Polylepis reticulata Hieron., Bot. Jahrb. Syst. 21: 312. 1896.
publication ID |
https://dx.doi.org/10.3897/phytokeys.203.83529 |
persistent identifier |
https://treatment.plazi.org/id/90B8A195-AB78-BA4D-367A-16149660A9AE |
treatment provided by |
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scientific name |
Polylepis reticulata Hieron., Bot. Jahrb. Syst. 21: 312. 1896. |
status |
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20. Polylepis reticulata Hieron., Bot. Jahrb. Syst. 21: 312. 1896.
Figs 55 View Figure 55 , 56 View Figure 56
Polylepis brachyphylla Bitter, Bot. Jahrb. Syst. 45: 616. 1911. Type. Ecuador. Bolivar: Volcán Chimborazo, slopes towards Guaranda, 4 November 1856, Remy s.n (holotype: P!; isotypes: GOET!, P!).
Polylepis nitida Bitter, Bot. Jahrb. Syst. 45: 615. 1911. Type. Ecuador. Tungurahua: Minza at Volcan Tungurahua, 3600 m, Stübel 287 (holotype: B destroyed; photos at F!, GH, MO!, NY, US!).
Type.
Ecuador. Pichincha: near Las Calderas de Pasachoa and Rumiñaqui, Stübel 20a. (holotype: B destroyed; photos at F!, GH, NY, US).
Description.
Trees 2-15 m tall. Leaves slightly congested at the branch tips, imparipinnate with (1-)2-3(-4) pairs of lateral leaflets, obtrullate in outline, (1.3-)2.6-4.2(-7.0) × (1.5-)2.2-3.6 cm; rachises densely tomentose, points of leaflet attachment with a tuft of long, lanate hairs; stipular sheaths apically acute with spurs, densely lanate on the outer surfaces; leaflets elliptic to obovate in outline, second pair from the terminal leaflet the largest, one of this pair (0.9-)1.2-2.2 × 0.4-1.1 cm; margin entire or slightly crenate at the apex with 3-5 teeth, apically deeply emarginate, basally unequally cordate; upper leaflet surfaces glabrous; lower leaflet surfaces densely tomentose with whitish hairs 0.6-1.5 mm long. Inflorescences pendant, (2.3-)4.3-13.8) cm long, bearing 5-9 flowers; floral bracts 3.1-8.0 mm long, narrowly triangular, densely lanate on the outer surface; rachises tomentose. Flowers 6.2-12.2 mm diam.; sepals 4, ovate, green, densely tomentose outside; stamens 7-9, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 2.6-3.9 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely villous; 3.4-6.0 × 2.9-8.3 mm including spines. Diploid and hexaploid.
Distribution, habitat and ecology.
Polylepis reticulata is found in Carchi, Imbabura, Pichincha, Tungurahua, Bolivar, Chimborazo, Azuay and Loja (Ecuador) (Fig. 61 View Figure 61 ). It grows under relatively dry conditions (except at Llanganates and Pasochoa that are more humid) in Andean forest at 2750-4600 m elevation. It often co-occurs with P. simpsoniae in Azuay. In Sacha Huayco Forest (Chimborazo) at 4000-4500 m elevation, the canopy layer is comprised almost completely by P. reticulata , with Gynoxys sp. dominating the subcanopy ( Osha 2000). At 4300 m elevation in Chimborazo Wildlife Production Reserve, trees of P. reticulata are, on average, 2.8 m tall (maximum 8.2 m) with diameters of 8.6 cm (maximum 16.9 cm) ( Castillo et al. 2017). In Cuenca (Ecuador), the minimum temperature threshold of vegetative growth activity was found to be close to 6 °C ( Saravia et al. 2016). In the same forest, mean annual litterfall is 3.77 Mg ha-1, representing 51% of the canopy leaf biomass, indicating a leaf life span of 1.98 years and a mean decomposition rate of 0.38 year-1 ( Pinos et al. 2017). Diameter growth rates measured over seven years average 1.2 mm/year and the average carbon sequestration rate is 2.6+/-0.3 Mg C. ha-1 year-1, which is high for this generally slow-growing genus ( Montalvo et al. 2018). Tree rings of P. reticulata are characterized by semi-ring porosity and minor changes in fiber wall thickness between latewood and earlywood. Tree ring width is more related to temperature than precipitation ( Alvites et al. 2019). Based on an AFLP study, clonal propagation affects the fine-scale genetic structure within a spatial distance of 3 m ( Peng et al. 2015). In a forest of P. reticulata , the soil had low microbiological quality with low biological activity, even though there were some beneficial nematodes representing a great advantage for the elimination of insect pests that could affect Polylepis ( Castillo et al. 2017). Hybrids with P. pauta have been found in Pichincha.
