro, sabana collinosa en el sector central de una altiplanicie en la Serranía del Vinilla, ca. 20 km SW of Mavaca, 2°26'N, 65°20'W, 420 m, 13 Jun. 1981 (fl,
that the corollas are 15–25 mm long, with tubes 15–18 mm long and lobes 4.5–7 mm long. An error occurred in that description, as the tube and lobe length do not add up to the minimal length given for the corolla (i.e., 15 mm), or, most likely, the minimal length supplied by Steyermark might refer to corollas not completely expanded. Steyermark’s inconsistency might be due to the fact that in *Sipaneopsis* the corolla lobes open before the tube is completely elongated (i.e., before anthesis), and they remain erect and slowly elongating while the corolla is still expanding, and they become perpendicular to the tube and finish their expansion only after the corolla tube is completely elongated. On the type specimens of *S. wurdackii* (Maguire *et al.* 37639), fully expanded corollas have tubes 16–18 mm long and lobes 5–6 mm long, adding up to corollas 21–24 mm long.

Seyermark (1967: 285; 1974: 273–274), in the key to species of *Sipaneopsis*, distinguished *S. maguirei* from *S. wurdackii* by the corolla lobes glabrous and papillose within (vs. densely strigillose within in *S. wurdackii*) and leaf blades 3–6 cm long (vs. 1–2.7 cm long). However, after careful examination of the type specimens and additional material from Venezuela, Delprete and Steyermark (2004f: 833–835) concluded that these characters are found on a continuous morphological and geographical gradient. For example, specimens of *Davidse et al.* 17391 (MO, US), identified by Steyermark as *S. wurdackii*, have linear leaves 2–4 × 0.3–0.4 cm, and corolla lobes glabrous and papillose within. Therefore, I confirm that the two taxa are conspecific.

**Distribution and ecology:** This is the most widespread and most collected species of the genus. Known from numerous gatherings in Venezuelan state of Amazonas and in the Brazilian state of Amazonas, one collection in the Brazilian state of Rondônia, and several collections in the Colombian departments of Guainía and Vaupés (Figure 81). Commonly found in Amazonian open field savannas, dominated by grasses and sedges, or shrubby savannas (Figure 80A), herbaceous layer with sparse shrubs and treelets, in Brazil called “campinas” or “campinaranas”, respectively; growing on white sand, on white sand intermixed with black granitic sand (“salt & pepper sand”), or on white sand intermixed with clay, sometimes associated with *Brocchinia prismatica* L.B. Sm., often in areas seasonally inundated; at 55–984 m altitude.

**Phenology:** Due to the extended flowering period, it is common to see flowers and fruits on the same inflorescence. Such specimens were collected the whole year around.

**Suggested conservation status: Least Concern (LC).** This species is amply distributed in Amazonian Colombia (Guainía and Vaupés), Venezuela (Amazonas) and Brazil (Amazonas and Rondônia), and is known from numerous collections. It grows in open fields or shrubby savannas within the Amazon Basin, in Brazil called “campinas” or “campinaranas” respectively, and in Colombia and Venezuela called “catingas”. It sometimes forms large populations with sparse individuals (pers. obs., e.g. Delprete et al. 7451–7453). Because of its wide extent of occurrence (EOO) and its numerous collections, this species belongs the Least Concern (LC) category, following IUCN criteria (IUCN 2012, 2019).

**Specimens examined: COLOMBIA. Guainía:** Mun. Puerto Inírida, comunidad de Caño Vitina, ecotono entre bosque y las sabanas naturales de arenas blancas, arbustal de hasta 6–8 m de altura, denso, alta radiación solar hasta el suelo, suelos con mal drenaje y encharcados en época de lluvias, alta presencia de flor de Inírida (*Guacamaya superba*), hierba de 35 cm, flores rosadas, 9 Jun. 2015 (fl), *A. Barona et al.* 862 (COAH); Mun. Inírida, Río Inírida, sabana de arena blanca, Caño Vitina (margen derecha del resguardo de Caranacoa), 10 km N de Inírida, hierba de 40 cm, tallos jóvenes y flores rosadas, 170 m, 3°56'N, 67°55'W, 8 Sep. 2002 (fl), *D. Cárdenas & M. Avellaneda* 13484 (COAH); Mun. Inírida, Sector del Caño Culebra, sabanas naturales de arena blanca, con presencia de Flor de Inírida (*Schoenocephalium tertifolium* y *Guacamaya superba*), hierba de 20 cm, flor rosada, 3°46'49"N, 67°46'49"W, 120 m, 3 Mar. 2005 (fl), *D. Cárdenas & G. López* 16331 (COAH); Mun. Inírida, sector comprendido entre Caño Inírida y Caño Culebra, sabanas naturales de arena blanca, con presencia de arbustales y sabanas abiertas, hierba de 45 cm, flor lila, 3°48'32"N, 67°51'29"W, 120 m, 4 Mar. 2005 (fl), *D. Cárdenas López* 16435 (COAH); Mun. Inírida, Comunidad de Guamal, arbustal (catinga) sobre arenas blancas y sabanas naturales y pequeños bosques, hierba de 25 cm, flor rosada, 3°52'20"N, 67°51'08"W, 100–150 m, 5 Jan. 2007 (fl), *D. Cárdenas & J.G. Ramírez* 20466 (COAH); Mun. Inírida, Comunidad de Guamal, arbustal (catinga) sobre
<figure><img src="image_83.png" /><figcaption>Figure 79.</figcaption></figure>
<figure><img src="image_84.png" /><figcaption>**Figure 80.** *Sipaneopsis maguirei* A. Habitat. White-sand savanna with sparse vegetation. B. Inflorescence, top view. Note flowers at various stages of the flowering process. Young corollas pink, with lobes already opened and basal portion of lobes with appendages appressed against each other. The other two corollas completely expanded and opened, with lobes perpendicular to the tube, and basal appendages free from each other. C. Same inflorescence of B, side view. Note flowers at various stages of the flowering process. Short corollas, pink, with lobes already opened. One corolla at intermediate stage, with tube completely elongated, and lobes ascending (not completely reflexed). The other two corollas opened, with lobes perpendicular to the tube, and basal appendages free from each other. And two corollas with lobes completely expanded and perpendicular to the tube. D. Infructescence with immature fruits. Photos taken by Piero Delprete on 7 January 2001, Mun. San Fernando de Atabapo, Amazonas State, Venezuela, where *Delprete et al. 7451-7453* were collected.</figcaption></figure>
arenas blancas y sabanas naturales y pequeños bosques, hierba de 25 cm, flor rosada, 3°52'20"N, 67°51'08"W, 100-150 m, 5 Jan. 2007 (fl), *D. Cárdenas & J.G. Ramírez 20469* (COAH); Mun. Inírida, Corregimiento de Cacagual, comunidad Cacagual, sabana de arenas blancas con arbustales y matorrales, sufrutice de 20 cm, tallos rojizos, flores rosadas, 3°31'43"N, 67°25'53"W, 11 May 2007 (fl), *D. Cárdenas et al. 20723* (COAH); Mun. Puerto Inírida, comunidad de Caño Vitina vegetación en sabana natural de arenas blancas, hierba de 30 cm, flores rosadas, 150 m, 3°43'48"N, 67°50'3"W, 1 Dec. 2013 (fl), *D. Cárdenas et al. 44045* (COAH); Mun. Inírida, comunidad de Caño Vitina, sabanas naturales de arenas blancas, vegetación a borde de bosque, hierba de 25 cm, flor morada, 3°43'48"N, 67°50'3"W, 150 m, 8 Jun. 2015 (fl) (fl), *D. Cárdenas et al. 44326* (COAH); Mun. Puerto Inírida, comunidad de Caño Vitina, sabanas naturales de arenas blancas, hierba de 15 cm, flores rosadas, 3°48'21"N, 67°49'19"W, 150 m, 9 Sep. 2014 (fl), *M. Rodríguez et al. 1514* (COAH); Mun. Inírida, via Inírida-Caño Vitina, Sítio Curva de Los Pepes, sabanas naturales de arenas blancas, dominadas por varias especies de Cyperaceae, Xyridaceae y Rapateaceae, hierba, ejes de la inflorescencia, brácteas y tallos morados, flores

rosadas, 110 m, 3°48'45"N, 67°52'20"W, 7 Apr. 2006 (fl), *N.R. Salinas et al. 601* (COAH). **Vaupés:** Mitú and vicinity [ca. 1°11'N, 70°10'W, ca. 180 m], lower Río Kubiyú, sandstone savanna, 1 Jul. 1976 (fl), *J.L. Zarucchi & M.J. Balicki 1796* (F, MO, US).

