Hydrangea weberbaueri Engl., Nat. Pflanzenfam. (ed. 2) 18a: 207. 1930., Engl., Nat. Pflanzenfam. (ed. 2) 18 a: 207. 1930.

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

publication ID

http://dx.doi.org/10.3897/phytokeys.171.56351

persistent identifier

http://treatment.plazi.org/id/4FA144A1-D935-53C7-A2CF-8074F5035EF7

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scientific name

Hydrangea weberbaueri Engl., Nat. Pflanzenfam. (ed. 2) 18a: 207. 1930.
status

 

9. Hydrangea weberbaueri Engl., Nat. Pflanzenfam. (ed. 2) 18a: 207. 1930.   Figures 14 View Figure 14 , 15 View Figure 15 , 16 View Figure 16

Type.

Peru. Amazonas: Cheto, ♀, inflorescence bud, flower buds, flowers, A. Weberbauer 4372 (holotype: B (destroyed), F neg. 4146); lectotype: F! (fragment)). Peru. Amazonas: Chachapoyas, Soloco, Paraje Olia, 1.44 km S of Quitachi, W of road to Pampa del Tío, 6°20'2.85"S, 77°41'2.91"W, 2304 m, 22 Jun. 2011, ♀, flowers, M.S. Samain et al. 2011-062 epitype, designated here: USM!; isoepitypes: GENT!, IEB!, MEXU!).

Description.

Root-climbing liana of up to 20 m high, functionally dioecious; free-growing branches quadrangular, slightly fissured; leaves decussate, petiole flattened adaxially, sulcate in young leaves (Fig. 15B View Figure 15 ), terete abaxially, color dark brown to black, glabrous, leaving a semicircular to triangular scar on the branch when leaves fall, 1.0-2.6 cm long; lamina (ob)ovate to lanceolate, 13-21 cm long, 6-12 cm broad, base cuneate, sometimes asymmetric, apex rounded to acuminate, leaf margin slightly glandular-dentate, venation brochidodromous, veins 6-10 pairs, adaxial leaf side with midvein protruding, primary veins lightly marked, secondary veins barely visible, moderately pubescent with white, stellate hairs, caducous, abaxially with protruding midvein and primary veins, secondary veins marked, opaque olive green, secondary veins in between the primary veins parallel to these and forming a distinctive regular pattern with veins arching towards the apex (Fig. 15A View Figure 15 ), sometimes, primary and secondary veins starting from nearly the same position on the midvein, densely pubescent with stellate reddish hairs, especially on the veins, caducous, sometimes nearly glabrous, acarodomatia numerous, not very conspicuous as they are hidden under the pubescence, in axils of midvein and primary veins, sometimes forming a small membrane partially covering the cavity; inflorescence axis pubescent with simple white hairs to glabrous (Fig. 15C View Figure 15 ), 9-47 cm long, with 3-5 decussate leaf pairs below the inflorescence, persistent, petiole 0.5-1 cm long, lamina 2-11.5 cm long, 1.1-7.5 cm broad, adaxially glabrous, abaxially pubescent with small stellate hairs, apex of the inflorescence axis woody, cylindrical, elongated bract scars visible, thickening at the top, 6-10 mm broad, 4-7 mm high in functionally female plants, 5-7 mm broad, 4 mm high in functionally male plants, inflorescence bracts cucullate, cream-colored, coriaceous, margin membranous and glabrous to densely pubescent with yellowish stellate hairs, veins darker, parallel, bracts increasing in size, lowermost bract 2.5-3 cm large, 2.4-2.5 cm broad, inflorescences lateral, decussate, 1-3 pairs of inflorescences per flowering branch (Fig. 16A View Figure 16 ), sometimes only one inflorescence developing, flowering branch continues growing vegetatively with the same vigor and periodicity above the inflorescences as below these, 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 53 cm apart (in between the main branch ramified and those branches also carried inflorescences that flowered at the same time as the last inflorescences, Samain et al. 2013-130), medulla central in the branch, disappearing in older branches, leaving a hole only, inflorescence umbellate, buds up to 4.1 cm broad and 3.6 cm high before opening, in flowering stage 8-22.5 cm diameter, 3.5-14 cm high, with 6-10 main axes in functionally male plants, 7-14 main axes in functionally female plants, partial inflorescences cymes, secondary and tertiary inflorescence axes with reddish stellate hairs, pubescence gradually increasing towards flower insertion; enlarged marginal flowers sometimes present (Fig. 16A-C View Figure 16 ), terminally placed in a cyme, sepals with marked veins, (2-)4, separate at the base, veins marked, palmate, reticulate, pistils 2, reduced, central part of the flower generally amorph, further characters not observed in detail, 1.5-3.7 cm diameter, pedicel 1-2.5 cm long; flower pedicel of reduced flowers 3-7 mm long in functionally male flowers, 3 mm long in functionally female flowers, receptacle triangular in functionally male flowers, rectangular in functionally female flowers, ovary inferior, calyx lobes 4, triangular, 0.2 mm long, petals 3-4, purple, valvate, cucullate, papyraceous, 2.2-2.5 mm long, 1.2-1.3 mm broad; functionally male flowers (Fig. 16D View Figure 16 ): hypanthium 1.1-1.8 mm diameter, stamens 8, well-developed, filaments 3 mm long, anthers 0.8-1.0 mm long, 0.6 mm broad, pistils 2, reduced, 0.5 mm long, stigmas not penicellate; functionally female flowers (Fig. 16E View Figure 16 ): hypanthium 1.0-1.8 mm diameter, stamens 8, reduced, filaments 0.1-0.4 mm long, anthers 0.1 mm long, 0.3-0.4 mm broad, pistils 2, 0.7 mm long, enlarging to 2-3 mm during fruit maturation, stigmas apically clavate and shortly penicellate; fruit a rectangular capsule, apically with a conspicuous border, ribs 4-6, pale reddish brown, 1.5 mm high, 2.5 mm broad above, 1.5 mm diameter, opening between the two pistils to release seeds, seeds not seen.

