Sassea urticae (L.f.) Klotzsch, Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 128 ( Klotzsch 1854 ) Casparya urticae (L.f.) A.DC, Begonia columnaris Benth., Plantas Hartwegianas Imprimis Mexicanas : 131 ( Bentham 1845 ) Sassea columnaris (Benth.) Klotzsch, Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 128 ( Klotzsch 1854 ) Casparya columnaris (Benth.)A.DC. 274 ( de Candolle 1864 Begonia columnaris var. glabra (Klotzsch) L.B.Sm. & B.G.Schub., Publications 187 ( Smith & Schubert 1941a Begonia trachyptera Benth., Plantas Hartwegianas Imprimis Mexicanas : 184 ( Bentham 1845 ) Stiradotheca trachyptera (Benth.) Klotzsch 127 ( Klotsch 1854 Stibadotheca trachyptera (Benth.) Klotzsch, Gattungen und Arten 1854: 251 ( Klotzsch 1855 ) 251 ( Klotzsch 1855 ) Casparya trachyptera (Benth.) A.DC., Prodromus Systematis Naturalis Regni Vegetabili 274 ( de Candolle 1864 Klotzsch, Gattungen und Arten 1854: 248 ( Klotzsch 1855 ) Klotzsch, Gattungen und Arten 1854: 254 ( Klotzsch 1855 ) Casparya columnaris var. glabra (Klotzsch) A.DC., Casparya urticae var. hispida A.DC., Prodromus Systematis Naturalis Regni Vegetabili 274 ( de Candolle 1864 Begonia monticola C.DC. 325 ( de Candolle 1908 Begonia torresii Standl. , Journal of the Washington Academy of Sciences 17: 313 ( Standley 1927 ) Begonia chiriquensis Standl., Annals of the Missouri Botanical Garden 27: 321 ( Standley 1940 ) Begonia urticae var. retusa L.B.Sm. & B.G.Schub. 37 ( Smith & Schubert 1946a ) Begonia glabra Ruiz ex Klotzsch Sassea glabra Klotzsch 127 ( Klotzsch 1854 Sassea glabra Klotzsch Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 127 ( Klotzsch 1854 ) Casparya coccinea Klotzsch Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 127 ( Klotzsch 1854 ) Begonia cucullata Ruiz ex Klotzsch Sassea glabra Klotzsch 254 ( Klotzsch 1855 Begonia columnaris Pav. ex A.DC. Casparya columnaris var. glabra (Klotzsch) A.DC. 274 ( de Candolle 1864 Stibadotheca urticae Klotzsch ex A.DC. Casparya trachyptera (Benth.) A.DC. 274 ( de Candolle 1864 Sassea hoffmanniana Klotzsch ex A.DC. Begonia urticae var. hispida A.DC. 275 ( de Candolle 1864 Begonia longirostris 193 ( Brako & Zarucchi 1993 The genus Begonia (Begoniaceae) in Peru Moonlight, Peter. W. Jara-Muñoz, Orlando A. Purvis, David A. Delves, Jay Allen, Josh P. Reynel, Carlos European Journal of Taxonomy 2023 2023-07-18 881 1 334 7SHXF 1568985 L.f. L. f. [687,930,265,291] Magnoliopsida Begoniaceae Begonia Plantae Cucurbitales 58 59 Tracheophyta species urticae   Figs 21B, 23   Supplementum Plantarum Systematis vegetabilium: 420 ( Linnaeus 1781). –  Sassea urticae(L.f.) Klotzsch, Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin1854: 128 ( Klotzsch 1854). –  Casparya urticae(L.f.) A.DC, Prodromus Systematis Naturalis Regni Vegetabilis15(1): 274 ( de Candolle 1864).  – Type: COUNTRY UNKNOWN • Locality unknown;  J. Mutiss.n.; lectotype: LINN [LINN1125.3], designated here; possible isolectotype: MA[ MA660695].   Smith (1790: t. 45) [as ‘urticaefolia’]; de Candolle (1864: 274); Walpers (1858: 938); Smith & Schubert (1946a: 33, 1952: 36, 1958: 61); Smith (1973: 212); Smith & Wasshausen (1979: 242, 1986: 29, 1989: 18); Vásquez et al.(2005: 112–125); Burt-Utley (2015: 90).     Begonia columnarisBenth., Plantas Hartwegianas Imprimis Mexicanas: 131 ( Bentham 1845). =   Sassea columnaris(Benth.) Klotzsch, Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin1854: 128 ( Klotzsch 1854). =  Casparya columnaris(Benth.)A.DC.(nom. illeg.; later homonym non Klotzsch), Prodromus Systematis Naturalis Regni Vegetabilis15 (1): 274 ( de Candolle 1864). =  Begonia columnarisvar. glabra(Klotzsch) L.B.Sm. & B.G.Schub., Publications of the Field Museum of Natural History, Botanical Series13: 187 ( Smith & Schubert 1941a).  – Type:  ECUADOR• Mountains of Loxa[ Loja];  K.T. Hartweg740; lectotype: K[ K000536702], designated here; isolectotypes: LD[ LD1404657], K, OXF.   de Candolle (1864: 274); Walpers (1858: 938); Smith & Schubert (1946a: 33).     