McDowell & Stobey 3853, Mutchnick et al. 156); however, *S. cururuensis* is known only from the type gathering from the Tapajos River Basin [ca. 5°18′N, 59°18′W], south of the Amazon River, and can be distinguished from *Neobertiera pakaraimensis* by the generic characters of *Sipaneopsis* (e.g., corolla opening before sexual parts mature, corolla lobes with a basal appendage covered with a tuft of hairs, and homostylous flowers).

In the description of *Neobertiera pakaraimensis*, Delprete (2015a) wrote that the corollas are “4.8–5 mm long” with tubes “4.5 mm long” and lobes “oblong, 3–3.2 × 1.1–1.2 mm”. While the corolla lobes shape and dimensions were correctly reported, the corollas of this species are 7–7.7 mm long with tubes 4–4.5 mm long.

**Distribution and ecology:** Endemic to Guyana (Figure 30), growing at elevations of 120–1125 m. It is known by four collections: two on the Pakaraima Mountain Range, eastern portion of the Roraima Massif, at 975–1125 m elevation, one on islands at Amatuk Falls, at 720 m, and one near the Eping River, at 120 m; growing at river margins inside wet forest.

**Phenology:** The collections with flowers in anthesis were made in October and February, and one collection with flowers and fruits in September.

**Suggested conservation status:** Endangered (EN). Delprete (2015) at the time of the original description of this species, was aware of only two nearby collection localities, and assigned this species to the category of Endangered (EN) following IUCN criteria. Currently, this rare species is known from three localities: two on the Pakaraima Mountain Range, at 975–1125 m elevation, one near Amatuk Falls on the Potaro River, at 720 m, and one near the Eping River, at 120 m. The size of the populations is unknown, and the Extent of Occurrence (EOO) is ca. 4,100 km². None of these localities are within protected areas, and no recent observations to assess their conservation status have been made. The basin of the Potaro River is subject to indiscriminate, often illegal, gold hunters' activities, either by mining, panning, or by dragging the bottom of rivers with dredgers (Delprete, pers. obs.). Because the localities where it has been collected are threatened by these human activities, this species is here confirmed to be Endangered (EN), following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** GUYANA: Kuyuni-Mazaruni Region, Eping River, common along river, 6°0′N, 60°10′W, 120 m, 6 Feb. 1991 (fl), *T. McDowell & A. Stobey* 3853 (BRG, CAY, MO, NY, U, US); Potaro-Siparuni

Region, Pakaraima Mountains, upper Ireng watershed, Sukabi River, E branch to Kukunang Falls, 5°7′N, 59°57′W, 720 m, 21 Oct. 1994 (fl), *P. Mutchnick et al.* 156 (MO, NY, US); Potaro-Siparuni Region, islands at Amatuk Falls [ca. 5°18′N, 59°18′W; Potaro River], 7 Sep. 2006 (fl, fr), *K.J. Wurdack et al.* 4290 (US).

## 6-5. Neobertiera palustris (A.Rich. ex DC.) Delprete, Phytotaxa 392: 228. 2019. (Figures 4K, 30, 34–35).

(≡) *Bertiera palustris* A.Rich. ex DC., Prodr. 4: 392. Sep. 1830. [A.Rich., Mem. Fam. Rubiacées 174. Dec. 1830; reimpr. in Mem. Soc. Hist. Nat. Paris 5: 254. 1834]. – *Sipanea palustris* (A.Rich. ex DC.) J.H.Kirkbr., comb. illeg., Brittonia 49: 360. 1997; non *Sipanea palustris* Seem. (1853 [= *Limnosipanea palustris* (Seem.) Hook.f.)].

**Type:** FRENCH GUIANA. "In paludosis, rivis sylvarum, inter Conana et Ynéri, mayo et junio floreus, augusto fructif." s.d. [May–Jun. 1781–1785], *L.C.M. Richard s.n.* (holotype P [barcode P02273205]; photo-P at F).

(=) *Neobertiera glomerata* Delprete, Phytotaxa 206: 124, figs. 2–4. 2015

**Type:** FRENCH GUIANA. Commune de Montsinyery, Route D-5, trail to Bagne des Annamites, ca. 1 km from Route D-5, forest seasonally flooded, plants growing at the edge of trail, with roots near the water, herb 40–70 cm tall, erect, rooting at basal nodes, leaves chartaceous, corolla white, 4°50′3"N, 52°31′2"W, 10 m, 18 Apr. 2012 (fl), *P.G. Delprete & G. Quenette* 11860 (holotype CAY [barcode CAY214218]; isotypes GB, K, L, MG, MO, NY, P, RB, US).

*Perennial herbs or subshrubs*, 0.4–1.5 m tall, erect or basally decumbent, sparsely branched; branches densely appressed strigose; rooting at basal nodes and internodes. *Stipules* ovate to broadly ovate, 2.5–4.5 × 2.5–4.5 mm, acuminate or bifid at apex, appressed-strigose outside, basal portion antrorse-pubescent with group of 7–11 colleters on each side and glabrous at distal portion inside, margins pubescent. *Leaves* petiolate; petioles 7–30 mm long, appressed-strigose-pubescent; blades elliptic, oblanceolate to obovate, 9–14.5 × 3–5.8 cm, acute-decurrent at base, acute at apex, chartaceous, drying dark olive-green to brown above and pale olive-green to olive-green below, sparsely appressed strigose above, appressed strigose below, midvein densely strigose above
and below; secondary veins 13-15 each side of mid-
rib. *Inflorescences* glomerulate to densely cymose dur-
ing anthesis, densely cymose after anthesis; peduncles
1.3-3.5 cm long; rachis stout, appressed white-pubes-
cent, 2.5-4 cm long (excluding corollas), with 2-3 pairs
of very short lateral branches; lower lateral branches
0.4-1.2 cm long (excluding corollas); lateral branches
cymose, condensed, terminating in flowering glomer-
ules; bracteoles subtending each flower, linear-lanceolate,
2.5-3 x 0.4-0.6 mm, densely appressed-strigose. Flow-
ers 5-merous, sessile to subpedicellate; pedicels, when
present, stout, to 1 mm long. *Hypanthium* obovoid
to broadly obovoid, 0.9-1.5 x 0.9-1.5 mm, appressed
strigose. *Calyx* lobes slightly unequal, lanceolate, 2.5-
3.5 x 0.4-1.1 mm, appressed strigose outside, and with
1 oblong, 0.2-0.25 mm long, colleter at each lobe sinus.
*Corolla* white, 11.8-13.3 mm long; tube narrowly cylin-
drical, slightly wider at mouth, 8-9 mm long, sparsely
appressed-sericeous outside, puberulent inside (hairs
moniliform), with a narrow line of sparse, yellow, mon-
iliform hairs at mouth inside; lobes 5, oblong-ovate to
oblong-ellipsoid, 3.8-4.3 x 2.3-2.5 mm, round at apex,
glabrous on both sides. *Long-styled flowers*: stamens
inserted 2-3 mm from base of corolla tube; filaments
0.5-0.7 mm long; anthers narrowly sagittate, 0.8-1 x
0.1-0.2 mm; style about the same length as corolla tube
(tip of branches barely exserted), 8-9 mm long, glabrous;
style branches narrowly oblong, 1.4-1.7 mm long, dense-
ly papillose when receptive. *Short-styled flowers*: stamens
inserted 3-4 mm from base of corolla tube; filaments
0.7-1.1 mm long; anthers linear, 1.4-1.5 x 0.2-0.3 mm,
acute at both ends; style much shorter than corolla tube,
4.5-5 mm long, glabrous; style included, branches nar-
rowly oblong, 0.5-0.6 mm long, densely papillose when
receptive. *Fruits* 3-4 mm diam., sparsely strigose-seri-
ceous. *Seeds* 0.3-0.4 mm long, black, microscopically
foveolate (barely visible at 50× magnification).

Notes:–Achille Richard described *Bertiera palustris*, which was first published in Candolle’s *Prodromus* (Sep. 1830: 392), and re-published three months later in Richard’s *Mémoire sur la famille des Rubiacées* (Richard Dec. 1830: 174), based on a collection made by his father, L.C.M. Richard, in French Guiana. Steyermark (1967: 321–322) excluded this taxon from *Bertiera* and explained, “I have examined a sheet in the Paris Herbarium corre-

sponding to the type specimen from Richard's Herbarium. This specimen has crustaceous capsules and globose non-angled seeds. Furthermore, the inflorescence is regularly branched and does not have a secondly arranged dichasial flowering axis as occurs in the other species of the genus. The specimen obviously pertains to another genus, but at the present time I am unable to assign it to a definite genus. The bifid stipules and herbaceous habit are, likewise, extraneous to the genus *Bertiera*.”

Robbrecht et al. (1993: 135) in a survey of *Bertiera* excluded *B. palustris* from the genus with the comment “doubtful, see Steyermark, Mem. New York Bot. Gard. 17: 321 (1967).”

Kirkbride (1997) wrote "In my opinion S. [Sipanea] wilson-brownei and B. palustris are the same species. The epithet palustris has priority over wilson-brownei, and so must be transferred to the genus Sipanea." Accordingly, he proposed the new combination Sipanea (sect. Sipanea subsect. Nudae Steyerm.) palustris (A. Rich. ex DC.) J.H. Kirkbr. However, the binomial Sipanea palustris was already published [Sipanea palustris Seem. (Seeman, 1853) = Limnosipanea palustris (Seem.) Hook.f.], and renders Kirkbride's combination illegitimate. He cited the type of Bertiera palustris as "(holotype: P-n.v. [Field Museum of Natural History Photograph 37285])," which is the specimen with barcode P02273205 at P.

A close examination of the holotype specimen of *Bertiera palustris* at P showed that all the features correspond to the recently described *Neobertiera glomerata* Delprete. Both Steyermark and Kirkbride did not notice that the fruits of this specimen are indehiscent and fall off as dispersal units, which excludes this specimen from *Sipanea*, which has dehiscent, permanent capsules, and places it in *Neobertiera*. Therefore, Delprete (2019b) published the new combination *Neobertiera palustris* (A. Rich. ex DC.) Delprete for this species.

Several collections of this species from French Gui-
ana have been previously been identified as "*Neobertiera
gracilis*", but they are *N. palustris*. *Neobertiera gracilis*
does not occur in French Guiana.

**Distribution and ecology:** Apparently endemic to French Guiana (Figure 30). Known from a few collections in coastal forests, mostly in sites along Route National 2, the historic collection “inter Conana et Yné-ri”, and one recent collection near the village of Tampak

**Figure 34.** *Neobertiera palustris*. A. Branch inflorescence and old infructescence. B. Node with stipule. C. Dissected long-styled flower, with stamens inserted at the middle of the corolla tube. D. Dissected long-styled flower, with stamens inserted at lower portion of the corolla tube. E. Dissected short-styled flower, with stamens inserted at the upper portion of the corolla tube. F. Fruit. A-C: drawn from *Delprete & Quenette 11860* (CAY); D: drawn from *Granville 2888* (CAY); E: drawn from *Feuillet 3635* (CAY); F: drawn from *Granville 2888* (UB). Illustration by Piero Delprete (Reproduced from fig. 3 of Delprete (2015a) with permission of copyright holder Magnolia Press).
<figure><img src="image_38.png" /><figcaption>Figure 34.</figcaption></figure>
<figure><img src="image_39.png" /><figcaption>**Figure 35.** *Neobertiera palustris*. A. Flowering branch. B. Node with stipule. C. Inflorescence. D. Close-up of inflorescence. Photos taken by Gwenaël Quenette on 18 April 2012, along Trail to Bagnes des Annamites, French Guiana, where *Delprete & Quenette 11860* was collected (Reproduced from fig. 4 (as *Neobertiera glomerata* Delprete) of Delprete (2015a) with permission of copyright holder Magnolia Press).</figcaption></figure>

(Commune de Camopi), Camopi River, Upper Oyapock River Basin. Growing in the undercanopy of primary forest, often near standing or slowly running water, at 10–90 m altitude.

**Phenology:** Specimens with flower buds were collected in February, with flowers in anthesis and no fruits

in March, and with flowers in anthesis and mature fruits in May and June.

**Suggested conservation status:** **Endangered (EN).**
This species is known from several localities in French Guiana. According to personal observations made in 2012 and 2014, the populations of this species are always
very small, with less than 50 individuals. Taking into
account this species relatively small populations, Del-
prete (2015) suggested the Endangered (EN) category for
this species, following IUCN criteria. Several personal
visits to the localities of occurrence of this species were
made in 2016 and 2018, but it was not found. Therefore,
this species is maintained in this category following
IUCN criteria (IUCN 2012, 2019).

**Specimens examined (paratypes):** FRENCH
GUIANA: Commune de Montsinery, Route D-5, trail
to Bagne des Annamites, ca. 1 km from Route D-5,
4°50'3"N, 52°31'2"W, 10 m, 19 Feb. 2014 (fl buds), P.G.
*Delprete & J.C. da Silva* 12293 (CAY, L, NY); Commune
de Camopi, near the village of Tampak, base of Mon-
tagne Couronné, 3°10'3"N, 52°32'16"W, 100 m, 25 Feb.
2014 (fl buds), P.G. *Delprete et al.* 12310 (CAY, K, MG,
NY, P, US); Route Nationale 2, P.K. 58, bord de piste
forestiere, 4°35'N, 52°20'W, 25 Mar. 1986 (fl), C. *Feuillet*
3635 (CAY, MO, P); Montagne Maripa, Route Nation-
ale 2, P.K. 75, 10 May 1979 (fl, fr), *J.J. de Granville* 2888
(CAY, NY, P, U, UB); Commune de Montsinery, along
the trail between D-5 and Bagne des Annamites, 8 Jun.
2008 (fl), *J.J. de Granville & M. Rome* 17592 (CAY, K,
MO, P, NY, US).

7. PTERIDOCALYX

*Pteridocalyx* Wernham, J. Bot. 49: 317. 1911.

Wernham, J. Bot. 51: 218. 1913; Steyerm., Mem. New York Bot. Gard. 17: 234 [key to genera], 289-290 [discussion under *Neobertiera*] 1967; Delprete, Phytotaxa 391: 81-91. 2019.

Type: *Pteridocalyx appunii* Wernham.

Shrub, single-stemmed. *Stipules* basally adnate to
the petioles, ovate to narrowly triangular, acuminate or
bifid at apex, strigose outside, strigose intermixed with
colleters on medio-basal portion inside. *Leaves* opposite,
petiolate; blades elliptic-lanceolate, chartaceous; domatia
absent. *Inflorescence* terminal, cymose, many-flowered,
ultimate branches scorpiod, more evident when fully
expanded; bracts and bracteoles small, lanceolate. *Flow-
ers* distylous, 5-merous. *Hypanthium* oblong-obovoid.
*Calyx* tube absent or extremely reduced, lobes small,
linear-lanceolate to very narrowly oblanceolate; with 1
or 2 colleters at each lobe sinus; with 1 or 2 expanded,
petaloid lobes (i.e., calycophylls) per flower. *Calycophylls*
stalked, blades ovate to narrowly ovate, white, palmately-
veined. *Corolla* hypocrateriform, white during anthesis,

turning yellowish-white after anthesis; lobes left-contort-
ed, oblong-ovate to ovate, margin entire. *Pollen* 3-col-
porate; exine foveolate-reticulate, perforate. *Short-styled
flowers:* stamens included, filaments inserted at distal
portion of corolla tube; anthers linear; style included,
much shorter than corolla tube. *Long-styled flowers:* stamens included, filaments inserted near the base of
corolla tube; anthers linear; style about the same length
of the corolla tube, with tips of branches barely exsert-
ed or exserted just beyond corolla mouth. *Ovary* with a
stalked placenta; stalk inserted at the distal most portion
of the septum near the roof of the locule, with a pend-
lulous, narrowly cylindrical extension; ovules inserted
on the pendulous placental extension. *Fruit* capsular,
dehiscing septicidally, thinly woody; calycophylls often
breaking off before full maturity capsule (i.e., before seed
dispersal). *Seeds* many, horizontally inserted along the
longitudinal placenta, small, globose; testa foveolate.

**Distribution and ecology:** A genus of a single species endemic to the Upper Potaro River Basin, central Guyana.

**Morphological observations:** Delprete (2019a) described the placenta of *Pteridocalyx* as “longitudinally adnate along the central septum”. However, further examination of ovaries from several additional specimens revealed that the placenta has instead a minute stalk inserted at the distalmost portion of the septum (i.e., near the roof of the locule), and terminates with a pendulous, narrow cylindrical extension almost as long as the locule on which are inserted numerous ovules. Because of these placental features and the presence of calycophylls, *Pteridocalyx* is unique within the Sipaneeae.

*Pteridocalyx* is similar to *Neobertiera* in having cymose inflorescences with scorpiod terminal branches, corolla lobes with a basal triangular area of glandular hairs inside, and ovary with a stalked placenta; the former differs from the latter by the presence of calycophylls (vs. lacking in *Neobertiera*), placenta with a stalk inserted at the distalmost portion of the septum, terminating with a pendulous cylindrical extension (vs. with a stalk inserted at the middle of the septum, terminating in a hemi-ellipsoid, oblong-ellipsoid or hemi-elliptic extension), oblong-ellipsoid to narrowly oblong, 6-14 × 2-3 mm, septicidal capsules (vs. globose, 1.2-4 mm diam., indehiscent fruits), and globose seeds (vs. tetrahedral). No phylogenetic studies are available to test the relationships between *Pteridocalyx* and members of the Sipaneeae (study in progress).

*Pteridocalyx* is also similar to *Dendrosipanea* by
being shrubs or treelets 1-2 m tall, with ovate-triangular
stipules acute or bifid at apex, terminal inflorescences, 5-merous, distylous flowers, hypocrateriform corollas with lobes left-contorted, and septicidal capsules. The former differs from the latter by the presence of 1-2 calycophylls per flower (vs. calycophylls absent in *Dendro-sipanea*); corolla tube pale brown or white and brownish at medial portion, lobes pink, with a basal white triangle of glandular hairs on adaxial side at base (the rest glabrous) forming a white star-like pattern with the 5 lobes (Figure 37D), at mouth (vs. corolla white to cream-white, sometimes tube green, lobes without a basal white triangle of glandular hairs on adaxial side); placenta with a stalk inserted at the distal most portion of the septum, with a pendulous, narrowly cylindrical extension almost as long as the locule, and the ovules inserted on the cylindrical extension (vs. placenta with a fleshy extension adnate along the entire length of the septum, and ovules irregularly peltate on the longitudinal placental extension); capsules oblong-ellipsoid to narrowly oblong, crustaceous (vs. oblong-turbinate, thinly woody); seeds globose, testa foveolate (vs. 3-5-angular or irregularly-shaped, testa reticulate).

7-1. *Pteridocalyx appunii* Wernham, J. Bot. 49: 318. 1911. (Figures 4L, 5A-B, 36-39).

Type: GUYANA. "Demerara" [Potaro-Siparuni Region, Upper Potaro River], Kaieteur Falls [5°10'N, 59°28'W], [day & month unknown] 1872 (fl, fr), C. Appun s.n. (holotype BM [barcode BM000099002]).

(=) *Pteridocalyx minor* Wernham, J. Bot. 51: 218. 1913.

Type: GUYANA. Potaro-Siparuni Region, Potaro River, "Sheenabowa" [Chenapou Village, 4°55'N, 59°34'W, ca. 470 m], Sep.-Oct. 1881 (fl), G.S. Jenman 1282 (holotype K [barcode K000173940]).

*Shrub* 1-2 m tall, with a central, thin stem, sparsely branched towards the top; young branches reddish, older stems and branches pale gray; young branches densely appressed strigose. *Stipules* ovate to narrowly triangular, 2.2-3 × 4-6 mm, acuminate or bifid at apex, sometimes entire at early stage and becoming bifid at apex at

later stage, apical teeth (when present) to 1.5 mm long, appressed strigose-pubescent outside, strigose-pubescent intermixed with 15-17 colleters at medio-basal portion and glabrous distally inside, margins short-strigose. *Leaves* with petioles 1.5-5 mm long, appressed-strigose-pubescent; blades elliptic to elliptic-lanceolate, 9-16 × 3-5.5 cm, acute to decurrent at base, acuminate at apex, chartaceous, shiny and dark green above and dull pale green below when fresh, pale brown above and olive-green below when dry, sparsely pubescent above, pubescent below, midrib and secondary veins sericeous-pubescent below; secondary veins 10-12 on each side of midrib. *Inflorescence* cymose when young (up to anthesis), expanding during and after anthesis, appressed-pubescent, with 1-3 pairs of lateral branches; lateral branches 2-6 cm long when fully expanded, dichotomously divided, with one flower at the base of each bifurcation, ultimate branches scorpiod, more evident when fully expanded; bracts subtending branchlets lanceolate, 2-4 × 0.5-1 mm, appressed-pubescent; bracteoles subtending flowers 1-2(-3), linear-lanceolate, 1-3 × 0.2-0.7 mm, appressed-pubescent. *Flowers* sessile or rarely sub-pedicellate, secundiflorous. *Hypanthium* oblong-obovoid, 1.5-2 × 1.1-1.3 mm, acute-decurrent at base, round to truncate at apex, appressed strigose-pubescent. *Calyx* tube absent (lobes free) or extremely reduced, to 0.2 mm long; smaller lobes white when fresh, cream-white to beige when dry, linear to linear-lanceolate, 2-4.5 × 0.4-0.7 mm long, sparsely appressed strigose-pubescent outside, margins hispidulous; with 1 or 2 colleters in each calycine sinus, oblong, 0.15-0.2 mm long. *Calycophylls* 1-2 per flower, pure white when fresh, cream-white to beige when dry, varying in size and shape within the same calyx; smaller calycophylls without a distinguishable stalk, linear, narrowly oblanceolate to narrowly oblong-oblanceolate, 4.5-9.5 × 0.8-1.5 mm, acute at apex, sparsely hispidulous along veins and sometimes at margins, with 3 veins (including midrib); larger ones with stalks 2-5 mm long, blades narrowly elliptic, elliptic to ovate, 4-13 × 5-9 mm, acute, obtuse to round at base, acute to obtuse at apex, sparsely hispidulous along veins and sometimes at margins, with 5-7 veins (including midrib). (Figs. 37, 39). *Corolla* 14-19 mm long, tube pale brown or white and brownish at medial portion, lobes pink, with a basal white triangle of glandular hairs inside; tube narrowly cylindrical, slightly wider at

**Figure 36.** *Pteridocalyx appunii.* A. Branch with inflorescence. B-C. Stipule shapes present on the same branch. D. Flower in anthesis. E. Dissected long-styled flower, with stamens inserted at basal portion of corolla tube. F. Dissected short-styled flower, with stamens inserted at distal portion of corolla tube. G. Mature capsule. A-E: drawn from *P.G. Delprete & P. Benjamin 12813* (CAY); F: drawn from *P.G. Delprete & P. Benjamin 12824* (CAY); G: drawn from *P.G. Delprete & P. Benjamin 12836* (CAY). Illustration by Piero Delprete (Reproduced from fig. 1 of Delprete (2019a) with permission of copyright holder Magnolia Press).
<figure><img src="image_40.png" /><figcaption>Figure 36.</figcaption></figure>
<figure><img src="image_41.png" /><figcaption>**Figure 37.** *Pteridocalyx appunii*. A. Habit. B. Inflorescences, side view. C. Inflorescence, top view. D. Corolla, top view. E. Inflorescence and infructescence from the same plant. Photos taken by Piero Delprete, Upper Potaro River, near Chenapou Village, Guyana, where *Delprete & Benjamin 12813* was collected. (Reproduced from fig. 2 of Delprete (2019a) with permission of copyright holder Magnolia Press).</figcaption></figure>
mouth, 10.5–13.5 mm long, 0.7–0.8 mm wide for most
length, 1.3–1.5 mm wide at mouth, appressed strigose-
sericeous outside, sparsely glandular-puberulent inside;
lobes oblong-ovate to ovate, 3.5–5.5 × 1.7–3 mm, obtuse
to acute at apex, glabrous outside, with a basal triangu-
lar area of glandular hairs inside (Figure 37B & D), the
rest glabrous. *Long-styled flowers*: stamens inserted at
3–4 mm from base of corolla tube; filaments 0.5–0.7
mm long; anthers linear, 1.8–2 × 0.2–0.3 mm, acute at
both ends; style about the same length as corolla tube
(tips of branches exserted) or barely exserted, 11–15 mm
long, glabrous; style branches linear, 1–1.2 mm long,
densely papillose when receptive. *Short-styled flowers*:
stamens inserted at 5.5–6 mm below corolla mouth;
filaments 0.3–0.5 mm long; anthers linear, 2.5–2.7 ×
0.2–0.3 mm, acute at both ends; style included, 6–6.5
mm long, glabrous; branches narrowly lanceolate, 1.5
mm long, acute, densely papillose when receptive. *Cap-
sules* oblong-ellipsoid to narrowly oblong, 6–14 × 2–3

mm, acute-decurrent at base, round to truncate at apex,
crustaceous, sparsely strigose-sericeous, pale green when
young, turning pale brown when dry. *Seeds* globose,
0.3–0.4 mm in diam., foveolate.

**Morphological observations:** Wernham (1911: 318) described the calyx tube of *P. appunii* Wernham as “3–4 mm long”; however, in the type specimen the calyx tube is either absent (lobes free) or extremely reduced, to 0.2 mm long, as in all other specimens studied.

Wernham (1913: 218) separated *Pteridocalyx minor*
Wernham from *P. appunii* by the stipules entire (vs.
divided at apex in *P. appunii*), calycophyll blades 4 × 5
mm (vs. 11–13 × 7–9 mm), and corolla tubes 16 mm long
(vs. 10–11 mm long). However, although he described *P.
minor* as having “corolla tube 16 mm long, 1 mm wide,
lobes 5.5 × 2 mm”, the holotype specimen at K has
corolla tube 11 mm long and lobes 3.5 × 1.5 mm, which
largely correspond to the measurements supplied by him

<figure><img src="image_42.png" /><figcaption>**Figure 38.** Distribution of *Pteridocalyx appunii*.</figcaption></figure>
for *P. appunii*, i.e., “corolla tube 10–11 mm long, lobes 4–5 × 2 mm.” (Wernham, 1911: 318).

In addition, recent personal observation of herbarium specimens and natural populations near the Chenapou Village, Upper Potaro River, Guyana, demonstrated that the stipules can be either entire or bifid at the apex on the same branch, often being entire when young and becoming bifid at an older stage.

Finally, petaloid calyx lobes (calycophylls) can be one or two per flower, sometimes with one or two additional less expanded lobes, and can vary in shape and size within the same inflorescence (Figures 37, 39), with blades ranging from narrowly elliptic, elliptic to ovate and 4–13 × 5–9 mm within the same inflorescence. The largest ones are found on the basal flowers and the smaller ones on the distal flowers of the same inflorescence branch. All morphological characters used by Wernham to distinguish *P. minor* from *P. appunii* are widely overlapping, and the two taxa are conspecific.

**Distribution and ecology:** Endemic to the Upper Potaro River basin, above and below Kaieteur Falls (Figure 38), central Guyana, at 200–500 m altitude. *Pteridocalyx appunii* is a shade-loving species, found in the undercanopy of tall forests, in alluvial or lateritic soils, either seasonally inundated, e.g., mora forest (dominated by *Mora excelsa* Benth.), or not inundated although constantly humid, e.g., bulletwood forest (dominated by *Manilkara bidentata* (A. DC.) A. Chev.).

**Phenology:** Flowering specimens were collected in June, September, October and November; the sole collection with mature capsules is the type of *Pteridocalyx appunii*, which was collected at an unknown date in 1872.

**Conservation status:** Endangered (EN). Delprete (2019a) wrote "Until recently *Pteridocalyx* was a genus known from only a few historical collections made in 1872, 1881 and 1907, preserved in BM and K. A gathering of 1992 remained unidentified until recently. All these collections are from the Upper Potaro River basin, above and below Kaieteur Falls, located within the Kaieteur National Park, central Guyana, which is an area botanically poorly collected. During my field expedition of 2016, I observed several populations of approximately 100–200 individuals along the trails bordering the Potaro River and its affluents, near Chenapou Village, in forests seasonally inundated or never inundated. Because of the difficult terrain in the forests away from the trails, the only populations that I examined are those bordering the rivers. Therefore, the AOO (Area of Occupancy) of this taxon is difficult to estimate, and the EOO (Extent of Occurrence) is less than 20,000 km². Although this monospecific genus is endemic to the Upper Potaro River basin, no imminent threats were detected in 2016, as the forests bordering the rivers surrounding Chenapou Village are fairly well preserved, except for selective logging of certain large trees, even though they do not have any official conservation status. In addition, the only way to access the area surrounding the Potaro River above Kaieteur National Park is by canoe, which renders impossible any considerable timber exploitation on this portion of the river, aside from sparse extraction by the local indigenous community, including the forests surrounding the

<figure><img src="image_43.png" /><figcaption>**Figure 39.** *Pteridocalyx appunii*. Calycophyll variation within the same inflorescence branch. **A.** Basal flower. **B–D.** Intermediate flowers, from base towards distal portion. **E.** Distal flower. **A–E:** drawn from Delprete & Benjamin *12813* (CAY). Illustration by Piero Delprete (Reproduced from fig. 3 of Delprete (2019a) with permission of copyright holder Magnolia Press).</figcaption></figure>
Chenapou Village. In conclusion, because of the small populations observed and the relatively restricted area of occurrence, this taxon is here treated as Vulnerable (VU) based on criterion D1 of IUCN (2012, 2017)."

As a result of a renewed analysis regarding the conservation status, here presented, it should be emphasized that the collection localities of this species are within a small area, including the most recent collections. Its Extent of Occurrence (EOO) is here calculated to be of ca. 174 km², which is slightly larger than criteria B1 for Critically Endangered (CE) and falls within the Endangered (EN) category. Therefore, although Delprete's (2019a) discussion and conservation status is recent, considering the EOO here calculated, and the vicinity of Chenapou Village, which impacts the populations of the species, *Pteridocalyx appunii* is here re-classified as Endangered (EN) following IUCN (2012, 2019) criteria.

