The genus Begonia (Begoniaceae) in Peru Author Moonlight, Peter. W. Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh, EH 3 5 LR, Scotland, UK. & Botany Department, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland. moonligp@tcd.ie Author Jara-Muñoz, Orlando A. Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá D. C., Colombia. oajaram@unal.edu.co Author Purvis, David A. Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh, EH 3 5 LR, Scotland, UK. dpurvis@rbge.org.uk Author Delves, Jay Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh, EH 3 5 LR, Scotland, UK. j.delves@westernsydney.edu.au Author Allen, Josh P. Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh, EH 3 5 LR, Scotland, UK. myrmeciaman@gmail.com Author Reynel, Carlos Facultad de Ciencias Forestales, Universidad Nacional Agraria-La Molina, Lima 12, Peru. reynel@lamolina.edu.pe text European Journal of Taxonomy 2023 2023-07-18 881 1 334 http://dx.doi.org/10.5852/ejt.2023.881.2175 journal article https://doi.org/10.5852/ejt.2023.881.2175 2118-9773 8178280 49. Begonia acerifolia Kunth in Humboldt et al. (1825) Figs 1C , 2F , 65A , 66 Nova Genera et Species Plantarum (quarto ed.), vol. 7 : 186, t. 644 ( Humboldt et al. 1825 ). – Type : ECUADOR – [ Prov. Loja ] • Loxam ; 1060 m a.s.l. ; A.J.A. Bonpland 3333 ; lectotype : P [ P00679517 ], designated by (2017: 221). de Candolle (1864: 306) ; Smith & Schubert (1952: 36) ; Smith & Wasshausen (1979: 246 , 1986: 48 ); Brako & Zarucchi (1993: 191) ; Moonlight & Fuentes (2022: 39) . Begonia erythrocarpa A.DC., Annales des Sciences Naturelles Botanique, Série 4 11: 121 ( de Candolle 1859 ) . – Type : BOLIVIA – Dept. La Paz : Prov. Larecaja • May 1847 ; H.A. Weddell 4729 ; lectotype: P [ P01900755 ], designated by Tebbitt (2017: 221) ; isolectotypes: G-DC ex P , P [ P01900754 ]. de Candolle (1864: 285) ; Smith & Schubert (1952: 37 , 1955: 114 ); Smith & Wasshausen (1979: 239 , 1986: 38 ); Brako & Zarucchi (1993: 192) ; Wasshausen et al. (2014: 384); Tebbitt (2017: 221) . Begonia dolabrifera C.DC. , Bulletin de L’Herbier Boissier II 2 (8): 324 ( de Candolle 1908 ). – Type : ECUADOR • Pallatanga ; [ 2°01′ S , 78°58′ W ]; Sep. 1891 ; L. Sodiro 594 ; holotype: G [ G00237358 , photo K , F , MO ]; isotypes: F [ V0052627F ], G , P [ P01900768 ], QPLS . Smith & Wasshausen (1979: 246) . Begonia griseocaulis Irmsch., Bibliotheca Botanica 116: 112 ( Irmscher 1937 ) . – Type : ECUADOR – Prov. Chimborazo • Tal des Rio Chanchan , oberhalb Huigra ; [ 2°16′ S , 78°57′ W ]; 1600 m a.s.l. ; 23 Sep. 1933 ; L. Diels 1173 ; holotype: B [ B101068556 ]. Smith & Wasshausen (1979: 244) ; Tebbitt (2017: 221) . Begonia pennellii L.B.Sm. & B.G.Schub., Publications of the Field Museum of Natural History, Botanical Series 13 (4/1): 186 ( Smith & Schubert 1941a ). – Type : PERU – Cusco Region • Above “Pillahuata” , Cerro de Cusilluyoc ; [ 13°10′ S , 71°36′ W ]; 2800–3100 m a.s.l. ; F.W. Pennell 14126 ; holotype: GH [ GH00068262 ]; isotypes: B [ B100365224 ], PH [ PH00007794 ]. Smith & Schubert (1950: 86 , 1955: 114 ); Tebbitt (2017: 221) . Begonia triramosa Irmsch., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 74: 613 ( Irmscher 1949 ) . – Type : ECUADOR – [ Prov. Chimborazo ] • Vicinity of Huigra , mostly on the Haciendade Licay ; [ 3°10′ S , 78°59′ W ]; 3 Sep. 1918 ; J.N. Rose & G. Rose 22493b ; holotype: US [ US00115477 , photo NY ]. Smith & Wasshausen (1979: 244 , 1986: 43 ); Quintana & León-Yáñez (2011: 201) ; Tebbitt (2017: 221) . Begonia pennellii subsp. lobato-ovata Irmsch ., Botanische Jahrbücher für Systematik,Pflanzengeschichte und Pflanzengeographie 776 (1): 85 ( Irmscher 1953 ) . – Type : PERU – Cusco Region : Prov. Calca • Dist. Lares , Lares valley above Mantoc ; [ 12°58′ S , 72°58′ W ]; 2400– 2500 m a.s.l. ; 8 Mar. 1929 ; A. Weberbauer 7906 ; lectotype: B [ B101068564 ], designated here ; isolectotypes: F [illustration B [ B101068565 ]], BM [ BM001191435 ], GH [ GH00068264 ], NY [ NY03091037 ], U [ U1144413 ], US [ US00222073 ]. Smith & Schubert (1950: 86 , 1955: 114 ); Irmscher (1953: 84) ; Golding & Karegeannes (1984: 498) ; Tebbitt (2017: 221) . Begonia pennellii subsp. lobato-ovata var. longiloba Irmsch ., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 76 (1): 85 ( Irmscher 1953 ) . – Type : PERU – Ayacucho Region : Prov. Huanta • Weg von Tambo über Osno zum Flusse Apurimac ; [ 12°44′ S , 73°58′ W ]; 2700–2800 m a.s.l. ; 1 Jun. 1910 ; A. Weberbauer 5622 ; lectotype: B [ B101068561 ], designated here ; isolectotypes: B [ B101068562 ], F [photo K , P , illustration B [ B101068563 ]], NY . Golding & Karegeannes (1984: 498) ; Tebbitt (2017: 221) . Begonia pennellii subsp. lobato-ovata var. longiloba forma macrantha Irmsch., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 76 (1): 86 ( Irmscher 1953 ). – Type : PERU – Píura Region : Prov. Ayabaca • Westabhänge der Anden über Frias ; [ 4°56′ S , 79°54′ W ]; 2900–3000 m a.s.l. ; 20 May 1912 ; A. Weberbauer 6402 ; holotype: B [ B100243083 ]. Golding & Karegeannes (1984: 498) ; Tebbitt (2017: 221) . Begonia lobato-peltata Irmsch., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 76 (1): 86 ( Irmscher 1953 ). – Type : PERU – Puno Region : Prov. Sandia • [ 14°18′ S , 69°27′ W ]; 2100–2300 m a.s.l. ; 10 Jul. 1903 ; A. Weberbauer 503 ; lectotype: B [ B100243008 ], designated by Tebbitt (2017: 222) ; isolectotypes: B [ B100243009 ], G . Smith & Wasshausen (1979: 244) . Begonia macbrideana Irmsch., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 76 (1): 87 ( Irmscher 1953 ). – Type : PERU – Junín Region : Prov. Tarma • Entre Huacapistana y Palca ; [ 11°14′ S , 75°32′ W ]; 2200 m a.s.l. ; A. Weberbauer 2011 ; lectotype: B [ B101068557 ], designated here ; isolectotypes: B [ B101068558 , B101068559 , B101068560 ]; MOL [ MOL00002991 ]. Smith & Wasshausen (1984: 468) ; Tebbitt (2017: 221) . Begonia xerophyta L.B.Sm. & Wassh., Phytologia 44 (4): 245 ( Smith & Wasshausen 1979 ). – Type : ECUADOR – Prov. Loja • Between Loja and San Lucas , very steep and dry slope ; ca 2100 m a.s.l. ; [ 3°45′ S , 79°15′ W ]; 8 Oct. 1955 ; E. Asplund 18036 ; holotype: S [ S04-762 ]. Smith & Wasshausen (1986: 44) ; Quintana & León-Yáñez (2011: 201) ; Tebbitt (2017: 221) . Etymology Named for the superficial resemblance of the species’ leaves to those of maples ( Acer L., Sapindaceae ). Selected