Begonia trachypteraBenth., Plantas Hartwegianas Imprimis Mexicanas: 184 ( Bentham 1845). =  Stiradotheca trachyptera(Benth.) Klotzschorth. var.,  Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin1854: 127 ( Klotsch 1854). =  Stibadotheca trachyptera(Benth.) Klotzsch, Gattungen und Arten1854: 251 ( Klotzsch 1855). =  Casparya trachyptera(Benth.) A.DC., Prodromus Systematis Naturalis Regni Vegetabili15 (1): 274 ( de Candolle 1864).  – Type:  COLOMBIA– [Prov. Cauca]• In sylvis juxta Pitayo, Prov. Popayán; [ 2°45′ N, 76°18′ W];  3200 ma.s.l.; 1843;  K.T. Hartweg1023; lectotype: K[ K000536703], designated here; isolectotypes: K, LD[ LD1418113].   Smith & Schubert (1946a: 33).    Casparya coccinea  Klotzsch, Gattungen und Arten1854: 248 ( Klotzsch 1855).  – Type:  PERU– [ Huánuco Region: Prov. Huánuco]• in Muña [ 9°40′ S, 75°49′ W];  H. Ruiz & J. A. Pavón s.n.; lectotype: B[ Fneg. 20855], designated here.   de Candolle (1864: 274); Walpers (1858: 935); Smith & Schubert (1941a: 187; 1946a: 33).    Sassea glabra  Klotzsch,  Gattungen und Arten1854: 254 ( Klotzsch 1855). =  Casparya columnarisvar. glabra(Klotzsch) A.DC., Prodromus Systematis Naturalis Regni Vegetabili15 (1): 274 ( de Candolle 1864).  – Type:  PERU• 1777–1788;  H. Ruizs.n.; lectotype: HALex Bex herb. Lamberti[ HAL0121733], designated here.   de Candolle (1864: 274); Walpers (1858: 939); Smith & Schubert (1941a: 187, 1946a: 33); Smith & Wasshausen (1989: 18).    Casparya urticaevar. hispidaA.DC., Prodromus Systematis Naturalis Regni Vegetabili15 (1): 274 ( de Candolle 1864).  – Type:  COSTA RICA• Prope Aguacate; 1857;  K. Hoffmann736; lectotype: B[ B100243045, Fneg. 20856], designated here;  COLOMBIA• In montibus ad orient Botogae;  I.F. Holton723; syntype: G-DC; isosyntypes: G-BOIS, K[ K000536700], NY•  In montibus juxta Botatam;  Oct. 1852;  I.F. Holton727; syntype: K[ K000536699]; isosyntypes: G-BOIS, G-DC, NY.   Standley (1937: 747); Smith & Schubert (1946a: 34).    Begonia monticolaC.DC.,   Bulletin de L’Herbier BoissierII8 (5): 325 ( de Candolle 1908).  – Type:  ECUADOR• Montibus Pichincha et Corazon;  A. Sodiro584; lectotype: G[F neg. 24195], designated here; isolectotype: G•  In silva subandina et subtropica, m. Pichincha et Corazon;  A. Sodiro592; syntype: Smith & Schubert (1946a: 34).     Begonia torresiiStandl.,  Journal of the Washington Academy of Sciences17: 313 ( Standley 1927).  – Type:  COSTA RICA– Prov. San José• Las Nubes; [ 9°59′ N, 83°58′ W];  21 Mar. 1924;  P.C. Standley38561; holotype: US[ US00115472].   Smith & Schubert (1946a: 34).    Begonia chiriquensisStandl., Annals of the Missouri Botanical Garden27: 321 ( Standley 1940).  – Type:  PANAMA– Prov. Chiriqui• trail from Cerro Punta to headwaters of río Caldera; [ 8°50′ N, 82°30′ W];  2250–2500 ma.s.l.;  14 Jan. 1939;  P.H. Allen1435; holotype: F[ V0077648F]; isotypes: GH[ GH00068149], MO[ MO-510547], NY[ NY00112265].   Smith & Schubert (1946a: 33).    Begonia urticaevar. retusaL.B.Sm. & B.G.Schub., Caldasia4: 37 ( Smith & Schubert 1946a).  – Type:  COLOMBIA–  Valle Department• Cordillera occidental, Los Farallones, filo de la cordillera, extremo N en el cerro Alto del Buey; [ 3°22′ N, 76°42′ W];  3500–3600 ma.s.l.;  11 Oct. 1944;  J. Cuatrecascas17942; holotype: GH[ GH00068296]; isotypes: COL[2: COL000003009, COL000003010], F[ V0077669F], P[ P01900836], US[ US00169383, US00115488].   Smith & Wasshausen (1989: 18).   Begonia glabraRuiz ex Klotzsch(nom. inval.; nom. rej. pro syn.   Sassea glabraKlotzsch(nom. inval.; nom. nud.)),  Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin1854: 127 ( Klotzsch 1854).    Sassea glabraKlotzsch(nom. inval.; nom. nud.),   Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin1854: 127 ( Klotzsch 1854).    Casparya coccineaKlotzsch(nom. inval.; nom. nud.),   Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin1854: 127 ( Klotzsch 1854).   