**Specimens examined:** GUYANA: Potaro-Siparuni Region, Potaro River, Holmia [ca. 4°58′N, 59°35′W], Nov. 1907 (fl), A.W. Bartlett 8763 (K); Potaro-Siparuni Region (Region 8), Upper Potaro River, Chenapou Village, trail from village to Karisparu Landing, tall Mora forest seasonally inundated, on alluvial soil, 4°58′40″N, 59°34′14″W, 450 m, 12 Jun. 2016 (fl), P.G. Delprete & P. Benjamin 12813 (BRG, CAY, K, MG, MO, NY, US), 12814 (B, BBS, BR, BRG, L, P, US), 12815 (BRG, CAY, K, GB, NY, P, US); Potaro-Siparuni Region, Upper Potaro River, Chenapou Creek (affluent of Potaro River), Chenapou Creek Mouth, secondary forest, on lateritic soil, 4°58′25″N, 59°34′47″W, 420 m, 13 Jun. 2016 (fl), P.G. Delprete & P. Benjamin 12824 (BRG, CAY, GB, L, MG, MO, NY, RB, W); Potaro-Siparuni Region, Upper Potaro River, Chenapou Village, Paramakatoi trail, parallel to Chenapou Creek, going from Chenapou Village to Paramakatoi Village, bulletwood forest on lateritic soil, never inundated, 4°57′26″N, 59°35′1″W, 440 m, 14 Jun. 2016 (imm fr), P.G. Delprete & P. Benjamin 12836 (BRG, CAY, G, MO, NY, RB, US); Potaro-Siparuni Region, Potaro River, left margin on portage trail on second set of rapids above Chenapou, 5°0′40″N, 59°38′22″W, 430–490 m, 18 Mar. 2014 (fl), F.A. Michelangeli et al. 2307 (MO, NY); Potaro-Siparuni Region, Micobe, mixed forest on brown and white sand, shady places, 5°20′N, 59°30′W, 200 m, 20 Oct. 1992 (fl), S. Tiwari & A. Mengarini 487 (BRG [3 sheets], CAY).

## 8. SIPANEA

*Sipanea* Aubl., Hist. Pl. Guiane 1: 147. 1775 K.Schum. in Martius et al., Fl. Brasil. 6(6): 247–252, fig. 122. 1889

Wernham, J. Bot. 55: 171–175. 1917; Steyerm., Mem. New York Bot. Gard. 17: 261–282. 1967; Steyerm. in Lasser and Steyermark, Fl. Venezuela 9(1): 357–384, figs. 58–60. 1974; Delprete and Steyerm. in Steyermark et al., Fl. Venez. Guayana 8: 828–832, fig. 640–647. 2004; Delprete in J.A. Rizzo, Fl. Goiás Tocantins 40(3): 1140–1152, fig. 106. 2010.

Type: *Sipanea pratensis* Aubl.

*Virecta* L.f., Suppl. 17. 1781

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

Non *Virecta* Smith in A.Rees, nom. superfl. (= *Virectaria* Bremek.).

*Ptychodea* Willd. ex Cham. & Schlechtdl., Linnaea 4: 168. 1829

Type: the two specific binomials published by Chamisso and Schlechtendal (1829: 128) are illegitimate, because are cited in synonymy under two validly published names in *Sipanea*.

From Chamisso & Schlechtendal (1829: 168): "Addimus synonyma Herbarii Willdenowianii ad *Sipaneas* Humbodtianas: *Sipanea dichotoma* H.B.Kh. 3. p. 110. est *Ptychodea pedunculata* Hb. Willd. n. 4123. – *Sipanea glomerata* H.B.Kh. l.c. est *Ptychodea sessiliflora* Hb. Willd. n. 4123."

**Note:** The citation "Hb. Willd. N. 4123" represents the genus number in the Willdenow Herbarium, where "4123" in the number of *Sipanea*, and is cited for both *P. pedunculata* and *P. sessiliflora*.

*Herbs*, erect, trailing, prostrate or decumbent, often rooting at basal nodes, sometimes mat-forming, rarely floating on water in inundated areas (*S. biflora*), or *sub-shrubs* (stems woody at base), erect or decumbent. *Stipules* free or basally adnate to the petioles, sometimes sheathing at base, truncate to broadly transversally elliptic, broadly to narrowly triangular, suborbicular, subulate, linear, rarely setaceous or reduced to a line between the petioles; adaxial side with a few to several colleters inserted on each side or irregularly spread near the base or at median portion or just below the margin, or inserted along the margin. *Leaves* opposite, sessile, subsessile to short-petiolate, rarely with axillary brachyblasts with numerous smaller leaves (e.g., *S. galioides*); blades ovate, narrowly ovate, rhombic-
ovate, elliptic, oblong-elliptic, oblong-lanceolate, linear-lanceolate to linear, membranaceous, chartaceous or thinly subcoriaceous; domatia absent. *Inflorescences* terminal, sometimes appearing axillary as a result of the reiteration of an axillary bud subtending the inflorescence, paniculate, cymose or corymbose, usually not frondose or rarely frondose, pauci- to multi-florous or uniflorous; in several species (e.g., *S. pratensis*, *S. glomerata*, *S. hispida*) inflorescences initially condensed, subcapitate, and in later stages the rachis elongates and the secondary branches expand from subcapitate to scorpiod at fruit maturity. *Bracteoles* subtending the flowers or opposing the flowers, commonly present or rarely absent. *Flowers* distylous, (4-5)5(-6)-merous. *Hypanthium* globose, subglobose, ovoid, obovoid, narrowly obovoid to broadly turbinate, glabrous or pubescent, sometimes with hairs basally tuberculate (e.g., *S. wilson-brownei*, *S. ovalifolia*, *S. prancei*). *Calyx* persistent; lobes free or rarely shallowly connate at base, oblong-lanceolate, lanceolate, linear to setaceous; with 1 or 2 colleters in each calycine sinus. *Calycophylls* absent. *Corolla* hypocrateriform, narrowly infundibuliform, narrowly campanulate to campanulate, white, pinkish-white, cream-white, pink, or pale purple, sometimes the lobes with a white basal triangular area forming a star shape at mouth with the five lobes; tube glabrous, puberulent, short-pubescent or hirsute, or lower half glabrous and distal portion pubescent outside, glabrous, or puberulent or pubescent at stamens insertion and the rest glabrous, or puberulent or pubescent throughout inside; corolla mouth glabrous or with sparse hairs (Sect. *Nudae*), or with a pubescent ring of white hairs (Sect. *Virecta*), or with a pubescent ring of yellow hairs (Sect. *Sipanea*) at orifice inside; lobes left-contorted, round, broadly ovate, obovate, elliptic-ovate or lanceolate, sometimes slightly asymmetrical, in several species (e.g., *S. galioides*, *S. praten sis*) varying in size and shape within the same population and/or on the same plant, round, obtuse or acute, sometimes mucronate, at apex. *Pollen* 3-colporate; exine foveolate-reticulate or finely foveolate, perforate. *Long-styled flowers:* stamens included or exserted beyond the corolla mouth; filaments inserted near the base, at about the middle, or at medio-distal portion of the corolla tube, anthers narrowly oblong-elliptic to linear; style about the same length as the corolla tube with tips of branches barely exserted or exserted just beyond corolla mouth. *Short-styled flowers:* stamens included, inserted at the distal or medio-distal portion of the corolla tube, anthers narrowly oblong-elliptic, very narrowly lanceolate or linear; style included, much shorter than corolla tube. *Ovary* with stalked placenta;

stalk inserted at median or medio-basal portion of the septum, with a narrowly oblong-elliptic extension (in certain species the stalk is very short, appearing as if the ovules are inserted on the central septum); ovules many per locule, inserted on the placental extension. *Fruit* capsular, dehiscing loculicidally, crowned by the permanent calyx, thinly woody to crustaceous, glabrous or pubescent, rarely with hairs basally tuberculate (e.g., *S. wilson-brownei*, *S. ovalifolia*). *Seeds* many per locule, perpendicular to the placental extension or acropetally imbricate, minute, irregularly globose, subglobose, ellipsoidal, polyhedral, tetrahedral, rhomboidal, or trigonous, faintly to strongly angular; testa reticulate-foveolate, smooth or rugulose-verrucose.

**Distribution:** A genus of 17 species, occurring in Nicaragua, Costa Rica, Panama, Colombia, Venezuela, the Guianas, Brazil, Ecuador, Peru, Bolivia and Paraguay.

## History of *Sipanea* sections

Schumann (1889: 249; 1891: 38-39) divided *Sipanea* into two sections: 1) Sect. *Eusipanea* K.Schum., characterized by multiflorous or rarely uniflorous inflorescences with elongating cincinni, and crustaceous-subwoody capsules, where he included *S. glomerata* and *S. praten sis*; 2) Sect. *Panisea* K.Schum., with 1-2(-3)-florous inflorescences, and papyraceous capsules, where he included only *S. biflora*.

Steyermark (1967, 1974) divided *Sipanea* into two sections:

1) Sect. *Sipanea*, characterized by “Stipules prominent, tapering to a long triangular or subulate-setaceous apex; mainly erect to ascending herbs with relatively stout firm stems, large leaf-blades, and usually 3-20- or more-flowered terminal or rarely axillary, once or twice dichasial branched inflorescences with monochasial branches; capsule usually thick-walled, difficult to break.” He subdivided this section into two subsections: 1a) Subsect. *Nudae*, with “Orifice of corolla mouth glabrous or at least not yellow villous” (*S. ayangannensis*, *S. stahelii*, *S. wilson-brownei*), and 1b) Subsect. *Luteovillosae*, with “Orifice of corolla within densely yellow-villous” (*S. glaberrima*, *S. glomerata*, *S. pratensis*, *S. hispida*, *S. glabrata*, *S. pubinoda*, *S. galioides*).

2) Sect. *Virecta*, characterized by “Stipules low, shallow, rounded, or broadly triangular, at most acute at apex, or obsolete; usually creeping, prostrate herbs with slender weak rooting stems, often prostrate vegetative stems, or with ascending to erect flowering axes; leaf blades relatively small and thin; inflorescences axillary or terminal, usually 1-3(-5)-flowered; capsules thin-
walled, easily broken or splitting.” He subdivided this section into two subsections: 2a) Subsect. *Cryptotricha*, with “Orifice of corolla within white-villous, the hair mass not projecting; one colleter at sinus between bases of calyx lobes; inflorescence mainly 2-3-flowered, axillary or terminal” (S. micrantha, S. ovalifolia, S. gleasonii, S. biflora, S. cowanii [=S. biflora]); and 2b) Subsect. *Phanerotricha*, with “Orifice of corolla within densely yellow-villous, the mass of hairs projecting beyond orifice; two squamellae [i.e., colleters] usually present at sinus of calyx-lobes; inflorescence mainly 1-2-flowered, falsely terminal” (S. acinifolia [= S. veris], S. veris)

In the molecular phylogenetic study of the tribe Sipaneeae of Delprete and Cortés B. (2004), nine taxa of *Sipanea* were included. The sections and subsections of *Sipanea* proposed by Steyermark (1967, 1974) were not supported, as species with different habits (i.e., prostrate or erect) were found side by side on the same monophyletic clades. On the other hand, species with similar corolla mouth vestiture were found grouping together, indicating a possible sinapomorphic character for monophyletic sections within the genus. For example, in the strict consensus tree from the combined trnL-F and ITS analysis, S. stahelii and S. wilson-brownei, both with glabrous corolla mouth, were found as sister taxa of one clade, although with low support (bootstrap value (BV) = 58). On the same tree, S. biflora, the sole species with white-pubescent corolla mouth included in the analysis, was found at the base of the sister clade, which in turn is a sister clade with a strongly supported clade (BV = 97) with S. pratensis, S. glomerata, S. galioides, S. acinifolia (= S. veris), S. hispida and S. veris, all with yellow-pubescent corolla mouth. However, the number of species sampled in *Sipanea* was about half the species recognized in the genus, and the phylogenies obtained could not be treated as conclusive. Therefore, further analyses with a larger species sampling is necessary to confirm these groupings.

Seyermark’s (1967, 1974) *Sipanea* sections were used in his dichotomous keys, which are practical for identifying species within the genus. However, in this case I prefer to recognize sections that are at least indicated to be monophyletic in the molecular phylogenetic study of Delprete and Cortés B. (2004). Therefore, in the present treatment three sections are recognized within *Sipanea*, which are characterized by the internal vestiture at the corolla mouth: 1) Section *Sipanea*, with a ring of yellow hairs; 2) Section *Virecta*, with a ring of white hairs; 3) Section *Nudae*, glabrous or with sparse white hairs. In the present treatment, species are organized according to these three sections, and within each section they are arranged in alphabetical order.

*Inflorescence expansion after anthesis*

After detailed study of numerous populations of *Sipanea glomerata*, *S. hispida*, and *S. pratensis* var. *dichotoma*, it was observed that the inflorescences are initially condensed, subcapitate (i.e., extremely shortly-branched), and later, during and after anthesis the rachis elongates and the secondary branches also elongate from subcapitate to scorpiod. Therefore, this inflorescence expansion is generally treated as a variation correlated with different stages of development, and might be observed even on the same individual. However, in *S. pratensis* two varieties with allopatric distributions and different fruit shapes are here recognized: var. *pratensis*, with inflorescences initially congested-cymose, appearing fasciculate, secondary branches that do not elongate in later stages, flowers and fruits congested-cymose, appearing fasciculate, without a subsecund monochasium of dichasial appearance, infructescences remain congested-cymose, appearing fasciculate, and capsules about as long as broad, 4-4.5 x 3-4.5 mm, strongly costate; var. *dichotoma*, with inflorescence axes that elongate to various lengths during and after anthesis, producing a subsecund monochasium of dichasial appearance, infructescences show some degree of expansion, often with scorpioid secondary branches (See *S. pratensis* for further discussion), and capsules longer than broad, (3-)4.5-8 x 1.5-3 mm, smooth or faintly costate.

*Sectional classification of *Sipanea*.*

**8A. *Sipanea* section *Sipanea***

Type: *S. pratensis* Aubl. [Seyermark (1967: 268) wrote that the type of *S. sect. Sipanea* is *S. wilson-brownei* Cowan, but the type of this section is *S. pratensis*]

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

Type: *Sipanea pratensis* Aubl.

*Sipanea* sect. *Sipanea* subsect. *Luteovillosae* Steyermark., Mem. New York Bot. Gard. 17: 261, 270. 1967.

Type: *Sipanea pratensis* Aubl.

*Sipanea* sect. *Virecta* subsect. *Phanerotricha* Steyermark., Mem. New York Bot. Gard. 17: 263, 266. 1967.

Type: *Sipanea acinifolia* Spruce ex Sprague (= S. veris S.Moore).
4.3 × 0.2-0.9(-1.2) cm, 5-8 times longer than wide; leaf secondary veins 2-3(-4) on each side of midrib ................................. 1. S. galioides

5. Stems commonly without axillary brachyblasts; leaf blades ovate, broadly elliptic, oblong-lanceolate, ovate-lanceolate, oblong-elliptic, lanceolate to narrowly oblong-lanceolate, 1.2-9.5 × 0.5-5 cm, 2-3 times longer than wide; leaf secondary veins 3-11 on each side of midrib..................................6

6. Stems densely hirsute or villous; leaf blades sparsely to densely strigose above; corollas (8.5-)9.5-16.5 mm long, tube (6-)7-12.5 mm long, lobes 2.5-5 × 1.5-3 mm ................................. 4. S. hispida

7. Stems sparsely to densely appressed or spreading pubescent or sericeous; leaf blades densely sparsely or sparsely to densely appressed pubescent to glabrate above; corollas 14-28 mm long, tube (8-)10-17 mm long, lobes (2.5-)4-11 × (2-)2.5 × 7 mm....................................................................7

8. Leaf blades densely sparsely or sparsely to densely appressed pubescent to glabrate above, densely sericeous or sparsely appressed pubescent or glabrous below, midrib and secondary veins appressed-strigose below; inflorescences 5-8-flowered, a single head subtended by two leaf-like bracts, or made of three pedunculate heads; hypanthium narrowly obovoid, 2.5-3 × 2-2.5 mm, densely sericeous; calyx lobes 7-13 mm long ............3. S. glomerata

9. Leaf blades appressed pubescent to densely short- to long-pubescent (hairs 0.3-2 mm long) above, densely appressed pubescent or villous or short- to long-hirsute (hairs 0.3-2 mm long) below; inflorescences 4-8-flowered, congested-cymose (appearing fasciculate), rarely with secondary branches to 4 mm long, sometimes subtended by 2 or 4 leaf-like bracts (var. pratensis), or 4-37-flowered, evidently branched during anthesis (var. dichotoma); hypanthium turbinate to narrowly obovoid, 0.7-1.5 × 0.5-0.9 mm, appressed pubescent to sericeous; calyx lobes 2.3-6 mm long ....................................................................................5. S. pratensis

10. Stems often with axillary brachyblasts bearing numerous smaller leaves; leaf blades linear-lanceolate to linear, 5-8 times longer than wide........................................1. S. galioides

11. Stems commonly without axillary brachyblasts; leaf blades ovate, ovate-lanceolate to rhombic-ovate, elliptic, oblong-elliptic, oblong-lanceolate, lanceolate, narrowly lanceolate to narrowly oblong-elliptic, 2-3 times longer than wide...9

12. Perennial herb or subshrub, terrestrial or epipetric; stems often woody at basal internodes; stipules 2-6 × 0.6-1.5 mm, appressed pubescent to sericeous outside; leaf blades 1.5-7.5 × 0.5-2.5 cm, chartaceous, secondary veins 3-8 on each side of midrib; inflorescences 4-8-flowered, congested-cymose (appearing fasciculate, rarely with secondary branches to 4 mm long), sometimes subtended by 2 or 4 leaf-like bracts (var. pratensis), or 4-37-flowered, evidently branched during anthesis (var. dichotoma) ................................. 5. S. pratensis

13. Delicate herbs, prostrate or trailing, rarely decumbent, often forming thick patches; stems thin, herbaceous; stipules 0.5 × 0.5-0.8 mm, strigulose or glabrate outside; leaf blades 0.5-2.6 × 0.2-1.7 cm, membranaceous to papyraceous, secondary veins 2-3 on each side of midrib; inflorescences 1-2-flowered..................................................6. S. veris

14. Stipules setaceous, 0.1 mm wide .................................12. S. setacea

15. Stipules broadly transversely elliptic to truncate, ovate, broadly ovate, broadly triangular, deltoid to triangular-lanceolate, 0.4-2 mm wide....................................................................11

16. Leaf blades 5-17 × 3-7 cm; corolla campanulate, white, 4-6.5 mm long, tube obconical, 2.5-3 mm long, lobes 1.5-3 × 1-1.5 mm ..................................................10. S. micrantha

17. Leaf blades (0.4-)1.2-5 × (0.35-)0.8-3 cm; corolla hypocrateriform, narrowly infundibuliform to narrowly campanulate, white, pinkish-white, pink, deep pink to pale purple, 4.3-21(-29) mm long, tube caniculate or cylindrical to narrowly obconical, 2.5-13(-15) mm long, lobes 1.5-8.5(-14) × 1.5-5.5(-7) mm ..................................................12

18. Hypanthia and capsules with hairs conspicuously tuberculate at base....................................................................................13

19. Hypanthia and capsules with hairs not tuberculate at base (slightly inflated at base in S. prancei)..................................14

20. Hypanthium 1.3-1.7 × 1-1.5 mm; calyx lobes setaceous, 4-4.5 × 0.1-0.2 mm; corollas narrowly infundibuliform, 12-14 mm long, tube 7.5-9.5 mm long, lobes 4-8 × 2.5 × 3 mm; inflorescences 2-3-flowered..................9. S. gleasonii

21. Hypanthium 0.7-0.8 × 0.7-0.8 mm; calyx lobes linear to linear-lanceolate, 2.5-4 × (0.1-)0.2-0.4 mm; corollas narrowly campanulate, 4.3-5.3 mm long, tube 2.8-3.6 mm long, lobes 1.5-1.8 × 1.2-1.3 mm; inflorescences 3-5-flowered ..................................................................................11. S. ovalifolia

22. Leaf blades ovate, rhombic-ovate or rarely oblong, (0.4-)1.2-5 × (0.35-)0.8-2.8 cm; stipules broadly triangular, 0.3-1.2(-2) × 0.7-0.9(-1.2) mm, glabrous or sparsely strigillose outside; leaf secondary veins (3-)4-6 on each side of midrib; calyx lobes linear-lanceolate to setaceous, 1-5 × 0.2-0.6 mm; corolla tube 5-13(-15) mm long, lobes 3-8.5(14) × 2-5.5(-7) mm; capsules subglobose, 3-5 × 3-5 mm ....................................................................................7. S. biflora

23. Leaf blades ovate-lanceolate, elliptic to elliptic-oblong, 0.7-2.3 × 0.4-0.9 cm; stipules deltoid to triangular-lanceolate, 2-3 × 0.5-2 mm, densely strigose outside; leaf secondary veins 2-4 on each side of midrib; calyx lobes linear-lanceolate, 4.5-5.5 × 0.5-0.7 mm; corollas tube 10-13 mm long, lobes 5.5-6.5 × 4-5 mm; capsules narrowly oblong-elliptic, 8-9 × 2-3 mm..................................8. S. carrenoi

24. Stipules reduced to a line between the petioles of the same node or very shallowly transversally ovate, 0.5-3 mm long, sometimes with a central seta..................................................16
15. Stipules deltoid, narrowly triangular to almost linear, or subulate (base triangular and medio-distal portion narrowly triangular to linear in *S. ayangannensis*), (2–3.5–5 mm long, without central seta.................................................. 17

16. Herbs or subshrubs, to 50 cm tall, terrestrial; stipules reduced to line between petioles and crowned by 5-7 colleters; leaf blades ovate to elliptic, 2-7.5 x 1-4.5 cm, secondary veins 5-7 on each side of midrib; calyx lobes linear to setaceous, 2.2-3 x 0.1-0.2 mm; corolla 8.5-11 mm long, tube 5.7-7.5 mm long, lobes ovate, 2.7-3.5 x 1.4-1.5 mm; capsules globose, 2.5-3 x 2.5-3 mm....................
.................................................................................. 14. *S. prancei*

16. Herbs, 15-20 cm tall, terrestrial or epipetric; stipules transversely oblong to reduced to a line between petioles; leaf blades narrowly obovate, 9-12 x 2.5-4.3 cm, secondary veins 8-9 on each side of midrib; calyx lobes very narrowly elliptic to almost linear, 7-10 x 0.6-0.8 mm; corolla 14-16.5 mm long, tube 9-10.5 mm long, lobes obovate, 5-6 x 3-4 mm; capsules globose to very broadly ellipsoid, 5.5-6 x 4-5.5 mm .................................................. 15. *S. saxicola*

17(15). Bracteoles opposing each flower linear to acicular, 0.8-2.5 x 0.1-0.3 mm; calyx lobes linear to acicular, 1.5-3 x 0.2-0.5 mm .................................................................... 16. *S. stahelii*

17. Bracteoles opposing each flower elliptic, narrowly elliptic, lanceolate, narrowly lanceolate to linear-lanceolate, 3.5-7 x 0.5-1.7 mm; calyx lobes oblong-lanceolate, oblanceolate, lanceolate to linear, 3.5-8 x 0.3-1.7 mm ................................. 18

18. Stipules subulate, base triangular and medio-distal portion narrowly triangular to linear, entire; leaf blades lanceolate-elliptic, 4-5.5 x 1.7-2.2 cm, acute at apex, secondary veins 5-7 on each side of midrib; corolla tube 14.5 mm long, lobes 10.4 x 4.3 mm; hypanthium with hairs not basally tuberculate.................................................. 13. *S. ayangannensis*

18. Stipules deltoid or narrowly triangular to almost linear, commonly entire or sometimes becoming bifid at apex at later stage, apical teeth to 2 mm long; leaf blades elliptic to elliptic-lanceolate, 2.5-10.5(-14.5) x 1-4(-5.5) cm, acuminate at apex, secondary veins 6-10 on each side of midrib; corolla tube (7-)9.5-15 mm long, lobes 4-9.5(-14) x 2-6(-8) mm; hypanthium with hairs basally tuberculate....
.................................................................................. 17. *S. wilson-brownei*

**8A. *Sipanea* section *Sipanea* Aubl.**

Type: *Sipanea pratensis* Aubl. [Steyermark (1967: 268)]

wrote that the type of *S. sect. Sipanea* is *S. wilson-brow-
*nei* Cowan, but the type of this section is *S. pratensis*]

8-1. *Sipanea galioides* Wernham, J. Bot. 55: 172. 1917.
(Figures 5C, 40-41).

Type: GUYANA. Roraima Region, Spelemoota, Ara-
poo River, 1 Dec. 1884, *E.F. Im Thurn* 29 (holotype
BM [barcode 000614317]; isotypes K [without bar-
code], US [barcode 00137725]; photo-K at NY). [Sec-
tion *Sipanea*].

Perennial herb 10-50 cm tall, prostrate, decumbent or ascending, with (1)3-20(-50) stems departing from a central taproot; stems simple (rarely a few secondary branches), to 200 cm long, usually reddish; young branches thin, 0.6-1.5 mm thick, terete, sparsely to densely antrorse strigose-hirtellous or antrorse sericeous (hairs white, 1-1.5 mm long). *Stipules* basally adnate to the petioles, sheathing, free portion truncate or broadly triangular to triangular at base and narrowly triangular to linear distally, or narrowly triangular, 1.5-3.3 x 0.6-1.5 mm, acute to acuminate, densely antrorse-strigose outside (hairs white, 1-1.5 mm long, longer hairs on the narrowly triangular lobe); glabrous, slightly thickened at margin inside; margin with 2 colleters on each side of the central lobe. *Leaves* sessile to subpetiolate, often with axillary brachyblasts with numerous smaller leaves; blades linear-lanceolate to linear, 1.3-4.3 x 0.2-0.9(-1.2) cm, 5-8 times longer than wide, acute to decurrent at base, obtuse to acute at apex, papyraceous to thinly subcoriaceous, dark green above and pale green below when fresh, olive-green when dry, sparsely strigose above, densely strigose or glabrous with midrib and secondary veins antrorse-strigose below; secondary veins 2-3(-4) on each side of midrib, obsolete above, prominent below; margins narrowly revolute. *Inflorescences* terminal, pedunculate, congested-cymose, commonly 5-flowered or rarely 1- or 3-flowered; lateral axes 5-10 mm long (excluding corollas), with 1 central flower and 2 flowers on each lateral axe; peduncles 0.7-3.7 cm long, sparsely to densely antrorse strigose-hirtellous or antrorse sericeous (hairs white, 1-1.5 mm long); bracts subtending inflorescence (when present) 2, leaf-like,

**Figure 40.** *Sipanea galioides*. **A.** Habit with short fertile branches, and long stoloniferous branches for vegetative reproduction. **B.** Sterile branch. **C.** Reproductive branch. **D.** Dissected short-styled flower, with stamens inserted at distal portion of corolla tube, and included style. **E.** Spinulose style branches of short-styled flower. **F.** Dissected long-styled flower, with stamens inserted at median portion of corolla tube, and exserted style. **G.** Corolla lobes variation within the same population. **H.** Capsule. **A, B:** drawn from *Delprete & Araujo 7430* (CAY); **C:** drawn from *Delprete & Araujo 7430* (CAY); **D-E:** drawn from *Delprete & Araujo 7412A* (CAY); **F-G:** drawn from *Delprete & Araujo 7412C* (CAY); **H:** drawn from *Steyermark 111276* (US). Illustration by Piero Delprete.
<figure><img src="image_44.png" /><figcaption>Figure 40.</figcaption></figure>
linear-lanceolate to linear, 3.3–6 × 0.4–0.8 mm, acute at apex, sparsely strigose above, densely strigose or glabrous with midrib and secondary veins antrorse-strigose below, or bracts absent; bracteoles subtending flowers linear-lanceolate to linear, 4–6 × 0.2–1 mm, densely antrorse-strigulose. *Flowers* distylous, 5(6)-merous, central flower sessile, those on lateral branches sessile to subsessile. *Hypanthium* turbinate to narrowly obovoid, 1.7–2 × 1–1.7 mm, acute at base, round at apex, sparsely to densely antrorse setose-strigulose. *Calyx* lobes equal or subequal, green to reddish-green when fresh, linear, 4–8 × 0.3–0.7 mm, acuminate at apex, densely white-strigose outside, margins strigose-ciliate, glabrous inside; with 1 colleter in each calcine sinus, linear, 0.2–0.4 mm long. *Corolla* hypocrateriform, (18.5–)22.5–27.5(–28) mm long, pink, pale pink, whitish-pink, or tube pink and lobes white during anthesis, turning whitish-pink to white after anthesis, with a dense ring of yellow hairs exserted beyond mouth; tube (13–)15–18 mm long, gradually expanding towards the mouth, 0.7–1.2 mm wide at base, 2–3 mm wide at mouth, glabrous at medio-basal portion and densely strigose, sometime only in lines, at medio-distal portion outside, glabrous at basal portion, densely puberulous above and densely yellow-pubescent at distal portion inside, the yellow pubescence exerted beyond corolla mouth; lobes 5(6), round, broadly ovate, obovate, elliptic-ovate to lanceolate, varying in shape and size within the same population (Figure 40G), sometimes unequal or of different shape on the same corolla, sometimes asymmetric, (5.5–)7.5–10.5 × 3–6(–7.5 mm), round, subobtuse to acute or apiculate at apex (Figure 40G), glabrous throughout. *Long-styled flowers* [Delprete & Araujo 7412A pro parte (corolla tube 14.5–15 mm long); Delprete & Araujo 7412C (corolla tube 13 mm long)]: stamens inserted at 7.5–8 mm from base, about the middle of corolla tube; filaments 0.7–1 mm long; anthers narrowly oblong-elliptic, 2.5–3 × 0.2–0.3 mm, acute at both ends; style exserted 1.5–3.5 mm beyond corolla mouth, 17–17.5 mm long, glabrous; style branches oblong, 1–1.3 mm long, round at tip, papillose-aculeate. *Short-styled flowers* [Delprete & Araujo 7412A pro parte (corolla tube 17.5 mm long)]: stamens inserted at 3.5 mm below corolla mouth; filaments 2.5 mm long; anthers narrowly oblong-elliptic, 3 × 0.2–0.3 mm, acute at both ends; style included, 12 mm long, glabrous; branches linear, 3 mm long, round at tip, papillose-aculeate. *Infructescences* cymose, slightly more expanded than inflorescences. *Capsules* cylindrical-oblong to oblong-turbinate, 5–7.5 × 2–4 mm, acute at base, round to acute at apex, thinly woody, faintly costate, densely to sparsely antrorse-strigulose or antrorse-hispid, (hairs 1–1.5 mm long), green to reddish-green when young,

turning beige to pale brown when dry. *Seeds* irregularly trigonous-rhomboid or subglobose, faintly or strongly angular, (0.4–)0.5–0.7 mm in diam., testa foveolate.