Conservation status.
The EOO for Polylepis reticulata is estimated as 33,103 km2, the AOO is assessed at 324 km2 and it is known from 40 locations. Most of the populations occur in protected areas. There are populations in the Cajas (Azuay), Llanganates (Tungurahua) and Yacuri (Loja) National Parks, Chimborazo Wildlife Production Reserve, Illinizas Ecological Reserve (Pichincha) and in part of Pasochoa Ecological Reserve. However, P. reticulata was categorized as Vulnerable VU (A4c) ( León-Yánez et al. 2011). Although the species occurs within national parks, its populations are heavily fragmented. We assess P. reticulata as Vulnerable (B1a+B2a, C1).
Notes.
Polylepis reticulata shows some degree of morphological differentiation between juvenile and adult leaves, with the young leaves having more lateral leaflet pairs (4 versus 1-3), slightly crenate margins (versus entire) and sparsely tomentose in the upper leaflet surfaces (versus glabrous). This differentiation was the basis for the recognition of three species by Bitter (1911) in what is now recognized to be a single species.
Polylepis reticulata is morphologically closest P. occidentalis , with which it shares similar leaflet size, margin types and number of lateral leaflets pairs. It differs by its longer lower leaflet surface hairs (0.6-1.5 mm), longer inflorescences ((2.3-)4.3-13.8 cm) and longer styles (2.6-3.9 mm), compared to the shorter lower leaflet surface hairs (0.3-0.6 mm), shorter inflorescences (2.4-6.7 cm) and shorter styles (1.5-2.0 mm) of P. occidentalis . Polylepis reticulata is also morphologically close to P. simpsoniae , with which it shares similar leaflet surface hair type and density. The most obvious differences between P. reticulata and this species are leaflet length (0.9-2.2 cm versus 0.9-1.6 cm), length of hairs on the lower leaflet surfaces (0.6-1.5 mm versus 0.5-0.7 mm) and longer inflorescences ((2.3-)4.3-13.8 cm versus (2.5-)3.7-3.9(-5.5) cm).
Specimens examined.