Begonia cucullataRuiz ex Klotzsch(nom. inval.; nom. rej. pro syn.   Sassea glabraKlotzsch), Gattungen und Arten: 1854: 254 ( Klotzsch 1855).   Begonia columnarisPav. ex A.DC.(nom. inval.; nom. rej. pro syn.   Casparya columnarisvar. glabra(Klotzsch) A.DC.), Prodromus Systematis Naturalis Regni Vegetabilis15 (1): 274 ( de Candolle 1864).   Stibadotheca urticaeKlotzsch ex A.DC.(nom. inval.; nom. rej. pro syn.   Casparya trachyptera(Benth.) A.DC.), Prodromus Systematis Naturalis Regni Vegetabilis15 (1): 274 ( de Candolle 1864).   Sassea hoffmannianaKlotzsch ex A.DC.(nom. inval.; nom. rej. pro syn.   Begonia urticaevar. hispidaA.DC.), Prodromus Systematis Naturalis Regni Vegetabilis15 (1): 275 ( de Candolle 1864).    Begonia longirostrisauct. non Benth.: Brako & Zarucchi, Monographs in Systematic Botany from the MissouriBotanical Garden, 45: 193 ( Brako & Zarucchi 1993).    Etymology Named for the superficial resemblance of the species’ leaves to those of stinging nettles (  UrticaL., Urticaceae).    Selected specimens examined   PERU•  J.A. Pavóns.n.; G, G-BOIS[2], OXF. –  Amazonas Region: Prov. Bongará • Cordillera Colán SE of La Peca; [ 5°34′ S, 78°24′ W];  2280–2410 ma.s.l.;  30 Sep. 1978;  P.J. Barbour3701; MO[ MO-2180412], US[ US00222330] • Dist. Yambrasbamba, Área de Conservación Privada Abra Patricia-Alto Nieva, Abra Patricia; 5°42’39.58–54.99”S, 77°48’30.96–27.60”W;  2658 ma.s.l.;  18 Jul. 2014;  Y.F. Denget al. 1487; USM. – Prov. Bagua• Bosque Berlín, reserva privada cerca Santa Clara; 5°54′08″ S, 78°24′55″ W;  2064 ma.s.l.;  6 Jul. 2018;  P.W. Moonlight1276; USM. –  Prov. Luya• Dist. Camporedondo, Tullanya, Pascana, La Palma; 6°04′35″ S, 78°21′45″ W;  2710 ma.s.l.;  7 Dec. 1996;  R. Vásquez& R. Rojas21967; MO[ MO-2078375], USM • Camporrendo, Tullanga, subiendo del campamento o Paseana “La Pampa”, hacia el Cerro Condorpuna; [ 6°36′ S, 77°53′ W];  2700– 3000 ma.s.l.;  7 Sep. 1989;  C. Díaz& J. Campos3831; MO[ MO-1643487], USM. – Prov. Rodríguez de Mendoza• Dist. Vista Alegre, La Fortaleza camino a las construcciones Chachapoyas; 6°07′10.4– 06′56″ S, 77°18′46.4–19.34″ W;  2020–2393 ma.s.l.;  22 May 2008;  V. Quipuscoa S., L. Cáceres M., I. Treviño Z.& J. Estela R.3943; HUT. – Prov. Chachapoyas• South side of the Molinopampas-Diosan pass; [ 6°10′ S, 77°41′ W];  2700–3100 ma.s.l.;  8 Aug. 1962;  J.J. Wurdack1633; US[ US00222331], USM • Dist. Leymebamba, middle eastern Calla-Calla slopes,  3–5 kmSE of km 422 Leymebamba- Balsas road; [ 6°45′ S, 77°53′ W];  3000–3200 ma.s.l.;  22 Aug. 1962;  J.J. Wurdack1758; G[2], NY, P[ P05586561], US[ US00222329], USM • Dist. Leymebamba, río El Jardín; 6°56′44″ S, 77°41′54″ W;  3090 ma.s.l.;  30 Jun. 2009;  R.W. Bussmann, J. Gruhn, R. Wagster, J.E. Briones, O. Espinoza& W. Espinoza15933; HUT. –  Piura Region: Prov. Huancabamba• Carmen de la Frontera, alturas de Nueva York; [ 5°02′ S, 79°17′ W];  3160 ma.s.l.;  28 Jul. 2006;  A. Cano, W. Mendoza& N. Valencia16825; USM[3] • Cerro Chinguela, a 20 kmde Zapalache; [ 5°08′ S, 79°23′ W];  3100–3200 ma.s.l.;  24 Apr. 2006;  A. Cano, N. Valencia& I. Salinas16488; USM • El Batán (Chinguela-El Carmen); [ 5°06′ S, 79°21′ W];  2200 ma.s.l.;  15 Sep. 1981;  A. Sagástegui A.10233; HUT, MO[ MO-2180410], NY. –  Cajamarca Region: Prov. San Ignacio• San José de Lourdes, Cerro Parco; 5°04′38″ S, 78°53′30″ W;  2250 ma.s.l.;  9 Nov. 2000;  R. Vásquez, J. Campos, G. Calatayud& M. Huamán26606; MO[ MO-1835905], USM. –  Prov. Cutervo• Dist. San Andrés de Cutervo, Parque Nacional de Cutervo, Saucedal pasando por Chorro Blanco; [ 6°09′ S, 78°41′ W];  2250 ma.s.l.;  3 Aug. 1988;  C. Díaz& H. Osores2954; MO[ MO-098014] • Parque Nacional de Cutervo, trail from Chorro Blanco to San Andrés; 6°10′ S, 78°45′ W;  2100–2200 ma.s.l.;  15 Sep. 1991;  A.H. Gentry, C. Díaz& R. Ortiz74851; MO[ MO-098017], USM • Dist. San Andrés de