**VENEZUELA. Amazonas:** Depto. Casiquiare, Caño San Miguel, Sector Las Tinajas y el Caño Iqueven, sabanas (dominancia de Rapateaceae) con bosque bajo sobre suelos quarzosos, 2°39'N, 66°45'W, 160 m, 25 Apr. 1991 (fl), *G. Aymard 9188* (MO, NY); Depto. Atabapo, SE bank of the middle part of Caño Yagua at Cucurbital de Yagua, inundated sabaneta, 3°36'N, 66°34'W, 120 m, 8 May 1979 (fl), *G. Davidse et al. 17391* (MO, US); Mun. San Fernando de Atabapo, Indigenous Community of Minicia Nuevo, sabaneta inundavel, at the moment of collection with dry soil, exposed white sand intermixed black granitic sand ("salt & pepper sand"), vegetation dominated by sparse shrubs 2-3 m tall, 3°55'N, 67°26'W, 120 m, 7 Jan. 2001 (fl, imm fr), *P.G. Delprete et al. 7451* (CAY, NY, TFAV, VEN), 7452 (CAY, MO, NY, TFAV, VEN), 7453 (CAY, MG, TFAV, VEN); Depto. Atabapo, sabanas y bosques ribereños en los alrededores de Canaripó, en la margen izquierda (S) del Bajo Río Ven-

<figure><img src="image_85.png" /><figcaption>**Figure 81.** Distribution of *Sipaneopsis maguirei*.</figcaption></figure>
tuari, ca. 20 km al E de la confluencia con el Río Orinoco, sabana inundada con hummocks ("tatucos") por debajo de morichal al S de la pista de aterrizaje, 4°3'N, 66°49'W, 98 m, 30 May 1978 (fl), O. Huber 1910 (NY); Depto. Atabapo, tercera sabana al pié del Cerro Yapacana, hierba común al borde de los motículos en la sabana anegadiza, cáliz rojo carmesí, corolla rosada, 3°38'N, 66°52'W, 100 m, 3 Jun. 1978 (fl), O. Huber 2021 (NY); Depto. Atures, sabana en la ribera N (derecho),

del Río Ventuari, frente al caserío Carmelitas, 4°13'N, 66°30'W, 100 m, 22 Aug. 1978 (fl), O. Huber 2420 (NY); Depto. Atabapo, sabana de Minicio, en la ribera S (Izquierda) del Río Orinoco, 3°56'N, 67°32'W, 23 Aug. 1978 (fl), O. Huber 2512 (NY); Depto. Atabapo, sabanas en las cabeceras del Caño Yagua, 3°57'N, 66°27'W, 120 m, 26 Aug. 1978 (fl), O. Huber 2691 (NY); Depto. Atabapo, pequeña sabanita distante 15 min al SE del Sítio Chipital, ubicado en la ribera izquierda (S) del Caño Yagua, ca.

<figure><img src="image_86.png" /><figcaption>**Figure 82.** *Sipaneopsis maguirei*, flowering process. **A, B.** Flower at initial stage of anthesis, with corolla lobe basal appendages appressed against each other. **C, D.** Longitudinal section of flower at intermediate stage of anthesis. **C.** Longitudinal section. **D.** Longitudinal section of distal portion of corolla tube and basal portion of lobes, with appendages appressed against each other. **E, F.** Longitudinal section of flower at successive intermediate stage of anthesis. **E.** Longitudinal section, with tube and lobes further elongated. **F.** Longitudinal section of distal portion of corolla tube and basal portion of lobes, with appendages appressed against each other. **G, H.** Longitudinal section of flower at final stage of anthesis, with mature anthers and style branches not receptive (still appressed to each other), allowing visitors and pollinators to access the corolla tube and nectar disk. **G.** Longitudinal section of flower, with lobes spreading perpendicularly to the tube, and erect appendages not touching each other. **H.** Longitudinal section of distal portion of corolla tube and basal portion of lobes, with appendages not touching each other. Drawing based on field observations, photographs and pickled flowers (*Delprete et al. 7452*). Drawn by Piero Delprete (Reproduced from fig. 3 of Delprete (2017) with permission of copyright holder Magnolia Press).</figcaption></figure>
15 km (el línea recta) río arriba desde la boca, 3°29'N, 66°41'W, 120 m, 6 Dec. 1978 (fl), *O. Huber & S.S. Tillett* 2900 (NY); Depto. Atabapo, extensas sabanas al NE y E del Cerro Cucurito, ribera SE del medio Caño Yagua, sabana de *Brochinia prismatica*, 3°36'N, 66°34'W, 120 m, 17 Jan. 1979 (fl), *O. Huber* 3111 (NY); Depto. Atures, sabana ubicada al pié SE del Cerro Moriche, 4°43'N, 66°22'W, 120 m, 19 Feb. 1979 (fl), *O. Huber* 3199 (NY); Depto. Atabapo, Caño Caname (aflente derecho (oriental) del Medio Río Atabapo), sabanita al N del Medio Caño Caname, ca. 18 km E de la boca, 3°40'N, 67°22'W, 100 m, 29 Apr-4 May 1979 (fl), *O. Huber et al.* 3748 (NY); Depto. Atures, sabana ubicada a ca. 5 km al S del Río Autana y ca. 15 km SSW del Cerro Autana, 4°45'N, 67°35'W, 100 m, 26 Jun. 1979 (fl, fr), *O. Huber* 3830 (NY); Depto. Atures, sabana ubicada a ca. 8 km al E del Río Orinoco, ca. a la altura de Morganito, 5°6'N, 67°46'W, 90 m, 3 Jul. 1979 (fl), *O. Huber* 4109 (NY); Depto. Atabapo, sabanas a ca. 30 km al W de la Seranía El Tigre, en la region del Alto Caño Yagua, 3°51'N, 66°27'W, 130 m, 29 Feb. 1980 (fl), *O. Huber* 4854 (NY); Depto. Atabapo, sabana a ca. 10-12 km W de Esmeralda, en la ribera derecho (N) del Río Orinoco, 3°11'N, 65°37'W, 125 m, 7 Mar. 1980 (fl, fr), *O. Huber* 5053 (NY); Depto. Casiquiare, sabana y caatinga abierta a ca. 2-3 km al SE del Bajo Río Guasacavi, 3°8'N, 67°30'W, 90 m, 10 Mar. 1980 (fl), *O. Huber* 5127 (NY); Depto. Atabapo, pequeña sabana ubicada a ca. 2 km al S del Medio Río Puruname, al SE del caserío de Puruname, 3°20'N, 66°32'W, 100 m, 22 Jul. 1980 (fl, fr), *O. Huber & S. Tillett* 5456 (INPA, NY); Depto. Atures, sabana ubicada en la ribera izquierda (S) del Río Guayapo medio, 4°14'N, 67°22'W, 100 m, 27 Jul. 1980 (fl), *O. Huber & S. Tillett* 5499 (INPA, NY); Depto. Atures, sabanas ubicadas a ca. 4 km al N del Río Pipapo Medio, ca. 5 km al W del Macizo del Sipapo, 4°36'N, 67°12'W, 120 m, 28 Jul. 1980 (fl, fr), *O. Huber & S. Tillett* 5589 (NY); Depto. Sasiquiare, extensas sabanas ubicadas a ca. 20 km al NW de Yavita, en las cabeceras del Caño Pimicín, 3°1'N, 67°33'W, 120 m, 11 Feb. 1981 (fl), *O. Huber & E. Medina* 5945 (NY); Río Orinoco, Savanna N. 3 at NW base of Cerro Yapacana [ca. 4°0'N, 66°35'], 130 m, 1 Jan. 1951 (fl), *B. Maguire et al.* 30549 (F, NY, US, paratypes of *S. maguirei*); Río Atabapo, between San Fernando de Atabapo and Caño Temi, savanna 1 km W of Cacagual, 130 m, 19 Nov. 1953 (fl, fr), *B. Maguire et al.* 36280 (NY, P, R, paratypes of *S. maguirei*); Río Guainía, Sabana El Venado, on L bank of Caño Pimichín [ca. 2°74'N, 67°56'W], frequent, 120-140 m, shrub to 0.2 m, corolla pink, 23 Nov. 1963 (fl), *B. Maguire et al.* 36353 (NY, paratype of *S. wurdackii*); Río Guainía, Sabana El Venado, on L bank of Caño Pimichín [ca. 2°74'N, 67°56'W], frequent, 140

m, 14 Apr. 1953 (fl), *B. Maguire & J.J. Wurdack* 35564 (F, NY, paratype of *S. wurdackii*); Cerro Yapacana, savanna 4, 3°45'N, 66°50'W, 140 m, forming herbaceous dense mat, 14 Mar. 1987 (fl), *K.J. Systma et al.* 5122 (NY); Savannas near Santa Cruz, a small village on the Río Atabapo 1 km N of the confluence of the Río Temeni and the Río Atacavi, 17-18 Nov. 1979 (fl), *W.W. Thomas & G.K. Rogers* 2680 (NY); area of San Fernando de Atabapo (4°2'N, 67°42'W), at the confluence of Atabapo and Guaviare Rivers with the Orinoco, Caño La Vieja, 10 km from Pozo Lucas and 20 km from San Fernando, along rd to Santa Barbara, 27 May-3 Jun. 1974 (fl), *S.S. Tillett* et al. 745-319 (NY).

