Hydrangea trianae Briq., Annuaire Conserv. Jard. Bot. Geneve 20: 403. 1919., Briq., Annuaire Conserv. Jard. Bot. Genève 20: 403. 1919.

Samain, Marie-Stephanie, Granados Mendoza, Carolina & Martinez Salas, Esteban Manuel, 2021, On Hydrangea peruviana, an endangered species from Ecuador, and Hydrangea oerstedii, very common in Costa Rica and Panama, and seven threatened Central and South American Hydrangeas, which have been confounded with these, PhytoKeys 171, pp. 91-153: 91

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Hydrangea trianae Briq., Annuaire Conserv. Jard. Bot. Geneve 20: 403. 1919.


8. Hydrangea trianae Briq., Annuaire Conserv. Jard. Bot. Geneve 20: 403. 1919.   Figures 13 View Figure 13 , 14 View Figure 14

H. lehmannii   Engl., Nat. Pflanzenfam. (ed. 2) 18a: 207. 1930. Type. Colombia. Valle del Cauca: Cali, 4 Oct. 1900, ♀, fruits, F.C. Lehmann 9074 (lectotype, designated by Freire-Fierro 2004, pg. 33, second-step designated here: K! [K000486136]; isolectotypes: B [destroyed, F neg. 4144], K! [K000486135])

Hydrangea platyphylla   Briq., Annuaire Conserv. Jard. Bot. Genève 20: 401. 1919. Type. Colombia. Tolima: Mariquita, ♂, old flowers, J.J. Linden 894 (lectotype, designated by McClintock 1957, pg. 238 [as “holotype”], second-step designated here: G! [G00439775, F neg. 27369]; isotypes: F [V0066625F]!, G! [G00439776, G00223929], GENT! [BR0000005119332, BR0000005119660] (photos of both on one sheet in COL!, GH!, K!, NY!), K! [K000486132]!


Colombia. Tolima: Quindío, ♂, flowers, J.J. Triana s.n. (holotype: G-DC n.v. (photo in GENT!), isotypes: BM!, BR [BR0000005119011, BR0000005119974, BR0000005119646]!., COL [COL000369449] !, F! [V0066627F], MEXU! [acc. # 14858], MICH!)