Distribution.

Hydrangea weberbaueri   is known from Colombia, Ecuador and Peru (Fig. 14 View Figure 14 ).

Habitat.

This species grows in tropical rainforest and mountain cloud forest at elevations between 974-3500 m.

Phenology.

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

Vernacular names and use.

The following vernacular names are indicated on the specimen Ellemann 75381 (AAU, LOJA, MO, QCA): bejuco matapalo (Spanish), yura huanutic caspic (Quichua). Its use as fuel wood is also indicated on the label of these specimens. This is the only species of this study of which vernacular name and use have been recorded.

Notes.

Not to be considered a synonym of H. peruviana   from which it can be distinguished by the notorious regular venation pattern with primary veins and secondary veins parallel to each other, all of them arching towards the apex.

No duplicate material of the original type collection has been located in the herbarium MOL of the Universidad Nacional Agraria La Molina in Lima, Peru, nor any other herbarium where we have searched. The fragment held in F that is the obligate lectotype is fragmentary and does not have the diagnostic features of this taxon, therefore we designate an epitype here for diagnostic purposes. The epitype specimen was collected approximately 8.5 km south of the original type location, which at the time of our field work was the nearest location to Cheto with primary forest. Unfortunately, satellite images ( Google Earth 2020) show that the forest where the epitype has been collected was destroyed for land use change less than five years after our visit.

A Weberbauer collection with the same number as the original type is present in the herbarium USM, but this is an entirely different species (with branched inflorescences) which does not belong to this group. The label on the USM specimen is not the original Weberbauer label, suggesting this represents a labeling error.

The leaf venation in the specimens from Colombia does slightly differ from that in the material from Ecuador and Peru, although it is still very recognizable. Therefore, we decided to include these in H. weberbaueri   , although future studies on Colombian hydrangeas may well show it concerns a distinct and new species.

According to our molecular study, Hydrangea weberbaueri   is not closely related to any of the species mentioned here, but it occurs in a clade which is sister to the one with H. goudotii   and H. trianae   (Granados Mendoza et al. unpublished results).

Preliminary conservation status.

Although this species has an EOO of nearly 660,000 km2, it is Endangered according to the IUCN categories and criteria ( IUCN 2012), with an AAO of 224 km2, as well as an extensive reduction in both EOO and AOO resulting from severe habitat destruction and fragmentation.

Additional specimens examined.