specimens examined PERU – Piura Region : Prov. Ayabaca • Montaña de Cuytas , 8 km NE of Ayabaca ; 4°32′ S , 79°44′ W ; 2400–2410 m a.s.l. ; A.H. Gentry , C. Díaz & R. Ortiz 75123 ; MO [ MO-1910846 ], USM • Cerro Chacas ; [ 4°36′ S , 79°41′ W ]; 2820 m a.s.l. ; 27 Mar. 2006 ; M. Farfán Sandoval 25 ; HUT • Dist. Ayabaca , Las Lomas to Ayabaca road , above Chinchinpampa ; 4°40′ S , 79°45′ W ; 2710 m a.s.l. ; 28 May 2015 ; M.C. Tebbitt & A. Daza 843 ; MOL . – Prov. Morropón • Dist. Chalaco , Chalaco village , road Chalaco up to emsata ; 5°01′20″ S , 79°47′10″ W ; 2730 m a.s.l. ; 3 Dec. 2002 ; K. Hoenselaar 512 ; U • Dist. Chalaco , Vista Alegre , Bosque Mija ; 5°04′07.2″ S , 79°43′07.7″ W ; 2920 m a.s.l. ; I. Sánchez V. , A. Cordova A. 12146 ; CPUN . – Prov. Huancabamba • [ 5°22′ S , 79°34′ W ]; 1 Nov. 1876 ; É.F. André 4396 ; K [2], NY • Above Canchaque on the Huancabamba pass ; 5°22′39″ S , 79°33′32″ W ; 2371 m a.s.l. ; 27 Jan. 2016 ; P.W. Moonlight & A. Daza 120 ; E [ E00885880 ], MO , MOL • Cerro Porculla 3–11 km W of Continental Divide down W slope of Cerro ; [ 5°52′ S , 79°31′ W ]; 12 Jun. 1966 ; G. Edwin , J. Schunke V. 3763 ; US [ US00222015 ], USM . – Amazonas Region : Prov. Chachapoyas • Ladera del Cerro Tambo Viejo , road between Balsas and Leymebamba ; 6°47′ S , 77°56′ W ; 2810 m a.s.l. ; 24 May 2015 ; M.C. Tebbitt & A. Daza 831 ; E , MOL . – Cajamarca Region : Prov. Cutervo • Gruta Blanca , Parque Nacional San Andrés ; [ 6°14′ S , 78°45′ W ]; 2400 m a.s.l. ; 24 Feb. 1985 ; S. Llatas Q. 1170 ; US [ US00424990 ]. – Prov. Chota • Dist. La Paccha , Rajopampa ; [ 6°26′ S , 78°45′ W ]; 2600 m a.s.l. ; 21 Jul. 1993 ; J. Cabanillas S. 756 ; CPUN • Near Las Palmas , ca 24km NE of Chota ; 6°29′25″ S , 78°37′25″ W ; 2789 m a.s.l. ; 17 Apr. 1993 ; M.O. Dillon , I. Sánchez V. & M. Sánchez M. 6330 ; CPUN , F • Cuchuli-Chetilla , Chalamarka ; [ 6°30′ S , 78°30′ W ]; 2600 m a.s.l. ; 2 Aug. 2011 ; L. García Llatas 7913 ; HUT . – Prov. Santa Cruz • Dist. Pulan , El Cedro ; [ 6°46′ S , 78°55′ W ]; 2600 m a.s.l. ; 31 Jan. 2008 ; L. Santa Cruz 2106 ; USM • Dist. Catache , upper río Zaña valley , ca 5 km above Monte Seco on path to Chorro Blanco ; [ 6°51′ S , 79°06′ W ]; 1500–2000 m a.s.l. ; 16–18 Mar. 1986 ; M.O. Dillon , A. Sagastegui A. , D. Dillon , P. Alcorn , J. Santisteban , S. Leiva , C. Téllez & M. Guzmán 4390 ; MO [ MO-1643551 ] . – Prov. Hualgayoc • Monte Seco , 6°47′33″ S , 78°30′10″ W ; Aug. 1949 , J.J. Soukup 3876 ; COL , US [ US002222016 ]. – Prov. San Miguel • Bosque de Quellahorco , noreste de la localidad de Tongod ; [ 6°43′ S , 78°47′ W ]; 2700 m a.s.l. ; 14 Sep. 1991 ; I. Sánchez V. & A. Briones 5800 ; CUZ • Dist. La Florida , above La Florida ; 6°53′ S , 79°03′ W ; 1787 m a.s.l. ; M.C. Tebbitt & A. Daza 847 ; MOL . – Junín Region : Prov. Tarma • Huacapistana ; [ 11°14′ S , 75°31′ W ]; 2135 m a.s.l. ; Oct. 1943 ; C. Sandeman 4583 ; K , OXF . – Cusco Region : Prov. Calca • Dist. Lares , Suyo ; 12°28′ S , 71°35′ W ; 2634 m a.s.l. ; 16 Jun. 2005 ; L. Valenzuela , E. Suclli , I. Huamantupa , J. Farfán , N. Anaya , H. Coasaca & J. Tito 5654ª ; MO [ MO-1664124 ] • Dist. Yanatile , above intersection of roads coming