**Notes:** Wernham (1917) in the introduction of the article where he described *Sipanea galioides* wrote "The types of all the novelties described are in the National Herbarium [BM]." Therefore the specimen *Im Thurn 29* at BM, barcode 000614317, is the holotype of this name.

The type specimen at US has the typewritten label "Mt. Roraima Expedition, British Guiana, Oct.-Jan., 1884-5" with the handwritten note "Sipanea pratensis Aublet, N. 29, Spelemonta [sic! Spelemoota], Arapoo R." which corresponds entirely with the information supplied on the holotype specimen label and in the original publication. On the same sheet an additional handwritten label reports the collector as "E. Jenman." Because collection date and locality are identical to those of the holotype, the US specimen should be cited as "E.F. *Im Thurn 29*" and is an isotype.

*Sipanea galioides* is similar to *S. pratensis*, the former differing from the later by the combination of the following characters: stems commonly simple or rarely branched, often with axillary brachyblasts topped by numerous smaller leaves (vs. commonly branched, without brachyblasts in *S. pratensis*), leaf blades papyraceous to thinly subcoriaceous, with margins narrowly revolute, and 2-3(-4) secondary veins on each side of midrib (vs. chartaceous, margin planar, with 3-8 secondary veins on each side of midrib), inflorescences (1-3)3-5-flowered (vs. congested, 4-8-flowered, or evidently branched, 4-37-flowered), hypanthium 1.7-2 × 1-1.7 mm, sparsely to densely antrorse setose-strigulose (vs. 0.7-1.5 × 0.5-0.9 mm, appressed pubescent to sericeous), corolla tube (13-)15-18 mm long (vs. (8-)10-17 mm long). In addition, *Sipanea galioides* is restricted to Venezuela (Amazonas and Bolívar) and Guyana, at (125-)700-2200 m altitude, while *S. pratensis* is widely distributed in Colombia, Venezuela, the Guianas, and northern Brazil (Roraima, Amapá, Pará, Amazonas, and Maranhão), and is an extremely variable species in terms of habit, leaf shape, inflorescence architecture, flower size, and fruit shape, found in extremely different environments from near sea level to 1200 m altitude.

**Phenology:** Individuals of this species are commonly found with flowers and fruits on the same inflorescence. Flowering and fruiting mostly during the rainy season, from November-December through May.

**Distribution and ecology:** Restricted to Venezuela (Amazonas and Bolívar) and Guyana (Figure 41). Pros-
trate, decumbent or ascending herb (stems sometimes woody at basal nodes), usually growing in open dry, sandy savannas, savannas with sparse trees, white-sand savannas, rocky savannas, seasonally flooded area with Maurititia flexuosa, bare laterritic slopes of savannas, borders of dwarf forests, riparian forests, open tepui slopes, or on rocky outcrops; at (125–)700–2200 m altitude.

**Suggested conservation status:** Least Concern (LC). This species is known from numerous localities in Venezuela, mostly in the state of Bolívar, at altitudes of 700–2200 m, and a couple of collections in the state of Amazonas at about 125 m altitude, as well as several collections from Guyana, at altitudes of (80–)700–1830 m. In Venezuela, most collection localities are within the Canaima National Park, state of Bolívar. In Guyana,

several collections are from the Mount Roraima Massif, which includes the Pakaraima Mountains, and is contiguous with the Canaima National Park. During a botanical expedition in Venezuela in December 2000, several healthy populations of this species (*Delprete & Araujo 7412A, 7412C, 7414, 7427*) were personally observed in the municipalities of Gran Sabana and Cavanayen, which are both currently included within the Canaima National Park. The number of individuals in each population was not reported. Considering that this species is known by numerous collections, and many of the localities are within a well-preserved national park, without any imminent threat, this species is positioned in the Least Concern (LC) category, following IUCN criteria (IUCN 2012, 2019).

<figure><img src="image_45.png" /><figcaption>**Figure 41.** Distribution of *Sipanea galioides* (circles) and *S. glaberrima* (star).</figcaption></figure>
**Specimens examined:** VENEZUELA. Amazonas:
Mun. Guainía, along road from Maroa to Yavita, ca. 700 m from Yavita, 2°54'42"N, 67°26'55"W, evergreen, moist forest, decumbent herb, many-branched from base, corolla turning from white to pinkish, 19 Feb. 1998 (fl, fr), *P. Acevedo-Rodriguez et al. 10266* (MO, PORT, US); Ríos Sipapo y Autana, 1952 (fl), *Herbier S.M. Roi Léopold III N.* 332 (BR); Río Negro, rock outcrop and forest around it, Mamurividi, Zona 10, Estación Climatológica of Ministerio del Ambiente on Río Pasimoni, 1°32'N, 66°32'W, 125 m, 3 Apr. 1984 (fl, fr), *R. Liesner 17201* (CAY [2 sheets], MG). **Bolívar:** Distr. Roscio, sabana inundable en la base del Cerro Roraima, 5°13'N, 60°45'W, 2200 m, 26 Mar. 1984 (fl), *G. Aymard & J. Luteyn 2432* (NY, PORT, VEN); Calzeta de la Botella, 150 m, 14 Apr. 1957 (fl), *A.L. Bernardi 6505* (NY [2 sheets]); La Gran Sabana, km 145 along hwy, 2 km S of La Cuidadela, open savanna dominated by *Trachypogon, Echinolaena*, and *Paspalum*, soil with top 20 cm sand and organic material mixture, 1 m of white sand, below which is yellow sand, 1300 m, 3 Dec. 1983 (fl), *G. Davidse 4680* (MO, NY, VEN); Mun. Gran Sabana, Parque Nacional Canaima, Tarotá, 5°47'8"N, 61°36'55"W, 1320 m, 1 May 2002 (fl), *F. Delascio et al. 18203* (NY, VEN); Mun. Gran Sabana, Parque Nacional Canaima, savanna with herbaceous vegetation dominated by Graminaceae, Cyperaceae, Orchidaceae and *Lycopodium*, white sand soil, full sunny exposure, growing on mounds and in areas with low, slowly flowing water, 5°58'N, 61°24'W, 1300 m, 20 Dec. 2000 (fl), *P.G. Delprete & A.C. Araujo 7412A* (CAY, pickled flowers only); Mun. Cavanayen, 2 km E of village, disturbed herbaceous-shrubby vegetation near small creek, herbaceous layer dominated by Graminaceae, Cyperaceae, Xyridaceae and sparse Melastomataceae, small population of sparse individuals, perennial herb, erect, suberect to prostrate with axillary brachyblasts with smaller leaves, 1200 m, 5°35'N, 61°44'W, 21 Dec. 2000 (fl), *P.G. Delprete & A.C. Araujo 7412C* (CAY, pickled flowers only); Mun. Gran Sabana, ca. 15 km S of Kama-Meru Camp, open field frequently burned, with some degree of moisture dominated by Cyperaceae and Poaceae, 5°20'N, 61°13'W, 1170 m, 22 Dec. 2000 (fl), *P.G. Delprete & A.C. Araujo 7414* (CAY, MO, VEN); Mun. Gran Sabana, 2 km N of San Rafael de Kamoiren, km 174, area seasonally inundated with white-sand soil, dominated by Melastomataceae, Rapateaceae, Piperaceae and *Drosera*, 5°39'N, 61°24'W, 1300 m, 27 Dec. 2000 (fl, fr), *P.G. Delprete & A.C. Araujo 7427* (CAY, GB, NY, VEN), *7430* (BR, CAY, P, US, VEN); km 119–122 S of El Dorado, 1100 m, 16 Mar. 1974 (fl), *A. Gentry, G. Morillo & B. de Morillo 10462* (MG, NY); Distr. Roscio, sabana arbustiva sobre cuestas de piedemonte al WSW del Ilu-

Tepui, en la region del Río Caraurín medio, 5°18'N, 61°0'W, 1150 m, 27 Jun. 1983 (fl), *O. Huber & C. Alarcón 7701* (NY, VEN); Distr. Roscio, extensas sabanas abiertas sobre altiplanicie en la region del Alto Río Arabapó, ca. 10 km SW del Uei-Tepui (Cerro El Sol), 4°56'N, 60°41'W, 1100 m, 21 Jan. 1985 (fl), *O. Huber 9998* (NY, VEN); Alto Caroni, near Sta. Elena de Uairen, 25 Apr. 1946 (fl, fr), *T. Lasser 1457* (NY, US, VEN); Kavanayen, en sabanas, Guayana Venezolana, 30 May 1946 (fl, fr), *T. Lasser 1893* (NY); 17 km E of El Pauji by rd. and 64 km W of Santa Elena by road, 4 km N of hwy, Río las Ahallas, 4°30'N, 61°30'W, 850 m, 1 Nov. 1985 (fl), *R. Liesner 19267* (MO, NY, VEN); Dist. Piar, 2–5 km W of Amaruay-Tepui, savannas between Tepui and Río Acanán Guarumo, 5°56'N, 62°17'W, 470 m, 4 May 1986 (fl), *R. Liesner 20594* (MO, NY); Gran Sabana, 5 km S of San Ignacio de Yuruani, wet savanna on slope, 4°59'N, 61°10'W, 850 m, 9 May 1988 (fr), *R. Liesner 24412* (MO, NY); km 121 along El Dorado-Santa Elena rd., wet dwarf forest, 1310 m, 16 Nov. 1978 (fl, young fr), *J.L. Luteyn et al. 6264* (NY, US, VEN); Gran Sabana, savanna 1–5 km NW of Misión at Kavanayen, 1300 m, 4 Apr. 1952 (fl), *B. Maguire 33733* (NY, US); road S of El Dorado, km 98, along edge of rain forest, 450 m, 18 Dec. 1979 (fl), *J. Pruski & J.A. Steyermark 1368* (NY, US, VEN); Gran Sabana, dry, open, rocky savanna on Roraima sandstone near km marker 164 S of El Dorado, 20 Dec. 1979 (fl), *J. Pruski & J.A. Steyermark 1441* (NY); Gran Sabana, vía Kavanayen, El Jardin (Guamu-Loma de Guamu), entre el campamento Parupa y Cavanayen, ca. 65 km E del Fuerte Luepa, 1350 m, 7 Oct. 1984 (fl, fr), *N. Ramírez 976* (NY, VEN); vicinity of road campamento 150 at km 150 in valley of savanna of Río Uarama below Uarama-Tepui, NE of Luepa [ca. 5°50'N, 61°30'W], 1220 m, 24–25 Apr. 1960 (fl), *J.A. Steyermark & S. Nilsson 537* (NY, VEN), *694* (NY, VEN); Mount Roraima, forested SW-facing quebrada near Rondón Camp, 2040 m, 25 Sep. 1944 (fl), *J.A. Steyermark 58649* (NY); dense forest along Río Karuai, between Santa Teresita de Kavanayén and base of Ptari-Tepui, 1220 m, 18 Nov. 1944 (fl), *J.A. Steyermark 60297* (NY, US, VEN); Carretera El Dorado hacia Santa Elena de Uairen, km 198 al S de El Dorado, 1200–1400 m, 7–10 Dec. 1972 (fl, young fr), *J.A. Steyermark et al. 106626* (NY, US, VEN); km 121 S of El Dorado on rd. to Sta. Elena, bordering dwarf forest, 1200 m, 29 Dec. 1974 (fl, young fr), *J.A. Steyermark 111276* (US); Gran Sabana, Las Delicias, Sta. Elena, Feb. 1946 (fl), *F. Tamayo 2769* (US).

**GUYANA:** Potaro-Siparuni Region, Surama, 4°10'N, 59°5'W, 80 m, 1 Dec. 1995 (fl), *D. Clarke 698* (MO, US); Cuyuni-Mazaruni Region, Utshi River to Great Falls trail, on Kamarang River, 5°40'N, 61°6'W, 910 m, 30 Jan. 1996 (fl), *D. Clarke 876* (CAY, MO, NY, U, US); Cuyu-
ni-Mazaruni Region, Pakaraima Mountains, savanna between Koatse River and Chinowieg Village, white-sand savanna, 5°27′N, 60°4′W, 700–800 m, 12 Nov. 1992 (fl), *T.W. Henkel & B. Hoffman* 217 (MO, US); Mount Roraima Expedition, 1000 m, fall of 1894 (fl), *J.J. Quelch & F. McConnell* 195 (K), 198 (K); Mount Roraima, stony slopes on sandy soil, Philipp Camp, 5200–6000 ft [1585–1830 m], 7 Nov. 1927 (fl, fr), *G.H.H. Tate*, 288 (NY); Atkinson, St. Cuthbert's Trail, 4.5 mi from Highway, fringe of xeromorphic scrub/palm marsh woodland, 26 Feb. 1969 (fl), *U.G. Bio 106* [*University of Guyana, Biology, course 106*] 74 (NY).

### 8-2. *Sipanea glaberrima* (Bremek.) Steyerm., Mem. New York Bot. Gard. 17: 270. 1967. (Figures 41–42).

(≡) *Sipanea pratensis* var. *glaberrima* Bremek., Recueil Trav. Bot. Néerl. 33: 707. 1936 [reimpr. Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 35: 707. 1936].

Type: SURINAME. Upper Sipaliwini River, Great Savannah, 2°N, 56°W, 23 Oct. 1935 (fl), *H.E. Rombouts* 223bis (holotype U [barcode 0006310]; photo-U at NY). [Section *Sipanea*].

*Herbs*, height unknown, erect, entirely glabrous, exep corolla throat; young branches thin, 1.3–1.6 mm thick, terete, glabrous. *Stipules* basally adnate to the petioles, free portion truncate and mucronate, or broadly triangular at base and narrowly triangular distally, 1.5–2 × 1.3–1.5 mm, distal triangular portion 0.7–0.9 mm long, slightly thickened at margin inside; margin with 3 colleters on each side of the central lobe. *Leaves* short-petiolate; petioles 2.5–3 mm long, glabrous; blades elliptic to narrowly ovate, 3–3.7 × 1.3–1.5 cm, obtuse to acute and shortly decurrent at base, acute at apex, chartaceous, olive-green when dry, glabrous throughout; secondary veins 7 on each side of midrib, obsolete above, prominent below. *Inflorescences* terminal, pedunculate, dichotomously branched, 25-flowered, 6.7 × 2.5 cm (excluding corollas), lateral axes 2.8 cm long, scorpiod; peduncle 3.9 cm long; bracts subtending secondary branches leaf-like, elliptic, 9.5–10.5 × 6 mm, acute; bracteoles subtending flowers narrowly lanceolate, 1–7 × 0.2–0.6 mm. *Flowers* sessile, 5-merous; distyly not observed. *Hypanthium* turbinate, 1–1.5 × 0.6 mm, acute at base, round at apex. *Calyx* lobes subequal, linear-lanceolate, 3.5–4 × 0.1–0.3 mm, acuminate; with 1 colleter in each calyxine sinus, linear, 0.3 mm long. *Corolla* hypocrateriform, 15.3–15.5 mm long, color unknown, with a dense ring of yellow hairs exserted beyond mouth; tube 12.5 mm

long, gradually expanding towards the mouth, 0.7 mm wide at base, 1.5 mm wide at mouth, glabrous at basal 5 mm, densely puberulous above (up to stamen insertions) and densely yellow-pubescent at distal portion inside, the yellow pubescence exerted beyond corolla mouth; lobes broadly obovate to oblong-elliptic, 3–3.5 × 2.3–2.5 mm, obtuse at apex. *Short-styled flowers (Rombouts 223bis* (corolla tube 12.5 mm long)): stamens inserted 3 mm below corolla mouth; filaments 0.2–0.3 mm long; anthers narrowly oblong-elliptic, 2–2.2 × 0.2 mm, acute at both ends; style included, 6 mm long; branches linear, 2–3 mm long, acute at apex. *Long-styled flowers* unknown. *Infructescences* dichotomously branched, with scorpioid secondary branches, lateral axes 2.8 cm long. *Capsules* oblong, 5–5.8 × 2.7–3 mm, round at base, obtuse at apex, thinly woody, strongly costate, turning beige when dry. *Seeds* irregularly tetrahedral, faintly angular, 0.4–0.5 mm in diam., testa foveolate, rugulose.

**Notes:**—Bremekamp (1936: 707) described *Sipanea pratensis* var. *glaberrima* Bremek., and wrote "This variety is known in one specimen only, a plant collected by Mr. Rombouts in the Great Savanna (Lat. 2°N, Long. 56°W) in the midst of plants belonging to the *forma typica*." Steyermark (1967: 270) upgraded Bremekamp's variety to species rank, and stated "This is a distinct species, easily recognized by having all parts of the plant glabrous," and provided a complete species description.

*Sipanea glaberrima* (Bremek.) Steyerm. is unique within the genus by being completely glabrous, except for the pubescence at corolla throat. It is most similar to *S. pratensis* var. *dichotoma*, with which is sympatric, because of the erect habit, branched inflorescences, flower size, and oblong, costate capsules. As the two taxa are sympatric, a varietal rank for *S. glaberrima* is incompatible with the variety concept adopted in the present treatment (secondary variation with a morphological-geographical correlation, where sympatric varieties are not allowed; see section Species and Variety Concepts, above); therefore, it is here treated as a distinct species. Aside from being completely glabrous, *S. glaberrima* differs from *S. pratensis* by the stipules truncate and mucronate, 0.5–2 mm long, glabrous (vs. broadly triangular at base and very narrowly triangular to linear distally, 2–6 mm long, appressed pubescent to sericeous outside).

**Distribution and ecology:** Only known by the single type specimen collected in southernmost Suriname, in the Great Savanna of Sipaliwini, at 2°N, 56°W (Figure 41). Growing in sunny exposure, in the midst of a population of *S. pratensis* var. *dichotoma*.
<figure><img src="image_46.png" /><figcaption>**Figure 42.** *Sipanea glaberrima*. A. Branch with terminal inflorescence. B. Node with stipule and petioles. C. Dissected short-styled flower, with stamens inserted at distal portion of corolla tube, and included style. D. Capsule. A–D: drawn from *Rombouts 223bis* (U, holotype). Illustration by Piero Delprete.</figcaption></figure>
vate, 7-8 × 3.5-5.5 mm, round to subacute at apex, glabrous throughout. *Long-styled flowers* [Ll. *Williams 13016* (corolla tube 12 mm long]): stamens inserted at 4.5-5.5 mm from base of corolla tube; filaments 0.5 mm long; anthers linear, 2.5 × 0.2 mm, acute at both ends; style exserted 2-3 mm beyond corolla mouth, 14-15 mm long, glabrous; style branches obovate, 1-1.3 mm long, round at apex. *Short-styled flowers* [Holt & Blake 799 (corolla tube 15 mm long), Zarucchi 1675 (corolla tube 17 mm long)]: stamens inserted at 4-4.5 mm below corolla mouth; filaments 0.3-0.5 mm long; anthers linear, 2.5-3 × 0.2-0.3 mm, acute at both ends; style included, 6-9 mm long, glabrous; branches linear, 2.3-2.8 mm long, acute at apex. *Infructescence* a condensed single head or tricotomous with three condensed heads, similar to inflorescence. *Capsules* oblong-ovoid to narrowly ovoid, 6-9 × 3-4.5 mm, round at base, acute at apex, crustaceous, densely gray-sericeous (hairs 0.4-1 mm long), pale green when young, turning beige when dry. *Seeds* irregularly tetrahedral, subangular, 0.6-0.8 mm in diam., testa foveolate, rugulose-verrucose.

**Notes:** After exhaustive searches in P, P-JU and P-Bonpl., no original specimen of *Sipanea glomerata* Kunth could be found. See discussion under *Sipanea pratensis* var. *dichotoma* for further information regarding the complex history of the specimens collected by Bonpland and Humboldt and described by Kunth. An original specimen of *S. glomerata* was found at B-W, with barcode B-W 04123-010. This specimen has two handwritten labels: "Sipanea glomerata H.B.Kth. 3 p. 310. - ! Schum." and "germin. an 8 [Germinal (22 March-20 April) 1800], orinoco, No. 862. Pent. 1ia [Pentandria monogynia], ποσχύδης - psonatur (W.) - in Guiana d. Aturès - (Humboldt)." On the lower right corner of the sheet it is handwritten "Humboldt. W." and on the upper right corner is handwritten "Pt. [Ptychodea] sessiliflora." On the sheet is mounted a branch with numerous leaves, and an inflorescence with a flower in anthesis. Aside with the branch are present several inflorescence fragments, including one loose flower in anthesis. This specimen is here designated the lectotype of *S. glomerata*. This specimen is also here designated as the lectotype of the illegitimate superfluous name "*Ptychodea sessiliflora*" that was cited by Chamisso & Schlechtendal (1829: 168) as a synonym of *Sipanea glomerata*. The image of this specimen is avail-

able at: https://ww2.bgbm.org/Herbarium/specimen.cfm?Barcode=BW04123010.

Humboldt donated to Kunth about 3000 original specimens from the Bonpland and Humboldt collection (Stauffer et al. 2012), which Kunth took with him to Berlin, after he finished writing the *Genera Plantarum*. Kunth's private herbarium comprised nearly 70,000 specimens, which were all destroyed during WWII in 1943. Francis Macbride, of the Field Museum, took photographs of type specimens in numerous European herbaria before they were destroyed during WWII. One of these photographs is of a specimen of *S. glomerata*, which has the label handwritten by several authors "Sipanea glomerata Kunth in Humb. et Bonpl. 3 p. 398. Ptychodea sessiliflora Willd. herb. ! Schum., Hb. Kunth, Ex herb. Humb." Most likely, that specimen was part of the original material used by Kunth to describe this species, however the label does not indicate any locality or collection number, which renders impossible to connect this image to the specimen of *Bonpland & Humboldt 862* present in B-W. Macbride's photograph of this specimen is available at: https://fm-digital-assets.fieldmuseum.org/36/440/60.jpg

Steyermark (1967: 281-282; 1974: 364-365) distinguished the typical variety of *Sipanea glomerata* from var. *paucinervia* Steyerm. by having "upper surface of leafblades densely sericeous with crowned appressed hairs" (vs. "glabrous or sparsely appressed-pilose with more scattered hairs" in var. *paucinervia*), "leaf blades 2.5-4.5 cm broad" (vs. 1-2.2 cm broad), and "lateral nerves of leafblades 7-11 on each side" (vs. "5-6"). Delprete and Steyermark (2004e: 829-830) did not recognize any variety in this species, without any comment. After a detailed study of ample material from throughout its geographic range, I confirmed that the characters used by Steyermark to differentiate the two varieties are widely overlapping and no varieties are here recognized in this species.

**Distribution and ecology:** Distributed in the Amazon Region of Colombia, Venezuela, Surinam, and Brazil (Mato Grosso, Roraima, Pará, and Amazonas) (Figure 44). Often found in large stands, in open savannas with sparse trees and shrubs, or on forested and open igneous granitic outcrops, gravelly savannas, sometimes in secondary vegetation (in Brazil called "capoeira"), in sunny or semi-shady exposures, in sandy, granitic or lateritic soils; at 80-200 m altitude.

**Figure 43.** *Sipanea glomerata.* A. Branch with terminal young inflorescence. B. Node with sheathing stipule and leaf bases. C. Dissected long-styled flower, with stamens inserted at medio-distal portion of corolla tube, and exserted style. D. Dissected short-styled flower, with stamens inserted at distal portion of corolla tube, and included style. E. Capsule. A–C: drawn from *Williams 13016* (US); D: drawn from *Holt & Blake 799* (US); E: drawn from *Maguire et al. 36046* (US). Illustration by Piero Delprete.
<figure><img src="image_47.png" /><figcaption>Figure 43.</figcaption></figure>
<figure><img src="image_48.png" /><figcaption>**Figure 44.** Distribution of *Sipanea glomerata*.</figcaption></figure>

**Notes:** *Sipanea glomerata* is similar and has been often confused with *S. hispida*. The former can be distinguished from the latter by the stems sparsely to densely appressed or spreading, pubescent or sericeous (vs. densely hirsute or villous in *S. hispida*), leaf adaxial side densely sericeous or sparsely to densely appressed pubescent to glabrate (vs. sparsely to densely strigose), leaf abaxial side densely sericeous or sparsely appressed pubescent or glabrous (vs. densely villose-sericeous), inflorescence a single head usually subtended by two leaf-like bracts, or trichotomous and made of three pedunculate heads (vs. cymose, dichotomous or rarely trichotomous), calix lobes 7–13 × 0.7–1.2 mm in flowers, enlarging to 15–17 mm long in fruits (vs. 4–6 × 0.3–0.6 mm), corollas 19–25 mm long, white (vs. 8.5–15(–16.5) mm long, pink), corolla tubes 12–17 mm long (vs. 6–10(–12.5) mm long), and corolla lobes 7–8 × 3.5–5.5 mm (vs. 2.5–5 × 1.5–3 mm).

**Phenology:** Individuals of this species are commonly found with flowers and fruits. Flowering and fruiting the whole year around.

**Uses:** In Venezuela (Amazonas: Sanariapo), an infusion made with the whole plant of this species is used for stomach problems (Ll. *Williams* 16002).

**Suggested conservation status:** Least Concern (LC). This species has been collected in numerous localities in Colombia, Venezuela, Surinam, and northern Brazil. It is often found in large stands in secondary vegetation and along roadsides (pers. obs. in Venezuela and Brazil). Considering that this species is known by numerous collections, and is often found as a ruderal in secondary vegetation, it is positioned in the Least Concern (LC) category, following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** COLOMBIA: Guainía: Immediately S of Casuarito, lajas along the Río Orinoco, in cracks on laja and grassy pockets, 5°40′N, 67°37′W, 90 m, 22 Jun. 1984 (fr), *G. Davidse & J.S. Miller* 26391 (NY); Río Negro, San Felipe and vicinity, below confluence of Río Guainía and Río Casiquiare, 1°50′N, 67°0′W, 200 m, 26 Oct. 1952 (fl), *R.E. Schultes et al.* 18065 (NY. US), 18094 (US). **Vaupés:** Río Vaupés, Cerro de Mitú, 20 Aug. 1960 (fr), *R.E. Schultes & I. Cabrera* 22716 (NY); Mitú and vicinity [ca. 1°11′N, 70°10′W], granitic slope of Cerro de Mitú, side facing Mitú, 28 May 1976 (fl), *J.L. Zarucchi* 1675 (MO, US). **Vichada:** Corregimento Cumaribo, carretera Cumaribo-Santa Rita, camino entre
Palmarito y Chaparral, en morichal, 4°25'W, 69°48'W, 150 m, 12 Dec. 1993 (fl, fr), C. Sastre et al. 9082 (P), 9083 (COL); "prope Maypures, flumen Orenoco" [Maipures, Río Orinoco, ca. 5°12'N, 67°50'W], Jun. 1854 (fl), R. Spruce 3665 (BR, P).

**VENEZUELA: Amazonas:** Bords de l'Orenoque, savane Atures, s.d. (fr), *J. Chaffanjon 171* (P); Mission Camp, ornamental plant with white flowers, 4 Apr. 1950 (fl), *H.M. Curran 209* (NY); Depto. Atures, ca. 23 km NE of Puerto Ayacucho along the rd to El Burro, laja and savanna at its base, 5°51'N, 67°29'W, 80-150 m, 17 Apr. 1979 (fl), *G. Davidse & O. Huber 15248* (MO, NY, VEN); Depto. Atures, ca. 4.5 km N of Puerto Venado, just E of Samariapo, 5°15'N, 67°47'W, 85 m, 25 Apr. 1979 (fl), *G. Davidse et al. 16766* (MO, US, VEN); near Puerto Ayacucho, 100 m, 30 Sep. 1960 (fl), *E. Foldats 3529* (NY); bords de l'Orenoque, entre Atures et San Fernando de Atabapo, 17 May 1887 (fr), *A. Gaillard 7* (P); 22-23 km N of Samariapo on rd. to Puerto Ayacucho, 100 m, forest edge, 5°39'50"N, 67°37'25"W, 29 Jun. 1975 (fl), *A. Gentry & P. Berry 14571* (BM, MO); Puerto Ayacucho, 100 m, May 1931 (fl), *E.G. Holt & E.R. Blake 799* (NY, US); alrededores de Puerto Ayacucho, ca. 12 km S, laja granitica, derecho de la carretera Puerto Ayacucho-Samariapo, 5°34'N, 67°35'W, 80-250 m, 20 Jul. 1977 (fl), *O. Huber & S.S. Tillett 923* (NY); Great Rapids of the Río Orinoco, Puerto Ayacucho, crystalline laja 1-1.5 km E of Hotel Amazonas, 100-120 m, 7 Nov. 1953 (fl, fr), *B. Maguire et al. 36046* (NY, US); sobre laja granitica, ca. 3.5 km al N de Puerto Ayacucho, margen de la carretera Puerto Ayacucho-El Burro, 80 m, 14 Oct. 1988 (fl, fr), *G.A. Romero 1693* (NY, S, TFAV), 17 Oct. 1988 (fl, fr), 1756 (NY); transecto entre bosque inundado (rebalse) y Cerro Caribe (afloramiento granítico al N de San Juan de Ucata), 4°20'N, 67°49'W, 70-100 m, 22 Oct. 1989 (fl, fr), *G.A. Romero et al. 2168* (NY); Depto. Atures, Montaña Fría, entre poblaciones de Puerto Ayacucho y Sanariapo, 180 m, 26 Jul. 1967 (fl), *L. Ruiz-Terán 4327* (MERF, MG); Depto. Atures, forested areas and igneous outcrops alog Río Coromoto, at Tobogán de la Selva, 35 km SE of Puerto Ayacucho, 5°22'N, 67°33'W, 150 m, 14 May 1980 (fl), *J.A. Steyermark et al. 122563* (MO, NY); Depto. Atures, Tobogán de la Selva, Río Coromoto, 35 km SE of Puerto Ayacucho, among igneus exposures, 5°27'N, 67°33'W, 7 Sep. 1985 (fl), *J.A. Steyermark et al. 131518* (MO, NY, VEN); Puerto Ayacucho, orilla del Riachuelo El Bagre, 100 m, 20 May 1940 (fl), *Ll. Williams 13016* (US); sabana abierta de Sanariapo [ca. 5°14'N, 67°47'W], parte inferior, 120 m, 3 Jul. 1942 (fl), *Ll. Williams 16002* (US).

**SURINAME:** Fluv. Corantjin [Courantyn River], Apihollo Savanna, 29 Jan. 1911 (fl), *J.F. Hulk 56* (U); Via secta

ab Moengo [ca. 5°37'N, 54°24'W], tapoe ad Grote Zwiebelzwamp, km 6.7 in savanna, 5 Oct. 1948 (fl), *J. Lanjouw & J.C. Lindeman 657* (U); ibid., km 15.7-15.9 in savanna, 15 Oct. 1948 (fl), *J. Lanjouw & J.C. Lindeman 844* (U).