Root-climbing liana of up to 20 m high, functionally dioecious; free-growing branches many-ribbed, slightly angular, old branches quadrangular, with erect, white stellate hairs (Fig. 13A View Figure 13 ); leaves decussate, petiole terete, sometimes abaxially flattened, color dark green, sometimes drying black, pubescent with small, erect, whitish to yellowish stellate hairs, 1.5-4 cm long, leaving a semicircular scar on the branch when leaves shed; lamina spoon-shaped, obovate to elliptic, (slightly) asymmetric, coriaceous, 15-29 cm long, 8.5-18 cm broad, with varying length-width ratios, base cuneate, acute or decurrent, apex rounded to acuminate, leaf margin (slightly) dentate, venation brochidodromous, veins 8-9 pairs, adaxial leaf side with protruding midvein and primary veins, secondary veins slightly protruding, primary veins join to form submarginal vein, dark green, slightly pubescent with white stellate hairs, especially near the margin, abaxially with protruding veins, midvein notoriously protruding, opaque olive green, pubescent with stellate white hairs, primary veins branching towards the margin, ending in a submarginal vein very near the margin, secondary and tertiary veins forming a reticulate network, connecting the primary veins, acarodomatia numerous, consisting of an often small simple cavity in axils of midvein and primary veins, sometimes covered by a membrane; inflorescence axis in the axils of kataphylls of 1-1.4 cm long or fully grown leaves, pubescent with erect, white, stellate hairs (Fig. 13C View Figure 13 ), 6-25 cm long, many-ribbed, with 2 opposite rapidly shedding leaf or kataphyll pairs along the axis, petiole of the leaves 5-10 mm long, lamina obovate to lanceolate, (slightly) asymmetric, 2-7 cm long, 1.4-4.3 cm broad, margin glandular dentate, small, appressed, stellate, abaxially and adaxially with white hairs along the midveins, similar pubescence between the veins but not appressed and larger, veins 6 pairs, all protruding, in the case of kataphylls, these initially protecting the small inflorescence buds (Fig. 13A View Figure 13 ), followed by elongation of the axis occurring between the kataphylls and the inflorescence bud, apex of the inflorescence axis woody, quadrangular, elongated bract scars visible, thickening at the top, 7-8 mm broad, 2 mm high in functionally female plants, 6-7 mm broad, 2-3 mm high in functionally male plants, inflorescence bracts cucullate, dark green, coriaceous, abaxially (densely) pubescent with small, erect, whitish and yellowish stellate hairs, inflorescences lateral, opposite, 1-2 pairs of inflorescences per flowering branch, flowering branch continues growing rapidly during inflorescence development, with up to three decussate leaf pairs above the inflorescences, with appressed, whitish and yellowish stellate hairs, dense when young, rapidly shedding, inflorescence axes with basal lignified parts of inflorescences of previous years visible in well-collected specimens, allowing to observe growth and flower periodicity, these rests 2.5-5 cm apart, 7 cm when inflorescence originate in the axils of the regular leaves, with none or one decussate leaf pair (or the scars of these leaves) in between, inflorescence umbellate, buds up to 4 cm broad and 4.5 cm high before opening, in flowering stage 7-19 cm diameter, 4-12 cm high, with 5-9 main axes in functionally male plants, 4-8 main axes in functionally female plants, partial inflorescences dichasia, secondary and tertiary inflorescence axes with whitish and/or reddish stellate hairs; enlarged marginal flowers generally present (but e.g. absent in Samain et al. 2011-064) (Fig. 13B View Figure 13 ), terminally placed in a cyme, 1.8-4 cm diameter, sepals with marked palmate veins, fully separated or fused at the base, sometimes one or two reduced or even absent, pistils very rudimentary, pedicel 1-3.5 cm long; flower pedicel of reduced flowers 1-2 mm long in functionally male flowers, 1-5 mm long in functionally female flowers, receptacle semiglobose in functionally male flowers, broadly semiglobose in functionally female flowers, ovary inferior, calyx lobes 4, triangular, translucid, 0.2-0.5 mm long, 0.6-1 mm broad, petals 4, adaxially red, abaxially purple with white margin, valvate, cucullate with a small mucron at the apex, coriaceous, 1.5-3 mm long, 1-1.5 mm broad; functionally male flowers (Fig. 13C View Figure 13 ): hypanthium 1.5-2 mm diameter, 1.5 mm high, stamens 8, well-developed, filaments 1.2-2 mm long, anthers 1-1.5 mm long, 0.5-1 mm broad, purple, pistils 2, very reduced, 0.5 mm long, stigmas not penicellate; functionally female flowers (Fig. 13D View Figure 13 ): hypanthium 1.2-2.2 mm diameter, 1.5 mm high, stamens 8, very reduced, filaments 0.2-0.5 mm long, anthers 0.2-0.4 mm long, 0.1-0.2 mm broad, pistils 2(-3), 1.5-2 mm long, broad at the base, narrower in the middle and broadening towards the apex, enlarging up to 3.2 mm during fruit maturation, stigmas apically clavate and shortly penicellate, 0.5 mm long; fruit a semiglobose capsule, apically with a conspicuous border, dark reddish brown, 2.5 mm high, 4-4.5 mm broad above, 3-4 mm diameter, the part around the pistils thickening upwards up to 1 mm during fruit maturation, opening between the two pistils to release seeds, seeds not seen.


Hydrangea trianae   is a widespread species occurring in Colombia, Ecuador and Peru (Fig. 14 View Figure 14 ).


Hydrangea trianae   is known from cloud forest and remnants of this vegetation type at elevation between 400-3680 m. Moreover, it has been noted to occur in disturbed or late secondary forests.


This species has been collected with flowers and fruits throughout the year.


Hydrangea trianae   should not be considered a synonym of H. peruviana   , from which it can be distinguished by the notoriously coriaceous leaves, the abaxial reticulate network of secondary and tertiary veins connecting the primary ones, and the abaxial white pubescence.