**BRAZIL. Amazonas:** Mun. Barcelos, Serra do Aracá, Platô da Serra do Aracá, Acampamento da Grotinha, vegetação sobre afloramento rochoso e sob solo arenoso, 0°52'4"N, 63°19'56"W, 984 m, 22 Aug. 2014 (fl), *F.N. Cabral et al.* 1235 (INPA); [Mun. Humaitá], Estrada do Estanho, rd to Igarapé Preto, ca. 60 km SE of Transamazonian Hwy, 2 Jul. 1979 (fl), *C.E. Calderon et al.* 2750 (INPA, US); Mun. Novo Aripuanã, Parque Estadual do Matupiri, savana arbustiva, inundação lençol raso periodicamente, solo arenoso, ca. 4°49'55"S, 61°0'12"W, 55 m, 28 Nov. 2012 (fl), *F.M. Costa & M.L.G. Mari* 1920 (INPA); Maués, Campina Socoró, campina arbustiva alagável, p orte baixo, com densa cobertura de gramíneas baixas, 4°17'2"S, 58°34'40"W, 7 Aug. 2015 (fl), *L.O. Demarchi et al.* 523 (INPA); Mun. Lábrea, Parque Nacional Mapinguari, Campos do Rio Coti (acesso aéreo, em elicóptero), campinaranas gramíneo-lenhosa e arbustiva recém queimadas, 8°55'47"S, 65°23'45"W, 110 m, 26 Jul. 2014 (fl), *L.L. Giacomin & Y.F. Gouvêa* 2131 (INPA); N margin of Rio Aracá, just above Igarapé Sauadaua, savanna on white sand, subshrub 30 cm tall, 26 Jul. 1985 (fl), *G.T. Prance et al.* 29861 (INPA, NY); Canutama, Parque Estadual do Matupiri, campina gramíneo-lenhosa, solo hidromórfico areno-argiloso, 5°0'45"S, 61°16'15"W, 29 Oct. 2010 (fl), *E.M.B. Prata et al.* 789 (INPA); Canutama, Reserva de Desenvolvimento Sustentável do Matupiri, campina gramíneo-lenhosa, solo hidromórfico areno-argiloso, 4°50'46"S, 60°31'46"W, 2 Nov. 2010 (fl), *E.M.B. Prata et al.* 930 (INPA); Borba, Campina da Catuquira, vegetação savânica encharcável, com Poaceae, Rapateaceae, Cyperaceae, Humiriaceae, *Bactris campestris*, 4°54'13"S, 61°6'15"W, 30 m, 11 Jul. 2007 (fl), *P.L. Viana et al.* 3042 (INPA); Parque Nacional do Jaú, Campina do Patauá, caminho para a cabeça do espermatozoide [ca. 2°15'S, 62°38'W], campo queimado, campina arbustiva, 5 Sep. 1998 (fl), *A. Vicentini & J.G. Loco* 1391 (INPA, MO). **Rondônia:** Igarapé Preto, SC-20-XA-Ponto.30 [ca. 9°25'S, 62°30'W], 30 Jun. 1975 (fl), *J.C. Silva* 120 (MG).
9-6. *Sipaneopsis morichensis* Steyerm., Mem. New York Bot. Gard. 17: 286. 1967. (Figures 75, 83).

Steyerm. in Lasser & Steyermark, Fl. Venez. 9: 275-277, fig. 45. 1974.

Type: VENEZUELA. Amazonas: Cerro Moriche [ca. 7°18'N, 63°23'W], Río Ventuari, 800 m, in montane caño savanna, shrub to 2 m tall, flowers pink, 14

Jan. 1951 (fl), B. Maguire, R.S. Cowan & J.J. Wurdack 30878 (holotype NY [barcode 00133333]; isotypes MICH [barcode 1108305], MO [barcode 1811402], NY [barcode 00133332], VEN [Acc. No. 65195]).

*Shrubs* to 2 m tall; stems erect; young branches stout, 4-5 mm in diam., villous, hairs ochroleucous. *Stipules* free or shallowly adnate to petioles, free portion broadly triangular at base, topped by 1-4 linear lobes

<figure><img src="image_87.png" /><figcaption>**Figure 83.** *Sipaneopsis morichensis*. A. Branch with terminal inflorescence. B. Flower at final stage of anthesis and flower bud. C. Longitudinal section of hypanthium, ovary, disk and calyx. D. Node with stipule and basal portion of petioles. E. Stamens, dorsal and ventral view. F. Dissected flower at final stage of anthesis, with stamens at median portion of corolla tube, and detail of moniliform hairs. Illustration by Bruno Manara (Reproduced with permission from Flora de Venezuela vol. 9(1), fig. 45. 1974).</figcaption></figure>
8-10 mm long, villous, hairs ochroleucous, spreading or acropetally oriented outside, white-sericeous, with 6-7 colleters inserted just below the margin inside, with tips exserted; margins villous. *Leaves* with petioles 2-5 mm long, densely villous; blades broadly lanceolate, oblong-elliptic to narrowly ovate, 4.5-8.5 x 1.5-2.7 cm, acute to obtuse at base, acute at apex, chartaceous to subco-riaceous, dark green above, ochroleucous-villous below when fresh, drying brown above and beige-villous below, sulcate and hispidulous with midvein and secondary veins ochroleucous-strigose above, densely ochroleu-cous-villous below; with 8-11 secondary veins on each side, depressed above, prominent below, strongly ascending, terminating near the margin. *Inflorescences* congest-ed-cymose, branched portion reduced during anthesis, 27-39-flowered, 3.5-4.5 cm tall, 2-3 cm broad (including peduncle and flowers), trichomously branched; terminal branchlets 3-7-flowered; rachis villous, hairs ochroleu-cous, spreading or acropetally oriented; peduncle 1-2.5 cm long, densely ochroleucous hirsute; bracts subtend-ing the inflorescence subulate, 4-11 mm long, canescent outside, sometimes leaf-like, 2.5-2.7 x 0.8-1.4 cm. *Flow-ers* secundiflorous, sessile to subpedicellate, pedicels to 0.5 mm long, villous, hairs ochroleucous, spreading or acropetally oriented; subtended by one bracteole; bracte-oles lanceolate to narrowly lanceolate, decreasing in size towards the tip, dimensions 3.5-5.5 x 1-1.5 mm, villous, hairs ochroleucous, spreading or acropetally oriented outside, glabrous inside. *Hypanthium* broadly obovoid, 1.5-1.7 x 1.5-1.7 mm, white-sericeous-strigose, hairs both acropetal and spreading. *Calyx* lobes equal or sube-qual, lanceolate, 2-2.5 x 0.5-0.7 mm, acute at apex, gray or ochroleucous sericeous outside (hairs 0.5 mm long), glabrous inside; with 1-2 colleters in each calycine sinus, narrowly ellipsoid, acute at apex, 0.25 mm long. *Corol-la* hypocrateriform, pink, 14.5-18 mm long when fully elongated (lobes perpendicular to the tube); tube cylin-drical and slightly wider at mouth, 11-12 mm long when fully elongated, 1.5 mm wide at base, 3-3.5 mm wide at mouth, densely grayish-strigose outside, glabrous at basal 3-3.5 mm and papillose-villosulous from filament insertion to corolla mouth inside; lobes left-contorted, subequal, oblong to narrowly obovate, 4.5-6 x 2-3.5 mm when fully expanded (perpendicular to the tube), round or ovate at apex, white-sericeous or appressed-pilose outside, glabrous and papillose inside. *Stamens* included, inserted 4-5 mm below corolla mouth; filaments 1.5 mm long; anthers oblong-lanceolate, 1.5 mm long, acute at top, round at base. *Style* included, 6.5-10 mm long, glabrous; style branches lanceolate, ca. 1 mm long. *Infructescences* a compound dichasium, 8.5-9 cm long, villous, hairs ochroleucous, spreading or acropetal-

ly oriented; terminal branches scorpiod, 2.5-3.5 cm long. *Mature fruits and seeds* unknown.

**Distribution and ecology:** Known from only two collections from Cerro Moriche, state of Amazonas, Venezuela, at 300-800 m elevation (Figure 75).

**Phenology:** The two flowering collections were made in January.

**Suggested conservation status:** Critically Endan-gered (CR). A species only known by two collections made in 1951 on Cerro Moriche, ca. 7°18'N, 63°22'W, at 300-800 m, state of Amazonas, Venezuela. Cerro Moriche is located not far from the Río Ventuari, and is an important center of biodiversity with a high degree of endemism. As a result of the same expedition to this locality in January 1951, the botanists B. Maguire, R.S. Cowan and J.J. Wurdack discovered numerous new taxa that were subsequently described [e.g., *Axonopus arun-dinaceus* G.A. Black (1957: 251), *Brocchinia cowanii* L.B. Smith (1957: 293), *Caraipa llanorum* subsp. *cordifolia* Kubitzki (1978: 104), *Cassia tetraphylla* var. *ventuaren-sis* Irwin (1964: 95), *Clidemia morichensis* J.J. Wurdack (1964: 183), *Clusia annularis* Maguire (1961: 23), *Gon-gylolepis paniculata* Maguire & Phelps (1953: 157), *Ilex savannarum* var. *morichei* Edwin (1965: 147), *Macrolo-bium discolor* var. *egranulosus* R.S. Cowan (1953: 292), *Navia crispa* L.B. Smith (1953: 378), *Navia hohenber-gioides* L.B. Smith (1957: 298), *Navia semiserrata* L.B. Smith (1957: 309), *Orthaea thibaudioides* Maguire, Stey-ermark & Luteyn (1978: 173), *Perama galioides* forma *cinerea* Steyermark (1964: 243), *Phoradendron micros-tachyum* Kuijt (2003: 301), *Remijia argentea* Steyermark (1972a: 267), *Rudgea morichensis* Steyermark (1967: 424), *Siphanthera cowanii* J.J. Wurdack (1958: 97), *Stegolepis hitchcockii* subsp. *morichensis* Maguire (1965: 83)], including the present species.

