Faramea humicapiens Delprete & J. G. Jardim, 2025

2. Faramea humicapiens Delprete & J. G. Jardim sp. nov.

Figs 3 View Figure 3 , 4 View Figure 4 , 5 View Figure 5

Type

FRENCH GUIANA • Maroni River Basin, Gobaya Soula, rive droite, layon vers les Monts Atachi-Bacca, entre 0–1.5 km de l’Itany [River], camp 1 ; 3°37’N, 53°58’W; 100 m; 6 Jan. 1989; fl.; Granville et al. 10434; holotype: CAY [Sheet 1 of 2, CAY 074825 About CAY , Sheet 2 of 2, CAY 074826 About CAY ] GoogleMaps .

Diagnosis

Faramea humicapiens is similar to F. tinguana in being erect, monocaulous, woody plants up to 1 m tall, with large, sessile or subsessile leaves in apical rosettes, gathering leaf litter at their bases, and short, axillary, 1–3 - flowered inflorescences. The former differs from the latter in having sessile or subsessile inflorescences with 1, 2, or 3 flowers (vs pedunculate, with peduncles 6–8 mm long, cymose, commonly 3 - flowered in F. tinguana ), bracteoles subtending the hypanthia lanceolate to narrowly lanceolate, 6–12 × 0.7–3 mm, with entire margin, acute to acuminate at apex (vs subtending the pedicels, narrowly ovate to oblong, 2.5–3.5 × 0.7–1.5 mm, with fringed margins, often with 2–3 apical lobes 0.3–0.5 mm long), flowers sessile to subsessile, with pedicels (when present) to 1.5 mm long (vs with pedicels 2–4.5 mm long), calyces 1.3–1.5 mm long (vs 2.8–3 mm long), corollas 17–20.5 mm long, with tubes 12.5–14 mm long and lobes 3.5–6.5 × 1.5–1.8 mm (vs 27–29.5 mm long, with tubes 16–17 mm long and lobes 11–12.5 × 3.5–4 mm), and anthers 2.5 mm long (vs 4.5 mm long).

Description

Single-stemmed plant, 20–80 cm tall, erect; rarely the basal portion of the stem leaning on the ground, with a few adventitious roots; stem woody, terete or laterally compressed, sometimes with a central longitudinal groove, glabrous. Stipules free or shallowly connate at base, persistent, long-aristate, 12–25 mm long, glabrous, base deltoid to narrowly triangular, 4–9 × 4–6 mm, margins thickened-cartilaginous, arista 8–17 mm long. Leaves subsessile to short-petiolate; petioles 2–5 mm long, basally thickened, glabrous; blades narrowly long-obovate, oblong-obovate to narrowly spatulate, often narrow-oblong at basal portion, (8 –) 16–42 × (3.5 –) 5–17 cm, cordate at base, acuminate at apex, acumen 1–1.5 cm long, dark green above, pale green below, drying olive-green, subcoriaceous, glabrous throughout; secondary veins 12–22 on each side; domatia absent. Inflorescences axillary, sessile to subsessile, 1–3 - flowered. Bracts subtending flowers lanceolate to narrowly lanceolate, 6–12 × 0.7–3 mm, membranaceous, margin entire, acute to acuminate at apex. Flowers 4 - merous, sessile or subsessile, pedicels (when present) up to 1.5 mm long. Hypanthium obovoid, 1–1.1 × 0.8 mm, glabrous. Calyx cupular, 1.3–1.5 mm long, truncate, glabrous. Corollas hypocrateriform, 17–20.5 mm long, glabrous throughout, white or pale purplish-white during and after anthesis; tube 12.5–14 mm long, cylindrical, 1–1.5 mm wide throughout; lobes lanceolate, 3.5–6.5 × 1.5–1.8 mm, acute at apex. Long-styled flowers stamens included, subsessile, inserted 8.5 mm below corolla mouth; anthers narrowly oblong, 2.5 × 0.3 mm, round at both ends; style included, as long as corolla tube (lobe at corolla mouth), bilobed, lobes narrowly lanceolate, 1 mm long, acute at apex. Short-styled flowers stamens partially exserted, subsessile, inserted near the corolla mouth; anthers oblong, 2.5 × 0.4–0.5 mm, apiculate at both ends; style included, much shorter than corolla tube, 5–6 mm long, bilobed, lobes narrowly lanceolate, 2–2.5 mm long, apiculate. Fruits sessile or subsessile, or with pedicel (when present) up to 1.5 mm long, globose to subglobose, 10–15 mm in diam., purple, bluish-violet or dark blue at maturity when fresh; 9.5–12 mm diam., dark brown when dry.