Hydrangea lehmannii   was synonymized with H. peruviana   by McClintock (1957), whereas she considered H. platyphylla   as a synonym of H. oerstedii   .

With respect to the type material of Hydrangea lehmannii   , McClintock (1957) cited the K material as an isotype, albeit without designating a specific sheet, and this was also cited by Freire-Fierro (2004).

With respect to the type material of H. platyphylla   , McClintock (1957) cited G material as holotype, which effectively lectotypified the name. However, there are two specimens of this collection in G, therefore requiring second-step lectotypification, which is done here.

One of the F specimens (V0066626F), which supposedly is an isotype of H. platyphylla   , does not belong to this species based on leaf venation, probably due to a labeling error; however, it is not possible to propose any other identification as the material is very limited.

Briquet (1919) distinguished H. trianae   and H. platyphylla   based on leaf shape and venation, as well as size of cymes, sepals of enlarged marginal flowers and styles, which can be contributed to the sex of the single collections of both species he used for his descriptions, the specimens of H. trianae   being functionally female and the ones of H. platyphylla   which are functionally male.

The specimen of Triana 4668 in NY does not correspond with H. trianae   or any of the other species treated here, in contrast with the specimen with the same number in COL. Triana’s numbers do not correspond to field collection numbers as currently used, but instead are generic numbers based on the Endlicher system ( Kirkbride 1982; Aymard 2017).

According to our molecular study, Hydrangea trianae   is closely related to H. goudotii   (Granados Mendoza et al. unpublished results).

Preliminary conservation status.

Although this species has an EOO of nearly 726,300 km2, it is Endangered (B2abiii) according to the IUCN categories and criteria ( IUCN 2012), with an AOO of 372 km2, as well as an extensive reduction in both EOO and AOO as a consequence of habitat destruction and fragmentation.

Additional specimens examined.