The label of the type specimen (*Maguire et al. 30878*) reports *Sipaneopsis morichensis* Steyerm. as a shrub growing near a small mountain creek at 800 m altitude, at the top of Cerro Moriche. The label of the paratype specimen (*Maguire et al. 30868*) describe it as a frequent shrub growing on the east flank of Cerro Moriche, between 300 and 800 m. The size of the popu-lations was not reported on the specimen labels. Cerro Moriche is part of the “Monumento Natural Macizo Guao-Sipapo y Cerro Moriche” (Guao-Sipapo and Cerro Moriche Massif Natural Monument), a protected area that was established in 1990. Because of its rarity, small extent of occurrence (EOO) of < 100 km², and small area of occupancy (AOO < 10 km²), this species is treat-
ed as Critically Endangered (CR) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** VENEZUELA. Amazonas: Cerro Moriche [ca. 7°18′N, 63°22′W], frequent in E slope, 300–800 m, 13 Jan. 1951 (fl), *B. Maguire et al.* 30868 (NY, US, VEN, paratypes).

9-7. *Sipaneopsis pacimoniensis* Steyerm., Mem. New York Bot. Gard. 17: 289. 1967. (Figures 75, 84).

Steyerm. in Lasser & Steyermark, Fl. Venez. 9: 282. 1974.

**Type:** VENEZUELA. Amazonas: Infrequent in savanna on R bank of Río Pacimoni [Río Pasimoni, ca. 2°0′N, 66°43′W], 50 km above mouth, 100–140 m, 7 Feb. 1954 (fl), *B. Maguire, J.J. Wurdack & G. Bunting* 37570 (holotype NY [barcode 00133334]).

*Perennial herbs* 10–12 cm tall, single-stemmed, erect; young branches thin, densely appressed-canescent. *Stipules* free at base, triangular, 2.5–3 mm long, densely antrorse-strigose outside, inside not seen. *Leaves* shortly petiolate; petioles 1–2 mm long, densely antrorse strigulose; blades oblong-elliptic to narrowly ovate, 1.5–3 × 0.7–1.2 cm, acute to obtuse at base, acute to subacute at apex, chartaceous, blade colors when fresh unknown, drying dark reddish brown above and below, glabrous with midvein strigulose above, densely strigulose at midvein and secondary veins below; with 3–5 secondary veins on each side, ascending; margins strigulose-ciliate. *Inflorescences* cymose, 2–3-flowered; peduncle 1–2 cm long, canescent. *Flowers* 5-merous, sessile to subpedicellate, pedicels to 1.5 mm long, hispidulous; subtended by one bracteole linear-lanceolate, 3–4 mm long, acuminate, hispidulous outside, inside unknown. *Hypanthium* broadly subturbinate, 1.5 × 1.5 mm, densely hispidulous. *Calyx* lobes subequal, lanceolate, 1.7–2 × 0.7 mm, acute at apex, strigose-canescent outside, glabrous inside; with 1 colleter in each calycine sinus, narrow cylindroconical, acute at apex, 0.2 mm long. *Corolla* hypocrafteriform, pink, 27.5–28.5 mm long when fully elongated; tube cylindrical and gradually wider near the mouth, 19 mm long when fully elongated, 0.7 mm wide at base, 2.5 mm wide at mouth, canescent-sericeous or canescent-strigose, densely so near the mouth outside, glabrous at basal 3.5 mm and puberulent above inside; lobes left-contorted in bud, subequal, obovate, 8.5–9.5 × 4.5–5.5 mm when fully expanded (perpendicular to the tube), obtuse and abruptly mucronate at apex, appressed-strigulose outside, glabrous inside.

*Stamens* included, inserted 4 mm below corolla mouth; filaments 3 mm long, papillose; anthers oblong-lanceolate, 1.8 mm long, acute at top, round at base. *Style* included (not fully elongated), 11.5 mm long, glabrous; style branches unknown. *Infructescences, fruits and seeds* unknown.

**Notes:** Steyermark (1967: 289) wrote that *Sipaneopsis pacimoniensis* Steyerm. "differs from *S. foldatsii* in the dwarf stems, shorter internodes, shorter leaves, much larger corolla lobes, glabrous interior of corolla-lobes, and fewer lateral nerves of the leaf-blades." Steyermark (1967: 289; 1974: 282) also wrote that the corolla of *S. pacimoniensis* is 26 mm long, with a tube 17 mm long and lobes 8–9.5 mm long; however, the total of the tube plus the maximal length of the lobes add up to more than 26 mm. This species is only known from the holotype specimen preserved at NY, which has a single flower in full anthesis. Careful measurements of this specimen revealed that the corolla is 27.5–28.5 mm long, with tube 19 mm long and lobes 8.5–9.5 mm long.

Delprete and Steyermark (2004f) in the key to species of *Sipaneopsis* differentiated *S. pacimonensis* from *S. foldatsii* by the petioles 1–2 mm long (vs. 2–4 mm long in *S. foldatsii*), leaf blades 1.5–3 cm long, glabrescent except the midrib strigillose above, strigose below (vs. 3–7.5 cm long, densely hispid-strigose above and below), with 3–5 secondary veins on each side of the midrib (vs. 6–10 each side), and corolla lobes 8–9.5 mm long, glabrous adaxially (vs. 3.5–6.5 mm long, strigose-hispidulous adaxially).

**Distribution and ecology:** Only known from the type specimen, collected in a savanna on the right bank of Río Pacimoni, state of Amazonas, Venezuela, at 100–140 m altitude (Figure 75).

**Phenology:** The sole flowering collection was made in February.

**Suggested conservation status:** Critically Endangered (CR). A species only known from the holotype specimen collected in 1954 at about 50 km above the mouth of the Río Pacimoni, ca. 2°0′N, 66°43′W, at 100–140 m, state of Amazonas, Venezuela. This herb is 10–12 cm tall and reported as infrequent in a savanna near the Río Pacimoni. The size of the populations was not reported in the specimen label, and no recent observation has been made in the area to assess its conservation status. Because of it is known from a single collection locality, which is not within a protected area, this species is Critically Endangered (CR) following IUCN criteria (IUCN 2012, 2019).
<figure><img src="image_88.png" /><figcaption>**Figure 84.** *Sipaneopsis pacimoniensis*. A. Habit. B. Node with stipule and basal portion of petioles. C. Flower in anthesis, side view. D. Longitudinal section of flower at intermediate stage of anthesis, with corolla lobes not completely reflexed, and style not completely elongated. A–D: drawn from *Maguire et al. 37570* (NY, holotype). Illustration by Piero Delprete.</figcaption></figure>
9-8. *Sipaneopsis rupicola* (K.Schum.) Steyerm., Mem. York Bot. Gard. 17: 285, fig. 29g-m. 1967. (Figures 85-86).

Steyerm. in Lasser & Steyermark, Fl. Venez. 9: 274-275. 1974. - *Rondeletia rupicola* K.Schum. in Martius et al., Fl. Bras. 6(6): 222. 1889.

Type: VENEZUELA. Amazonas: "ad flumina Casiquiare, Vasiva et Pacimoni," s.d. [1853-1854] (fl), *R. Spruce* 3392 (B†; lectotype BR [barcode 00532837], here designated; isolectotypes C [without barcode], E [barcode 00285371], G [barcode G00436652], GH [barcode 00094363], L [barcode 1214530], LD [Acc. No. 1214530], K n.v., NY [barcode 00133193], P [barcode P00729443], RB [2 sheets, barcodes 00560059 and 00543699]; frag at F [Acc. No. 587152]; photo-B at F and NY; photo-K and photo-US at NY).

(=) *Rondeletia rupicola* var. *chiribiquetana* R.E. Schult., Bot. Mus. Leafl. 14: 46. 1949, **syn. nov.**

Type: COLOMBIA. Vaupés: Macaya-Ajaju River confluence, Mount Chiribiquete [ca. 1°19'N, 72°45'W], quartzitic base, summit, 1700-2100 feet [520-640 m], 15-16 May 1943 (fl, fr), *R.E. Schultes* 5448 (holotype GH n.v.; isotypes BR [barcode 00532837], COL [barcode 000004687]; UC [barcode 925799], US [barcode 00137636]).