Cutervo, El Pajonal, alrededores las grutas da los Guacharos; [ 6°14′ S, 78°45′ W];  2500 ma.s.l.;  J. Cabanillas S.60; CPUN. – Prov. Chota• a 1 kmde Paraguay( Querocota-La Granja); [ 6°21′ S, 79°04′ W];  2250 ma.s.l.;  7 Aug. 1994;  S. Leiva G., P. Chuna& J. Cadle1394; MO[2: MO-1641563, MO-1641564], NY • Rejopampa, al O del Pueblo de Paccha; [ 6°26′ S, 78°45′ W];  2600 ma.s.l.;  21 Jun. 1993;  I. Sánchez V.6568; CPUN. –  San Martín Region: Prov. Rioja• Bosque Protección Alto Mayo, unnamed trail from Centro de Interpretación Venceremos (km 382.6) to km 381 on highway 5N, Cerretera Fernando Belaúnde Terry; 5°39′57″ S, 77°44′54″ W;  1700–1800 ma.s.l.;  5 Jun. 2010;  J.L. Clark, M. Chocce, L. Clavijo, M. Overstreet& J. Pérez11847; US[ US001862879], USM • Bosque Protección Alto Mayo, Trocha Kovachi; 5°42′02″ S, 7°44′27″ W;  1707 ma.s.l.;  2 Feb. 2016;  P.W. Moonlight& A. Daza154; E[ E00885556], MO, MOL • Dist. Nuevo Cajamarca, cerca poblado Miraflores, Bosque Protección Alto Mayo; 5°54′ S, 77°23′ W;  1420 ma.s.l.;  3 Nov. 1996;  I. Sánchez V.& M.O. Dillon8479; CPUN, F[ V0086776F], US[ US00673159]. – Prov. Mariscal Cáceres• Above Gran Pajaten ruins; 7°S, 77°W,  2950 ma.s.l.;  22 Jul. 1985;  K. Young1250( US[ US00222175] • ibid.;  24 Jul. 1985;  K. Young1292; HUT • Above timberline Puerta del Monte; [ 7°39′ S, 77°29′ W];  22 Nov. 1985;  K. Young2094; MOL, US[ US00222326] • Puerta de Monte, Parque Nacional río Abiseo; 7°38′56.8″ S, 77°24′58.4″ W;  3140 ma.s.l.;  5 Aug. 2000;  A. Cano et al.10840; USM. –  Huánuco Region: Prov. Huánuco• In Muña; [ 9°40′ S, 75°49′ W];  J.A. Pavóns.n.; G-DCex G-BOIS);  Cerros al este del río Huallaga, entre Muña y el tambo de las vacas; [ 9°34′ S, 75°49′ W];  3100 ma.s.l.;  1 Jun. 1913;  A. Weberbauer6723; MOL[3], US[ US00222044] • Muña; [ca 9°40′ S, 75°49′ W; 1785],  H. Ruiz& J.A. Pavóns.n.; MA[2, MA813495, MA813496]. –  Pasco Region: Prov. Oxapampa• Dist. Huancabamba, Parque Nacional Yanachaga-Chemillén, Sector Tunqui, camino hacia María Puñis; 10°16’31.26”S, 75°30’58.72”W;  1895 ma.s.l.;  18–21 Oct. 2008;  M. Cueva, A. Peña, R. Rivera& M. Moens231; HOXA, USM • Dist. Huancabamba, Parque Nacional Yanachaga-Chemillén, Abra Yanachaga; 10°22′46″ S, 75°27′43″ W;  2870–3200 ma.s.l.;  12 Jun. 2003;  R. Vásquez et al.28129; HOXA, MO[ MO-1102451], US[ US000843968] • Parque Nacional Yanachaga-Chemillén, Sector San Alberto, alrededores Refugio El Cedro; 10°32′43″ S, 75°21′30″ W;  2483 ma.s.l.;  26–27 Apr. 2009;  M. Cueva621; HOXA, USM. –  Cusco Region: Prov. La Convención• In deep valley just below Camp 5 and the land-slide; 12°37′ S, 73°32′ W; ca  2700 ma.s.l.;  7 Jul. 1968;  T.R. Dudley10853; US[ US00222325].    Description Caulescent herb, to 50 cmhigh. Stemerect, branching; internodes to 4.5 cmlong, to 2 mmthick, succulent, green to red, tomentose. Stipuleslate deciduous, lanceolate, 4–7 × 2–4 mm, apex obtuse, translucent, green, glabrous, margin entire, aciliate. Leaves> 5, alternate, basifixed; petiole 0.3–2 cmlong, pale green to red, glabrous to tomentose; blade subsymmetric, lanceolate, to 8 × 3 cm, membranaceous, apex acuminate, base dimidiate, the large side rounded, the smaller side cuneate to rounded, margin undulate, serrulate, or irregularly serrate, ciliate, upper surface pale to dark green, glabrous to pilose, lower surface pale green to vivid red, glabrous throughout to glabrous and densely pilose on the major veins, veins pinnate, with 5–10 secondary veins on the larger side, 4–7 secondary veins on the smaller side. Inflorescences1–5 per stem, bisexual, axillary, pendulous erect, cymose, with 2 branches, bearing up to 3 staminate flowers and 2 pistillate flowers, protandrous; peduncle to 3 cmlong, pale green to red, glabrous to tomentose, bracts deciduous, ovate to obovate, 3–6 × 1–4 mm, translucent, white to pale green, glabrous, apex rounded to truncate, margin entire, aciliate. Staminate flowers: pedicels