Distribution

Endemic to French Guiana (Fig. 3 View Figure 3 ).

Ecology

Growing in undercanopy of tall dense ombrophilous forests, in soil rich in organic material and significant accumulation of leaf litter, at 5–400 m altitude.

Phenology

Specimens with flowers were collected in October, December, January, and February, with immature fruits in April and May, and with mature fruits in February, April, August, September, October, and November.

Vernacular name

French Guiana: Uwakaya (Wayapi, Grenand 824 A).

Etymology

The specific epithet is derived from the Latin words “ humus- ” (“ ground ”) and “ - capiens ”. In this sense, “ capiens ” is the present participle of the transitive verb “ capere ” (capio, capis, cepi, captum), which means “ to hold ”. Hence, the specific epithet means “ holding humus ”, referring to the ability of this plant to accumulate organic material at the leaf bases.

Preliminary IUCN conservation assessment

Vulnerable: VU B 1 ab (i, ii, iii, iv). This species is most likely endemic to French Guiana. It has an AOO of 28 km 2 and an EOO of 16,536 km 2, indicating that it could be classified as EN and VU, respectively. However, according to the following considerations, we suggest to treat it as vulnerable. The vast majority of the collections are from three main regions, along the coast, from the Oyapock River Basin, and from the interior of the country, corresponding with 10 locations. The region along the coast is suffering from considerable human pressure due to selective logging and agricultural activities. The Oyapock River Basin is subject to important human pressure, due to increasing local populations. Most collections were made in the Saül area, a region of free adhesion of the French Guiana Amazonian Park, and with significant human pressure due to growing human population, increasing agricultural pressure, frequent tourist visitation, and nearby gold mining exploitation. In total, we studied 28 gatherings present in CAY, MO, NY, P, and US. Most of them are only represented by unicate specimens, and only four of them by two duplicates. This fact is here interpreted as the evidence of a species present in eleven main localities, but locally extremely rare. Exhaustive searches were undertaken in three localities (Mt. de Kaw, 4°31’N, 52°12’W; Oyapock River basin, Crique Gabaret, 3°55’N, 51°55’W; Crique Probert, 3°52’N, 51°48’W), in January 2024, during the flowering season of the species, but it was not found. All the above information is leading us to conclude that, although it is reported from several localities, this species is locally very rare, hence prone to be locally obliterated by human activities. Therefore, we classify this species as Vulnerable (VU) under IUCN criteria B 1 b ( IUCN 2012; IUCN Standards and Petitions Committee 2019), because it is interpreted as occurring in several small subpopulations (or a sole individual) and under considerable human pressure.

Additional specimens examined (paratypes)

