Drymonia quadrangulata Clavijo & J. L. Clark 2024, sp. nov.

Drymonia quadrangulata Clavijo & J. L. Clark sp. nov. Figs 2, 3 Diagnosis. Differs from Drymonia killipii by quadrangular to strongly angulate stems in cross-sections vs. terete; stems strigose apically vs. hirsute; blades 12.0– 26.7 cm long vs. 20.5–45.0 cm long; corolla lobes white to yellow vs. royal purple to maroon; and fruits indehiscent globose berries vs. bivalved fleshy capsules. Type. Ecuador. Imbabura: Ibarra. Parroquia: Lita. Unpaved road heading south, near Rocafuerte (between San Gerónimo and La Carolina), accessed via the Lita-San Lorenzo highway (KM 98), road recently developed through the SolGold mining concession (Alpala); 0.7526389°N, 78.373889°W; 1550–1650 m; 23 Jul 2022; J. L. Clark, Álvaro Pérez, Francisco Tobar & Russell Clark 17079 (holotype: QCA; isotypes: COL, CUVC, E, F, G, MO, NY, QCNE, SEL! [barcode: SEL 086219], US). Description. Terrestrial herb or shrub, 0.6–1.5 m tall. Stem scandent basally and then erect, usually branched, adventitious roots usually absent, quadrangular in cross-section to strongly angulate, sometimes winged, 4.2–10.5 mm in diameter, herbaceous to succulent, green to green with maroon spots, smooth, strigose apically, becoming glabrescent with age, lenticels sometimes present, internodes 2.2–10.2 cm long, reduced toward the apex. Leaves opposite, decussate, equal to subequal in a pair; petiole (2.2 –) 5.2–15.0 cm long, green with maroon spots, terete in cross-section, grooved, flattened at base, pairs of petiole bases fused together forming a perfoliate-like flap or wing, 1–2 mm wide, petiole enations present at base of petiole, strigose in apical leaves, glabrescent in basal leaves; blade elliptic to ovate, 12.0–26.7 × 5.2–18.5 cm, coriaceous, green adaxially, light green suffused with maroon abaxially, brown-maroon when dried, apex acute to acuminate, base obtuse, rounded or subcordate, margin serrulate to serrate, minutely strigose to glabrescent adaxially, glabrescent abaxially, 5–7 pairs of main lateral veins, minutely puberulent, reticulated, evident on both surfaces but more so abaxially. Inflorescence axillary, a reduced pair-flowered cyme with 1–6 flowers per inflorescence; bracts usually caducous, 7.0–16.9 × 2.0– 3.7 mm, light green suffused with maroon to mostly maroon, lanceolate to oblong, apex acute, base obtuse, margin entire, glabrescent adaxially, strigulose abaxially; peduncle absent. Flowers non-resupinate; pedicel erect to perpendicular 9.5–42.0 mm long, green, strigulose, enations scattered along the pedicel. Calyx green, green with reddish or maroon margins, or mostly maroon, membranaceous, persistent in fruit, calyx lobes 5, 4 nearly equal, nearly free, fused at the base for 0.7–2.0 mm, with margins overlapping at least half their length, lanceolate, apex attenuate, base cordate, margins entire to minutely serrulate, glabrescent adaxially, glabrescent but strigulose at base abaxially, ventral and lateral lobes 25.4–53.2 × 10.3–36.0 mm, dorsal lobe smaller, 25.0–37.6 × 8.4–19.0 mm. Corolla zygomorphic, protandrous, oblique to perpendicular relative to calyx, tubular, 42.7–64.6 mm long; tube gibbous at base, slightly constricted above base and wider at the middle, 30.8–49.5 mm long, 5.8–9.9 mm wide at constriction above base, 10.7–16.6 mm wide at the middle, outside white and puberulous, inside light yellow and glabrescent, with glandular trichomes toward throat on dorsal surface, nectary chamber 4.7–10.2 mm long; throat 9.1–15.1 mm in diameter, outside white and puberulous, inside yellow, ventrally darker and thickened forming a groove, dorsally with glandular trichomes; corolla lobes 5, subequal, white to yellow, orbicular, apex rounded, margin incised to short laciniate, glabrous adaxially, glabrous to strigulose abaxially, ventral lobe larger, straight to spreading, 14.2–23.9 × 12.2–33.4 mm, lateral lobes spreading, 12.1–23.1 × 12.7–25.0 mm, upper lobes, 10.3–22.3 × 10.3–21.6 mm. Androecium of 4 didynamous stamens, included, filaments 29.0–35.0 mm long, adnate to the corolla tube for 5.1–14.0 mm, white, glabrous, coiling after anthesis, staminode absent; anthers oblong, sagittate, coherent by the lateral walls, dehiscence by basal pores that develop into longitudinal slits, 4.2–7.0 × 0.7–2.0 mm. Gynoecium with a single dorsal nectary gland, oval, apex irregularly acute to obtuse, 1.7–2.7 mm long, white, glabrous; ovary superior, 4.2–10.0 × 2.1–6.7 mm, ovate, yellow to yellow-green, puberulous to velutinous; style included, 23.5–35.1 mm long, puberulous to velutinous, reddish at base, white apically; stigma stomatomorphic, white. Fruit a berry, 6.8–17.0 × 5.5–16.5 mm, globose, yellow at maturity. Seeds numerous, 0.8–1.0 × 0.4–0.5 mm, brown, fusiform, longitudinally ridged. Phenology. Collected with flowers throughout the year. Fruits collected in May and June. Etymology. The specific epithet is in reference to the quadrangular to strongly angulate stem cross-section which is occasionally winged (Figs 2 