Begonia bracteosa A.DC.

Moonlight, Peter. W., Jara-Muñoz, Orlando A., Purvis, David A., Delves, Jay, Allen, Josh P. & Reynel, Carlos, 2023, The genus Begonia (Begoniaceae) in Peru, European Journal of Taxonomy 881, pp. 1-334 : 69-78

10. Begonia bracteosa A.DC.

Figs 4A , 5B , 6A , 24B , 26 , 27

Annales des Sciences Naturelles Botanique, Série 4 11: 132 ( de Candolle 1859).

– Type: COUNTRY UNKNOWN • s.c., s.n.; lectotype: K [ K000536745 ], designated here.

de Candolle (1864: 331); Smith & Schubert (1941a: 186); Irmscher (1949: 583); Vásquez et al. (2005: 112–125); Brako & Zarucchi (1993: 191).

Begonia subciliata A.DC. Annales des Sciences Naturelles Botanique, Série 4 11: 132 ( de Candolle 1859).

– Type: PERU • J.A. Pavón s.n.; lectotype: G ex G-BOIS [ F neg. 8518], designated here; syntype: PERU • J.A. Pavón s.n.; G ex G-BOIS. Syn. nov.

de Candolle (1864: 331); Smith & Schubert (1941a: 186); Irmscher (1949: 586); Brako & Zarucchi (1993: 195).

Begonia roezlii Regel, Gartenflora 25: 194 ( von Regel 1876).

– Type: lectotype: plate 871 in Gartenflora 25, 1876, designated here. Syn. nov.

Begonia roezlii Lynch (nom. illeg.; later homonym non Regel), The Gardeners’ Chronicle, New Series II 11: 566 ( Lynch 1879). – Begonia lynchiana Hook.f. Curtis’s Botanical Magazine 110: t. 6758 ( Hooker 1884).

– Type: neotype: plate CDII in The Garden, 24: 162 ( Lynch 1883), designated here. Syn. nov.

Begonia subciliata var. persicina Irmsch., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 74: 588 ( Irmscher 1949).

– Type: PERU – Junín Region: [Prov. Chanchamayo] • Pichis Trail , Eneñas ; [10°45′ S, 75°13′ W]; 1600 m a.s.l.; 28 Jun.–8 Jul. 1929; E.P. Killip & A.C. Smith 25613; lectotype: US [ US00115461 ], designated here; isolectotype: NY [ NY03091030 ]. – [Pasco Region: Prov. Oxapampa] • Pichis Trail , between San Nicolas and Azupizú; [10°41′ S, 74°55′ W]; 650–900 m a.s.l.; 6 Jul. 1929; E.P. Killip & A.C. Smith 26118; syntype: F; isosyntypes: NY [ NY03091031 ], US [ US00222023 ]. Syn. nov.

Brako & Zarucchi (1993: 195); Vásquez et al. (2005: 112–125); León & Monsalve (2006: 169).

Begonia tribracteata Irmsch., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 74: 589 ( Irmscher 1949).

– Type: PERU – Ayacucho Region • Ccarrapa, between Huanta and río Apurimac; [12°43′ S, 73°55′ W]; 1500 m a.s.l.; 5–17 May 1929; E.P. Killip & A.C. Smith 22326; lectotype: F [ V0042332F ], designated here; isolectotypes: NY [ NY01085486 ], US [ US00115474 ]. Syn. nov.

Brako & Zarucchi (1993: 195); León & Monsalve (2006: 169).

Begonia suprafastigiata Irmsch., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 74 ( Irmscher 1949: 590).

– Type: PERU – Cusco Region: Prov. Calca • Lares valley above Mantoc ; [12°57′ S, 72°05′ W]; 2700 m a.s.l.; 8 Mar. 1929; A. Weberbauer 7907; lectotype: B [ B100243054 ], designated here; isolectotypes: F [ V0042330F ], NY [2: NY03091032 , NY03091033 ], US [ US00955808 ]. Syn. nov.

Vásquez et al. (2005: 112–125).

Begonia ciliata Pav. ex A.DC. (nom. inval.; nom. rej. pro syn. Begonia subciliata A.DC. ), Prodromus Systematis Naturalis Regni Vegetabilis 15 (1): 331 ( de Candolle 1864).

Begonia cyathophora auct. non Poepp. & Endl.: R.Vásquez et al., Arnaldoa 12 (1–2): 112–125 ( Vásquez et al. 2005).

Begonia suspinulosa auct. non Irmsch.: R.Vásquez et al., Arnaldoa 12 (1–2): 112–125 ( Vásquez et al. 2005).

