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 60. Begonia foliosa Kunth in Humboldt et al. (1825) Figs 1F , 5D , 7B , 76A , 77 Nova Genera et Species Plantarum (quarto ed.), vol. 7 : 183 ( Humboldt et al. 1825 ). – Lepsia foliosa (Kunth) Klotzsch, Gattungen und Arten 1854: 182( Klotzsch 1855 ) . – Type : COUNTRY UNKNOWN • Location unknown; F.W.H.A.v. Humboldt & A.J.A. Bonpland s.n. ; lectotype : P [ P00307185 ], first stage designated by Smith & Schubert (1946c: 191) ; second stage designated by Smith & Wasshausen (1979: 238) ; isolectotypes : B [ BW17572010 ], F ex P [ V0052628F ], P [2: P01900750 , P01900751 ]. Walpers (1858: 899) ; de Candolle (1864: 375) ; Smith (1973: 216) ; Smith & Wasshausen (1986: 12 , 1989: 35 ); Dorr (1999: 258) . Begonia elegans Kunth in Humboldt et al. (1825) , Nova Genera et Species Plantarum (quarto ed.), vol. 7 : 182 ( Humboldt et al. 1825 ). – Casparya elegans (Kunth) Klotzsch, Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 127 ( Klotzsch 1854 ) . – Type : COLOMBIA – [Dept. Cauca ] • Popayán ; [ 12°27′ S , 76°37′ W ]; A.J.A. Bonpland 2082 ; lectotype: P [ P00307183 ], designated here , first stage designated by Dorr (1999: 258) ; isolectotypes : P [ P01900756 , P00307184 ]. de Candolle (1864: 291) ; Smith & Schubert (1946c: 196) ; Smith & Wasshausen (1986: 12) . Begonia fuchsioides Hook. , Curtis’s Botanical Magazine 73: t. 4281 ( Hooker 1847 ). – Tittelbachia fuchsioides (Hook.) Klotzsch, Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 126 Klotzsch 1854 ). – Type : lectotype : t. 4281 in Hook., Curtis’s Botanical Magazine 73: t. 4281 ( Hooker 1847 ), designated here . de Candolle (1864: 291) ; Smith & Schubert (1946c: 197) ; Smith & Wasshausen (1989: 36) ; Dorr (1999: 262) . Begonia miniata Planch. & Linden, Flore des Serres et des Jardins de l’Europe, Sér. I 8: 105 ( Planchon & Linden 1853 ) . – Begonia fuchsioides var. miniata (Planch. & Linden) A.DC., Prodromus Systematis Naturalis Regni Vegetabilis 15 (1): 291 ( de Candolle 1864 ) . – Begonia fuchsioides forma. miniata (Planch. & Linden) Voss , Vilmorin’s Blumengärtnerei 1: 360 ( Voss 1894 ) . – Begonia foliosa var. miniata (Planch & Linden) L.B.Sm. & B.G.Schub. , Caldasia 4 (18): 196 ( Smith & Schubert 1946c ). – Type : lectotype : plate ‘ Begonia miniata ’ in Planch. & Linden, Flore des Serres et des Jardins de l’Europe, Sér. I 8: 105 ( Planchon & Linden 1853 ), designated here . Smith & Wasshausen (1989: 36) ; Brako & Zarucchi (1993: 192) ; Dorr (1999: 261) . Fig. 76. Distribution of Begonia sect. Lepsia (Klotzsch) A.DC. in Peru and surrounding countries. A . B. foliosa Kunth (red). B . B. guaduensis Kunth. (red). Black lines indicate country borders, grey lines indicate major administrative divisions, blue lines indicate rivers, and shading indicates elevation. Begonia multiflora Benth., Plantas Hartwegianas Imprimis Mexicanas 1845: 185 ( Bentham 1845 ) . – Type : COLOMBIA – Dept. Cundinamarca • near the village of Tena , Prov. Bogotá ; [ 4°39′ N , 74°23′ W ]; Sep. 1845 ; A. Hartweg s.n. ; holotype: K [ K000994431 ]. Dorr (1999: 261) . Lepsia microphylla Klotzsch, Gattungen und Arten 1854: 182 ( Klotzsch 1855 ) . – Begonia microphylla (Klotzsch) A.DC., Prodromus Systematis Naturalis Regni Vegetabilis 15 (1): 375 ( de Candolle 1864 ). – Begonia foliosa var. microphylla (Klotzsch) Dorr., Harvard Papers in Botany 4 (1): 261 ( Dorr 1999 ) . – Type : VENEZUELA • “in silvis