*Subshrubs* to *shrubs*, 0.3-2 m tall, erect; young branches stout, strigulose. *Stipules* free at base, triangular, 4-5 mm long, entire and caudate, or base triangular topped by 2-3 acute lobes, or base reduced to a line 0.5-0.7 tall with 6-7 linear lobes, central one longer, 3-4 mm long, the lateral ones shorter, 1.2-1.7 mm long, or two longer ones intercalated with shorter ones, densely strigulose outside, white-sericeous with 10-11 colleters inserted just below the margin inside with tips exerted; margins strigulose. *Leaves* with petioles 3-4 mm long, sericeous-strigose; blades elliptic, narrowly ovate, narrowly elliptic to elliptic-lanceolate, rarely lanceolate, 2.7-6.5 × 0.8-1.5 cm, acute to obtuse at base, acute or acuminate at apex, subcoriaceous, margins narrowly revolute, dark green above and pale green below when fresh, drying dark brown above and pale brown, glabrous or sparsely strigose and sulcate with midvein

and secondary veins white-strigose above, glabrous or sparsely to densely strigose, or densely sericeous-strigose with midvein and secondary veins white- or ochroleucous-strigose below; with 7-9 secondary veins on each side, ascending. *Inflorescences* cymose, a compound dichasium with scorpiod secondary branches, 21-47-flowered, 3-4 cm long (including peduncle and flowers), 2.5-3 cm broad during anthesis (expanding after anthesis), with 2-3 main branches, each branch 1-3-furcate, ultimate branchlets 3-6-subsecundiflorous; peduncle 1.5-3 cm long, densely strigose. *Flowers* sessile or with pedicels 0.5 mm long; except for central flower with pedicel 1.5-2 mm long; subtended by bracteoles linear-lanceolate, decreasing in size towards the tip of the branch, 2-3.5 × 0.1-0.3 mm, acuminate, strigulose outside, glabrous inside. *Hypanthium* subglobose, 1.2-1.5 × 1 mm, strigose-sericeous. *Calyx* lobes subequal, lanceolate, 1-1.5 × 0.5-0.7 mm, acute at apex, sparsely to densely strigose outside, glabrous inside; with 1 colleter in each calcine sinus, narrowly conical (acute at apex), 0.2 mm long. *Corolla* hypocrateriform, pink, 14.5-16.2 mm long when fully expanded; tube cylindrical and slightly wider at mouth, 11-11.5 mm long when fully expanded, 0.7 mm wide at base, 1.5 mm wide at mouth, sericeous-strigose, densely so near the mouth outside, glabrous at basal 3.5-4 mm, puberulent above and papillose near the mouth inside; lobes left-contorted or irregularly imbricate, 1 or 2 external, subequal, oblong-obovate, 3.5-4.7 × 2.2-2.5 mm when fully expanded (perpendicular to the tube), obtuse to round at apex, sparsely strigulose outside, glabrous inside. *Stamens* included, inserted at 4.5 mm below corolla mouth; filaments 1.5-2 mm long; anthers oblong-lanceolate, 1-1.5 mm long, round at base, acute at top. *Style* as long as the corolla tube, with branch tips barely exerted, glabrous; style branches lanceolate, ca. 1 mm long. *Infructescences* a compound dichasium, 6.5-7.5 cm long, terminal branches scorpiod, 3-3.5 cm long. *Immature fruits* subglobose-urceolate, 3 × 2.5 mm (fide Steyermark, 1974: 275), strigose. *Seeds* 8-10 per locule, those inside immature fruits subglobose, 0.7-1 mm diam. (fide Steyermark, 1974: 275), rugose-foveolate. *Mature fruits* and *seeds* unknown.

**Morphological observations:** Steyermark (1967: 285-286; 1974: 274-275) described the corolla of *S. rupicola* as being 11 mm long, with tube 8-8.5 mm long

**Figure 85.** *Sipaneopsis rupicola.* **A–B.** Habit variation. **A.** Habit from Mount Chiribiquete (Colombia). **B.** Habit from Aracaruara (Colombia). **C–E.** Stipule variation. **F–G.** Leaf shape variation and detail of vestiture on abaxial side. **H.** Flower bud (note lobes already opened at early stage). **I.** Flower in anthesis. **A, D, E, F, H, I** from *Gutierrez & Schultes* 654 (US); **B, G** from *Sastre & Raichel* 5150 (P); **C** from *Williams* 14897 (US). Illustration by Piero Delprete.
<figure><img src="image_89.png" /><figcaption>Figure 85.</figcaption></figure>
and lobes 3–3.5 × 1.7–2 mm, and style only 4 mm long; however, these measurements were made on flowers not completely expanded. The corollas and style are much larger when fully expanded, with corolla tubes 11–11.5 mm long and lobes 3.5–4.7 × 2.2–2.5 mm, and style is as long as the corolla tube.

**Notes:** Schumann (1889: 222 [in *Flora Brasiliensis* the page is erroneously numbered as “224”]) published *Rondeletia rupicola* K. Schum. citing “*Sipanea rupicola* Spruce Ms.! in Pl. *exsicc. n. 3392*” in synonymy. Art. 46.4 of the *Code* (Turland et al. 2018) states that “When a validly published name or its final epithet is taken up from and attributed to the author of a different “name” that has not been validly published, or one at a different rank likewise not validly published, only the author of the validly published name is cited (except as provided in Art. 46.7).” Therefore, “*Sipanea rupicola* Spruce” is not a name, and should not be referred to as such. It is here cited between quotation marks to show that it is not a name (see Art. 36.1, Exs. 1-8).

Schultes (1949: 46) separated *Rondeletia rupicola* var. *chiribiquetana* R.E. Schult. from the typical variety by the leaves elliptic-lanceolate, 4–5 × 1–1.3 cm, acute at apex (vs. long-lanceolate, 5–8 × 1.5 cm, acuminate at apex in the typical variety), bullate above, mid-

rib and secondary veins densely sericeous-strigillose (vs. glabrous or sparsely to densely strigose), and larger corolla lobes. Steyermark (1967: 285–286) and Delprete and Steyermark (2004f: 833–835) synonymized the two varieties without any comment. A detailed comparison of the specimens from throughout the geographic range of the species confirmed that the characters used by Schultes to differentiate the two varieties are found on a gradient. For example, examination of several collections demonstrated that leaf size is variable within the populations present on Cerro de Chiribiquete. Also, in the populations of Cerro de Chiribiquete the leaf abaxial side is densely sericeous-strigillose, while in those from the Colombian and Venezuelan lowlands the leaf abaxial is glabrous or sparsely to densely strigose, which could indicate a possibility of the recognition of two varieties. However, the collections from near Araracuara and Inírida (Colombia lowlands) have ovate-elliptic blades densely strigose below, which are here interpreted as an intermediate form between the two putative varieties. Also, corolla lobes in *Sipaneopsis* are already open in flower bud and gradually expand during the flowering process, reaching the full length only at final stage, when they are completely reflexed and perpendicular to the corolla tube. Corolla size and vestiture is consistent among all the collections examined. As a result of the

<figure><img src="image_90.png" /><figcaption>**Figure 86.** Distribution of *Sipaneopsis rupicola*.</figcaption></figure>
above analysis, no variety is here recognized in *Sipaneopsis rupicola*.

Several herbarium specimens of *Sipaneopsis rupicola* have been incorrectly identified as *S. maguirei*, although these species differ in many characters. In *S. maguirei* the stems are thin, ascending or reclining (rarely decumbent), the stipules are entire (i.e., not lobed), broadly to narrowly triangular, the inflorescences are congested, 2-8-flowered, and fully elongated corollas are 21-30 mm long (tubes 15-22 mm long, lobes 4.5-9 mm long). Whereas, in *S. rupicola* the stems are stout, erect, the stipules can be entire and caudate or reduced at base and with 2-3 or 6-7 linear lobes, the inflorescences are cymose, compound dichasia with scorpioid secondary branches, 21-47-flowered, and fully elongated corollas are 14.5-16.2 mm long (tubes 11-11.5 mm long, lobes 3.5-4.7 mm long).

A few herbarium specimens of *Sipaneopsis rupicola* from Colombia have been incorrectly identified as *S. foldatsii*. In *S. foldatsii*, the stems and the leaves are densely appressed-strigose, the hypanthia are broadly turbinate, densely hispid, and fully expanded corollas are 20.5-24 mm long (tube 14-16 mm long, lobes 6.5-8 mm long). Whereas, in *S. rupicola*, the stems are strigulose and the leaves are glabrous or sparsely strigose (rarely densely strigose below), the hypanthia are subglobose, strigose-sericeous, and fully expanded corollas are 14.5-16.2 mm long (tube 11-11.5 mm long, lobes 3.5-4.7 mm long).

**Distribution and ecology:** Known from several collections in Amazonian Colombia and Venezuela (Figure 86). Growing on sandstone mountains (Tepuis), or on rocky or white-sand savannas, at 100-640 m altitude.

**Phenology:** Specimens with flowers were collected in January, February, March, April, July, September and November. Specimens with flowers and immature fruits were collected in May, June and September.