Colombia. Cundinamarca: Quebrada la Aguadita, 2 km arriba de la carretera a Fusagasugá, 1800 m, 28 Nov 1960, ♀, fruits, Idrobo 4186 (COL); Cordillera Oriental, municipio San Bernardo, Vereda de Venecia, margen izquierda del río Chorrera, afluente del Sumapáz, 2450 m, 21 Jul 1981, floral buds, Jaramillo et al. 6844 (COL); Sasaima, vereda San Bernardo, quebrada La María y Río Dulce, 1800 m, sterile, Jan 1958, sterile, García-Barriga 15757 (COL); Santander: municipio Suaita, corregimiento San José de Suaita, carretera a La Veterana, bosque al lado de la carretera, vereda San Imidio, 6°9'44.5"N, 73°25'09.4"W, 1800-1910 m, 6 Apr 2003, ♀, fruits, Betancur et al. 10122 (COL, HUA).

Ecuador. Azuay: the eastern Cordillera, 1-8 km, north of the village of Sevilla de Oro, 8000-9000 ft, 27 Jul-12 Aug 1945, inflorescence buds, Camp E-4464 (AAU, F, MO); Sevilla de Oro, ca 1.7 airline km E of Osorancho (El Cisne), N Sevilla de Oro, 2°45'42"S, 78°37'38.1"W, 2804 m, 8 Jun 2012, sterile, Granados Mendoza et al. 2012-22 ( GENT, HOXA, IEB, MEXU, QCNE); same data as preceding, 2°45'42.1"S, 78°37'37.4"W, 2804 m, 8 Jun 2012, sterile, Granados Mendoza et al. 2012-23 ( GENT, HOXA, IEB, MEXU, QCNE); same data as preceding, 2°45'42"S, 78°37'37.3"W, 2805 m, 8 Jun 2012, sterile, Granados Mendoza et al. 2012-24 ( GENT, HOXA, IEB, MEXU, QCNE); ca 1.8 airline km E of Osorancho (El Cisne), N Sevilla de Oro, 2°45'43.3"S, 78°37'35.9"W, 2813 m, 8 Jun 2012, inflorescence buds, Granados Mendoza et al. 2012-25 ( GENT, HOXA, IEB, MEXU, QCNE); Loja: Río Jipiro near Hda. Jipiro, about 7 km northeast of Loja, 03°58'S, 79°07'W, 8000 ft, 14-16 Jun 1944, ♂, flower buds, flowers, Prieto AP-27 (AAU, NY); between Nudo de Sabanillas and Río Cachiyacu at Tambo Cachiyacu, 3000-3500 m, 17 Oct 1943, ♂, flowers, Steyermark 54551 (F, US); Loja-Zamora, 12 km from Loja, on the finca of Dr. David Espinosa, 03°55'S, 79°09'W, 2400-2600 m, 17 Nov 1988, ♀, fruits, Ellemann 75381 (AAU, LOJA, MO, QCA); old road Loja-Zamora, El Tiro, 2600 m, 10 Oct 2004, ♂, flowers, Homeier 1399 (LOJA, QCNE); Loja, 3.5 Km S of Nudo de Sabanillas on road Yangana-Valladolid, 4°26'54.2"S, 79°8'51.3"W, 2798 m, 2 Jun 2012, sterile, Granados Mendoza et al. 2012-11 ( GENT, HOXA, IEB, MEXU, QCNE); Morona Santiago: Parque Nacional Sangay-Tinguichaca, 2°13'559632"S, 78°26'364956"W, 2870 m, 29 Aug 2011, sterile, Duchicela et al. 759 (QCA); Morona, ca. 4.5 airline km WNW of Plan de Milagro on road Plan de Milagro-Gualaceo, 3°0'14.6"S, 78°30'29.2"W, 2158 m, 