Ecuador. Azuay: Chaucha, Páramo de Soldados, 02°53'S, 079°18'W, 3700-4000 m, 28-29 August 1985, Lægaard 55114C (AAU!). Cuenca, Páramo de Cajas, 3650-3890 m, s.d., Boeke 631 (MO!, QCA!); Parque Nacional El Cajas, 02°50'S, 079°12'W, 3500 m, 19 September 1983, Brandbyge 42269 (MO!); Along the río Matadero, west Cuenca, 03 March 1945, Camp E-2000 (MO!); along the río Matadero, west Cuenca, 3500 m, 03 March 1945, Camp E-2000 (MO!, NY, US!); Area Nacional de Recreación Cajas, Laguna Toreadora and Virgen Machay, 02°46'S, 079°13'W, 4000-4400 m, 18 December 1995, Clark 1794 (MO!, QCNE); Parque Nacional El Cajas, Laguna Toreador, 02°53'S, 079°08'W, 3500 m, 19 April 1998, Clark 4992 (AAU!, MO!); Area Nacional de Recreación “Cajas” Sayausi-Molleturo road W of Cuenca, 02°53'S, 079°59'W, 3910 m, 21 June 1989, Dorr 6391 (AAU!); Parque Nacional de Cajas, WNW of Cuenca, 02°52'S, 079°09'W, 3450 m, 28 December 1979, Holm-Nielsen 20927 (AAU!); Parque de Recreación Cajas, 4000-4100 m, 02 September 1983, Jaramillo 7185 (AAU!, GB, QCA!); Parque Reacreacional Cajas, Strasse von Cuenca nach Morlleturo, 3900 m, 14 March 1991, Kessler 2746B (GOET!); Cuenca-Molleturo road ca. 5 km W of pass in Las Cajas, 02°49'S, 079°16'W, 3350 m, 01 May 1992, Lægaard 102646 (AAU!, GOET!, QCA!); Parque Nacional de Las Cajas, along road Soldados-Angas, in pass app. 13 km from Soldados, 02°53'S, 079°17'W, 4000 m, 02 May 1992, Lægaard 102691 (AAU!, GOET!, QCA!); 102696 (AAU!, QCA!); Páramo de las Cajas W of Cuenca, 02°47'S, 079°14'W, 4000-4150 m, 02 September 1984, Lægaard 52870 (AAU!, QCA!); 52878 (AAU!, MO!, QCNE); 52885 (AAU!, MO!, QCNE); 52888 (AAU!); 52891C (AAU!); Páramo de Soldados SW of Cuenca, 02°53'S, 079°17'W, 3700-3800 m, 24 October 1984, Lægaard 53243 (AAU!); Páramo de Soldados, SW of Cuenca, 02°53'S, 079°18'W, 3750-3850 m, 03 March 1985, Lægaard 53800 (AAU!, MO!); Páramo de Cajas, W of Cuenca, along new road, ca. 6-8 km W of pass, 02°48'S, 079°17'W, 3800 m, 31 March 1985, Lægaard 53935 (AAU!, QCA!); Páramo de Soldados, 02°53'S, 079°18'W, 3700-4000 m, 28-29 August 1985, Lægaard 55098 (AAU!); 55098C (AAU!); 55098D (AAU!); 55113 (AAU!); 55114A (AAU!); 55114B (AAU!); Páramo de Soldados-Angas, at highest point of road, 02°52'S, 079°17'W, 4000 m, 14 February 1988, Lægaard 70122 (AAU!, MO!); Area Nacional Recreacional Cajas, sector Llavinco, 02°50'S, 079°20'W, 3450-3550 m, 09 January 1991, León 2606 (USM!); Parque Nacional Cajas, km 32.1 desde redondel Cuenca-Molleturo, sendero este alrededor de Laguna Toreadora, 3850-3870 m, 08 January 2003, Ulloa 1013 (BRIT, HA, K, MO!, QCNE, US!); 1014 (HA, MO!, NY, QCNE); Parque Nacional Cajas, km 28 redondel Cuenca-Molleturo, sendero izquierdo Laguna Patoquinoas, 3730-3800 m, 12 January 2003, Ulloa 1130 (HA, ILLS, MO!); Parque Nacional Cajas, km 35 Cuenca-Molleturo, Centro de Información, Laguna Toreadora sendero oeste, 3880-3890 m, 16 January 2003, Ulloa 1234 (HA, ILLS, MO!, QCNE). Molleturo, Páramo de las Cajas, W of pass, 02°46'S, 079°15'W, 3700 m, 27 August 1985, Lægaard 55047 (AAU!, MO!, QCA!); Area Nacional Recreacional Cajas, sector La Toreadora, 02°50'S, 079°20'W, 3950 m, 12 January 1991, León 2624 (USM!); Area nacional Recreacional Cajas, sector La Toreadora, 02°50'S, 079°20'W, 3800 m, 13 January 1991, León 2651 (QCA!, USM!); Carretera Sayausi-Molleturo km 10-31, Parque Recreacional Las Cajas, 02°50'S, 079°15'W, 3350-3950 m, 28 November 1992, Romoleroux 1482 (AAU!, QCA!). San Antonio, 3-5 km W of pass at Las Cajas, W of Cuenca, 02°40'S, 079°14'W, 3800-4000 m, 22 October 1984, Lægaard 53199 (AAU!); Las Cajas, near Laguna Toreadora, 02°43'S, 079°12'W, 3900 m, 13 September 1983, Larsen 45141 (AAU!, QCA!). San Joaquin, Páramo de Soldados SW of Cuenca, 02°53'S, 079°17'W, 3700-3800 m, 24 October 1984, Lægaard 53235; 53235B; 53235D; 53235F; 53235H; 53245; 53246 (AAU!); between Cuenca and Soldados, 02°57'S, 079°10'W, 2800-3000 m, 28 August 1985, Lægaard 55081 (AAU!); at Soldados, 02°57'S, 079°14'W, 3200-3400 m, 28 August 1985, Lægaard 55084 (AAU!); Páramo de Soldadas-Angas, at highest point of road, 02°52'S, 079°17'W, 3950-4050 m, 14 February 1988, Lægaard 70101 (AAU!); Páramo de Soldadas-Angas, at highest point of road, 02°52'S, 079°17'W, 3200-3300 m, 14 February 1988, Lægaard 70101A (AAU!); 70101B (AAU!); 70101C (AAU!); 70101D (AAU!). Sayausi, Parque Nacional de Cajas, WNW of Cuenca, 02°52'S, 079°09'W, 3450 m, 28 December 1979, Holm-Nielsen 20926 (AAU!); Páramo de las Cajas W of Cuenca, 02°47'S, 079°14'W, 4000-4150 m, 02 September 1984, Lægaard 52879; 52886; 52890; 52891; 52891A; 52891B; 52891D (AAU!); Cuenca-Area Recreacional Las Cajas, 26-30 km de Cuenca, 02°50'S, 079°06'W, 3360-3500 m, 17 November 1991, Romoleroux 1187 (AAU!, QCA!). Zurucucho, Canton Cuenca, 3200 m, 11 August 1978, Boeke 2622 (AAU!); ca. de la laguna La Toreadora, 3850 m, 26 December 1997, Burnham 3007 (QCA!); Dudahuaycu, Mazán, 3300 m, 28 May 1994, Chacón 211 (QCA!); Cantón Cuenca, collection made along río Patul from the Comuidad Baute Laguna Patul, watershed of río Patul, 3500-4200 m, 05 February 2001, Clark 6229 (QCA!); Parque Nacional Cajas, morenas ca. de la Laguna Toreadora, 4025 m, 15 November 2000, Endara 520 (QCA!); Parque Nacional Cajas, morrenas cerca de la Laguna de Soldados, 3830 m, 17 November 2000, Endara 564 (QCA!); ca. 10 km W of Sayausid, 3300 m, 04 November 1982, Harling 20218 (AAU!, GB); above Sayausi at first bridge over Río Tomebamba, 3200 m, 03 March 1985, Harling 22671 (AAU!, GB, QCA!); Páramo de Cajas, versant