Begonia cyathophora Poepp. & Endl. pro parte in L. B.Sm. & B. G.Schub., Publications of the Field Museum of Natural History, Botanical Series 13 (4/1): 186, ( Smith & Schubert 1941a).

Begonia albomaculata C.DC. pro parte in Brako & Zarucchi, Monographs in Systematic Botany from the Missouri Botanical Garden 45: 191 ( Brako & Zarucchi 1993).

Begonia cyathophora Poepp. & Endl. pro parte in Brako & Zarucchi, Monographs in Systematic Botany from the Missouri Botanical Garden 45: 191 ( Brako & Zarucchi 1993).

Begonia lophoptera Rolfe pro parte in R.Vásquez et al., Arnaldoa 12 (1–2): 112–125 ( Vásquez et al. 2005).

Etymology

Named for the large bracts that protect the developing inflorescence in this species.

Selected specimens examined

PERU • H. Ruiz & J.A. Pavón s.n.; G ex G-BOIS, possible isolectotype of Begonia subciliata A.DC • H. Ruiz & J.A. Pavón s.n.; G-DC ex G-BOIS, possible isolectotype of Begonia subciliata A.DC. – San Martín Region: Prov. Huallaga • Abajo de La Morada ; 6°57′ S, 77°32′ W; 2000–2200 m a.s.l.; 12 Aug. 1997; V. Quispuscoa S. & J. Bardales 1013; HUT, US [ US00673154 ] • Dist. Saposoa, al Noroeste de Añazco Pueblo ; [6°50′ S, 77°28′ W]; 2000–2100 m a.s.l.; 5 Sep. 2000; V. Quispuscoa S., S. Leiva G., Y. Díaz V. et al. 2357; HUT. – Prov. Mariscal Cáceres • La Morada y el río Huayabamba ; [7°01′ S, 77°19′ W]; 1900–2200 m a.s.l.; 29 Jun. 1995; V. Quipuscoa S. 84; HUT. – Huánuco Region: Prov. Marañón • Centro poblado Chipaco , sector Nueva Sena ; 9°14’05.21”S, 76°29’42.77”W; 1450 m a.s.l.; 24 Jul. 2021; P. González & P. Arista 10062; USM. – Prov. Leoncio Prado • Dist. Hermilío Valdizán: La Divisoriam km 194 ; [9°11′ S, 75°48′ W]; [ca 1600 m a.s.l.]; 13 Aug. 1945; C.A. Ridoutt 12997; MO [ MO-1642862 ], USM • La Divisoria, ca 20 km NNE of Tingo María on road to Pucallpa; [9°12′ S, 75°48′ W]; ca 1600 m a.s.l.; 11 Jul. 1981; M.O. Dillon 2633; F, USM • Arroyo Bravo , about 40 km from Tingo María on highway to Pucallpa; [9°13′ S, 75°51′ W]; ca 1350 m a.s.l.; 1 Nov. 1949 – 5 Jan. 1950; H.A. Allard 20378; US [ US00222063 ]. – Prov. Huánuco • Dist. Mariano Damaso Beraun: Fundo Honolulo , entre Cayumba y Tingo María; [9°27′ S, 75°59′ W]; 600–650 m a.s.l.; 8 Aug. 1947; R.A. Ferreyra 2272; USM [2] • Carpish entre Huánuco y Tingo María; [9°30′ S, 75°56′ W]; 800–900 m a.s.l.; 15 Jul. 1948; R. Ferreyra 4222; MO [ MO-2264385 ], MOL, USM • Cuesta de Carpish; [9°42′ S, 76°06′ W]; 1600 m a.s.l.; C. Vargas 160; USM. – Ucayali Region: Prov. Padre Abad • Dist. Padre Abad, cerca a río Chino ; [8°58′ S, 75°42′ W]; 1400–1600 m a.s.l.; 11 Jun. 1976; J. Schunke V. 9221; MO [2: MO-1642856 , MO-1642857 ], NY, US [2: US00222051 , US00222054 ] • km 212 ; [9°09′ S, 75°47′ W]; 12 Aug. 1943; C.A. Ridoutt 13112; USM • [Dist. Hermilío Valdizán], Divisoria, 6 Sep. 1947; [9°10′ S, 75°47′ W]; 1600 m a.s.l.; F. Woytkowski 34491; BM, G, MO [ MO-1642852 ], US [ US00222059 ]. – Pasco Region: Prov. Oxapampa • Dist. Pozuzo, Parque Nacional Yanachaga-Chemillén , carretera cerca a la Quebrada Misho; 10°10′ S, 75°34′ W; 1210 m a.s.l.; 14 Apr. 2003; A. Monteagudo, G. Ortiz & R. Francis 4993; E [2: E01007256 , E01007257 ], HOXA [2], MO [ MO-2991519 ] • Along road Chatarra-Cacazu ; 10°32′ S, 75°04′ W; 890 m a.s.l.; 13 Jul. 2003; H. van der Werff, R. Vásquez, B. Gray, R. Ortiz & N. Davila 18427; HOXA, MO [ MO-1102987 ], NY, US [ US00843931 ] • Dist. Palcazu, Bosque de Protección San Matias-San Carlos ; 10°44′ S, 74°55′ W; 900–1600 m a.s.l.; 7 Jul. 2002; R. Vásquez & A. Monteagudo 27716; HOXA, MO [2: MO-3009218 , MO-3009238 ], USM . – Junín Region: Prov. Tarma • 38.4km NE of Tarma on Carretera 20B to Oxapampa; [11°14′ S, 75°31′ W]; 2036 m a.s.l., 13 Nov. 1979; J. Jones 9106; MO [ MO-1642850 ], NY • Huacapistana ; [11°15′ S, 75°31′ W]; 1800–2400 m a.s.l.; 5–8 Jun. 1929; E.P. Killip & A.C. Smith 24097; NY, US [ US00222027 ] • ibid.; E.P. Killip & A.C. Smith 24110; NY, US [ US00222026 ]. – Prov. Chanchamayo • Colonia Perene ; [10°57′ S, 75°13′ W]; ca 680 m a.s.l.; 14–26 Jun. 1929; E.P. Killip & A.C. Smith 24988; F, NY, P [ P05587391 ], US [ US00222322 ] • Valley of río Tulumayo, ca 10 km S of San Ramón; ca 11°15′ S, 75°20′ W; 800–900 m a.s.l.; A.H. Gentry, D. Smith & N. Jaramillo 41526; MO [ MO-2154626 ], US [ US00672866 ], USM • ca 8.5 km NW of San Ramón on dirt road along E side of río Oxabamba valley; 11°30′50″ S, 75°24′20″ W; 1096 m a.s.l.; 22 Jun. 2014; P.W. Moonlight & A. Daza 25; E [ E00724450 ], MOL, USM. – Prov. Jauja • Road from Monobamba to Jauja ; 11°24′54″ S, 75°20′40″ W; 1977 m a.s.l.; 17 Feb. 2016; P.W. Moonlight & A. Daza 265; E [ E00885551 ], MOL. – Prov. Satipo • 20 km SW of Satipo on road to Concepción; [11°22′ S, 74°44′ W]; 1000 m a.s.l.; 25 May 1979; D.C. Wasshausen & F. Enarnación 1102; G, K, US [ US00222342 ], USM • Road Satipo- Comas , above Mariposa; 11°26′27.6″ S, 74°46′54.4″ W; 1430 m a.s.l.; 1 Oct. 2007; R.T. Pennington & A. Daza 1993; E [ E00567573 ], MOL • Road from Comas to Satipo ; 11°30′39″ S, 74°50′52″ W; 2550 m a.s.l.; 13 Feb. 2016; P.W. Moonlight & A. Daza 232; E [ E00885604 ], MO [ MO-3254811 ], MOL. – Ayacucho Region: Prov. La Mar • Along roadside between Machente and Rosario; [12°39′ S, 73°51′ W]; 1100 m a.s.l.; 15 Sep. 1976; D.C. Wasshausen & F. Encarnación 665; K, US [ US00222343 ], USM • Between El Tambo and Ayna; [12°45′ S, 73°59′ W]; 2375 m a.s.l.; 3 Feb. 1974; T. Plowman & E. Wade Davis 4687; USM • Dist. Anco; [12°58′ S, 73°39′ W]; 2150 m a.s.l.; 20 Jun. 2001; J. Roque & C. Arana 3011; USM. – Cusco Region: Prov. Quispicanchis • Tio to Huarayaca ; 1960 m a.s.l.; 28 Jan. 1943; J.H. Vargas López 3156; MO [ MO-2264402 ]. – 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 5671; MO [ MO-2153592 ], US [ US00932206 ] • Road