7 Jun 2012, ♀, fruits, Granados Mendoza et al. 2012-21 ( GENT, HOXA, IEB, MEXU, QCNE); Napo: area of the Yanayacu biological station, 5 km SW of Cosanga, 00°36'S, 77°53'W, 2100 m, 27 Sep 2005, ♀, fruits, Homeier 1680 (QCNE); Pichincha: Reserva Florística - Ecológica " Río Guajalito", km 59 de la carretera antigua Quito - Sto. Domingo de los Colorados, a 3.5 km al NE de la carretera, estribaciones occidentales del Volcán Pichincha, 0°13'53"S, 78°48'10"W, 1800-2200 m, 9 Aug 1985, ♂, flowers, Jaramillo & Zak 7900 (NY, QCA, US); same data as preceding, 2200 m, 25 Apr 1987, sterile, Grijalva 317 (QCA); road from Chiriboga to Santo Domingo, 1910 m, 3 May 1985, ♂, flowers, Stein 2680 (AAU, NY, QCNE); Cordillera Oriental, Oya Cachi, 3200 m, 26 Oct 1945, ♂, old flowers, Acosta 11144 (F); Sucumbios: Gonzalo Pizarro, near km 14 of Vía a casa de Máquinas of Bosque Protector los Cedros, 0°7'21.9"S, 77°26'42.1"W, 974 m, 9 Jul 2012, sterile, Granados Mendoza et al. 2012-103 ( GENT, HOXA, IEB, MEXU, QCNE); Tungurahua: Cordillera el Encanto, ca. 6 airline km SSW of Río Negro, 1°27'33.4"S, 78°13'7"W, 1815 m, 13 Jun 2012, floral buds, Granados Mendoza et al. 2012-35 ( GENT, HOXA, IEB, MEXU, QCNE); same data as preceding, 1°27'34.6"S, 78°12'53.7"W, 1810 m, 13 Jun 2012, inflorescence buds, Granados Mendoza et al. 2012-36 ( GENT, HOXA, IEB, MEXU, QCNE); same data as preceding, 1°27'42.5"S, 78°12'52"W, 1806 m, 13 Jun 2012, inflorescence buds, Granados Mendoza et al. 2012-37 ( GENT, HOXA, IEB, MEXU, QCNE); Zamora-Chinchipe: Carretera Loja-Zamora, km 15-18, falda oriental, 3°58'S, 79°7'W, 23 Sep 1982, floral buds, Balslev 3181 (AAU, MO, NY, QCA); Zamora, 3.7 km on foot path S Romerillos Alto, 4°14'49.9"S, 78°56'9.1"W, 1681 m, 4 Jun 2012, ♂, inflorescence buds, flowers, Granados Mendoza et al. 2012-16 ( GENT, HOXA, IEB, MEXU, QCNE); ca. 4.8 km on foot path S Romerillos Alto, 4°15'0.6"S, 78°55'40.4"W, 1801 m, 4 Jun 2012, sterile, Granados Mendoza et al. 2012-17 ( GENT, HOXA, IEB, MEXU, QCNE); same data as preceding, 4°15'0"S, 78°55'39.9"W, 1795 m, 4 Jun 2012, sterile, Granados Mendoza et al. 2012-18 ( GENT, HOXA, IEB, MEXU, QCNE); same data as preceding, 4°15'0.2"S, 78°55'39.4"W, 1796 m, 4 Jun 2012, ♂, flowers, Granados Mendoza et al. 2012-19 ( GENT, HOXA, IEB, MEXU, QCNE); El Pangui, ca. 5.7 km SSE of Tundayme on dirt road to military base El Condor, 3°36'19.6"S, 78°28'21.1"W, 1092 m, 7 Jun 2012, sterile, dry inflorescences, Granados Mendoza et al. 2012-20 ( GENT, HOXA, IEB, MEXU, QCNE).