**BRAZIL. Amazonas:** Rio Negro, Tapuruquara [ca. 0°24'S, 65°2'W], capoeira, 25 Apr. 1948 (fl, fr), *G.A. Black 48-2481* (IAN); Barcelos, Serrinha, afloramento rochoso, próximo do Rio Aracá, 0°25'3"N, 63°22'58"W, 59 m, 20 Aug. 2014 (fl, fr), *F.N. Cabral et al. 1217* (INPA); Rio Negro, Uaupés, Taracuá [ca. 0°41'N, 69°32'W], arredores, 3 Mar. 1959 (fl, fr), *P. Cavalcante 738* (MG); Mun. Barcelos, margem do Rio Araça, pouco abaixo da foz do Rio Jauari, 0°25'N, 63°25'W, 20 Jul. 1985 (fl), *I. Cordeiro 245* (INPA, MG, NY, US); Barcelos, Rio Aracá, acima da comunidade de Bacuquara, afloramento de granito [inselberg] localmente chamado Serrinha, 0°11'20"N, 63°10'5"W, 110 m, 17 Apr. 2014 (fl), *R.C. Forzza et al. 7961* (INPA, RB); Rio Negro, Tapurucoara [ca. 0°24'S, 65°2'W], 6 Apr. 1947 (fl, fr), *R.L. Frões 22076* (IAN); Rio Uaupés, local Ipanoré, uma hora de motor de popa acima de Taraquá, Estrada para Urubuquara, ca. 0°18'N, 68°40'W, caatinga, 17 Nov. 1987 (fl), *M.L. Kawasaki 171* (INPA); Basin of Rio Negro, Tapuruquara [ca. 0°24'S, 65°2'W], beside road to airport, capoeira, 16 Oct. 1971 (fl), *G.T. Prance et al. 15315* (NY, S, US); Rio Negro, Tupuruquara, terra firme, solo argiloso, erva ereta muito comum no local, 20 May 1973 (fl), *M.F. Silva et al. 1740* (INPA, US); Tyler-Duida Expedition, Rio Negro, Santa Izabel [now Santa Isabel do Rio Negro, ca. 0°24'S, 65°2'W], 8 Sep. 1928 (fl), *G.H.H. Tate 98* (NY, US); Mun. Barcelos, Serra do Aracá, platô, solo arigiloso e pedregoso, subarbusto, 0°55'18"N, 63°26'48"W, 28 Aug. 2001 (fl), *A. Vicen tini et al. 1913* (INPA); Upper Rio Negro River, 1907-1908 (fr), *L. Weiss & H. Schmidt s.n.* (NY [2 sheets, 02690184 and 02690185]). **Pará:** Mon. Óbidos, Rio Parú de Oeste (Tiriós), campo, 21 Jun. 1960 (fl, fr), *P. Cavalcante 794* (IAN, MG); Região dos Tiriós, Rio Paru do Oeste [ca. 1°31'S, 56°02'W], 16 Mar. 1962 (fl), *E.J. Fittkau & D. Coel hlo s.n.* (INPA No. 12773) (INPA, RB), 17 Mar. 1962 (fl), s.n. (INPA 12805) (INPA). **Roraima:** Local Anauá [ca. 0°57'N, 61°22'W], 7 Nov. 1973 (fl, fr), *B.G.S. Ribeiro 391* (IAN); Morro Tepequem [ca. 3°45'N, 61°45'W], 3 Feb. 1975 (fl, fr), *B.G.S. Ribeiro 692* (IAN), 698 (IAN).

8-4. *Sipanea hispida* Benth. ex Wernham, J. Bot. 55: 173. 1917. (Figures 45-46).

K.Schum. in Martius et al., Fl. Brasil. 6(6), fig. 122 (as *Sipanea pratensis*). 1889.

Type: BRAZIL. Amazonas: Rio Negro, São Gabriel da Cachoeira [ca. 0°7'N, 67°5'W], Jan.-Aug. 1852,
R. *Spruce 2051* (first-step lectotype, designated by Steyermark (1967: 279), **second-step lectotype** BM [barcode 000614346], **here designated**; isolectotypes E [barcode E00505240], F [Acc. No. 767786], FI-Webb [barcode FI004807], G [2 sheets; barcodes G00436761, G00436762], K [without barcode], LD [barcode 1220003], M [barcode M-0189372], NY [barcode 00133313], P [barcode P00748149], RB [Acc. No. 17401, barcode 00543715]; frag F [Acc. No. 635362]; photo-K at NY). [Section *Sipanea*].

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

Type: COLOMBIA. [Villavicencio, Susumuco, 400 m, 1000 m; see Kirkbride (1982: 305)], s.d. (fl, fr), *J.J. Triana* 1776 (holotype BM [barcode 000614328]; iso types E [barcode 00285373], P [barcode P00748116 "entre Susumuco y Villa Vicencio"], US [Acc. No. 1481047, barcode 00137730]; photo-US, photo-K and photo-BM at NY).

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

Type: BRAZIL. Minas Gerais: on a dry bank near Sabara, Sep. [1840] (fl, fr), *G. Gardner* 5009 (holotype, BM [barcode 000614348]).

(=) *Sipanea pratensis* var. *major* Hassl., Repert. Spec. Nov. Regni Veg. 14: 170. 1915

(=) *Sipanea hispida* Benth. ex Wernham var. *major* (Hassl.) Steyerm., Mem. New York Bot. Gard. 17: 280. 1967

Type: PARAGUAY. Upper Apa River, Nov. [year unknown], *E. Hassler* 7733 (**lectotype**, G [2 sheets, barcode G00307684], **here designated**; isolectotypes, [4 sheets, G00307683, G00307685, G00307685, G00307726], BM [barcode 000549052], MPU [barcode MPU021334], NY [barcode 00133326], P [3 sheets, barcodes P00729402, P00753869, P00753870], S [Acc. No. 10-27662]).

(=) *Sipanea glabrata* Wernham, J. Bot. 55: 173. 1917

Type: BRAZIL. Mato Grosso: "Santa Anna da Chapada" [now Chapada dos Guimarães], 28 Jun. 1902 (fl, fr), *A. Robert* 333 (**lectotype**, BM [barcode 000614333], **here designated**; photo at NY).

Perennial herbs or subshrubs (basal internodes woody), 0.1-1.2 m tall, terrestrial, erect; young branch-

es terete not geniculate, green or rarely reddish, densely hirsute or villous (hairs long, spreading). *Stipules* adnate to the petioles, sheathing, free portion broadly to shallowly triangular at base and narrowly triangular to linear distally, 4-13 × 1-2.5 mm, densely strigose or villous outside, glabrous and slightly thickened at margin inside; with 5-6 colleters inserted just below the base of the central lobe inside or 6-7 colleters inserted just below the margin inside; margin densely strigose or villous. *Leaves* sessile to short-petiolate; petioles to 5 mm long, densely antrorse or spreading strigose or villous; blades ovate, ovate-lanceolate, narrowly elliptic to oblong-elliptic, 1.2-7.5 × 0.6-3 cm, acute to round at base, acute at apex, chartaceous to membranaceous, dark green above and pale green below when fresh, olive-green when dry, lamina sparsely to densely strigose above, densely villose-sericeous below, midrib and secondary veins densely antrorse or spreading strigose or villous; secondary veins 5-7 on each side of midrib, slightly impressed above, prominent below. *Inflorescences* cymose, dichotomous or rarely trichotomous, with a solitary branch at axil of each branching, 5-20-flowered, slightly expanding after anthesis, 5-16(-24) cm long, peduncle 1-8 cm long, densely hirsute or villous, simple, without later branches, or with 1-3 pairs of lateral branches; lateral branches 3-14(-18) cm long when fully expanded, dichotomously branched, ultimate branches scorpiod, 3-7-flowered; bracts subtending branchlets lanceolate, 7-10 × 0.7-2 mm, densely hirsute or villous; bracteoles opposing each flower narrowly triangular to linear, 3-7 × 0.4-0.7 mm, densely hirsute or villous. *Flowers* heterostylous, 5-merous, commonly sessile, the central ones at base of inflorescence bifurcations with pedicels to 1.5 mm long, glabrous or puberulous. *Hypanthium* obconical to obovoid, 2.5-5 × 1.5-2 mm, acute at base, round at apex, densely antrorse hispid or sericeous, hairs slightly enlarged-tuberculate at base. *Calyx* lobes subequal to unequal, green when fresh, linear to lanceolate, 4-6 × 0.3-0.6 mm, sparsely antrorse hispidulous to sericeous; with 1-2 colleters in each calcine sinus, linear, 0.3-0.4 mm long. *Corolla* hypocrateriform, (8.5-)9.5-16.5 mm long, pink, with a wide ring of yellow hairs at tube distal portion inside and at corolla mouth; tube (6-)7-12.5 mm long, narrowly cylindrical at lower portion and slightly expanded at point of stamen insertions, 1-1.2 mm at base, 1.5-2 mm wide at mouth, sparsely sericeous outside, glabrous or puberulent below stamen insertion and densely antrorse-pubescent in area surrounding stamens inside; lobes oblong to obovate, 2.5-5 × 1.5-3 mm, round at apex, reflexed at anthesis, glabrous, margins villous-pubescent. *Medium-styled flowers* (in corolla tubes 7.5-9 mm long): stamens inserted
<figure><img src="image_49.png" /><figcaption>**Figure 45.** *Sipanea hispida*. **A.** Habit. **B.** Longitudinal section of hypanthium, ovary and calyx. **C.** Inflorescence branchlet, with flower bud and capsule. **D.** Node with stipule and petioles. **E.** Seeds. **F.** Stamen. **G.** Dissected corolla, and detail of hairs at distal and medio-basal portion of the corolla tube. Illustration by Bruno Manara (Reproduced with permission from Flora de Venezuela vol. 9(1), fig. 58. 1974).</figcaption></figure>

below the tips of the style branches, at 3–3.5 mm (3/10 to 1/2 of the corolla tube) from base; filaments 0.5–0.7 mm long; anthers linear, 2.1–2.5 × 0.2–0.3 mm, round or acute at base, acute at apex; style included, as long

as the middle of the anther or as the upper tip of the anthers, 4–5 mm long (4.5/10 to 7/10 of the corolla tube), glabrous; style branches linear, 1–1.5 mm long, acute at apex. *Short-styled flowers* (in corolla tubes 7.5–12.5 mm
long): inserted above or at the same height of the tips of the style branches, at 3-8.5 mm (2/5 to 7/10 of the corolla tube) from base; filaments 0.5-0.7 mm long; anthers linear, 2-2.5 × 0.2-0.3 mm, round or acute at base, acute at apex; style included, shorter than the points of stamen insertions, style 2-7.5 mm long (2/5 to 3/5 of the corolla tube), glabrous; branches linear, 1-1.5 mm long, acute at apex. *Infructescence* cymose, dichotomous or rarely trichotomous, slightly more expanded than the inflorescence. *Capsules* elliptic-oblong to oblong, 6-11 × 3-5 mm, acute at base and at apex, crustaceous or thinly woody, pale green when young, turning pale brown-beige when dry, densely antrorse hispid or hirsute, hairs slightly enlarged at base. *Seeds* trigonous or rhomboid-suborbicular, 0.2-0.5 mm in diam., compressed, angular or irregularly tetrahedral, testa foveolate.

**Notes:** Schumann (1889) included "S. hispida" Spruce ms." (later validly published by Wernham (1917), see below), along with several other species, in a very broadly delimited *Sipanea pratensis*. In fact, Schumann's figure of S. pratensis (Schumann, 1889, tab. 122) is instead S. hispida. Schumann's erroneous and much too broad delimitation of S. pratensis caused great confusion among Rubiaceae specialists. For additional information about the confusion between these two species, see discussion under *Sipanea pratensis*.

Hassler (1915: 170) described *Sipanea pratensis* var. *major* Hassl., citing the gathering Hassler 7733, without indicating the herbarium of deposit. At G are present six sheets of Hassler 7733. The two sheets with barcode G00307684 have the following letter glued on the first sheet "Geneve, le 22 mars 1909. Cher Monsieur, Après avoir examiné à nouveau votre *Sipanea* et l'avoir comparé aux échantillons que nous avons, j'ai constaté que votre numéro 5691 est à peu près identique à R. Spruce, nr. 2051 auquel Schumann a mis le nom de S. dichotoma H.B.K., nom qu'il a ensuite relégué dans la synonymie de son S. pratensis, qui deviant ainsi une espéce excessivement polymorphe. Votre bien dévoué, Aug. de Candelle." The text of this letter shows the confusion present in those days between *Sipanea pratensis* and *S. hispida*, and might also explain why Hassler positioned his new variety in *S. pratensis*. The two sheets with barcode G00307684 are here treated as a single specimen with multiple preparations (Art. 8.3 of the ICN; Turland et al. 2018), which is here designated as the lectotype of S. pratensis Aubl. var. major.

Wernham (1917) in the introduction of the article where he revised *Sipanea*, wrote "The types of all the novelties described are in the National Herbarium [BM]." Therefore, the specimen Gardner 5009 at BM,

barcode 000614348, is the holotype of *S. brasiliensis* Wernham (1917: 174), and the specimen *Triana 1776* at BM, barcode 000614328, is the holotype *Sipanea trianae* Wernham.

Wernham (1917) along with the description *Sipanea glabrata* Wernham cited two gatherings, collected by A. Robert near the town of Santa Anna da Chapada (now Chapada dos Guimarães), Mato Grosso, Brazil, with collection numbers 333 and 364. Both collections are mounted on the same BM sheet, with a different barcode number.

Steyermark (1967: 282) commented for *Sipanea glabrata* Wernham, "I have examined the holotype and paratype collections deposited in the British Museum (BM)." However, as he did not annotate these specimens it is impossible to know which he intended for "holotype" and which for "paratype". Therefore, the specimen Robert 333, with barcode BM 000614333, is here designated as the lectotype of this taxon because it is a more complete specimen, with flowers and fruits. Also, Wernham separated *S. glabrata* from *S. hispida* by the "nearly glabrous leaves, the obsolete stipules, and the very small corolla-limb, barely half a centimeter in diameter." However, the leaves of both Robert 333 and 364 have veins densely appressed pubescent above and below, while the lamina are either glabrous or sparsely puberulent. All the characters used by Wernham to distinguish *S. glabrata* overlap with the morphological variation of *S. hispida*, and the two taxa are here treated as synonymous.

Steyermark (1967: 279) cited the type of *S. hispida* as "Rio Negro, prope São Gabriel da Cachoeira, Amazonas, Brazil, Jan-Aug. 1852, R. Spruce 2051 (lectotype)," which is one of the gatherings cited by Wernham (1917), although without citing the herbarium of deposit; therefore, Steyermark's citation should be treated as a first-step lectotypification. The specimen Spruce 2051 at BM, barcode 000614346, is here designated the second-step lectotype of this name.

Steyermark (1967: 279-280) wrote: "*Sipanea hispida* sometimes approaches pubescent extremes of *S. pratensis* var. *dichotoma*, but the two have a different type of inflorescence; in *S. hispida* the ultimate branches of the inflorescence have the fruits much more widely separated and more strongly hispid with enlarged tuberculate bases [although they only slightly enlarged at base], whereas in *S. pratensis* var. *dichotoma* the fruits are much closer together, and much less hispid with shorter more erect and relatively fewer trichomes. Moreover, in *S. hispida* the capsules are longer, and the pubescence of the vegetative parts are denser, longer, and more villous. Where the ranges of two taxa approach one another, apparently hybrid populations are encountered. Thus,
<figure><img src="image_50.png" /><figcaption>**Figure 46.** Distribution of *Sipanea hispida*.</figcaption></figure>

same inflorescence. In order to assess the presence of heterostyly in this species, the flowers of more than 50 collections were examined. In all flowers studied, the style is included and shorter than the upper tip of the anthers. Also, two forms were detected: short-styled flowers, with stamens inserted above or at the same height of the tips of the style, at 2/5 to 7/10 from the base of the corolla tube; and medium-styled flowers, with stamens inserted below the tips of the style, at 3/10 to 1/2 from the base of the corolla tube.

collected the whole year around, with flowering peak during the rainy season.

**Suggested conservation status: Least Concern (LC).** This species has been collected in numerous localities in Colombia, Venezuela, Peru, Bolivia, Brazil, and Paraguay. In Brazil it is very common in the Cerrado Biome, in savanna formations in the Amazon Basin, in the states of Acre, Amazonas, Distrito Federal, Goiás, Maranhão, Mato Grosso, Minas Gerais, Pará, Rondônia, Roraima, and Tocantins. It is also found forming large
stands in secondary vegetation and along roadsides (pers. obs. in Brazil). Considering that this species is known by numerous collections, and is also found as a ruderal species, it is Least Concern (LC), following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** **COLOMBIA. Amazonas:**
Corregimiento de Araracuara [ca. 0°36'S, 72°23'W], trocha al Río Yarí, al NE del corregimiento, en pié de monte, 200-300 m, 18 Apr. 1986 (fl, fr), G. Galeano et al. 911 (COL); La Pedrera [ca. 1°19'S, 69°34'W], ca. 2 km en el camino entre La Pedrera y Taparacá, borde del camino en sitio abierto, 230 m, 11 Mar. 1990 (fl), G. Galeano et al. 2039 (NY); Araracuara, entre el campamento y el aeropuerto, 220 m, 3 Aug. 1977 (fl, fr), J.M. Idrobo 8874 (COL); Río Igara-Parana, prairie a bovides entretenue regulierment, sur sable et grès (La Chorrera [ca. 1°26'S, 72°47'W]), 19 Sep. 1973 (fl, fr), C. Sastre 2259 (COL, P). **Caquetá:** Mun. Valparaiso, vereda La Esmeralda, tierra firme, 1°16'20"N, 75°42'46"W, 220 m, 18 Oct. 2010 (fr), D. Cárdenas et al. 40542 (NY). **Cundinamarca:** Ubalá B [ca. 4°44'N, 73°32'W], Inspección de Policia San Pedro de Jagua, Vereda Soya, marge derecha Río Zaguea, s.d. (fl, fr), J.L. Fernandez et al. 16310 (COL); Cáqueza [ca. 4°24'N, 73°56'W], 1810 m, 23 May 1940 (fl, fr), H. Garcia Barriga 8465 (COL). **Guaviare:** San José del Guaviare [ca. 2°34'N, 72°38'W], sabana, 240 m, 4 Nov. 1939 (fl), J. Cuatrecasas 7412 (COL, US). **Guainía:** San Felipe, across river from San Carlos de Río Negro (Venezuela), on grassy Wall of fort, 1°55'N, 67°5"W, 120 m, 12 Nov. 1977 (fl, fr), R. Liesner 3395 (COL, MO). **Meta:** E del Río Manacacías, potrero sobrepastoreado, 5 Jun. 1963 (fl), J. Blydenstein 992 (COL); San Juan de Arama [ca. 3°22'N, 73°52'W], Campamento Los Micos, 350 m, 9 Aug. 1972 (fl, fr), E. Echeverry & R. Jaramillo M. 2261 (COL); Las Lagartijas, plateau between Río Papa-mene and Río Duda, Colombia-Uribe trail, 7 km SW of Uribe, clearing on hilltop, 3°12'N, 74°29'W, 1100 m, 19 Dec. 1942 (fl, fr), F.R. Fosberg 19418 (P, US); Llanos Orientales, alrededores de Villavicencio [ca. 4°8'N, 73°37'W], 450 m, cerca de Peralonso, 24-28 Jul. 1946 (fl), R. Jaramillo et al. 289 (COL, US); Villavicencio, grassy llano, 450 m, s.d., F.W. Pennell, 1450 (MO, NY); chemin longeant le Río Guatiquia, 15 km en amont de Villavicencio, 4 Mar. 1971 (fl, fr), C. Sastre 920 (COL, P); Mun. Puerto Lopez [ca. 4°5'N, 72°57'W], Hacienda Llano Grande, laguna, 7 Mar. 1971 (fl), C. Sastre & P. Pinto 1007 (COL, P); Ranch Menegua, ca. 100 km S and E of Villavicencio, 15 Jul. 1945 (fl, fr), H. Schiefer 800 (US [2 sheets]); frente al Hato Yopare, Carimagua, 200 m, 5 Oct. 1995 (fl), P. Torrijos O. & R. Serna I. 102 (COL); Andes de Bogotá, Pusumuco y Villavicencio, [400 800-1000 m, Jan. 1856 (fl, fr), J.J. Triana

3248 (COL, P). **Vaupés:** Mun. Mitú, Comunidad Indígena de Acaricuara, Paca, sector de sabanetas y catingas de 6 a 8 m de altura, com presencia de varillales, sobre arenas blancas, 0°42'29"N, 70°14'14"W, 200 m, 2 Jun. 2009 (fr), D. Cárdenas et al. 22627 (COAH, NY); Mun. Mitú, alrededores de Mitú, camino hacia el Cerro Urania, bosque de rebalse intervenido, 8 Jan. 2000 (fl, fr), R. Cortés 1579 (NY); Mun. Mitú, camino entre Bogotá Cachivera y Acaricuara, bosque de catinga alta sobre arena blanca, 16 Jan. 2000 (fl, fr), R. Cortés 1649 (NY [2 sheets]); Río Kuduyarí, Pacuativa, 380 m, 19-20 Oct. 1952 (fl), H. García-Barriga 14901 (COL [2 sheets]); Bajo Vaupés, Mitú y sus alrededores, 8 Mar. 1944 (fr), G. Gutierrez & R.E. Schultes 943 (NY), 14 Mar. 1944 (fl), 1010 (COL); Río Vaupes, a Mitu, 320 m, 7-8 Nov. 1952 (fr), H. Humbert & A. Fernandez 27241 (P); Río Vaupés, Mitú and vicinity, 250 m, 8 Sep. 1951 (fl, fr), R.E. Schultes & I. Cabrera 13957 (NY, S, US); Río Kuduyarí (tributary of Río Vaupés), middle and lower course, 230-270 m, 16 Oct. 1952 (fl, fr), R.E. Schultes & I. Cabrera 17864 (NY [2 sheets], US); Río Vaupés, Mitú and vicinity, 1°0'N, 70°5'W, 9 May 1953 (fl, fr), R.E. Schultes & I. Cabrera 19251 (NY, US); Río Vaupés, Mitú and vicinity, Sep.-Oct. 1966 (fl, fr), R.E. Schultes 24242 (COL). **Vichada:** Mun. El Viento [ca. 4°35'N, 71°4'W], Centro de Las Gaviotas, sabana no inundable, 170 m, 20 Jul. 1995 (fl), F. Cortés P. & H. Dueñas G. 322 (COL); near Saracuré, Región bocas Río Muco y San José de Ocuné [ca.4°14'N, 70°19'W], 13 May 1945 (fl, fr), F. di Giovanni s.n. (COL 000410421); hacienda de Hector Pérez, Saracuré, on the Río Vichada, ca. 40 km W of San José de Ocuné, dry grassy llanos, 100 m, 18 Jan. 1944 (fl, fr), F.J. Hermann 10959 (US).

**VENEZUELA. Amazonas:** Alto Orinoco, La Esmeralda [3°10'N, 65°32"W], rather rare in the Savannas along the river, 14 Jul. 1951 (fl, fr), L. Croizat 84 (NY [2 sheets]); Dto. Atures, near Puerto Ayacucho, ca. 3 km N of town, sabana arenosa, on L side of rd towards El Burro, 5°43'N, 67°30'W, 85 m, 29 Jan. 1978 (fl, fr), O. Huber & J. Cerda 1480 (NY [2 sheets], VEN); Dto. Atures, faldas orientales del Cerro Yaví, Laguna de Maguari, 5°45'N, 65°48'W, 300 m, 17 Aug. 1978 (fl, fr), O. Huber & J. Cerda 2261 (NY, VEN). **Bolívar:** Morichal 1 km E of ranch house, 6°45'N, 63°30'W, 420 m, s.d., J.J. Wurdack & N.G.L. Guppy 34 (NY). **Carabobo:** El Paito, 1 Sep. 1951 (fl), Hermano Gines 4538 (US). **Miranda:** Path from Petare [ca. 10°28'N, 66°46'W] to La Guayrita, 900 m, 11 Dec. 1938 (fl, fr), A.H.G. Alston 5477 (BM); Los Teques [ca. 10°20'N, 67°2'W], in German Plantations (Parque de los Bárbaros), 1400-1500 m, 19 Oct. 1917 (fl), H. Pittier 7507 (NY, US, VEN). **Yaracuy:** El Amparo, 7 km N of Salom [ca. 10°11'N, 68°28'W], 1100-1200 m, 1 Dec. 1974 (fr), J.A. Steyermark et al. 111209 (NY, US, VEN).
**ECUADOR. Morona-Santiago:** Cantón Gualaquiza, Parroquia Bomboiza, road to Nueva Tarqui on S side of Río Cuyes, -3.441583°, -78.61631°, 850 m, 30 Jan. 2019 (fl, fr), *C. Persson et al. 3284* (GB). **Zamora-Chinchipe:** Road Zumba - La Balsa, km 23, regularly burned pastures, 4°57'S, 79°8'W, 900 m, 25 May 2001 (fl), *J.E. MadSEN & C. Rosales 8140* (AAU).

**BRAZIL: Acre:** Km 4 from Rio Branco [ca. 9°58'S, 67°48'W], on Rio Branco-Brasiléia road, roadside, 16 Oct. 1980 (fl), *S.R. Lowrie et al. 560* (INPA, NY, UB); Mun. Plácido de Castro, km 20, rod. AC-40 (Plácido de Castro-Rio Branco), beira da estrada, terra firme, 10°15'45"S, 67°36'46"W, 4 Feb. 2000 (fl), *I.S. Riveiro et al. 327* (NY). **Amazonas:** Estrada Manaus-Caracaraí, Reserva Biológica do INPA, capoeira, solo argiloso, 1978, *A. Anderson s.n.* (INPA 142512); São Gabriel da Cachoeira, 50 m N do Porto de Camanaus, terra firme, areia branca, 0°7'S, 67°4'W, 70 m, 7 Jan. 2003 (fr), *A. Antonelli 245* (GB, R); Rio Negro, São Gabriel (Rio Uaupés), campo aberto, 30 Apr. 1948 (fl, fr), *G.A. Black 48-2453* (IAN); Mun. Presidente Figueiredo, Balbina, Estrada de acesso ao antigo porto, vegetação aberta, degradada, 1°0'N, 59°0'W, 1 Aug. 2006 (fl), *J.G. de Carvalho-Sobrinho & K.M. Silva 825* (INPA); Mun. Humaitá, Estrada Porto Velho-Humaitá, BR-319, km 675, campo, 7°31'S, 63°10'W, 70 m, 5 Apr. 1976 (fl, fr), *M.R. Estrela & W. Bellusci 101-5476* (INPA); ibid., 6 Apr. 1976 (fl, fr), *M.R. Estrela & W. Bellusci 106-6476* (INPA); Jauareté [ca. 0°36'N, 69°11'W], Vaupés, Rio Negro, 22 Oct. 1945 (fr), *R.L. Frões 21251* (IAN, NY); Rio Negro, Camanáos, 22-23 Dec. 1930 (fl, fr), *E.G. Holt & E.R. Blake 570* (NY, RB); Mun. Humaitá, Fazenda Flávio Neri, S of BR-319, km 658, 7°31'S, 63°10'W, 27 Dec. 1979 (fl, fr), *A. Janssen, & I. Gemtchujnicov 96* (INPA, MG, RB, UB); Rio Negro, Umarituba [ca. 0°40'N, 67°15'W], 11 Sep. 1928 (fl, fr), *P. Luetzenburg 22180* (R); Rio Negro, Comunidade Aparecida, opposite mouth of Igarapé Tuarí, near abandoned portion of Perimetral Norte, open secondary vegetation along river, 0°20'N, 67°18'W, 6 Nov. 1987 (fl, fr), *P.J.M. Maas et al. 6937* (INPA); Caatinga do Porto de Camanaus, campina alta, 19 Oct. 1978 (fl, fr), *P.I. Madison, S. Braga & H. Kennedy PFE-447* (INPA); Rio Negro, acima de São Gabriel da Cachoeira, terra firme, capoeira, 21 Jul. 1979 (fl, fr), *L.A. Maia et al. 722* (INPA); Pr. Rio Negro, s.d. [Nov. 1819-Jan. 1820], *C.F.P. Martius s.n.* (Herb. Reg. Monac. Duplum 186) (FI-Webb); Estrada Manaus-Porto Velho, BR-319, km 530, 21 Apr. 1976 (fl, fr), *O.P. Monteiro & J. Ramos 835* (INPA); Lábrea [ca. 7°15'S, 64°47'W], na estrada do aeroporto, capoeira baixa de terra firme, 19 Jun. 1976 (fl, fr), *C.D.A. Mota, s.n.* (INPA 60348); Rio Purus, km 1-4 rd. Lábrea-Humaitá, 5 Nov. 1968 (fr), *G.T. Prance et al. 8187* (INPA,

