Carapichea guianensis Aubl.

Subshrubs, shrubs, or small trees. Raphides present. Stipules interpetiolar, free or shortly connate at base, entire to bifid at apex, rarely multifid (C. ipecacuanha), lacking dorsal appendages, persistent on distal nodes, marcescent, and eventually falling off through fragmentation, leaving a well-developed scar or a persistent hardened basal portion. Leaves opposite, rarely ternate (C. tillettii), short- to long-petiolate; blades ovate, elliptic, obovate, oblong or narrowly elliptic, papyraceous to coriaceous; secondary veins strongly to weakly prominent; tertiary veins reticulate or subparallel; domatia absent, or present as a continuous line of pubescence along midrib. Inflorescence terminal, rarely pseudoaxillary, few- to many-flowered, variously capitate, subcapitate, spiciform, or thyrsoid and branched to 1 or 2 orders; bracts reduced to very large, free to variously connate, sometimes forming an involucre. Flowers bisexual, usually heterodistylous, protandrous, (4)5-merous. Hypanthium ovoid to obovoid. Calyx tube extremely reduced or cup-shaped, truncate, undulate or lobed, persistent, lobes (when present) small, broadly to narrowly triangular. Corolla tubular, funnelform to hypocrateriform, actinomorphic, white, yellow, orange, or salmon-pink, tube glabrous or pubescent inside; lobes valvate, oblong-ovate, margin entire, acute at apex, thickened or sometimes with horn-like extensions (C. araguariensis). Stamens included, partially exserted or well exserted beyond corolla mouth; filaments inserted at the basal (C. urniformis), median, or distal portion of the corolla tube, short, equal, glabrous; anthers narrowly elliptic or narrowly oblong or linear, round at base, round or apiculate at apex, dorsifixed near middle. Pollen aperturate. Ovary 2-locular, placenta reduced. Ovules 1 per locule, basally inserted, erect. Disk usually bilobed to the base, sometimes undivided and cylindrical or laterally 5-sulcate (C. adinantha). Style included or partially exserted, glabrous; branches 2, oblong or linear. Fruits drupaceous, fleshy, variable in colour (yellow, orange-red, red, purple, maroon, blue, white or cream-white, or black at maturity) with 2 fibrous to woody pyrenes. Pyrenes vertical, plano-convex, ovate to elliptic in outline, dorsal side multi-costate, ventral side longitudinally sulcate or rarely flat.

The main diagnostic characters of Carapichea are the marcescent stipules lacking dorsal appendages, lack of ethanol-soluble pigment in the seed testa, and aperturate pollen of a generalized form, corresponding to Types XIV and XVI of Johansson (1992). The genus is rather variable in inflorescence and pyrene characters, as described by Taylor and Gereau (2013). Most species of Carapichea have a bipartite disk. This character was not emphasized by previous authors and is often not even mentioned in their descriptions, though it is well depicted on illustrations of C. tillettii (Steyermark 1972: 496, figure 70), C. urniformis (Steyermark 1972: 557, figure 74), and C. vasivensis and C. pacimonica (Steyermark 1974, figures 234, 240). Among species investigated for this character, the only exceptions are C. cardenasiana, C. ipecacuanha, and C. panurensis, which have an entire disk, and C. adinantha, which has an undivided, laterally 5-sulcate disk. We were not able to verify the shape of the disk in C. fimbriflora, C. lucida, C. maturacensis, and C. verrucosa. Among related genera, bipartite disks also occur in some species of Notopleura (see above) and in two recently described species of Rudgea Salisb.: R. glomerulata Zappi & O.Lachenaud and R. itoupensis O.Lachenaud (Lachenaud et al. 2022).

Position within the tribe and delimitation of Carapichea. Carapichea has been included in the Palicourea complex of the tribe Psychotrieae s.l. by Andersson (2002) and Lachenaud (2019). This complex has been treated as the tribe Palicoureeae by Robbrecht and Manen (2006), Razafimandimbison et al. (2014), and Taylor and Bruniera (2018), and includes Carapichea, Chassalia Comm. ex Poir., Eumachia DC. [= Chazaliella E.M.A.Petit & Verdc., Margaritopsis C.H.Wright], Geophila D.Don, Hymenocoleus Robbr., Notopleura, Palicourea Aubl. s.l., Puffia Razafim. & B.Bremer, and Rudgea. Phylogenetic studies strongly support the monophyly of Carapichea, albeit with limited sampling (Andersson 2002; Razafimandimbison et al. 2014; Bruniera 2015). The genus appears either as sister to Eumachia (Andersson 2002; Bruniera 2015; as Margaritopsis) or as sister to a clade including Eumachia, Chassalia, Geophila, Puffia, and Hymenocoleus (Razafimandimbison et al. 2014) with low support in all cases.

According to the present circumscription, Carapichea includes at least 22 species that are found mostly in the Amazon basin and the Guianas, with two species (C. affinis and C. ipecacuanha) extending to Central America, one (C. ipecacuanha) extending to the Cerrado Biome of Central Brazil, and a single species (C. lucida) endemic to the Atlantic Forest of Brazil. The species occur in lowland or lower montane forest habitats (up to 1570 m but mostly below 1000 m), either on drained or seasonally flooded soils. Eight species occur in the Guianas; a ninth one, C. araguariensis, is to be expected there and has been included in the treatment below.

No infrageneric taxa are formally recognised in Carapichea, but the species may be arranged in six groups (Table 1), separable by characters in the key below, and largely based on those proposed by Taylor and Gereau (2013). We have usually kept the group names used by these authors, although their Pacimonica group is better called the Ligularis group now that C. pacimonica is a synonym of C. ligularis. Their Altsonii group is not maintained here, since, as discussed above, three of its species are transferred to Notopleura, while the last one, C. urniformis, is included in the Carapichea group of which it has all the key characters. Also transferred to the Carapichea group are C. tillettii (see Notes under this species) and C. fimbriflora. The latter species, from Brazil, is very poorly known, but its leaf venation matches the Carapichea group, rather than the Ligularis group in which it was placed by Taylor and Gereau (2013).

PERU – Loreto • near Iquitos, Mishayacu; 100 m; Feb.–Mar. 1930; fl.; Klug 988; holotype: F [No. 612612]; isotypes: NY [00133128], US [0013820].

Figure 2. 

Carapichea adinantha. A–B. Detail of inflorescence with buds and one open flower. C. Fruiting plant. D. Infructescence with mature fruits. E. Infructescence with mature fruits, some eaten by birds. A–B from Gonzalez et al. 3281; C–E from Gonzalez 1368. Photographs by Sophie Gonzalez.

Shrub or small tree up to 4 m tall, sparsely branched; terminal branchlets terete or slightly laterally compressed, 2.5–3.0 mm in diam., glabrous, soon covered with a greyish-beige thin bark. Stipules free, interpetiolar, 2–5 mm long, bifid at apex, each lobe deltoid to lanceolate, 1–2 mm wide at base, acute to acuminate, glabrous. Leaves with petioles 1.5–3.0 cm long, glabrous; blades elliptic, oblong-elliptic oblong-lanceolate, 19–34 × 5.5–15 cm, acute at base, acuminate at apex, chartaceous to papyraceous when dry, entirely glabrous, drying olive-green; midrib and secondary veins slightly prominent on the upper side; secondary veins 9–12 on each side of the midrib, ascending and forming loops away from the margin; intersecondary veins 2–3 between each couple of secondary veins, terminating far from the margin; tertiary veins reticulate, sparse. Inflorescences terminal, erect, spiciform, 7–20 cm long, with (6–)7–9 multiflorous glomerules, rachis minutely puberulous; peduncle 1.0–8.7 cm long; bracts subtending each glomerule shallowly ovate to broadly deltoid, 1–2 mm long. Flowers 5-merous, sessile (heterostylous?). Hypanthium glabrous or sparsely puberulous. Disk 5-sulcate laterally. Calyx cupuliform, 1.0–2.2 mm long, minutely puberulous, margin truncate, undulate or shallowly denticulate, ciliate. Corolla hypocrateriform, white; tube cylindrical, 4.5–7.0 mm long, 1.5–1.7 mm wide, granular-puberulous outside, glabrous inside; lobes imbricate, 3.5–4.5 × 1.4–1.7 mm, reflexed at anthesis, internally appendiculate, granular-puberulous outside, glabrous inside; appendages linear, 1.5–2.0 mm long. Stamens inserted at 1.0–1.5 mm from base of corolla tube, included; filaments 1.1–1.3 mm long; anthers linear, 2.3–2.5 × 0.2–0.3 mm. Style as long as corolla tube (branch tips barely exserted), ca 4.5–7.0 mm long, style branches linear, 1.3–1.5 mm long. Fruits drupaceous, fleshy, red to pink when immature (one collection reported as “white”), turning dark blue to black at maturity, interior spongy-white, ellipsoid to subglobose, 13–14 × 9–10 mm when fresh, 7–16 × 5–12 mm when dry, glabrous, sessile. Pyrenes 2, thin-walled, plano-convex, elliptic to narrowly elliptic in outline, 9–10 × 7–8 mm, dorsal side smooth (i.e. without ridges), ventral side flat (without longitudinal furrow), with a pore with a raphal plug, sometimes with short basal marginal slits. Seeds with a deep T-shaped ventral groove in cross-section.

This species occurs sporadically in Amazonian Peru, Colombia, Brazil (Acre, Amapá, Amazonas, Pará, Rondônia), and French Guiana.

In undercanopy of terra firme primary and secondary forest, often at river margins, on clay soil, at 100–200(–300) m in elevation.

FRENCH GUIANA • Approuague River Basin, Rivière Kourouaï; 4°15’N, 52°01’W; 120 m; 9 Jul. 2008; fr.; Gonzalez 1368; BR, CAY, L, MO, NY, P, US • Monts Tumuc-Humac, Sommet en Cloche; 2°13’40”N, 54°28’10”W; 12 Mar. 2015; fl.; Gonzalez 3281; BR, CAY.