to 12 mmlong, glabrous; tepals 4, spreading to reflexed, outer 2 ovate, 4–8 × 2.5–4 mm, apex rounded, usually pink, orange, or red, rarely white, glabrous, margin entire, aciliate, inner 2 elliptic to obovate, 5–8 × 2–4 mm, apex rounded, the same colour as the other tepals, glabrous, margin entire, aciliate; stamens 6–10, projecting, the same colour as the tepals, filaments 0.5–1 mmlong, free, anthers ellipsoid, 1.5–2 × 0.1 mm, dehiscing via lateral slits, connectives not extending, symmetrically basifixed. Pistillate flowers: pedicels to 5 mmlong; bracteoles lacking; tepals 5, subequal, deciduous in fruit, spreading to reflexed, ovate, 5–6 × 2–4 mm, apex rounded, usually pink, orange, or red, rarely white, glabrous, margin entire, aciliate; ovary body obdeltoid, 2–3 × 3–6 mm, pale green, white, pink, orange, or red, glabrous to tomentose, equally 3-winged, wings horn-shaped, sometimes ascending ca 1.5–2 × 3–6 mm; 3-locular, placentae branches divided, bearing ovules on both surfaces; styles 3, the same colour as the tepals, free, 2–5 mmlong, 2–3 times divided, stigmatic papillae in a spirally twisted band. Fruiting pedicelto 10 mmlong. Fruit bodyovoid, to 6 × 6 mm, drying light brown, wings same shape as in ovary, expanding to 12 × 6 mm, column at the apex of the ovary elongating to 18 mm.    Proposed conservation assessment Locally common and widespread in Peruand as far north as Costa Rica. Its EOO is> 1 900 000 km 2and includes numerous protected areas. We assess  B. urticaeas Least Concern (LC).    Notes Ruiz and Pavón collected a number of specimens of  B. urticaeand it appears from two unpublished illustrations of this species housed in Madrid(MA-AJB04-D-1400, MA-AJB04-D-1398) that they intended to publish some of this material as  B. roseaand  B. coccinea. Ruiz and Pavón never completed an account of the Begoniaceaefor their  Flora Peruvianaet Chilensisso these remain as unpublished names.   Synonymy notes We clarify here the correct author string for the name  B. columnarisvar. glabra. The name  Sassea glabraKlotzschwas first published as a nom. nud. ( Klotzsch 1854) before being published as a valid and legitimate name a year later ( Klotzsch 1855). In 1864, Alphonse Pyramus de Candolle treated this name as a variety of his  C. columnarisas  C. columnarisvar. glabra( de Candolle 1864: 274). It is clear de Candolle based this name upon Klotzsch’s  S. glabrabecause he cited the protologue of this name. Later, Smith & Schubert (1941a: 187)transferred the name  C. columnarisand its varieties into the genus  Begoniabut cited the authorship of  B. columnarisvar. glabraas “(A.DC.) L.B.Sm. & B.G.Schub.”. This is not correct, as the author of the basionym was Klotzsch, so the correct author string should be “  B. columnarisvar. glabra(Klotzsch) A.DC.”.   Fig. 23.  Begonia urticaeL.f. A. Habit. B. Inflorescence, showing staminate and pistillate flowers, red variant. C. Inflorescence, showing staminate flowers and developing fruit, pink variant. D. Fruit. All photographs by P.W. Moonlight from P.W. Moonlight & A. Daza 1276(A, D) in Bagua Province, Amazonas Region and 73(B, C) in Cutervo Province, Cajamarca Region.   Typification notes The protologue of  B. urticaeincludes the citation ‘Haec Dr. Mutis’ ( Linnaeus 1781), implying that Carl Linnaeus the younger used material collected during Mutis’s expedition to modern day Colombiato describe the species. Previous authors have struggled to locate the material used; for example, Smith & Wasshausen (1979)stated it may be housed in M herbarium. Following his death, Linnaeus the younger’s collections were sold to the Linnean society of London by his widow. This collection contains a single duplicate collected by Mutis labelled as ‘  urticae’ and matching Linnaeus the younger’s description (LINN1125.3). We designate this sheet as the lectotypeof  B. urticaeherein. There is a single specimen of the