MG, NY, S, US); Rio Negro, São Felipe, km 2 da Perimetral Norte, mata de caatinga, 21 May 1975 (fl, fr), *N.T. Silva 3862* (IAN), 3863 (IAN); Mun. Humaitá, Estrada Humaitá-Jacarecanga, km 150, 60 km S, campo natural, solo arenoso, 21 Jun. 1982 (fl, fr), *L.O.A. Teixeira et al. 1256* (INPA, MG, NY, RB); Uaupes, terreno aberto, 26 Dec. 1964 (fr), *S. Vogel 332* (INPA). **Distrito Federal:** Planaltina, CPAC, rod. BR-020, próximo ao Banco de Proteínas, 47°42'W, 1175-1200 m, 17 Apr. 1985 (fl), *S.P. Almeida 1001* (IBGE); Parque Nacional de Brasília [ca. 15°38'S, 48°1'W], 4 Feb. 1992 (fl), *M.A.G. Barros et al. 2234* (IBGE, UB), 2278 (IBGE, UB); Centro Olímpico da UnB, 7 Jun. 1985 (fl-fr), *L. Branco s.n.* (UB); Bacia do Rio Descoberto, caminho para a cidade Eclética, 19 May 1999 (fl, fr), *K. Calago & S. Silva 144* (CEN); Brasília, 15 Jun. 1979 (fr), *F. Chagas e Silva 188* (IBGE); Brasília, próximo do MBTC, 6 Jul. 1985 (fl), *C.O. Córdova s.n.* (UB); Barragem do Paranoá, 15°45'S, 47°47'W, 31 May 1983 (fl-fr), *C. Cunha 14* (UB); Brazlandia ["Brasilândia"], prox. Brasília, 20 Dec. 1960 (fl, fr), *R. Delforge s.n.* (RB108700) (RB); Parque Nacional de Brasília, campo limpo, 14 Dec. 2006 (fl), *E.B.A. Dias et al. 264* (CEN), 326 (CEN), 342 (CEN); Chacara Tororó, 26 Jun. 1985 (fl), *A.M. Dos Santos, s.n.* (UB); Brasília, área do Zoobotânico, 17 Jan. 1967 (fl, fr), *A.P. Duarte 10189* (RB, US); Fazenda Santa Cecília, Núcleo Rural Rajadinha II, mata de galeria inundável, solo argiloso, 15°45'59"S, 47°38'12"W, 992 m, 22 Nov. 2012 (fl), *M.H. Fernandes & B.R. Teixeira 275* (CEN); Incra, Núcleo Rural, 9 Nov. 1971 (fl), *M.B. Ferreira 587* (HEPH); Convenio Florestal de Brasília, 29 May 1960 (fl, fr), [without initials] *Gomes 773* (RB); Bacia do Rio São Bartolomeu, 20 Dec. 1979 (fl), *E.P. Heringer et al. 2963* (IBGE, MG, NY, UEC), 9 Jan. 1980 (fl-fr), *3090* (IBGE, MG, UB), 17 Mar. 1980 (fr), *3826* (IBGE, MG, NY, US); foz do Córrego Torto com o lago, 17 Jul. 1979 (fl-fr), *E.P. Heringer 6941* (IBGE); Rio São Bartolomeu, 25 Oct. 1965 (fl), *E.P. Heringer 10765* (IAN, NY [2 sheets], RB, UB); Córrego Papuda, 5 Jun. 1979 (fl), *A.E. Heringer Salles 98* (IBGE, NY); E of Lagoa Paranoá, 975 m, 9 Dec. 1965 (fl), *H.S. Irwin et al. 11144* (IAN, NY, UB, US); Córrego Landim, ca. 25 km N of Brasília, 850 m, 27 Jan. 1966 (fl), *H.S. Irwin et al. 12066* (IAN, NY, S, UB); Cachoeira Piripiripau, ca. 15 km S of Planaltina, 20 Feb. 1970 (fl, fr), *H.S. Irwin et al. 26440* (NY [2 sheets], RB); 1 km S of Planaltina, ca. 15°38'S, 47°40'W, ca. 900 m, 26 Jan. 1972 (fl), *J.H. Kirkbride 1697* (C, NY, P, R, UB, US); pastagem perto do Lago Paraim, 15°7'S, 47°0'W, 800 m, 24 May 1980 (fl-fr), *J.H. Kirkbride 3485* (UB); perto da DF-18, ca. 16 km E da BR-251, 16°1'S, 47°26'W, 15 Oct. 1980 (fl), *J.H. Kirkbride 3655* (NY, RB, UB[2 sheets]); Córrego bananal, 15°35'S, 47°55'W, 900 m, 19 Apr. 1983 (fl-fr), *J.H. Kirkbride 5229*
(INPA, NY, RB, UB, US), D.A.M. Netto 14 (NY, UB), M.L.M. Thomé s.n. (UB); Parque Nacional de Brasília, entorno da antiga pista de ultra leve, Cerrado ralo, 15°44'53"S, 47°55'34"W, 4 Jan. 2015 (fl), C.R. Martins 1403 (CEN); Parque Nacional de Brasília, margem da Lagoa da Meditação, solo compactado, 15°44'1"S, 47°55'39"W, 30 Nov. 2016 (fl), C.R. Martins 2318 (CEN); Córrego Santa Rita, divisa com estado de Goiás, 19 Mar. 1992 (fl), E. Melo & F. França 619 (UB); Brasília, Fundos da QI 29 Lago Sul, entrada pelo condomínio Morada Sul, 17 Mar. 1992 (fl–fr), A.E. Morbeck & D. Alvarenga 51 (IBGE); Córrego Bananal, a esquerda da pastagem e Estrada DF-02, 15°25'S, 47°55'W, 19 Apr. 1983 (fr), Novato et al. 14 (NY); Centro Olímpico da UnB, entre a quadra de futebol e o lago, 18 Nov. 1981 (fl), M.A.F. Oliveira s.n. (UB); Brasília, Rio São Bartolomeu, 28 Mar. 1963 (fl, fr), E. Pereira 7420 (RB); Brasília, Parque Flor-estal, 12 Apr. 1963 (fl–fr), J.M. Pires 9007 (UB); Fundação Zoobotânica, 30 Apr. 1963 (fl–fr), J.M. Pires et al. 9514 (RB, UB); 5 km N de Brazlândia, 24 Apr. 1982 (st), J.P. Pires 235 (CEN); Reserva Biológica de Águas Emendadas, campo úmido próximo ao Córrego Fumal, 10 Dec. 1982 (fl), A.E. Ramos 168 (HEPH, INPA, UFMT); Lagoa Bonita, ca. 30 km NE de Brasília, 21 Jan. 1983 (fl), A.E. Ramos 178 (CEN, HEPH, INPA, UFMT); Parque Nacional de Brasília, 15°53'S, 47°56'W, 28 Nov. 1990 (fl), P.C.M. Ramos 340 (UB); Samambaia, 15°52'S, 48°3'W, 1150 m, 22 Apr. 1996 (fr), J.M. Rezende 405 (CEN); Brasília, Parque Nacional, 13 Apr. 1963 (fl, fr), J. Sacco 2014 (R [3 sheets]); DF-100 sentido BsB - Formosa, Fazenda Só Frango, ao lado da Fazenda Itapeti, 15°56'S, 47°22'W, 27 Nov. 2002 (fl), A.A. Santos, J.B. Pereira & J.M. Rezende 1708 (CEN); Brasília, Parque Nacional, 13 Apr. 1963 (fr), E. Santos & J. Secco 1781 (NY); Parque Nacional de Brasília, Trilha Cristal Água, Cerrado sensu stricto perturbado, 18 Jan. 2007 (fl, fr), J.R. Santos, C.A.F. Neiva & L.M.G. Nogueira 969 (CEN); Centro Olímpico da UnB, May. 1983 (fl–fr), A.F. Neto 5 (UB), 1984, M.N. Sato 23 (UB); Brasília, 15 Jul. 1979 (fr), F.C. Silva 188 (NY, UB); Reserva Ecológica do IBGE, 15°57'S, 47°52'W, 25 Apr. 1988 (fr), M.A. da Silva et al. 581 (IBGE, RB, US); Reserva Ecológica do IBGE/ Fazenda Água Limpa, UnB, cerca ao S da reserva, 15°57'S, 47°53'W, 8 Jan. 1990 (fl), M.A. da Silva & E.C. Lopes 924 (IBGE); Mun. Guará, Condom. Lúcio Costa, Reserva Ecologica do Guará, 15°48'26"S, 47°59'3"W, 1120 m, 22 Nov. 2001 (fl), M.A. da Silva 6049 (IBGE, UFG); Brazlân-dia, Fazenda Velho Barreiro, 24 Jun. 1985 (fl–fr), H.B. Souza s.n. (UB); Centro Olímpico da UnB, beira do Lago Norte, 15 Jun. 1983 (fl–fr), J.L. Sousa 15 (UB); Reserva Ecológica do IBGE, 15 Dec. 2015 (fl, fr), V.C. Souza, T.B. Flores, G.D. colletta, R.L.G. Coelho & M.A.S.S.C. Souza

39639 (RB, SP); Parque Mansões de Recreio Estrela Dalva 7, 40 km em linha reta ao S do centro de Brasília (8 km em linha reta ao N de Luziânia), 7 Dec. 1987 (fl), *Taxonomy Class of the UnB 1236* (UB); Brasília, margem esquerda do Lago Paranoá, capoeira, campo sujo, latos-solo vermelho, 20 Jun. 2001 (fl, fr), *Taxonomy Class of the UnB s.n.* (INPA); Universidade de Brasília, Campus Universitário, centro Olímpico, 13 Jun. 1984 (fl, fr), S.C. Ulhoa 22 (NY, UB), 1984 (fl, fr), M.N. Sato 23 (NY); Fazenda Sucupira, após a área de estudo do SPSB, campo sujo, 2 Mar. 2007 (fl, fr), G.D. Vale, J.B. Pereira & C.G. Fontes 352 (CEN, RB); Planaltina, 3 km W da DF-130, próximo ao Córrego Saco dos Pilões, afluente do Rio São Bartolomeu, 15°44'S, 47°41'W, 1100 m, 20 May 1992 (fr), B.M.T. Walter et al. 1425 (CEN, NY, RB).
**Goiás:** Mun. Goiás, Serra do Brumado, cerrado, 30 Mar. 1996 (fl, fr), M.B. Alcântara et al. s.n. (NY, UFG 18972); Serra dos Pireneus, ca. 15 km (straight line) N of Corumbá de Goiás, 1230 m, 16 Mar. 1973 (fl), W. Anderson 10463 (MO, NY, UB, US); Goiania, Dec. 1936 (fl, fr), A.C. Brade 15462 (RB); prope urbem Goyaz [now town of Goiás, also known as “Goiás Velho”, 15°56'S, 50°8'W], s.d. (fl, fr), W.J. Burchell 6497 (BR); Mun. Mineiros, Parque Nacional das Emas [ca. 18°S, 53°W], próx. à ponte E.J. Amaral, 15 Feb. 1995 (fl, fr), I.B. Cassimiro 29 (NY, UFG); Mun. Luziânia, Hidrelétrica de Corumbá III, área de influência, 16°40'35"S, 47°59'25"W, 868 m, campo limpo úmido, 11 Dec. 2007 (fl), C.H.G. Cezare et al. 574 (UB); Mun. Caldas Novas [ca. 17°44'S, 48°37'W], 3 km da Alternativa 15, 580 m, 27 Apr. 1993 (fl–fr), S.P. Cordovil et al. 286 (CEN, NY); Mun. Cocalzinho, BR-414, direção para Niquelândia, 100 m antes do trevo para Brasília, floresta densa, 1180 m, 15°45'30"S, 48°45'14"W, 16 Dec. 2004 (fl), P.G. Delprete & L.F. da Silva 9171 (NY, UB, UFG); Mun. Cocalzinho, Parque Estadual dos Pireneus, formação rupestre desde a base até o topo do Morro Cabeludinho, 15°48'40"S, 48°48'54"W, 1280–1320 m, 10 Dec. 2005 (fl, fr), P.G. Delprete et al. 9395 (CAY, NY, RB, SPF, UB, UFG); Mun. Pirenópolis, Serra dos Pireneus, margem da estrada sentido Vereda do Cristal, 15°49'22"S, 48°53'20"W, 1040 m, 18 Mar. 2006 (fl), P.G. Delprete et al. 9545 (CAY, HTO, NY, RB, UB, UFG); Mun. Cocalzinho, Parque Estadual dos Pireneus, ca. 700 m depois do portal do parque do lado de Cocalzinho, campo limpo estacionalmente encharcado, 15°47'56"S, 48°49'7"W, 1170–1180 m, 15 May 2006 (fl), P.G. Delprete & L.B. Bosquetti 9780 (BR, RB, UFG); Mun. Pirenópolis, Serra dos Pireneus, estrada para Vereda dos Cristais, entre o mata-burro e a casa de Fernando Maduenho, campo limpo, 15°49'19"S, 48°53'22"W, 950–1000 m, 16 Mar. 2007 (fl), P.G. Delprete et al. 10022 (MBM, UB, UFG); Mun. Pirenópolis, Serra
dos Pireneus, em direção do Portal São Lázaro, entre a
Cachoeira do Coqueiro e a Cachoeira da Santa, área de
nascentes, com solo constantemente encharcado,
15°47'29"S, 48°54'1"W, 1020 m, 27 May 2007 (fl), P.G.
*Delprete & V.L. Gomes-Klein* 10202 (UFG); Mun.
Pirenópolis, Serra dos Pireneus, estrada para a subida da
serra desde Pirenópolis, primeira estrada a direita em
direção da Cachoeira da Andorinha, bosque estacional
semicaducifolio na beira do riacho, 15°50'23"S,
48°55'36"W, 800 m, 19 Aug. 2007 (fl), P.G. *Delprete et al.*
10301 (UFG); Mun. Pirenópolis, Serra dos Pireneus,
estrada para a subida da serra desde Pirenópolis, primei-
ra estrada a direita em direção da Cachoeira da
Andorinha, vegetação na beira do riacho, 15°50'23"S,
48°55'36"W, 800 m, 21 Dec. 2007 (fl), P.G. *Delprete &
Estudantes da UFG* 10513 (CAY, NY, RB, UB, UFG);
Mun. Caldas Novas, estr. para Alternativa 4, ca. 5,7 km
da estr. UHE-Corumbá-Caldas Novas, 17°54'S, 48°3'W,
9 Feb. 1993 (fl–fr), *T.A.B. Dias et al.* 289 (CEN); Mun.
Caldas Novas, estr. para UHE-Corumbá-Caldas Novas,
ca. 13 km, 17°58'S, 48°35'W, 12 Feb. 1993 (fl), *T.A.B.
Dias et al.* 390 (CEN); estr. Brasília-Unaí, 17 Oct. 1965
(fl, fr), *A.P. Duarte* 9350A (RB [2 sheets], US); Mun.
Mossâmendes, Fazenda do Jander (Estância Quinta da
Serra), Serra Dourada, cerrado típico, 16°2'S, 50°3'W, 4
Feb. 2009 (fl, fr), *J.E.Q. Faria et al.* 409 (CEN); Mun.
Mineiros, Parque Nacional das Emas, 22 Dec. 1991 (fl),
*H.D. Ferreira* 3021 (UFG); Mun. Pirenópolis, brejo próx.
do córrego, 8 Jan. 1999 (fl), *H.D. Ferreira* 3671 (UFG),
3672 (UFG); Mun. Luziânia, Sítio Novo, 24 Nov. 1971
(fl–fr), *M.B. Ferreira* 939 (HEPH); Mun. Corumbá de
Goiás, ca. 4 km da cidade, estrada para Aparecida, 1060
m, 15°54'S, 48°45'W, 13 Mar. 2002 (fl, fr), *M.L. Fonseca
et al.* 3182 (CEN, IBGE, RB); “Goyaz,” Nov. 1839 (fl), G.
*Gardner* 3224 (BM, FI-Webb); without locality, 1841 (fl),
3224 (K); Planalto de Goyaz, Retiro do Bananal, 16 Feb.
1898 (fl, fr), A. *Glaziou* 21519 (BR, MPU, P [3 sheets], S);
Mun. Mossâmendes, Serra Dourada, rd. Goiânia-Goiás,
17 Mar. 1994 (fl, fr), *V.L. Gomes-Klein et al.* [additional
collectors not cited] 2210 (NY, UFG); Mun. Jataí, Bom
Sucesso, 17 Nov. 1973 (fl, fr), *G. Hatschbach & C. Koc-
zicki* 33340 (C, MBM, MO); Mun. Uruaçú [ca. 14°S,
49°W], rod. para Niquelândia, Fazenda Guaporé, 550 m,
19 Jan. 1992 (fl, fr), *G. Hatschbach et al.* 56231 (C, MBM,
MO); Mun. Catalão, rod. BR-050, Córrego Capão
d'Antas [ca. 18°10'S, 47°56'W], 6 Feb. 1994 (fl), G.
*Hatschbach et al.* 59871 (FLOR, MBM); Samambaia, Rio
Corumbá, 8 Jan. 1967 (fl–fr), *E.P. Heringer* 11263 (MG,
RB, UB); Mun. Caldas Novas, 12 km W of Caldas
Novas, on rd Caldas Novas-Morrinhos, on side rd lead-
ing to Hotel Pousada Rio Quente, 17°44'S, 48°45'W, 23
Dec. 1974 (fl–fr), *E.P. Heringer & G. Eiten* 14200 (NY,

UB, US); Serra de Caldas Novas, Termas do Rio Quente,
4 Jan. 1977 (fl), *E.P. Heringer* 16714 (IBGE, UB); Mun.
Corumbá de Goiás, 26 Mar. 1978 (fl), *E.P. Heringer,
A.E.H. Salles & F.C. Silva* 17017 (IBGE, UB, UEC p.p.
[UEC 118192]); Luziânia, 4 Feb. 1983 (fl), *E.P. Heringer
18435* (IBGE, RB, US); 23 km E of Caiapônia, on rd to
Montevideu, 4 Feb. 1959 (fl, fr), *H.S. Irwin* 2576 (NY,
US); Serra do Caiapó, ca. 48 km S of Caiaponia, on rd to
Jataí, 800–1000 m, 17°12'S, 51°47'W, 23 Oct. 1964 (fl),
*H.S. Irwin & T.R. Soderstrom* 7264 (NY); Serra dos Pire-
neus, ca. 20 km S of Corumbá de Goiás, ca. 20 km of
Corumbá de Goiás, 1000 m, 3 Dec. 1965 (fl), *H.S. Irwin
et al.* 11011 (IAN, NY, UB); Serra dos Pireneus, 30 km N
of Corumbá de Goiás, on rd to Niquelândia, in Valley of
Rio Corumbá, ca. 1150 m, 18 Jan. 1968 (fl), *H.S. Irwin et
al.* 18816 (IAN, NY, UB); Serra Dourada, lower slopes,
pasture, ca. 30 km SE of Goiás Velho, 14°S, 50°W, 650
m, 22 Jan. 1966 (fl, fr), *H.S. Irwin, R. Souza & R. Reis
dos Santos* 11978 (NY); ca. 15 km SE of Guará, on rd to
Tupirama, ca. 500 m, 21 Mar. 1968 (fl), *H.S. Irwin et al.*
21605 (IAN, NY, RB, UB, US), 21612 (IAN, NY, RB, UB,
US); Contraforte Central, ca. 24 km NE of Catalão, 875
m, 22 Jan. 1970 (fl), *H.S. Irwin* 25101 (UB); ca. 15 km S
of Niquelândia [ca. 14°28'S, 48°27'W], ca. 750 m, 22 Jan.
1972 (young fr), *H.S. Irwin, W.R. Anderson et al.* 34756
(NY, UB, US); ca. 17 km S of Niquelândia, ca. 750 m, 23
Jan. 1972 (young fr), *H.S. Irwin et al.* 34901 (NY, UB,
US); Abadiana, Fazenda S. Mateus, 11 Apr. 1968 (fl),
*A.M. Lima* 69 (RB, UB); Mun. Jataí, localidade Queixa-
da, 10 Dec. 1948 (fl), *A. Macedo* 1466 (NY, S, US); 20 km
from Jataí on rd to Goiânia, 1 Oct. 1963 (fl), *B. Maguire
et al.* 56988 (MG, NY, UB); Mun. Luziânia, Santo
Antônio do Descoberto, 27 Mar. 1980 (fl), *R.C. Mendon-
ça* 101 (IBGE); Mun. Leopoldo de Bulhões, 16°34'S,
48°43'W, 20 Mar. 1989 (fl), *R.C. Mendonça* 1320 (IBGE,
UB, US); Mun. Pirenópolis, Parque Estadual dos Pire-
neus, 28 Feb. 2004 (fl, fr), *S.C. Miranda* 1003 (UFG);
Mun. Aparecida de Goiânia, Chacara Jatobá, 26 Dec.
2002 (fl), *J.F.B. Pastore* 203 (CEN); Mun. Corumbá de
Goiás, Serra de Salto Corumbá, 15°50'59"S, 48°45'41"W,
988 m, 15 Feb. 2016 (fl, fr), *J.R.L. Paz* 49 (UB); km 18 N
of Aragarças, ca. 15°51'S, 52°15'W, 14 Jan. 1968 (fl),
*D. Philcox & A. Ferreira* 4037 (NY, P, RB, UB); Villa Boa
[now town of Goiás], s.d. (fl), *J.E. Pohl s.n.* (BR 824753)
(BR); “Megaponte” [sic! Meiaponte, now town of
Pirenópolis], s.d., *J.E. Pohl* 1130 (878d) (NY, W); Mun.
Padre Bernardo, Fazenda Lagoa Santa, 18 Nov. 1993 (fl,
fr), *C.E.B. Proença* 1059 (UB); Paraúna, caminho para
ponte de pedra, Ponto 7, 17°10'22"S, 50°50'4"W, 673 m,
17 Jan. 2013 (fr), *C.E.B. Proença et al.* 4327 (UB, UC,
UEG); Mun. Luziânia, Canteiro de obras do AHE
Corumbá IV, margem direita do Rio Corumbá, 860 m,
16°20'S, 48°11'W, 9 Dec. 2002 (fl, fr), J.M. Rezend et al. 691 (CEN); rd. Goiânia-Inhumas, km 14, transição cerrado-cerradão, 17 Apr. 1968 (fl), J.A. Rizzo & A.M. Barbosa 468 (UFG); Mun. Goiânia, rod. para Guapó, etr. BR-19, 30 Dec. 1968 (fl), J.A. Rizzo & A.M. Barbosa 3229 (NY, UB); Mun. Goiânia, saída Sul, 31 Dec. 1968 (fl–fr), J.A. Rizzo & A.M. Barbosa 3268 (NY, UFG, UB); mun. desconhecido, esquerda do Ribeirão Dourado, próx. à cabeceira, 29 Jan. 1969 (fl, fr), J.A. Rizzo & A.M. Barbosa 3419 (UFG); Mun. Goiânia, rod. GO-07 (Goiânia-Guapó), 31 Jan. 1969 (fl–fr), J.A. Rizzo & A.M. Barbosa 3617 (UB); Mun. Morrinhos, rod. para Caldas Novas, Córrego Samambaia, 28 Mar. 1970 (fl, fr), J.A. Rizzo & A.M. Barbosa 4918-A (UFG), 25 Apr. 1970 (fl, fr), 5081 (UFG), 25 Apr. 1970 (fl, fr), 5102 (UFG), 28 Nov. 1970 (fl), 5759-A (UFG), 6 Mar. 1971 (fl, fr), 6023-A (NY, UFG); Mun. Goiandira, Fazenda do Chapéu, 18°0'S, 48°8'W, 22 Jan. 2005 (fl), J.A. Rizzo et al. 12860 (UFG), 26 Feb. 2005 (fr), 12983 (UFG); Mun. Pirenópolis, Terra Nostra, 6 May 2003 (fl), A.S. Rodrigues, J.R. Pará, M.M. Santiago & S.M. Delgado 115 (CEN); “Province de Goyaz”, without locality, s.d. [1816–1821], A. Saint-Hilaire Catal. C1 No. 729 (P [2 sheets]); Mun. Niquelândia, 1.5 km após a ponte sobre o Rio Tocantinzinho, sentido Colinas do Sul-Niquelândia, 17 Dec. 1999 (fr), A.A. Santos et al. 612 (CEN); Mun. Chapa dos Viadeiros, Rio dos Brancos, prox. Estrada entre São Gabriel de Goiás e São João da Aliança, campo rupestre, 18 Mar. 1976 (fl, fr), J. Semir 4261 (NY); Mun. Caldas Nova, alternativa 4 regua de leitura da vazão da hidrometria, 660 m, 17°56'S, 48°33'W, 27 Feb. 1996 (fl, fr), G.P. Silva et al. 3379 (CEN); Mun. Cristalina, margem do Rio Arrepentido, a NW da área de empréstimo, 850 m, 16°13'S, 47°20'W, 6 Mar. 2002 (fr), G.P. Silva et al. 6038 (CEN); Mun. Santo Antônio do Descoberto, margem direita do Rio Descoberto, próx a Barra com o Córrego Engenho das Lages, 900 m, 16°5'S, 48°16'W, 17 Feb. 2003 (fl, fr), G.P. Silva et al. 7100 (CEN); Mun. Niquelândia, Niquelândia, Avenida Bernardo Sayao, lote 51, 14°4'S, 48°4'W, 600 m, 6 Apr. 1988 (fl–fr), L.A. Skorupa & J.N. Silveira 341 (CEN); nos corrados abertos perto de Goyaz [now town of Goiás], Jan. 1893 (fl, fr), Ule, E. 430 (P); Mun. Niquelândia, estrada de terra para Barro Alto, 7 km após a entrada, 14°34'S, 48°41'W, 30 Nov. 1999 (fl), S.M. Verboonen et al. 135 (CEN). **Maranhão:** Mun. Imperatriz, behind Rodobrás station on S side of city of Imperatriz, opem meadow with thick grass and herbs 1–1.5 m tall, 5°33'S, 47°28'W, 6 Jan. 1970 (fl, fr), G. Eiten & L.T. Eiten 10207 (US); Mun. São Pedro dos Crentes, margem esquerdo do Riberirão da Mata, Estrada Estreito-São Pedro dos Crentes, km 75, mata de galeria, solo argiloso, 6°54'17"S, 46°55'43"W, 280 m, 9 Jan. 2008 (fl, fr), G.P. Silva & G.A.

Moreira 12469 (CEN); Mun. Carolina, Estrada Carolina-Barra do Ouro, km 4, margem esquerda do Rio Lages, mata ciliar, solo arenoso, 7°22'33"S, 47°27'57"W, 150 m, 18 Jan. 2008 (fl, fr), G.P. Silva & G.A. Moreira 12760 (CEN); Mun. Carolina, Parque Nacional Chapada das Mesas, acesso no km 596 da BR-230, 19 km E em Estrada vicinal, Ribeirão Cancela, solo arenoso, 7°6'53"S, 47°17'13"W, 192 m, 12 Apr. 2016 (fl, fr), M.F. Simon et al. 2929 (CEN). **Mato Grosso:** Mun. Aripuanã, margem MT-420, 5 km da cidade, 10°12'29"S, 59°24'26"W, floresta ombrofila alterada, 3 Jul. 1997 (fr), G.F. Árbocz et al. 4056 (UFMT); estrada Ranchão da Lagoa Engenho Velho, 15°47'11"S, 56°4'17"W, 140 m, 24 Feb. 1977 (fr), S. Assunção & A. Duarte s.n. (BR 824589) (BR); Cuiabá, margem esquerda do Ribeirão dos Barbados, muito frequente no Cerrado, solo argilo-arenoso, 5 Mar. 1977 (fl, fr), S. Assunção & A. Duarte 652 (UFMT), 653 (INPA); Chapada dos Guimarães [ca. 15°27'S, 55°45'W], Colégio Evangélico de Burití, margem do Rio Monjolinho, 6 May 1983 (fl, fr), J. Barcia et al. 1289 (R [2 sheets], US), 1294 (R [2 sheets]); Itauba, Resgate de Flora da UHE Colider, Lot G de supressão, 21 L 0681275, 8779614, 270 m, Floresta do Planalto dos Parecís, região ecotono entre floresta amazonica e cerrado, 30 Jan. 2015 (fl, fr), A.Z. Bronholi et al. 11-240 (RB); Chapada dos Guimarães, São Vicente, Cachoeirinha, 22 Mar. 1983 (fl, fr), L. Carreira et al. 613 (MG, NY); Mun. Poconé, Estrada nova para Boqueirão, Cerrado, 3 Apr. 1982 (fl, fr), C.N. da Cunha & A.L. Prado 473 (UFMT); Chapada dos Guimarães, proximo à Cachoeira da Noiva, 1 Mar. 1983 (fl, fr), C.N. da Cunha et al. 748 (MG, UFMT); Alto Paraguai, Região Sul, Estrada Cuiabá-Barra do Bugres, km 47, Fazenda São Pedro, caminho para subida da serra do Limboso, 2 Apr. 1983 (fl), C.N. da Cunha et al. 886 (MG, UFMT); rd. from Juruena to Aripuanã, 94 km NW of Juruena, 26 May 1998 (fl), Dubs, B. 2451 (K, S, UFMT, US); Rio Branco, Fazenda Rio Vermelho, próximo à Vila Boa União, 28 Nov. 1984 (fr), M. Emmerich et al. 5530 (IAN, R [3 sheets], US); Itaúba, Resgate de Flora de UHE Colíder, Floresta do Planalto do Parecís, região de ecótono entre floresta amazonica e cerrado, 21 L 0672676, 8783487, 383 m, 9 May 2017 (fl, fr), M.E. Engels et al. 5255 (RB); Chapada dos Guimaraes, arredores da cidade, cerrado, solo argilo-arenoso, 15°17'25"N, 55°48'15"W, 793 m, 10 Feb. 1978 (fl, fr), F. Ehrendorfer 2405 (UFMT); Mun. Batagunçu, Porto XV, 14 Feb. 1970 (fl, fr), G. Hatschbach 23537 (C, INPA, NY); Mun. Terenos, Cachoeirão, Cerrado junto a córrego, 13 Aug. 1970 (fl, fr), G. Hatschbach 24610 (NY); Mun. Rio Verde, Serra da Pimenteira, 28 Aug. 1973 (fl, fr), G. Hatschbach 32445 (C, US), 8 Feb. 1975 (fl, fr), 35962 (C, NY); Mun. Poconé, Rod. BR-070, km 584, 4 May 1995 (fl, fr), G. Hatschbach
et al. 62218 (MBM, US); Cuiabá, Cox. da Ponte, campos humidos, May 1911 (fl, fr), F.C. Hoene 4800 (R); Serra do Roncador, wet campo and gallery margin, ca. 60 km N of Xavantina, 550 m, 25 May 1966 (fl), H.S. Irwin et al. 15982 (NY); Campina, Serra do Amolar, 7 May 2003 (fl, fr), L. Junior et al. 333 (UFMT); 11 km S de Ponta Porá, 21 Feb. 1968 (fl, fr), A. Krapovickas et al. 14115 (C, P); Santa Cruz, Rio Jamary, Dec. 1918 (fl, fr), J.G. Kuhlmann 2364 (R [2 sheets], RB); Rio Garças, barranco pedregoso, 6 May 1958 (fl, fr), A. Lima 58-3133 (RB [2 sheets]); Capão Secco, prope S. Anna da Chapada [now Chapada dos Guimarães], in nemoribus vel uliginosis, 14 Jan. 1894 (fl), C.A.M. Lindman A2731 (S [2 sheets]); Mun. Varzea Grande, lot. 13 de Setembro, 3 Feb. 1994 (fl, fr), M. Macedo & R. Godinho s.n. (UFMG 3596) (INPA); Cuyabá [now Cuiabá], 27 Dec. 1893 (fl), G.A. Malme s.n. (S), 29 May 1894 (fl, fr), G.A. Malme s.n. (S [2 sheets]); Mun. Rosário Oeste, Sesc Serra Azul, antiga Fazenda Santo Antonio, matas da trilha a Cachoeira da Serra Azul, 14°29'44"S, 55°42'20"W, 343 m, 26 May 2015 (fl, fr), G. Martinelli et al. 18506 (CEN, RB); Xavantina, marsh, 1 km from Olária, 4 km from Xavantina, 14°38'S, 52°14'W, 5 Mar. 1968 (fl), D. Philcox & A. Ferreira 4476 (NY); Mt. Santo Antonio de Laverger, mata alagada, 25 Mar. 1982 (fl, fr), A.L. Prado & H. Freitas Leitão Filho 329 (UFMT); Cuyabá [now Cuiabá], Jan. 1827 (fl, fr), L. Riedel 764 (Herb. Hort. Petropol. 137) (BR); "Santa Anna da Chapada" [Now Chapada dos Guimarães], 28 Jun. 1902 (fr), A. Robert 364 (BM 000614334, paratype of S. glabrata), 364b (K); Poconé, Cotia [ca. 15°58'S, 56°28'W], entre Livramento e Poconé, km 39, em frente à Fazenda Assombrada (de seu Gigi), cerrado aberto, 19 Oct. 1983 (fl), N. Saddi, C.T. Rizzini & [initials unknown] Mattos Filho 3383 (RB); Mun. Chapada dos Guimarães, rodovia entre Cuiabá e Chapada, 21 Nov. 1982 (fl), J.U. Santos & Rosário, C.S. 420 (HAMAB, IAN, INPA, MG, UFMT); Salobre, Jan.-Feb. 1940 (fl), M. Santos s.n. (R 150197); Faz. Mandioré, Brasil/Bolivia, Serra do Amolar, 2 Apr. 2003 (fl, fr), M.C.V. Santos et al. 288 (UFMT); Colíder, Resgate de Flora da UHE Colider, Lote B de Supressão, [ca. 10°59'S, 55°46'W], 9 Dec. 2014 (fl), S. Sardelli et al. 237 (RB); Colíder, Resgate de Flora da UHE Colíder, lote B de supressão, [ca. 10°59'S, 55°46'W], 259 m, 19 Dec. 2014 (fl), L.F. Sardelli et al. 265 (RB); Mun. Rio Branco, Fazenda Rio Vermelho, prox. à vila Boa União, 28 Nov. 1984 (fl, fr), J.D. Silva & M. Emmerich 1002 (CEN); Mun. Luciara, 8-16 km N de Luciara, Estrada em direção da fazenda Três Pontas do Araguaia, 140 m, savanna parquet, beira da estrada, 19 Mar. 1997 (fr), V.S. Souza et al. 14628 (UFMT); Mun. Alta Floresta, Fazenda Pontal, ca. 90 km SSW (em linha reta) de Alta Floresta, transição savanna florestada-floresta onbrófila aberta, 10°33'19"S,