oriental; zone très humide; relique forestière dégradée, 3650 m, 10 April 1989, Huttel 1668 (QCA!); Páramo de Cajas, versant oriental, zone très humide, 3350-3950 m, 14 April 1988, Huttel 995 (QCA!); carretera Cuenca-Sayausid-Area de Recreación Cajas, 3900 m, 16 August 1987, Jaramillo 9898 (MO!); Parque Nacional Cajas, near headquarters on cuenca-Guayaquil road, 3900 m, 12 March 1991, Kessler 2746 (QCA!); Parque Recreacional Cajas, Strasse von Cuenca nach Molleturo, 3900 m, 14 March 1991, Kessler 2746A (GOET!); Páramo de Soldados, 3700 m, s.d., Lægaard 55096 (QCA!); 55114 (QCA!); at bridge 2 km below Soldados, 3200 m, s.d., Lægaard 70082 (QCA!); at bridge 2 km below Soldados, 3200 m, s.d., Lægaard 70083 (QCA!); s.d., León 2527 (QCA!); colección en parcela del bosque, arriba del lago Dos Chorreras, 3940 m, 05 July 1995, León 3541 (QCA!); arriba de laguna Chorreras, 3940 m, 05 July 1995, León 3542 (AAU!, QCA!); colección en bosque en la base del valle de Cajas cerca del hotel Dos Chorreras y en la ladera SO de la propiedad Dos Chorreras, 3400 m, 06 July 1995, León 3577 (QCA!); Vertiente oriental, 3100 m, 07 July 1995, León 3611 (QCA!); Área recreacional Cajas, 4000 m, 21 September 2000, Lizarzaburu 24 (QCA!); Fierroloma. Zorrocucho, ribera de Río Taitachugo, 3400 m, 20 May 1997, Minga 125 (QCA!); Totorococha-Mazan valley, 3500-4000 m, 12 September 1987, Ramsay 507 (QCA!); Parque Nacional Cajas, 3850-3900 m, 10 September 2001, Romoleroux 4028 (QCA!); Area de Recreación Cajas. km 1 de la via Miguir-Naranjal, 3500, 16 August 1987, Romoleroux 404 (QCA!); Hacienda Gulag, Masan, 3000-3200 m, 17 August 1987, Romoleroux 417 (QCA!); carretera Soldados-Angas, 3620 m, 04 April 2007, Romoleroux 4459 (QCA!); Bosque San Luis, 3939 m, 20 August 2008, Romoleroux 5108 (QCA!); 5127 (QCA!); carretera Molleoto Naranjal, parches de bosque fuera del parque nacional Cajas, 3939 m, 21 August 2008, Romoleroux 5157 (QCA!); Baños, 3300 m, 17 July 2006, Salgado 165 (QCA!); Cajas, 3725-3830 m, 24 June 1999, Smeets 432 (QCA!); Río Machangara, NW Cuenca, 3300-3400 m, 17 September 1967, Sparre 18568 (AAU!, S); s.d., Valverde 912 (MO!); carretera Sayausí-Quinona-Tres cruces, Area recreacional Cajas, 3500-4050 m, 10 April 1987, Zak 1894 (AAU!). Bolívar: upper west mountain Chimborazo, 3500-4000 m, 19 June 1887, Lehmann 4457 (MO!). Cañar: Vía Queseras-Rumiloma (Fund. Cordillera Tropical), 3353 m, 03 October 2006, Salgado 1181 (QCA!). Carchi: Espejo, parche de bosque alrededor de laguna Rasococha, 00°44'N, 078°04'W, 3600 m, 09 November 1993, Palacios 11897 (AAU!, MO!, QCNE). Chimborazo: Pungala, Near Alao, along Río Alao, 01°52'S, 078°30'W, 3200-3400 m, 21-22 September 1985, Lægaard 55295 (AAU!). Riobamba, Reserva Faunistica Chimborazo, 01°32'41"S, 