Humidis” ; Dec.; J.W.K. Moritz 1263 ; lectotype: B [ B100243005 ], designated here , first stage designated by de Candolle (1864: 375) ; isolectotypes: B [ B100243006 ], BM [ BM000832010 ], BR [ BR0000006956646 ], P [ P00482243 ]). Walpers (1858: 899) ; Smith & Schubert (1946c: 188) ; Smith (1973: 216) ; Smith & Wasshausen (1989: 37) . Lepsia poeppigiana Klotzsch, Gattungen und Arten 1854: 247 ( Klotzsch 1855 ) . – Begonia poeppigiana (Klotzsch) A.DC., Prodromus Systematis Naturalis Regni Vegetabilis 15 (1): 376 ( de Candolle 1864 ). – Type : PERU • “ Peruv. Subandin .”, E.F. Poeppig s.n. ; lectotype: B [ B100243078 , F neg. 20823], first stage designated by de Candolle (1864: 376) ; second stage designated by Dorr (1999: 259) ; isolectotype: P [ P00482230 ] • ibid.; E.F. Poeppig 1241 ; syntypes: OXF , G-BOIS , G-DC • [Prov. Huánuco ]: ad Cuchero ; 9°30′ S , 75°56′ W ]; E.F. Poeppig s.n. ; syntype: F [ V0042327F ]. Walpers (1858: 899) ; Smith & Schubert (1941a: 198) ; Smith & Wasshausen (1984: 469) . Begonia jamesoniana A.DC., Annales des Sciences Naturelles Botanique, Série 4 11: 139 ( de Candolle 1859 ). – Type : ECUADOR • “ad Quito” ; 2440 m a.s.l. ; W. Jameson 305 ; lectotype: G-BOIS [ F neg. 8516] designated by Dorr (1999: 259) ; isolectotypes: BM [ BM001191432 ], G , K [ K000536733 ], OXF [ OXF00058702 , OXF00058703 ], TCD [ TCD0005566 ]. de Candolle (1864: 376) ; Smith & Schubert (1946b: 86) . Begonia putzeysiana A.DC., Annales des Sciences Naturelles Botanique, Série 4 11: 139 ( de Candolle 1859 ) . – Begonia foliosa var. putzeysiana (A.DC.) L.B.Sm. & B.G.Schub. , Caldasia 4 (18): 192 ( Smith & Schubert 1946c ). – Type : VENEZUELA – Prov. Trujillo • 1980 m a.s.l.; Apr. 1846 ; Funck & Schlim 821 ; lectotype: G [F neg. 24205], designated by Dorr (1999: 259) . de Candolle (1864: 375) ; Smith (1973: 216) . Begonia foliosa var. rotundata L.B.Sm. & B.G.Schub. , Caldasia 4 (18): 192 ( Smith & Schubert 1946c ). – Type : COLOMBIA – Dept. Santander • vicinity of California , eastern Cordillera ; [ 7°21′ N , 72°57′ W ]; 2100 m a.s.l. ; 11–27 Jan. 1927 ; Killip & A.C. Smith 16761 ; holotype: GH [ GH00068234 ]; isotypes: NY [ NY00118610 ], US [ US00115315 ]. Dorr (1999: 259) . Begonia foliosa var. australis L.B.Sm. & B.G.Schub. , Caldasia 4 (18): 194 ( Smith & Schubert 1946c ). – Type : COLOMBIA – Dept. Antioquia • La Sierra , 18 km N of Medellin ; [ 6°23′ N , 75°35′ W ]; ca 2000 m a.s.l. ; Jan. 1931 ; Archer 1516 ; holotype: US [ US00115314 ]; isotype: MEDEL [ MEDEL000064 ]. Smith & Schubert (1950: 86) ; Brako & Zarucchi (1993: 192) ; Dorr (1999: 262) . Begonia microphylla var. major L.B.Sm., Phytologia 33 (7): 441 ( Smith 1976 ) . – Begonia foliosa var. major (L.B.Sm.) Dorr, Harvard Papers in Botany 4 (1): 260 ( Dorr 1999 ) . – Type : VENEZUELA – Lara State • Dist. Morán , forest along the río Tocuyo , south of Humocaro Alto ; 9°33′ N , 70°02′ W ; 1300 m a.s.l.; 13 Oct. 1974 ; holotype: VEN [ VEN109268 ]; isotype: US [ US00115395 ]. Smith & Wasshausen (1989: 38) . Begonia microphylla Willd. ex Klotzsch (nom. inval.; nom. rej. pro syn. Lepsia microphylla Klotzsch ), Gattungen und Arten 1854: 182 ( Klotzsch 1855 ) . Begonia splendens hort. ex A.DC. (nom. inval.; nom. rej. pro syn. Begonia puyzeysiana A.DC. ), Prodromus Systematis Naturalis Regni Vegetabilis 15 (1): 375 ( de Candolle 1864 ). Begonia foliosa Poepp. ex A.DC. (nom. inval.; nom. rej. pro syn. Begonia puyzeysiana