Quebrada-Alto Lacco ; 12°37′22″ S, 72°14′40″ W; 2800 m a.s.l.; 30 Apr. 2006; H. van der Werff, L. Valenzuela, E. Suclli & A. Carazas 21180; K, MO [ MO-2184528 ], US [ US00951227 ] • 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 21014; MO [ MO-1839263 ]. – Prov. Urubamba • Dist. Ollantaytabmbo, road from Ollantaytambo to Quebrada Honda; 13°01′28″ S, 72°19′29″ W; 2573 m a.s.l.; 7 Aug. 2014; P.W. Moonlight & A. Daza 99; E [ E00724461 ], MOL, USM • Urubamba valley , Machu Picchu ; [13°10′ S, 72°32′ W]; ca 1525 m a.s.l.; 9 May 1939; E.K. Balls 6818; E [ E00299515 ], K [2], US [ US00222025 ] • Dist. Machu Picchu, Aobamba ; 13°12′ S, 72°33′ W; 2383 m a.s.l.; 16 Apr. 2003; L. Valenzuela, G. Calatayud & I. Huamantupa 1821; MO [ MO-492891 ]. – Prov. Paucartambo • 6–10 km from Puente Manto Jan. towards Amparaes along the main road between Amparaes and Quebrada Honda; 12°58′12″ S, 72°03′40″ W; 2600 m a.s.l.; 8 Nov. 2017; T. Särkinen & M.Y. Correa 5303; E [ E01053435 ], USM • Dist. Kosñipata, San Pedro ; 13°07′41″ S, 71°34′39″ W; 2210 m a.s.l.; 28 Oct. 2007; R. Vásquez, C. Davidson, S. Davidson, J. Farfán, E. Suclli & A. Peña 32943; MO [ MO-2131220 ] • Wayqecha Biological Station , río Kosñipata watershed, near the road Paucartambo-Shintuya; 13°10′30″ S, 71°35′12″ W; 2200– 3100 m a.s.l.; 31 May 2010; J.L. Clark, L. Clavijo & M. Overstreet 11738; NY [ NY02862132 ]. – Prov. La Convención • Dist. Quellouno , Túpac Amaru ; 12°25′46″ S, 72°29′10″ W; 1131 m a.s.l.; 17 Sep. 2007; I. Huamantupa, G. Calatayud, J. Tito, B. Rado & R. Ayerbe 10358; MO [ MO-2132037 ]) • Dist. Quillabamba, Colcapampa , Maranura ; [12°56′ S, 72°40′ W]; 1210–1435 m a.s.l.; 29–30 Dec. 1986; P. Núñez, W. Cruz & M. Cruz 6775; MO [ MO-1835955 ], USM • Dist. Santa Teresa, road from Santa Teresa to Totora ; 13°12′58″ S, 72°37′07″ W; 2005 m a.s.l.; 6 Aug. 2014; P.W. Moonlight & A. Daza 87; E [ E00724446 ], MOL, USM. – Puno Region: Prov. Carabaya • Dist. Ollachea, entre Clavana y San Gabán; [13°32′ S, 70°27′ W]; 1000 m a.s.l.; Jun. 1982; C.N. Ochoa & A. Salas 14853; NY • Ollachea to San Gabán ; [13°37′ S, 70°27′ W]; 1000–2000 m a.s.l.; 17–24 Jul. 1978; M.O. Dillon, J. Aronson, A. Herra & P. Berry 1121; MO [2: MO-2264399 , MO-2264400 ], NY, US [ US00222350 ]. – Prov. Sandia • 2–6 km from Oconeque; [14°04′ S, 69°04′ W]; 1800–2100 m a.s.l.; 22–26 May 1942; R.D. Metcalf 30581; MO [ MO-1835961 ], US [ US00222028 ] • Dist. Alto Inarnbari, Sector Mancuari ; 14°06′54.5″ S, 69°15′44″ W; 1375 m a.s.l.; 26 Mar. 2000; P. Zegarra, I. Marnani, V. Ayala, N. Marnari & J.P. Marnani 50; MOL. – Cultivated • Grown in University of California Botanical Garden from seed from the ruins of Machu Picchu , collected during the University of California Botanical Garden Expedition to the Andes, accession 36.1773; 1939; J. West 6416; E [ E00299514 ], MO [2: MO-1643584 , MO-1835945 ], USM.

