Xenophyllum humile (Kunth) V.A.Funk, Novon 7(3): 239. 1997.

Calvo, Joel & Moreira-Munoz, Andres, 2020, Taxonomic revision of the Andean genus Xenophyllum (Compositae, Senecioneae), PhytoKeys 158, pp. 1-106 : 1

2. Xenophyllum humile (Kunth) V.A.Funk, Novon 7(3): 239. 1997.

Werneria humilis Kunth, Nov. Gen. Sp. [ed. fol.] 4: 150. 1818. Type. Ecuador. ["in summis Andibus Quitensium" according to the ind. loc.], [without date], F.W.H.A. Humboldt & A.J.A. Bonpland s.n. (lectotype: P-00320181 (digital image!), designated as “holotype” by Funk (1997a: 239); isolectotypes: B-W-16433-01-0 (digital image!), F-972259 (fragment! [almost nothing left]), P s.n.!).

Werneria humilis var. lindenii Wedd., Chlor. Andina 1: 82. 1856 [ “Lindenii”]. Type. Colombia. Tolima: [nevado del] Tolima, limes inferior nivis perpetuae, [without date], J. Goudot s.n. (lectotype: P-02088543 (digital image!), designated here; isolectotype: P-02088544 (digital image!)), syn. nov.

Werneria humilis var. β Wedd., Chlor. Andina 1: 82. 1856, nom. inval. ( Turland et al. 2018, ICN Art. 24.1 and 32.1c).

Werneria lehmannii Hieron., Bot. Jahrb. Syst. 28(5): 647. 1901 [ “Lehmannii”]. nom. illeg. ( Turland et al. 2018, ICN Art. 53.1), replaced name, non W. lehmannii Klatt 1894. Werneria articulata S.F.Blake, Contr. U.S. Natl. Herb. 22: 651. 1924, replacement name. Werneria humilis f. articulata (S.F.Blake) Rockh., Bot. Jahrb. Syst. 70: 294. 1939. Type. Ecuador. Imbabura/Pichincha: páramo de Mojanduleur [Mojanda], Otavalo, 3400-4000 m, [without date], F.C. Lehmann 6230 (lectotype: K s.n.!, designated by Funk (1997a: 238); isolectotype: US-00037320 (right hand specimen, fragment!)).

Werneria humilis var. fontii Cuatrec., Trab. Mus. Nac. Ci. Nat., Ser. Bot. 27: cuadro 22. 1934 [ “Fontii”], nom. nud. ( Turland et al. 2018, ICN Art. 38.1).

Werneria fontii Cuatrec., Trab. Mus. Nac. Ci. Nat., Ser. Bot. 29: 42. 1935 [ “Fontii”]. Xenophyllum fontii (Cuatrec.) V.A.Funk, Novon 7(3): 239. 1997. Type. Colombia. Tolima: vert. merid. mont. Tolima, San Juan del Agua, 4200 m, 15 May 1932, J. Cuatrecasas 2862 (lectotype: MA-246359!, designated as “holotype” by Funk (1997a: 239); isolectotypes: F-844457!, K-000527741 (digital image!), MA-246359b!, MA-246359c!), syn. nov.

Oresigonia brevifolia Willd. ex Rockh., Bot. Jahrb. Syst. 70: 293. 1939, nom. inval. pro syn. ( Turland et al. 2018, ICN Art. 36.1).

Werneria humilis var. angustifolia Cuatrec., nom. nud. in sched. ( Turland et al. 2018, ICN Art. 38.1).

Description.

Suffruticose plant, forming dense mats or hummocks, with rhizome-like stems up to 15 cm long covered with lanate indumentum and old leaves, rather erect, simple or branched from the base. Stems 1-3 cm tall (aerial part), lanate. Leaves stellate-imbricate to subimbricate, extending into a sheath-like base that bears long silky trichomes, usually with the leaf lamina spreading at nearly 90° from the sheath-like base; leaf laminas 2.5-10.8 × 0.5-1.5 mm, linear, obtuse to truncate, usually callous-like tipped at the apex, entire, elliptical to almost terete in cross section, glabrous, 1-nerved above (sometimes barely visible), 1-nerved beneath (sometimes barely visible), fleshy, drying coriaceous, matte, finely papillose. Capitula radiate, erect, sessile to subsessile (rarely with a short peduncle up to 5 mm long). Involucres 4.6-9.3 × 2.9-9 mm, cupuliform; involucral bracts (8-)10 to 13, 3.1-6.1 × 1-2.1 mm, obtuse at the apex, greenish to dark-purplish. Ray florets (8-)10 to 13; corollas 4.2-12.2 × 0.4-2.1 mm, 4 to 7-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets (8-)22 to 39; corollas 3.1-5 mm long, whitish, usually purple-tipped; style branches truncate with a crown of sweeping trichomes, purplish. Achenes 2-2.8 × 0.4-0.6 mm, cylindrical, 7 to 9-ribbed, glabrous; pappus 2.1-7.9 mm long, barbellate, whitish, rarely partially rose-colored. Chromosome number unknown. Figs 2A , 7 .

Additional iconography.

Funk (1997a: 237, fig. 1A); Funk (1997b: 33, fig. 3B, C); Beltrán (2016: 358, fig. 2C, as photo).

Distribution and habitat.

Northwestern Colombia to northern Bolivia. Bolivia (La Paz), Colombia (Antioquia, Caldas, Cauca, Cundinamarca, Huila, Meta, Nariño, Quindío [expected], Risaralda, Tolima, Valle del Cauca), Ecuador (Azuay, Bolívar, Cañar [expected], Carchi, Chimborazo, Cotopaxi, Imbabura, Loja, Morona-Santiago, Napo, Orellana, Pichincha, Sucumbíos, Tungurahua, Zamora-Chinchipe), Peru (Amazonas [n.v.], Ancash, Cajamarca, Junín, La Libertad [expected], San Martín). This species grows in marshes, grasslands, rocky outcrops, and exposed places of the paramo and humid puna ecoregions, between elevations of (2700-)3000-4800 m (Fig. 6 ).

The species is very common along the paramos of Colombia and Ecuador, but it becomes very scattered towards its southern limit. No collections have been studied from the region comprised between central Peru ( Junín) and northern Bolivia (La Paz).

Phenology.

Flowering nearly all year round.

Etymology.

The epithet humile means low-growing, close to the ground.

Notes.

Xenophyllum humile (Kunth) V.A.Funk is a morphologically polymorphic species with rhizome-like stems covered with lanate indumentum and marcescent leaves, obtuse to truncate leaf apex (usually callous-like tipped), (8-)10 to 13 involucral bracts, and (8-)10 to 13 ray florets with white corollas. The leaves are usually bent backwards due to a sort of articulation at the inflexion point between the leaf lamina and the sheath-like base. It is also characterized by its glabrous achenes, although we studied a single specimen from El Cajas (Azuay, Ecuador) displaying achenes with indumentum (Ulloa & Minga 1393, HA).