A.DC. ), Prodromus Systematis Naturalis Regni Vegetabilis 15 (1): 376 ( de Candolle 1864 ). Etymology The epithet derives from the Latin word ‘ foliosus ’, meaning ‘leafy’. This refers to the dense foliage of the species. Specimens examined PERU • A. Matthews s.n. ; BM . – Amazonas Region : Prov. Utcubamba • Dist. Bagua Grande , entre Vista Hermosa y Santa Clara , camino a la reserva Bosque Berlín ; 5°53′42″ S , 78°25′56″ W ; 2037 m a.s.l. ; 5 Feb. 2017 ; A. Orejuela , M. Cueva & J. Castillo 2708 ; E [ E01053106 ], USM . – Prov. Bongará • Dist. Cuispes , trail from Cuispes to Catarata Yumbilla ; 5°55′00″ S ; 5°55′38″ W ; 2231 m a.s.l. ; 5 Jul. 2018 ; P.W. Moonlight 1271 ; USM • Yambrasbamba , 1860–2000 m a.s.l. ; [ 5°45′ S , 77°54′ W ]; 2–26 Mar. 1967 ; S.S. Tillett 673-293 ; US [2: US00222364 , US00222365 ], USM • Lower portion of Shipasbamba-Pomacocha trail ; [ 5°53′ S , 77°58′ W ]; 1600–1900 m a.s.l. ; 29 Jun. 1962 ; J.J. Wurdack 1106 ; US [ US00222133 ], USM . – Prov. Chachapoyas • Matthews s.n. ; G , G-BOIS • Viewpoint on trail from Gocta Village to Gocta Falls ; 6°01′57″ S , 77°53′35″ W ; 2150 m a.s.l. ; 2 Jul. 2018 ; P.W. Moonlight 1242 ; USM . – Prov. Rodríguez de Mendoza • Dist. Vista Alegre , entre Vista Alegre y río Salas ; 6°10′ S , 77°19′ W ; 1470– 1525 m a.s.l. ; 30 Jun. 1998 ; I. Sánchez V. , M. Dillon & N. Zapata 9562ª ; CPUN , F , US [ US00673144 ] • 5 km sureste de Achamal , 6°31.85′ S , 77°24.5′ W ; 1400 m a.s.l. ; V. Quipuscoa S. , S. Leiva G. , Y. Díaz V. & M. Starup J. 1955 ; HUT , US [ US00689820 ]. – Cajamarca Region : Prov. Cutervo • Ladera entre San Andrés y la ruta a Santo Tomás , a 1 km de San Andrés ; [ 6°14′ S , 78°43′ W ]; 2100 m a.s.l. ; 12 Oct. 1987 ; I. Sánchez V. 4471 ; CPUN , MO [ MO-2258827 ]. – Huánuco Region : Prov. Huánuco • ad Cuchero ; 9°30′ S , 75°56′ W ]; E.F. Poeppig 1241 ; G-DC , OXF • Near Riachuelo Chontalagua ; [ 9°31′ S , 75°56′ W ]; 815 m a.s.l. ; 16 Oct. 1936 ; Y. Mexia 8298 ; BM , F , G , MO [ MO-2258823 ], NY , U , US [ US00222271 ]. – Prov. Leoncio Prado • Vicinity of Tingo María , north side of bridge over río Chinchao on Huánuco-Tingo María road ; [ 9°30′ S , 75°57′ W ]; 18 Jul. 1963 ; M.E. Mathias & D. Taylor 5905 ; K , MO [ MO-620235 ] • Cayumba , Tingo María ; [ 9°30′ S , 75°57′ W ]; 15 Mar. 1954 ; G.G. Smith & C. Arvalo 307 ; MO [2: MO-2603906 , MO-2603907 ]. Description Caulescent herb, to 2 m high. Stem erect, branching; internodes to 6.5 cm long, to 6 mm thick, succulent, pale green to brown or red, glabrous. Stipules late deciduous, triangular, 2.5–9 × 0.5–1.5 mm , apex acuminate, mucronate, translucent, pale green to brown, glabrous, margin entire, aciliate. Leaves > 5 per stem, alternate, basifixed; petiole 0.3–0.7 cm long, red, glabrous; blade sub-symmetric, elliptic to oblanceolate, to 3.5 × 1.2 cm , succulent, apex acute to obtuse, base cuneate, margin entire to serrulate or serrate, ciliate, upper surface green, glabrous, lower surface pale green, glabrous, veins pinnate, indistinct, with 2–4 secondary veins on the larger side, 1–3 secondary veins on the smaller side. Inflorescences 1–3 per stem, bisexual, axillary, erect, cymose, with up to 6 branches, bearing up to 64 staminate flowers and 32 pistillate flowers, protandrous; peduncle to 3 cm long, green to red, glabrous, bracts deciduous, lanceolate, 1–2.5 × 0.5–1.5 mm , white, translucent, glabrous, apex acuminate, mucronate, margin entire, aciliate. Staminate