same species collected by Mutis in MA herbarium (MA660695), which may be a duplicate of this collection. Bentham described  B. columnarisBenth.and  B. trachypteraBenth.based upon collections made by Karl Theodor Hartweg ( Bentham 1845: 131, 184). Hartweg’s main collections were deposited at Kew, which is also where Bentham was based so it is appropriate to designate lectotypesat K. The type collection of  B. columnarisis K.T. Hartweg 740and there are two collections in K, including a barcoded specimen (K000536702). This specimen was included in the Kew negative collection (#11506) so has been used as a de facto type for many years, so we designate it as the lectotypeof  B. columnaris. There is only a single sheet of K.T. Hartweg 1023at Kew (K000536703), which is the type collection of  B. trachyptera, so we designate this as the lectotypeherein. Klotzsch described  C. coccineaKlotzschbased upon material collected by Ruiz and Pavón collected in Muña ( Klotzsch 1855: 248). A specimen in Berlinherbarium matches this material and has been determined as “  Casparya coccineaKl.” Photographs of this specimen are found in several international herbaria used as the de facto type. We designate this in B specimen as the lectotypeof this name. The protologue of  Sassea glabraKlotzschcited material in “Herb. Ruizii Mss.” collected in the Andes of Muña and labelled as “  Begonia cucullata” ( Klotzsch 1855: 254). Despite a thorough search of European herbaria, we have not been able to find any specimens collected by Ruiz that were also collected in Muña. A specimen in HAL herbarium however is labelled as “  Begonia cucullata” and has a complex history. It was incorporated into the Lambert herbarium, most likely between 1816 and 1824. Lambert’s herbarium was sold in 1842, and this specimen was probably sold to Berlinas part of Lot 102, where it was worked on by Klotzsch. At a later, unclear date, the specimen became part of HAL herbarium where it now has the barcode HAL0121733. Upon incorporation into HAL herbarium, the specimen was remounted and given a new typed label. It seems likely that some data, perhaps including the locality, was lost in this process. As this specimen was definitely seen by Klotzsch and probably is the specimen cited in his locality, we designate this as the lectotypeof  S. glabraherein. The protologue of  C. urticaevar. hispidaA.DC.included reference to two specimenscollected in Colombiaby I.F. Holton and material collected in Costa Ricaby Hoffmann, variously distributed in herbaria in Berlin, Kew, and Geneva ( de Candolle 1864: 274). We designate a collection of C. Hoffmann 736in Berlin (B100243045) as the lectotypeof  C. urticaevar. hispidaherein as it has both flowers and fruit. This sheet was also photographed by J.F. Macbride (photo #20856) in the 1930s and the photograph distributed to various USand European herbaria, so has been used as a de facto type for almost a century. In the protologue of  B. monticolaC.DC., Anne Casimir Pyramus de Candollecited collections of A. Sodiro 584and  592in‘herb Candolle’ in Geneva( de Candolle 1908: 325). The de Candolle herbarium usually refers to the herbarium of the Prodromusin Geneva, which is now referred to as G-DC but there are no sheets of either collection in G-DC. We have however seen two duplicates of A. Sodiro 592in the general collection in Geneva(G), both of which have labels headed ‘Herbier de Candolle’. Of these, one was photographed by J.F. Macbride (photo #24195) and has therefore been used as a de facto type. This sheet is an excellent collection, including flowers, fruits, and a whole plant so we designate it the lectotypeof  B. monticolaherein.   Identification notes   Begonia urticaeis the most common Peruvian member of B.sect.  