The species shows high variability in leaf size and density, which led some authors to describe infraspecific taxa or even to propose distinct species. Werneria fontii Cuatrec. was described from Tolima (Cordillera Central, Colombia) mainly on the basis of the compact habit of the plants thriving in this area. These plants are certainly smaller than typical forms and display short leaves with the leaf lamina barely spreading from the sheath-like base. Specimens also collected in the paramos of Tolima were previously used by Weddell (1856) to describe a variety that he characterized as having flesher leaves than typical variety, which is in line with the remarks stated by Cuatrecasas (1935) under W. fontii . However, these small and compact plants are not exclusive from this region. We studied similar forms from Farallones de Cali (Cordillera Occidental, Colombia), Nariño (southern Colombia), Tungurahua (Ecuador), and from Ancash (central Peru). On the other hand, some specimens from the paramos of Chingaza and Sumapaz (Cordillera Oriental, Colombia) display unusual narrow leaves that Cuatrecasas named in sched. as W. humilis var. angustifolia (Cuatrecasas 25628, US-01269081; Cleef 1316, US-01324876). From label information, it can be understood that these plants grow almost submerged in the water, which would explain the singular morphology of these forms. The Bolivian populations, which represent the southern limit of distribution of X. humile , rather exhibit a morphology similar to the typical forms. The puzzling pattern of the different morphologies along the whole distribution area makes unadvisable the recognition of more than a single species.

Xenophyllum humile shows morphological affinities with X. funkianum , X. rigidum , X. roseum , and X. sotarense (see comments under these species). Some specimens from Cundinamarca (Colombia) have the rhizome-like stems covered with a dense lanate indumentum and with scarce leaves along them (e.g., Pedraza & Franco 452, COL; Cleef & Florschütz 5469, COL, US). They might be confused with X. crassum subsp. orientale (Cuatrec.) J.Calvo, although their distributions areas do not overlap and X. humile has shorter and narrower leaves, shorter involucral bracts, and smaller capitula. We also studied a few specimens containing material mixed with Werneria pygmaea Gillies ex Hook. & Arn., a strictly rosettiform species that can be readily differentiated because it grows in lax clumps or solitary plants and its leaves are acute to obtuse at the apex and have the leaf lamina barely spreading from the sheath-like base (vs. leaves obtuse to truncate at the apex or callous-like tipped, usually with the leaf lamina spreading at nearly 90° from the sheath-like base in X. humile ); see Cuatrecasas & Idrobo 27005 (COL) and Soderstrom 1293 (US).

An isolectotype of the name Werneria humilis Kunth is kept at the Willdenow Herbarium. This specimen was sent by Humboldt to Willdenow as stated at the bottom right of the sheet. It is not numbered but contains individuals identical to those from the lectotype at P.

Regarding the name Werneria fontii , it should be noted that the three specimens at MA are numbered with the same herbarium number, i.e., MA-246359. The isolectotypes, therefore, are indicated as MA-246359b and MA-246359c in order to differentiate them from the lectotype.

Additional specimens examined.