Description

Caulescent herb, to 4 m high. Stem erect, branching; internodes to 14 cm long, to 12 mm thick, succulent, pale green to red, glabrous or rarely sparsely tomentose. Stipules deciduous, elliptic to ovate, 18–44 × 6–18 mm, apex acute to truncate, translucent to transparent at the margins, pale green, glabrous, margin entire, aciliate. Leaves> 5, alternate, basifixed; petiole 2–16 cm long, pale green to red, glabrous or rarely pilose; blade, asymmetric, ovate, to 25 × 15 cm, succulent, apex obtuse to short acuminate, base obliquely cordate, basal lobes not overlapping or overlapping only at the petiole apex, sinus to 35 mm deep, margin denticulate or denticulate to serrate or dentate, sometimes with short, triangular lobes to 1 cm long at the end of the major veins, ciliate, upper surface green, sometimes flushed red around the petiole apex, glabrous, lower surface glabrous or rarely pilose, lower surface green, sometimes flushed red around the petiole insertion, flushed red around the major veins, or completely red, glabrous or rarely sparsely pilose, veins palmate but with one primary vein, 7–9 veined from the base, with 1–3 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 at least 100 staminate flowers and 100 pistillate flowers, protandrous; peduncle to 20 cm long, white, pale green, or red, glabrous or rarely sparsely pilose, bracts deciduous, ovate to elliptic, 7–35 × 1.5–23 mm, translucent to transparent at the margins, white, pink, or red, glabrous, apex acute to truncate, margin entire, aciliate. Staminate flowers: pedicels to 20 mm long, glabrous; tepals 2, spreading, broadly ovate, 6–17 × 6–18 mm, apex rounded, white, pink, or red, glabrous, margin entire, aciliate; stamens 20–50, spreading, yellow, filaments 0.5– 2 mm long, free, anthers ellipsoid, 1.5–2.5 × 0.25–0.5 mm long, dehiscing via lateral slits, connectives extending to 0.5 mm, symmetrically basifixed. Pistillate flowers: pedicels to 20 mm long; bracteoles lacking; tepals 2, deciduous in fruit, spreading, ovate, 5–14 × 5–15 mm, apex rounded, white, pink, or red, glabrous, margin entire, aciliate; ovary body ovoid, 3–13 × 2–6 mm, pale green, white, pink, or red, glabrous or rarely pilose, unequally 3-winged, wings semi-circular to triangular, largest 7–16 × 4–20 mm, smallest 3–14 × 1–6 mm; 3-locular, placentae branches divided, bearing ovules on both surfaces; styles 3, yellow, free, 4–6 mm long, once-divided, stigmatic papillae in a twice-spirally twisted band. Fruiting pedicel to 28 mm long. Fruit body ovoid, to 18 × 9 mm, drying brown, wings same shape as in ovary, the largest expanding to 25 × 20 mm, the smallest to 18 × 9 mm.

Proposed conservation assessment

Widespread and abundant throughout its range, and frequently found in secondary forest, forest edges, and heavily disturbed areas. Its EOO is> 21 500 km 2 and includes several protected areas. We assess B. bracteosa as Least Concern (LC), which replaces the DD assessments of B. tribracteata Irmsch. and B. subciliata var. persicina Irmsch. ( León & Monsalve 2006).

Notes

The specimens we recognise here as B. bracteosa form a complex that includes the types of seven separate basionyms and several combinations of these basionyms. Plants in this complex are extremely common from Huánuco Pasco to Cusco Regions and are present but less abundant within San Martín and Puno Regions, and La Paz Department in Bolivia. All specimens from this complex are similar in that they are succulent, upright herbs with large, deciduous stipules; transversely ovate leaves; large inflorescences with> 20 staminate and> 20 pistillate flowers; staminate and pistillate flowers both with 2 tepals; pistillate flowers that lack bracteoles and have twice-spirally twisted styles; and staminate flowers with 20–50 stamens. They differ however in their indumentum; the size, texture, colour, and margins of their leaves; the size and colour of their flowers; the shape and size of their fruits; and how succulent or woody their stems are.

Within this complex, the variation in these characters is largely, but not completely, geographically structured. For example, specimens from Pasco and Junín Regions tend to have the largest leaves with the smoothest texture while specimens from Ayacucho and Cusco Regions tend to have the thickest, woodiest stems and smallest leaves. Characters also vary, however, between plants in the sun and shade, or among plants across the species’ elevational range. Furthermore, we can find no consistent breaks in any of these characters nor links between any character states. We therefore synonymise all six basionyms and their combinations with B. bracteosa .

