Polylepis ochreata, (Wedd.) Bitter, Bot. Jahrb. Syst.

4. POLYLEPIS OCHREATA (Wedd.) Bitter View in CoL ,

Bot. Jahrb. Syst. 45: 598. 1911. Acaena ochreata Wedd. , Chlor. And. 2: 240. 1855.

TYPE: ECUADOR. Prov. Pichincha, W slopes of Cerro Pichincha , 3600 m, May 1856, Jameson 73 (lectotype, designated by Simpson (1979, p. 28): P; isolectotypes: A!, G!, GH!, US!; photographs in F!, MO!, US!).

Polylepis ochreata var. integra Bitter , Bot. Jahrb. Syst. 45: 598, fig. 4. 1911.

TYPE: ECUADOR. Prov. Imbabura, Volcán Mojanda, Mar. 1901, Sodiro s.n. (holotype: FI not seen; isotype: GOET!).

Polylepis stuebelii Hieron. , Bot. Jahrb. Syst. 21: 313. 1896.

TYPE: ECUADOR. Prov. Napo, E slope of Cerro Quilindana ~ near Bambasacha , 3700 m, Stübel 204 (holotype: B destroyed, photograph in F!, MO!, NY!, US!).

Polylepis subintegra Benoist View in CoL , Bull. Soc. Bot. France 81: 326. 1934.

TYPE: ECUADOR. Prov. Pichincha, W slopes of Cerro Pichincha , Taurichupa , 4000 m, 28 Nov 1930 Benoist 3356 (Holotype: P!).

Trees 4–7 m tall. Leaves strongly congested at the branch tips, imparipinnate with 5–8 pairs of leaflets, obtrullate in outline, 4.3–7.0 X 3.4–4.0 cm; rachises sparsely sericeous, points of leaflet attachment with a tuft of long, straight whitish hairs; stipular sheaths apically acute with spurs, glabrous to sparsely sericeous in the upper surface; leaflets narrowly elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 1.8–2.7 X 0.5–0.7 cm; margin crenate or slightly crenate, coriaceous, apically emarginate, basally unequally cordate; upper leaflet surfaces glabrous or sparsely sericeous with hairs on the midveins; lower leaflet surfaces sparsely sericeous with whitish hairs 0.9–1.2 mm long, most conspicuous on the veins. Inflorescences pendent, 5.8–20.0 cm long, bearing (16–)21–50 flowers; floral bracts 7.6–8.1 mm long, narrowly triangular, densely sericeous on the outer surface; rachises sericeous. Flowers 4.7–8.3 mm diam; sepals 4, ovate, green, densely sericeous outside; stamens 9–15, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.9–2.5 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; 1.7–4.1 X 1.8–5.0 mm, including spines.

Distribution and Habitat — Polylepis ochreata is distributed in the Andes of Ecuador in Nari~ no, and southernmost Colombia. It occurs at 2500–4030 m elevation in humid montane forest habitats. It often co-occurs with P. pauta , with which it hybridizes extensively ( Romoleroux 1996). Locally, these two species also overlap with P. incana , and then triple hybrids have been found ( Fig. 2 View FIG ).

Conservation Status — Polylepis ochreata is preliminarily assessed as LC or NT, based on an area of occupancy (AOO) estimated at 140 km 2 and an extent of occurrence (EOO) of 39,588 km 2 with 24 known locations.

Notes —We argue that this taxon should be reinstated at species level based on its distinctive morphology characters, including having 5 or more leaflet pairs, glabrous to sparsely sericeous leaf rachises and leaflet margins, and 21–50 flowers per inflorescence. Polylepis ochreata is most similar to P. albicans and P. argentea , with which it shares the elliptic leaflet shape, emarginate apices, and subcordate bases of the leaflets. However, the three species differ in number of leaflet pairs, with P. ochreata having 5–8 pairs, P. albicans 4–5, and P. argentea 3. Polylepis ochreata has (16)21–50 flowers per inflorescence, whereas P. albicans has 18–21, and P. argentea 5–6(–9).