Bolivia. La Paz: Murillo, bajando de la cumbre 16.7 km hacia Unduavi, pasando Pongo, 16°19'S, 67°57'W, 28 Apr 1991, S.G. Beck 18750 (LPB, USM); Murillo, entre Pongo y Unduavi, mina 50, subiendo hasta la mina San Luis, 16°17'S, 67°55'W, 28 Oct 1994, S.G. Beck et al. 21519 (LPB); Nor Yungas, San Luis, al NE en línea recta a 3.5 km del poblado de Pongo, 16°18'S, 67°55'W, 13 Nov 2016, F. Zenteno et al. 19280 (LPB). Colombia. Antioquia: páramo Frontino, cerro de Campanas, 6°27'N, 76°7'W, 29 Oct 1976, J.D. Boeke & J.B. McElroy 294 (US); Caldas: páramo del Ruiz, higher slopes, 29 Aug 1957, H.G. Barclay 5238 (COL); Villamaría, Nevado del Ruiz, a orillas de la carretera, 23 Sep 1980, L.A. Camargo 7406 (COL); cordillera Central, vertiente occidental, nevado de Santa Isabel, cabeceras del río Otún, 4°46'N, 75°25'W, 25 Nov 1946, J. Cuatrecasas 23219 (CONC); cordillera Central, vertiente occidental, páramos del nevado del Ruiz, 5 May 1940, J. Cuatrecasas 9283 (US); S side of Nevado el Cisne near laguna Verde, 4°50'N, 75°21'W, 28 Jan 1986, V.A. Funk 8091 (COL, US); S side of Nevado el Cisne near laguna Verde, 4°50'N, 75°21'W, 28 Jan 1986, V.A. Funk 8093 (COL, US); Cordillera Central alrededores del refugio del Ruiz, carretera El Silencio, 4°50'N, 75°22'W, 7 Oct 1978, O. Rangel, H. Sturm & S. Zuluaga 1734 (COL); Manizales, sector del Cisne, 4°51'N, 75°21'W, Oct 2010, W.G. Vargas 22353 (ICESI); Cauca: Puracé, P.N.N. Puracé, entre Pilimbalá y el volcán Puracé, 2°22'N, 76°24'W, 14 Nov 2004, J. Betancur & M. Gutiérrez 11215 (COL); Popayán, volcán Sotará, antecima del volcán, 2°6'N, 76°35'W, 30 Nov 2017, J. Calvo, T. Buira & C. Velasco 7622 (CAUP); cordillera Central, vertiente occidental, cabeceras del río Páez, páramos entre Perro Muerto y la laguna del Páez, 2°59'N, 76°2'W, 4/5 Dec 1944, J. Cuatrecasas 19095 (F); S del P.N. Puracé, inmediaciones de la laguna La Magdalena, sector Valencia, 1°55'N, 76°36'W, 15 Sep 1987, A. Duque 559 (COL); páramo de Las Papas, El Boquerón y alrededores de la laguna La Magdalena, 1°55'N, 76°36'W, 16 Oct 1958, J.M. Idrobo & H.G. Barclay 4039 (COL); Totoró, vereda Santa Teresa, páramo Carga Chiquillo, 2°26'N, 76°23'W, 9 Jul 2002, E.L. Muñoz 915 (CAUP); páramo de Moras, between Mozoco and Pitayó, Tierra Adentro, 2°41'N, 76°14'W, Feb 1906, H. Pittier 1418 (US); Totoró, corregimiento de Gabriel López, trayecto a la laguna la Herradura, 2°30'N, 76°13'W, 17 Apr 2009, B.R. Ramírez & J.F. Álvarez 21060 (CAUP); Jambaló, Pitayó-Quichaya, área especial de manejo Cresta de Gallo, 2°42'N, 76°19'W, 3 Mar 2000, G. Reina 472 (CAUP); Cundinamarca: P.N.N. Sumapaz, Nazaret, Santa Rosa, 4°11'N, 74°11'W, 2 Oct 1999, M. Ángel et al. 9 (COL); Cordillera Oriental al sur de Usme, páramo de Chisacá, near edge of laguna Grande ( Piñuelal), 4°18'N, 74°7'W, 16 Nov 1958, H.G. Barclay & P. Juajibioy 6220 (COL); La Calera, P.N.N. Chingaza, lagunas de Buitrago, 4°45'N, 73°49'W, 24 Jun 2017, L. Camelo, J. Castillo & J. Bernal 205 (COL); Bogotá, localidad de Usme, El Carrizal, alrededores de la laguna El Pato, 4°18'N, 74°9'W, 17 Jan 2013, N. Castaño & A. Orejuela 1429 (COL); páramo de Palacio, lagunas de Buitrago y alrededores, 4°45'N, 73°49'W, 16 Dec 1971, A.M. Cleef 294 (COL, US); páramo de Sumapaz, Chisacá, cabeceras del río S. Rosa, aprox. 1 km al S de la laguna Larga, 4°17'N, 74°12'W, 25 Aug 1972, A.M. Cleef 5277 (US); cabeceras de la quebrada Chuza entre la mina de cal del páramo de Palacio y el páramo de Chingaza, laguna Seca y alrededores, 1 km approx. al NW de la laguna, 4°40'N, 73°47'W, 19 Sep 1972, A.M. Cleef & P.A. Florschütz 5469 (COL, US); macizo de Bogotá, eastern drainage, páramo de Palacio, quebrada de Casarreales, 14 Dec 1959, J. Cuatrecasas, M.T. Murillo & R. Jaramillo 25628 (COL, MA, US); páramo de Chisacá, around the laguna de Chisacá, 4°17'N, 74°12'W, 29 Dec 1959, J. Cuatrecasas & R. Jaramillo 25746 (COL); macizo de Bogotá-macizo de Sumapaz, vertiente oriental, Media Naranja, 4°11'N, 74°15'W, 4 Jan 1969, J. Cuatrecasas & J.M. Idrobo 27005 (COL [mixed with Werneria pygmaea ]); La Calera, páramo Chingaza, laguna Seca, 4°40'N, 73°46'W, 28 Dec 1999, A. Díaz s.n. (COL); páramo de Chisacá, entre las lagunas y Nazareth, 25 Aug 1977, S. Díaz-Piedrahita 1131 (COL); Fómeque, P.N.N. Chingaza, inmediaciones del campamento permanente del área administrativa del parque, sendero Suasié, 12 Nov 2015, D. Giraldo-Cañas et al. 6026 (COL); páramo de Chisacá, laguna Verde, 16 Sep 1952, T. Hammen 506 (COL); Fómeque, páramo de Chingaza, laguna del Medio, 4°30'N, 73°44'W, 15 Dec 1963, G. Huertas & L.A. Camargo 5831 (COL); páramo de Chisacá, 5 Apr 1960, L.E. Mora 1101 (COL); P.N.N. Sumapaz, vereda Santa Rosa, alrededores de la laguna de los Tunjos, 4°16'N, 74°12'W, 7 Aug 1998, P. Pedraza et al. 274 (COL); laguna de Chisacá, 4°17'N, 74°12'W, 13 Dec 1998, P. Pedraza & P. Franco 452 (COL); pico el Zapato, 4°14'N, 74°11'W, 14 Jan 1999, P. Pedraza, C. Pedraza & A. Varón 614 (COL); páramo de Chisacá, alrededores de las lagunas de Chisacá (hoya del río Magdalena), 4°17'N, 74°12'W, 10 Feb 1961, P. Pinto & J. Hernández 485 (COL); Sumapaz, a 2 km de la división de la carretera a San Juan, 3 Oct 1978, O. Rangel 1655 (COL); Pasca, páramo de Chisacá al SE de la laguna de los Tunjos, 4°16'N, 74°12'W, 7 Nov 1987, R. Sánchez 727 (COL); páramo de Siberia, 24 Feb 1952, M. Schneider 1182 (COL); páramo de Chisacá, 4°17'N, 74°12'W, 5 Oct 1966, T.R. Soderstrom 1293 (US [mixed with W. pygmaea ]); macizo de Sumapaz, delante de las lagunas de Chisacá, 4°17'N, 74°11'W, 30 Sep 1963, L. Uribe Uribe 4492 (COL); páramo de Sumapaz, lagunas de Chisacá, 4°17'N, 74°11'W, 2 Dec 1968, L. Uribe Uribe 6206 (COL); Fómeque, P.N.N. Chingaza, lagunas de Buitrago, 4°45'N, 73°49'W, 12 Aug 2000, F. Zapata 66 (COL); Huila: vía al Nevado del Huila, 2°47'N, 76°1'W, 8 Dec 1993, C. Barbosa 9586 (COL); páramo de Las Papas, colinas al SE de la laguna La Magdalena sobre el cerro La Carona y El Boquerón, vertiente del Magdalena, 1°55'N, 76°35'W, 8 Sep 1958, J.M. Idrobo, P. Pinto & H. Bischler 3109 (COL); Meta: páramo de Sumapaz, Cerro Nevado del Sumapaz, pico del Nevado, exposición SE, 3°56'N, 74°6'W, 30 Jan 1972, A.M. Cleef 1316 (US); páramo de Sumapaz, Cerro Nevado del Sumapaz, pico del Nevado, lado NE, 3°56'N, 74°6'W, 11 Jan 1973, A.M. Cleef 7628 (COL, US); macizo de Sumapaz, entre el plano del Nevado y la laguna del Sorbedero, 3°56'N, 74°8'W, 13 Jul 1981, S. Díaz-Piedrahita et al. 2792 (COL); Nariño: Pasto, volcán Galeras, 1°13'N, 77°21'W, 1 Oct 1965, C.E. Acosta 12 (COL); Pasto, volcán Galeras, 1°13'N, 77°21'W, 1 Oct 1965, C.E. Acosta 13 (COL); in monte ignivomo Azufral, 18 May 1876, E. André 3281 (US); Cumbal, volcán de Chiles, en la antena vía El Laurel, Maldonado-Carchi, 0°49'N, 77°55'W, Jul 2012, F. Ávila 2247 (UDBC); Cumbal, volcán de Chiles, en la antena vía El Laurel, Maldonado-Carchi, 0°49'N, 77°55'W, Jul 2012, F. Ávila 2252 (UDBC); Cumbal, volcán de Chiles, en la antena vía El Laurel, Maldonado-Carchi, 0°49'N, 77°55'W, Jul 2012, F. Ávila 2259 (UDBC); Cumbal, volcán de Chiles, en la antena vía El Laurel, Maldonado-Carchi, 0°49'N, 77°55'W, Jul 2012, F. Ávila 2272 (UDBC); Mallama-Piedrahancha, páramo del Infiernillo, km 97 vía Pasto-Tumaco, vereda Pueblo Viejo, entre la cabaña principal de la reserva y el sitio "Las Lagunetas", 1°3'N, 77°46'W, 8 Jan 2009, L.M. Caballero et al. 31 (COL); macizo del volcán Galeras, E slope, around La Torre, 1°12'N, 77°21'W, 7 Feb 1965, J. Cuatrecasas & L.E. Mora 26899 (COL, US); La Cruz, corregimiento de La Estancia, volcán Doña Juana, 1°28'N, 76°55'W, 16 Nov 2001, S.L. Díaz Ibarra 2087 (CAUP); volcán de El Galeras (N slope) above Pasto, 1°12'N, 77°21'W, 18 Oct 1944, J.A. Ewan 16323 (US); camino de herradura entre Túquerres y el volcán Azufral, 8 Jan 1952, A. Fernández & L.E. Mora 1136 (COL); Cumbal, en la base del cerro nevado o volcán, 28 Oct 1955, A. Fernández 2897 (COL); al N de Yacuanquer, 1°9'N, 77°23'W, 4 Jan 1943, M. Garganta 488 (COL); volcán Galeras near Pasto, 1°12'N, 77°21'W, 11 Jan 1981, A. Gentry et al. 30523 (COL); Túquerres, volcán Azufral, E slopes, ca. 9-13 km W of Túquerres, 1°5'N, 77°40'W, 12 May 1989, J.L. Luteyn, J. Fuertes & O. Rangel 12826 (COL, US); Túquerres, volcán Azufral, 1°5'N, 77°40'W, 10 Feb 1962, L.E. Mora 1881a (US); Túquerres, volcán Azufral, 1°5'N, 77°40'W, 10 Feb 1962, L.E. Mora 1883 (US); Túquerres, volcán Azufral, 1°5'N, 77°43'W, 30 Sep 1979, L.E. Mora 7536 (COL); Pasto, P.N.N. Galeras, alrededores del volcán Galeras, 1°15'N, 77°26'W, 2 Sep 1999, P. Pedraza & M. Alvear 667 (COL); Pasto, volcán Galeras, 1°13'N, 77°21'W, 6 Aug 1977, P. Pinto et al. 1829 (COL); volcán Galeras, páramo, 1°13'N, 77°20'W, 22 Oct 1968, T. Plowman 1954 (US); alrededores de Cumbal, 20 Jul 1952, R. Romero-Castañeda 3279 (COL); Pasto, volcán El Galeras, 4 Jun 1946, R.E. Schultes & M. Villarreal 7955 (COL, CONC, F); Cumbal, volcán Chiles, 0°49'N, 77°56'W, 23 Oct 1978, H. Sturm 115 (COL); Túquerres, laguna Verde, 1°5'N, 77°43'W, May 1853, J.J. Triana 2803.1 (COL); Risaralda: Cordillera Central, páramo del Quindío, 4°44'N, 75°23'W, 15 Aug 1922, F.W. Pennell & T.E. Hazen 10020 (US); Tolima: at top of Cordillera Central, Florida-Herrera camino, 3°19'N, 76°2'W, 16 Nov 1944, E.L. Core 1587 (US); Santa Isabel, paso de la Cordillera Central por la quebrada de África, 4°45'N, 75°22'W, 5 Feb 1980, S. Díaz-Piedrahita & R. Jaramillo 1798 (COL); páramo de Ruiz, 16/17 Dec 1917, F.W. Pennell 3039 (NY, US); Valle del Cauca: Los Farallones, ca. filo de la cordillera, al N del cerro Pance, vert. del Pacífico, 3°21'N, 76°42'W, 26 Jul 1991, E. Calderón 43A (COL); alto de Los Farallones, 3°20'N, 76°41'W, 27 Feb 1979, C. Downey 131 (COL). Ecuador. Azuay: P.N. Cajas, cerca de la laguna Toreadora, 2°46'S, 79°13'W, 15 Nov 2000, L. Endara & M. Nonhebel 496 (QCA); P.N. Cajas, ca. 30 km W of Sayausí, at pass NE toward highest peak, 2°46'S, 79°14'W, 23 Oct 1995, V.A. Funk & X. Montezuma 11431 (QCA); P.N. Cajas, ca. 30 km W of Sayausí, at pass NE toward highest peak, 2°46'S, 79°14'W, 23 Oct 1995, V.A. Funk & X. Montezuma 11432 (HA, QCA); P.N. Cajas, ca. 30 km W of Sayausí, at pass NE toward highest peak, 2°46'S, 79°14'W, 23 Oct 1995, V.A. Funk & X. Montezuma 11433 (QCA); 5 km W of Soldados on Cuenca-San Joaquín-Angas rd., ca. 100 m up slopes N of rd. toward large laguna, near laguna Estrellas Cocha, 2°54'S, 79°15'W, 24 Oct 1995, V.A. Funk & X. Montezuma 11442 (QCA); 5 km W of Soldados on Cuenca-San Joaquín-Angas rd., ca. 100 m up slopes N of rd. toward large laguna, near laguna Estrellas Cocha, 2°54'S, 79°15'W, 24 Oct 1995, V.A. Funk & X. Montezuma 11443 (US); 5 km W of Soldados on Cuenca-San Joaquín-Angas rd., ca. 100 m up slopes N of