BRAZIL – Acre • Mun. Cruzeiro do Sul, Rio Branco, ca 25 km from Rio Juruá, secondary forest with canopy 30–35 m tall; 7°45’S, 72°17’W; 240 m; 22 Nov. 2001; fl.; Delprete et al. 8045; NY, UFACPZ, additional duplicates at UFACPZ to be distributed • Mun. Cruzeiro do Sul, road Cruzeiro do Sul–Rio Branco, road to Canela Fina, ca 7 km from 3-way junction at 5 km N of Cruzeiro do Sul; 7°32’44”S, 72°44’31”W; 220 m; 23 Nov. 2001; fl.; Delprete et al. 8078; NY, UFACPZ, additional duplicates at UFACPZ to be distributed. – Amapá • Parque Nacional Montanhas do Tumucumaque, cabeceiras do Rio Amaparí, margem esquerdo do Rio Anacuí, trilha 1, floresta de terra firme; 6 Mar. 2006; fl.; Hamada et al. 139; INPA, MO n.v. – Amazonas • Distrito Agropecuário, Fazenda Porto Alegre, Reserva 3402 (Cabo Frio) of the WWF/INPA MCS project; 2°25’25”S, 59°54’38”W; 50–125 m; 1 Apr. 1989; fr.; Aquino et al. 3; INPA • Basin of Rio Juruá, near mouth of Rio Embira (tributary of Rio Tarauacá); 7°30’S, 70°15’W; 6 Jun. 1933; fr.; Krukoff 4682; K, M, MO • Basin of Rio Juruá, near mouth of Rio Embira; 17 Jun. 1933; fr.; Krukoff 4900; M, MO, NY, US. – Pará • Igarapé das Pedras, Cataractam Furnas, Rio Tapajóz; 30 Dec. 1917; fl.; Ducke s.n. (MG 16859, RB 23125); B destroyed (photo at F), RB. – Rondônia • ca 4 km ENE along rd to São Sebastião off BR-364, terra firme forest; 24 May 1984; fr.; Frame 160; INPA, MO n.v., NY • Estrada Porto Velho–Cuiabá, BR-364, km 171, mata de terra firme; 6 Feb. 1983; fr.; Freitas et al. 13; INPA • Itapuã do Oeste, Floresta Nacional do Jamari, parcela convenio NYBG-RON; 9°15’6”S, 62°54’38”W; 23 Apr. 2015; fr.; Medeiros et al. 1692; MO n.v., NY, RON.

COLOMBIA – Caquetá • Araracuara, trocha Yari; 0°25’S, 72°20’W; 200 m; 23 Jan. 1989; fl.; Gentry & Sánchez 65001; COL, MO n.v.

PERU – Loreto • Maynas Prov., Dtto. Amazonas, Explornapo Camp, cerca de Sucusari, a lo largo del Río Napo; 3°20’S, 72°55’W; 100–140 m; 15 Feb. 1991; fl.; Pipoly et al. 12979; MO • Chacra Canamá, Requena; 7 Dec. 1962; fl.; Schunke Vigo 6243; K • Prov. Requena, Jenaro Herrera; 4°50’S, 73°45’W; 170 m; 5 Jul. 1981; fr.; Vásquez et al. 2192; MO • carretera Nauta–Iquitos km 5; 4°29’S, 73°35’W; 28 Mar. 1987; fr.; Vásquez & Arevalo 8984; P • Iquitos, Asociación Agraria Paujil; 4°10’S, 73°20’W; 2 Jul. 1988; fr.; Vásquez & Jaramillo 10837; P • Allpahuayo, Estación Experimental del Instituto de Investigaciones de la Amazonia Peruana; 2 Jun. 1990; fr.; Vásquez & Jaramillo 14029; P.

This species has recently been collected in French Guiana and the adjacent Brazilian state of Amapá, which represents a major eastern range extension. Although this species is known from numerous specimens, most of them are fruiting and we were able to dissect only a few flowers. In these flowers, the stamens are inserted at 1.0–1.5 mm from base of corolla tube and are included, while the style is as long as the corolla tube, with the branch tips barely exserted. Though we have not seen a short-styled form, we suspect the species may be heterostylous, as many of its congeners.

BRAZIL – Amapá • Araguari River, Camp 12; 1°11’N, 52°08’W; 30 Sep. 1961; fl.; Murça Pires et al. 51371; holotype: NY [00132598].

Shrub, 0.5–1.0 m tall, or treelet 2–5 m tall, glabrous; terminal branchlets terete, 2–4 mm in diam., soon covered with a greyish bark. Stipules shortly sheathing, truncate to very broadly ovate, 1–4 × 4.0–5.5 mm, entire and obtuse at apex, glabrous, soon corky and fragmenting. Leaves with petioles 0.7–2.5 cm long, glabrous; blades elliptic, to oblong-oblanceolate, 11–25 × 3.5–7.5 cm, acute-decurrent at base, acute and acuminate at apex, acumen narrowly triangular, 0.5–1.5 cm long, subcoriaceous to coriaceous when fresh, drying papyraceous to subcoriaceous, grey-green to olive brown, glabrous throughout; secondary veins 13–25 on each side of midrib, weakly ascending and hardly more prominent than the intersecondary veins, arching at 0.5–1.0 mm from the margin; intersecondary veins (1–)2–3 between each couple of secondary veins, terminating far from the margin; tertiary veins obsolete; domatia absent. Inflorescence compact-paniculate at early stage, expanding and becoming obviously paniculate at later stage, many-flowered, pedunculate; peduncles 4.5–9.5 cm long, glabrous or with distal portion puberulous, drying olive green to pale brown; secondary branches verticillate, 2–4 per node, 0.7–3.0 cm long, glabrous to puberulous, terminating into cymules; outer branches of cymules with bracts; bracts 2–5 in each cymule, subequal to unequal, narrowly elliptic to linear, longer ones 7–17 × 1–2 mm, shorter ones 4.0–4.5 × 0.7–1.2 mm, persistent or tardily caducous, drying olive green to brown, glabrous. Flowers 5-merous, (heterostylous?), sessile or with pedicel < 0.5 mm long during anthesis, elongating to 1–5 mm long at fruiting stage. Hypanthium narrowly obovoid, 0.7–1.0 mm long, glabrous. Disk bilobed to the base, 0.5 mm long, glabrous. Calyx cupular, 0.7–1.3 mm long, truncate or minutely denticulate, glabrous. Corolla infundibuliform, 14.5–16 mm long, glabrous, white (“orange” according to Pires et al. 51371), tube narrowly obconical, 11–12 mm long, 1.5 mm wide at base, 3.5–4.0 mm wide at mouth, glabrous outside and inside; lobes lanceolate, 3.0–3.5 × 1.2–1.3 mm, acute at apex, glabrous, bearing dorsal linear cornicula 0.3–0.7 mm long. Stamens inserted just below the corolla mouth, filaments 0.5 mm long, anthers subsessile, half-exserted, narrowly oblong, 2.3–3.0 × 0.3–0.6 mm. Style glabrous, barely exserted, 14.5–16.5 mm long. Fruits elliptic to ovoid, 6.5–10 × 5.5–8.5 mm, slightly costate when dry, green when young, orange or yellow at maturity. Pyrenes plano-convex, ellipsoid in outline, 6-9 × 4.5–5.5 mm, dorsal side with 3 prominent longitudinal ridges, ventral side with a shallow longitudinal groove. Seeds with a deep T-shaped ventral furrow.

Only known from northern Brazil (Amapá, Pará, and Amazonas states); to be expected in French Guiana.

In understory of moist, non-flooded forest, at 50–125 m elevation.

The flowering type specimen from Amapá state was collected in September; in Amazonas state, flowering specimens were collected in February and March, and fruiting specimens in February and August.

BRAZIL – Amazonas • Reserva Florestal Ducke, Estrada Manaus–Itacoatiara, km 26, floresta de campinarana; 2°53’S, 59°58’W; 13 Feb. 1996; fl., fr.; Campos et al. 481; INPA, MO, NY • Mun. Manaus, Distrito Agropecuário da SUFRAMA, Rod. BR-174, km 72, depois 6 km W da BR, Fazenda Dimona, mata de terra firme sobre latosolo amarelo; 2°19’S, 60°5’W; 50–125 m; 25 Mar. 1992; fl.; Dick 62; INPA • estrada Manaus–Caracaraí, km 125, igarapé da Lage, terra firme; 13 Feb. 1974; fr.; Loureiro et al. s.n. (INPA 47925); INPA • ibid., 14 Feb. 1974; fl.; Loureiro et al. s.n. (INPA 47947); INPA • Distrito Agropecuário, Reserve 1501 (km 41) of the Smithsonian/INPA Biological Dynamics of Forest Fragments Project; 2°24–2°25’S, 59°43’–59°45’W; 15 Jul. 1990; fr.; Mori & Costa Lima Assunção 21383; K • Reserva Florestal Ducke, Manaus–Itacoatiara km 26; 2°53’S, 9°58’W; 16 Jun. 1994; fr.; Ribeiro & Assunção 1325; K • Manaus, km 68 da Estrada Manaus–Itacoatiara, sub-bosque de terra firme; 19 Mar. 1963; fl.; Rodrigues 4985; INPA • Reserva Florestal Ducke, Estrada Manaus–Itacoatiara, km 26, floresta de baixo, solo arenoso; 2°53’S, 59°58’W; 3 Feb. 1995; fl.; Vicentini et al. 840; INPA, K, MG, MO, NY, U • ibid., 31 May 1995; fr.; Vicentini et al. 984; K. – Pará • Mun. Anajas, opposite to town of Anajas, on Rio Anajas; 31 Oct. 1984; fl.; Sobel et al. 4943; MG, MO n.v.