Colombia. Antioquia: Argelia, km 17 of road Sonsón-Argelia (2 km past fork in road to Nariño), 2160 m, 3 Oct 1987, ♀, fruits, Zarucchi et al. 6164 (HUA, MO); Frontino, Correg. Nutibara, Nutibara-La Blanquita road, region of Murrí, Alto de Cuevas, 6°45'N, 76°20'W, 1700-1800 m, 19 Apr 1988, ♂, old flowers, Luteyn et al. 12047 (HUA, K, MO); Urrao, Parque Nacional Natural las Orquideas, vereda Calles, 06°32'N, 76°19'W, 1350-1450 m, 5 Dec 1993, ♀, inflorescence buds, flowers, Pipoly et al. 17737 (MO); Jardín, km 20 of road Jardín-Riosucio (dpto. Caldas), ca 15 km SSE of Jardín, Alto de Ventanas, 29 Oct 1988, 2700-2790 m, ♂, flower buds, Zarucchi et al. 6962 (HUA, MO); road between Jardín and Río Sucio, ca 9 km from Jardín, 2300-2400 m, 29 Jan 1989, ♂, flowers, MacDougal & Roldán 3581 (HUA, MO); Frontino, region of Murrí, road between Nutibara and La Blanquita, 19.2 km from centro de Nutibara, 1560 m, 11 Feb 1989, ♂, old flowers, MacDougal et al. 3929 (CR, HUA, MO); Frontino, Corregimiento Nutibara, Región Murí, márgen de río, 1500 m, 11 Jul 1986, ♀, old inflorescences, Acevedo et al. 1260 (HUA, NY, US); along road between Caramanta and Supia (Risaralda), ca 4 mi S of Caramanta, 2030 m, 27 Jan 1990, ♀, flowers, Croat 70038 (MO); Medellín y Guarne, Parque Ecológico Piedras Blancas, laguna de Guarne, 2350 m, 26 Aug 1995, ♂, old inflorescences, Roldán & Medina 2385 (HUA, MO); Parque Ecológico Piedras Blancas, sector Lajas, 2350 m, 23 Sep 1995, ♂, flowers, Fonnegra et al. 5643 (HUA, MO); Parque Nacional Natural "Las Orquideas", sector Calles, margen derecha del río Calles, 1250 m, 30 May 1988, ♀, fruits, Cogollo et al. 3087 (COL, MO); Urrao, Corregimiento Encarnación, sitio El Río, 1 hora de camino del Páramo de Frontino, 3120 m, 7 Apr 1989, inflorescence buds, Callejas et al. 7774 (HUA, NY); same data as preceding, 3000-3150 m, 8 Apr 1989, ♀, flower buds, flowers, Callejas et al. 7835 (HUA, MO); Envigado, Vereda "El Escobero", Costa Occidental Cerro "San Luis", 2100-2250 m, 15 May 1996, ♂, inflorescence bud, flower buds, flowers, Correa & Carona 1073 (COL, HUA, MO); cerca de San José de San Andrés, 1 May 1948, inflorescence buds, floral buds, Correa & Velasquez 45 (US); bosque bajo de la cumbre cerca de Santa Elena, camino entre Medellín y Río Negro, 2300-2500 m, 3 Aug 1945, ♀, fruits, Medina 239 (US); municipio de Caldas, Alto de San Miguel, 2200 m, 8 Jul 1996, ♂, flowers, Monsalve 38 (F); municipio de Peque, 7°2'6.7"N, 75°58'27"W, 2500 m, 19 Nov 1995, ♀, fruits, Benítez et al. 584 (COL); municipio de Peque, vereda Romeral, 6°59'18.4"N, 75°58'02.5"W, 2650 m, 16 Nov 1995, floral buds, Benítez et al. 452 (COL); municipio Caldas, Reserva "Alto de San Miguel", cuenca alta del río Medellín, 6°5'N, 75°38'W, 2100 m, ♂, old flowers, Roldán et al. 2516 (HUA); municipio de Caldas, vereda La Corrala, Finca "La Zarza", Alto del Gallinazo, ca. 2600 m, 4 Feb 1986, inflorescence buds, Albert de Escobar y Stein 6238 (HUA); municipio de Urrao, Finca La Clara, cerca del río Mina y La Clara, 2340 m, 6 Dec 1984, ♀, fruits, Orozco et al. 1782 (COL); camino de Urrao al páramo de Frontino, 