rd. toward large laguna, near laguna Estrellas Cocha, 2°54'S, 79°15'W, 24 Oct 1995, V.A. Funk & X. Montezuma 11444 (US); W of Cuenca on gravel road between Soldados and Balao (hwy 25), laguna Estrellas Cocha, 6 km W of Arch at Soldados Park entrance, 2°54'S, 79°15'W, 18 Apr 2018, V.A. Funk & J.M. Bonifacino 14028 (US); W of Cuenca on gravel road between Soldados and Balao (hwy 25), laguna Estrellas Cocha, 6 km W of Arch at Soldados Park entrance, 2°54'S, 79°15'W, 18 Apr 2018, V.A. Funk & J.M. Bonifacino 14031 (US); carretera Cuenca-Angas, entre Soldados y Angas, 3 Aug 1983, J. Jaramillo & V. Winnerskjold 5497 (QCA); parque de recreación Cajas, 2 Sep 1984, J. Jaramillo 7202 (QCA); P.N. Cajas, road Cuenca-Sayausí-Molleturo, km 38.4, from the pass to the top of ridge north of pass, 2°46'S, 79°14'W, 4 Jan 2000, P.M. Jørgensen, C. Ulloa & E. Narváez 2098 (HA, QCNE); P.N. Cajas, road Cuenca-Sayausí-Molleturo, km 38.4, from the pass to the top of ridge north of pass, 2°46'S, 79°14'W, 4 Jan 2000, P.M. Jørgensen, C. Ulloa & E. Narváez 2117 (HA, QCNE); arriba laguna Dos Chorreras, llamado "upper meadow", 2°46'S, 79°9'W, 2 Jul 1995, B. León & K. Young 3481 (QCA); Cuenca, Sayausí, Tres Cruces, 2°46'S, 79°14'W, 12 Apr 2013, D. Minga & A. Verdugo 2567 (HA); eastern cordillera, between Oña and the río Yacuambi, 3°34'S, 79°5'W, 10/19 Sep 1945, F. Prieto 285 (COL, RB); Cajas N.P., E side of cerro Amarillo, 2°45'S, 79°15'W, 14 Jul 1997, P. Sklenář & V. Sklenářová 2556 (QCA); vicinity of Toreador [laguna Toreadora], between Molleturo and Quinoas, 2°46'S, 79°13'W, 15 Jun 1943, J.A. Steyermark 53216 (RB); P.N. Cajas, Cuenca-Molleturo km 28, sendero Patoquinoas-Totoras, 2°47'S, 79°12'W, 29 Aug 2003, C. Ulloa & D. Minga 1393 (HA); carretera Cuenca-San Joaquín-Angas, entre Soldados y Angas, 2°55'S, 79°15'W, 8 Oct 1990, R. Valencia & J.C. Matheus 395 (QCA); Bolívar: Guaranda, subcuenca del río Chimbo, cerro Tililag, 1°36'S, 78°52'W, 23 Jan 1982, V. Pasaca 1397 (LOJA); Carchi: páramo del Ángel, 21 Aug 1957, H.G. Barclay 5017 (COL); páramo del Ángel, high ridge, 22 Aug 1957, H.G. Barclay 5104 (COL); páramo del Ángel, on high ridge, 23 Aug 1957, H.G. Barclay 5137 (COL); along road from El Chical to Tulcán, at summit of road midway through paramo del Ángel, 0°48'N, 77°56'W, 15 Oct 2012, T.B. Croat et al. 104336 (QCNE); loma Paneíllo, 7.7 km W of Tufiño on road to Maldonado, 0°45'N, 77°50'W, 11 Nov 1988, L.J. Dorr & L.C. Barnett 6035 (QCA, QCNE); roas between Tulcán and Maldonado, S of volcán Chiles, 0°47'N, 77°58'W, 12 Mar 1985, B. Eriksen 59003 (QCA, QCNE); paramo El Ángel, between towns of El Ángel and Tulcán near the pass, 4 Mar 1992, V.A. Funk & M. Gavilanes 11071 (QCA, QCNE); paramo El Ángel, between towns of El Ángel and Tulcán near the pass, 4 Mar 1992, V.A. Funk & M. Gavilanes 11072 (QCA, QCNE); volcán Chiles, rd. from Tulcán to Maldonado, 32 km W of the bridge at the W edge of Tulcán, 5 Mar 1992, V.A. Funk & M. Gavilanes 11075 (QCA, QCNE); volcán Chiles, rd. from Tulcán to Maldonado, 38 km W of the bridge at the W edge of Tulcán, 5 Mar 1992, V.A. Funk & M. Gavilanes 11078 (QCA, QCNE); El Ángel-Tulcán, around laguna El Voladero, 0°42'N, 77°53'W, 9 Aug 1990, P.M. Jørgensen et al. 92321 (COL, QCA, QCNE); páramo El Ángel, laguna S de El Voladero, 0°37'N, 78°26'W, 18 Dec 1990, S. León 1116 (QCA); páramo El Ángel, laguna S de El Voladero, 0°37'N, 78°26'W, 1 Sep 1983, P. Mena 192 (QCA); páramo El Ángel, laguna S de El Voladero, 0°37'N, 78°26'W, [without date], P. Mena 43 (QCA); Espejo, reserva ecológica El Ángel, sitio de lagunas El Voladero, 0°42'N, 77°53'W, 31 Oct 1993, W. Palacios 11636 (QCNE); S slopes of volcán Chiles, 0°49'N, 77°57'W, 21 Oct 1987, P.M. Ramsay & P.J. Merrow-Smith 866 (QCA, QCNE); El Ángel, 19 Aug 1999, M. Smeets & M. Lind van Wijngaarden 736 (QCA, QCNE); Espejo, asociación 23 de Julio, 0°43'N, 77°55'W, 3 Aug 2003, D. Suárez, M. Chinchero & M. Cabascango 1325 (QCNE); Espejo, parroquia La Libertad, humedal Potrerillos, 0°42'N, 77°52'W, 3 Aug 2011, H. Valles & S. Chimbolema 373 (QCA); Chimborazo: eastern slope of Mount Chimborazo, 1°27'S, 78°46'W, 24 Jul 1939, E. Asplund 7819 (LIL); Cordillera Oriental, páramo de la laguna Negra, al N y al E de Alao, 1°46'S, 78°26'W, 11 Aug 1959, H.G. Barclay & P. Juajibioy 8782 (COL); Colta, vía Ambrosio Lazo, El Puyal, 1°41'S, 78°48'W, 2 May 2013, J. Caranqui, W. Haro & F. Salas 2182 (QCA); Alausí, laguna de Ozogoche, 2°13'S, 78°37'W, 10 Jul 2013, J. Caranqui 2439 (QCA); comunidad de Ambrosio Lazo, sector Patococha-Loma Caparina, 1°44'S, 78°53'W, 5 Jun 2009, D. Cárate et al. 609 (QCA); comunidad de Ambrosio Lazo, quebrada Cóndor, sector Yurak Rume, 1°43'S, 78°54'W, 6 Jun 2009, D. Cárate et al. 646 (QCA); comunidad de Ambrosio Lazo, Cóndor Alto, 1°43'S, 78°52'W, 7 Jun 2009, D. Cárate et al. 675 (QCA); nev. Chimborazo, cerca al refugio, 1°29'S, 78°48'W, 13 Jul 2009, D. Cárate et al. 950A (QCA); SW slope of volcano Chimborazo, 1°1'S, 78°47'W, 27 Jun 2012, N. Morueta-Holme, K. Engemann & P. Sandoval 84 (QCA); El Altar, N side of the volcano, on the ridge below the Canoningo peak, 1°41'S, 78°24'W, 19 Aug 1995, P. Sklenář & V. Kostečková 1039 (QCA); cerro Yanaurcu, on the W side of the N ridge of the mountain, 2°14'S, 78°30'W, 29 Oct 1995, P. Sklenář & V. Kostečková 148-17 (QCA); W side of the Chimborazo volcano, 1°28'S, 