E, 3). Distribution and preliminary conservation assessment. Drymonia quadrangulata has been recorded from the western slopes of the Andes in southwestern Colombia (Cauca and Nariño) and northern Ecuador (Carchi, Esmeraldas, Imbabura, and Pichincha), between 250 and 2300 m (Fig. 4). It grows in the premontane rainforest, in the transition between the Tropical Andes and the Tumbes-Chocó-Magdalena Hotspots (Mittermeier et al. 2004), in a region characterized by high precipitation, which in some areas may reach more than 7000 mm per year (FELCA 2024). In Colombia, this species has been documented in protected areas such as the National Natural Park Munchique, and the Natural Reserves Río Ñambí and La Planada, with few collections in their surroundings. In Ecuador, it has also been found in protected areas such as the Mache-Chindul Ecological Reserve, the Dracula Natural Reserve (Fundación Ecominga), and the Bosque Protector Mashpi (Mashpi Lodge). According to GeoCAT (Bachman et al. 2011), the following values were calculated: EOO = 34,836 km 2 and the AOO = 132 km 2. Based on the IUCN Red List Categories and Criteria (2012) and updated criteria in the IUCN Standards and Petitions Committee (2024), the AOO satisfies criterion B 2 for Endangered (<500 km 2), but the other criteria do not support a threatened category. Given the large EOO and the presence of several populations in protected areas, Drymonia quadrangulata is preliminarily assessed as being of Least Concern (LC). Comments. Drymonia quadrangulata is morphologically similar to D. killipii (Fig. 1), D. chiribogana (Fig. 5), and D. lanceolata (Fig. 6). These taxa possess foliaceous calyx lobes, nearly free, covering at least half of the corolla tube and sometimes the entire tube. The calyx lobe apices are acute to attenuate, with margins folded longitudinally outwards, and tubular-infundibuliform corollas. Drymonia quadrangulata is often collected, but misidentified in most herbaria as D. killipii, a rarely collected narrow endemic in Colombia. However, D. quadrangulata can be distinguished from D. killipii by its strigose indumentum in vegetative and reproductive structures that becomes glabrescent with age (Fig. 2 E) vs. hirsute, becoming glabrescent with age (Fig. 1 C), stem quadrangular to strongly angulate (Figs 2 E, 3), sometimes winged vs. terete (Fig. 1 C), petiole of (2.2 –) 5.2–15.0 cm long vs. 2.0– 6.4 cm long, blade base obtuse to subcordate (Figs 2 F, 3) vs. cuneate (Fig. 1 A), inter-secondary and tertiary venation reticulated (Fig. 2 F) vs. obscure or suppressed (Fig. 1 A), pedicel and calyx lacking glandular trichomes vs. presence of glandular trichomes, calyx lobes lanceolate (Fig. 2 C) vs. ovate (Fig. 1 A), corolla lobes white to yellow (Fig. 2 B – D) vs. royal purple to maroon, suffused with lemon-yellow toward the throat (Fig. 1 B), corolla lobes margin incised to short laciniate (Fig. 2 A – D) vs. subentire (Fig. 1 B), and indehiscent berry fruits (Fig. 2 H) vs. bivalved fleshy capsules. Additionally, Wiehler (1977) noted the fragrant lemon scent of D. killipii, while no scent has been recorded for D. quadrangulata. Drymonia quadrangulata can be distinguished from D. chiribogana (Fig. 5) by its terrestrial habit vs. epiphytic, stem quadrangular to strongly angulate (Fig. 2 E) vs. terete to subquadrangular (Fig. 5 A), stem surface smooth when dried vs. papyraceous, blade green with midvein green adaxially vs. green with veins whitish or silvery, blade glabrescent abaxially vs. with glandular trichomes sunken into the epidermis, inter-secondary and tertiary venation reticulated (Figs 2 F, 3) vs. obscure or suppressed, calyx lobes lanceolate (Fig. 2 C) vs. broadly ovate (Fig. 5 D), corolla of 4.3–6.5 cm long (Fig. 2 A – D) vs. 3.2–4.3 cm long (Fig. 5 D), corolla lobes white to yellow (Fig. 2 B – D) vs. magenta, with red or yellow toward the throat (Fig. 5 B), corolla lobes margin incised to short laciniate (Fig. 2 A – D) vs. subentire to incised (Fig. 5 B, D), and berry fruit (Fig. 2 H) vs. bivalved fleshy capsule (Fig. 5 C). While Drymonia quadrangulata shares a terrestrial habit, quadrangulate to angulate stems, and berry fruits with D. lanceolata (Fig. 6), it can be differentiated by the stem surface smooth when dried vs. papyraceous, blade base obtuse to subcordate (Fig. 2 F) vs. cuneate to attenuate (Fig. 6 A), veins minutely puberulent abaxially vs. strigose to tomentose, calyx lobes nearly free with margins folded longitudinally outwards vs. free with margins flat (Fig. 6 D), corolla lobes incised to short laciniate (Fig. 2 A – D) vs. long-fimbriate (Fig. 6 B, D), and stomatomorphic stigma vs. deeply bilobed. The morphological differences between Drymonia quadrangulata, D. killipii, D. chiribogana, and D. lanceolata are summarized in Table 1. Specimens examined. Colombia – Cauca. • El Tambo; PNN Munchique, camino a López de Micay, entre El Boquerón y La Cueva; Bosque Pluvial premontano; 2.76816667°N, 76.974361°W; 900–1050 m; 14 Jun 2017; fl; L. Clavijo, et al.