No records of this species are known from the Guianas to date, but due to its occurrence in the Brazilian state of Amapá, it may well be found in adjacent French Guiana, and is therefore included in this treatment. Carapichea araguariensis resembles C. ligularis but has laxer inflorescences with the basal bracts inserted at the end of the secondary branches (vs at the end of the peduncle). The bracts are also usually smaller and more distinctly unequal than in C. ligularis, and the corolla tube is longer. This species also resembles C. necopinata from Brazil (Amazonas state), but the latter has an inflorescence with only three glomerules and outer bracts 20–25 mm long, while in C. araguariensis the inflorescence has at least five glomerules and outer bracts 7–17 mm long. See also the notes under C. sp. A below. Due to the rarity of flowering specimens, it is not known whether the flowers are heterostylous. In the few flowers that we analyzed, the stamens are included and the style is barely exserted, which is consistent with a long-styled form.

FRENCH GUIANA • pente NE des Monts Galbao, à 10 km au SW de Saül; 500–600 m; 11 Mar. 1975; fl.; Granville 2383; holotype: VEN n.v.; isotype: CAY [CAY024926, CAY024927].

Shrub 0.5–3.0 m tall, sparsely branched; terminal branchlets terete, 2.0–2.5 mm in diam., glabrous, soon covered with a pale grey corky bark. Stipules triangular, 2–5 × 1.2–4.5 mm, free, entire or shortly bilobed for < 0.5(–1) mm, glabrous outside, villose at the base inside, becoming corky and soon damaged. Leaves with petioles 0.6–2.5 cm, glabrous; blades elliptic or slightly oblanceolate, 11–22.5 × 3.5–7.2 cm, long-attenuate and decurrent on the petiole at base, acuminate at apex, subcoriaceous when fresh, papyraceous when dry, entirely glabrous, drying blackish or dark grey; midrib and secondary veins prominent on the upper side; secondary veins 7–10 on each side of the midrib, strongly ascending, arching near the margin (almost reaching it); with 2–3 subsecondary veins between each pair of secondary veins; tertiary veins reticulate, barely visible when fresh, prominent when dry. Inflorescences terminal, involucrate heads, 14–20-flowered; peduncle 1–3 cm long, erect, glabrous; involucre whitish outside and dark green inside, very shortly connate basally, with two decussate pairs of bracts, both pairs with a basal stalk 8–11 × 7–10 mm; the outer pair 27–44 × 9–25 mm, with the medio-distal portion broadly ovate to ovate; the inner pair slightly smaller than the outer bracts, 24–36 × 5–15 mm, with the medio-distal portion broadly to narrowly ovate to lanceolate, acute to obtuse at apex; both pairs entirely glabrous, persistent at fruiting stage. Interfloral bracts much smaller than the involucral ones, narrowly elliptic, obovate to narrowly spathulate, obtuse at apex, gradually decreasing in size towards the center of the inflorescence, the largest ones 10–12 × 1.5–5 mm, glabrous, caducous or sometimes persistent at fruiting stage. Flowers 5-merous, heterostylous, sessile. Hypanthium obovoid, 1.0–1.3 × 0.7–0.8 mm, glabrous. Disk bilobed to the base, cylindrical, 0.7–1.0 mm long. Calyx shortly cupuliform, 0.3–0.4 mm long, truncate or shallowly denticulate, glabrous. Corolla hypocrateriform, white; tube narrowly cylindrical, slightly wider at mouth, (15–)16–19 × 2.0–2.5 mm, glabrous outside, shortly pubescent at stamens insertion inside; lobes oblong-lanceolate, 2.0–3.5 × 1.0–1.2 mm, reflexed, acute at apex, glabrous throughout. Short-styled flowers: stamens inserted at distal 1/4^th from the base of the tube; filaments ca 3 mm long; anthers with only the tips exserted, ca 3 × 0.3 mm; style included, ca 13 mm long; style branches 2, linear, ca 2 mm long. Long-styled flowers: stamens inserted just above the middle of the tube; anthers subsessile, fully included, 2.0–2.4 × 0.2 mm; style just reaching the corolla mouth or exserted beyond the mouth , 16–17 mm long; style branches linear, 1.0–1.2 mm long. Fruits ellipsoid, 8–10 × 5.5–6.0 mm when dry, orange to red, turning black at full maturity, glabrous, sessile. Pyrenes planoconvex, narrowly elliptic in outline, 8–9 × 3.5–4.0 mm, dorsal side with 3–4 faint ridges, ventral side with a deep narrow excavation for the whole length, ± C-shaped in cross-section, opening by 4 dorso-basal slits running along the ridges. Seeds entire, C-shaped in cross-section.

Figure 3. 

Carapichea galbaoensis. A. Stem with leaves, stipules, and inflorescences. B. Nodes with stipules. C. Flower bud and two floral bracts. D. Corolla at anthesis with partially exserted anthers, lateral view. E. Stamens, adaxial and lateral view. F. Infructescence surrounded by bracts, top view. G. Fruit (left) and seed (right). A–E from Mori & Pipoly 15511; F–G from Marshall & Rombold 102. Drawn by Bobbi Angell. Reproduced from Mori et al. (2002: figure 279) with permission.

Figure 4. 

Carapichea galbaoensis. A. Inflorescence with open flowers, top view. B. Inflorescence with open flowers, side view. C. Flowering branch. D. Habit. A from plant not collected, Monts La Fumée, French Guiana; B–C from plant not collected, Mont Galbao, French Guiana; D from Lachenaud 1689. Photographs by Sébastien Sant (A–C) and Olivier Lachenaud (D).

This species is found mostly in French Guiana, where it is locally frequent in the interior (occurring in most ranges of the Inini-Camopi chain, and in the Approuague basin north of the Nouragues Mountains) with a single record from the adjacent Brazilian state of Amapá.

This species is locally frequent in the understory of terra firme rainforest, at 100–800 m elevation.

This species flowers during the small dry season (March–early April) and produces fruits from mid-April to September.

FRENCH GUIANA • Saül, layon blanc vers Limonade; 3°37’N, 53°12’E; 3 Aug. 1987; Allorge 311; P • Mont Bakra; 3°18’N, 52°57’W; 475 m; 16 Apr. 1993; fr.; Cremers 13141; CAY [2 sheets], U • Mont Chauve; 3°49’N, 52°44’W; 150 m; 16 Apr. 1997; Cremers & Crozier 15010; CAY • ibid.; 120 m; 21 Apr. 1997; fl.; Cremers & Crozier 15134; CAY • Saül, sur le layon vers La Fumée, 500 m avant le plateau La Douane; 3 Feb. 1981; imm. fr.; Fournet 62; CAY, P • Monts Galbao, à 10 km WNW de Saül; 10 May 1973; Granville 1584; CAY, COL, U • Saül, 1 km environ au S des Monts La Fumée; 6 Mar. 1975; fl.; Granville 2354; CAY • Saül, layon ORSTOM du plateau La Douane aux Monts La Fumée; 16 Jan. 1976; fl.; Granville 2664; CAY [2 sheets], P • Saül, antenne est La Fumée/Nouvelle France, pk 1.2; 5 Mar. 1977; fl.; Granville 2806; CAY • Sommet Tabulaire, versant sud; 500 m; 24 Aug. 1980; fr.; Granville 3596; CAY [2 sheets], P • Massif des Emerillons, zone sud; 250 m; 8 Sep. 1980; fr.; Granville 3773; CAY • tracé ORSTOM La Fumée, à 1,2 km environ au N du Plateau La Douane; 3 Apr. 1982; fl.; Granville 5050; CAY • env. 13 km SSW Saül, 2–3 km W of Crique Limonade, Montagne Périnette; 31 Mar. 1983; fl.; Granville 5435; CAY [2 sheets], P • Montagne Bellevue de l’Inini, zone centrale; 750–800 m; 23 Aug. 1985; fl., fr.; Granville 7761; CAY, INPA, MG, MO n.v., P, U • Station des Nouragues; 4°03’N, 52°42’W; 100 m; 6 Aug. 1989; fr.; Granville et al. 11071; CAY • Pic Coudreau, Monts Bakra; 3°18’N, 52°57’W; 710 m; 18 Apr. 1993; fl.; Granville & Cremers 11788; CAY • Monts Bakra, 1,5 km à l’ouest du Pic Coudreau; 3°18’N, 52°57’W; 520 m; 16 Jun. 2002; fr.; Granville et al. 14804; CAY • Saül, Monts La Fumée; 3°40’N, 53°10’W; 200–300 m; 26 Jul. 1987; fr.; Hahn 3634; CAY, U • Piste Sophie, Saül; 11 Sep. 1962; F. Hallé 818; P, U • Station des Nouragues; 4°03’N, 52°42’W; Jul. 1996; fr.; Hequet 222; CAY • Saül, Les Eaux Claires; 3°39’N, 53°12’W; 200–400 m; 5 Sep. 2000; fr.; Junikka & Nieminen 3031; CAY • Saül, Monts La Fumée; 17 Mar. 2014; Lachenaud 1689; BR, CAY • Montagne des Nouragues; 4°03’N, 52°42’W; Jun. 1989; fr.; Larpin 539; CAY [2 sheets] • Commune de Saül; 13 Jul. 1977; fr.; Moretti 752; CAY [2 sheets] • Saül, Monts La Fumée; 3°37’N 53°12’W; 200–400 m; 30 Aug. 1982; fr.; Mori & Boom 14803; CAY, NY • Saül, La Fumée Oeste; 3°37’N, 53°12’W; 200–400 m; 5 Apr. 1983; fl.; Mori & Pipoly 15511; CAY, NY • Saül, La Fumée Mountain; 3°37’N, 53°12’W; 200–350 m; 22 May 1986; fr.; Mori & Pennington 18097; CAY, P, U • Station des Nouragues; 4°03’N, 52°42’W; 20 Jul. 1989; fr.; Sabatier & Prévost 2547; CAY, U • Station des Nouragues, chemin ouest; 9 Mar. 1996; fl.; Solano K 282; CAY.

