Drymonia clavijoiae J. L. Clark, 2025

Drymonia clavijoiae J. L. Clark sp. nov.

Figs 1 View Figure 1 , 2 View Figure 2

Diagnosis.

Differs from all other congeners by the presence of a conspicuous gelatinous residue often covering the inflorescence and flowers. The elongated tubular and laterally compressed flowers in Drymonia clavijoiae are similar to those in D. coccinea , but D. clavijoiae is also distinguished by an elongated inflorescence axis that often exceed 20 cm in length (versus consistently less than 10 cm in D. coccinea ). Additionally, the inflorescences in D. coccinea lack a gelatinous residue, which is a defining feature of D. clavijoiae .

Type.

Ecuador • Zamora-Chinchipe: parroquia Los Encuentros, Estación Experimental El Padmi (Universidad de Loja), northern outskirts of El Padmi, 5 km from the town of Los Encuentros , 3°43'S, 78°39'W, 850–915 m, 3 June 2007, J. L. Clark & Gesneriad Research Expedition Participants 9943 (holotype: US-2 sheets! [barcodes US 00961563 & US 00961562 ]; isotypes: AAU, COL, MO, NY, QCNE, SEL! [barcode SEL 065837 ]) GoogleMaps .

Description.

Elongate scandent subwoody nomadic lianas with leaves in the subcanopy (ca. 10 m above ground) and flowers produced near the forest floor along a leafless portion of the stem. Stems elongate and subwoody, terete in cross section, 4–9 mm in diameter. Leaves opposite, equal in a pair; petiole 1–5 cm long, green, terete in cross-section; blade oblong to ovate, 8–22 × 2–9 cm, coriaceous, adaxially and abaxially green, apex acute to acuminate, base acute, margin entire, 5–7 pairs of secondary veins, sparsely pubescent with single-celled trichomes abaxially, adaxially glabrous. Inflorescences covered in a gelatinous residue, produced along a leafless region of stem near ground, of a pair-flowered cyme that elongates from displaced bracteoles, often reaching 30 cm in length, each inflorescence branch subtended by a pair of persistent bracts; each bract uniformly puberulent, oval and red, ca. 1.5 × 1.5 cm; inflorescence with one mature flower open at a time. Flowers tubular and laterally compressed; pedicels 5–8 mm long. Calyx white to white suffused with red, uniformly puberulent on outside and glabrous on inside, lobes 5, nearly free, fused at the base for 2–4 mm, overlapping, imbricate, and clasping corolla tube, each broadly ovate, apex rounded, base broadly ovate, margins entire, ventral and lower lobes ca. 2.5 × 1.8 cm, the dorsal lobe slightly smaller, ca. 1.4 × 1.3 cm. Corolla tube zygomorphic, protandrous, oblique relative to calyx, 4–5 cm long, gibbous at base, constricted laterally throughout, 1.0– 1.3 cm wide, outside uniformly puberulent, inside mostly glabrous with minute glandular trichomes in the upper region of the throat, throat elliptic in cross section and nearly constricted laterally, lobes 5, subequal, margins entire to serrulate, lobes reflexed, 8–11 × 9–12 mm, upper lobes always yellow, lower and lateral lobes yellow with brown spots or uniformly yellow. Androecium of 4 didynamous stamens, included, filaments broad and flat, ca. 3.5 cm long, adnate to the corolla tube base for 4 mm, white, glabrous; anthers oblong, sagittate, coherent by the lateral walls, initially dehiscent by basal pores that later develop into longitudinal slits, 4.2–6.0 × 0.7–2.0 mm. Gynoecium with a single bilobed dorsal gland; ovary superior, 4.0–5.0 × 4.0–5.0 mm, cone-shaped, puberulent; style stout, included, 3.2 cm long; stigma stomatomorphic. Fruit a bivalved fleshy capsule, valves light red and reflexed when mature, each valve 1.3 × 1.3 cm. Seeds numerous, 0.8–10.0 × 0.4–0.5 mm, light brown, fusiform, ridged.

Additional specimens examined.