78°52'W, 5 Jul 1999, P. Sklenář 7508 (QCA, QCNE); Cotopaxi: P.N. Cotopaxi, faldas N de Rumiñahui, 0°40'S, 78°30'W, 28 Oct 1982, H. Balslev, J. Brandbyge & L. Coloma 3347 (QCA); around the Illiniza peaks, 4-10 mi. W from town of Magdalena, 0°39'S, 78°40'W, 2 Apr 1991, R. Bensman 370 (QCNE); entre Aucacocha y Tambo, 31 Dec 1983, J. Jaramillo 6297 (QCA); paramo de Quispicacha, E slope of loma Pucyucuchu, 1°5'S, 78°50'W, 24 Oct 2006, P. Sklenář 9086 (QCA, QCNE); paramo de Quispicacha, E slope of loma Pucyucuchu, 1°5'S, 78°50'W, 23 Oct 2006, P. Sklenář 9125 (QCA); paramo de Quispicacha, summit plateau of loma Pucyucuchu, 1°5'S, 78°50'W, 25 Oct 2006, P. Sklenář 9268 (QCA); Imbabura: Otavalo, reserva ecológica Cotacachi-Cayapas, SW base of Cotacachi mountain, 0°25'N, 78°20'W, 28 Apr 1996, J.L. Clark 2534 (QCNE); laguna de Mojanda, 0°8'N, 78°16'W, 13 Aug 1976, B. Øllgaard & H. Balslev 8749 (US); cúspide cerro Fuya-Fuya, al S de Otavalo, 0°8'N, 78°17'W, 27 Jan 1980, J. Jaramillo & F. Coello 2078 (QCA); Cotacachi, slopes of volcán Cotacachi, 0°35'N, 78°20'W, 11 Oct 1987, P.M. Ramsay & P.J. Merrow-Smith 759 (QCA, QCNE); nevado Cotacachi, on the SW side of the SE ridge of the volcano, 0°21'N, 78°21'W, 9 Sep 1995, P. Sklenář & V. Kostečková 120-1 (QCA); cerro Imbabura, on the E side of the volcano, 0°15'N, 78°10'W, 5 Jun 1995, P. Sklenář & V. Kostečková 31-20 (QCA); Loja: Loma del Oro cerca de Yacuambi, cerca de las lagunas de Condorcillo, 3°34'S, 79°4'W, 25 Oct 1997, R. Bussmann & S. Lange s.n. (HUTPL); Cajanuma, Podocarpus, camino a las lagunas del Compadre, 4°7'S, 79°9'W, 13 Dec 2017, J. Calvo 7681 (HUTPL); Jimbura-Zumba road, km 17, at small lake, 4°44'S, 79°25'W, 27 Jul 1990, P.M. Jørgensen, C. Ulloa & M. Gavilanes 92210 (QCA, QCNE); lagunas N of km 17 on road Jimbura-Zumba, 4°43'S, 79°26'W, 13 Aug 2001, S. Laegaard et al. 21588 (QCNE); Podocarpus N.P., along the trail from Nudo de Cajanuma towards lagunas Compadre, ca. 3-4 km from the refuge, 4°7'S, 79°9'W, 19 Aug 2004, P. Sklenář 8369 (QCA); Morona-Santiago: 24 km E of Gualaceo on rd. to Limón, at pass up rd. to tower, 3°0'S, 78°39'W, 25 Oct 1995, V.A. Funk & R.X. Zamora 11450 (QCA); cerros Yuibug-Pailacajas, E side of the mountain ridge, 1°45'S, 78°27'W, 30 Jul 1997, P. Sklenář & V. Sklenářová 2903 (QCA); W side of the rocky mountain ridge to the E of cerro Yuibug, 1°44'S, 78°27'W, 1 Aug 1997, P. Sklenář & V. Sklenářová 3099 (QCA); Napo: Archidona, Sumaco Napo-Galeras N.P., Sumaco, crater, 0°34'S, 77°38'W, 16 Mar 1996, J.L. Clark 2219 (QCNE); rd. from Quito to Baeza, just at the pass, 0°19'S, 78°12'W, 22 Feb 1992, V.A. Funk et al. 11024 (QCA, QCNE); SW-slope 1.5 km from cerro Quilindaña, 0°47'S, 78°21'W, 1 Apr 1979, L. Holm-Nielsen 16403 (QCA, US); NW slope of Antisana, N of lago Mauca-Machay, 0°26'S, 78°9'W, 2 Nov 1979, L. Holm-Nielsen 20787 (QCA); cordillera de los Llanganates, loma between río Muyu, río Toro and río Verde Grande, 5 km WNW of cerro Hermoso, 1°12'S, 78°19'W, 9 Nov 1980, L. Holm-Nielsen & J. Jaramillo 28223 (QCA); cordillera de los Llanganates, W shoulder of cerro Hermoso, 1.5 km W of the summit, 1°13'S, 78°17'W, 11 Nov 1980, L. Holm-Nielsen & J. Jaramillo 28475 (QCA); cordillera de los Llanganates, shoulder of cerro Hermoso, 1.5 km W of the summit, 1°13'S, 78°18'W, 12 Nov 1980, L. Holm-Nielsen & J. Jaramillo 28722 (QCA); Antisana, Jan 1865, J. Isern 43 (MA); carretera Pifo-Papallacta, páramo de Guamaní, 0°19'S, 78°12'W, 15 Jan 1981, J. Jaramillo 4129 (QCA); páramo de Guamaní, N of paso de La Virgen, 0°17'S, 78°10'W, 10 Jun 1984, S. Laegaard 52249 (QCA, QCNE); Archidona, volcán Sumaco, segunda cumbre cerca al cráter, 0°32'S, 77°37'W, 19 Sep 2012, P. Lozano et al. 79 (QCA); 13.5 km W of Papallacta, 26 Mar 1972, B. MacBryde & J.D. Dwyer 1207 (QCA); Pisayambo, laguna Cochas Negras, 1°6'S, 78°19'W, 14 Jan 1999, B. Merino & Á. Sánchez s.n. (LOJA [mixed with X. crassum subsp. crassum ]); Llanganati, below summit of Pan de Azúcar, 1°9'S, 78°18'W, 15 May 1982, B. Øllgaard et al. 38552 (QCA); Llanganati, ridge between Pan de Azúcar and Las Torres de Llanganati, 1°9'S, 78°17'W, 15 May 1982, B. Øllgaard et al. 38573 (QCA); páramo de Soguillas, near Las Torres de Llanganati, old crater ridge, 1°8'S, 78°15'W, 16 May 1982, B. Øllgaard & L. Holm-Nielsen 38755 (QCA); reserva ecológica Oyacachi, 0°13'S, 78°8'W, 16 Dec 2008, K. Romoleroux, D. Cárate & L.E. López 5355 (QCA); NE side of Antisana, 0°27'S, 78°8'W, 1 Nov 2007, P. Sklenář & E. Rejzková 10707 (QCA); volcán Antisana, W side of the mountain, 0°30'S, 78°10'W, 21 Jul 1997, P. Sklenář & V. Sklenářová 2810 (QCA); NE side of volcán Antisana, 0°27'S, 78°8'W, 17 Aug 1997, P. Sklenář & V. Sklenářová 3403 (QCA); NE side of volcán Antisana, 0°27'S, 78°8'W, 17 Aug 1997, P. Sklenář & V. Sklenářová 3422 (QCA); NE side of volcán Antisana, 0°27'S, 78°8'W, 18 Aug 1997, P. Sklenář & V. Sklenářová 3435 (QCA); NE side of volcán Antisana, 0°27'S, 78°8'W, 18 Aug 1997, P. Sklenář & V. Sklenářová 3538 (QCA); Quijos, reserva ecológica Antisana, páramo de Guamaní, carretera Pifo-Papallacta, La Virgen, 0°20'S, 78°12'W, 24 Jul 1998, H. Vargas, E. Narváez & W. Quizhpe 1936 (QCNE); Tena, P.N. Llanganates, vía Salcedo-Tena, de laguna Chaloa Cocha desvío a Rayo Filo, 0°57'S, 78°23'W, 20 Sep 1998, H. Vargas, E. Narváez & S. Orellana 2679 (QCNE); Orellana: N side of cerro