FRENCH GUIANA • Saül, Monts La Fumée ; 3°37’N, 53°12’W; 200–400 m; 24 Aug. 1982; fr.; Boom & Mori 1565; NY GoogleMaps • Commune de Régina, Bassin de l’Approuague ; 75 m; 5 Dec. 1994; fl.; Bordenave 1291; CAY • Camp Caïman, Asarco, Mt. de Kaw ; 4°31’N, 52°12’W; 10 m; 24 Jan. 2000; fl.; Bordenave & Raes 5140; CAY, U GoogleMaps • Région de Saül, Savane-Roche [Inselberg] Dachine ; 3°28’N, 53°13’W; 150 m; 5 Apr. 1997; imm. fr.; Cremers & Crozier 14730; CAY GoogleMaps • Nouragues, chablis au [parcelle] OX ; [ca 4°4’N, 52°43’W]; 20 Oct. 1995; fr.; Eockle 119; CAY GoogleMaps • Saül, forêt primaire; [3°37’N, 53°13’W]; 17 Oct. 1984; fr.; de Foresta 660; CAY GoogleMaps • Saül, Circuit Plateau La Douane ; [3°35’N, 53°12’W]; 30 Sep. 1974; fr.; Granville B- 5177; CAY GoogleMaps • Saül, tracé ORSTOM de la Crique Limonade à ca 2 km du village ; [3°34’N, 53°13’W]; 3 Nov. 197; fr.; Granville 2274; CAY GoogleMaps • Saül, Roche Bateau, sur la Crique Nouvelle France, ca 6 km E du village ; [3°35’N, 53°10’W]; 6 Nov. 1974; fr.; Granville 2292; CAY GoogleMaps • Nord du Massif des Emerillons, colline entre la Haute Approuague et la Crique des Emerillons ; [3°15’N, 53°5’W]; 20 Sep. 1980; fr.; Granville 3941; CAY, P GoogleMaps • Saül, sommet des Monts La Fumée ; 410 m; 12 Oct. 1982; fr.; Granville 5131; CAY [2 sheets] • Région de Saül, colline à ca 18 km au S de Saül ; 4 Apr. 1983; imm. fr.; Granville 5549; CAY • Mont Galbao, entre Crique Mana et le sommet ; 3°36’N, 53°17’W; 350 m; 12 Jan. 1986; fl.; Granville et al. 8647; CAY, US GoogleMaps • Oyapock River basin, Crique Gabaret, Saut Plat ; 3°55’N, 51°55’W; 5 m; 14 Apr. 1988; fr.; Granville 10293; CAY GoogleMaps • Camopi River Basin, Saut Petit Continent ; 2°54’N, 52°51’W; 90 m; 13 May 1992; st.; Granville 11670; CAY GoogleMaps • Oyapock River Basin, Roche Touatou ; 2°57’N, 52°32’W; 140 m; 22 May 1995; imm. fr.; Granville & Cremers 13035; CAY GoogleMaps • Région de Saül, 3 km E de Saint-Eloi, sur le layon vers Pic Matécho ; 3°44’N, 53°14’W; 200 m; 21 Sep. 2000; fr.; Granville & Crozier 14339; CAY GoogleMaps • Projet Routier entre Saint-Georges et le pont transfrontalier sur l’Oyapock, zone Crique Probert ; 3°52’50”N, 51°48’46”W; 6 m; 20 Dec. 2005; fl.; Granville & Crozier 17220; CAY, MO GoogleMaps • Mont Galbao, entre Crique Mana et le sommet ; 3°36’N, 53°17’W; 350 m; 13 Jan. 1986; fl.; Granville et al. 8647; CAY GoogleMaps • Région de Saül, 3 km E de Saint-Eloi, sur le layon vers le Pic Matecho ; 3°44’N, 53°14’W; 200 m; 21 Sep. 2000; fr.; Granville et al. 14339; CAY GoogleMaps • Trois Sauts ; [2°14’N, 52°52’W]; 14 Mar. 1975; st.; Grenand 824 B; CAY GoogleMaps • Saül, parcelle Belvédère ; 9 Feb. 1974; fr.; Jacquemin 1445; CAY • Saül, Les Eaux Claires, ca 300 m N from Les Eaux Claires to Belizon, ca 50 m on the right side from the bridge ; 3°40’N, 53°13’W; 200–400 m; 4 Sep. 2000; fr.; Junikka & Orava 3014; CAY GoogleMaps • Saül, trail from large fig to Creek Limonade ; 3°37’N, 53°12’W; 250–300 m; 18 Aug. 1988; fr.; Mori et al. 19043; NY GoogleMaps • Saül, vicinity of Eaux Claires, Sentier Botanique, from 450 to 1060 m (just past Creek Tortue) beyond entrance ; 3°37’N, 53°12’W; 250–350 m; 11 Feb. 1993; fr.; Mori et al. 22914; NY GoogleMaps • Yaroupi River, rive droite, Saut Tainoua ; [2°37’N, 52°42’W]; 17 Apr. 1970; imm. fr.; Oldeman B- 2998; CAY GoogleMaps • Yaroupi River, rive gauche, Saut des Polissoirs ; [2°36’N, 52°42’W]; 23 Apr. 1970; fr.; Oldeman B- 3063; CAY GoogleMaps • Inini River Basin, Znieff Atachi-Bakka ; 3°39’35”N, 53°52’06”W; 90 m; 7 Oct. 2011; fl.; Silland 105; CAY GoogleMaps .