Alphonse Pyramus de Candolle described B. bracteosa and B. subciliata A.DC. on the same page ( de Candolle 1859: 132) and distinguished them based upon their leaf margins (crenate and ciliate in B. bracteosa ; undulate to crenate in B. subciliata ); and their stipule shape and deciduousness (ovate and deciduous in B. bracteosa ; elliptic and late-deciduous in B. subciliata ). The two species were also distinguished by the presence of ciliate, bracteoles in B. bracteosa . This was however a misinterpretation of the type specimen, which contains no bracteoles but ciliate bracts. This mistake was corrected by Irmscher (1949: 586) who maintained B. bracteosa and B. subciliata as separate species, separating them by their leaf margins and texture, the shape of their fruit wings, and the shape of their stipule apex. These characters are not consistent, so we treat these two species as the same taxon. Priority for names published in the same publication is established based on the first designation as either name as accepted. This has not occurred with respect to B. bracteosa or B. subciliata , so we select the former as the accepted name because B. subciliata has previously been treated as a synonym of B. cyathophora ( Smith & Schubert 1941a) .

Begonia roezlii Regel was described by Eduard August von Regel from material grown from seed in the Saint Petersburg botanical garden ( von Regel 1876: 194). The seeds were sent to Saint Petersburg by the distinguished orchid collector Benedict Roezl who collected them in Peru. Roezl’s Peruvian travels included Chanchamayo Province and it is perhaps here that he encountered this species. Indeed, the type illustration designated shows a plant very similar in colouration to specimens we have collected in Chanchamayo (e.g., P. M. Moonlight & A. Daza 34). This illustration falls within our concept of B. bracteosa , so we synonymise this name herein.

Three years following Regel’s description of B. roezlii Regel, Robert Irwin Lynch published B. roezlii Lynch , which is a later homonym ( Lynch 1879: 566). In 1844, Joseph Dalton Hooker published a nom. nov. for this name as B. lynchiana Hook.f. ( Hooker 1884) . We have not been able to locate any type material of B. roezlii Lynch (see Typification notes below) and the description is not sufficient for identification, but illustrations of the original introduction clearly fit within our circumscription of B. bracteosa ( Lynch 1883) . We therefore synonymise both B. roezlii Lynch and B. lynchiana Hook. f. with B. bracteosa .

Irmscher (1949: 588) described B. subciliata var. persicina to encompass specimens of his concept of B. subciliata from Junín Region with unusually large inflorescences and flowers. These specimens were also unusual in that they have bright red flowers, which, to our knowledge, is only found in our concept of B. bracteosa in a small area of Junín Region. We have collected members of this species with particularly large and bright red flowers in this area ( P. W. Moonlight & A. Daza 36). Most other specimens of this species from Junín and Pasco Region have equally large flowers so synonymise this variety with B. bracteosa .

Irmscher (1949: 589) described B. tribracteata to accommodate specimens that were otherwise similar to B. subciliata and B. bracteosa but had three bracteoles subtending the ovary. Irmscher described the bracteoles as unequal, with two large bracteoles (ca 3 × 2 mm) and one residual bracteole. Our examination of the types of this name revealed no bracteoles. Rather, the pistillate flowers are in bud and their subtending peduncle has not yet elongated. With a <1 mm long peduncle, it appears as if the flower is subtended by a pair of bracteoles, but these are instead the bracts at the node below the flower. The third bract may be a misinterpretation of the inflorescence node itself. The types of B. tribracteata have unusually thick stems that appear woody but fall within our concept of B. bracteosa . We synonymise B. tribracteata with B. bracteosa herein.

Irmscher also described B. suprafastigiata Irmsch. in his revision of the begonias of South America ( Irmscher 1949: 590). While he included it in B. sect. Cyathocnemis , he noted that it resembled members of B. sect. Ruizopavonia in its elongated stems and inflorescences. The type gathering was also noted to reach 3 m tall and climb through other vegetation. We have collected this at the type locality ( P. W. Moonlight & A. Daza 89) and plants vary from relatively slender and elongated when growing through other plants to relatively robust when growing in the open. This is true of other populations in Cusco Region (e.g., P. W. Moonlight & A. Daza 79) and we do not consider these differences enough to separate species so we synonymise B. suprafastigiata with B. bracteosa .

Typification notes

Alphonse de Candolle cited material of B. bracteosa seen in “h. Hook.” In the species’ protologue ( de Candolle 1859: 132). This citation is repeated in de Candolle’s later revision of the Begoniaceae ( de Candolle 1864: 331) and by Smith and Schubert in their Flora or Peru Begonia account ( Smith & Schubert 1941a). Most of Hooker’s herbarium is now incorporated in Kew herbarium. The only sheet in this herbarium that includes a determination by de Candolle is K 000536745. This specimen has flowers and a single fruit so we designate it as the lectotype of B. bracteosa herein. The origins of this specimen are unclear, and it may be impossible to determine from where in Peru it originates.