flowers : pedicels to 6 mm long, glabrous; tepals 4, slightly projecting, outer 2 ovate, 3–6 × 2–3 mm , apex obtuse, white flushed pink, glabrous, margin entire, aciliate, inner 2 oblanceolate, 2.5–5 × 1–2 mm , apex rounded, white flushed pink, glabrous, margin entire, aciliate; stamens 30–50, projecting, yellow, filaments 1–3 mm long, fused into an irregularly branching 2–2.5 mm long column, anthers ovoid, ca 0.25–0.5 × 0.5 mm long, dehiscing via lateral slits, connectives extended to 0.1 mm , symmetrically basifixed. Pistillate flowers : pedicels to 12 mm long; bracteoles 2, positioned directly beneath the ovary, ovate, 2.5–3 × 1.5 mm , apex obtuse, translucent, white, glabrous, margin entire, aciliate; tepals 5, subequal, deciduous in fruit, spreading, ovate, 5–10 × 3–6 mm , apex obtuse, white flushed pink, glabrous, margin entire, aciliate; ovary body ovoid, ca 3 × 2 mm , white flushed pink, glabrous, unequally 3-winged, largest wing triangular 4–5 × 4–5 mm , smallest rib-like ca 0.5 mm wide; 3-locular, placentae branches entire, bearing ovules on both surfaces; styles 3, yellow, free, 2–4 mm long, once-divided, stigmatic papillae in a spirally twisted band. Fruiting pedicel to 15 mm long. Fruit body ovoid, to 9 × 5 mm , drying brown, the largest wing same shape as in ovary, expanding to 12 × 12 mm , the smallest triangular, to 8 × 4 mm , or rib-like, to 2 mm wide. Fig. 77. Begonia foliosa Kunth. A . Habit. B . Stipule , adaxial view. C . Leaf, abaxial view. D . Leaf, adaxial view. E . Staminate flower, side view. F . Staminate flower, front view. G . Largest tepal of the staminate flower. H . Smallest tepal of the staminate flower. I . Androecium, front view. J . Pistillate flower, side view. K . Pistillate flower, front view. L . Cross section of ovary. M . Largest tepal of the pistillate flower. N . Smallest tepal of the pistillate flower. O . Gynoecium, front view. All photographs taken by P.W. Moonlight in the living collections of the Royal Botanic Garden Edinburgh (A–I, Accession 20180920a, grown from seeds collected as part of P.W. Moonlight 1242 ; J–O, Accession 19480286d, of unknown origin). Proposed conservation assessment While only locally common in Peru , B. foliosa is common throughout much of its range. It has an EOO of ca 1 000 000 km 2 , which includes numerous protected areas. We assess B. foliosa as Least Concern (LC). Notes The label of the specimen V. Quipuscoa S., S. Leiva G., Y. Díaz V. & M. Starup J. 1955 states it was collected in Saposoa District, Huallaga Province, in San Martín Region . The GPS and locality description on the label both refer to localities in Amazonas Region . We do not know of any specimens of B. foliosa collected in San Martín . Typification notes In the protologue of B. foliosa , the author cited material of an unknown origin collected by Humboldt and Bonpland ( Humboldt et al. 1825: 183 ). There are three specimens in Paris herbarium (P) that match this description, a single specimen in Berlin (B) and a fragment of one of the Paris specimens in the herbarium of the Field Museum in Chicago (F). Smith & Schubert (1946c) cited the type as in Paris, which constituted an effective first stage lectotypification, ruling out both the Berlin and Chicago specimens as types . The second stage of lectotypification was carried out by Smith & Wasshausen (1979) because they cited material in P that has a photograph in US as the type