Casparya, which can be recognised by its three-horned rather than three-winged fruits. Species of this section are also unusual in that their androecium and gynoecium are the same colour as their tepals, which are white, pink, orange, or red in  B. urticae, rather than yellow. It can be distinguished from  B. hirtaand  B. condorensis sp. nov.by its much smaller staminate (≥ 8 mmlong vs ≤ 9 mmlong) and pistillate (≥ 6 mmlong vs ≤ 9 mmlong) tepals.    Distribution and ecology Known from Peru, Ecuador, Colombia, Venezuela, Panama, and Costa Rica. Within Peru, collected in Amazonas, Piura, Cajamarca, San Martín, Huánuco, Pasco, and CuscoRegions ( Fig. 21B). Found within northern Peruvian relict montane forest at an elevation of 2100–3200 ma.s.l., and within lower, middle, and upper montane Forest elevations of 1420–3370 ma.s.l. Typically collected as an epiphyte or lithophyte or more rarely as a terrestrial herb. It is only found within the most humid of forests or humid microhabitats, such as cave mouths or streams. The red, pink, or orange flowers of the species suggest it is pollinated by hummingbirds, but the flower lacks other obvious adaptations for bird pollination. The horned fruits of  B. urticaeappear adapted for dispersal by water drops. 4517461630 MA J. Mutis 58 59 MA660695 1 4517461499 K K. T. Hartweg Ecuador Mountains of Loxa 58 59 K000536702 1 Loja 4517461496 LD, K, OXF 58 59 LD1404657 1 4517461511 K K. T. Hartweg Colombia 3200 2.75 Prov. Popayan 1307 -76.3 In sylvis juxta Pitayo 58 59 K000536703 1 Cauca 4517462624 [635,1054,1310,1336] K, LD 58 59 LD1418113 1 4517462453 B Peru -9.666667 1299 -75.816666 58 59 1 Huanuco 4517461475 HAL, B H. Ruiz Peru Lamberti 58 59 HAL0121733 1 4517461346 B, F K. Hoffmann Costa Rica Prope Aguacate Prope Aguacate 58 59 B100243045 1 4517461670 G-DC I. F. Holton Colombia COLOMBIA 58 59 1 4517461934 [632,1160,1853,1880] G-BOIS, K, NY I. F. Holton Colombia COLOMBIA 58 59 K000536700 1 4517461307 1852-10 K I. F. Holton In montibus juxta Botatam 58 59 K000536699 1 4517462465 [1000,1398,1890,1916] 1852-10 G-BOIS, G-DC, NY I. F. Holton In montibus juxta Botatam 58 59 K000536699 1 4517462133 G A. Sodiro Ecuador 59 60 Montibus Pichincha et Corazon 58 59 1 4517461302 [452,633,265,291] G A. Sodiro Ecuador Montibus Pichincha et Corazon 59 60 1 4517462100 A. Sodiro In silva subandina et subtropica 59 60 1 4517461349 1924-03-21 US P. C. Standley Costa Rica 9.983334 Las Nubes 1298 -83.96667 Las Nubes 59 60 US00115472 1 San Jose 4517461794 1939-01-14 F P. H. Allen Panama 2375 8.833333 trail from Cerro Punta to headwaters of rio Caldera 1300 -82.5 59 60 V0077648F 1 Chiriqui 4517462340 1939-01-14 GH, MO, NY P. H. Allen Panama 2375 8.833333 trail from Cerro Punta to headwaters of rio Caldera 1300 -82.5 59 60 GH00068149, MO-510547, NY00112265 1 Chiriqui 4517462044 1944-10-11 GH J. Cuatrecascas Colombia 3550 3.3666668 Cordillera occidental, Los Farallones, filo de la cordillera, extremo N en el cerro Alto del Buey 1307 -76.7 59 60 GH00068296 1 Valle Department 4517461785 1944-10-11 COL, F, P, US J. Cuatrecascas Colombia 3550 3.3666668 Cordillera occidental, Los Farallones, filo de la cordillera, extremo N en el cerro Alto del Buey 1307 -76.7 59 60 COL000003009, COL000003010, V0077669F, P01900836, US00169383, US00115488 1 Valle Department 4517461699 [189,1246,1818,1845] G, G-BOIS J. A. Pavon Peru Prov. Bongara Prov. Bongara 59 60 1 Amazonas Region 4517462490 1978-09-30 MO, US P. J. Barbour 2345 -5.5666666 Cordillera Colan SE of La Peca 1305 -78.4 59 60 MO-2180412, US00222330 2 4517462733 2014-07-18 USM Y. F. Deng 2658 Area de Conservacion Privada Abra Patricia-Alto Nieva 59 60 1 Yambrasbamba 4517462068 2018-07-06 USM P. W. Moonlight 2064 -5.902222 Bosque Berlin 21 -78.415276 Bagua 59 60 1 4517461769 1996-12-07 MO, USM R. Vasquez & R. Rojas Camporedondo 2710 60 61 -6.076389 Tullanya 21 -78.362495 Prov. Luya 59 60 MO-2078375 1 4517462335 1989-09-07 MO, USM C. Diaz & J. Campos 2850 -6.6 Camporrendo 1304 -77.88333 60 61 MO-1643487 1 4517461976 2008-05-22 HUT