The protologue of B. subciliata cites material from Pavón’s herbarium labelled as “ Begonia ciliata ” ( de Candolle 1859: 132) . Pavón’s personal herbarium was bought by Edmond Boissier from the heirs of Pavón in 1844. Boissier’s herbarium was held in Geneva and frequented by de Candolle, who described several species from the collection. It seems likely that the protologue was referring to material held by Boissier. De Candolle’s later revision of the Begoniaceae ( de Candolle 1864) cites material of B. bracteosa in the Boissier herbarium, which provides further support for this hypothesis. There are two sheets in G that were incorporated from the Boissier herbarium in 1944. Both sheets have minimalist labels suggestive of Pavón’s personal herbarium. One of these ( G) has a determination for B. subciliata in de Candolle’s hand while the second has “ Begonia ciliata del Peru ” in Pavón’s hand ( G, F neg. 8518). Both are equally good material, and we chose the latter as a lectotype of B. subciliata herein because the F photograph means it has been used as a de facto type for many years. The lectotype of B. subciliata is without locality but Ruíz’s diaries ( Dahlgren 1940: 198) contain a reference to a description of “ B. ciliata ” written in Huánuco. The plant may have been collected in Huánuco or at any earlier point during their expedition. The expedition travelled extensively before this date so further clarity is difficult. An illustration in MA (MA-AJB04-D-1395) matching this material is also labelled as ‘ Begonia ciliata ’, providing a direct link between a specimen and an illustration made during the expedition.

Any specimens that could be considered original material of B. roezlii Regel would most likely be housed in LE herbarium, but we have been unable to locate these despite a thorough search by herbarium staff. The protologue however includes an excellent illustration of the species ( von Regel 1876: 194) so, as it is the only known original material used by Regel, we designate the illustration of B. roezlii as the lectotype of this name herein.

Begonia roezlii Lynch was described from plants sent to Lynch by M. Benary of Erfurt and Rev. Law of Sheffield. These plants were grown from seed sent by Roezl, though Lynch wrote that the seeds originated in Mexico ( Lynch 1879: 566). The original description of this plant is valid but not sufficient for identification. Fortunately, Lynch (1883) later published a short article in The Garden to advertise the species, which was accompanied by an excellent illustration. This illustration clearly fits into our circumscription of B. bracteosa and it is extremely unlikely that the seed originated in Mexico as B. bracteosa is only found in Peru and Bolivia and B. sect. Cyathocnemis is endemic to South America ( Moonlight et al. 2018). Roezl collected both in Mexico and Peru in 1873 so it seems likely the seeds were mislabelled. We have been unable to locate any specimens that could be considered original material of B. roezlii Lynch. If these existed, they would most likely be deposited in CGE, but a thorough search of the herbarium failed to uncover any such specimens. We therefore designate the illustration of B. roezlii in Lynch’s later article ( Lynch 1883) as the neotype of this species herein. This also serves as the neotype of Joseph Dalton Hooker’s 1884 nom. nov. of B. roezlii , B. lynchiana Hook.f.

The protologue of B. subciliata var. persicina includes reference to material of E. P. Killip & A. C. Smith 25613 and 26118 held in US and F herbaria, respectively ( Irmscher 1949: 588). It is therefore appropriate to designate a lectotype from these duplicates. The material of E. P. Killip & A. C. Smith 25613 at US ( US 00115461) is in flower; thus, we designate it the lectotype of B. subciliata var. persicina herein. The protologue of B. tribracteata cites material of E. P. Killip & A. C. Smith 22326 In F and US herbaria thus it is appropriate to designate a lectotype. The duplicate in US ( US 00115474) includes only a small section of a branch with a leaf and a detached inflorescence. The duplicate in F is much more complete, including two branches, a full infructescence, and staminate flowers. We therefore designate the duplicate in F ( V 0042332 F) as the lectotype of B. tribracteata herin. Irmscher cited duplicates of A. Weberbauer 7907 in B and F herbaria in the protologue of B. suprafastigiata ( Irmscher 1949: 590) . The material held in Berlin has flowers and fruits whereas the duplicate in Chicago is sterile. We therefore designate the duplicate in B ( B 100243054) as the lectotype of B. suprafastigiata herin.