of B. foliosa . There is only a photograph of one of the three P specimens in US , so this constitutes an unambiguous reference to a single specimen so an effective second stage lectotypification. Kunth described B. elegans Kunth from material collected by Humboldt and Bonpland ( Humboldt et al. 1825: 182 ), and in this publication a locality was specified as ‘La Vega de San Lorenzo, inter Popyan et almaguer’ in modern day Colombia . This description matches the collection A.J.A. Bonpland 2082 , which is represented in P by three collections. Smith & Schubert (1946c) also suggest there is a fragment of the type at F, but we have not seen this material. Dorr (1999) cited a sheet at P with a photograph in US as the holotype of B. elegans but also cited a second specimen in P with a photograph at US as an isotype. It is therefore not possible to determine which of these two specimens Dorr considered the lectotype , but as their citation narrowed the type down to two specimens it constitutes an effective first stage lectotypification. We designate one of these two specimens (P00307183) as the lectotype of B. elegans and therefore affect the second stage of lectotypification. Bentham described B. multiflora Benth. from material collected by A. Hartweg in Bogotá Province “Juxta pagum Tena” ( Bentham 1845: 185 ). There is a specimen in Kew herbarium that was collected by Hartweg and has a label with an English translation of this locality. This specimen is also labelled as B. multiflora and has the citation of the protologue handwritten in Bentham’s handwriting. It is clear that the protologue was referring to this specimen so we consider it the holotype of B. multiflora . In the protologue of B. fuchsioides Hook. , the author cited material collected by Mr. Purdie in the Ocaña mountains of New Granada in modern day Colombia , and material grown by Mr. Veitch in Exeter ( Hooker 1847 : t. 4281). We have seen no material at Kew collected by Purdie, but there is one collection in Kew herbarium identified as B. fuchsioides in J.D. Hooker’s handwriting [K000102452]. The only information on the label of this specimen is that it was of cultivated origin. This may have been grown in Veitch’s collection thus be a syntype of B. fuchsioides but this is unlikely as a stamp shows the sheet did not enter J.D. Hooker’s herbarium until 1867, twenty years after the protologue was published. It is more likely that this specimen was grown in Kew, perhaps from a cutting of Veitch’s material. As we cannot find a specimen that was definitively cited in the protologue, we instead designate the illustration in the protologue as the lectotype of B. fuchsioides and therefore also Tittelbachia fuchsioides (Hook.) Klotzsch. The protologue of B. miniata Planch & Linden cites living material introduced into Brussels by M. Linden directly from Colombia ( Planchon & Linden 1853: 105 ). No herbarium material was cited in the protologue, and we have been unable to locate any specimens collected by Linden in Colombia that could reasonably be considered as original material. Dorr (1999) was also unable to locate any herbarium material suitable for consideration as a potential lectotype . We therefore designate the excellent illustration in the protologue as the lectotype of B. miniata and its combinations. It would be reasonable to argue that the authors of B. miniata did not intend to describe a new species, which