from Amparaes , Quebrada Honda , and Lares, along road to Amparaes ; 12°58′ S , 72°03′ W ; 3660 m a.s.l. ; M.C. Tebbitt & A. Daza 797 ; E [ E01059298 ], MOL • Road between Calca and Colca , descending to Colca ; 13°01′48″ S , 72°02′10″ W ; 2000–2600 m a.s.l. ; 27 Apr. 2006 ; H. van der Werff , L. Valenzuela , E. Suclli & A. Carazas 20995 ; MO [ MO-2184526 ], US [ US00951225 ]. – Prov. La Convención • Dist. Huayopata , San Luis ; 12°23′ S , 72°44′ S , 1047– 3200 m a.s.l. ; 11 Apr. 2003 ; W. Galiano , P. Núñez , E. Suclli , E. Acurio & A. Rodríguez 4825 ; MO [ MO-492722 ] • Dist. Santa Ana , “Quillabamba” , 12°50′04.5″ S , 72°47′20.8″ W ; 2009 m a.s.l. ; 16 Mar. 2017 ; A. Orejuela , J. Castillo & M. Suarez 2865 ; E [ E01053422 ], USM • Dist. Vilcabamba , Vilcabamba , Pajonal , 13°07′27″ S , 72°58′55″ W ; 3470–3540 m a.s.l. ; 31 May 2002 ; W. Galiano , L. Valenzuela , E. Suclli , I. Huamantupa & F. Carazas 4108 ; MO [ MO-300885 ]. – Prov. Quispicanchi • Dist. Santa Ana , Madre Selva ; 12°53′49″ S , 72°45′02″ W ; 1500 m a.s.l. ; L. Valenzuela , E. Suclli & G. Calatayud 3090 ; MO [ MO-1102916 ] • Marcapata , en los alrededores de la mina cerca de la ciudad de Marcapata , en la carretera Cusco-Maldonado ; 13°25′25″ S , 70°54′16″ W ; 3200 m a.s.l. ; P. Núñez & N. Núñez 8200 ; MO [ MO-1835951 ] • 33 km past the peak on road from Ocongate to Marcapata , just before Opispata ; [ 13°35′ S , 70°59′ W ]; 3110 m a.s.l. ; 22 Jul. 1978 ; J. Aronson & P. Berry 525 ; MO [ MO-1642893 ], US [ US00672874 ]. – Prov. Paucartambo • Entre Esperanza y Pillahuata , km 133 de la carretera Paucartambo-Pilcopata ; [ 13°10′ S , 71°36′ W ]; 2650 m a.s.l. ; 18 Feb. 1990 ; A. Cano & K. Young 3045 ; HUT , USM • Pillahuata , Parque Nacional Manu ; [ 13°10′ S , 71°36′ W ]; 2700–2900 m a.s.l. ; 24 Mar. 1992 ; A. Cano 5272 ; USM • Pilco ; [ 13°11′ S , 71°37′ W ]; 3000 m a.s.l. ; 19 Jan. 1953 ; F. Woytkowski 339 ; MOL . – Prov. Urubamba • Dist. Ollaytaytambo , road from Quillabamba to Ollaytaytambo ; 13°05′07″ S , 72°22′52″ W ; 3173 m a.s.l. ; 7 Aug. 2014 ; P.W. Moonlight & A. Daza 96 ; E [ E00835900 ], MOL • Dist. Machu Picchu , Alcamayo , nacientes del río , 13°09′ S , 72°30′ W ; 2900 m a.s.l. ; I. Huamantuba , J. Farfán & G. Huallparimachi 3227 ; MO [ MO-1102543 ], US [ US00843963 ] • Huiñayhuayna ; [ 13°11′ S , 72°32′ W ]; 2300 m a.s.l. ; C.M. Ochoa 1 ; MO [ MO-1835894 ] • Huiñaihuaina ; [ 13°12′ S , 72°32′ W ]; 3000 m a.s.l. ; 22 Jul. 1948 ; J.C. Vargas Calderón 7243 ; US [ US00222070 ]. – Puno Region : Prov. Carabaya • Dist. Ollanchea , road directly below Ollanchea ; 13°37′ S , 70°28′ W ; 1949 m a.s.l. ; 9 Jan. 2015 ; M.C. Tebbitt & A. Daza 810 ; E [ E01059297 ], MOL . – Prov. Sandia • San Gabán , below Chichacori ; [ 13°49′ S , 70°29′ W ]; 2970 m a.s.l. ; 6 Jun. 1954 ; R.B. Fisher 31 ; BM • 8 km N of Limbani ; [ 14°05′ S , 69°41′ W ]; 3000 m a.s.l. ; 19 May 1942 ; R.D. Metcalf 30543 ; G , MO [ MO-1642892 ], US [ US00222074 ] • Entre Sandia y Cuyucuyo ; [ 14°26′ S , 69°31′ W ]; 3100–3200 m a.s.l. ; 16 May 1966 ; R. Ferreyra 16771 ; MO [ MO-1642864 ], USM [2]. Description Caulescent, rhizomatous herb, to 200 cm high. Rhizome ellipsoid, 7–20 × 0.75–3 cm , branching, with> 1 growing point per branch. Stem erect, branching; internodes to 20 cm long, to 30 mm thick, succulent, becoming woody at maturity, pale green to red or brown, glabrous to tomentose. Stipules persistent on the rhizome, deciduous on the stem, ovate, 15–18 × 8–14 mm , apex acute, opaque, pale green flushed red, glabrous, margin entire, aciliate. Leaves 1–5 per stem, alternate, basifixed or peltate; petiole 5.5–60 cm long, pale green to red, glabrous to tomentose; blade subsymmetric, ovate to orbicular in outline, to 30 × 23 cm , succulent, apex acute, base peltate to cordate, basal lobes overlapping to not overlapping, sinus to 45 mm deep, margin with 6–8 triangular lobes around the lamina, serrulate to serrate, ciliate, upper surface pale green to dark green, occasionally with white patches along the major veins, glabrous to hirsute, lower surface green to red, moderately pubescent, veins peltate or palmate but with one primary vein, 6–9 veined from the base, with 2–4 secondary veins on the larger side, 1–3 secondary veins on the smaller side. Inflorescences 1 per stem, bisexual, axillary, erect, cymose, with up to 3 branches, bearing up to 8 staminate flowers and 8 pistillate flowers, protandrous; peduncle to 53 cm long, green flushed red, glabrous, bracts deciduous, ovate, 11–16 × 9–10 mm , opaque, green to brown, glabrous, apex obtuse, margin entire, aciliate. Staminate flowers : pedicels to 30 mm long, glabrous; tepals 4, spreading, outer 2 ovate to broadly ovate, 21–24 × 18–24 mm , apex obtuse to rounded, white, sometimes flushed green or pink, glabrous, margin entire to serrate, aciliate, inner 2 elliptic to broadly obovate, 7–25 × 8–18 mm , apex acute to rounded, white, sometimes flushed green or pink, glabrous, margin entire, aciliate; stamens 50–100, spreading, yellow, filaments 1–2.5 mm long, fused at the base, anthers obovoid to obovoid-cuboid, 1–2 × ca 1 mm long, dehiscing via lateral slits, connectives not extending, symmetrically basifixed. Pistillate flowers : pedicels to 40 mm long; bracteoles absent; tepals 5, subequal, deciduous in fruit, spreading, elliptic to broadly ovate, 15–19 × 7–17 mm , apex obtuse to rounded, white, sometimes flushed white or pink, glabrous, margin entire, aciliate; ovary body ellipsoid to ovoid, 5–15 × 5–9 mm , pale green flushed pink, glabrous, unequally 3-winged, the largest rectangular 14–24 × 12–21 mm , the smallest rib-like, to 1 mm wide; 3-locular, placentae branches divided, bearing ovules on both surfaces; styles 3, yellow, fused at the base, 5–7 mm long, once-divided, stigmatic papillae in a spirally twisted band. Fruiting pedicel to 40 mm long. Fruit body ovate, to 15 × 19 mm , drying brown, the largest wing expanding to 25 × 25 mm , the smallest expanding to 2 mm wide. Fig. 65. Distribution of the acerifolia group of Begonia sect. Knesebeckia (Klotzsch) A.DC. in Peru and surrounding countries. A . B. acerifolia Kunth (red). B . B. arrogans Irmsch. (red) and B. ludwigii Irmsch. (blue). C . B. monadelpha (Ruiz ex Klotzsch) A.DC. (red) and B. serotina A.DC. (blue). D . B. huancabambae Moonlight sp. nov. (red) and B. wollnyi Herzog (blue). Black lines