BRAZIL – Amapá • Near Cachoeira Macacoara; 0°53’N, 53°21’W; 27 Aug. 1961; fr.; Egler & Irwin 46695 [Irwin’s collection number]; U, US.

Carapichea galbaoensis is closely related to C. guianensis and has been considered a synonym of the latter (Delprete 2001, 2003; Boom and Delprete 2002; Taylor and Gereau 2013) but is here reinstated as a distinct species. The two taxa are almost identical in vegetative characters (although C. galbaoensis has usually shorter stipules), fruits, pyrenes, and inflorescence structure, but the shape and dimensions of the involucral bracts differ, as well as the position of the stamens and style, and especially the length of the corolla tube, which is much longer in C. galbaoensis. Differences between them are summarised in Table 2 and illustrated in Figs 3, 5.

Figure 5. 

Carapichea guianensis. A. Habit. B. Detail of lower leaf surface. C. Inflorescence with long-styled flowers. D. Inflorescence with short-styled flowers. E. Infructescence. A from plant not collected, Mont Itoupé, French Guiana; B–C from Lachenaud 1725; D from plant not collected, Camopi, French Guiana; E from plant not collected, Bellevue de Papaïchton, French Guiana. Photographs by Sébastien Sant (A, D–E) and Olivier Lachenaud (B–C).

FRENCH GUIANA • “ad ripam amnis Galibiensis” [margin of the Galibi Creek]; s.d. [Apr.–May 1763]; Aublet s.n.; lectotype: P-JJR [8:266], designated by Lanjouw and Uittien (1940: 149); isolectotype: BM [BM001009105]; see Delprete (2015).

Shrub 0.4–1.5 m tall, sparsely branched; terminal branchlets terete, 1–2 mm in diam., glabrous, soon covered with a buff to pale grey corky bark. Stipules 5–9 × 1.5–3.0 mm, free, triangular to lanceolate, entire or shortly bifid for up to 3 mm, glabrous outside, villose at the base inside, becoming corky and soon damaged. Leaves with petioles 0.8–3.0 cm long, glabrous; blade elliptic, 10–18 × 3.3–7.5 cm, attenuate and decurrent on the petiole at base, narrowly acuminate at apex, rather thick when fresh (becoming papyraceous when dry), entirely glabrous, drying dark grey-green to blackish; midrib and secondary veins prominent on the upper side; secondary veins 6–9 on each side of the midrib, strongly ascending, curving towards the margin and almost reaching it; with 2–3 intersecondary veins between each pair of secondary veins; tertiary veins reticulate, rather inconspicuous in the fresh state, prominent when dry. Inflorescences in terminal involucrate heads, few-flowered; peduncle 1–3 cm long, erect to patent, glabrous; involucre green on both sides, sometimes with a reddish-brown basal part, consisting of two very unequal decussate pairs of bracts shallowly connate at base, the outer pair 19–42 × 3–10 mm, the inner pair 5–20 × 2–6 mm, both pairs with a basal stalk 4–6 mm long (that of the inner pair much narrower) and the medio-distal portion narrowly ovate to lanceolate, acute to round at apex, glabrous, persistent at fruiting stage; interfloral bracts much smaller than the involucral ones, elliptic, obovate to spathulate, 4–6 × 0.8–3.0 mm, obtuse to truncate at apex, glabrous, persistent in the fruiting stage. Flowers 5-merous, heterostylous, sessile. Hypanthium obovoid, 1.0–1.2 × 0.7 mm, glabrous. Disk bilobed to the base, 0.5–0.6 mm long. Calyx shortly cupuliform, truncate or undulate, 0.2 mm long, glabrous. Corolla hypocrateriform, white; tube infundibuliform, 5–8 mm long, 0.8–1.0 mm wide at base, 2.6–3.7 mm at mouth, glabrous outside, shortly pubescent inside around the insertion of the stamens; lobes ovate to deltoid, 2–3 × 1.5–2 mm, reflexed, glabrous throughout. Short-styled flowers: stamens inserted at distal 1/3^rd of the tube, exserted beyond corolla mouth; filaments ca 3 mm long; anthers 1.3–2.3 × 0.3–0.5 mm; style included, 5–6 mm long; style branches linear, 1.5–1.7 mm long. Long-styled flowers: stamens inserted at distal 1/3^rd of the tube, included; anthers subsessile, oblong-elliptic 1.5–1.8 × 0.3–0.4 mm; style exserted well beyond the mouth, 8.5–10 mm long, style branches obovate, 0.7–1.3 mm long. Fruits ellipsoid, 7–9 × 5–7 mm when dry (8–10 × 5–8 mm when fresh), vermillion, turning black at maturity, glabrous, sessile. Pyrenes plano-convex, narrowly elliptic in outline, 9 × 3.0–3.5 mm, dorsal side with 4 hardly distinct ridges, ventral side with a deep narrow excavation for the whole length, ± C-shaped in cross-section, opening by 4 dorso-basal slits running along the ridges. Seeds entire, C-shaped in cross-section.

Endemic to the Guiana Shield, occurring in French Guiana, Suriname, Guyana, and the contiguous Brazilian states of Amapá and Pará; locally common, but apparently absent from the range of C. galbaoensis (except on the Montagne Bellevue de l’Inini).

Commonly encountered in mature terra firme rainforest, or sometimes in seasonally flooded forest, from near sea level to 600 m elevation.

Flowering collections were made in January (once), March to early May (main flowering season), and August (once). Fruiting specimens were collected from April to November.

This species is called carapiche (hence the generic name) by the Karipuna (“Garipons”; Aublet 1775) and tapiiwapa’a by the Wayampi (Grenand 270).

GUYANA • Rupununi District, Kuyuwini landing, Kuyuwini River; 2°05’N, 59°15’W; 10 Oct. 1992; fr.; Jansen-Jacobs et al. 2844; CAY, U.

SURINAME • Tumuc Humac Mountains, Talouakem, Litani River; 2°31’N, 54°45’W; 6 Aug. 1993; fr.; Acevedo-Rodriguez et al. 5924; CAY, U, US • Upper Litany River; 2°27’N, 54°48’W; 30 Jul. 1993; fr.; Granville et al. 11895; B n.v., BBS, CAY, MO n.v., P, U, US • Sipaliwini, vicinity of airstrip along Ulemari River, 71 km up Ulemari River from its confluence with Litani River; 3°6’17”N, 54°32’28”W; 150 m; 29 Apr. 1998; fr.; Hammel et al. 21736; MO n.v., U • Sipaliwini, 99 km up Ulemari River from its confluence with Litani River; 2°58’18”N, 54°33’14”W; 150 m; 8 Apr. 1998; fr.; Hammel et al. 21391; MO n.v., U • Lely Mts, SW Plateau; 27 Sep. 1975; fr.; Lindeman et al. 472; CAY, K, U • road Afobaka–Brownsweg, N of Brokopondo Lake; 10 Nov. 1974; Maas et al. 2335; U • flum. Saramacca sup.; Mar. 1903; Pulle 212; U • between Saramacca R. and Goliath Mt.; 9 Jun. 1956; J.P. Schulz 7702; U • opposite Gansee, right bank of Suriname R.; 12 May 1964; van Donselaar 1310; U • W bank of Marowijne Creek (= Gran Creek) near Gran Dam; 20 May 1966; van Donselaar 3435; U • Brokopondo District, W. of Brokobaka; 2 Nov. 1966; van Donselaar 3830; U • ad rivulum Palaime, flum. Sipaliwinin trib.; 2°8’N 56°12’W; 23 Feb. 1963; Wessels Boer 719; U • S of Tafelberg at the margin of Kappelsavanna; 10 Jun. 1963; Wessels Boer 1532; K, U.