Additional Specimens Examined — Colombia. — NARINO ~: Cumbal, Volcán Nevado Cumbal , N- NE slopes (from the Vereda “Las Vuertas”), 05 Mar 1999 – 09 Mar 1999, D. Stanc´ ık 2752 ( COL) ; Vereda Quilizmal, Laguna de Cumbal , 02 Nov 2008, R. Cortés & B. Devia 2550 ( COL) . Ecuador. — AZUAY: W. Jameson s.n. ( MO) ; Along drinkwater-canal on W-side of Atacazo , c. 5 km S of Campamento, 00°20’S 078°38’W, 24 Oct 1984, S. Laegaard 53256 ( AAU) GoogleMaps ; Bol´ıvar: Guaranda. Parroquia Salinas . Recorrido entre los Arrayanes y Pambabuela . Bosque secundario. Bosque Húmedo Montano , 01°22 ' 06"S 079°03 ' 47"W, 10 Feb 2005, H. Vargas L et al. 4696 ( AAU, K, MO, GoogleMaps QCNE, US).— CARCHI: Km 31 west of Tulcán on road to Maldonado, 00°52’N 077°55’W, 21 Jun 1984, C. A. Todzia & J. W. Grimes 2485 (MO); E. Asplund 17037 (S); Cantón Montúfar. Loma El Corazón (Breta~ na), al sureste de Huaca, al este de la Colonia Huaque~ na. R´ıo Minas. Bosque muy húmedo Montano. Bosque primario; suelos volcánicos, 00°35’N 077°42’W, 9 Apr 1989, G. A. Tipaz 35 (AAU, MO); Carretera Tulcán - Tufi~ no - Maldonado, 12 Oct 1986, K. Romoleroux 189 (AAU, QCA); a 33 km de Tulcán, K. Romoleroux 173 (AAU, QCA); Southern slopes of Volcan Chiles, 00°49’S 077°57’W, 21 Oct 1987, P. M. Ramsay & P. J. Merrow-Smith 911 (QCNE); R. Bensman 418 (MO, WIS); Road Tulcan - Maldonado, near Volcán Chiles. Polylepis forest and paramo, 00°48’N 077°56’W, 16 Aug 1985, S. Laegaard 54967D (AAU, QCA), 54967E (AAU, QCA), 54967F (AAU, QCA); Volc. Los Chiles, along road 9 km W of Tufi~ no. Espeletia páramo with a small remain of mountain forest, 00°49’N 077°57’W, 10 Mar 1992, S. Laegaard 101661 (AAU, GOET, QCA); Road Tulcán - Maldonado, near Volcán Chiles. Polylepis forest and paramo, 00°48’N 077°56’W, 16 Aug 1985, S. Laegaard 54966 (AAU, MO, QCA); S slopes of volcan Chiles, 14–16 km W of Tufi~ no on road to Maldonado; 0–1 km S of the road, 00°47’N 077°57’W, 18 Jan 1988, U. Molau et al. 2536 (AAU, GB, MO, QCA); Tulcán, 33.4 km W of Tulcán on road to Maldonado, Páramo de Chiles on Colombia border, 00°48 ' 19"N 077° 53 ' 03"W, 22 Sep 1979, A. H. Gentry & E. Schupp 26342 (AAU, MO, QCA); Camino Tufi~ no, Sitio Agua Hediondas, en la base del Volcán Chiles, l´ ımite con Colombia. Bosque muy húmedo montano. Bosque secundario. Suelos volcánicos. 00°48’N 077°54’W, 08 Nov 1993, W. A. Palacios 11847 (AAU, MO, QCNE); Carretera entre Tulcán y Maldonado, Faldas del Volcán Chiles; punto más alto del cruce de carretera. Bosque de páramo, 00°45’N 077°59’W, 19 May 1991, W. A. Palacios & D. Rubio 7349 (AAU, MO). COTOPAXI: Quebrada Faldiguera. Hcda. El Pongo. cuadrat. 50x 10 m, 00°41’S 078°45’W, 16 Feb 1991, P. M. JØrgensen et al. 93000 (AAU, MO, QCA).— IMBABURA: W slopes of Volcán Cayambe, 27 Jul 1967, B. Sparre 17789 (AAU, S); Laguna Mojanda, camino, foret ˆd’ altitude, 00°08’N 078°15’W, 1 feb 1996, F. Billiet & B. Jadin 6762 (BR, MO); Road from Otovalo to Laguna Mojanda, collections made along road, 00°10’N 078°20’W - 00°15’N 078°20’W, 22 May 1989, J. F. Smith 1992 (MO); Laguna Mojanda, at the southern part of Laguna Negra, forested slopes near the lake, 00°08’N 078°15’W, 29 Jun 1983, J. S. Brandbyge 42200 (AAU, MO, NY, QCA); K. Romoleroux 339 (QCA); V´ıa hacia la laguna de Mojanda, 02 Nov 1987, K. Romoleroux et al. 475 (AAU, QCA); At road Chauasqui - Merced de Buenos Aires, km 20, near pass, 00°33’N 078°17’W, 10 Dec 1984, S. Laegaard 53475 (AAU, MO, QCA); Otavalo, Forested path to Laguna Mojanda (La v´ıa antiguo a Mojanda por el cementario), Paramo, 00° 10 ' 00"N 078° 15 ' 00"W, 31 Dec 00, J.L. Clark 5820 (QCA, US).