**Suggested conservation status:** **Endangered (EN).** A rare species known from several collections in Colombia and five in Venezuela. In Colombia, two collection were made in 1943 and 1949 (type of *Rondeletia rupicola* var. *chiribiquetana*) on Cerro de Chiribiquete, at 520-640 m, department of Vaupés. Cerro de Chiribiquete [ca. 1°31'N, 72°47'W] is a sandstone tepui located at the easternmost portion of the Guiana Shield, and is included within the Chiribiquete National Natural Park, which was declared a UNESCO World Heritage site in 2018. Three Colombian collections were made in the Vaupés department, in 1976, 1992 and 1994, near the locality of Araracuara [ca. 0°36'55'S, 72°20-23'W]. And two other

collections of *Sipaneopsis rupicola* where made in 1992 and 2007 in the municipality of Inírida, in the department of Guainía, in the same locality where *S. maguirei* grows.

In Venezuela, all the collections were made in the state of Amazonas. The first collection (type of *Rondeletia rupicola* K. Schum.) was made in 1853-1854 in a region cited on the specimen label as "Rivers Casiquiare, Vasiva and Pacimoni." Judging by the three rivers cited, this region is now included within the Alto Orinoco-Casiquiare Biosphere Reserve. Two additional collections were made in 1942 and 1959 near Caño San Miguel (ca. 2°39'N, 66°45'W and ca. 2°37'N, 66°40'W), and two other collections were made in 1954 and 1957 along the Casiquiare River, near Piedra Tururumeri and near Piedra Arauicaua, which are also within the Alto Orinoco-Casiquiare Biosphere Reserve.

Several localities where this species has been collected are within the Chiribiquete National Natural Park, in Colombia, and others within the Alto Orinoco-Casiquiare Biosphere Reserve, in Venezuela. The size of the populations was not reported in the specimen labels. Based on the collection localities, its EOO is of ca. 48,600 km². Although several localities are present within preserved areas, several other collection localities in non-protected areas near human communities that could be impact its natural environment, this species is treated as Endangered (EN) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** **COLOMBIA.** Caquetá: Río Caquetá, Araracuara [ca. 0°36'S, 72°23'W], camino a la Sabana, 2° Sabana de Tibeyes, suelo arenoso blanco, sobre rocas de arenisca, 5 Jan. 1976 (fl, fr), *C. Sastre* & *H. Raichel D. 5150* (P [2 sheets]); Mun. Puerto Santander, trocha a La Chorrera, 20 km SW de Araracuara, sabana de areniscas (aduche) (Bh-T), hierba de 50 cm, flor rosada, 0°55'S, 72°20'W, 160-260 m, 2 Apr. 1994 (fl), *Cárdenas et al. 4540* (COAH); Araracuara, areniscas de la Sabana de Aduche, arbusto de 2 m, flores rosadas, 25 Nov. 1992 (fl), *J. Murillo et al. 215* (COAH, COL). **Guainía:** Mun. Inírida, comunidad de Guamal, arbustal (catinga), sobre arenas blancas y sabanas naturales y pequeños bosques, hierba de 70 cm, flor roja, creciendo en sabana, 3°52'20"N, 67°51'08"W, 100-150 m, 5 Jan. 2007 (fl), *D. Cárdenas & J.G. Ramírez 20499* (COAH); Mun. Inírida, Río Atabapo, sector entre Maviso y comunidad de Chaquita, hierba de 80 cm, flores fucsia, en sabana, 4°2'15"N, 67°42'43"W, 9 May 2007 (fl), *D. Cárdenas et al. 20522* (COAH). **Vaupés:** Cerro de Chiribiquete [ca. 1°19'N, 72°45'W], a un lado del Río Macaya, terreno muy pedregoso, 17 Jan. 1944 (fl), *G. Gutiérrez & R.E. Schultes 654* (US).
**VENEZUELA. Amazonas:** Ríos Pacimoni-Yatua, Casiquiare, frequent in open areas on Piedra Turumeri, 110-220 m, 4 Feb. 1954 (fl), *B. Maguire et al.* 37512 (F, MO, NY); Ríos Pacimoni-Yatua, Casiquiare, locally frequent in clearing, Piedra Arauicaua, Río Yatua, 110-550 m, 27 Sep. 1957 (fl, imm fr), *B. Maguire et al.* 41603 (F, NY, RB, SP); Guainía, margenes y alrededor de las lajas, Caño San Miguel, cerca de la Boca del Caño Ichana [ca. 2°39'N, 66°45'W], 127 m, 26 Mar. 1942 (fl), *Ll. Williams* 14897 (F, US); Caño San Miguel, between Limoncito and Caño Ikebenie, ca. 70 river km from mouth [ca. 2°37'N, 66°40'W], 120-130 m, 29 Jun. 1959 (fl, fr) *J.J. Wurdack & L.S. Adderley* 43253 (NY, UB, US).

## 10. STEYERMARKIA

*Steyermarkia* Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 216. 1940.

Standley & Williams, Fl. Guatemala, Fieldiana, Bot. 24: 203-204, fig. 10. 1975; D.H. Lorence in G. Davidse et al., Fl. Mesoamericana 4(2): 284. 2012.

Type: *Steyermarkia guatemalensis* Standl.

*Subsessile perennial herbs or subshrubs*, with a rosette of leaves condensed near the ground; stem short, woody. *Stipules* persistent, free at base, lanceolate to linear-lanceolate, attenuate or rarely shortly bifid at apex. *Leaves* opposite, condensed in rosettes, short-petiolate; blades narrowly obovate or oblanceolate, membranaceous or chartaceous; domatia absent. *Inflorescences* axillary, long-pedunculate, not frondose, capitate, subtended by bracts. *Flowers* homostylous, 4-merous. *Hypanthium* turbinate, densely antrorsely hispid, hairs white, long, slightly enlarged-tuberculate at base. *Calyx* persistent; tube extremely reduced; lobes large, erect, narrowly ovate-lanceolate to linear-lanceolate, densely long-hispid-sericeous outside; with a linear colleter in each calycine sinus. *Calycophylls* absent. *Corolla* hypocrateriform, actinomorphic, pink; tube externally hirsute, internally glabrous throughout, puberulent at mouth; lobes left-contorted, ovate, margin entire, round at apex. *Stamens* included; filaments attached on the upper part of the corolla tube, free at base, slender, short, equal, glabrous; anthers subsessile, elongate, acute

at base, round at apex, dorsifixed near the base. *Pollen* 3-colporate; exine foveolate, perforate. *Style* partially exserted (branch tips exserted just above the corolla mouth), filiform; style branches oblong. *Ovary* with a stalked placenta; stalk inserted at the middle of the septum, with an oblong-hemielliptic extension; ovules many, inserted on the placental extension. *Fruit* capsular, dehiscing loculicidally, crustaceous. *Seeds* many per locule, horizontal, minute, globose.

**Geographic distribution:** A rare monotypic genus restricted to southern Mexico (Chiapas) and Guatemala.

**Notes:** Standley (1940) dedicated the name of this genus to Julian A. Steyermark (1909-1988), who collected the type specimens in Guatemala during the Stanley Field Expedition of 1939-1940. As Standley predicted, this monotypic genus is extremely rare. Later collections demonstrated that it occurs also in the contiguous Mexican state of Chiapas, and the only collection from Guatemala is the type gathering. Standley (1940) also indicated that *Steyermarkia* Standl. is most similar to *Sipanea*, from which it differs by being a plant reduced to a basal rosette of leaves, with large, 4-merous flowers, and globose seeds. On the other hand, *Sipanea* species are herbs or shrubs, erect or decumbent, with cauline leaves, with smaller, (4)5(6)-merous flowers, and angular seeds.

10-1. *Steyermarkia guatemalensis* Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 216. 1940. (Figures 5K-L, 87-88).

Standl. & Williams, Fl. Guatemala, Fieldiana, Bot. 24(9): 203-204, fig. 10. 1975.

Type: GUATEMALA. Izabal: Río Dulce, 2-4 miles W of Livingston, on S side (L side going up-river) [ca. 15°46'54"N, 88°46'54"W], shaded moist limestone bluffs near base, 1-3 m above sea level, 16 Apr. 1940 (fl), *J.A. Steyermark* 39520 (lectotype F [Acc. No. 1035008], designated by Lorence (1999: 170); isolectotypes F [Acc. No. 1035009], MO [Acc. No. 1204537]).

*Stems* short, unbranched, to 18 cm long; internodes very short. *Stipules* 1.5-2 cm long, densely white-hirsute.