V. Quipuscoa S. & L. Caceres M. & I. Trevino Z. & J. Estela R. Vista Alegre 2207 -6.1194444 La Fortaleza camino a las construcciones Chachapoyas -77.3127777 Rodriguez de Mendoza 60 61 1 4517461459 1962-08-08 US, USM J. J. Wurdack Chachapoyas 2900 -6.1666665 South side of the Molinopampas-Diosan pass 1305 -77.683334 60 61 US00222331 1 4517461843 1962-08-22 G, NY, P, US, USM J. J. Wurdack Leymebamba 3100 -6.75 middle eastern Calla-Calla slopes 1304 -77.88333 60 61 P05586561, US00222329 1 4517461813 2009-06-30 HUT R. W. Bussmann & J. Gruhn & R. Wagster & J. E. Briones & O. Espinoza & W. Espinoza Leymebamba 3090 -6.9455557 rio El Jardin 21 -77.69833 60 61 1 4517462251 2006-07-28 USM A. Cano & W. Mendoza & N. Valencia 3160 -5.0333333 Carmen de la Frontera 1305 -79.28333 Prov. Huancabamba 60 61 1 Piura Region 4517461908 2006-04-24 USM A. Cano & N. Valencia & I. Salinas 3150 -5.133333 Cerro Chinguela 1305 -79.38333 60 61 1 4517461847 1981-09-15 HUT, MO, NY A. Sagastegui A. 2200 -5.1 El Batan (Chinguela-El Carmen) 1305 -79.35 60 61 MO-2180410 1 4517462688 2000-11-09 MO, USM R. Vasquez & J. Campos & G. Calatayud & M. Huaman 2250 -5.0772223 San Jose de Lourdes 21 -78.89166 San Ignacio 60 61 MO-1835905 1 Cajamarca Region 4517462269 1988-08-03 MO C. Diaz & H. Osores San Andres de Cutervo 2250 -6.15 Parque Nacional de Cutervo 1305 -78.683334 Prov. Cutervo 60 61 MO-098014 1 4517462599 1991-09-15 MO, USM A. H. Gentry & C. Diaz & R. Ortiz 2150 -6.1666665 Parque Nacional de Cutervo 1305 -78.75 60 61 MO-098017 1 4517462655 CPUN J. Cabanillas S. Dist. San Andres de Cutervo 2500 -6.233333 El Pajonal, alrededores las grutas da los Guacharos 1305 -78.75 60 61 1 4517462678 1994-08-07 MO, NY S. Leiva G. & P. Chuna & J. Cadle Chota 2250 -6.35 Querocota-La Granja 1304 -79.066666 60 61 MO-1641563, MO-1641564 1 4517462087 1993-06-21 CPUN I. Sanchez V. 2600 -6.4333334 Rejopampa 1304 -78.75 60 61 1 4517461687 2010-06-05 US, USM J. L. Clark & M. Chocce & L. Clavijo & M. Overstreet & J. Perez 1750 -5.6658335 Bosque Proteccion Alto Mayo 21 -77.74833 Prov. Rioja 60 61 US001862879 1 San Martin Region 4517461646 2016-02-02 E, MO, MOL P. W. Moonlight & A. Daza 1707 -5.7005553 Bosque Proteccion Alto Mayo 21 -7.7408338 60 61 E00885556 1 4517462171 1996-11-03 CPUN, F, US I. Sanchez V. & M. O. Dillon Dist. Nuevo Cajamarca 1420 -5.9 Bosque Proteccion Alto Mayo 1305 -77.38333 60 61 V0086776F, US00673159 1 4517461323 1985-07-22 US K. Young 2950 -7.0 Above Gran Pajaten ruins 42801 -77.0 Mariscal Caceres 60 61 US00222175 1 4517462221 [429,940,1538,1565] 1985-07-24 HUT K. Young 60 61 1 4517461343 1985-11-22 MOL, US K. Young -7.65 Above timberline Puerta del Monte 1303 -77.48333 60 61 US00222326 1 4517461360 2000-08-05 USM A. Cano et al. 3140 -7.649111 Puerta de Monte 1 -77.41622 60 61 1 4517462247 1913-06-01 G-DC, G-BOIS, MOL, US J. A. Pavon & A. Weberbauer 3100 -9.566667 In Muna 1299 -75.816666 Huanuco 60 61 US00222044 1 Huanuco Region 4517461603 1913-06-01 G-DC, G-BOIS, MOL, US J. A. Pavon & A. Weberbauer 3100 -9.566667 In Muna 1299 -75.816666 Huanuco 60 61 US00222044 1 Huanuco Region 4517462144 MA H. Ruiz & J. A. Pavon -9.666667 Muna 1299 -75.816666 60 61 MA813495, MA813496 1 4517461537 2008-10-18 2008-10-21 2008-10-18 HOXA, USM M. Cueva & A. Pena & R. Rivera & M. Moens Huancabamba 1895 -10.27535 Parque Nacional Yanachaga-Chemillen, Sector Tunqui, camino hacia Maria Punis 1 -75.51631 Prov. Oxapampa 60 61 1 Pasco Region 4517462066 2003-06-12 HOXA, MO, US R. Vasquez et al. Huancabamba 3035 -10.379444 Parque Nacional Yanachaga-Chemillen 21 -75.461945 60 61 MO-1102451, US000843968 1 4517462714 2009-04-26 2009-04-27 2009-04-26 HOXA, USM M. Cueva 2483 -10.545279 Parque Nacional Yanachaga-Chemillen 21 -75.35833 60 61 1 4517461560 1968-07-07 US T. R. Dudley 2700 61 62 -12.616667 In deep valley just below Camp 5 and the land-slide 1293 -73.53333 Prov. La Convencion 60 61 US00222325 1 Cusco Region