FRENCH GUIANA • Crique Grand Laussat; 5°25’N, 53°37’W; 11 Jul. 2004; fr.; Barrabé 77; CAY, NY, P • RN2, pk 50; 11 Jun. 1984; fr.; Feuillet 1405; B n.v., CAY, MO, P, U, US • Rivière Itany × Koulé-Koulé; 20 Jul. 1985; fr.; Feuillet 2480; CAY • ibid.; 21 Jul. 1985; fr.; Feuillet 2504; CAY, P • Montagne de l’Inini; 3°30’N 53°30’E; 9 Apr. 1986; fr.; Feuillet 3730; CAY, MO n.v., P • ibid.; 11 Apr. 1986; fl.; Feuillet 3843; CAY • Eastern Plateau of Montagne Tortue; 13 Jun. 1988; Feuillet 9974; K, U • Riviére Kourouaï, Bassin de l’Approuague; 4°15’N, 52°1’W; 120 m; 8 Jul. 2008; fr.; Gonzalez 1361; CAY • Mont Itupé–sommet tabulaire – 15, versant W, layon D; 3°1’45”N, 53°5’28”W; 600 m; 15 Mar. 2010; fr.; Gonzalez 2158; CAY, MO, P, US • forêt à 1 km du saut “S” sur la rive droite du [fleuve] Grand Inini; 18 Aug. 1970; fr.; Granville C-1; CAY, COL, P, U • Monts Atachi Bacca; 5 Mar. 1971; fl.; Granville C-125; CAY, P • rive gauche du [Fleuve] Yaroupi, un peu au-dessus de Saut Ouaïmicouaré; 12 Apr. 1970; fl.; Granville 333; CAY [3 sheets], P, U • Crête des Monts Atachi Bacca; 5 Mar. 1971; fallen fl.; Granville 789; CAY, P • Route de Cacao à proximité de la Crique Boulanger; Mar. 1981; fl.; Granville 4428; CAY [2 sheets], P • Haute [Fleuve] Mana, lieu dit “Bellevue”, rive gauche; 18 Aug. 1981; fr.; Granville 4920; CAY • Sud de la Crique Martin, RN2, pk 22 environ; Apr. 1983; Granville 5559; fr. ; BR, CAY, P • Sud de la Crique Martin, RN2, pk 22 environ; Apr. 1983; Granville 5560; fl.; BR, CAY, P • Montagne de Kaw, versant nord à 2–3 km N de Camp Caïman; 1 Apr. 1984; fallen fl.; Granville 6710; BR, CAY, P, U • route de l’Est, à proximité de l’embranchement de la route de Cacao; 8 Apr. 1985; fl.; Granville 7256; CAY, P, U • Crique Gabaret, bassin de l’Oyapock, Saut Mérignan; 3°55’N, 51°48’W; 13 Apr. 1988; fl.; Granville 10279; CAY, P, U • Monts Atachi Bacca; 3°33’N, 53°55’W; 9 Jan. 1989; fl.; Granville et al. 10485; CAY, P, U • Camp Caïman, Montagne de Kaw; 4°32’N, 52°13’W; 11 Mar. 2004; fallen fl.; Granville & Bordenave 15828; CAY, P • Trois-Sauts en face du village Zidock; 21 May 1974; fr.; Grenand 270; CAY • savane-roche [inselberg] Virginie; 6 Apr. 2014; Lachenaud 1725; BR, CAY, MO • route de Cacao juste avant la scierie; 4 May 2014; Lachenaud 1792; BR, CAY, MO • Relais de Patawa, on the Montagne de Kaw road; 9 May 2001; fl., fr.; Mori et al. 25351; CAY, NY • Fleuve Oyapock, chemin Maripa, layon ORSTOM du km 1 au km 2; 8 Jun. 1970; fr.; Oldeman T-850; CAY • Oyapock, rive française face à Mariaflor, station de jaugeage hydrologique ORSTOM; 19 May 1965; fr.; Oldeman 1290; CAY, COL, P, U • rive gauche du [Fleuve] Yaroupi, Saut Ouaimicouaré; 15 Apr. 1970; fl; Oldeman B-3082; CAY • Rivière Grand Inini, au Saut Equerre; 18 Aug. 1970; fl.; Oldeman B-3508; CAY, P • Route de l’Est, km 53; 25 Aug. 1986; fr.; Prévost 2163; CAY [2 sheets], U • s.loc.; s.d. [1781–1785]; L.C.M. Richard s.n.; P • Abattis Cotica, sur le Maroni; 26 Aug. 1961; Schnell 11468; U.

BRAZIL – Amapá • Serra do Navio, Orebody slopes and trail to Serra do Viado; 16 Nov. 1954; fl.; Cowan 38335; K, NY, U, US • ibid.; 17 Nov. 1954; fl.; Cowan 38384; U • ibid.; fl.; Cowan 38416; U • immediately east of Colonia Agricola do Oiapoque, about 4 km N of mouth of Cricu River; 3°43’N, 51°55’W; 14 Aug. 1960; fl., imm. fr.; Irwin et al. 47516; F, IAN, K, NY [2 sheets]. – Pará • Rio Trombetas, 4 km S of Cachoeira Porteira; 6 Jun. 1974; fr.; Campbell et al. P22528 [Prance’s collection number]; INPA, NY, U, US.

Notes. The chaotic taxonomic history of this species has been summarized in Delprete (2003). It has often been confused with C. ligularis. Steyermark (1972), in particular, reduced it to a variety of the latter (as Psychotria ligularis var. carapichea). It is very strange that he arrived to this conclusion, since C. guianensis and C. ligularis are quite different in leaf venation (reticulate in the former and parallel in the latter) and bract shape (narrower and lacking an unguiculate basal part in C. ligularis) (Figs 5, 6). Delprete (2001) showed that they represent separate species. Taylor and Gereau (2013), in their revision of the genus, also recognized the two species as distinct, and even placed them in different groups, chiefly on account of their strikingly different leaf venation. On the other hand, C. guianensis closely resembles C. galbaoensis, which was until now considered a synonym (Delprete 2001, 2003; Taylor and Gereau 2013) and is here re-established as a distinct species, and C. squamelligera. The three species are almost identical in vegetative characters, and their differences are summarised in Table 2 (see also Figs 3, 4). The dimensions of the involucral bracts are rather variable in C. guianensis. Two collections from French Guiana (Feuillet 3730 and 3843) have the bracts unusually broad, resembling C. galbaoensis, but their basal stalk is shorter than in the latter species, and the corolla is typical for C. guianensis. At the other extreme, Wessels Boer 719 from Suriname and Schnell 11468 from French Guiana have exceptionally narrow bracts, and also differ from the rest of the material in their shortly corniculate corolla lobes; they are provisionally referred to C. guianensis but might prove to be distinct.

Figure 6. 

Carapichea ligularis. A. Habit, with two inflorescences. B. Detail of lower leaf surface. C. Inflorescence with open flowers. D. Infructescence with immature fruits. A from plant not collected, Montagne des Gouffres, French Guiana; B from Lachenaud 1793; C–D from plants not collected, Saül region, French Guiana. Photographs by Sébastien Sant (A, C–D) and Olivier Lachenaud (B).

FRENCH GUIANA [as “Guiana”] • s.loc.; s.d.; fl.; Martin s.n.; holotype: BM [BM001009103].

Description. Subshrub or shrub, 0.2–2(–4) m tall, single-stemmed or weakly branched (or rarely trees 7–10 m tall in the Brazilian and Venezuelan Amazon); terminal internodes terete, 2–3(–4) mm in diam., glabrous. Stipules shallowly sheathing, glabrous; sheath truncate to shallowly elliptic; lobes broadly triangular to broadly ovate, 2–5 mm long, persistent. Leaves with petioles 0.3–3.3 cm long, glabrous; blades elliptic, oblanceolate to narrowly oblong-elliptic, (4–)8.5–21 × (1.5–)2–6.5 cm, acute-decurrent at base, acute and long-acuminate at apex, acumen narrowly triangular, 0.5–2.5 cm long, sometimes falcate, papyraceous to coriaceous, drying pale olive-green to dark grey-green, glabrous throughout (except sometimes the domatia); secondary veins 14–26 on each side of the midrib; intersecondary veins 2–3(–4) between each couple of secondary veins, terminating far from the margin; tertiary veins barely visible or obsolete; domatia absent, or present as a row of hairs on each side of the midrib below. Inflorescence capitate to subcapitate (shortly branched, ramifications ≤ 0.5 cm when present); peduncle 1.5–7.0 cm long, glabrous to shortly pubescent; bracts inserted at distal end of peduncle and sometimes also on inflorescence axes, shallowly connate or free at base, (2–)4–8, usually decussate, narrowly lanceolate to linear, (7–)12–30 × (1–)2–4 mm, pale green or pale orange-green, glabrous, persistent. Flowers 5-merous, heterostylous, sessile to subsessile, the pedicels elongating to 2–5 mm long at fruiting stage. Hypanthium obovoid, 0.6–1.0 mm long, glabrous. Disk bilobed to the base, 0.5 mm long, shorter than or as long as the calyx. Calyx cupular, glabrous, tube 0.5–1.0 mm long, truncate or with short triangular lobes < 0.5 mm long. Corolla hypocrateriform, orange to salmon colored, 7.5–13(–14) mm long, glabrous; tube subcylindrical, gradually wider towards the mouth, 6–10.4(–11) mm long, 0.8–1.0 mm wide at base, 1.5–1.6 mm wide at mouth, glabrous outside and inside; lobes oblong-ovate, 2–4 × 0.8–0.9 mm, acute at apex, not corniculate, glabrous. Short-styled flowers: Stamens inserted at distal 1/3^rd of the tube, anthers partially exserted beyond corolla mouth; filaments ca 0.3 mm long; anthers 2.3–2.5 × 0.5 mm; style included, 3.5–4.0 mm long; style branches narrowly oblong, 1.5 mm long. Long-styled flowers: stamens inserted at about the middle of the corolla tube, included; filaments 1.5 mm long, glabrous; anthers narrowly oblong, 2.5 × 0.3–0.4 mm, acute at both ends; style exserted just beyond the corolla mouth, 12.5 mm long (corolla tube 10.5 mm long), style branches obovate, ca 1 mm long. Fruits ellipsoid to oblong-ovoid, 6–12 × 4–9 mm, smooth (slightly costate when dry), orange to orange-red. Pyrenes plano-convex, elliptic in outline, 7–9 × 3–5 mm, dorsal side with 3 ridges, ventral side with a very shallow longitudinal groove. Seeds with a deep T-shaped ventral furrow.

This species occurs in southeastern Venezuela (state of Amazonas), French Guiana, and eastern Amazonian Brazil (states of Amapá, Pará, Roraima, and Amazonas). It is locally common, at least in French Guiana. It could be expected in Suriname and Guyana, but we have seen no collections of the species from these countries; the type specimen, reported to be from Guyana (Delprete 2001: 399; Taylor and Gereau 2013: 120) was actually collected in French Guiana (see Notes below).

In understory of non-flooded forest, usually on superficial soils (e.g. lateritic crusts, granitic boulders) at 100–800 m elevation.

Flowering specimens were collected from August to January; and fruiting specimens throughout the year.

VENEZUELA – Amazonas • Cerro da Neblina, Río Yatua, along Upper Río Yatua between mouth of Río Yacibo and Piedra Arauicaua; 1 Feb. 1954; fl.; Maguire et al. 37411; F, US.