— LOJA: Paramo of Fierro Urcu SW of Saraguro, 03°43’S 079°19’W, 21 Nov 1996, B. Øllgaard, G.P. Lewis, P. Lozano & B. Merino 2121 (AAU); Road Loja-Saraguro, 8.5 km N of San Lucas, track to Fierro Urco, km 11, vegetation: tussock grass paramo on 30–40 cm deep black humid soil, with Polylepis forest in small declivity, 15 Jan 1998, G. P. Lewis & C. E. Hughes 3804 (AAU, MO); Road Loja-Cuenca, km 50, track to Fierro Urco, km 11, vegetation: Grass paramo with pockets of bosque andino, 03°41’S 079°17’W, 25 Oct. 1996, G. P. Lewis & P. Lozano 2724 (AAU, MO); Road Loja-Saraguro, km 52, track to fierro Urco, km 10. Vegetation: Polylepis forest in small declivity of tussock-grass paramo, 03°42’S 079°18’W, 17 Jan 1997, G.P. Lewis et al. 2932 (AAU, MO); Paramo of Fierro Urcu SW of Saraguro, 03°43’S 079°19’W, 21 Nov 1996, G. P. Lewis et al. 2121 (AAU); J. L. Jaramillo 7332 (QCA); 7335 (GB, QCA); Cerro Chinchilla “Parroquia Celén”, 19 Sep 1984, J. L. Jaramillo 7312 (AAU, GB, MO, NY, QCA); Road San Lucas-Saraguro km 9, turn off to Fierro Urcu, 11 km to the pass, disturbed, previously burnt grass páramo, with patchy Andean forest below. Dry season, 03° 43 ' 03"S 079°19 ' 25"W, 4 Nov 2000, P.M. JØrgensen et al. 2228; Fierro Urco, Saraguro-Loja, Km 12.4 turnoff towards Fierro Urco, Km 23.8,páramo frequently burned, 03°43 ' 10"S 079°19 ' 18"W, 06 Dec 1994, P.M. JØrgensen et al. 1240 (AAU, MO); Fierro Urco, grass paramo 12 km to the left (northbound) from the Panamerican highway, 03°43’S 079°19’W, 09 Jun 1999, P. Sklenár & S. Laegaard 7096 (AAU); Ca 10 km along road to Fierro Urco, in and near a small forest remain with Polylepis sericea , 03°41’S 079°01’W, 9 Aug 1998, S. Laegaard 19109 (AAU); Ca km 12 along paramo road to Fierro Urco, 03°43’S 079°19’W, 6 Sep 1999, S. Laegaard & P. Sklenár 20279 (AAU).— NAPO: Road Quito-Baeza, 7–8 km NW of Laguna de Papallacta (Páramo de Guaman´ı). Flat boggy depression, adjacent bunch grass páramo and Polylepis -forest, 00°19’S 078°08’W, 20 Jul 1976, B. Øllgaard & H. Balslev 8156 (AAU, MO, NY); Paso de Guaman´ı, quebrada, about +-4 km E Paso de Guaman´ı, on road to Papallacta, 26 Mar 1967, B. Sparre 15029 (AAU, S); Paramo de Papallacta, exact locality unknown, 12 Jan 2015, M. Kessler s.n. (Z); 3 km E of Paso de la Virgen on road Pifo - Papallacta, 00°20’S 078°11’W, 02 Jun 1985, S. Laegaard 54448 (AAU, MO, QCA); N-side of Volcán Pichincha above Hacienda Yanacocha, Polylepis forest, 00°07’S 078°34’W, 02 Jun 1985, S. Laegaard 54457 (AAU, MO, QCA). PICHINCHA: Volcán Pichincha, N slopes, road to Hda.Yanacocha from pass on Quito-Nono road, km 7–11.2,rocky banks in scrub forest, 00°07’S 078°33’W, 10 Dec 1991, B. Øllgaard 99187 (AAU); Reserva Yanacocha, Trocha “Inca” 1–600m, 00°06 ' 44"S 078°34 ' 24"W, 11 Jun 2011, C. Ulloa et al. 2171 (MO, QCA); E. Asplund 17103 (S); NW side of Pichincha, F. Fagerlind & P. Wibom s.n. (S); Camino Yanacocha NW of Volcan Pichincha, Montane cloud forest, 00°05’S 078°33’W, 10 Mar 1981, H. Balslev et al.2049 (AAU,MO, NY, QCA); I. Holmgren 