**Figure 87.** *Steyermarkia guatemalensis*. *A.* Habit of plant in bloom. *B.* Hypanthium and calyx (note partially exserted calycine colleters). *C.* Corolla partially dissected showing stamens and exserted style. *D.* Detail of outer surface of corolla. *E.* Young flower bud. *F.* Seed. *G.* Cross section of hypanthium and ovary. Upper right corner: Sketch of Dr. Julian Steyermark. Illustration by M. Pahl. (Reproduced with permission from Flora of Guatemala vol. 24, part 11, fig. 10. 1975).
<figure><img src="image_91.png" /><figcaption>Figure 87.</figcaption></figure>
sute outside, glabrous inside. *Leaves* with petioles 1-4 cm long, hirsute; blades 11.5-35 × 4-10 cm, attenuate at base, acute or acuminate at apex, chartaceous, dull green above, pale green and midrib white below, drying reddish brown above and olive green below, lamina densely long-hirsute and veins strigose above (hairs to 6 mm long), more densely hirsute below; with 15-20 secondary veins on each side of midrib. *Inflorescences* 10-12-flowered, with 3 head-like cymules, hirsute, rarely with an additional subterminal inflorescence; peduncle 9-16 cm long, long-hirsute, (hairs erect, beige, 3-4 mm long); bracts subtending the head narrowly lanceolate to narrowly ovate-lanceolate, 15-20 × 3-4 mm, long-attenuate, connate at base, with 2-3 colleters in each calycine sinus; bracteoles subtending the flowers narrowly lanceolate, 1.5-2 cm long, acuminate, long-hispid (hairs appressed antrorse, beige, 2-3 mm long) outside, glabrous inside. *Flowers* sessile or short-pedicellate. *Hypanthium* 1.5-2 × 1-1.5 mm. *Calyx* tube 0.5-1 mm long, densely long-hispid-sericeous outside (hairs white, 2-3 mm long), glabrous inside; lobes erect, 10-15 × 1-1.5 mm, acute at apex; with 1 colleter in each calycine sinus, linear, 0.7 mm long. *Corolla* 4.6-6 cm long; tube cylindrical and slightly wider at mouth, 3.5-4.5 cm long when fully expanded, 1.5 mm wide at base, 1.5-3 mm wide at mouth, with long-spreading hairs (3-4 mm long) outside, glabrous inside; lobes subequal, broadly ovate, 1.4-2 × 0.9-1.3 cm, round at apex, densely hispid, with long-

spreading hairs outside, glabrous and sparsely glandular at base inside. *Stamens* inserted near the base of corolla tube; filaments 4-6 mm long, glabrous; anthers narrowly oblong, 5-6 mm long, round at both ends. *Style* 3.5-4.5 cm long, glabrous; style branches oblong, 4-4.5 mm long, acute at tip. *Fruit* subglobose, 6.5-8 × 6.5-8 mm, densely long-hirsute. *Seeds* 90-110 per locule, globose to subglobose, 0.3-0.4 mm diam., foveolate, testa dark brown.

**Notes:** Standley (1940) along with the description of *Steyermarkia guatemalensis* Standl., cited as type *Steyermark 39520* preserved at F. However, at F are present two specimens of *Steyermark 39520*, which are kept in separate type folders and have different accession numbers. Lorence (1999: 170) cited the F specimen with Acc. No. 1035008 as the holotype, and cited the F specimen with Acc. No. 1035009 and a MO specimen (without citing the accession number) as isotypes. According to Art. 7.11 and 9.10 of the *Code*, Lorence's citation of the specimen with Acc. No. 1035008 as the holotype is a correctable error, and should be treated as an inadvertent lectotypification on this specimen.

**Distribution and ecology:** Restricted to the state of Chiapas, southern Mexico, and known by a single collection in northern Guatemala (Figure 88). Growing on rock walls, in tropical rain forest understory, from near sea level to 550 m altitude.

<figure><img src="image_92.png" /><figcaption>**Figure 88.** Distribution of *Steyermarkia guatemalensis*.</figcaption></figure>
**Phenology:** Specimens with flowers were collected in April, October and December; and with flowers and fruits in April and May.

**Conservation status:** Critically Endangered (CR) or Probably Extinct (EX). This species is known from a single collection from Guatemala and four collections from the contiguous Mexican state of Chiapas. The sole collection from Guatemala (the type) was collected in 1940 in the community of Izabal, near Río Dulce, at 2-4 miles W of Livingston [ca. 15°46'54"N, 88°46'54"W], in shaded moist limestone bluffs. Standley and Williams (1975: 203-204) wrote that the habitat of *Steyermarkia* in Guatemala "may have been destroyed where the plants were collected more than 30 years ago." In fact, the type specimen remains the sole record from this country, where it might be locally extinct.

In Mexico, this species was collected four times in the state of Chiapas. Two collections were made in nearby localities, one in 1973 in the municipality of Palenque, ca. 17°29'N, 92°2'W, on shaded cliff face, at 300 m, and the other in 1986 in the municipality of Chontal, 17°28'48"N, 92°4'48"W, on a vertical rock face in forest along a river, at 450 m. The other two collections were made in the municipality of Ocosingo, ca. 16°54'N, 92°5'W, in 1980 and 1981, in lower montane rain forest on a rock wall, at 550 m altitude. This species was last collected in 1986.

The extent of occurrence (EOO) of this species, counting the type locality in Guatemala, is of 11.700 km². However, as stated by Standley and Williams in 1975, its natural environment in Guatemala has most likely been obliterated. Therefore, if the Guatemalan locality is excluded from the calculation, its EOO in Mexico is of 243 km². Also, the size of the populations was not reported on the specimen labels, and none of the localities are within protected areas. In Mexico, the natural vegetation in the region between Ocosingo and Palenque is extremely fragmented due to agriculture and general human pressure. Because of the small area of occurrence, fragmented natural vegetation, and probably small populations, this species is treated as Critically Endangered (CR) following IUCN criteria (IUCN 2012, 2019). On a final sad note, considering that its last collection was made in 1986, and it is not cultivated in any garden, it is also possible that this species is Extinct (EX).

**Additional specimens examined:** MEXICO: Chiapas: Mun. Palenque [ca. 17°29'N, 92°2'W], on shaded cliff face, tropical rain forest adjacent to small cascading river at Agua Azul, 300 m, 24 May 1973 (fl, fr), *D.E. Breedlove* 35338 (CAS, MO, NY); Mun. Ocosingo [ca. 16°54'N, 92°5'W], lower montane rain forest, 70

km SW of Palenque, on road to Ocosingo along the Jol Uk'um, plant on rock wall, 550 m, 4 Dec. 1980 (fl), *D.E. Breedlove* 48293 (CAS, MO); Mun. Ocosingo [ca. 16°54'N, 92°5'W], lower montane rain forest, 70 km SW of Palenque, on road to Ocosingo along the Jol Uk'um, 550 m, 12 Apr. 1981 (fl, fr), *D.E. Breedlove* 50883 (CAS [2 sheets]); Mun. Chontal, rd between San Cristobal de las Casas and Palenque, 136 km NE of San Cristobal, Villa Paraiso, on vertical rock face in forest along river, 17°28'48"N, 92°4'48"W, 450 m, 10 Oct. 1986 (fl), *B.E. Hammel et al.* 15637 (BM, F, MO).

## ACKNOWLEDGMENTS

This project started more than two decades ago, and, due to my moves to different institutions and numerous concurrent long-term projects, it was stopped and restarted several times. This gave me the opportunity to visit numerous herbaria and undertake precious field work in many Neotropical countries. My first field expedition dedicated to the Sipaneeae took place in Venezuela in December 2000 – January 2001, for which I wish to thank Gerardo Aymard (then at UNELLEZ-Guanare, Herbario Universitario (PORT), Mesa de Cavacas, Venezuela) for help during the application for the collecting permit, and Ana Claudia Araujo (at that time doctoral student at the University of São Paulo, Brazil, currently at the Natural History Museum, London, England), who helped during field work and processing herbarium specimens; that expedition gave me the opportunity to observe the unique flower modality of *Sipaneopsis maguirei* in a population growing in a white-sand savanna near San Fernando de Atabapo. Following that field trip, I compiled sufficient material for a molecular phylogenetic study of the Sipaneeae, which was accomplished in collaboration with Rocío Cortes-Ballén (then a doctoral student under my supervision at City University of New York (CUNY), New York, USA, and now professor at the Universidad Distrital, Bogotá, Colombia).

Important advancements towards this project were made while I was working as a Visiting Scientist at the National Herbarium of the Netherlands, Utrecht University branch, when I started the treatment of the Rubiaceae for the series *Flora of the Guianas*, in 2004; funds for that visiting fellowship were provided by the Netherlands Organization for Scientific Research NWO (grant B 85-368), and additional funds were granted from the Alberta Mennega Stichting and the Van Eeden Foundation for work in 2006 and 2008. My profound gratitude goes to Marion Jansen-Jacobs (U) for the efficient coordination of the project and for answering
my daily questions, and the entire staff of the Nationaal Herbarium Nederland, Utrecht University branch, now integrated with Naturalis, Nationaal Herbarium Nederland, Leiden, for their friendly collaboration and for their kind support. I am also much in debt to Erik Simonis, Rob van Aubel, Erik van Marle, and Rob Bakker for help in the handling of specimens at U and the large loans from NY and US, deciphering the historical labels from the Guianas, help with bibliographic references, and routine herbarium activities. Paul Hiepko (1932–2019), Botanischer Garten und Botanisches Museum, Berlin-Dahlem, also contributed his professional assistance during my visit of the Willdenow Herbarium (B-W) at Berlin in 2004.