Colombia. • Cauca: cantón Santa Rosa, carretera Macoa-Pitalito , 928 m, 1°20'25.45"N, 76°32'12.51"W, 2 Aug 2024, J. L. Clark et al. 19102 ( COL, HEEA, SEL) GoogleMaps . Ecuador. • Morona-Santiago: cantón Limón Indanza, Cordillera del Condor, trail towards crest of the Cordillera del Condor from camp # 1 (10–15 km S / SE from the comunidad Warints) , 830–1200 m, 3°13'S, 78°15'W, 17 Dec 2002, J. L. Clark 7058 ( QCNE, US) GoogleMaps ; • cantón San Juan Bosco, road between San Juan Bosco and El Pangui, 27 km south of San Juan Bosco , 1591 m, 3°17'51"S, 78°33'27"W, 2 Jun 2007, J. L. Clark et al. 9904 ( ECUAMZ, US) GoogleMaps ; • road Limón to Mendez , Aug 1989, A. Hirtz & X. Hirtz 4413 ( SEL) ; • Cordillera de Cutucú, western slopes, along a trail from Lograño to Yaupi , 700 m, Nov 1976, M. T. Madison et al. 3151 et al. ( SEL, US) ; • Macas, along new road, west into the Andes, first 17 km westward, then ca. 12 km south on side road , 13 Apr 1988, H. Wiehler et al. 8805 ( SEL, US) ; • Cordillera de Cutucú, between Sucua and Patuca , 17 Apr 1988, H. Wiehler et al. 8858 ( SEL) . • Napo: cantón Archidona, Reserva Ecológica Antisana, comunidad Shamato, entrada por km 21 – Shamato, camino Sardinas – Shamato , 1700 m, 0°44'S, 77°48'W, 27 Apr 1998, J. L. Clark, E. Narváez & G. Alvarado 5232 ( QCNE, US) GoogleMaps ; • cantón Archidona, parroquia Catundo, buffer zone of Parque Nacional Sumaco Napo Galeras, comunidad Mushullakta , 900–970 m, 0°47'49"S, 77°35'10"W, 24 Feb 2003, J. L. Clark, N. Harris & L. Narváez 7206 ( F, K, MO, QCA, QCNE, SEL, US) GoogleMaps ; • Tiputini Biodiversity Station (Universidad San Francisco, Quito), Yasuní Biosphere Reserve, near Río Tiputini , 200 m, 0°38'11"S, 76°8'58"W, 24 Jun 2006, J. L. Clark, L. Bohs & I. Nenquimo 9473 ( ECUAMZ, MO, NY, SEL, US) GoogleMaps ; • Yasuní Biosphere Reserve, Tiputini Biodiversity Station (Universidad San Francisco, Quito), sendero Chichico to sendero Parahuaco , 200 m, 0°38'11"S, 76°8'58"W, 17 May 2007, J. L. Clark et al. 9571 ( ECUAMZ, MO, NY, SEL, US) GoogleMaps ; • Yasuní Biosphere Reserve, Tiputini Biodiversity Station (Universidad San Francisco, Quito), sendero Maquisapa to sendero Harpía , 200–250 m, 0°38'11"S, 76°8'58"W, 22 May 2008, J. L. Clark et al. 10233 ( ECUAMZ, SEL, US) GoogleMaps ; • cantón Archidona, parroquia Catundo, buffer zone of Parque Nacional Sumaco Napo Galeras, trail from the Comunidad Mushullakta to Lluya Patcha (campamento # 1) towards summit of Galeras , 1100–1200 m, 0°49'40"S, 77°33'49"W, 11 May 2011, J. L. Clark et al. 12055 ( ECUAMZ, SEL, US) GoogleMaps ; • cantón Tena, Jatun Sacha Biological Station (Fundacion Jatun Sacha), forest between main