FRENCH GUIANA • Route de l’Est, ca 25 km NW of Régina; 4°23’N, 52°19’W; 16 Mar. 1994; fr.; Andersson et al. 1989; BR, CAY • Cacao; 27 Oct. 1983; fl.; Billiet & Jadin 1877; BR • Route RN2 Cayenne-Régina, pk 67, Crique Tibourou; 4°29’N, 52°19’W; 10 Feb. 1993; fr.; Billiet & Jadin 5733; BR • Extension Nord-Ouest des Petites Montagnes Tortue; 4°20’29”N, 52°16’16”W; 30 m; 29 Sep. 2010; fl., fr.; Bordenave & Le Hir 9068; CAY, K, P, MO • Montagne Tortue, Plateau Est; 4°17’N, 52°21’W; 460 m; 5 Oct. 2010; fl.; Bordenave & Le Hir 9233; CAY • Upper Maroni River, layon de chasse au NE d’Antecume Pata, confluent de l’Itany et du Marouini; 19 Nov. 1977; fl.; Cremers 5077; CAY • Saül, piste du carbet; 31 Oct. 1984; fl.; Foresta 705; CAY • Monts Atachi Bacca, autour du Camp 3, entre les sommets 525 et 782 m; ca 400 m; 5 Mar. 1971; fr.; Granville C-128; CAY [2 sheets], P • Monts Atachi Bacca; 3 Mar. 1971; fr.; Granville 752; P • Rivière Petite Ouaqui, rive droite, au niveau de l’ancien village Hubert; 20 Jul. 1973; fr.; Granville 1872; CAY, P • Sommet Tabulaire, zone nord; 2 Sep. 1980; fr.; Granville 3710; P • Monts Bakra, 2,5 km à l’ouest du Pic Coudreau; 2 Oct. 1980; fr.; Granville 4065; CAY, P • Mont Bellevue de l’Inini, central zone, slope near the summit; 23 Aug. 1985; fr.; Granville 7762; CAY, F, INPA, MG, MO, NY, P, U • Région de l’Inini, Mount Atachi Bacca, NW summit, 6 km E of Gobaya Soula, near Camp 2; 10 Jan. 1989; fl.; Granville et al. 10541; CAY, MO n.v., P, U • Montagne de la Trinité, Bassin de la Mana, partie supérieure de la pente N du plateau tabulaire; 13 Mar. 1997; fr.; Granville 13318; CAY, K, MO, P, U, US • Montagne Lucifer, SW du plateau sommital; 4°46’30”, 53°56’30”W; 520 m; 18 Nov. 1999; fr.; Granville & Crozier 13869; CAY, MO • Monts Bakra, 1.5 km W du Pic Coudreau; 3°18’N, 52°57’W; 600 m; 18 Jun. 2002; fr.; Granville et al. 14864; B, CAY, MO, P • Monts Kotika, plateau latéritique sommital; 3°55’10”N, 54°11’10”W; 730 m; 23 Feb. 2005; fr.; Granville et al. 16922; B, CAY, MO, NY, P, U • Route Cayenne–Régina, km 52; 4°34.638’N, 52°23.873’W; 9 Jan. 2011; fr.; Lachenaud 1074; BR, CAY • route de Cacao; 4 May 2014; fr.; Lachenaud 1793; BR, CAY • Mont Itoupé, Sommet Tabulaire, Layon D, 2me sommet; 3°02’35”N, 53°05’18”W; 760 m; 31 Mar. 2010; fr.; Molino & Sabatier 2843; BR, CAY, MO • Montagnes de la Trinité; 4°36’30”N, 53°21’30”W; 12 July 2001; fl. buds, fr.; Poncy & Crozier 1458; CAY, NY, P, U, US • RN2 – Cayenne–Régina, km 85.5; 4°20’N, 52°18’W; 26 Jun. 2003; fl., fr.; Prévost & Sabatier 4742; CAY, MO • Rivière Itany, Crique Petit Marouini, près de son embouchure; 2 Sep. 1972; fr.; Sastre 1814; P.

BRAZIL – Amapá • Upper Jari River, foret primaire humide; 2°28’N, 54°46’W; 420 m; 20 Aug. 1993; fl.; Granville et al. 12387; BBS, CAY [2 sheets], US. – Amazonas • Rio Curicuriarí, afl. do Rio Negro; 21 Dec. 1931; fl.; Ducke s.n.; INPA [No. 16533] • Nova Prainha, Rio Aripuanã, RADAM/BRASIL, SB-20-ZD, Ponto 10; 9 Jul. 1976; fr.; C.D.A. Mota s.n.; INPA [No. 60607] • Mun. Nova Olinda, Rio Paca, tributari of the Rio Mari Mari, terra firme forest, tree to 7 m tall; 2 Jul. 1983; fr.; Todzia et al. 2299; INPA, MO, NY, US • Mun. Axinim, basin of Rio Abacaxis, lower Rio Paca, ca 1 km from its confluence with Rio Mari Mari, forest on terra firme, shrub 2 m tall; 4°07’S, 58°58’W; 1 Jul 1983; fr.; Zarucchi et al. 2917; INPA, MO, NY. – Pará • Ilha de Marajó, Rio Jipurú, afluente do Rio Anajas; 20 Oct. 1987; fl.; A. Tavares & J. Cardoso 227; INPA, NY. – Roraima • Mun. Rorainópolis, Reserva Popular Xixuaú-Xiparina, ilha subindo o Rio Xipariná; 0°55’41”S, 61°50’26”W; 25 Aug. 2010; fl, young fr.; Zappi et al. 2902; INPA, MIRR.

This species has often been confused with C. guianensis (see Notes under that name) but is very different in leaf venation and bract shape (Delprete 2001; Figs 5, 6). It is much more similar to C. araguariensis and C. sp. A, from which it differs by the characters mentioned in the key.

Delprete (2001: 401) separated Carapichea ligularis from C. pacimonica by the lower habit, being a small, single-stemmed shrub 0.5–1.5 m tall (vs shrub up to 2 m tall or tree up to 10 m tall), the presence of corniculate appendages on the abaxial side of the corolla lobes (vs appendages absent), the subcapitate to sparsely cymose inflorescences (vs capitate), and the larger ovoid fruits (vs smaller, subspherical fruits). None of these characters proves to be reliable: the variation in habit and fruit seems to be continuous, the corolla appendages are commonly present in flower buds and usually fall off during or shortly after anthesis or during the preparation of herbarium specimens (Piero Delprete pers. obs.) and the inflorescences are usually capitate or nearly so at anthesis and often develop short ramifications in the fruiting stage. Taylor and Gereau (2013: 114) also kept the two species separate, based on other characters: “outermost inflorescence bracts at base straight-sided and not sheathing; plants often drying with a gray cast” in C. ligularis vs “outermost inflorescence bracts at base widened and shortly sheathing; plants often drying yellowish brown” in C. pacimonica. After a detailed study of numerous specimens throughout the geographic range, we observed that the inflorescence bracts can be shallowly sheathing or free at base in duplicate specimens of the same collection, and the colour of the dry plants may vary from greyish to olive green to pale brown to dark brown (possibly depending on the drying method). Taking into account all the above observations, we treat these two names as synonyms.

Steyermark (1972: 589) regarded Psychotria necopinata Standl. as a synonym of P. pacimonica, but we follow Taylor and Gereau (2013) in recognizing the former as a distinct species, Carapichea necopinata, which is only known from the type and not recorded from the Guianas to date.

The typification of C. ligularis has been a source of confusion. In his original description, Rudge (1806: 29, pl. 45) did not cite any gathering. Delprete (2001: 399) designated the lectotype as “Guyana, Rudge s.n. (P)”, which was followed by Taylor and Gereau (2013: 120). However, this lectotypification is erroneous since no such specimen exists at P, and Rudge never travelled to South America; the material that he used to describe the taxa in his Plantarum Guianae Rariorum Icones et Descriptiones was in fact collected by Joseph Martin in French Guiana. The fate of Martin’s specimens was described by Stearn and Williams (1957), and summarized by Stafleu and Cowan (1983: 971–972). Succinctly, in 1803, France and England were at war, and the specimens collected by Joseph Martin in French Guiana, originally intended for the Museum of Natural History of Paris, were confiscated by two British privateers and brought to London. About 400 of these specimens were bought by the British Museum (BM), and 772 were bought by Rudge and included in his own herbarium (Stafleu and Cowan 1983: 971–972). Also, Rudge gave a partial set of these specimens to Banks, whose herbarium became the founding collection of BM. After Rudge’s death, his widow donated his herbarium to BM in 1847. Therefore, most of Martin’s collections converged at BM, although some of his specimens are also reported to be at FI or FI-Webb (Stafleu and Cowan 1983: 971–972). These specimens have the penciled information “Guiana. Martin”, which may give the false impression that they were collected in modern day Guyana, while they actually came from French Guiana. After an exhaustive search for original material of S. ligularis, we were unable to find any specimen at FI and FI-Webb. However, a specimen with the indication “Guiana, Martin” penciled on the upper corner of the sheet, exists at BM [BM001009103], and being the sole original material, should be regarded as the holotype.

Müller (1881: 338), in the original description of Psychotria pacimonica, cited a single gathering, Spruce 3445, but he did not indicate the herbarium of deposit. Taylor and Gereau (2013: 120) assumed that the holotype is in M, but no specimen of Spruce 3445 was found there after an exhaustive search (Andreas Fleischmann pers. comm.). Specimens with this collection number are found in G, K, P, and W; the P sheet, which is the only one with open flowers, and has a label handwritten by Müller, is here designated as lectotype.

The photograph of the fruits published by Campos and Brito (1999: 627) as Psychotria pacimonica appears to represent another species, probably not a Carapichea.