Sumaco, 100 m NW of campsite, 0°32'S, 77°37'W, 24 Apr 1979, L. Holm-Nielsen, J. Jaramillo & T. Vries 17109 (QCA); N side of cerro Sumaco, 100 m NW of campsite, 0°32'S, 77°37'W, 24 Apr 1979, L. Holm-Nielsen, J. Jaramillo & T. Vries 17126 (QCA); N side of cerro Sumaco, upper part of the loma NW of campsite, 0°32'S, 77°37'W, 25 Apr 1979, L. Holm-Nielsen, J. Jaramillo & T. Vries 17204 (QCA); S side of crater of cerro Sumaco, 0°33'S, 77°37'W, 26 Apr 1979, L. Holm-Nielsen, J. Jaramillo & T. Vries 17312 (QCA); Sumaco, 0°32'S, 77°37'W, Jun 1865, J. Isern 313 (MA); S side of cerro Sumaco, 100-200 m S of the main crater, 0°33'S, 77°37'W, 29 Apr 1979, B. Løjtnant & U. Molau 12942 (QCA); Pichincha: vicinity of Quito, Rucu Pichincha, 0°9'S, 78°33'W, 31 Aug 1939, E. Asplund 8612 (LIL); Cordillera Oriental, entre Pifo y el boquerón de cerro Corrales, páramo de Guamaní, 0°19'S, 78°12'W, 15 Aug 1959, H.G. Barclay & P. Juajibioy 8900 (COL); flanco SW del volcán Sincholagua, 0°35'S, 78°21'W, 1 Jun 1985, J. Bosco 145 (QCA); faldas SE volcán Guagua Pichincha, proximidades del refugio, 0°10'S, 78°35'W, 25 May 1985, J. Bosco & M. Marcillo 72B (QCA); entre Pifo y Papallacta, 0°19'S, 78°12'W, 26 Nov 1977, S. Castroviejo, M. Costa & E. Valdés-Bermejo 1053 (MA); Rumiñahui, bosque protector Pasochoa, línea de cumbre del Pasochoa, 0°22'S, 78°27'W, 17 Nov 1990, C.E. Cerón & R. Alarcón 12307 (QCNE); Cayambe, laguna San Marcos, 0°7'N, 77°57'W, 27 Dec 1999, B. Cuamacás & E. Gudiño 473 (QCNE); Quito, parroquia Lloa, sector NE del volcán Guagua Pichincha, 0°10'S, 78°35'W, 12 Aug 2014, D. Fernández et al. 1739 (QCNE); Quito, parroquia Lloa, sector NE del volcán Guagua Pichincha, 0°10'S, 78°35'W, 12 Aug 2014, D. Fernández et al. 1743 (QCNE); Quito, volcán Atacazo, 0°22'S, 78°35'W, 28 Jun 2000, D. Fernández, M. Cerna & P. Villacrés 325 (QCNE); Atacazo, 0°21'S, 78°37'W, Sep 1928, F.G. Firmín 531 (US); road from Cayambe city to Mt. Cayambe, dirt/rock road high on Mt. Cayambe, W slope, 0°0', 78°1'W, 24 Apr 2018, V.A. Funk & J.M. Bonifacino 14094 (US); páramo de Guamaní, carretera Quito-Pifo-Papallacta, 0°23'S, 78°9'W, 20 Oct 1990, E. Guerrón 13 (QCA, QCNE); volcán Cayambe, S slope of the vulcanic cone, valley near road to refugio, 0°3'S, 78°0'W, 18 Jun 1980, L. Holm-Nielsen 24224 (QCA); volcán Cayambe, S slope of the vulcanic cone, near the refugio, 0°2'S, 77°59'W, 18 Jun 1980, L. Holm-Nielsen 24245 (QCA); volcán Iliniza, NE slope below the refugio, 0°39'S, 78°42'W, 13 Aug 1980, L. Holm-Nielsen, B. Øllgaard & C. Sperling 24899 (QCA); volcán Iliniza, NE slope below the refugio, 0°38'S, 78°42'W, 13 Aug 1980, L. Holm-Nielsen, B. Øllgaard & C. Sperling 24959 (QCA); volcán Iliniza, NE slope below the refugio, 0°38'S, 78°42'W, 13 Aug 1980, L. Holm-Nielsen, B. Øllgaard & C. Sperling 24982 (QCA); route de Tufiño à Maldonado, 10 km après Tufiño, 6 Jul 1988, C. Huttel 1385 (QCA); Andium quitensium, crescit in jugis Andium, Dec 1860, W. Jameson s.n. (QPLS); vía Chillogallo-San Juan, partidero desde la población de San Juan hacia faldas del Atacazo "antenas militares", 0°20'S, 78°36'W, 13 Jul 1980, J. Jaramillo & M. Lascano 3142 (QCA); NE Pasochoa, 0°27'S, 78°28'W, 30 Jul 1980, J. Jaramillo, R. Narváez & F. Coello 3168 (QCA); 1 km SE of Cayambe on road to hacienda Piemonte, 10 May 1990, R.M. King, P.M. Peterson & E.J. Judziewicz 10052 (QCA, QCNE); páramo de Guamaní, paso de la carretera Quito-Baeza, 0°19'S, 78°12'W, 25 Aug 1985, B.B. Larsen & B. Dall 190 (QCA); páramo de Guamaní, carretera Pifo-Papallacta, km 27, 0°19'S, 78°12'W, 13 Jan 1991, S. León 1135 (QCA); páramo de Guamaní, carretera Pifo-Papallacta, km 27, 0°19'S, 78°12'W, 13 Jan 1991, S. León 1136 (QCA); in m. Antisana, Sep 1897, L. Mille 469 (QPLS); upper SE slopes of Guagua Pichincha, between the refuge and the crater rim, 0°11'S, 78°36'W, 9 Jan 1988, U. Molau, B. Eriksen & B.B. Klitgaard 2403 (QCA, QCNE); faldas occidentales del volcán Antisana, 0°28'S, 78°12'W, 4 Mar 1984, L. Muñoz 339 (QCA); volcán Cayambe, N slopes, along road to the antenna, 0°5'N, 77°59'W, 9 Jul 1980, B. Øllgaard et al. 34196 (QCA); road Olmedo-laguna San Marcos, E of the pass, 0°7'N, 77°59'W, 10 Jul 1980, B. Øllgaard et al. 34456 (QCA); páramo de Guamaní, laguna de Hoyas, 0°15'S, 78°12'W, 8 Aug 1987, P.M. Ramsay & P.J. Merrow-Smith 186 (QCA, QCNE); páramo de Guamaní, 0°15'S, 78°12'W, 7 Oct 1987, P.M. Ramsay & P.J. Merrow-Smith 726 (QCA, QCNE); W side of a mountain ridge ca. 2 km to the W from cerro SaraUrcu, 0°6'S, 77°57'W, 29 Aug 1995, P. Sklenář & V. Kostečková 100-11 (QCA); W side of a mountain ridge ca. 2 km to the W from cerro SaraUrcu, 0°6'S, 77°57'W, 30 Aug 1995, P. Sklenář & V. Kostečková 111-7 (QCNE); nevado Cayambe, SW slopes left from the road to the refuge, 0°1'N, 78°1'W, 1 Jul 1995, P. Sklenář & V. Kostečková 49-16 (QCA); nevado Cayambe, SW slopes left from the road to the refuge, 0°1'N, 78°1'W, 1 Jul 1995, P. Sklenář & V. Kostečková 51-9 (QCNE); Cotopaxi volcano, on the N side of the mountain, 0°39'S, 78°25'W, 28 Jun 1999, P. Sklenář 7328 (QCA, QCNE); NE slopes of Rucu Pichincha, 0°10'S, 78°34'W, 18 May 1995, P. Sklenář & V. Kostečková 8-1 (QCNE); in pasc. andin. m. Rucu Pichincha, Jul 1871, L. Sodiro 61/3 (QPLS); cerro Atacazo, pendiente N cumbre Atacazo, 0°20'S, 78°36'W, 18 Jun 1983, B. Treiber de Espinosa 101 (QCA); volcán Cayambe, cerca del refugio, 