cabin and Río Napo, 23 km east of Puerto Napo, 8 km east of Misahuallí , 485 m, 1°4'12.24"S, 77°37'46.46"W, J. L. Clark 17485 ( ECUAMZ, SEL, US) GoogleMaps ; • La Joya de los Sachas, comunidad de Pompeya, lado sur del Río Napo, km 2 carretera de Maxus , 230 m, 0°28'S, 76°40'W, 26 Aug 1993, M. Aulestia 356 ( MO, QCNE, SEL) GoogleMaps ; • 6 km south of Coca, towards Auca oil fields , Jul 1982, L. Besse, H. Kennedy & R. Baker 1568 ( SEL, US) ; • Coca, road to Las Aucas , 23 Jul 1977, J. D. Boeke 2217 ( NY, SEL) ; • Río Jivino, Limoncocha , 13 Mar 1968, G. Harling, G. Storm & B. Strom 7631 ( SEL) ; • Cañon de los Monos, ca. 12 km north of Coca (Puerto San Francisco de Orellana) , 250 m, 4 Feb 1974, G. Harling & L. Andersson 11725 ( SEL) ; • Coca (Puerto San Francisco de Orellana), trail along Río Payamino, rastrojos , 250 m, 12 Feb 1974, G. Harling & L. Andersson 11927 ( SEL) ; • Río Sumino, tributary of the Río Napo, ca. 5 km northeast of Santa Rosa , 8 Sep 1968, H. Lugo S. 204 ( SEL, US) ; • Hongota at Río Mishuallí, ca 6 km east of Tena , 3 Apr 1969, H. Lugo S. 1007 ( SEL) ; • Santa Rosa at Río Napo , 27 Arp 1972, H. Lugo S. 1950 ( SEL) ; • Lago Agrio , 4 Feb 1973, H. Lugo S. 3158 ( SEL) ; • Coca (Puerto Francisco de Orellana) , 17 Jan 1973, H. Lugo S. 2791 ( SEL) ; • environs of Limoncocha , 250 m, 16 Jun 1978, M. T. Madison, T. C. Plowman & L. Besse 5331 ( SEL) ; • Río Aguarico, environs of Dureno, downriver from Lago Agrio , 457 m, 1 Aug 1974, T. Plowman, C. Sheviak & E. W. Davis 4035 ( SEL, US) ; • along road from Napo to Puyo on way to Hacienda Dos Ríos, just outside Tena, above Mission Evangelica, near Río Tena , 3 Aug 1971, H. Wiehler 71116 ( SEL) ; • woods southeast of Tena , 4 Aug 1971, H. Wiehler 71126 ( SEL, US) ; • along road Hollin towards Loreto , 25 Apr 1983, H. Wiehler et al. 93187 ( SEL) . • Pastaza: cantón Pastaza, parroquia Simón Bolívar, km 38 on the Puyo-Macas highway, near turn off towards Palora , 1000 m, 1°42'6"S, 77°50'36"W, 25 Jun 2003, J. L. Clark & J. Katzenstein 8326 ( QCNE, US) GoogleMaps ; • cantón Mera, Sumak Kawsay In Situ reserve, ecological corridor Llanganates-Sangay, Río Anzu watershed, trail to parcela # 2 , 1211–1459 m, 1°24'25.91"S, 78°4'12.67"W, 9 Jan 2024, J. L. Clark 17835 ( QCA, SEL) GoogleMaps ; • parroquia Veracruz, finca de Ursula Gelchsheimer, between Puyo and Macas, via a Taculín, entrada al Calvario, Río Bobonaza watershed , 609 m, 1°31'4.4"S, 77°50'21.03"W, 12 Jan 2024, J. L. Clark, H. X. Garzón-Suárez 17882 ( QCA, SEL) GoogleMaps ; • cantón Mera, Río Anzu, 10–15 km north of Shell via dirt road , 1211 m, 1°24'39.75"S, 78°3'12.35"W, 14 Jan 2024, J. L. Clark, L. Jost & H. X. Garzón-Suárez 17912 ( QCA, SEL) GoogleMaps ; • Puerto Sarayacu, at Río Bobonaza , 20 Mar 1971, H. Lugo S. 1718 ( SEL) ; • along road from Puyo to Macas, about 19 km outside of Puyo , near pond, 30 Apr 1979, H. Wiehler et al. 79195 ( SEL) ; • Río Anzu, road Mera-Río Anzu , 1000 m, 12 Feb 1990, A. Hirtz 4586 ( SEL) . • Sucumbíos: Sacha Lodge, 3 km NW of the village Añangu, near the Napo river , 200 m, 0°30'S, 76°26'W, 8 Jun 1995, J. L. Clark, L. Demattia & T. Miller 1060 ( QCNE, US) GoogleMaps ; • San Rafael , 25 Apr 1998, H. Wiehler et al. 98150 ( SEL) . • Tungurahua: cantón Baños, parroquia Río Negro, patch of forest near Baños-Puyo road, western side of Río Topo , 1200 m, 1°24'16"S, 78°11'17"W, 27 Jun 2003, J. L. Clark & J. Katzenstein 8402 ( QCNE, US) GoogleMaps . • Zamora-Chinchipe: cantón Zamora, Jambo Bajo, eastern border of Podocarpus National Park, Fundación Maquipucuna, permanent plot 1, sector Nororiental, property of Sr. Jorge Acacho , 1100 m, 4°5'S, 78°55'W, 4 Nov 1996, J. L. Clark, P. Conza, P. Walter & M. Zapata 3204 ( MO, QCNE, US) GoogleMaps ; • cantón Nangaritza, parroquia Zurmi, comunidad Centro Shaime (along Río Nangaritza), forest 2–4 km NW of Centro Shaime , forest on limestone outcrop, 1000 m, 4°18'6.3"S, 78°41'1.9"W, 14 Dec 2001, J. L. Clark, A, Lucia, M. Terry & R. Chuinda 6492 ( QCA, QCNE, SEL, UNA, US) GoogleMaps ; • cantón Nangaritza, parroquia Zurmi, comunidad Centro Shaime (along Río Nangaritza), forest 2–4 km NW of Centro Shaime , 1000 m, 4°18'6.3"S, 78°41'1.9"W, 14 Dec 2001, J. L. Clark, A, Lucia, M. Terry & R. Chuinda 6498 ( QCA, QCNE, US) GoogleMaps ; • cantón Zamora, buffer zone near the eastern border of Parque Nacional Podocarpus, near entrance of Copalinga Ecolodge , 955 m, 4°5'34"S, 78°57'38"W, 4 Jun 2007, J. L. Clark et al. 9980 ( ECUAMZ, US) GoogleMaps ; • south of Yanzatza 850 m, 3 Feb 1987, C. Luer, J. Luer & A. Hirtz 12612 ( SEL) . Peru. • San Martín: cantón Rioja, bosque Proteción Alto Mayo (BPAM), near Puente Aguas Verdes, confluence of Rios Aguas Verdes and Serranoyacu, km 397 on Highway 5 N, carretera Fernando Belaunde Terry , 1170 m, 5°39'57"S, 77°44'54"W, 5 Jun 2010, J. L. Clark et al. 11884 ( SEL, US, USM) GoogleMaps ; • cantón Rioja, Bosque de Proteción Alto Mayo (BPAM), near Puente Aguas Verdes, confluence of Rios Aguas Verdes and Serranoyacu, KM 397 on Highway 5 N, carretera Fernando Belaunde Terry , 1170 m, 5°39'57"S, 77°44'54"W, 7 Jun 2010, J. L. Clark et al. 11922 ( SEL, US, USM) GoogleMaps ; • cantón Rioja, distrito Pardo de Miguel, Aguas Verdes , 1234 m, 5°41'4.69"S, 77°39'28.25"W, 3 Jun 2024, J. L. Clark, J. Flores, L. Valenquela & R. Rojas 18803 ( HOXA) GoogleMaps .