Shrub, up to 1 m tall, glabrous; terminal internodes terete, 3–4 mm in diam. Stipular sheath truncate to shallowly ovate, 2–4.5 mm long, glabrous, persistent. Leaves with petioles 1.6–2.7 cm long, glabrous; blades narrowly oblong-elliptic, oblong-lanceolate to oblong-oblanceolate, 15–25 × 3.5–6.5 cm, acute-decurrent at base, acute and acuminate at apex, acumen narrowly triangular, 1.2–1.3 cm long, drying papyraceous and brown, glabrous throughout; secondary veins 17–22 on each side of midrib, with 2–3 intersecondary veins between each pair of secondary veins; tertiary venation reticulate; domatia absent. Inflorescence terminal, long-pedunculate, distally short-trichotomous, peduncle 8.2–8.5 cm long, glabrous, drying brown; branches 4–6 mm long, thick-fleshy, puberulous, terminating into cymules; each cymule subtended by 1–2 bracts; bracts unequal within each cymule, narrowly lanceolate to narrowly oblong-lanceolate, the longer ones 14 × 5.2 mm, the shorter ones 4.0 × 2.6 mm, persistent, drying brown. Hypanthium narrowly obovoid, 2.5 mm long. Calyx cupular, 0.7 mm long, undulate, glabrous. Corolla unknown (fallen off). Style 4 mm long, lobes narrowly elliptic (after corolla has fallen off). Fruits unknown.

Only known by two gatherings from the Kamoa Mountains, Guyana.

Understory of dense forest on brown sand with rocky outcrops, at 240–400 m elevation.

The two flowering collections were collected in November.

GUYANA • Upper Takutu-Upper Essequibo Region, Kamoa Mountains, 0–2 km N of camp on Kamoa River; 1°31’N, 58°49’W; 240 m; 11 Nov. 1996; fl.; Clarke 3081; MO, US • Upper Takutu-Upper Essequibo Region, Kamoa Mountains, 1.5 km S of Kamoa River; 1°31’N, 58°49’W; 400 m; 12 Nov. 1996; fl.; Clarke 3136; MO, US.

This probably new taxon is only known from two gatherings, both of which have inflorescences but no corollas left (one specimen still has a fully elongated style) and no fruits. It closely resembles C. araguariensis and C. necopinata, both of which are so far only known from Brazil. The differences with the former species are indicated in the key. From the latter species, which is still only known from the type (Taylor and Gereau 2013: 120), it may be separated by its smaller bracts (the largest ones up to 14 mm long vs 20–25 mm long in C. necopinata), longer inflorescence peduncle (8.2–8.5 cm long vs 5.0–5.5 cm long), shorter calyx (0.7 mm long vs 1.5–2.0 mm long) and leaves with 17–22 (vs 22–24) secondary veins on each side of the midrib. With so little material, it is difficult to assess the significance of these characters, and better collections are required to establish the identity of this taxon.

FRENCH GUIANA • sur la rivière Comté, en forêt sur la Montagne Soufflet; 12 Jun. 1975; fr.; Granville B-5288; holotype: VEN [No. 289722]; isotypes: CAY n.v., P [P06800571]

Shrub up to 1.5 m tall, branched; terminal branchlets terete, 1.5–2.5 mm in diam., glabrous, soon covered with a greyish-brown corky bark. Stipules free, narrowly triangular to lanceolate, 6–8 × 1.5–4.0 mm, entire, glabrous outside, villose at the base inside, becoming corky and soon damaged. Leaves with petioles 0.5–1.4 cm, glabrous; lamina elliptic, 8–18.5 × 1.8–6.7 cm, attenuate and decurrent on the petiole at base, acuminate at apex, slightly coriaceous when dry, entirely glabrous, drying dark brown or dark olive-green; midrib and secondary veins prominent on the upper side; secondary veins 6–9 on each side of the midrib, strongly ascending, curving towards the margin and almost reaching it; 2–3 intersecondary veins between each pair of secondary veins; tertiary veins reticulate, dense and conspicuous in the dry state. Inflorescences terminal, apparently erect, in involucrate heads, few-flowered; peduncle 4–5 cm long, glabrous; involucre orange, consisting of two unequal, decussate pairs of almost free bracts, the outer pair broadly obovate, 12–17 × 9–12 mm, the inner pair narrowly elliptic, 11 × 2.5–4.0 mm, both pairs erect, obtuse at apex, entirely glabrous, persistent in the fruiting stage; interfloral bracts numerous and much narrower than the involucral ones, linear, acute, ca 9 × 0.5 mm, glabrous, persistent in the fruiting stage. Flowers unknown. Calyx (in fruit) shortly cupuliform, truncate, ca 1 mm long, glabrous, persistent on fruit. Disk bilobed to the base, ca 1 mm long. Fruits narrowly ovoid, 12 × 5 mm when dry, glabrous, sessile. Pyrenes plano-convex, narrowly elliptic to oblong in outline, 11–14 × 4–7 mm, acute at apex, dorsal side nearly smooth with 3 very vague indications of ridges, ventral side with a deep longitudinal narrow excavation, ± C-shaped in cross-section, opening by 3 dorso-basal slits running along the ridges. Seeds entire, C-shaped in cross-section.

Distribution. Endemic to northeastern French Guiana, only known from the type specimen collected on Montagne Soufflet.

Ecology. The type label only reports that the specimen was collected in forest, without indicating the elevation.

The only known specimen, with fruits, was collected in June.

Notes. This species is only known from the fruiting type specimen. Presumably due to the incompleteness of the material, Steyermark (1984) noted that its immediate relationships were not evident. However, it closely resembles C. guianensis and C. galbaoensis in the marcescent stipules and in the shape, texture, and venation of the leaves – the three species being virtually indistinguishable in vegetative state – as well as in the shape and mode of opening of the pyrenes. The three species also share inflorescences with two pairs of involucral bracts – although Steyermark (1984) described and illustrated only one pair of involucral bracts in C. squamelligera, two pairs are actually present – and the flowers are surrounded by numerous bracteoles. They can be separated by the characters summarized in Table 2. The above description is mostly based on the P isotype; the holotype in VEN was seen as a photograph only, and an isotype from CAY cited in the original description was sent on loan to VEN in 1981 but has not been returned.

GUYANA • Upper Mazaruni River basin, Partang River, ridge of Merumé Mountains; 1140 m; 1 Jul. 1960; fr.; S.S. Tillett et al. 43946; holotype: NY [0013844], isotypes: COL [COL000004671], F [No. 1704833, 1704834], K [K000432817, K000432818], NY [0013845], P [P02428007], US [00131318, 00146632], VEN [No. 82283].

Shrub or treelet 1–3 m tall; terminal branchlets terete to slightly quadrangular, 4.0–5.5 mm in diam., glabrous, soon covered with a buff corky bark. Stipules shallowly sheathing at base, 3.5–8.0 × 5 mm, broadly ovate, glabrous, bilobed or irregularly fimbriate at apex, with lobes 1–5 mm long, soon corky and marcescent. Leaves opposite or ternate; petioles 1.5–3.0 cm long, glabrous; blades narrowly elliptic or oblong-elliptic, 14–25 × 5–10 cm, acute to decurrent at base, acute and long-acuminate at apex, acumen narrowly triangular to linear, 1.0–2.5 cm long, coriaceous, drying olive green above and yellowish green below, glabrous throughout; primary and secondary veins prominent on both sides; secondary veins 8–12 on each side of midrib, curving towards the margin and almost reaching it; tertiary venation densely and prominently reticulate in the dry state; domatia absent. Inflorescence thyrsoid, rather narrowly pyramidal, long-pedunculate (expanded at fruiting stage); peduncles 7.5–12.5 cm long, glabrous, drying brown; secondary branches whorled, 3–5 per node, spreading, 0.4–2.0 cm long (0.8–4.0 cm at fruiting stage), glabrous except fringes of hairs at nodes, terminating into cymules; cymules with 15–22 flowers; bracts 4–5 around each cymule, ovate to triangular, 1–4 × 0.7–2.0 mm, obtuse to rounded at apex, slightly concave, persistent, drying brown, glabrous except a row of hairs at the base inside. Flowers 5-merous, heterostylous, sessile. Hypanthium narrowly obovoid, 0.5–1.0 mm long, glabrous. Disk bilobed to the base, ca 1 mm long. Calyx cupular, 0.7–1.0 mm long, truncate or minutely denticulate, glabrous. Corolla hypocrateriform, 8–10 mm long, white, greenish-white or brownish-white; tube narrowly obconical to almost cylindrical, 6.0–6.5 mm long, 1.0–1.5 mm wide at base, 1.5–4.0 mm wide at mouth, glabrous outside, pubescent in distal portion inside; lobes lanceolate to triangular, 1.5–3.5 × 1.0–1.3 mm, acute at apex, glabrous. Short-styled flowers: stamens exserted, filaments 2.5 mm long, anthers narrowly oblong, 1.8 mm long; style included, 2.5 mm long, glabrous. Long-styled flowers: stamens included; style exserted, 8 mm long, shortly bifid, glabrous. Fruits ellipsoid to ovoid or subglobose, 7–10 × 5–8 mm, costate when dry, dark red or maroon (probably when immature) to purplish-black. Pyrenes plano-convex, elliptic to oblong in outline, 6–9 × 4.5–6.0 mm, dorsal side 3–4-costate, ventral side longitudinally sulcate. Seeds entire, C-shaped in cross-section.

Endemic to western Guyana, Potaro-Siparuni and Cuyuni-Mazaruni Regions, on Merume Mountains, Mount Ayanganna, and Mount Wokomung, which are the easternmost extensions of the Pakaraima Mountains.

Growing in dense scrub forest on tepui sandstone, at 1070–1570 m elevation.

Flowering specimens were collected in June and July, and fruiting specimens in February, March, June, and July.