0°0', 78°1'W, 6 Oct 2010, C. Ulloa et al. 2416 (QCA); Sucumbíos: parroquia El Playón de San Francisco, 0°36'N, 77°40'W, 16 Oct 2008, D. Reyes et al. 3688 (QCNE); Tungurahua: páramo del Caryhuayrazo, lado SW, entrando por el arenal del Chimborazo, 1°24'S, 78°48'W, 1 Aug 1988, C.E. Cerón, M. Cerón & G. Viteri 4397 (QCNE); páramo del Caryhuayrazo, lado SW, entrando por el arenal del Chimborazo, 1°24'S, 78°48'W, 1 Aug 1988, C.E. Cerón, M. Cerón & G. Viteri 4408 (QCNE); páramo del Caryhuayrazo, lado SW, entrando por el arenal del Chimborazo, 1°24'S, 78°48'W, 1 Aug 1988, C.E. Cerón, M. Cerón & G. Viteri 4423 (QCNE); Santiago de Píllaro, páramos de Pisayambo, alrededor de la laguna de Pisayambo, 1°5'S, 78°23'W, 9 Oct 1998, E. Cueva 207 (QCNE); cordillera de los Llanganates, Achiriqui, páramo de Jaramillo, 12.7 km NW of cerro Hermoso, 1°9'S, 78°21'W, 7 Nov 1980, L. Holm-Nielsen & J. Jaramillo 28012 (QCA); cordillera de los Llanganates, N end of the loma between río Muyu and río Verde Grande, 5.5 km WNW of cerro Hermoso, 1°12'S, 78°19'W, 9 Nov 1980, L. Holm-Nielsen & J. Jaramillo 28183 (QCA); cordillera de los Llanganates, N end of the loma between río Muyu and río Verde Grande, 5.5 km WNW of cerro Hermoso, 1°12'S, 78°19'W, 9 Nov 1980, L. Holm-Nielsen & J. Jaramillo 28186 (QCA); cordillera de los Llanganates, S side of laguna Verde at cerro Hermoso, 1.8 km from the summit, 1°14'S, 78°18'W, 11 Nov 1980, L. Holm-Nielsen & J. Jaramillo 28423 (QCA); cordillera de los Llanganates, páramo de Jaramillo, 12 km NW of cerro Hermoso, 1°9'S, 78°21'W, 14 Nov 1980, L. Holm-Nielsen & J. Jaramillo 28781 (QCA); páramo de Jaramillo, 2 Nov 1984, J. Jaramillo 7359 (QCA); Santiago de Píllaro, P.N. Llanganates, desde el río Millín hasta la colina Ashpachaca, 1°8'S, 78°22'W, 12 Oct 1998, E. Narváez & W. Quizhpe 302 (QCNE); Santiago de Píllaro, P.N. Llanganates, W of cerro Hermoso, near saddle between headwaters of río Verde and río Topo, 1°11'S, 78°19'W, 12 Nov 1999, D. Neill et al. 11959 (QCNE); Santiago de Píllaro, P.N. Llanganates, SW ridge of cerro Hermoso, 1°13'S, 78°17'W, 14 Nov 1999, D. Neill et al. 12077 (QCNE); Santiago de Píllaro, P.N. Llanganates, SW ridge of cerro Hermoso, 1°13'S, 78°17'W, 14 Nov 1999, D. Neill et al. 12095 (COL, QCNE); Santiago de Píllaro, P.N. Llanganates, SW ridge of cerro Hermoso, 1°13'S, 78°17'W, 14 Nov 1999, D. Neill et al. 12112 (COL, QCA, QCNE); Ambato, ladera NW del cerro Carihuairhazo, 1°23'S, 78°45'W, 2 Feb 2012, K. Romoleroux & G. Peyre 5756 (QCA); P.N. Llanganates, 1°7'S, 78°21'W, 15 Feb 2009, S. Salgado 686 (QCA); volcán Tungurahua, N side of the mountain, trail to the summit, 1°27'S, 78°27'W, 8 Aug 1997, P. Sklenář & V. Sklenářová 3246 (QCA); cerro Hermoso, SW ridge of the mountain, 1°14'S, 78°18'W, 6 Sep 1997, P. Sklenář & V. Sklenářová 3604 (QCA); cerro Hermoso, SW ridge of the mountain, 1°14'S, 78°18'W, 6 Sep 1997, P. Sklenář & V. Sklenářová 3632 (QCA); Patate, P.N. Llanganates, faldas del cerro Pan de Azúcar, en el trayecto páramo de Soguillas-cerro Pan de Azúcar, 1°9'S, 78°17'W, 13 Oct 1998, H. Vargas, J.C. Ronquillo & N. Granda 2837 (QCNE); Patate, P.N. Llanganates, faldas del cerro Pan de Azúcar, en el trayecto páramo de Soguillas-cerro Pan de Azúcar, 1°9'S, 78°17'W, 13 Oct 1998, H. Vargas, J.C. Ronquillo & N. Granda 2838 (QCNE); Patate, P.N. Llanganates, alrededores de la laguna Pan de Azúcar, 1°9'S, 78°17'W, 14 Oct 1998, H. Vargas, J.C. Ronquillo & N. Granda 2869 (QCNE); Zamora-Chinchipe: cordillera de Sabanilla, cerca de la carretera Jimbura-Zumba, alrededor de la laguna Negra, 4°42'S, 79°25'W, 22 Oct 1996, R. Bussmann & S. Lange s.n. (HUTPL, QCNE). Peru. Ancash: cordillera Blanca, quebrada Illanro, nevado Portachuelo, north side to west of glacier, 9°2'S, 77°35'W, 13 Jul 1979, M. Gibby & J.A. Barrett 185 (BM); Carhuaz, Huascarán N.P., quebrada Ulta, below Ulta pass, 9°7'S, 77°31'W, 28 Jul 1985, D.N. Smith 11323 (F, MO, USM); Cajamarca: surroundings of Cajamarca, Jan 1986, B. Becker & F.M. Terrones 303 (LPB); surroundings of Cajamarca, 29 Jan 1986, B. Becker & F.M. Terrones 328 (LPB); Tingo, minas Conga, 6°55'S, 78°21'W, 3 Oct 2005, A. Granda 2491 (MOL); Junín: Jauja, encima de la hacienda Runatullu, 11°39'S, 74°58'W, 23 Apr 1913, A. Weberbauer 6628 (MOL, USM); San Martín: Mariscal Cáceres, valle al oeste del campamento Chochos en el P.N. del río Abiseo, 7°37'S, 77°28'W, 28 Jun 1996, A. Cano et al. 7388 (USM).

Figures

Figure 2. Main habit types in Xenophyllum A, B species forming dense mats or hummocks C, D species forming clumps of somewhat distantly spaced stems A X. humile (Ecuador, Pichincha, pr. Papallacta) B X. sotarense (Colombia, Cauca, Sotara Volcano) C X. juniperinum (Chile, Antofagasta, Aucanquilcha Volcano) D X. digitatum (Bolivia, Potosi, cordillera Kari Kari). Pictures by Joel Calvo.

Figure 6. Distribution map of Xenophyllum acerosum (left hand) and X. humile (right hand).

Figure 7. Xenophyllum humile A habit B stem apical part and capitulum C detail of involucral bract apex D adaxial leaf surface E ray corolla and style F style G disc floret (frontward bristles removed) H disc corolla, stamens, and style (vertically sectioned). All details drawn from Funk & Montezuma 11443 (US) except for A (drawn from Ollgaard & Balslev 8749, US) and G, H (drawn from Funk 8093, US). Illustration by Alice Tangerini.