664 (S); Yanococha, faldas noroccidentales, 22 Mar 1987, J. L. Jaramillo 9573 (AAU, NY,QCA); 9588 (AAU, QCA); SW-slopes of Volcán Atacazo, Remnants of forest and open parts with tussuck grasses and shrubs, 00°20’S 078°35’W, 28 Oct 1984, J. S. Brandbyge 42837 (AAU, MO, QCA); SW-slopes of Volcán Atacazo, Remnants of forest, mainly Polylepis , Oreopanax , Escallonia , Gynoxys , 00°20’S 078°35’W, 11 Oct 1984, J. S. Brandbyge 42817 (AAU,MO, QCA); 03 Nov 1987, K. Romoleroux 476 (AAU, QCA); 243 (QCA); 305 (QCA); 245 (NY, QCA); Carretera a San Juan - Atacazo, km 1–12, 00°20’S 078°35’W, 02 Sep 1990, K. Romoleroux 1060 (AAU, QCA); 350 (QCA); K. Romoleroux & A. Freire 1495 (AAU, QCA); Carretera Quito-Nanegalito-Santa Ana del Tablón, desv´ıo Had. Yanacocha km 1–10 desde el desvio, 00°07’S 078°34’W, 06 Dec 1992, K. Romoleroux & A. Freire 1495A (AAU); Laguna Grande de Mojanda, 00°08’N 078°16’W, 30 Jun 1988, K. Romoleroux & S. Laegaard657 (AAU,QCA), 658 (AAU,QCA);Yanacocha.Sector La Despensa. Trasecta de 40 X 20, 00°07 ' 52"S 078° 35 ' 06"W, 14 Feb 2009, K. Romoleroux et al. 5413 (MO, QCA); Corazón, 00°31 ' 53"S 078°39 ' 36"W, L. Sodiro s.n. (AAU); Northern slopes of Cerro Corazón, 2–4 km W of along on the road to Hacienda El Pongo, Montane scrub along road, 00°28’S 078°36’W, 13 May 1979, L. B. Holm-Nielsen 18007 (AAU, MO); Volcán Atacazo, W slope, 17 km from San Juan. Polylepis forest, 00°20’S 078°38’W, 25 Aug 1980, L.B. Holm-Nielsen & E. Asanza C. 25115 (AAU), 25148 (AAU), 25169 (AAU); Volcán Atacazo, Luis A.Mille 364 ( US); R. Benoist 4549 (S);West-side of Volcán Atacazo, along drinkwater-canal, 00°20’S 078°38’W, 11 Aug 1984, S. Laegaard 52639 (AAU,MO,QCA);Páramo de Mojanda, at Laguna Negra and S-side of Laguna Grande, 00°08’N 078°16’W, 14 May 1985, S. Laegaard 54316A (AAU, QCA); Nside of Volcán Pichincha above Hacienda Yanacocha, Polylepis forest, 00°07’S 078°34’W, 04 Jun 1985, S. Laegaard 54459 (AAU), 54467 (AAU, MO), 54476 (AAU, MO, QCA); Along drinkwater-canal on W-side of Atacazo, c. 5 km S of Campamento, 00°20’S 078°38’W, 28 Oct 1984, S. Laegaard 53259 (AAU), 53260 (AAU); Páramo de Mojanda. at Laguna Negra and S-side of Laguna Grande, 00°08’N 078°16’W, 14 May 1985, S. Laegaard 54331 (AAU, MO, QCA), 54338 (AAU, MO, QCA), 54343 (AAU, MO, QCA); Along drinkwater-canal on Wside of Volcán Atacazo, 00°20’S 078°38’W, 24 Nov 1985, S. Laegaard 55665 (AAU, GOET, MO, QCA); At highest pass on road Mojanda-Tabacundo, 00°07’N 078°15’W, 4 Aug 2001, S. Laegaard et al. 21538B (AAU); about 500 m S of (above) Laguna Grande de Mojanda, 00°08’N 078°16’W, 30 Dec 1987, U. Molau & B. Eriksen 2294 (AAU, GB, QCA); carretera Quito-San Juan-San José de la Victoria, en la cercan´ıa de San José en vegetación de borde de carretero, 00°17 ' 53"S 078°38 ' 20"W, 24 Dec 1987, V. Zak & J. L. Jaramillo 3265 (AAU,GB,MO); carretera Quito-Nono-Tandayapa,desviación a Yanacocha en la localidad de Guanto-pugro, en la hacienda “ Alto Perú ”, en restos de vegetación andina, estribaciones N.O. del Volcán Pichincha, 00°05’S 078°35’W, 17 Nov 1987, V. Zak & J. L. Jaramillo 2946 (AAU, GB, MO).— TABACUNDO: Lagunas Mojanda, manchas de vegetación arbórea en páramo, suelos de origen volcánico, 00°07’N 078°16’W, 30 Jul 1992 – 31 Jul 1992, W. A. Palacios et al. 10210 (AAU, MO), 10239 (AAU, MO).