indicate country borders, grey lines indicate major administrative divisions, blue lines indicate rivers, and shading indicates elevation. Fig. 66. Begonia acerifolia Kunth. A . Habit. B . Leaf , adaxial surface. C . Leaf, abaxial surface. D . Inflorescence. E . Bracts of varying ages, back view. F . Bracts of varying ages, front view. G . Staminate flower, front view. H . Staminate flower, back view. I . Tepals of the staminate flower, back view. J . Androecium, side and front view. Photographs taken by D.A. Purvis (A–H) and P.W. Moonlight (I–L) in the living collections of the Royal Botanic Garden Edinburgh (Accession 20141063, grown from seeds collected as part of P.W. Moonlight & A. Daza 96 ). Proposed conservation assessment Assessed as Least Concern (LC) by Tebbitt (2017) . Typification notes Begonia pennellii subsp. lobato-ovata Irmsch. was described based upon duplicates of the collection A. Weberbauer 7906 held in Chicago and Berlin ( Irmscher 1953: 85 ). No specimen was cited as a holotype , so it is appropriate to designate a lectotype . We designate the duplicate held in Berlin [B101068564] as the lectotype , as it is an excellent and highly complete specimen. The variety B. pennellii subsp. lobato-ovata var. longiloba Irmsch. was described in the same publication based upon duplicates of A. Weberbauer 5622 held in Berlin and Chicago herbaria ( Irmscher 1953: 85 ). There are two excellent sheets of this collection held in Berlin , so we choose one [B101068562] as the lectotype of this name. The protologue of B. Macbrideana Irmsch. cited specimens of the collection A. Weberbauer 2011 held in Berlin herbarium ( Irmscher 1953: 87 ). There are three duplicates of this specimen held in Berlin , so it is appropriate to designate a lectotype form among this material. We choose B101068557 as the lectotype as it is the most complete of the three duplicates. Identification notes Begonia acerifolia is a variable species and the second most common species in Peru with large (> 10 × 10 cm ) leaf laminae with lobate leaves after B. parviflora . The latter species is easy to distinguish from B. acerifolia as a tall (> 2 m high) herb with> 100 flowers per inflorescence (vs up to 16) and leaves with an indumentum that is rough to the touch. Individuals with peltate leaves are unmistakable but care should be taken to ensure candidate specimens with basifixed leaves do not have red flowers ( B. monadelpha ), a ring of trichomes at the apex of the petiole ( B. ludwigii or B. huancabambae sp. nov. ), fewer than 40 stamens per staminate flower ( B. arrogans Irmsch. ), or fruits with subequal wings ( B. wollnyi ). Distribution and ecology Known from Ecuador , Peru , and Bolivia . In Peru , it has been collected in Piura , Amazonas, Cajamarca , Junín , Cuzco , and Puno Regions ( Fig. 65A ). Found in upper and middle montane forest and northwest Peruvian relict montane forest at an elevation of 1500 to at least 3660 m a.s.l. Some collections may have been made as high as 4600 m a.s.l. in elevation, but the stated elevation on the specimen labels is unusually broad. Populations of B. acerifolia vary from fully deciduous to evergreen.