2400 m, 19 May 1985, ♂, flower buds, flowers, Cárdenas 128 (HUA); Argelia, vereda La Vibora, antiguo camino de Sonsón a Argelia, 2200-2700 m, 28 Aug 1989, ♂, flowers, Hoyos y Avendaño 1300 (HUA); municipio Ríonegro, vereda Yarumales   , 20-30 km SE de Medellín en la vía a Rionegro, 6°15'N, 75°28'W, 2140-2410 m, 12 Nov 1990, ♀, fruits, Callejas & Roldán 9628 (HUA); municipio de Urrao, Páramo de Frontino, El Río, 3020 m, 6 Jan 1985, ♀, fruits, Londoño et al. 669 (HUA); Páramo de Frontino, Urrao - zona situada entre el 15 y la Esperanza, 2980-3680 m, 18 May 1985, ♀, fruits, Rentería 4082 (HUA): Medellín, corregimiento de Santa Elena, Sector de Piedras Blancas, 6°17'N, 75°32'W, 2400 m, May 1998, ♂, flowers, Cardona y Alzate 551 (HUA); Medellín, corregimiento de Santa Elena, Reserva Natural Montevivo, sector Casapalo, 6°12'48"N, 75°29'32.2"W, 2500 m, 30 Sep 2002, ♀, fruits, David & Idarraga 291 (HUA); municipio de Caldas, vereda La Corrala, Finca "La Zarza", Camino La Zarza - Alto del Gallinazo, ca. 2440 m, 10 Nov 1987, ♂, old flowers, Albert de Escobar et al. 8022 (HUA); municipio de Urrao, Páramo de Frontino, 6°29'N, 76°7'W, 2890-2250 m, 16 May 1987, ♂, old flowers, Estrada et al. 108 (HUA); municipio de Medellín, corregimiento de Santa Elena, vereda El Llano y Perico, cabeceras y recorrido Q. San Pedro, 2550 m, 8 May 1996, ♀, fruits, Giraldo et al. 785 (HUA); Caldas: Vereda la Corrala, Finca La Zarza, al lado del camino entre la cascada y el Alto del Gallinazo, 2550 m, 27 Apr 1987, floral buds, Albert de Escobar 7576 (HUA, UC); Chocó: municipio San José del Palmar, hacia el galápago, 1380 m, 11 Nov 1985, ♀, fruits, Lozano et al. 4920 (COL); Cundinamarca: San Francisco, hacienda "La Laja", 2880 m, 30 May 2004, ♂, flowers, Parra et al. 465 (COL); Huila: municipio de la Argentina, Quebrada del pueblo, 1850 m, 25 Sep 1984, ♀, fruits, Lozano et al. 3981 (COL); Nariño: Mpio. de Ricaurte, Reserva Indígena Gualcalá, Santa Fé, camino al Río Gualcalá, 1°18'N, 77°54'W, 1100-1200 m, 18 Dec 1995, inflorescence buds, floral buds, Ramírez & González 9135 (QCA). Quindío: Parque Nacional de Cocora, Quebrada Santa Lucía, 3000 m, 08 May 1990, ♀, flowers, Franco 3095 (NY); Risaralda: municipio de Pereira, corregimiento La Florida, SFF Otún Quimbaya, 4°44'17"N, 75°34'01"W, 1900 m, 22-28 Feb 2004, floral buds, Alzate et al. 1962 (F); municipio de Pereira, orilla del camino entre Ceylan y el Cedral, 2150 m, 4 Dec 1989, ♀, fruits, Franco et al. 2929 (COL); Mpio. Pereira, Parque Nat. Reg. UCUMARI, 2200-2450 m, ♀, fruits, González 1635 (COL); municipio de Pereira, Parque Regional Ucumari, vereda la Pastora, camino de el Ceilán a El Cedral, borde de la quebrada Miraflores, 2340 m, 30 Jul 1989, floral buds, Galeano 222 (COL); municipio de Apía, vereda "La Cumbre", 2285 m, 24 Feb 1983, ♀, fruits, Torres et al. 2192 (COL); Tolima: Quindio, El Roble, old inflorescence axis, Triana 358 (US); same data as preceding, ♂, flowers, Triana 359 (US); same data as preceding, ♂, flowers, Triana 2805 (US); Valle del Cauca: San Antonio, about 25 km W of Cali, 1800 m, 25 Jan 1988, ♂, flowers, van