Identification notes

There are more than ten large (> 50 cm tall) species of Peruvian Begonia with an upright stem and transversely ovate leaves that lack lobes but, of these, B. bracteosa is by far the most common and widespread. It can therefore be treated as the default name for any such Begonia in Peru, as long as specimens do not have any of the following characters: a tuber (see B. piurensis ); bracteoles or> 2 tepals in the pistillate flower (see B. sect. Hydristyles and the stenotepala group of B. sect. Cyathocnemis ); multifid styles (see B. sect. Hydristyles); <20 flowers in an inflorescence (see B. longinqua Moonlight sp. nov.); bracts fused into a cyathium (see B. cyathophora ); or aciliate leaf margins (various species of B. sect. Cyathocnemis or B. sect. Hydristyles).

Distribution and ecology

Known from Peru and northern Bolivia. Within Peru, collected in San Martín, Huánuco, Ucayali, Pasco, Junín, Ayacucho, Cusco, and Puno Regions ( Fig. 24B ). Found in lower, middle, and upper Montane Forests and rarely Amazonian Forest at elevations from 600 to 2900 m a.s.l. Begonia bracteosa is usually found on the edges of montane forest or on cliffs and banks, including in disturbed areas. Like other members of B. sect. Cyathocnemis , the species flowers throughout the year but most frequently from June to November.

Figures

Fig. 4. Staminate tepal number in Peruvian Begonia. A. A single, equal pair of tepals, B. bracteosa A.DC. B. An outer pair of tepals held perpendicular to a smaller, inner pair of tepals, B. buddleiifolia A.DC. C. Ten tepals in a member of the octopetala group of B. sect. Eupetalum (Lindl.) A.DC., B. pleiopetala A.DC. Photographs taken by D.A. Purvis in the living collections of the Royal Botanic Garden Edinburgh.

Fig. 5. Staminate tepal projection in Peruvian Begonia. A–B. Spreading tepals. C–D. Projecting tepals. A. B. glabra Aubl. B. B. bracteosa A.DC. C. B. stenotepala L.B.Sm. & B.G.Schub. D. B. foliosa Kunth. Photographs taken by D.A. Purvis & P.W. Moonlight in the living collections of the Royal Botanic Garden Edinburgh.

Fig. 6. Styles and stigmas in Peruvian Begonia. A–C. Bifid styles. D. Reniform styles. E–F. Irregularly multifid styles. A. Three styles and stigmas of B. bracteosa A.DC. B. Three styles and stigmas of B. glabra Aubl. C. Single style and stigma of B. yuracyacuensis Moonlight sp. nov. D. Three styles and stigmas of B. pedemontana Moonlight sp. nov. E. Three styles and stigmas of B. obtecticaulis Irmsch. F. Single styles and stigma of B. pleiopetala A.DC. Photographs taken by D.A. Purvis & P.W. Moonlight in the living collections of the Royal Botanic Garden Edinburgh.

Fig. 24. Distribution ofBegonia sect.Cyathocnemis (Klotzsch)A.DC. in Peru and surrounding countries. A. B. amoeboides Moonlight (red) andB. imbrexiformis Moonlight sp. nov. (blue).B. B. bracteosaA.DC. (red) and B. longinqua Moonlight (blue). C. B. cyathophora Poepp. & Endl. (red), B. serratistipula Moonlight sp. nov. (blue) and C. vargasii Moonlight sp. nov. (green). D. B. lamolina Moonlight (red), B. lophoptera Rolfe (blue) and B. obtecticaulis Irmsch. (green). Black lines indicate country borders, grey lines indicate major administrative divisions, blue lines indicate rivers, and shading indicates elevation.

Fig. 26. Begonia bracteosaA.DC. A. Habit.B. Leaf, adaxial surface.C. Leaf, abaxial surface.D. Stipule, abaxial surface. E. Stipule, adaxial surface. F. Inflorescence with staminate and developing pistillate flowers. G. Developing inflorescence. H. First inflorescence bract, abaxial surface. I. Staminate flower, side view. J. Staminate flower, front view. K. Androecium, side view. L. Pistillate flower, side view. M. Pistillate flower, front view. N. Cross section of ovary. O. Pistils, side view. All photographs taken by D.A. Purvis in the living collections of the Royal Botanic Garden Edinburgh.

Fig. 27. Begonia bracteosa A.DC. A. Habit. B. Inflorescence. C. Developing inflorescence. D. first bract of inflorescence. E. Leaf, adaxial surface. F. Leaf, abaxial surface. G. Stipule, abaxial surface. H. Stipule, adaxial surface. I. Cross section of ovary. J. Pistillate flower, side view. K. Pistillate flower, front view. L. Pistils, side view. M. Staminate flower, side view. N. Staminate flower, front view. O. Androecium, side view. All photographs taken by D.A. Purvis in the living collections of the Royal Botanic Garden Edinburgh.