Peru. Amazonas: Bongará, Jumbilla, Bosque de Protección Alto Mayo, surroundings of Chichilac, 4.79 km NNE of Jumbilla, 5°51'51.33"S, 77°47'16.7"W, 2462 m, 10 Dec 2013, ♀, old fruits, Samain et al. 2013-130 ( GENT, IEB, MEXU, USM); same data as preceding, 4.65 km NNE of Jumbilla, 5°51'56.7"S, 77°46'55.6"W, 2649 m, 30 Jun 2011, ♂, flowers, Samain et al. 2011-103 ( GENT, IEB, MEXU, USM); same data as preceding, 4.57 km NNE of Jumbilla, 5°51'51.9"S, 77°47'16.7"W, 2397 m, 30 Jun 2011, ♀, flower buds, flowers, Samain et al. 2011-097 ( GENT, IEB, MEXU, USM); Bosque de Protección Alto Mayo, surroundings of Chichilac, 4.78 km NNE of Jumbilla, 5°51'52.51"S, 77°47'0.86"W, 2570 m, 10 Dec 2013, ♀, fruits, Samain et al. 2013-133 ( GENT, IEB, MEXU, USM); Bongará, Yambrasbamba, carretera Chiclayo-Tarapoto, km 370.2, antes de Puente Nieva, 5°41'6.1"S, 77°46'56.5"W, 2031 m, 9 Dec 2013, ♂, flower buds, flowers, Samain et al. 2013-127 ( GENT, IEB, MEXU, USM); margen del Río Nieva, km 381 de la Carretera Marginal, 5°41'S, 77°47'W, 1070 m, 11 Jul 1999, ♀, flower developing into fruits, Sánchez Vega 10031 (F); Florida, 4-8 km E of Pomacocha on road to Rioja, 5°45'S, 77°53'W, 2100-2200 m, 9 Feb 1984, ♂, flowers, Gentry 45191 (AMAZ, CAS, COL, MEXU, MO); Chachapoyas, Leymebamba, Hacienda San Cristobal del Yeso, 3.08 km NW of Leymebamba, 6°41'34.9"S, 77°49'36.3"W, 2766 m, 20 Jun 2011, ♂, flowers, Samain et al. 2011-046 ( GENT, IEB, MEXU, USM); same data as preceding, Hacienda San Cristobal del Yeso, 2.7 km NNW of Leymebamba, 6°41'39.33"S, 77°49'29.23"W, 2738 m, 8 Dec 2013, ♀, fruits, Samain et al. 2013-124 ( GENT, IEB, MEXU, USM); alrededor de la Laguna de Los Cóndores, parte sur, 6°51.20'S, 77°40.96'W, 2500-2700 m, 16 Aug 1998, ♀, fruits, Quipuscoa 1218 (F, NY, QCNE); San Francisco de Daguas, 2.4 km NNE of Pipus, Chachapoyas-Molinopampa road, km 30, 6°12'32.3"S, 77°43'22.8"W, 2110 m, 21 Jun 2011, ♀, fruits, Samain et al. 2011-056 ( GENT, IEB, MEXU, USM); Pampa del Tio, camino entre Quitachi y (las) Tinas, 2000 m, 5 Jul 1991, ♂, flowers, Díaz 4609 (USM); Cajamarca: Cutervo, San Andrés de Cutervo, 3.9 km SE of San Andres Cutervo, road Cutervo-Socota-Chiple, 6°16'05.6"S, 78°41'35.4"W, 2513 m, 8 Jul 2011, inflorescence buds, Samain et al. 2011-141 ( GENT, IEB, MEXU, USM); same data as preceding, 3.92 km SE of San Andrés de Cutervo, road Cutervo-Socota-Chiple, 6°16'05.5"S, 78°41'34.5"W, 2518 m, 8 Jul 2011, ♀, inflorescence buds, Samain et al. 2011-143 ( GENT, IEB, MEXU, USM); La Pucarilla, entre Socota y San Andrés, 2500 m, 3 Nov 1991, ♂, flowers, Sánchez 5947 (F, MO, UC); Bosque Cutervo, Parque Nacional de Cutervo, NW corner of Cordillera Tarros, Chorro Blanco sector, ca. 10 km WNW of San Andres de Cutervo, 6°12'S, 78°46'W, 2650 m, 4 Nov 1990, ♂, inflorescence buds, flowers, Dillon 6152 (F, UC); Cutervo National Park, Chorro Blanco area, 15 km N of San Andres de Cutervo, 6°10'S, 78°40'W, 2400-2650 m, 13 Sep 1991, ♀, flower buds, flowers, Gentry 74745 (F, USM); Parque Nacional Cutervo, arriba del Saucedal pasando por Chorro Blanco, 2250 m, 3 Aug 1988, ♂, inflorescence buds, flower buds, flowers, Díaz 2943 (USM); Cutervo National Park, 12 km NE of San Andres Cutervo, transect 5, 6°10'S, 78°40'W, 2240 m, 11 Sep 1991, sterile, Gentry 74670 (MO, USM); Madre Mia, entre el Suro y la Flor al NO del Parque, 2400 m, 25 Jun 1992, ♂, inflorescence buds, flower