Important observations of several species of *Sipanea* and *Limnosipanea* in Central-Western Brazil, in the states of Goiás, Tocantins and Mato Grosso, were made during 2004–2008, while I was working as a Visiting Scientist at the Federal University of Goiás (UFG), with a fellowship from the Conselho Nacional de Pesquisa (CNPq) of the Brazilian Government (grant 309885/2003–5). Special help during that period was provided by Juliana Costa da Silva (now my wife), who accompanied me during numerous field expeditions and herbarium studies in this region.

Field work in French Guiana was made while working at the IRD, starting from 2009 to present; during this period, I received help from Gwenaël Quenette and Juliana Costa da Silva, who accompanied me during local field excursions, provided digital images, and helped processing herbarium and pickled specimens of *Neobertiera palustris* from a population growing along the trail to the Bagne des Annamites, French Guiana, in 2013–2014. Funds for field work, during which I had the occasion to study an additional population of *N. palustris* in the surroundings of the village of Camopi, upper Oyapock River basin, French Guiana, were provided by the LABEX CEBA (Laboratory of Excellence - Center for the Study of Biodiversity in Amazonia; Project TEKO); during this expedition, I was accompanied by Guillaume Odonne (CNRS), Damien Davy (CNRS), and Sophie Gonzalez (IRD), who are here acknowledged for help during field work.

In Suriname, important observations on populations of *Sipanea pratensis* var. *dichotoma* and *S. veris* were made during a multidisciplinary expedition, in April–May 2014, to the surroundings of the Sipaliwini village, coordinated by Antoine Fouquet (CNRS, Laboratoire Evolution et Diversité, Toulouse, France), who also obtained the collecting permits for the entire team. At the Sipaliwini village we were hosted by the Trio Indig'enous tribe, which is here acknowledged. During that

expedition I was helped by the personnel of the Herbarium (BBS) University of Suriname, Paramaribo, especially Gunovaino Marjanom for collecting and processing herbarium specimens, and Dorothy J. Traag, at that time Herbarium Curator, for local coordination as well as help for obtaining the collecting and export permits. Funds for that expedition were provided by the LABEX CEBA (Project PHYLOGUYANA).

Field work in Guyana in 2016, during which I studied natural populations of *Pteridocalyx* in the surroundings of the Chenapou Village, *Chalepophyllum guyanense* and *Sipanea veris* in the Kaieteur National Park, Upper Potaro River. Funds for that expedition were provided by the LABEX CEBA (Laboratory of Excellence - Center for the Study of Biodiversity in Amazonia). During that expedition I was accompanied by Paul Benjamin, excellent field guide and based at the Chenapou Village, of the Patamona Indigenous people, and Sebastien Cally, at that time doctoral student studying opilions at the University of Toulouse, France, who are here acknowledged for their help during field work. Thanks are also due to the staff of the Biodiversity Center of the University of Guyana, Georgetown, especially Kaslyn Holder-Collins and Elford Liverpool for help in gathering field supplies, drying facilities, and their valuable collaboration. Thank are also due to the Environmental Protection Agency (E.P.A.) of Guyana, and especially Diana Fernandes for her assistance in the process of obtaining the collecting permit (No. 060716 BR004) and the export permit (No. 062716 SP007).

The following members of the *Muséum National d'Histoire Naturelle* of Paris are here acknowledged: Odile Poncy and Cécile Aupic, for help with historical herbaria (P-JJR, P-JU, P-AD), Sovanmoloy Hul, for her supervision during the study of Rubiaceae specimens of the general herbarium (P), and Gérard G. Aymonin (1934–2014), for conversations about Aublet specimens, help with taxonomic matters, facilitating access to historical collection, supplying historical literature, and identifying the authors of historical handwritings on herbarium specimens.

I am also grateful to the various herbarium curators, especially Johnatan Gregson, of the Natural History Museum of London (BM) for help searching for Aublet specimens. Significant help was provided by Lynda Brooks and Janet Ashdown for studying Aublet Rubiaceae specimens in the Smith Herbarium (LINN-SM) of the Linnean Society of London.

The contributions of Nicolas Fumeaux and Laurent Gautier are here recognized, during the consultation of the Rubiaceae specimens in the general herbarium (G) and in the Candolle herbarium (G-DC) of the Conserva-
toire et Jardin Botanique de la Ville de Genève, and Fer-
nand Jacquemoud (G) for digital images of specimens in
the Candolle Herbarium (G-DC). Funds for that project
were provided by the Institut de Recherche pour le Dével-
opplement (IRD). I am also much in debt to Bruno Wall-
nöfer (W), Curator of Vascular Plant Collections of the
Naturhistorisches Museum, Wien, Austria, for placing
digital images of types on the website of his institution.

Additional support was provided by the Herbarium
of the Montpellier University (MPU), during 2008–2009,
for providing working space and particularly to Peter
Schäfer, then Herbarium Curator, for information about
the historical collections preserved in this institution.

Gerardo Aymard is here acknowledged for help find-
ing the coordinates of remote collection localities in
Venezuela, as well as figuring out if they are included
within protected areas.

Funds for visiting the IAN, INPA, MG, and NY her-
baria in 2018 were provided by the LABEX CEBA (Labo-
ratory of Excellence - Center for the Study of Biodiver-
sity in Amazonia) and by the UMR AMAP – CIRAD,
CNRS, INRA, IRD, Université de Montpellier – of the
IRD (Institut de Recherche pour le Développement). I
would also like to thank Michael Hopkins, INPA her-
barium curator, for assistance in the herbarium and for
hosting me in his private residence during my stay in
Manaus in 2018.

I am extremely grateful to the directors and cura-
tors of the following herbaria for loans of material and/
or for providing working space during my visits: B,
B-W, BBS, BM, BR, BRB, C, CAS, CAY, CEN, COL, E,
F, FI, FT, G, G-DC, HEPH, HTO, IAN, IBGE, INPA, K,
L, LINN-SM, MBM, MG, MO, MPU, NY, NX, P, P-AD,
P-BONPL, P-JJR, P-JU, R, RB, U, UB, UFG, UFMT,
UMP, US, and VEN.

Rebecca Peters, Herbarium Curator of the Califor-
nia Academy of Sciences, San Francisco, U.S.A., is here
acknowledged for sending digital images of specimens
of *Steyermarkia guatemalensis* in the CAS herbarium.
Thalia Morales, Herbarium Curator (VEN), Universi-
dad Central de Venezuela, Caracas, Venezuela, is here
acknowledged for sending digital images of numerous
specimens preserved at VEN.

I wish to thank the staff members of the projects
Jstor Global Plants (https://plants.jstor.org/), Reflora –
Brazilian Plants: Historic Rescue and Virtual Herbarium
for Knowledge and Conservation of the Brazilian Flora
(http://reflora.jbrj.gov.br/reflora/PrincipalUC/Princi-
palUC.do), and INCT – Herbário Virtual da Flora e
Fungos (http://inct.splink.org.br/) for making millions of
images of herbarium specimens available to the general
public.

My profound thanks are due to Ib Friis, of the Nat-
ural History Museum of Denmark, Copenhagen, who
provided important help interpreting Rottbøll’s sev-
eral publications and re-publications, and figuring out
which of them included the first description and illus-
tration of *Rondeletia biflora*, as well as valuable infor-
mation on Vahl’s herbarium, and consulting Vahl’s card
catalogue on the accession *Virecta biflora*. I also wish
to thank Henning Knudsen, of the same institution, for
searching the copper plate engraving and the drawing
commissioned by Rottbøll of *Rondeletia biflora* in the
library archives.

Important help regarding nomenclatural issues was
supplied by Joseph H. Kirkbride Jr., John H. Wiersema
(both at the Smithsonian Institution, Washington, D.C.,
USA), Kanchi Gandhi (Harvard University, Cambridge,
Massachussetts, U.S.A.), and John McNeill (Royal
Botanic Garden, Edinburgh, Scotland), who made consid-
erable contributions in solving numerous nomenclatural
problems, interpreting the *Code* (Turland et al. 2018),
and for their assistance in understanding the correc-
tion of the term isotype. Last, but not less important, I
wish to express my gratitude to the Joseph H. Kirkbride
and Gerardo Aymard (UNELLEZ-Guanare, Herbario
Universitario (PORT), Mesa de Cavacas, Venezuela;
Compensation International Progress S.A., Ciprogress-
Greenlife, Bogotá, Colombia) for the careful revision
of the final version of manuscript and their significant
comments and corrections.

A special thank is due to the staff of the CAY
Herbarium – Herbier IRD de Guyane, https://dx.doi.
org/10.23708/herbier-guyane-ird – for their help in han-
dling the loans of herbarium specimens from numerous
institutions.

Finally, I wish to thank the Centro Studi Erbario
Tropicale (Herbarium FT of the University of Firenze,
Italy) and Riccardo M. Baldini, Editor in Chief of Webbia
Journal, for sponsoring the publication of this monograph.

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