56°15'54"W, 19 Apr. 1997 (fr), V.S. Souza et al. 15109 (UFMT); Mun. Alta Floresta, Fazenda Brasil, ca. 80 km SSW (em linha reta) de Alta Floresta, 10°32'S, 56°20'W, 21 Apr. 1997 (fr), V.S. Souza et al. 15225 (UFMT); Mun. Matupá, Sítio Amutum, ca. 50 km NE de Amutá, 9°49'S, 54°40'W, contato floresta estacional-floresta ombrófla, 26 Apr. 1997 (fr), V.S. Souza et al. 15742 (UFMT); Mun. Tapurah, Fazenda São Jacó, 12°30'S, 56°15'W, contato floresta estacional-floresta ombrófla, 8 Jun. 1997 (fl), V.S. Souza et al. 17423 (UFMT); Mun. Aripuanã, Estrada velha Aripuanã-Castanheira, arredores de Serra Morena, 10°17-19'S, 59°12-18'W, savanna arborizada sobre solo arenoso (fl), V.S. Souza et al. 18488 (UFMT); Mun. Maracapu, Fazenda Santo Antonio, Propriedade do Dr. Alfredo Neder, 590 m, 29 Dec. 1973 (fr), D. Sucre 10580 (RB [4 sheets], US). **Mato Grosso do Sul:** Corumbá, povoado do Amolar, Capoeirinha, 18°1'22"S, 57°30'15"W, 29 Nov. 2010 (fl, fr), R. Borges et al. 1083 (RB); Bataguaçu [Bataguassu, ca. 21°42'S, 52°25'W], beira da Estrada várzea aluvional do Río Paraná, brejo, 20 Nov. 982 (fr), I. Cordeiro et al. 982 (UFMT); Mun. Três Lagoas, Retiro dos Telhados, 30 km da cidade, 20°47'S, 51°41'W, 21 May 1964 (fr), J. Correa Gomes Jr. 1752 (UB); Mun. Três Lagoas, Córrego da Onça, Faz. Iamaguti, 22°S, 49°W, 7 Jul. 1964 (fl, fr), J. Correa Gomes Jr. 2076 (UB); Mun. Aquidauana [ca. 20°28'S, 55°47'W], Piraputanga, 18 Feb. 1970 (fl, fr), G. Hatschbach 23793 (US); Coxim [ca. 18°30'S, 54°45'W], campos humidos, May 1911 (fl, fr), F.C. Hoene 4297 (R), 4298 (R); Mun. Paranaíba, Rod. MS-306, margem do Córrego Fundo, local brejoso, 22 Mar. 2001 (fl, fr), M.D. Moraes et al. 607 (CEN); Mun. Tacuru [ca. 23°37'S, 55°0'W], ca. 10 km W of Tacurú, 8 Feb. 1994 (fl, fr), T.M. Pedersen 15974 (C); Mun. Bataguaçu, local Rio Pardo, rodovia MS-276, 5 km N de Bataguaçu, 21°40'42"S, 52°24'8"W, 22 Apr. 2002 (fl), V.J. Pott et al. 5385 (CEN, HMS); Corumbá, Serra do Amolar, Estrada do Taquaral, floresta estacional semidecidua, 18°2'58"S, 57°29'53"W, 320 m, 29 Nov. 2010 (fl, fr), A. Quinet et al. 2153 (RB); Mun. Campo Grande, edge of brejo, ca. 15 km E os Campo Grande, 4 Jul. 1946 (fr), J.R. Swallen 9608 (US). **Minas Gerais:** Lagoa Santa, s.d. (fl, fr), [without initials] Becker 229 (RB); Lagoa Santa [ca. 19°37'S, 43°53'W], Dec. 1937 (fl, fr), C.E.M. Burret & A.C. Brade 15972 (RB); Carassa [Serra do Caraça, ca. 20°5'S, 43°29'W], 1843 (fl, fr), P. Clausen 134 (P); without locality, 1841 (fl), P. Clausen 621 (FI-Webb); without locality, 1838 (fl, young fr), P. Clausen 642 (C, P [2 sheets]); without locality, s.d. (fl, fr), P. Clausen 720 (414) (FI-Webb); Uberlândia, Estação Ecológica do Panga [ca. 19°9'-11'S, 48°23'-24'W], Cerrado secundário, herbáceo, 10 Apr. 1992 (fr), FEEP 13 (NY), 15 May 1992 (fr), FEEP 100 (NY), 26 Jun. 1992 (fl, fr), FEEP 230 (NY); São
Roque de Minas [ca. 20°14'S, 46°21'W], ca. 1 km from the town, towards Serra da Canastra, campo cerrado, 12 Jan. 1994 (fl, fr), A.M. *Giulietti et al. CFCR 13576* (NY); Santa Luzia do Rio das Velhas [now Santa Luzia, ca. 19°46'S, 43°51'W], vieille route de Sabara, 25 Nov. 1893 (fl, fr), A.F.M. *Glaziou 20358* (C, P, R); Mun. Perdizes, Estação Ambiental Galheiro, Península, 13 Feb. 2004 (fl), E.K.O. *Hattori et al. 200* (UB); Mun. Perdizes, Estação Ambiental Galheiro [ca. 19°21'S, 47°17'W], Céu Cavalo, 13 Feb. 2004 (fl), E.K.O. *Hattori et al. 216* (UB); Mun. Lagoa Santa, barrancas humidas, Nov. 1915 (fl, fr), F.C. *Hoene 6456* (R); Mun. São Roque de Minas, Serra da Canastra, 3 km from São Roque de Minas to Araxá, 20°19'S, 46°31'W, 2 Nov. 1996 (fl–fr), A. *Ibáñez García & S. Bridgewater A480* (UB); wet campo, ca. 35 km NE of Patrocinio [ca. 18°56'S, 46°59'W], 1000 m, 29 Jan. 1970 (fl), H.S. *Irwin et al. 25636* (UB); ca. 12 km N of Patrocinio, cerrado and gallery forest, 1000 m, 31 Jan. 1970 (fl), H.S. *Irwin, E. Onishi, S.F. da Fonseca, R. Souza, R. Reis dos Santos & J. Ramos* 25766 (NY, RB, UB, US); ca. 2 km N of Paracatú, rocky hillside, 700 m, 3 Feb. 1970 (fl), H.S. *Irwin et al. 25926* (UB); São Gonzalo do Abaeté [ca. 18°20'S, 45°49'W], 7 Jan. 1989 (fl, fr), A. *Krapovickas & C.L. Cristobal 42819* (C); Riacho Fundo [now Santana do Riacho, 19°10'S, 43°42'W], Mar. 1835 (fl, fr), P. *Lund s.n.* (C); prope Curvello, Mar–Apr. 1835 (fl, fr), *Lund, P. s.n.* (C); Cachoeira Dourada, margem do Rio Parnaiba, 14 Aug. 1945 (fl, fr), H.L. *Mello Barreto s.n.* (RB [2 sheets]); Mun. Lagoa Santa, Santa Luzia, vallo em capoerinha, 17 Dec. 1937 (fl, fr), H.L. *Mello Barreto & A.C. Brade 10158* (R); Mun. Lagoa Santa, vallo no cerrado, 6 Jan. 1940 (fl), H.L. *Mello Barreto 10515* (R, US); Mun. Perdizes, Estação Ambiental Galheiro, cerrado próximo da caixa d'água, 20 Dec. 2002 (fl), S. *Mendes et al. 361* (UB); Mun. Caeté, Campo dos Coelhos, entre União e Curralinho, campo pedregoso, 26 Nov. 1942 (fl), G. *Mendes Magalhães 2299* (US); Mun. Monte Carmelo [ca. 18°43'S, 47°29'W], Alagoas, Brejo Seco, 3 Jun. 1940 (fr), J.E. *de Oliveira 45* (IAN, US); Mun. São Roque de Minas, Vargem Grande, 20°20'9"S, 46°25'20"W, 6 Jan. 2008 (fl), C. *Proença & S.A. Harris 3473* (UB); without locality, Nov. 1828 (fl, fr), L. *Riedel 876* (Herb. Hort. Petropol. 138) (P, RB); Serra do Curral, Nova Lima, BH, 13 Mar. 1955 (fr), L. *Roth 1729* (RB); Mun. Unai, mata abaixo do vertedouro, mata cilliar de encosta, solo argilo-arenoso, 16°13'13"S, 47°19'29"W, 25 Jun. 2002 (fr), A.A. *Santos et al. 1289* (CEN); Mun. Ibiá, cerrado, rd. MG-230, 2 km from BR-262, 19°34'51"S, 46°29'47"W, 2 Mar. 1989 (fl, fr), B.M.T. *Walter et al. 42* (US); Mun. Pratinha, cerrado, dirt rd. connecting BR-262 and town of Pratinha, 5 km from BR-262, 19°38'4"S, 46°28'39"W, 2 Mar. 1989 (fl, fr), B.M.T. *Walter et al. 67* (SP, UB, US);

Mun. Cabeceira Grande, região da ponte sobre o Rio Preto, 29 km from jct with BsB/Unai/Palmital, towards Palmital, divisa DF/MG, campo sujo, 16°2'15"S, 47°18'34"W, 850 m, 14 Feb. 2002 (fl, fr), B.M.T. *Walter et al. 5026* (CEN, US); Lagoa Santa, s.d. (fl), E. *Warming s.n.* (C [2 sheets]); Lagoa Santa, Oct. 1863 (fl), E. *Warming s.n.* (P P00748135); Lagoa Santa, 26 Nov. 1863 (fl), E. *Warming s.n.* (C). **Pará:** São Geraldo do Araguaia [ca. 6°24'S, 48°32'], campo cerrado, morro 3, margem da Cachoeira do Isidoro, 15 Jun. 1995 (fl, fr), M.N. *Bastos & M.R. Cordeiro 2205* (IAN, MG); without locality, “planta que nasce em terra prêta misturada com areia e barro, Fl. Março et Abril”, “Brésil, Pará, donné par l’herbier du Muséum lusitanique”, s.d. (fr), *Collector Unknown s.n.* (P [P00729409]); Mun. Anajás [ca. 0°59'S, 49°56'W], Rio Anajás, Fazenda Santa Quitéria, 20 Aug. 1982 (fr), M.R. *Cordeiro & N.A. Sena 1717* (IAN); Mun. Anajás, cidade de Anajás, ao lado do aeroporto, 20 Aug. 1982 (fr), M.R. *Cordeiro & N.A. Sena 1755* (IAN); Prainha [ca. 1°48'S, 53°28'W], Rio Marapy, Campo Alto, 15 May 1903 (fl), A. *Ducke s.n.* (MG 3575) (MG); Mun. Conceição do Araguaia [ca. 7°33'S, 49°42'W], campos gerais, 24 Jun. 1953 (fl, fr), R.L. *Frões 29913* (IAN); Reserva Florestal de Gorotire (Kayapó Indian Reservation), surroundings of Gorotire Village at Rio Fresco, 7°47'S, 51°7'W, 200 m, 19 Jan. 1983 (fl, fr), G.K. *Gottsberger & D.A. Posey 11-19183* (US); Rio Tiriós, Taperamba, 21 May 1962 (fl, fr), E. *Oliveira 1973* (IAN, UB); Rio Tiriós, lado esquerdo, parte norte ocidental do Pará, 24 May 1962 (fl, fr), E. *Oliveira 2002* (IAN); Mun. Conceição do Araguaia, 2 km W of town along hwy PA-287, 8°15'S, 49°18'W, 300 m, 24 Feb. 1980 (fl, fr), T. *Plowman et al. 9077* (IAN, INPA, MG, NY, UB, US); Mun. Palestina do Pará, inicio da mata do Río Gameleira, mata, solo-areno-argiloso, 6°7'8"S, 48°25'2"W, 160 m, 18 Apr. 2004 (fr), G.P. *Silva et al. 8794* (CEN); Marajó Island, Cuanta do Anajás, Rio Anajás, between Anajás and Vista Alegre, 0°57'S, 49°48'W, 4 Nov. 1987 (fr), G.T. *Prance et al. 30292* (HAMAB, INPA, MG); Rio Tocantins, Jacundá [ca. 4°27'S, 49°6'W], barranco da beira do rio, varzea alta, 17 May 1977 (fl, fr), M.G. *Silva & R. Bahia 3104* (INPA, MG, RB, UB); Mun. Breves, Comunidade São Pedro, capoeira baixa, 1°33'41"S, 50°27'0"W, 30 Jul. 2011 (fl, fr), M.G.C. *Souza & M.R. Pietrobom 1213* (IAN). **Rondônia:** Margins of forest on terra firme, along Rio Pacaás Novos [ca. 10°S, 63'W], between the first and second cachoeiras, 220 m, 18 Mar. 1978 (fr), W.R. *Anderson 12174* (INPA, NY, UB, US); Ouro Preto do Oeste [ca. 10°44'S, 62°13'W], mata da CEPLAC, estrada para Embratel, 11 Jul. 1986 (fl, fr), J. *Augusto et al. 1738* (R [3 mounted sheets, 2 unmounted]); Cerejeiras, ramal de acesso ao Parque Estadual de Corumbiara, beira de estrada antropizada, 13°7'26"S,
61°12'16"W, 204 m, 20 Apr. 2013 (fl, fr), A.P.B. Avaroma & F.C.P.S. Justiniano 679 (RB); Porto Velho, Fazenda Papai e Mamãe, campo cerrado, 9°39'28"S, 65°9'60"W, 132 m, 7 May 2013 (fr), N.C. Bigio et al. 850 (RB); Estrada Guajará Mirim-Abunã, trecho entre km 45–60, linha 4A do INCRA, 1 Feb. 1983 (fl, fr), L. Carreira et al. 476 (INPA, MG, NY); BR-364, area brejosa com buriti, 12°23'3"S, 60°37'24"W, 348 m, 19 Jan. 1999 (fl), R. Farias et al. 304 (HUEFS, UB); Mun. Vilhena, Fazenda Vilhena do Pensamento, 12°18'18"S, 60°23'13"W, cerradão, solo arenoso, 25 May 1997 (fl, fr), I.S. Miranda & P.J.D. Silva 1576 (IAN, MG); Basin of Rio Madeira, track between Mutumparaná and Rio Madeira, 3 Jul. 1968 (fl, fr), G.T. Prance et al. 5518 (INPA, MG, NY, S, US); Madeira-Mamoré Railroad, km 217–9, 2–4 km E of Abunã, savanna, 18 Nov. 1968 (fl), G.T. Prance et al. 8592 (INPA, MG, NY, P, S, US); Vilhena, arredores do aeroporto [ca. 12°41'S, 60°5'W], campo cerrado, 4 Jan. 1979 (fl), M.G. Silva & A. Pinheiro 4139 (INPA, MG, NY, UB); Mun. Ariquemes, Mineração Mibrasa, Setor Alto Candeias, km 128, 10°35'S, 63°35'W, SW of Ariquemes, 11 May 1982 (fl, fr), L.O.A. Teixeira et al. 359 (INPA, MG, NY, RB, US), 13 May 1982 (fl, fr), 419 (MG, NY, RB, US). **Roraima:** along Mucaí-Caracaraí road, BR-174, km 17, 8 Nov. 1977 (fl, fr), L. Coradin & M.R. Cordeiro 1012 (IAN, NY, US); Mun. São Luiz do Anauá, Estrada Perimetral Norte (BR-210), 2 km da cidade de São Luiz, 1°5'N, 60°10'W, afloramento rochoso, 26 Aug. 1987 (fl), C.A.C. Ferreira 9228 (INPA, NY); Ilha de Maracá, sandy savanna at Santa Rosa at the E side of Island, 3°20'N, 61°25'W, 8 Oct. 1987 (fl, fr), J. Pruski et al. 3424 (INPA, NY); Fazenda Canada, margem direita do Rio Uraricuera, campo alagado temporariamente, 7 Feb. 1979 (fl), N.A. Rosa 3071 (MG, NY). **Tocantins:** Mun. Palmas [ca. 10°11'S, 48°20'W], Bosque de Biodiversidade, Campus Unitins, 1 Dec. 1999 (fl), Arnaldo 1893 (HTO, UFG); Mun. Porto Nacional, Faz. Ribeirão Manduca, margens da Carreira Comprida [agora submersa], cerrado fechado, 14 Dec. 1999 (fl), Arnaldo 2175 (HTO, UFG); Mun. Santa Rosa, Rod. Santa Rosa-Silvanópolis (TO-050), ca. 15 km de Santa Rosa, floresta de galeria com solo encharcado, 11°13'20"S, 48°10'39"W, 250 m, 3 Nov. 2005 (fl), P.G. Delprete et al. 9242 (HPL, HTO, UFG); Mun. Darcinoópolis, L margin of Rio Totantins, Córrego Pequi, gallery forest, 6°50'2"S, 47°32'5"W, 160 m, 19 Apr. 2008 (fr), J.B. Pereira et al. 13083 (CEN); Mun. Couto de Magalhães, Estrada de Pequizeiro a Couto de Magalhães, 30 km antes de Couto de Magalhães, mata secundária, 11 Jan. 1974 (fl), J.A. Rizzo 9538 (UFG, UB); Mun. Barra do Ouro, Fazenda Campo Grande, ca. 15 km do Rio Manoel Alves, plintossolo, 7°30'45"S, 47°32'56"W, 180 m, 4 Mar. 2005 (fr), G.P. Silva et al.

9783 (CEN); Mun. Ipueiras, próx. Rio Tocantins, margem da lagoa, 11°14'50"S, 48°27'31'W, 6 Dec. 2001 (fl), E.A. Soares 1842 (HTO); ca. 15 km S of Araguaína, ca. 300 m, 16 Mar. 1968 (fl), H.S. Irwin et al. 21275 (NY, UB, US). **State Unknown:** without locality, s.d. (fl, fr), Collector Unknown s.n. (BR 824748) (BR); without locality, s.d. (fl, fr), Collector Unknown s.n. (ex herbario Musei imperialis lusitaniae communicata) (P [P00729401]); “Bresil”, without locality, Collector Unknown s.n. (Fi-Webb 080085); “Bresil”, without locality, [Viagem Filosófica], s.d., A.R. Ferreira s.n. (ex herb. Lusitanic.) (Herb. Lus. M. Geoffroy) (P P00748147); without locality, s.d. (fl), A. Glaziou 21519 (C); without locality, s.d., A. Glaziou 19725 (C, P); “Andrequecé,” Oct. 1855 (fl), G.T. Reinhardt s.n. (C); “Brésil central”, 1844 (fr), M.A. Weddell 1984 (P).

**PERU. Junín:** San Ramón [ca. 11°7'S, 75°21'W], 900–1300 m, 9 or 12 Jun. 1929 (fl), E.P. Killip & A.C. Smith 24788 (NY, US); Pichis Trail, Porvenir, 1500–1900 m, 3–4 Jul. 1929 (fl), E.P. Killip & A.C. Smith 25909 (NY, US); Puerto Yessup [ca. 10°27'S, 74°54'W], 400 m, 10–12 Jul. 1929 (fl), E.P. Killip & A.C. Smith 26270 (NY, US). **Loreto:** Florída, Río Putumayo, at mouth of Río Zubineta, 180 m, forest clearing, Oct.–Dec. 1931 (fl, fr), G. Klug 2285 (BM, NY, S, US). **Pasco:** Prov. Oxapampa, Central Selva, Palcazu Valley, Izcozacin, above PEPP Project Camp, forestry experimental strip in old secondary forest, clay soil, 31 Mar. 1986 (fr), J. Salick 7302 (NY). **San Martín:** Zepelacio, near Moyobamba [ca. 6°2'S, 76°58'W], 1200–1600 m, mountain forest, Jan. 1934 (fr), G. Klug 3543 (NY, S, US); Mount La Campana, 1 Sep. 1925 (fl), D. Melin 253 (S); Otto. Lámas [ca. 6°25'S, 76°32'], bosque bajo, orilla de camino, 1000 m, 6 Apr. 1972 (fl, fr), J. Schunke-Vigo 5349 (NY [2 sheets], US); Tarapoto [6°29'S, 76°22'W], 1855–1856 (fl), R. Spruce 4602 (BM, NY); San Roque, 1350–1500 m, Jan–Feb. 1930 (fl, fr), Ll. Williams 7409 (US); Río Negro, open places among ferns, 1400 m, 18 Jan. 1961 (fr), F. Woytkowski 6224 (NY).

**BOLIVIA. Beni:** Prov. Ballivian, Isla de Espiritu, Rio Yacuma, 200 m, sabana húmeda, 13 Apr. 1981 (fl, fr), G. Beck 5376 (UB). **La Paz:** Mapiri Region, San Carlos, 850 m, 1 Apr. 1927 (fl, fr), O. Buchtien 1433 (BR, C, NY, S); Mapiri, Jul.–Aug. 1892 (fl, fr), M. Bang 1488 (BM, NY [2 sheets], R, US); Mapiri [ca. 15°18'S, 68°13'W], 830 m, May 1886 (fl, fr), H.H. Rusby 2461 (BM, NY [3 sheets], P, US). **Pando:** Nicolás Suárez, cerca de Porvenir [ca. 11°5'S, 68°54'W], en los entornos y dentro de un curiche, 21 Jan. 1983 (fl, fr), J. Fernández Casas & Susanna 8366 (NY). **Santa Cruz:** Prov. Chiquitos, ca. 10 km W of Roboré to San Pedro, then NE onto serrania, from 0.5–5 km, mosaic of Cerrado, dry forest and large rock outcrops,
18°15'S, 59°49'W, 400-600 m, 5 Feb. 1995 (fl, fr), J.R. Abbott & B. Mostacedo 16094 (NY); between Río Pirai and Río Cuchi, campos, 550 m, Jan. 1911 (fr), T. Herzog 1438 (S); Prov. Núflo de Chávez, between kms 15 and 30 from Ascensión de Guarayos on rd to Urubicha, road-side at edge of deciduous woodland rich in Orbignya palms, 15°30'S, 63°5'W, 4 Aug. 1983 (fl, fr), M.J.G. Hopkins et al. 141 (NY); Santa Cruz [Santa Cruz de la Siera, 17°47'S, 63°12'W], Dec. 1891 (fl), S. Moore 801 (BM); Prov. Guarayos [formerly part of Prov. Núflo de Chávez], 11 km N of Ascensión de Guarayos on rd. to Urubichá, forest on flat, somewhat swampy Quaternary river or lake deposit, 15°47'S, 63°9'W, 190 m, 13 Jul. 1991 (fl, fr), M. Nee 41709 (NY); Prov. Ichilo, 3 km SW of El Hondo, "potrerillo", Cerrado savanna vegetation S of the buildings, sandy soil, 17°40'20"S, 63°27'25"W, 400 m, 12 Feb. 1994 (fl), M. Nee 44897 (NY); Santa Cruz, Jun. 1865 (fl, fr), R.W. Pearce s.n. (BM); Prov. Sara, campos de Buena Vista [ca. 17°40'S, 63°44'W], 500 m, 27 Mar. 1921 (fl), J. Steinbach 5481 (NY); Prov. Sara, campos de Buena Vista, 450 m, 8 Mar. 1925 (fl, fr), J. Steinbach 6965 (BM, NY, S); San José [now San José de Chiquitos, 17°51'S, 60°45}], 600 m, 2 Feb. 1902 (fr), R.S. Williams 383 (BM, NY); Santa Cruz, Chiquitos, Santiago de Chiquitos, ca. 1 km del pueblo, alrededores del campo militar, cerrado con arbustos y arboles disperses, suelo arenoso y afloramientos rocosos (campo rupestre), pocas plantas el lugar más húmedo, arenoso, 18°21'5"S, 59°36'8"W, 590 m, 12 Apr. 2008 (fl, fr), J.R.I. Wood et al. 24433 (UB).

**PARAGUAY. Amambay:** Entre Bellavista y San Carlos [San Carlos del Apa], estero húmedo próximo al Apa, 12 Feb. 1982 (fl), J. Fernandez Casas & J. Molero 6268 (NY); Zwischen Río Apa und Río Aquidaban, Caballero-rué, 1908-1909 (fl, fr), K. Fiebrig 4799 (BM); Parque Nacional Cerro Corá, 22°40'S, 56°5'W, open savannah dominated by Anadenanthera, Gochnatia, Styrax, many myrtaceous shrubs, etc., 1 Nov. 1983 (fl, fr), W. Hahn 1769 (US); Cerro Corá, Arroyo Aceite, 40 km de P.J. Caballero, sobre la barranca del arroyo, 24 Feb. 1968 (fl), A. Krapovicas et al. 14179 (C); Parque Nacional Cerro Corá, 22°39'S, 56°3'W, open campos, S side of Cerro Tujá-og, 20 Feb. 1982 (fl, fr), J.C. Solomon et al. 7137 (MO, US); Bella Vista, 15 Dec. 1983 (fl, fr), R. Vanni et al. 293 (C); Parque Nacional Cerro Corá, 300 m, 21 Jan. 1981 (fl, fr), M. Vavrek 134 (MO, US). **Canindeyú:** Serra de Maracayú [ca. 24°20'S, 55°3'W], yerbales, 1898-1899 (fl, fr), E. Hassler 5961 (BM, MPU, NY, P); 65 km S of Capitán Bado, on frontierline [ca. 23°16'S, 55°32'W], cerrado-pantanal, 550 m, 29 Mar. 1983 (fr), J.E. Simonis 227 (UB). **Concepción:** Arroyo Trementina, 15 km E of Paso Barreto, gallery forest "chaco" type, 23°2'39"S, 57°0'41"W, 210 m, 1 Jul. 1994 (fl, fr), E.M. Zardini &

L. Guerrero 39905 (NY); Yaguareté Forest, inundated savanna on black soil, 23°47'48"S, 56°6'42"W, 210 m, 18 Jan. 1996 (fl, fr), E.M. Zardini & L. Guerrero 44323 (P). **San Pedro:** Distr. Lima, Estancia Carumbé [ca. 25°33'S, 56°40'W], 24 Nov. 1969 (fl, fr), T.M. Pedersen 9383 (C, P); Alto Paraguay, Primavera, 6 Jun. 1958 (fl, fr), A.L. Woolston 991 (C [2 sheets], MG, NY, S, US). **Province Unknown:** Without locality, s.d. (fr), S.G. Weddell 3272 (P [2 sheets]), 3274 (NY).

**URUGUAY:** "Banda Oriental del Uruguay", s.d. [1816-1821] (fl), A. Saint-Hilaire Catal. C2 No. 2278 (P [2 sheets, P00729293, P00729294]).

**Hybrid forms between S. hispida and S. pratensis var. dichotoma:** Intermediate forms, as already indicated as "introgression" by Steyermark (1967: 275-276), are found in a few specimens from Colombia, where both taxa occur, are here interpreted as hybrid forms between S. hispida and S. pratensis var. dichotoma. Two gatherings from Colombia show intermediate characters between these two taxa, as spreading pubescence, typical of S. hispida (vs. appressed pubescent in S. pratensis) and corolla tubes 11-14 mm long, typical of S. pratensis var. dichotoma (vs. 6-10(-12.5) mm long in S. hispida). This combination of intermediate characters is present in the following gatherings:

**COLOMBIA: Cundinamarca:** Caqueza, 1810 m, 23 May 1940 (fl-fr), H. García Barriga 8466 (COL, US). **Meta:** Los Llanos, Río Metica, Puerto López, 380 m, 14 Oct. 1938 (fl-fr), J. Cuatrecasas 3567 (US).

8-5. **Sipanea pratensis** Aubl., Hist. Pl. Guiane 1: 148, tab. 56. 1775. [Section *Sipanea*] (Figures 5E-F, 47-50).

For synonyms and types see under varieties.