Notes on Neotropical litter-gathering Rubiaceae

Single-stemmed plants with large sessile, subsessile, or short-petiolate leaves in apical rosettes, collecting leaf litter, are present in numerous Rubiaceae genera. In the Neotropics, there are several species of Rubiaceae reported to be litter-gathering. One example is the recently described Rudgea quisquiliae Bruniera & Torres-Leite ( Torres-Leite et al. 2016: 192, figs 1–2), from the mountain slopes of the Brazilian state of Espírito Santo. Aside from those characteristics, R. quisquiliae is similar to Faramea tinguana and F. humicapiens in having axillary (which is exceptional in Rudgea Salisb. ), congested inflorescences, and differs from Faramea by the characteristic stipules with numerous appendages, and white fruits (vs aristate stipules and commonly blue fruits). The ecological preference of R. quisquiliae , F. tinguana , and F. humicapiens are also similar and typical of litter-gathering species, as they grow in the undercanopy of dense ombrophilous forest on mountain slopes.

Another Brazilian litter-gathering species of Rudgea , growing in ombrophilous forest, is R. macrophylla Benth. ( Bentham 1850: 456) , which is also found on the mountain slopes of the state of Rio de Janeiro. Rudgea macrophylla was personally observed growing sympatrically with Faramea tinguana on the Serra do Tinguá, and displays a rather variable habit, which changes with the aging of individuals (Fig. 6 View Figure 6 ). Young individuals of R. macrophylla are single-stemmed woody plants 0.5–1 m tall, topped by a rosette of litter-gathering leaves 20–40 cm long (Fig. 6 B View Figure 6 ). As the individuals continue to grow, they become sparsely branched treelets 1.5–3 m tall. The older, ramified individuals maintain the large, ascending leaves in dense rosettes at the apex of each branch, which continue to collect leaf litter even when they become 3 m tall (Fig. 6 A View Figure 6 ). Mantovani and Vieira (1993) studied the leaf surface of R. macrophylla , and did not report any foliar anatomical character that could be associated with the litter-gathering habit of this species.

Taylor (2002: 564–566, fig. 1 D) published Faramea accumulans C. M. Taylor , a species found on San Blas Island, Panama, in wet forests at 50–180 m altitude. She described it as a tree up to 5 m tall, entirely glabrous, with elliptic-oblong leaves, 28–44 × 13–24 cm, and axillary, cymose inflorescences. She also added “ This species apparently accumulates detritus in the clasping leaf bases that surround the stem, and the specific epithet refers to this habit. ”

The litter-collecting Psychotria dressleri (Dwyer) C. W. Ham. ( Hamilton 1998: 223) occurs in Panama and Colombia. Because of the bi-dentate stipules and several other morphological characters, this species should probably be transferred to Palicourea Aubl. , as currently broadly delimited (e. g. Taylor 2015 a, 2015 b; Delprete and Lachenaud 2018). Psychotria alfaroana Standl. ( Standley 1928: 273) , a species found in Costa Rica, Panama, and Colombia, also has bicuspidate stipules, and large, litter-gathering leaves in apical rosettes. Morphological and phylogenetic studies are necessary to eventually support the transfer of these two species to Palicourea s. l. If these transfers are supported, they would be two litter-gathering species of Palicourea .