der Werff & Giraldo 9746 (MEXU, MO); same data as preceding, van der Werff & J. Giraldo 9747 (CR, MO); Cordillera Occidental, vertiente occidental, Hoya del Río Digua, lado izquierdo, Piedra de Moler, 900-1180 m, 19-28 Aug 1943, ♀, fruits, Cuatrecasas 14974 (F); Cali, forests of West Cordillera above Cali, 2000-2200 m, June 1900, ♀, young fruits, B.T. 623 (NY); Finca Zingara, km 18 de la carretera Calia   -Buenaventura, km 4 vía a Dapa, corregimiento de la Elvira, cordillera Occidental, 1900 m, 23 Jan 1994, ♀, flowers, Giraldo-Gensini & Agredo 130 (MO); same data as preceding, 26 Feb 1995, ♀, fruits, Giraldo-Gensini & Agredo 611 (MO); same data as preceding, 25 Aug 1996, ♀, fruits, Giraldo-Gensini & Corrales 765 (MO); Cuenca del Río Cali, cercanías de Peñas Blancas, 10 Jan 1963, sterile, López Figueiras 8139 (US); Cordillera Occidental, vertiente oriental, Hoya del río Cali, río Pichindé, entre los Cárpatos y El Olivo, 2920-2025 m, 5 Aug 1946, inflorescence buds, Cuatrecasas 21935 (F, P, US); Cordillera Occidental, Los Farallones, extremo N, vertiente oriental, Almorzadero, bosques, 2980 m, 13 Oct 1944, ♂, flowers, Cuatrecasas 18113 (F, US); Yumbo, Finca La Samaria, NE of Darien, near Lago Calima (reservoir), 1700-1750 m, 14 Feb 1984, ♀, flowers, Juncosa 2181 (MO); Peñas Blancas, los Cárpatos, 1600-1700 m, 9 Sep 1988, inflorescence buds, Franco et al. 2509 (COL, MO); Bosque de San Antonio, W of Cali, near television tower, 2250 m, 15 Jul 1984, ♀, fruits, Gentry 48079 et al. (MO); Buga, Carretera a El Retiro, a 1 km de carretera Buga-El Placer ( desvío a Retiro está a 12 km antes de El Placer), Cordillera Central, vertiente occidental, al lado de carretera, 2300 m, 15 Sep 1991, ♂, flowers, Silverstone-Sopkin & Giraldo-Gensini 6400 (MO); Restrepo, Río Calima, Cusumbo, 680 m, 21 Feb 1989, ♀, fruits, Devia & Prado 2581 (US); El Silencio, Hacienda Himalaya, Cordillera Occidental W of Yumbo, transect 1, 1860 m, 2 Jan 1989, ♂, flowers, Gentry et al. 65414 (MO); Argelia, Vereda Las Brisas, 2050-2200 m, 21 Jan 1983, ♀, fruits, Díaz P. 3854 (COL, MO); Manizales   , vereda La Esperanza, Reserva Torre Cuatro, cerca de la quebrada La Siberia, 5°1'41"N, 75°23'10"W, 2650-2750 m, 28 Mar 1999, ♂, old flowers, Alvear et al. 314 (COL); Manizales   , 1851-1857, ♂, old flowers, Triana s.n. (K); Neira, Cementos Caldas, 5°9'45"N, 75°29'43.1"W, 2203 m, 29 Jul 2004, ♀, fruits, Mancera 553 (COL); Manizales   , Reserva Río Blanco, 2600 m, 28 Feb 2003, sterile, Sanin 39 (HUA); Manizales   , Río Blanco, El Paso nivel, 2500 m, 10 Jul 2003, sterile, Sanin 958 (HUA). Cauca, Parque Nacional Munchique, El Tambo, vereda La Romelia, la Gallera, 1950 m, 25 Jul 1993, ♀, fruits, Velayos et al. 6891 (COL); El Tambo, Parque Nacional Munchique, vereda La Romelia, la Gallera, 2385 m, 26 Jul 1993, ♂, old flowers, Velayos et al. 6917 (COL). With three localities: Quindio: Mariquita, Roble, 2080 m; Antioquia, Manizales   , 2140 m: Buenaventura, Cordillera Occidental, 2190 m, Jul 1833, ♂, old flowers, Triana 4668 (COL); same data as preceding, floral buds, Triana 1866 (photo in F of a specimen in G).