would mean B. miniata would be considered an invalid name. This is because the authors express doubt about whether B. miniata is a species or a variety of B. fuchsioides . The authors do however express the strong opinion of M. Putzey, the minister of justice of Brussels and ‘amateur émérite’ of Begonia , that B. miniata cannot be confused with B. fuchsioides . We consider this as proof that the authors did accept B. miniata as a new species, even if they did so under political pressure. Begonia miniata was soon relegated to a variety of B. fuchsioides by A.P. de Candolle (1864: 291) and then to a form by Voss (1894) . Smith & Schubert (1946b: 146) then regonised B. miniata as a variety of B. foliosa . The protologue of Lepsia microphylla Klotzsch cites the collection J.W.K. Moritz 1263 , collected in Merida , Venezuela , but does not specify an herbarium ( Klotzsch 1855: 182 ). There is material of this collection in several European herbaria and an unnumbered specimen at K (K000322986) collected by Moritz that may be a duplicate. In 1864, A.P. de Candolle transferred the species to the genus Begonia as B. microphylla (A.DC.) Klotzsch ( de Candolle 1864: 375 ) and in doing so cited material of the type collection at Berlin, which constitutes an effective first stage lectotypification. Dorr (1999) later transferred this taxon to a variety of B. foliosa and erroneously cited the Berlin material as destroyed, but it survived the war in Irmscher’s personal herbarium. Both specimens at Berlin are excellent, and we designate B100243005 as the lectotype of L. microphylla because it has a good selection of fruits and both staminate and pistillate flowers. Klotzsch described L. poeppigiana Klotzsch using material collected in Peru by Eduard Friedrich Poeppig ( Klotzsch 1855: 247 ). There are two Peruvian collections by Poeppig that match Klotzsch’s description: E.F. Poeppig s.n. made in “Peruv. Subandin” and deposited in Berlin and Paris; and E.F. Poeppig 1241 made in Cuchero and deposited in Oxford, Chicago, and G-BOIS in Geneva, with a further fragment in G-DC in Geneva. Klotzsch usually included full specimen details in his protologues, so if he had cited E.F. Poeppig 1241 , we believe the collection number and detailed locality (i.e., Cuchero) would have been included in the locality. When A.P. de Candolle transferred L. poeppigiana into the genus Begonia in 1864, he cited material of E.F. Poeppig s.n. in Berlin herbarium and herb Petropolis. This constitutes an effective first stage lectotypification of L. poeppigiana . Dorr (1999) later cited the Berlin specimen as the holotype . This constituted an effective second stage lectotypification as Dorr cited F neg. 20823, which unambiguously shows the specimen B100243078. Identification notes Begonia foliosa is easily recognised as a frequently branching glabrous herb with small (< 5 cm long), pinnately nerved leaves. It differs from all other pinnately nerved Peruvian Begonia species in having <4 secondary veins on both sides of the leaf lamina. Distribution and ecology Known from Venezuela , Colombia , and Ecuador . Within Peru , collected in Amazonas, Cajamarca , and Huánuco Regions ( Fig. 76A ). Found in lower and middle montane forest at an elevation of 815– 2231 m a.s.l. Begonia foliosa can grow in relatively seasonal montane forest due to its thick, succulent stem. The species has been collected in flower throughout the year.