Palicourea woronovii (Standl.) Bruniera & C. M. Taylor (in Taylor et al. 2015: 2, fig. 2), is another litter-gathering species from Colombia. It was originally described by Standley (1930: 155) as Rudgea woronovii Standl. , as having succulent stems, laciniate stipules 3–4 cm long, and large, oblong-ovate leaves, ca 30 × 12.5 cm, abruptly acuminate at apex. Bruniera and Taylor (in Taylor et al. 2015: 2–3), following the observation of additional specimens, corrected Standley’s stipules description as being “ two-lobed, with the lobes relatively broad and glandular-fimbriate. ” They also added that “ It has the habit of a ‘ rubbish-bin’ plant (fig. 2 B), with a low (0.5–1 m tall) unbranched habit and detritus held at the stem nodes; in most such plants detritus accumulates in the axils of the subsessile or sessile leaves, but in Palicourea woronovii the detritus is held by the enlarged stipules and the leaf axils. ”

An additional litter-gathering species was recently described from the Brazilian state of Rio de Janeiro. Palicourea goytacaz Torres-Leite (in Torres-Leite et al. 2025: 294, fig. 1–2) was described as “ Shrubs up to 3 m tall, much branched, with litter-trapping habit, i. e., with the leaves grouped at the apex of the branches and favoring litter accumulation. [...] Leaves with blades (18 –) 24–30 (– 36) × (6 –) 10–15 (– 19) cm, smaller at at the base of the inflorescence 6–15 × (2.5 –) 3–5 (– 7.5) cm, obovate to elliptic, base cordate, subcordate or acute [...] ”, mentioning its habitat as “ understory of a dense ombrophilous forest near a watercourse. ”

Zona and Christenhusz (2015) presented a worldwide review of litter-gathering plants, analysed the terminology present in literature, and provided the following definition “ Litter-trappers can be defined as those plants that, by virtue of their growth habit and morphology, trap or channel falling debris (leaf litter, twigs, flowers, fruits, bird droppings, dead animals, etc.) and use the nutrients derived from this detritus for their own growth. ” For the family Rubiaceae , they stated that litter-trapping species are “ primarily African, despite the fact that Rubiaceae are abundant throughout the American and Asian tropics. ” In tropical Africa, litter-gathering Rubiaceae species are more numerous than in the rest of the World. Lachenaud and Jongkind (2013) described two new African litter-gathering species of Psychotria L. , P. blydeniae O. Lachenaud & Jongkind and P. tetragonopus O. Lachenaud & Jongkind , which are both single-stemmed woody plants, either 30–50 cm tall, or 120–150 cm tall, respectively, with shortly petiolate large leaves. In the same article, those authors cited about a dozen litter-gathering Rubiaceae occurring in Africa, namely, one species of Chassalia Comm. ex Poir. , two species of Chazaliella E. M. A. Petit & Verdc. , one species of Coffea L. , one species of Ixora L. , one species of Oxyanthus DC. , and several species of Pavetta L. and Psychotria . Lachenaud (2019: 68–70, table 3) augmented the number of African litter-gathering Rubiaceae to a total of 44, listing one species of Bertiera Aubl. , two species of Chassalia , one species of Coffea , two species of Eumachia DC. , one species of Ixora , one species of Oxyanthus , five species of Pavetta , and 31 species of Psychotria (many of them newly described). All those species share a similar habit and ecological preference as those from the Neotropics, by growing in dense ombrophilous forests, and by being mostly 30–150 cm tall, usually single-stemmed or rarely sparsely branched, with large leaves in apical rosettes.

Zona and Christenhusz (2015: 572–573, table 2), listed 25 species of litter-gathering Rubiaceae , in several genera from Africa, one species of Ixora from mainland Asia, five species of Morinda L. from the Samoa Islands, and six species from the Neotropics. For the Neotropics, they listed the following Rubiaceae : Pentagonia macrophylla Benth. ( Costa Rica, Panama, Colombia, and Ecuador), Pentagonia wendlandii Hook. f. ( Costa Rica, Panama, and Colombia), Psychotria alfaroana (see discussion above), Psy. chitariana Dwyer & C. W. Ham. ( Costa Rica), Psy. dressleri (see discussion above), and Psy. insueta (Dwyer) C. W. Ham. ( Panama). In conclusion, adding Rudgea quisquiliae , R. macrophylla , Faramea accumulans , F. tinguana and F. humicapiens , and Palicourea woronovii and P. goytacaz , to Zona and Christenhusz’s list, there are at least 13 litter-gathering species of Rubiaceae in the Neotropics.