078°53'05"W, 4100 m, 01 June 2006, Caranqui 1541 (CHEP, MO!); 3350-3500 m, 01 March 1944, Acosta-Solís 7524 (F!); 3750 m, s.d., Acosta-Solís 7696 (F!); 4000 m, s.d., André 3933 (F!, US!); San Juan, Chimborazo, 4350 m, 05 November 2013, Cardoso 4262 (QCA!); San Juan. Pasguazo Sambrano, 4100 m, 31 October 2013, Cardoso 4266 (QCA!);carretera Penipe a Utuñac Paramito, 2800 m, 07 April 2008, Jaramillo 26496 (QCA!); Sector Utuñac, Pijipamba, 3250-3330 m, 03 December 2008, Jaramillo 28123 (QCA!); Sector Utuñac, Pijipamba, 2800 m, 04 December 2008, Jaramillo 28132 (QCA!); E of Cordillera, 3400 m, s.d., Rimbach 14 p. p. (F!, NY, US!); 3400 m, s.d., Rimbach 156 (US!); s.d., Rimbach 498 (S); Vía Chambo-Guayllabamba-Aloa, 2900-3000 m, 09 August 1987, Romoleroux 366 (NY, QCA!). Imbabura: 4000-4300 m, s.d., Hirsch E-113 (NY). Loja: Manu, Cerro de Arcos W of road Manu-Zaruma, 03°34'S, 079°28'W, 3250-3600 m, 14 September 1999, Lægaard 20601 (AAU!, MO!). Napo: 3 km E of Paso de la Virgen on road Pifo-Papallacta, 3950-4050 m, 06 February 1985, Lægaard 54450 (QCA!); Parque Nacional Llanganates, 12 February 2009, Salgado 820 (QCA!). Pichincha: Cotogchoa, Volcán Pasochoa, Páramo and forest remains, 00°27'S, 078°30'W, 3500 m, 16 September 1985, Lægaard 55269A (AAU!, MO!); 55278 (AAU!, MO!); Bosque Protector Pasochoa, 30 km SE de Quito, 00°27'S, 078°28'W, 2850-3900 m, 08 September 1986, Romoleroux 161; 171 (AAU!, MO!, QCA!). El Chaupi, road El Chaupi-Iliñiza, 00°37'S, 078°40'W, 3200-3700 m, 20 June 1985, Lægaard 54547 (AAU!, QCA!); 54548 (AAU!, MO!); Volcán Iliñiza, N-side, 00°38'S, 078°41'W, 4200-4300 m, 20 June 1985, Lægaard 54554; 54555 (AAU!, MO!, QCA!). Rumiñahui, Bosque Protector Pasochoa, 00°22'S, 078°27'W, 3870-3900 m, 17 November 1990, Cerón 12323 (MO!). Rumipamba, Volcán Pasochoa, 00°27'S, 078°30'W, 2900-3300 m, 16 September 1985, Lægaard 55265 (AAU!, GOET!, MO!). Tambillo, Volcán Pasochoa, above house of Fundación Natura, 00°27'S, 078°31'W, 2800-3300 m, 27 April 1985, Lægaard 54176 (AAU!, MO!, QCA!); Volcán Pasochoa, 00°27'S, 078°30'W, 3500 m, 16 September 1985, Lægaard 55273 (AAU!, MO!). s.d., Holmgren 962 (S); Volcán Iliñiza, N-side, 4200-4300 m, 20 June 1985, Lægaard 54557 (QCA!); Volcán Pasochoa, Páramo and forest remains, 3500 m, 16 September 1985, Lægaard 55266; 55269B; 55274 (QCA!); Bosque Protector Pasochoa, 30 km SE de Quito, 2850 m, 29 October 1988, Paz 105 (MO!, QCA!); Faldas del Pasochoa, colección al Sur-Oeste, 3800 m, 03 March 1980, Raza 167 (AAU!); Bosque Protector Pasochoa, 30 km SE de Quito, 2850-3900 m, 17 July 1986, Romoleroux 141 (AAU!, QCA!); 162; 164; 166; 295 (AAU!, QCA!); Pasochoa, 3556 m, 20 August 2008, Romoleroux 5367 (QCA!); Volcán Rumiñahui, 4000 m, 19 December 2008, Salgado 595 (QCA!). Tungurahua: Mocha, Parroquia Pinguili, Reserva Faunístic del Chimborazo, pared de valle en U, 2 km antes de la Base del Lado Oriental del Caryguayrazo, cerro Pocacochas, páramo de Salasaca entrando por Mocha, 01°24'S, 078°45'W, 3860 m, 02 July 1992, Cerón 19203 (MO!). Pilaguin, Sabanza, about 5 km SE of Yatzapuzan, 01°19'S, 078°46'W, 3950-4050 m, 21 December 1982, Brandbyge 42024 (AAU!, MO!); Río Negro, Cordillera de Llanganates, loma between Río Topoand Río Verde Grande, 3.5 km NW of Cerro Hermoso, 01°12'S, 078°18'W, 4000 m, 10 November 1980, Holm-Nielsen 28297 (AAU!); Cordillera de Llanganates, NW of saddle between Río Topo and Río Verde Grande on W slope of Cerro Hermoso, 2.5 km from summit, 01°13'S, 078°18'W, 3950 m, 10 November 1980, Holm-Nielsen 28330 (AAU!); 28371 (AAU!); Cordillera de los Llanganates, at Río Verde Grande at base of Cerro Hermoso, 2 km WSW of the summit, 01°13'S, 078°18'W, 3800 m, 11 November 1980, Holm-Nielsen 28403 (AAU!); 28641 (AAU!); Cordillera de los Llanganates, loma 2.5 km W of Cerro Hermoso, 01°13'S, 078°18'W, 3800 m, 12 November 1980, Holm-Nielsen 28662 (AAU!); Quinuales, 01°13'18"S, 078°18'16"W, 3890 m, 15 February 2010, Romoleroux 5525 (MO!, QCA!, QCNE); 5529 (MO!, QCA!, QCNE); 5533 (MO!, QCA!, QCNE). Río Verde, Cordillera de los Llanganates, at Río Verde Grande at base of Cerro Hermoso, 2.5 km SW of the summit, 01°14'S, 078°18'W, 3700 m, 12 November 1980, Holm-Nielsen 28538 (AAU!); 28599 (AAU!); 28609 (AAU!). San Fernando, Quebrada Huarcusacha, 01°15'S, 078°47'W, 3800-3900 m, 27 January 1983, Brandbyge 42040 (AAU!, MO!); Near Calamaca, app. 20 km W of Ambato along old road Ambato-Guaranda, 01°16'S, 078°48'W, 3400-3900 m, 22 June 1985, Lægaard 54569 (AAU!, QCA!); 54574 (AAU!, MO!, QCA!); 54576 (AAU!, MO!, QCA!). Santiago de Píllaro, Parque Nacional Llanganates, slopes of metamorphic rock, west of Cerro Hermoso, near saddle between headwaters of Río Verde and Río Topo, 01°11'40"S, 078°19'34"W, 3950 m, 12 November 1999, Neill 11944 (MO!, QCNE); 3500 m, s.d., Acosta-Solís 8736 (F!); 4200 m, s.d., Asplund 8464 (S, WIS); Páramo de Yatsaputzan, 3600 m, 05 February 2010, Caranqui 1912 (QCA!); Tamboloma, 3721 m, 29 April 2012, Caranqui 2052 (QCA!); Minza mountain Travesia de Utañag, 3600 m, s.d., Jameson s.n (US!, K); 3400 m, s.d., Penland 420 (F!, US!); Entrando por Baquerizo Moreno hacia el sector de Lagartococha, 3270 m, 01 March 2015, Pérez 8424 (QCA!); Carretera Mocha-Hda. Atillo, 3000 m, 25 April 1988, Romoleroux 567 (AAU!, MO!, QCA!); 570 (AAU!, QCA!); Sendero El Triunfo-Los Llanganates, localidad El Playón, 3600-3800 m, 02 March 1989, Romoleroux 681 (AAU!, MO!, QCA!); 3200 m, s.d., Rimbach 26 (F!); s.d., Stübel 201 (MO!).
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