Ecuador. Carchi: prominent hillcrest directly N of Lita, on N side of Río Mira and just to E of Río Baboso, on steep W-facing slope, 760 m, 7 Aug 1994, sterile, Boyle 3476 (MO); along Chical-Pailon-San Marcos trail, 01°06'N, 78°18'W, 600-1200 m, 18-19 Nov 1983, ♂, flowers, Kvist et al. 48670 (AAU, QCA, QCNE); El Pailon, ca. 45 km above Maldonado along a footpath to Tobar Donoso, 0°49'S, 78°9'W, 800 m, 1 Dec 1979, ♀, fruits, Madison & Besse 7226 (QCA); Esmeraldas: Quininde, Bilsa Biological Station, Montañas de Mache, 35 km W of Quininde, 5 km W of Santa Isabel, slope between banana plantation and stream SE of Station, 400-600 m, 9 Dec 1994, ♀, fruits, Bass & Pitman 320 (MEXU, MO, QCNE); same data as preceding, 31 Dec. 1994, ♀, flowers, Pitman & Marsh 1133 (MO, QCNE); San Lorenzo, Alto Tambo, Frente Finca del Sr. Lalama, a 1.5 km del sector de El Cristal, 650 m, 13 May 1992, ♂, flowers, Quelal & Luteyn 489 (MO, QCNE); Imbabura: Cordillera Occidental, La Union, lower Intag Valley, 4600 ft, 20 Sep 1944, ♀, flowers, Drew E-684 (NY, US); Los Ríos: Quevedo, Centinela-La Pirámide, vía Santa Domingo de los Colorados-Quevedo entrando por Patricia Pilar km 41, 650 m, 25 Feb 1992, ♂, flowers, Quelal & Tipaz 347 (MO, QCNE); Collections from path following ridge line at El Centinela at crest of Montañas de Ila on road from Patricia Pilar to 24 de Mayo at km 12, Patricia Pilar is at Km 45 on road from Sto. Domingo to Quevedo, 600 m, 4 Feb 1983, ♀, fruits, Dodson 13654 (MO, QCNE); same data as preceding, 6 Apr 1980, inflorescence bud, Dodson s.n. (MO); Napo: Quijos, along road 1 km SE of Cosanga, 00°35'S, 77°52'W, 2000 m, 31 Jul 1990, floral buds, Webster & Richerson 28512 (QCNE); El Chaco, San Juan Chico, camino hacia el Río Oyacachi, 00°17'S, 77°52'W, 2000 m, ♀, fruits, Alvarez 97 et al. (QCNE); Baeza-Sardinas Alto, col. en borde de carretero, al NO de Sardinas Alto, a 1.5 km, 00°22'S, 77°52'W, 1780 m, 15 Dec 1993, inflorescence buds, Freire-Fierro & Yánez 2693 (QCA, QCNE); hills above R. San Juan, at confluence with R. Oyacachi, ca 10 km W of El Chaco, 00°17'S, 77°51'W, 1800-2000 m, ♀, fruits, Ståhl et al. 2354 (QCA); Napo-Pastaza, Mera, 26 Mar 1940, ♀, fruits, Lugo 126 (C, MO); Pichincha: along road and trail from Maquipucuna Lodge to Ecolodge Santa Lucia, 2 km N of Maquipucuna entrance, 1400 m, 15 Mar 2006, ♀, fruits, Croat 95938 (MO); Maquipucuna, 5 km E of Nanegal, Transect # 5, 1550 m, 10 Feb 1991, ♀, flowers, Gentry & Valencia 73174 (AAU, MO, QCNE); Quito, Nanegal, Bosque protector Maquipucuna, 5 km airline SE of Nanegal, disturbed rainforest above Río Tulambi, 00°07.5'N, 78°38.5'W, 1500 m, 31 Aug 1993, ♂, old flowers, Webster et al. 30003 (QCNE); Reserva Biológica Maquipucuna, Nanegal, colectada en primer lindero, camino del Inca, 1600 m, 9 Jan 1992, ♂, flowers, Tipaz et al. 593 (AAU, MO, QCNE); Maquipucuna Tropical Reserve, western slopes of Andes, northern boundary of reserve, 10 km north of Nanegalito, 1200 m, 2 Dec 1988, inflorescences, floral buds, Neill et al. 8643 (MO, QCNE); Reserva Biológica Maquipucuna, 1300 m, 31 Jan 1991, ♀, flowers, Alvarado 406 (COL, MO, QCNE); along road and trail from Maquipucuna Lodge to Ecolodge Santa Lucia, 2 km N of Maquipucuna entrance, 00°07'19"N, 78°37'06"W, 1400 m, 15 Mar 2006, ♀, fruits, Croat et al. 95939 (MO, QCNE); Mindo, on western slopes of Andes, 1200 m, 16 Apr 1994, ♂, flowers, Neill & Asanza 10349 (MO, QCNE); 11 km W of Tandapi, trail along Chictoa River, tributary of Río Pilatón, 1350-1550 m, 26 Oct 1974, ♀, fruits, Gentry et al. 12073 (MO, QCA); Tungurahua: Baños, 2.8 airline km SSW of Río Negro, near El Coral, 1481 m, 1°25'59.5"S, 78°12'1.5"W, 13 Jun 2012, sterile, Granados Mendoza et al. 2012-38 ( GENT, HOXA, IEB, MEXU, QCNE); ca. 2 km E of Río Verde on touristic path before tunnel on road to Baños-Mera, 1629 m, 1°23'41.5"S, 78°16'53.2"W, 16 Jun 2012, sterile, Granados Mendoza et al. 2012-44 ( GENT, HOXA, IEB, MEXU, QCNE); Zamora-Chinchipe, Zamora, Jamboe Bajo, Eastern border of Podocarpus National Park, 1100 m, 5 Nov 1996, sterile, Clark 3283 (MO); Westkordillere, 1000 m, 28 Sep 1934, sterile, Heinrichs 741 (NY).