*Perennial herb or subshrub* (basal internodes woody), 0.3-1.5 m tall, terrestrial or epipetric on inselberg or granitic outcrops, basal portion sometimes woody, prostrate, decumbent, ascending or erect, often forming large patches (var. *pratensis*) or large populations (var. *dichotoma*); stems to 1.8 m long, rarely or often geniculate, sometimes rooting at basal nodes; young branches thin to stout, 0.6-3.3 mm thick, terete or sometimes slightly quadrangular in older stems, sparsely to densely, appressed or spreading, pubescent or sericeous (hairs whitish or beige, 0.8-2 mm long). *Stipules* adnate to the petioles, sheathing, free portion truncate to broadly triangular at base and very narrowly triangular to linear distally, 2-6 × 0.6-1.5 mm, appressed pubescent to sericeous outside (hairs whitish or beige,
1.5-2 mm long); glabrous inside, slightly thickened at margin inside; with 2-3 colleters inserted at margin on each side of the central lobe; margins pubescent to sericeous. *Leaves* sessile to short-petiolate; petioles (when present) to 5 mm long, hirsute-pubescent, villous, appressed-pubescent or strigose-sericeous (hairs 1-2 mm long); blades broadly ovate, ovate, ovate-elliptic, elliptic, oblong-elliptic, oblong-lanceolate, lanceolate, narrowly lanceolate to narrowly oblong-elliptic, 1.5-7.5 × 0.5-2.5 cm, round, obtuse to acute at base, rarely decurrent, acute to acuminate at apex, chartaceous, dark green above and pale green below when fresh, olive-green when dry, appressed pubescent to densely short- to long-pubescent above, (hairs 0.3-2 mm long) above, densely appressed pubescent or villous or short- to long-hirsute below (hairs 0.3-2 mm long); secondary veins 3-8 on each side of midrib, obsolete or slightly impressed above, prominent below. *Inflorescence* terminal, sessile or pedunculate, congested-cymose, appearing fasciculate, rarely with secondary branches to 4 mm long, 4-8-flowered, sometimes subtended by 2 or 4 leaf-like bracts, with heads 0.4-1.5 × 0.3-1.5 cm (corollas excluded) during anthesis (var. *pratensis*), or evidently branched during anthesis (var. *dichotoma*), 4-37-flowered, 1-10 × 0.8-7 cm (excluding corollas) during anthesis, with lateral axes 0.5-7 cm long, more or less elongating during and/or after anthesis, 1-10 cm long (excluding corollas), scorpiod; peduncles (when present) to 7.5 cm long, sparsely to densely, appressed or spreading, pubescent or sericeous (hairs whitish or beige, 1.5-2 mm long), bracts subtending secondary branches (when present) lanceolate, oblong-lanceolate, narrowly lanceolate to linear, 2.5-7 × 0.5-1.2 mm. *Flowers* distylous, (4)5(6)-merous, sessile to subsessile; pedicels (when present) to 0.7 mm long. *Hypanthium* turbinate to narrowly obovoid, 0.7-1.5 × 0.5-0.9 mm, acute at base, round at apex, appressed pubescent to sericeous. *Calyx* lobes free, equal, subequal or unequal, lanceolate, narrowly oblong-lanceolate to linear, 2.3-6 × 0.2-0.6 mm, acuminate at apex, green to yellowish-green when fresh, sparsely to densely appressed pubescent to sericeous, sometimes almost glabrous outside, margins long-ciliate, glabrous inside; with 1-2 colleters in each calcine sinus, linear, 0.2-0.4 mm long. *Corolla* hypocrateriform, 14-28 mm long, pale to dark pink, or tube pink and lobes white during anthesis, turning paler to almost white after anthesis, with a dense ring of yellow hairs exerted beyond mouth; tube (8-10)10-17 mm long, gradually expanding towards the mouth, 0.4-0.7 mm wide at base, 1.5-3 mm wide at mouth, glabrous at medio-basal portion and sparsely to densely appressed pubescent at medio-distal portion or completely glabrous outside, glabrous at basal portion,

densely puberulous above and densely yellow-pubescent at distal portion inside, the yellow pubescence exerted beyond corolla mouth; lobes round, broadly elliptic, elliptic, obovate, oblong-elliptic to oblanceolate, sometimes varying in shape and size within the same population, sometimes unequal or of slightly different shape within the same corolla, (2.5-)4-11 × (2-)2.5-7 mm, round, obtuse to acute, rarely apiculate when subobtuse or acute, or indented when obtuse or round at apex, glabrous throughout. *Long-styled flowers*: stamens inserted at 6-7 mm from base or about the middle of corolla tube; filaments 0.5-0.8 mm long; anthers narrowly oblong-elliptic to linear, 1.8-2.5 × 0.2-0.3 mm, acute at both ends; style exserted 1-2.5 mm beyond corolla mouth, (9.5-)15-16.5 mm long, glabrous; style branches narrowly oblong to oblong-ovate, (0.6-)1-1.5 mm long, round at apex. *Short-styled flowers*: stamens inserted at 2.5-5.5 mm below corolla mouth; filaments 0.2-0.7 mm long; anthers linear to narrowly oblong-elliptic, 1.7-3 × 0.2-0.3 mm, acute at both ends; style included, 6-8 mm long, glabrous; branches linear, 1.5-3 mm long, round or acuminate at apex. *Infructescences* with mature fruits congested-cymose, appearing fasciculate, rarely with secondary branches to 4 mm long (var. *pratensis*), or dichotomously branched, more or less expanded, with scorpioid secondary branches (0.5-)1-10 cm long (var. *dichotoma*). *Capsules* nearly as broad as long, subglobose, ovoid to obovoid, 4-4.5 × 3-4.5 mm, round or acute at base, round at apex, strongly costate when dry (var. *pratensis*), or longer than broad, cylindrical-oblong to oblong-narrowly ovoid, (3-)4.5-8 × 1.5-3 mm, round to acute at base, round at apex, commonly smooth or faintly to strongly costate when dry (var. *dichotoma*); thinly woody, sparsely to densely pubescent to sericeous (hairs 0.5-1.5 mm long) to almost glabrate (very sparsely pubescent), pale green when young, turning beige to pale brown when dry. *Seeds* irregularly tetrahedral, faintly to strongly angular, (0.4-)0.5-0.7 mm in diam., testa foveolate, rugulose-verrucose.

**Distribution and ecology:** Distributed in Colombia, Venezuela, the Guianas, and northern Brazil (Roraima, Amapá, Pará, Amazonas, Maranhão); an extremely variable species in terms of habit, leaf shape, inflorescence architecture, flower size, and fruit shape; prostrate, decumbent, ascending or erect herb or shrub, often forming extended patches (var. *pratensis*) or large populations (var. *dichotoma*) in open savannas, white-sand savannas, brown-sand savannas, on granitic inselbergs, granitic outcrops, swampy margins in savannas, seasonally inundated savannas, open field savannas with *Trachypogon*, or savannas with sparse
Steyermark (1967, 1974) characterized the typical variety of *Sipanea pratensis* by “no part of the inflorescence elongating, the flowers or fruits all fasciculate-congested [i.e., reduced-cymose], without a subsecund monochasial or dichasial appearance, [...] capsules subglobose to ovoid, mainly 4-4.5 mm long, nearly as broad as long,” while he distinguished var. *dichotoma* by the “lateral axis of inflorescence more or less elongating, producing a subsecund monochasial or dichasial appearance, [...] capsules cylindrical-oblong, 4-8 mm long, longer than broad.” (Steyermark, 1967: 271). After a detailed study of several hundred specimens of S. pratensis throughout its geographic distribution, these characters are here confirmed to be diagnostic of these two varieties, and are correlated with geographical ranges; therefore these two varieties are here recognized.

Aside from *Sipanea pratensis*, in several other species, a certain degree of expansion of the inflorescence has been observed from condensed during anthesis to much expanded starting from the final stage of anthesis to fruit maturity, as, for example in *S. glomerata* and *S. hispida*, which have young inflorescences condensed to subcapitate, expanding into cymose with scorpiod secondary branches when the capsules reach full maturity. However, this variation in inflorescence expansion is present in various degree throughout the range, and has no correlation with any geographic distribution. Therefore, in several species the degree of inflorescence expansion is treated as a developmental stage, and no varieties are recognized in *S. glomerata* and *S. hispida*.

Bremekamp's (1936) *Sipanea pratensis* var. *glaberima*, which was treated as a distinct species by Steyermark (1967), is here maintained as such because the plant is completely glabrous, except for the corolla throat pubescence, although it is known only from the type specimen collected "in the midst of a population" of *S. pratensis* var. *dichotoma*. For additional information see discussion under *S. glaberima* (Bremek.) Steyerm.

Finally, there is a certain tendency for the populations of *Sipanea pratensis* growing on inselbergs and white-sand savannas to have a prostrate-decumbent habit and smaller leaves (i.e., approaching *Sipanea galioides* in general habit); however, these morphological features intergrade with those of nearby populations with ascending or erect habit and larger leaves, occurring at the edges of these formations, and in other open populations occurring in open fields on lateritic soils. Therefore, variation in habit and leaf size are interpreted as morphological variation influenced by ecological factors, and leaf shape and size cannot be used as taxonomic characters for supporting subspecific ranks within this species.

Key to varieties of *Sipanea pratensis*

1. Inflorescences congested-cymose, appearing fasciculate, or rarely with secondary branches to 4 mm long during anthesis, sometimes subtended by 2 or 4 leaf-like bracts (i.e., pherophylls), no part of the inflorescence elongating during and/or after anthesis (without a subsecund monochasial of dichasial appearance); infructescences congested, appearing fasciculate, rarely with secondary branches to 4 mm long; capsules nearly as broad as long, subglobose, ovoid to obovoid, 4-4.5 × 3-4.5 mm, strongly costate [Guyana, Suriname, French Guiana, Brazil (Amapá, Pará), mostly costal areas] .8-5a. *S. pratensis* var. *pratensis*

1. Inflorescence congested-cymose, appearing fasciculate, or obviously dichomously branched during anthesis, expanding after anthesis, with lateral axes 0.5-7 cm long, excluding corollas, more or less elongating during and/or after anthesis, scorpiod, producing a subsecund monochasial of dichasial appearance; infructescences dichotomously branched, more or less expanded, with scorpioid secondary branches (0.5-)1-10 cm long; capsules longer than broad, cylindrical-oblong to oblong-narrowly ovoid, (3-)4.5-8 × 1.5-3 mm, smooth or faintly costate [Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas, Maranhão, Pará, Roraima)]. 8-5b. *S. pratensis* var. *dichotoma*

8-5a. *Sipanea pratensis* Aubl. var. *pratensis*, Hist. Pl. Guiane 1: 148, tab. 56. 1775. (Figures 5E-F, 47-49).

Type: FRENCH GUIANA. "in pratis Caïennae & Guianae" [open fields of Cayenne and French Guiana] and "se trouve en abondance dans les savanes qui sont autour de la ville de Caïenne [...] presque toujours en fleur & en fruit," [it is found in the savannas that surround the city of Cayenne [...] almost always in flower and fruit], s.d. [1762-1764], J.B.C. Aublet s.n. (Lectotype P-JJR 8: 281, designated by Lanjouw & Uittien (1940: 158); isolectotype BM [barcode BM000614306], specimens with N. "1" [excluding the specimen with N. "2"; see Nomenclatural Notes below]; photo-BM at NY). [Section *Sipanea*].

(=) *Sipanea coppenamensis* Bremek., Recueil Trav. Néerl. 31: 258. 1934.

Type: SURINAME. Coppename River, near Turcotabbetje, 26 Jul. 1921 (fl), J.W. Gonggrijp 58 (B.W. 5326) (holotype U [2 sheets, barcodes 0006308 and 0006309]; photo-U at NY).

*Perennial herb* 0.3-1 m tall, terrestrial or epipetric (on inselberg or granitic outcrops), prostrate, decum-
bent, ascending or erect, often forming extended patches; stems often geniculate, basal portion prostate, rooting at nodes; young branches thin, 0.6-1(-1.3) mm thick, terete or sometimes slightly quadrangular in older stems), sparsely to densely, appressed or spreading, pubescent or sericeous (hairs whitish or beige, 1.5-2 mm long). *Leaves* with blades ovate-elliptic, elliptic, oblong-elliptic, oblong-lanceolate, lanceolate to narrowly lanceolate, 1.5-4.5 x 0.5-1.8 cm, obtuse to acute, sometimes decurrent, at base, acute to acuminate at apex; secondary veins 3-7 on each side of midrib, obsolete above, prominent below. *Inflorescences* sessile or pedunculate, congested-cymose (appearing fasciculate), or rarely with secondary branches to 4 mm long, sometimes subtended by 2 or 4 leaf-like bracts; heads 0.4-1.5 x 0.5-1.5 cm (corollas excluded) during anthesis, 4-8-flowered, no part of the inflorescence elongating during and/or after anthesis, without a subsecund monochasial of dichasial appearance; peduncles (when present) to 4.5 cm long, sparsely to densely, appressed or spreading, pubescent or sericeous (hairs whitish or beige, 1.5-2 mm long); external bracts (when present) 4-7, lanceolate, oblong-lanceolate to narrowly lanceolate, 2.5-5.5 x 0.5-1.2 mm, acute to acuminate, sparsely to densely antrorse-pubescent outside, sometimes absent; bracteoles subtending flowers (when present) lanceolate to narrowly lanceolate, 2-2 x 0.4-0.7 mm, appressed pubescent to sericeous outside (hairs whitish or beige, 1.5-2 mm long), sometimes absent. *Hypanthium* 1-1.5 x 0.7-0.9 mm, acute at base, round at apex, appressed pubescent to sericeous. *Calyx* lobes narrowly lanceolate to linear, 2.5-5 x 0.3-0.6 mm. *Corolla* 17.5-25.5 mm long; tube (10-)12.5-17 mm long, 0.4-0.5 mm wide at base, 1.5-2.8 mm wide at mouth; lobes broadly elliptic, elliptic, obovate, oblong-elliptic to oblanceolate, (sometimes varying in shape and size within the same population), sometimes unequal or of slightly different shape within the same corolla, 5-9 x 2.5-5.5 mm, round, obtuse to acute, rarely apiculate when subobtuse or acute, or indented when obtuse or round at apex. *Long-styled flowers* [Delprete 12882 (corolla tube 13 mm long), Granville et al. 15909 (corolla tube 12.5 mm long)]: stamens inserted at 6 mm from base or about the middle of corolla tube; filaments 0.7-0.8 mm long; anthers linear, 2.3-2.4 mm long; style

exserted 1-2 mm beyond corolla mouth, 14-18 mm long; style branches narrowly oblong to oblong-ovate, 1-1.3 mm long, round at apex. *Short-styled flowers* [Delprete 12882 (corolla tube 13 mm long), Delprete 12886 (corolla tube 10 mm long)]: stamens inserted at 2.5-3.5 mm below corolla mouth; filaments 0.2-0.3 mm long; anthers linear, 1.7-2.8 x 0.2-0.3 mm, acute at both ends; style included, 6-8 mm long, glabrous; branches linear, 1.5-3 mm long, round at apex. *Infructescences* with mature fruits congested-cymose, appearing fasciculate, rarely with secondary branches to 4 mm long. *Capsules* subglobose, ovoid to obovoid, nearly as broad as long, 4-4.5 x 3-4.5 mm, round at base, strongly costate when dry, sparsely pubescent to sericeous (hairs 0.5-1 mm long), pale green when young, turning beige to pale brown when dry.

**Nomenclatural notes:** A short biography of Aublet and full typification of the Rubiaceae taxa described by him, including *Sipanea pratensis*, was recently published by Delprete (2015b). Aublet (1775), for each species supplied the localities and habitat where they grew, without citing any collection number. Therefore, according to the *Code* (Turland et al. 2018), Aublet's specimens cannot be treated as single gatherings, and lectotypes or isolectotypes should be designated. Lanjouw and Uittien (1940) reported the discovery of an additional set of Aublet's specimens within the Jean-Jacques Rousseau herbarium (P-JJR). Most importantly, in Section 6 of their article, entitled (translated from French) "Types of genera and species of Fusée Aublet in the herbarium of Jean Jacques Rousseau in possession of Henri Denaiffe at Carignan (Ardennes), France," they listed all the Aublet's specimens found in this herbarium. As the title of this section begins with "Types", according to Art. 9.10 of the *Code* their statement should be treated as an error to be corrected, and the types citations in this section should be recognized as lectotypifications. There are two Aublet specimens of *S. pratensis*, one at P-JJR and the other at BM. Lanjouw and Uittien (1940: 158) cited the P-JJR specimen as "Hb. Denaiffe vol. VIII, no. 281: une branche portant des fleurs ressemblant à la planche [one branch with flowers, resembling the plate]." On sheet P-JJR 8: 281 is mounted one much-branched speci-

**Figure 47.** *Sipanea pratensis* var. *pratensis*. **A.** Habit, with flowers in anthesis and capsules. **B.** Node with sheathing stipule and leaf bases. **C.** Dissected short-styled flower, with stamens inserted at distal portion of corolla tube, and included style. **D.** Dissected long-styled flower, with stamens inserted at medio-distal portion of corolla tube, and exserted style. **E-I.** Corolla lobes variation within the species. **J.** Infructescence with mature capsules. **K.** Mature capsule. **A-B, D:** drawn from Delprete 12882 (CAY); **C:** drawn from Delprete 12886 (CAY); **E:** drawn from Hoff 5068 (CAY); **F:** drawn from Billiet & Jadin 5751 (CAY); **G:** drawn from Albertini 18 (CAY); **H:** drawn from Billiet & Jadin 4403 (CAY); **I:** drawn from Moretti 1191 (CAY); **J:** drawn from Delprete 12885 (CAY); **K:** drawn from Rova et al. 2018 (CAY). Illustration by Piero Delprete.
<figure><img src="image_51.png" /><figcaption>Figure 47.</figcaption></figure>
men, with numerous leaves and several inflorescences with flower buds and flowers in anthesis. At the upper-left corner there is a label, handwritten by Aublet, which states "Classis V, pentandria monogynia, *Sipanea* pratensis, hist. guia. franc. 148, T. 56." Therefore, sheet P-JJR 8: 281 is the lectotype of *S. pratensis*.

Steyermark (1967: 272) cited the type of *Sipanea pratensis* as follows: "Photo of type from BM, (1) Guiana, Aublet, (2) Cayenne, von Rohr." Two collections are mounted on the BM sheet, barcode 000614306, with most of the branches collected by Aublet, and one branch collected by Julius von Rohr. Aublet's specimens are identified with No. 1, and are represented by one large, much-branched specimen with several inflorescences with flowers in anthesis and flower buds (mounted at the center-left of the sheet), one branch with an inflorescence and a couple of flowers in anthesis (mounted near the right margin of the sheet), and a small sterile branch with a couple of nodes (mounted on the upper right corner of the sheet). Rohr's specimen, identified with No. 2, is a single branch (mounted on top of the distal nodes of the largest Aublet specimen) with two terminal inflorescences, one with flowers in anthesis, and the other with flower buds. At the bottom of the sheet, handwritten in pencil, probably by Solander, is "Sipanea pratensis Aublet!" And on the back of the sheet, handwritten by Solander is "Guiana. F. Aublet. 1, Cayenne. Jul. v. Rohr. 2", and there is a printed label saying "New York Botanical Garden, Neg. No. N.S. 553 [1952]." Following the type citation of Lanjouw and Uitten (1940: 158), the three specimens with No. 1 on the BM sheet are the isolectotype of *S. pratensis*. Whereas, the specimen with No. 2 was collected by Rohr and is not original material.

**Hybridization between varieties:** One gathering from the Upper Oyapock River, State of Amapá, Brazil, presents transitional characters between *S. pratensis* var. *dichotoma* and var. *pratensis*, and was collected in an intermediate locality of the distribution of the two varieties. In these specimens the inflorescences are congested during anthesis and slightly expanded, with lateral branches 6-9 mm long at fruit maturity (typical of var. *dichotoma*), while the fruits are globose and costate (typical of var. *pratensis*). This collection is:

**BRAZIL:** Amapá: Upper Oyapock River, Brazilian side ["rive brésilienne"], Roche Mon Pére [ca. 3°17'N, 52°12'W], 19 Feb. 1968 (fl, fr), R.A.A. Oldeman & C. Sastre 309 (CAY, P, U).

**Distribution and ecology:** Distributed in Guyana, Suriname, French Guiana, and the Brazilian states of

Amapá and Pará (Figure 49). Perennial herb, prostrate, decumbent or ascending (exceptionally erect), often forming extended patches in open savannas and at forest margins, at edges of and on inselbergs, road margins, and disturbed vegetation, in sunny exposures, on white or brown sand, and on granitic or lateritic soils; from near sea level to 750 m altitude.

**Phenology:** Due to the rapid phenology of this species it is common to find individuals with flowers and fruits. Flowering and fruiting the whole year around.

**Suggested conservation status:** Least Concern (LC). This variety is most common in French Guiana and in the contiguous Brazilian state of Amapá, and is less frequent in Guyana, Suriname, and the Brazilian state of Pará. It often forms large patches in open savannas and forest margins, at edges of and on inselbergs, and is also found as a ruderal species quite common at road margins, and in disturbed vegetation. Taking into account that is a ruderal species known from numerous collections, it is here positioned the Least Concern (LC) category, following IUCN criteria (IUCN 2012, 2019).

**Vernacular names:** FRENCH GUIANA: arajada (Arawak, *Capus* 215), ka?api?i (Wayapi, *Grenand* 1228), ka?api?ipilã, sə?əkɛa (Wayapi, *Sastre* 4458), Peruβia (Palikur, *Granville* 4295), Peruβia, winikei (Palikur, *Berton* 153), winikwei (Palikur, *Berton* 162).

**Uses:** One herbarium label (*Albertini* 18) reports that it is used in French Guiana to cure conjunctivitis in humans and cataracts in dogs.

**Specimens examined:** GUYANA: Potaro-Siparuni Region, Kato, 4°40'N, 59°55'W, 750 m, savanna and low forest vegetation on pale red clay soils, 11 Mar. 1989 (fl, fr), W. Hahn et al. 5587 (NY); East Coast Water Conservancy, SE of Georgetown, canal SE of Lamaha Stop-off, 27 Nov. 1919 (fl), A.S. Hitchcock 16981 (NY, S, US); without locality, s.d. (fl), *J.P.B. von Rohr s.n.* (C [2 sheets]); without locality, s.d. (fl), *J.P.B. von Rohr* 43 (C); without locality, s.d. (fl), *E. Rudge s.n.* (BM 000614350) (BM).

**SURINAME:** Doetabikinosoe, Marowyne River, Pedrosoengue-val, 1 Jul. 1939 (fr), G. Stahel 73 (NY, U).

**FRENCH GUIANA:** Piste de Saint-Elie, sideroad vegetation, 5°20'N, 53°0'W, 27 Apr. 1994 (fl), S. *Albertini* 18 (CAY); Haut Oyapock, petite ile entre Eureupou Crique et Riv. Motoura, Jan. 1948 (fl), *E. Aubert De la Rüe s.n.* (P P00729323); Haut Oyapock, Roche Sikini [ca. 3°15'N, 52°16'W], zone dénudé en foret, Vallée riv. Siki-
<figure><img src="image_52.png" /><figcaption>**Figure 48.** *Sipanea pratensis* var. *pratensis*. **A.** Habitat on roadside vegetation. **B.** Habit. **C.** Detail of inflorescences, top view. **D.** Detail of inflorescence, side view. **E.** Short-styled and long-styled flowers longitudinally dissected, from the same population. Photos taken by Piero Delprete on 19 April 2017, Commune de Matoury, RN 2, pk 10–11, French Guiana, where *Delprete 12882-12884* were collected.</figcaption></figure>
ni, 50 m, 16 Mar. 1949 (fl), E. Aubert De la Rüe s.n. (P [3 sheets, P00729314, P00729324, P00729326]); without locality, 1961 (fl, fr), A. Aubreville s.n. (P [P00719295]); N° 3, without locality, 1961 (fl), A. Aubreville s.n. (P [P00729316]); Savane Gallion, near Carrefour Gallion, 4°47'N, 52°25'W, 7 Jul. 2004 (fl), L. Barrabé & F. Crozier 55 (CAY, NY); Charvein, savane blanche, 26 Oct. 1913 (fl), R. Benoist 126 (P); Pariacabo, 10 Jul. 1914 (fl), R. Benoist 1414 (P); Oyapock River Basin, Crique Gabaret, 3°55'N, 51°48'W, 11 May 1997 (fl, fr), M.-E. Berton 153 (CAY); Oyapock River Basin, Commune de Saint-George, edge of private house, 3°53'N, 51°48'W, 10 May 1997 (fl, fr), M.-E. Berton 162 (CAY); Le Gallion (RN2), 18 Oct. 1981 (fl, fr), F. Billiet & B. Jadin 1034 (BR, CAY, U); Kourou, piste vers la Montagne de Singes, 24 Oct. 1981 (fl), F. Billiet & B. Jadin 1142 (BR, CAY); Kourou, savane récemment brulé, 27 Oct. 1982 (fl), F. Billiet & B. Jadin 1423 (BR); Piste FRG, km 87 de la RN1, PK 16, 5°11'N, 52°55'W, 13 Feb. 1988 (fl), F. Billiet & B. Jadin 4403 (AAU, BM, BR, CAY, RB); Route RN2 Cayenne-Regina, PK 67, 4°29'N, 52°19'W, 5 m, 10 Feb. 1993 (fl), F. Billiet & B. Jadin 5751 (BR, CAY, MO, P); Le Larivot, mangrove à Avicennia, 4°53'N, 52°20'W, 14 Nov. 1998 (fl), F. Billiet & B. Jadin 7099 (BR, CAY); Campo de Passoura, 21 Oct. 1954 (fl, fr), G.A. Black & Klein 54-17108 (IAN); Campo de Passoura, 25 Oct. 1954 (fl, fr), G.A. Black & [without initials] Klein 54-17216 (IAN); Savane Gabrielle, degrade d'Escole, 4°41'N, 52°18'W, 4 Aug. 1993 (fl, fr), P. Blanc & P. Birnbaum BB-93-42 (CAY, NY, U); trail to Bagne des Annamites, 31 Jan. 2003 (fl), G. Bourdy 3034 (CAY); Route de Monsinery, Kamuyene Kamuyene, 12 Apr. 2005 (fl), G. Bourdy 3124 (CAY); vicinity of Cayenne, 13 May 1921 (fl, fr), W.E. Broadway 186 (US), 398 (CAY, US); Savane incluse Agami, 5°15'28"N, 52°48'37"W, 8 Mar. 1995 (fl), L. Cadamuro & F. Solacroup 399 (CAY); Savane incluse Agami, 5°13'19"N, 52°48'59"W, 8 Mar. 1995 (fl, fr), L. Cadamuro 424 (CAY); Savane Nicole, 5°15'10"N, 52°46'41"W, 23 Mar. 1995 (fl, fr), L. Cadamuro & F. Solacroup 496 (CAY); Carrière Roche-Nicole, 23 Feb. 1994 (fl), L. Cadamuro 621 (CAY); Arawak Village of Sainte Rose de Lima, 23 May 1981 (fl), F. Capus 215 (CAY), 24 Sep. 1981 (young fr), 229 (CAY); Cayenne, s.d. (fl), Collector Unknown s.n. (FI-Webb 080083) (FI-Webb); Savanna W of Cayenne, 26 Oct. 1954 (fl), R.S. Cowan & B. Maguire 38016 (NY); Savane de Charvin, white sand overlaying laterite-clay, 19 Dec. 1954 (fl, young fr), R.S. Cowan 38869 (NY, U); Savane Bordelaise, Route du Tour de l'Ile, 25 Feb. 1977 (fl), G. Cremers 4349 (CAY, US); Massif des Emerillons, N inselberg, 350 m, 20 Sep. 1980 (fl, fr), G. Cremers 6741 (CAY [2 sheets], P [2 sheets], U, UB); Route du Tour de l'Ile de Cayenne, R.N. 2, PK 13,