GUYANA • Potaro-Siparuni Region, Ayanganna Slope; 2 Mar. 1960; fr.; R. Browne 118 (Forest Department of British Guiana No. 7942); NY • Potaro-Siparuni Region, Mount Ayanganna, E face, camp above first of four escarpments; 5°20’19”N, 59°56’46”W; 1070 m; 12 Jun. 2001; fl., fr.; Clarke et al. 9062; MO n.v., US; ibid., Clarke et al. 9063; MO n.v., US • Potaro-Siparuni Region, Mount Ayanganna, E face, plateau above third of four escarpments; 5°23’12”N, 59°58’36”W; 1570 m; 19 Jun. 2001; fr.; Clarke et al. 9350; MO n.v., US • Potaro-Siparuni Region, Mount Ayanganna, E face; 5°23’05”N, 59°58’33”W; 1545 m; 26 Jun. 2001; fl., fr.; Clarke et al. 9565; MO n.v., NY, US • Potaro-Siparuni Region, Mount Wokomung, easternmost pinnacle of massif; 5°05’34”N, 59°50’13”W; 1524 m; 30 Jun. 2003; fl. buds; Clarke et al. 10341; MO n.v., NY, US • Potaro-Siparuni Region, Mount Wokomung, base of fourth escarpment; 5°05’39”N, 59°50’36”W; 1375 m; 4 Jul. 2003; fr.; Clarke et al. 10508; MO n.v., NY, US • Potaro-Siparuni Region, Mount Wokomung, Little Ayanganna, summit of highest point of Mount Wokomung massif; 5°04’53”N, 59°50’26”W; 1660 m; 6 Jul. 2003; fl.; Clarke et al. 10588; MO n.v., NY, US • Potaro-Siparuni Region, Mount Wokomung, summit; 5°04’3”N, 59°51’42”W; 1560 m; 10 Jul. 2003; fl. buds; Clarke et al. 10713; MO n.v., US • Potaro-Siparuni Region, Mount Wokomung, summit; 5°04’03”N, 59°51’42”W; 1560 m; 10 Jul. 2003; fl. buds; Clarke et al. 10757; MO n.v., US • Potaro-Siparuni Region, Pakaraima Mountains, Mount Wokomung, summit plateau, from central plateau 1–2 km to escarpment; 5°04’N, 59°52’W; 1500–1530 m; 19 Feb. 1993; fr.; Henkel & Chin 1483; MO n.v., NY, US, Mount Ayanganna, east slope; 13 Mar. 2014; fr.; Radosavljevic et al. 146; P.

Notes. This species was placed by Taylor and Gereau (2013) in their Panurensis group, together with C. panurensis from Brazilian and Colombian Amazon, but the two differ in so many characters that a close relationship between them seems unlikely. In fact, C. panurensis has several aberrant characters within the genus: almost spiciform inflorescences (the lateral branches being extremely reduced), an entire disk, large mitriform stipules usually with a prominent midrib, and secondary leaf veins forming conspicuous loops far from the margin. On the other hand, C. tillettii has thyrsoid inflorescences, a bipartite disk, stipules smaller than those of C. panurensis and without prominent midrib, and secondary leaf veins looping near the margin. All these characters fit very well with Taylor and Gereau’s (2013) Carapichea group, where C. tillettii most closely resembles C. franquevilleana and C. klugii. It differs from these species (which are probably not distinct from each other) by its pyramidal inflorescence with ramifications shorter than rachis (vs not or hardly so), smaller and coriaceous bracts, corolla tube 6.0–6.5 mm long (vs 3 mm long), and sessile fruits (vs shortly pedicellate). As noted by Taylor and Gereau (2013), the leaves of C. tillettii may be opposite or verticillate, sometimes with both conditions on the same branch, and the stipules can be bilobed or irregularly 3–5-fid at apex. The locality, altitude, collection date, and field notes of the type collection have been wrongly cited both in the protologue (Steyermark 1972: 498) and in Taylor and Gereau (2013: 123). The type label actually reads “Tree to 3 m; flowers white; fruit dark red; occasional in wet forest along trail, ridge of Merume Mountain, elev. 1140 m.”. This species is not to be confused with Rudgea tillettii Steyerm. (Steyermark 1976: 416), which is a synonym of R. coussareoides (Standl.) C.M.Taylor, Bruniera & Zappi (Taylor et al. 2015). The latter somewhat resembles C. tillettii in general appearance, but its stipules are basally connate into a truncate sheath and bearing dorsal appendages (these soon caducous), its flowers are sparsely arranged (not densely crowded at the apex of inflorescence ramifications) and its disk is entire.

GUYANA • Upper Mazaruni River Basin, Mount Ayanganna, along NE side; 800–900 m; 2 Aug. 1960; fr.; S.S. Tillett et al. 45008; holotype: NY [NY00132855]; isotypes: F [No. 1704839], COL [COL000004672], K [K000432825], NY [00146649], US [00146649], VEN [No. 82281].

Shrub or small tree, up to 4 m tall, much branched; terminal branchlets terete or slightly quadrangular, 3–7 mm in diam., somewhat succulent, glabrous, soon covered with a buff corky bark. Stipules free, oblong-ovate, 7–12 × 7–9 mm, obtuse and entire when young (apical and distal nodes), later splitting into two lobes at older nodes, each lobe oblong-ovate, 4–9 mm long, shortly apiculate, soon corky, persistent or fragmenting. Leaves with petioles 1.0–3.5 cm long, glabrous; blades elliptic to oblong-elliptic, 12–22 × 5.5–10 cm, acute at base, acute at base, acuminate at apex, acumen narrowly triangular, 0.7–1.2 cm long, papyraceous to subcoriaceous, drying dark brown above and pale brown below, glabrous throughout; secondary veins 10–13 on each side of midrib, curving towards the margin and almost reaching it; tertiary veins rather densely and prominently reticulate in the dry state; domatia absent. Inflorescence capitate, short-pedunculate; peduncle 0.5–1.7 cm long, glabrous; head included in a large involucral structure, 5.5–9.0 cm long, green at base and pinkish-red at apex, formed by the basally fused bracts, glabrous; involucral basal portion urceolate, 2.5–5.7 × 2.3–3.5 cm, thick and fleshy, distal portion flaring, funnel-shaped, 2- to 4-lobed, lobes ovate to broadly ovate, 2.5–3.5 × 3.5–4.0 cm, membranaceous; internal bracts (when present) ligulate, orbicular to round, 1.7–2.5 × 1.0–1.7 cm; bracteoles ligulate to linear, 0.7–1.5 mm, glabrous. Flowers 5-merous, (heterostylous?), pedicellate; pedicels 1.0–3.5 mm long (5–7 mm long in mature fruits), glabrous. Hypanthium narrowly cylindrical, ca 1 mm long, glabrous. Disk bilobed to the base, ca 1 mm long. Calyx cupular, dentate, 1.0–1.7 mm long, glabrous; tube 0.7–1.0 mm long; teeth deltoid to narrowly triangular, 0.3–0.7 mm long. Corolla narrowly tubular, 18–31.5 mm long, glabrous, white; tube narrowly cylindrical, 17–30 mm long, 1.5–2.5 mm wide, glabrous outside, with a sericeous ring just above the base and glabrous above inside; lobes narrowly ovate, 1.0–1.5 × 0.7 mm, acute at apex, glabrous outside. Stamens inserted near the base of the corolla tube, included; anthers subsessile, linear, 3–4 × 0.2 mm, glabrous. Style barely exserted among the corolla lobes, 17–34 mm long, glabrous, branches oblong, 0.7–0.8 mm long. Fruits ovoid, 6–12 × 4.5–8.0 mm when dry (“20 × 15 mm” when fresh, fide Henkel & Hoffman 187), costate when dry, bright blue to bluish-purple. Pyrenes plano-convex, elliptic to ovate in outline, 5–10 × 3.0–6.5 mm, dorsal side 5-costate, ventral side longitudinally sulcate. Seeds with a deep ventral furrow, ± T-shaped in cross-section.

Endemic to Mount Ayanganna in western Guyana.

In understory of tall wet evergreen forest, on brown sandy lateritic soil, at 712–1100 m elevation.

One flowering specimen was collected in June, one specimen with flowers and young fruits was collected in November, and two specimens with mature fruits were collected in March and August.

GUYANA • Potaro-Siparuni Region, Mount Ayanganna, E face; 5°20’04”N, 59°55’30”W; 712 m; 4 Jun. 2001; fl., fr.; Clarke et al. 8963; MO n.v., NY, US • Cuyuni-Mazaruni Region, Pakaraima Mountains, toe slope on NW side of Mt. Ayanganna; 5°24’N, 59°57’W; 1000–1100 m; 8 Nov. 1992; fl.; Henkel & Hoffman 187; K, MO n.v., NY, U, US • Mount Ayanganna, E slope, vicinity of new airstrip; 5°18’08”N, 59°50’10”W; 689 m; 9 Mar. 2014; fr.; Radosavlijevic et al. 103; MO n.v., US.

Notes. This species is remarkable by its large, urn-shaped involucre, varying from bilobed to unequally 4-lobed, which is formed by the fusion of the external bracts. The involucre is reported to be red (Taylor and Gereau 2013) although on the type label it is more precisely described as “green at base, shading through brown to pinkish-red at tips”. Carapichea urniformis was placed by Taylor and Gereau (2013) in their Altsonii group, together with C. altsonii, C. nivea, and C. sandwithiana. However, as discussed above, it is quite different from these three species (which are here transferred to the genus Notopleura) in characters of the stipules, leaf venation, and pyrenes, and is better placed in Taylor and Gereau’s (2013) Carapichea group, alongside with C. guianensis and C. galbaoensis, which are quite similar in vegetative characters. Due to the small number of collections, it is not known whether the flowers of this species are heterostylous. In the few flowers that we analyzed, the stamens are included and the style is barely exserted, which is consistent with a long-styled form.