Peru. Amazonas: Bagua, Aramango, 8.86 km SE of Aramango, 1.23 km SW of Nueva Esperanza, 1806 m, 5°28'28.9"S, 78°23'08.4"W, 3 Jul 2011, sterile, Samain et al. 2011-110 ( GENT, IEB, MEXU, USM); Chachapoyas, Soloco, Paraje las Tinas, 5.26 km SE of Quitachi, E of road to Pampa del Tío, 2108 m, 6°21'06.45"S, 77°38'52.95"W, 22 Jun 2011, ♂, flowers, Samain et al. 2011-064 ( IEB, GENT, MEXU, USM); Cajamarca: San Ignacio, San José de Lourdes, localidad Estrella del Oriente, 1630 m, 2 Sep 1997, ♀, fruits, Campos 4326 & Rodriguez (F, MO, NY); Huánuco: Chinchao, 0.86 km ESE of Carpish, 330 m SE of road Huánuco-Tingo María, 2681 m, 9°42'24.3"S, 76°05'11.2"W, 17 Jul 2011, sterile, Samain et al. 2011-185 ( GENT, IEB, MEXU, USM); Chinchao, Carpish, 2626 m, 9°42'21.29"S, 76°05'14.46"W, 29 Nov 2013, ♂, inflorescence buds, flowers, Samain et al. 2013-109 ( IEB, GENT, MEXU, USM); same data as preceding, 9°42'20.41"S, 76°05'15.79"W, 2610 m, 29 Nov 2013, ♀, fruits, Samain et al. 2013-112 ( IEB, GENT, MEXU, USM); 0.85 km ESE of Carpish, 330 m SE of road Huánuco-Tingo María, 2700 m, 9°42'24.5"S, 76°05'11.6"W, 17 Jul 2011, sterile, old inflorescence axes, Samain et al. 2011-186 ( GENT, IEB, MEXU, USM); Pasco: Oxapampa, Valle de San Alberto, east of Oxapampa, above hydroelectric plant, at base of Cordillera Yanachaga, 1900 m, 1 Jan 1984, ♂, flowers, Foster 7703 et al. (F, MEXU, NY, USM); Oxapampa, Huancabamba, Sector Tunqui, Parque Nacional Yanachaga Chemillén, camino al valle del Palcazú, 10°16'24"S, 75°30'37"W, 2006 m, 15 Sep 2009, floral buds, Castillo et al. 1006 (HOXA); Oxapampa, Huancabamba, Parque Nacional Yanachaga Chemillen, parte media de la Quebrada Muchuy Mayo, sector Tunqui, 10°17'30"S, 75°31'05"W, 1800 m, 26 Oct 2007, functionally male, inflorescence buds, young flowers, Monteagudo et al. 15627 (HOXA); Oxapampa, sector San Alberto, 10°32'58"S, 75°21'58"W, 2353 m, 23 Dec 2007, ♀, fruits, Rojas et al. 5052 (HOXA); San Martín: Moyobamba, Jepelacio, Cascada Paccha, 1.15 km SW of San Miguel, road Jepelacio-Carrizal, 6°13'19.5"S, 76°55'14.9"W, 1229 m, 28 Jun 2011, ♀, fruits, Samain et al. 2011-092 ( IEB, GENT, MEXU, USM).