28 Mar. 1986 (fl), G. Cremers 9404 (CAY); Bassin de l'Approuague, Savane Roche (Inselberg) Virginie, 4°11'N, 52°9'W, 140 m, 12 Feb. 1991 (fl), G. Cremers & P. Petronelli 11865 (CAY, P, US); Bassin du Bas-Oyapock, Savane Roche (Inselberg) Quatorze de Juillet, 3°58'N, 51°52'W, 50 m, 16 Apr. 1991 (fl), G. Cremers 12163 (CAY, NY, P, U, US); Savane de Corossony, Piste de Saint-Elie, 5°23'N, 53°0'W, 10 m, 26 Mar. 1992 (fl), G. Cremers & M.N.C. Bastos 12872 (CAY, MG); Bassin de l'Approuague, Savane Roche (Inselberg) Anabelle, 4°0'N, 52°16'W, 120 m, 10 May 1997 (fl), G. Cremers 15250 (CAY); Savane de bord de Piste St. Elie, 7 Feb. 1990 (fl), DEA Paris VI 136 (CAY); Piste de St. Elie, 5°20'N, 53°2'W, 18 Sep. 2005 (fl), C. Delnatte & C. Girod 141 (CAY), 172 (CAY); Piste de St. Elie, PK 2, savane inondée, 5°22'N, 52°59'W, 8 m, 8 Jan. 2006 (fl), C. Delnatte et al. 509 (CAY); Macouria, Savane Marivat, locality Maya, 4°56'59"N, 52°26'30"W, 15 m, 1 Apr. 2006 (fl), C. Delnatte, S. Gonzalez, & F. Crozier 713 (CAY); Cayenne, Source de Baduel, 4°55'30"N, 54°18'40"W, 14 m, 24 Jun. 2006 (fl), C. Delnatte 859 (CAY); Commune de Roura, Reserve Naturelle Volontaire de Trésor, Route de Kaw, 4°36'N, 52°16'W, 30-200 m, savanna dominated by grasses and sedges with sparse mounts with shrubs 3-5 m tall, 8 Feb. 1999 (fl), P.G. Delprete & F. Crozier 7157 (CAY); Commune de Kourou, Savane des Pères, ca. 2 km from Station Radar-Télémesure of CSG, towards Kourou, right side, savanna with sparse trees of Byrsonima crassifolia and Curatella americana, mostly grassy field, seasonally inundated, at the moment with water 5-30 cm deep, 5°7'7"N, 52°39'4"W, 30 m, 5 Jul. 2009 (fl), P.G. Delprete 10586 (CAY, GB, MO, NY, US), 10605 (CAY, F, G, L, MO, US); Commune de Macouria, Savane de Matiti, open field savanna dominated by graminaceae and cyperaceae, with sparse shrubs and small trees, including Palicourea rigida, area slightly elevated, not flooded, on rocky-lateritic soil, 5°4'10"N, 52°35'47"W, 10-20 m, 12 Jul. 2009 (fl), P.G. Delprete 10651 (CAY, NY, US); Commune de Macouria, Savane de Matiti, savanna permanently flooded, with dense shrubs, mostly Melastomacaeae, vines and small herbs, at the moment with water 10-20 cm deep, 5°4'10"N, 52°35'47"W, 10 m, 12 Jul. 2009 (fl), P.G. Delprete 10678 (CAY, GB, MO, P, US); Commune de Montsinery, Savane de Monsinery, across the road from Centre de Diffusion Internationale de Guyane, grassy savanna with sparse treelets and shrubs, sometimes with sparse islets of shrubby vegetation, 4°53'39"N, 52°30'25"W, 20 m, 18 Jul. 2009 (fl), P.G. Delprete 10704 (CAY, L, K, MO, P, US); Commune de Ira-coubo, Savanne de Trou Poisson, road towards Kaja Nostra Hotel, grassy savanna dominated by poaceae and cyperaceae, with sparse trees and shrubs, seasonally
inundated, 5°25'27"N, 53°5'19"W, 10 m, 19 Jul. 2009 (fl), *P.G. Delprete 10746* (CAY, K, US); Commune de Iracoubo, Savanne de Trou Poisson, road continuing after Kaja Nostra Hotel, ca. 2 km after the hotel, open field grassy savanna dominated by poaceae and cyperaceae 10-40 cm tall, soil of fine white sand, 5°24'25"N, 53°5'43"W, 20 m, 30 Jan. 2010 (fl), *P.G. Delprete & O. Vrignaud 10845* (CAY, MO, P); Commune de Iracoubo, Savanne de Trou Poisson, road continuing after Kaja Nostra Hotel, ca. 1.5 km after the hotel, open field savanna with sparse grasses and many species of small herbs, soil of fine white sand, very impregnated with water due the abundant rainy season, 5°24'25"N, 53°5'43"W, 20 m, 27 Apr. 2012 (fl), *P.G. Delprete 11880* (CAY, K); Commune de Montsinery, road RD-5, ca. 1 km before the bridge on Crique Aguille, on road between Port Inini and Village Patawa, secondary vegetation at edge of roadside, 4°50'60"N, 52°30'58"W, 10 m, 12 Oct. 2014 (fl, fr), *P.G. Delprete 12619* (CAY); Commune de Kourou, Savane des Pères, ca. 2 km from Station Radar-Télémesure of CSG, towards Kourou, right side, open field herbaceous savanna with sparse trees of *Byrsonima crassifolia* and *Curatella americana*, seasonally inundated, at the moment soil moist but not inundated, 5°7'7"N, 52°39'4"W, 20-30 m, 21 Mar. 2015 (fr), *P.G. Delprete et al. 12625* (CAY); Commune de Sinnamary, Region of Priprit de Yiyi, Maison Nature, going E from Maison Nature, secondary vegetation resembling savanic vegetation (once forested), with sparse tall shrubs, 5°25'8"N, 53°2'48"W, 20 m, 18 Apr. 2015 (fl), *P.G. Delprete 12633* (B, CAY, MO, NY, P), *12634* (CAY, L, NY, P), *12635* (CAY, MO, NY, P), *12636* (CAY, MO, NY, P); Commune de Matoury, Route Nationale 2 (RN2), between PK 10 and PK 11, roadside vegetation with herbs and small shrubs, soil gravelly-lateritic, 4°48'48"N, 52°21'10"W, 10 m, 19 Apr. 2017 (fl), *P.G. Delprete 12882* (B, CAY, F, MO, P), *12885* (BR, CAY, L, MO, NY, P, S, US), *12886* (CAY, G, MO, NY, RB), *12887* (B, BR, CAY, F, G, GB, MG, MO, P, NY, RB, S, US); Commune de Saint George de l'Oyapock, roadside vegetation on Route Nationale 2 (RN2), at ca. 1 km outside Saint-George, 3°54'13"N, 51°48'47"W, 10 m, 28 May 2018 (fl), *P.G. Delprete 12892* (CAY, F, MO, NY, P); Commune de Regina, Savane Roche Virginie (Inselberg), vegetation growing at the edges and on organic substance accumulating on the granitic inselberg, 4°11'55"N, 52°9'5"W, 150 m, 19 Feb. 2019 (fl), *P.G. Delprete & J.C. da Silva 12902* (CAY, F, MO, NY, P, US), *12904* (CAY [2 sheets], F, L, INPA, MG, MO, NY, P, R, RB, UB, US); route Saint Laurent du Maroni-Mana, 21 km de Saint Laurent, savane, 21 Jan. 1974 (fl), *B. Descoings & C. Luu 20478* (P); Mts. de Kaw, Trésor Reserve, near main trail, 4°35'N, 52°16'W, 210-

378 m, 30 Jan. 2003 (fl), *R.C. Ek 1390* (CAY, U); Piste St. Elie, km 15.5, roadside vegetation, 25 Sep. 1982 (fl), *C. Feuillet 190* (CAY); Inselberg Borne N. 4 (border between French Guiana and Brazil), 13 Apr. 1983 (fl), *C. Feuillet 845* (CAY); Montagne de Kaw, Piste Roura-Kaw, PK 8, 4°33'N, 52°9'W, 26 Nov. 1985 (fl), *C. Feuillet 2882* (CAY, P); Piste St. Elie, Savane Corossony, 5°23'N, 53°0'W, 27 Dec. 1986 (fl), *C. Feuillet 4037* (CAY); Bassin de l'Arataye, Montagne des Nouragues, 4°3'N, 52°42'W, lisiere se savane roche (inselberg), 27 Aug. 1987 (fr), *C. Feuillet 4363* (CAY); E of Cayenne, Route de Montabo, PK 3.5, 14 Apr. 1982 (fl), *A. Fournet 147* (CAY, P); Sauts de l'Oiapoque [?], 1900 (fl), *F. Geay 889* (P); SW of Cayenne, near airport, open savanna, 50 m, 21 Feb. 1985 (fl), *A. Gentry & E. Zardini 50266* (CAY, MO); Savane Roche Malmaison [inselberg], 4°58'25", 52°26'20"W, 18 Sep. 2008 (fl, fr), *S. Gonzalez & M. Chaix 1440* (CAY); Savane de Tonate, 5 Feb. 1978 (fl), *J.J. de Granville 2855* (CAY, P, UB); Montagne de Kaw, Camp Caïman, vegetation secondaire sur laterite, 13 Jun. 1979 (fl), *J.J. de Granville & G. Cremers 2947* (CAY, P, U, UB); Massif des Emerillons, zone central, inselberg [ca. 3°15'N, 53°5'W], 300 m, 11 Sep. 1980 (fl), *J.J. de Granville 3817* (CAY [2 sheets], NY, P, U, UB); Region of Oyapock River estuary, SE slope of Montagne des Trois Pitons, inselberg, 250 m, 21 Jan. 1981 (fl), *J.J. de Granville 4295* (CAY); Inselberg à l'extremité NW des Monts de la Trinité [ca. 4°35'N, 53°21'W], 6 Aug. 1981 (fl), *J.J. de Granville 4775* (BR, CAY [2 sheets], P, UB); Montagnes de la Trinité NW inselberg, [ca. 4°40'N, 53°20'W], 350 m, 17 Jan. 1984 (fl) *J.J. de Granville et al. 6109* (CAY, MO, NY, P, U, US); Upper Camopi River Basin, Mont Belvédère [ca. 2°24'N, 53°6'W], inserberg on S slope, 300 m, 21 Nov. 1984 (fl), *J.J. de Granville 6926* (B, BR, CAY [2 sheets], G, MO, NY, P, U, US); Upper Marouini River Basin, Roche Koutou, 2°53'N, 54°4'W, 220 m, 15 Aug. 1987 (fl, fr), *J.J. de Granville et al. 9307* (CAY, US); Upper Marouini River Basin, Camp N. 3, Roche N. 1, Akouba Booka goo Soula, lisiere de savane roche (inselberg margin), 500 m SW, 2°36'N, 54°1'W, 160 m, 26 Aug. 1987 (fl, fr), *J.J. de Granville et al. 9729* (B, CAY, MO, NY, P, U, US); Upper Marouini River Basin, Roche N. 3 Monpé Soula, 2°39'N, 54°5'W, 200 m, 1 Sep. 1987 (fl, fr), *J.J. de Granville et al. 9916* (CAY [2 sheets], MO, NY, P, US); Savanne Renner, 5°20'N, 52°53'W, 3 m, 18 Apr. 1996 (fl, fr), *J.J. de Granville 14377* (CAY, MO, P); Monts d'Awara, central inselberg, 2°49'0"N, 53°22'0"W, 6 Jul. 2002 (fl, fr), *J.J. de Granville et al. 15060* (B, CAY, K, MO, NY, P); Ile de Cayenne, Crique Cabassou, 4°54'N, 54°18'W, 14 Aug. 2002 (fl, fr), *J.J. de Granville & F. Crozier 15619* (CAY, MO); Upper Wanapi River Basin, central dome of inselberg, 2°31'0"N, 53°49'20"W, 14 Apr. 2004 (fl, fr), *J.J. de Granville et al.*
15909 (CAY, MO, P, U); Mont Saint Marcel, zone SF du massif, inselberg, 2°23'N, 53°0'W, 300 m, 18 Jul. 2002 (fl), *J.J. de Granville et al. 15291* (B, CAY [2 sheets], K, MO, NY, P), 15319 (B, CAY [2 sheets], NY, P); Upper Marouini River, Inselberg D.Z., 2°36'30"N, 54°1'45"W, 200 m, 21 Jun. 2004 (fl, young fr), *J.J. de Granville & F. Crozier 16302* (CAY, MO, P, U), 180 m, 1 Jul. 2004 (young fr), *J.J. de Granville & F. Crozier 16568* (CAY, MO, P); Commune de Macouria, route de la Carapa, locality Maya, 4°56'59"N, 52°26'30"W, 9 m, 2 Aug. 2005 (fl, fr), *J.J. de Granville & O. Tostain 17038* (CAY, MO); Site Ariane 4, Crique Karouabo, C.S.G., 5°14'N, 52°47'W, 8 m, 11 May 2007 (fl, fr), *J.J. de Granville & O. Tostain 17343* (CAY); Savane Matiti, zone du projet d'implantation d'éoliennes, 5°5'15"N, 52°35'50"W, 4 m, 12 Aug. 2008 (fl, fr), *J.J. de Granville 17626* (CAY, MO); Trois Saut, Inselberg of Mont St. Marcel [ca. 2°23'N, 53°1'W], 16 Mar. 1976 (fl), *P. Grenand 1228* (CAY [2 sheets]); Savane de Montsinery, 23 Mar. 1982 (fl), *P. Grenand 2066* (CAY [2 sheets], P); Route Cayenne-Rochambeau, 17–21 Jul. 1962 (fl), *F. Hallé 441* (CAY [3 sheets], P, U, US); Trou Poisson, s.d. (fl), *V. Hequet 1150* (CAY); Savane Lambert, s.d. (fl), *V. Hequet 1240* (CAY); Ile de Cayenne, Route de Baduel, garden, 4°56'N, 52°19'W, 10 m, 23 Apr. 1987 (fl), *M. Hoff 5068* (CAY); Savane Matiti, 5°5'N, 52°37'W, 10 m, 13 Dec. 1988 (fl), *M. Hoff & J.J. de Granville 5474* (B, CAY); Savane Mamaribo, R.N. 1, 1 km E, 5°31'N, 53°22'W, 10 m, 26 May 1989 (fl), *Hoff, M. & G. Cremers 5618* (CAY); Kourou, Passoura, Parcelle I, 13 Apr. 1956 (fl), *J. Hoock s.n.* (P [P00729335]); Route de Sinnamary, 19 May 1956 (fl), *J. Hoock s.n.* (P [P00729338]); without locality, Relevé N. 1, 22 Mar. 1962 (fl, fr), *J. Hoock s.n.* (IAN, NY, P [P00729336], U, US); without locality, relevé n. 3, 6 Apr. 1962 (fl), *ORSTOM – IFAT s.n.* (P [7 sheets; P00729444–P00729450], R); Route de l’Est [RN2], 1 km after the bridge on the Conté River, 20 Apr. 1975 (fl), *H. Jacquemin 1625* (CAY); Kaw Mountains, Trésor, Favard Creek, savanna, 4°36'N, 52°18'W, 7 m, 8 Feb. 1996 (fl), *M.J. Jansen-Jacobs et al. 5114* (CAY, P, U); Kourou, Savane des Pères, 5°7'N, 52°39'W, 10 m, 24 Jul. 2000 (fl, fr), *G. Kallin 115* (CAY [3 sheets]); [Cayenne], Route de Montabo, Jardin des Alexandres, 4 Apr. 1986 (fl, young fr), *Kodjoed, J.F. 106* (CAY [3 sheets]); Savane Bordelaise, 25 Jan. 1983 (fl), *J.-N. Labat 212* (P); Kourou, Savane de Combi, pépinière du Cirad, 31 Dec. 2010 (fl), *O. Lachenaud 1014* (BR, CAY); Savane-roche Virginie, route Regina-St. George, 4°11'44"N, 52°9'8"W, 59 m, 7 Apr. 2014 (fl), *O. Lachenaud 1739* (BR, CAY, MO, P); Bassin de l'Approuague-Arataye, Montagne des Nouragues, 4°3'N, 52°42'W, 100 m, lisière se savane roche (inselberg), Feb. 1990 (fl), *D. Larpin 864* (CAY, P); without locality, 1792 (fl, fr), *J.B. Leblond s.n.* (P

P00729382); Savane Sinnamary, exploitation maraichere de Benoit Girard, s.d., *A. Leclerc 36* (CAY), 37 (CAY), 77 (CAY); Montagne des Cheveaux, S of Cayenne (30 km along rd), 4°43'N, 52°25'W, 50 m, 23 Jan. 1978 (fl), *A.J.M. Leeuwenberg 11628* (CAY, NY, WAG); Cayenne, 1857–1859 (fl), *E.J.F. Leguillou s.n.* (P [2 sheets; P00729305, P00729308]); without locality, s.d. (fl, fr), *J.P. Leprieur s.n.* (P [2 sheets, P00729395, P00729396]); without locality, s.d. [1835] (fl), *J.P. Leprieur s.n.* (BM [000614351], F Webb [080082], P [P00729393]); Mt. Sineri, 1838 (fl, fr), *J.P. Leprieur 36* (P); without locality, s.d. (fl), *J.B.L.T. Leschenault de la Tour 392* (P [2 sheets]); Kourou, Route de Tonate, 25 Jan. 1974 (fl), *J.P. Lescure 42* (CAY, P); Ile de Cayenne, Circuit du Lac du Rorota, 10 Feb. 1974 (fl), *J.P. Lescure 81* (CAY, P [2 sheets]); Terrain de l’IRAT, Cabassou, 5 Apr. 1976 (fl), *J.P. Lescure 599* (CAY [2 sheets], P, U); Cayenne, s.d. (fl), *J. Martin 77* (P); without locality, 1842 (fl), *E.M. Mélinon 42* (P); Ile de Cayenne, Cabassou, Station IRAT, 9 Dec. 1982 (fl), *H. Merlier GY-242* (CAY); Monts d’Awara, central inselberg, 2°48'59"N, 53°21'59"W, 24 Mar. 2006 (fl), *J.F. Molino & D. Sabatier 2252* (CAY); [Maroni River Basin], Sauts du Litany, Antekume Pata, 5 Aug. 1980 (fl), *C. Moretti 1191* (CAY); Savannas of Gallion, ca. 20 km SW of Cayenne, 4°46'N, 52°24'W, 26 Apr. 2001 (fl, fr), *S.A. Mori, E.F. Hecklaum, B.W. Keeley, T.A. Lobova & H.P. Pekham 25198* (CAY, NY, P); Nouragues Filed Station, ca. 7 km from Saut Pararé on the Arataï River, near Crique Cascade, 1.2 km SW of Camp Inselberg, 4°5'N, 52°41'W, 120 m, 13 Aug. 2004 (fl), *S.A. Mori, T. Lobova, N. Pitcairn, C. Geiselman & B. Vlásákova 25776* (CAY, NY); Ile de Cayenne, bord de route du Tour de l’Ile, Carrefour de la Route de Stoupan, prairies inundables, 24 Jan. 1968 (fl), *R.A.A. Oldeman & C. Sastre 2* (CAY, IAN, NY, P, U, US); Fleuve Oyapock, Savane Canot Fandé-Fandé, 5 Jul. 1969 (fl, fr), *R.A.A. Oldeman B-2473* (CAY [2 sheets], NY, P, U); fleuve Oyapock, savane roche (inselberg) Canari Zozo, 7 Jul. 1969 (fl), *R.A.A. Oldeman B-2484* (CAY [2 sheets], NY, P [2 sheets], U); Lower Oyapock River, downstream from Saut Fourmi, 3 Jun. 1970 (fl), *R.A.A. Oldeman B-3353* (CAY, NY, P); Ile de Cayenne, route forestiere près du pont des cascades, 16 Apr. 1971 (fl), *R.A.A. Oldeman B-3885* (CAY [2 sheets], P); Ile de Cayenne, base of Mt. Matoury, 30 Jul. 1965 (fl, fr), *R.A.A. Oldeman 1499* (CAY, P, U); Ile de Cayenne, S.M.A., near Camp Tigre, left side of Route de Cabassou, 8 Mar. 1971 (fl, fr), *R.A.A. Oldeman 3217* (CAY [2 sheets], P, U, US); savanne aux environs de Kourou, May–Jun. 1969 (fl, fr), *J. Petitbon 59* (P); without locality, 1826 (fl, fr), *P.A. Poiteau s.n.* (P [3 sheets; P00729387–P00729389]; Ile de Cayenne, Cabassou, Station IRAT, 14 Feb. 1983 (fl), *M.F. Prévost 1252* (CAY, P, UB); Station de
la Piste de St. Elie (ECEREX), PK 15.7, 5°17'N, 53°3'W,
24 Mar. 2000 (fl), M.F. *Prévost 3830* (CAY, NY, P); Piste
de St. Elie, 20 Mar. 1994 (fl), Y. *Purwanto 79* (CAY); Cay-
enne, école d'agriculture de Suzini, entre les monts
Bourda et St. Martin, 29 Feb. 1976 (fl), A. *Raynal & C.*
*Tirel 18302* (NY, P, U); Savane du Gallion, ca. 20 km SSE
de Cayenne, 4 Mar. 1976 (fl), A. *Raynal & C.* *Tirel 18452*
(NY, P, U); in pratis Guyane, s.d. [1781-1785] (fl), L.C.M.
*Richard s.n.* (P P00729306); in pratis, campestribus, s.d.
[1781-1785] (fl), L.C.M. *Richard s.n.* (P P00729383); with-
out locality, s.d. [1781-1785] (fl), L.C.M. *Richard s.n.* (P
P00729384); old airstrip at Le Gallion, ca. 10 m SW of
Rochambeau airport along RN2, 4°47'N, 52°24'W, 8
Mar. 1994 (fl), J.H.E. *Rova et al. 1901* (CAY, GB, NY, S);
Route de St.-Elie, savanna along rd at km 1.5, 5°22'N,
53°0'W, 15 Mar. 1994 (fl), J.H.E. *Rova et al. 1980* (CAY,
GB, S), 1981 (CAY, GB, S); Savane de Organabo, 5 km E
of Organabo on RN 1, 5°32'N, 53°25'W, 20 Mar. 1994
(fl), J.H.E. *Rova et al. 2018* (CAY, GB, S); Cayenne, 1859
(fl), P.A. *Sagot s.n.* (P [2 sheets] P00729301, P00729304);
Cayenne, 1859 (fl), P.A. *Sagot 1246* (P [P00729303]); Bas-
sin de l'Arataye, Montagne des Nouragues, S slope,
4°3'N, 52°42'W, vegetation herbacea de Savane roche

(inselberg), 16 Jan. 1988 (fl, fr), C. *Sartou 168* (CAY), 8
Jan. 1988 (fl, fr), C. *Sartou 191* (CAY); Savane de Kourou,
savane à *Byrsonima crassifolia*, 7 Jul. 1972 (fl, fr), C. *Sas-
tre 1328* (CAY [2 sheets], IAN, NY, P, U); Maroni River,
Saut Lassédédé (between Beligui and Loca), 17 Apr. 1975
(fl), C. *Sastre et al. 3822* (CAY, P); Haut Oyapock, Mont
Saint Marcel, inselberg, [ca. 2°25'N, 53°0'W], 300-450
m, 20 Mar. 1976 (fl), C. *Sastre 4458* (CAY, P, US); Savane
Changement, 6 km from Sinnamary, PK 79, 16 Apr. 1976
(fl), C. *Sastre 4772* (CAY [2 sheets], P); Matiti, savane
seche a *Byrsonima verbascifolia*, sur sable gris, 12 Aug.
1961 (fl), R. *Schnell 11067* (P); Pariacabo, 14 Aug. 1961
(fl), R. *Schnell 11123* (P [2 sheets]); Stoupan, savane de la
ferme Petit, Jul. 1948 (fl), *Service Forestier 3034* (U);
route Larivot, Jan. 1950 (fl), *Service Forestier 3493* (P);
Mont Joly, Aug. 1950 (fl), *Service Forestier 3591* (P); La
Madelaine, Jan. 1951 (fl), *Service Forestier 3680* (P); with-
out locality, s.d. (fl), *Service Forestier 3685* (P); Region
Littorale, Digue Yiyi, 5°25'N, 53°4'W, 2 m, 26 May 1992
(fl), D. *Toriola-Marbot & M. Hoff 92* (CAY); RN 1,
Savane Yiyi, 5°24'N, 53°2'W, 7 m, 25 Jun. 1992 (fl, fr), D.
*Toriola-Marbot 281* (CAY); Station de l'Arataye, Appr-
ouage River Basin, 4°2'N, 52°42'W, 200 m, 26 Mar. 1987

<figure><img src="image_53.png" /><figcaption>**Figure 49.** Distribution of *Sipanea pratensis* var. *pratensis* (squares) and *S. pratensis* var. *dichotoma* (circles).</figcaption></figure>
(fl), *J.F. Villiers 4022* (CAY); Arataye River, 60 km WSW of Regina, 6 km N of Saut Pararé, 14 Feb. 1988 (fl), *J.F. Villiers 4334* (CAY); Inselberg Roche-Dachine, 3°28′N, 53°13′W, 220 m, 2 Apr. 1997 (fl), *J.F. Villiers & C. Sartou 6001* (CAY [3 sheets], NY, P); Inselberg Mont Chauve, 3°49′N, 53°44′W, 265 m, 13 Apr. 1997 (fl), *J.F. Villiers & C. Sartou 6094* (CAY, P); end of road overlooking Kaw, 4°30′N, 52°4′W, 13 Sep. 1987 (fr), *A.L. Weizman & W. Hahn 311* (NY, U); “Guyane”, s.d. (fl), *Collector Unknown 174* (FI).

**BRAZIL. Amapá:** Rod. Perimetral Norte, km 162, beira da estrada, 5 Jun. 1992 (fl), *O. Alencar & B. Santos 007* (HAMAB); entre km 105 e Local Areal, 5 km E de Porto Grande [ca. 0°42′N, 51°24′W], posto ao Norte, 18 Oct. 1979 (fl, fr), *D.F. Austin et al. 7110* (INPA, MG, NY, US); Macapá [ca. 0°22′N, 51°3′W], campina, 30 Oct. 1949 (fl), *G.A. Black 49-8251* (IAN); Oiapoque, esquerda do Campo de Aviação [3°51′N, 51°47′W], 7 Oct. 1949 (fl), *G.A. Black 49-8444* (IAN, P, U); Mun. Amapá, Fazenda Tucunaré, campo com manchas de campina, apresentando ilhas espalhadas, 19 Apr. 1950 (fl), *G.A. Black & I. Lobato 50-9462* (IAN), *50-9469* (IAN, INPA); 6 km N of Porto Grande, Cerrado close to Cerrado-mata transition zone, 0°44′N, 51°20′W, 6 Aug. 1993 (fl, fr), *S. Bridgewater et al. S121* (HAMAB, INPA); Porto Grande, BR-210, área antropizada, 0°40′29″S, 51°30′1″W, Apr. 2014 (fl), *A.L.R. Cardoso 1305* (MG); Mun. Mazagão, próximo a entrada de Vila Santa Clara, 28 May 2001 (fl), *N.V. Costa Neto & O.T. Silveira 511* (HAMAB); Mun. Calçoene, cerrado na margem do Rio Cassiporé, 3°46′N, 51°9′W, 31 Aug. 2002 (fl), *Costa S.V. Neto & M.S. Silva 1019* (HAMAB); Mun. Tartarugalzinho, Rio Tartarugalzinho [ca. 1°30′N, 50°54′W], erva de 30 cm, 12 Aug. 2004 (fl), *S.V. Costa Neto et al. 1561* (HAMAB); Serra do Navio, Rio Amapari, forest of lower slope of Observatorio Ore Body, 70–300 m, 8 Nov. 1954 (fl), *R.S. Cowan 38152* (NY); Serra do Navio, Rio Amapari, Terezinha-Reservatorio trail, wet lowland forest, 75 m, 13 Nov. 1954 (fl), *R.S. Cowan 38317* (IAN, NY, RB, U); Mun. Macapá, km 43, área de baixada, 20 Mar. 1982 (fl, fr), *M. Dantas 948* (IAN); Amapá [2°3′N, 50°47′W], campo, 27 Jun. 1904 (fl, fr), *A. Ducke s.n. (MG 4804)* (MG); Oiapoque, Clevelandia, pasto, 23 Apr. 1960 (fl, fr), *W.A. Egler 1405* (MG, NY); Rio Oiapoque, Roche Mon Pére [ca. 3°17′N, 52°12′W], among loose rocks in soil-filled depression on large granitic outcrop, 3°33′N, 52°5′W, 17 Aug. 1960 (fl, fr), *W.A. Egler 47652* (Irwin collection number) (IAN, MG, NY, US); Porto de Santana [ca. 0°2′S, 51°51′W], 29 Jan. 1961 (fl), *M. Emmerich & A.G. Andrade 694* (R); Porto Grande, Rodovia Perimetral Norte, trecho Porto Grande-Serra do Navio, localidade Mungubas, ca. 59 km da BR-156, na entrada

de Porto Grande, 0°38′55″S, 51°50′51″W, 89 m, 21 Jul. 2010 (fl, fr), *C.N. Fraga 2955* (MG, RB); Mun. Macapá, Rio Macacoraí, 6 Jul. 1951 (fl), *R.L. Frões & G.A. Black 27265* (IAN, U); Rio Araguarí, perto da Cachoeira do Paredão, terra firme, 25 Jul. 1951 (fl), *R.L. Frões & G.A. Black 27691* (IAN); “Guyana Brasileira”, Counany [ca. 2°51′N, 51°7′W], 18 Oct. 1895 (fl), *J. Huber Herb. Amaz. 1103 (RB 15627)* (RB); Rio Oiapoque, Cachoeira Cacheri, immediately W of mouth of Cricú River, wet sandy soil, 3°43′N, 51°58′W, 15 Aug. 1960 (fl), *H.S. Irwin & W.A. Egler 47543* (IAN, MG, NY [2 sheets], US); Rio Araguarí, 20 minutes downriver from Porto Platón, 0°44′N, 51°22′W, 16 Sep. 1961 (fl, fr), *H.S. Irwin et al. 50965* (IAN, NY); Cuxibi, a 150 km de Serra do Navio, entre pastos, 15 Jun. 1966 (fl), *A. Lourteig 1783* (P, R, US); Rio Oiapoque, second-growth woodland along rd between Oiapoque and Clevelandia, frequent, 19 Jul. 1960 (fl), *B. Maguire et al. 47066* (NY); Mun. Calçoene, BR-156, 12 km W of Calçoene, 2°28′N, 51°0′W, 8 Dec. 1984 (fl, fr), *S.A. Mori & R. Cardoso 17266* (HAMAB, MG, US); Mun. Calçoene, BR-156, vicinity of government road camp “Carnot”, 53 km WNW of Calçoene, 2°33′N, 51°16′W, 10 Dec. 1984 (fl), *S.A. Mori et al. 17334* (HAMAB, MG, NY); São Joaquim do Pacuí, margem de estrada, 28 Feb. 1981 (fl), [without initials] *Mozart s.n.* (HAMAB No. 2172); Mazagão, BR-156, as proximidades de Vila de Santa Clara, cerrado, 22 Sep. 2001 (fl, fr), *S.V. Neto & G.A.R. Melo 591* (HAMAB); Pracuúba, BR-156, Flexal, beira da estrada em campo aberto, 1°42′N, 50°53′W, 26 Feb. 2001 (fl), *L.A. Pereira, F. Cesarino, S.M. Alfaia, & J.P. Reis 98* (HAMAB, INPA); Amapá, BR-156, crescendo na beira da estrada, 2°3′N, 50°53′W, 13 Mar. 2001 (fl), *L.A. Pereira & M.A.C. Santos 166* (HAMAB, INPA); Mazagão, Estrada do Jari, BR-210, solo arenoso de campo aberto, 0°20′S, 51°52′W, 23 Mar. 2001 (fl), *L.A. Pereira & J.P. Reis 295* (HAMAB, IAN); Mun. Pracuúba, área de campo cerrado, subarbusto 20 cm altura, 16 Jun. 1983 (fl), *L.C. Pinto, Jonas & H.P. Belo 21* (HAMAB); Rio Oiapoque, near Mt. Carupina, 3°33′N, 51°37′W, 10–30 m, 16 Oct. 1960 (fl, fr), *J.M. Pires & L.Y.T. Westra 48878* (Irwin collection number) (IAN, NY, US); Rio Araguari, Porto Platon, 18 Sep. 1961 (fl), *J.M. Pires 51049* (Irwin collection number) (IAN, MG, NY); Mun. Macapá, km 38 para Matapi, 2 Feb. 1955 (fl, fr), *J.M. Pires & N.T. Silva 4795* (IAN); road to Amapá, km 48, 0°17′N, 51°5′W, 7 Jul. 1962 (fl), *J.M. Pires & P.B. Cavalcante 51977* (Irwin collection number) (IAN, MG, NY, US); Colonia do Torrão, forest at border of igarapé, 2°25′N, 51°15′W, 28 Aug. 1962 (fl), *J.M. Pires & P.B. Cavalcante 52622* (Irwin collection number) (IAN, MG, NY); km 104 da BR-156-Macapá, campo cerrado, 5 May 1979 (fr), *B.V. Rabelo s.n.* (Museu Ângelo Costa Lima N. 1412) (MG);