Phenology.

Collected with flowers throughout the year. Collected with fruits in June.

Etymology.

The specific epithet honors Dr. Laura Vibiana Clavijo Romero, a preeminent botanist whose doctoral dissertation significantly advanced our understanding of Drymonia systematics. Clavijo’s extensive, collections-based research has greatly enriched our knowledge of Andean plant diversity, with a particular focus on Colombia and members of the flowering plant family Gesneriaceae . Currently serving as the director of the National Herbarium of Colombia ( COL) and as faculty member at the Universidad Nacional de Colombia - Sede Bogotá, Dr. Clavijo has established herself as a leading authority on Drymonia . Her name has become synonymous with the study of this genus, and the specific epithet commemorates her invaluable contributions to the systematics and taxonomy of this group.

Distribution.

Drymonia clavijoiae is the most widespread of the four species described here (Table 1 View Table 1 ). It is relatively common on the eastern slopes of the Andes, especially in Ecuador, southern Colombia, and northern Peru in premontane wet forests. It grows in the shade of forest edges and is often observed along roads in secondary and primary forests.

Comments.

Among the four species described, Drymonia clavijoiae is the most widespread and most frequently collected. Its corolla tube is consistently tubular, yellow, and laterally compressed (Figs 1 View Figure 1 , 2 View Figure 2 ). Most corolla lobes are yellow with brown spots (Fig. 1 A View Figure 1 ), although some are uniformly yellow and spot-free (Fig. 1 B View Figure 1 ). This species is characterized by clustered bracts, elongate inflorescence axes, and the presence of a gelatinous residue covering the flowers and inflorescences.

Drymonia clavijoiae resembles D. coccinea , a species with an Amazonian distribution that includes the Amazon Basin and the Guiana Shield. The type locality for D. coccinea is French Guiana, based on a collection by Jean Baptiste Christophe Fusée Aublet (Aublet s. n., BM [barcode 000645637]) from Cayenne ( Aublet 1775). Specimens from Amazonian regions (including the Guiana Shield) often exhibit compact inflorescences on the same stem region as the foliage. In contrast, D. clavijoiae typically has inflorescences and fruits near the ground (Fig. 1 E View Figure 1 ), while its foliage is positioned in the subcanopy (Fig. 1 B View Figure 1 ). Occasionally, both inflorescences and foliage are adjacent, but it is more common to find the flowers and fruits on a region of the scandent shoot near the forest floor and the leaves in the subcanopy. A tissue sample of D. clavijoiae (J. L. Clark 6492) was initially identified as D. coccinea and included in a phylogenetic analysis where it was strongly supported with other members of Drymonia that shared laterally compressed corollas (fig. 3 in Clark et al. 2015).

Documenting Drymonia clavijoiae presents significant challenges due to the gelatinous residue or secretions covering its inflorescences. This sticky gelatinous exudate causes floral parts to adhere to newspaper during specimen preparation, often resulting in a hardened mass of compressed floral material and paper. Preparing the type specimen (J. L. Clark et al. 9943) required meticulous care, including multiple changes of newspaper throughout the drying process. Some specimens of this species were labeled with the herbarium epithet “ umecta, ” acknowledging the gelatinous residue and its medicinal use by natives of the region ( Wiehler 1995).

The ecological significance of the gelatinous exudate in D. clavijoiae is unknown, but the presence of glandular structures and the production of secretory products has been the topic of several studies. Structural and functional roles of colleters or multicellular secretory structures were reviewed across 60 flowering plant families, where their presence was investigated for ecological roles in pollination, protection from pathogens and herbivores, or by reducing transpiration ( Thomas 1991). It is noteworthy that Gesneriaceae was not listed among the 60 families by Thomas (1991) as having colleters.

Secretions on inflorescence bracts are well-known in members of the Bromeliaceae ( Benzing 2000) . A recent study by Ballego-Campos et al. (2023) investigated secretions, with an emphasis on trichomes in the floral bracts of 52 species of Tillandsioideae ( Bromeliaceae ). Other studies of secretions in Bromeliaceae have suggested that the gelatinous exudate acts as a sticky trap to protect against herbivory by insects, thus reducing damage to floral structures ( Monteiro and Macedo 2014). It is likely that the gelatinous exudate in D. clavijoiae could play a similar ecological role in reducing herbivory, but further studies are needed.