buds, flowers, Sánchez Vega 6329 (F); Bosque Cutervo, Parque Nacional de Cutervo, NW corner of Cordillera Tarros, Chorro Blanco sector, ca. 10 km WNW of San Andres de Cutervo, 6°12'S, 78°46'W, 2600 m, 5 Nov 1990, ♂, flowers, Dillon 6184 (F); Hualgayoc, 2 km. above Palmito. Hacienda Taulis, between the Casa Hacienda and Palmito, 2575 m, 31 Aug 1964, ♂, flowers, Hutchinson 6395 (GH, F, US); San Ignacio, Huarango, poblado Selva Andina, 05°03'37"S, 078°45'13"W, 1798 m, 18 Apr 2007, ♀, fruits, Perea & Mateo 2929 (MO, QCNE); San Ignacio, 12.83 km SW of San Ignacio, 3.05 km SSW of Alto Ihuamaca, 5°12'52.8"S, 79°05'47.9"W, 2129 m, 6 Jul 2011, ♀, fruits, Samain et al. 2011-127 ( GENT, IEB, MEXU, USM); Nuevo Mundo-Pisaguas, 5°10'S, 68°32'W (the correct coordinates are 78°32'W), 1550 m, ♀, fruits, 13 Nov 1997, Campos 4632 (F, MEXU, MO, NY, USM); Pasco: Oxapampa, Parque Nacional Yanachaga Chemillén, parte alta del Refugio el Cedro, 10°32'S, 75°21'W, 2450-2680 m, 20 Mar 2003, inflorescence buds, Monteagudo 4762 (F, HOXA, USM); Oxapampa, sector Chacos parte media, 1 0°38'42.9"S, 75°17'30"W, 2740 m, 4 Nov 2004, floral buds, Monteagudo et al. 7534 (HOXA); Oxapampa, camino a Paucartambo (a la altura de Mesapata), < 1800 m, 21 Mar 2005, ♀, flower buds, fruits, Arias et al. 261 (HOXA); Oxapampa, Huancabamba, camino hacia la Laguna San Daniel, Granapazú-Sector San Daniel, Parque Nacional Yanachaga-Chemillén, 10°26'15.6"S, 75°27'9.9"W, 2200 m, 12 Sep 2005, sterile, Ortiz et al. 950 (HOXA); Oxapampa, Huancabamba, Parque Nacional Yanachaga-Chemillen, sector San Daniel, 10°26'35S, 75°26'16W, 2200-2500 m, 6 Mar 2006, ♂, flowers, Vásquez et al. 30989 (HOXA); Oxapampa, Huancabamba, sector Quebrada Yanachaga (paralela a la quebrada), 10°24'08"S, 75°29'18"W, 2097 m, 14 Sep 2004, sterile, Perea & Mateo 1682 (HOXA); Oxapampa, El Abra, camino Oxapampa-Villa Rica, 10°40'30"S, 75°18'21"W, 2365 m, 24 Apr 2004, ♂, flowers, Rojas et al. 2242 (HOXA); San Martín: Mariscal Cáceres, Huicungo, Río Abiseo National Park, 7°S, 77°W, 2700 m, 22 Jul 1985, ♀, flowers, Young 1230 (F, NY); near La playa base camp, Río Abiseo National Park, 7°27'N, 77°2'W, 2650 m, 30 Aug 1985, ♂, inflorescence buds, flowers, Young 1512 (MO); Río Abiseo National Park, ridge top on hill E of that with Gran Pajaten ruins, 2550 m, 14 Aug 1986, ♀, fruits, Young 4141 (USM); trail between La Playa camp and Papayas camp, Rio Abiseo National Park, 7°S, 77°W, 2650-2750 m, 25 Jul 1987, ♀, flower buds, flowers, Young 5014 (F, USM); Pardo Miguel, Caserío Jorge Chávez, km 398 de Carretera Marginal, 5°40'S, 77°43'W, 1400 m, 1 Jul 1999, ♀, fruits, Sánchez Vega 9966 (F); Rioja: Pardo Miguel, 7.90 km E of El Progreso, Chiclayo-Tarapoto road, km 369.7, 5°41'18.8"S, 77°47'12.0"W, 2063 m, 23 Jun 2011, ♀, flower buds, flowers, Samain et al. 2011-067 ( GENT, IEB, MEXU, USM); same data as preceding, 7.94 km E of El Progreso, Chiclayo-Tarapoto road, km 369.7, 5°41'16.9"S, 77°47'11.9"W, 2132 m, 23 Jun 2011, ♀, inflorescence buds, flower buds, flowers, fruits, Samain et al. 2011-068 ( GENT, IEB, MEXU, USM); carretera Chiclayo-Tarapoto, km 377.5, 5°35'59.8"S, 77°45'18.7"W, 1867 m, 9 Dec 2013, ♀, fruits, Samain et al. 2013-129 ( GENT, IEB, MEXU, USM); Buenos Aires, along road Pedro Ruiz-Rioja, 5°42'9"S, 77°53'6"W, 2000 m, 21 Mar 1998, ♀, inflorescence buds, flower buds, flowers, van der Werff 15329 (MEXU, USM).