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 58. Begonia maynensis A.DC. Figs 71B , 74 Annales des Sciences Naturelles Botanique, Série 4 11: 126 ( de Candolle 1859 ). – Type : PERU – San Martín Region : [ Prov. Tarapoto ] • in andibus Maynensium ; [ 6°27′ S , 76°20′ W ]; Aug. 1856 ; R.E. Spruce 4859 ; lectotype : G-DC [F neg. 7345], designated by Smith & Wasshausen (1979: 246) ; isolectotype : K [ K000006031 ]. de Candolle (1864: 310) ; Smith & Schubert (1941a: 194 , 1952: 39 ); Smith & Wasshausen (1979: 246 , 1986: 49 ); Brako & Zarucchi (1993: 193) ; Vásquez et al. (2005: 112–125) ; Moonlight et al. (2017b: 190) . Begonia longimaculata Irmsch., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 76 (1): 89 ( Irmscher 1953 ) . – Type : PERU • W. Lobb s.n. ; holotype: W [ W1889-0149133 ]. Brako & Zarucchi (1993: 193) ; León & Monsalve (2006: 167) ; Moonlight et al. (2017b: 190) . Etymology The duplicate of B. maynensis in G-DC used by Alphonse Pyramus de Candolle to describe the species is labelled as from ‘In sylvis humidis andium Maynensibus’ (but see Typification notes). The species was named for this locality. Specimens examined PERU – Loreto Region : Prov. Maynas • Alto río Arabela , headwaters of the western tributary of the río Curaray , a Napo tributary ca 70 km SE of the Ecuador border ; 2°08′13″ S , 75°08′58″ W ; 160–270 m a.s.l.; 28 Aug. 2006 ; N. Dávila , R.B. Foster , G. Nuñez , C. Vriesendorp & I. Mesones 2833 ; F [ V0387176F ] • Indiana , Inique , 3°30′ S , 72°58′ W ; 115 m a.s.l.; 16 Dec. 1987 ; R. Vásquez & N. Jaramillo 10171 ; MO [ MO-2228056 ], USM • Mishuyacu , near Iquitos ; [ 4°00′ S , 73°09′ W ]; ca 100 m a.s.l.; Feb.–Mar. 1930 ; G. Klug 1065 ; NY , US [ US00222191 ]. – Prov. Alto Amazonas • Puranchim , río Sinchiyacu ; 2°50′ S , 76°55′ W ; 200 m a.s.l.; 30 Feb.–1Apr. 1987 ; W.H. Lewis , M. Elvin-Lewis , D. Fast & J.R. Campos 13392 ; MO [ MO-2340409 ]. – Prov. Datem del Marañon • Washintsa and vicinity , río Huallaga ; 3°20′ S , 76°20′ W ; 320 m a.s.l.; 13–16 Nov. 1986 ; W.H. Lewis , M. Elvin-Lewis , J. Campos & D. Fast 11799 ; MO [ MO-2340416 ], USM . – Prov. Alto Amazonas • Balsapuerto , lower río Huallaga basin ; [ 5°50′ S , 76°34′ W ]; 350–550 m a.s.l.; 29 Aug. 1929 ; E.P. Killip & A.C. Smith 28476 ; NY , US [ US0222189 ] • Dist. Teniente César Lopez Rojas , Shucushuyacu ( río Huallaga ); 6°02′ S , 75°50′ W ; 250 m a.s.l.; 13 Sep. 1981 ; R. Vásquez & N. Jaramillo 2452 ; MO [ MO-2228060 ], USM . – Prov. Manseriche • Pongo de Manseriche ; 4°26′01″ S , 77°34′18″ W ; 500 m a.s.l.; 28 Nov. 1997 ; R. Rojas , N. Peña & M. Correa 706 ; HUT , MO [2: MO-1642430 , MO-1642431 ], US [ US00672903 ], USM • Above Pongo de Manseriche ; [ 4°27′ S , 77°35′ W ]; 200 m a.s.l.; 21 Nov. 1931 ; Y. Mexia 6130 ( BM , G , MO [ MO-1642420 ], NY , U , US [ US00222193 ]. – Prov. Mariscal Ramón • Castilla , margen izquierda del río Yavari , entre Colonia Angamos y Lago Preto ; 19M 0178075 9500324; 9 Apr. 2003 ; H. Beltrán , R.B. Foster , N. Pitman , R. Garcia , C. Vriesendorp & M. Huite 5699 ; USM . – Amazonas Region : Prov. Condorcanqui • río Cenepa region , quebrada Nahem ; 15 Jul. 1974 ; R. Kayak 1081 ; MO [ MO-286165 ] • río Cenepa region , camino de jutui entsa ; [ Ajuntaienntsa ]; [ 3°52′ S , 78°18′ W ]; 215–245 m a.s.l., 30 Jul. 1984 ; R. Kayak 1385 ; MO [2: MO-286161 , MO-286162 ], NY • Dist. El Cenepa , Puerto Mori , río Comaina ; 4°23′ S , 78°21′ W ; 800 m a.s.l.; R. Vásquez , N. Jaramillo , R. Apanu , R. Apanu & V. Bermeo 18984 ; HUT , MO [ MO-286173 ], US [ US00672902 ] • Trail to Sasa , S of Huampami and E of Cenepa ; [ 4°38′ S , 78°12′ W ]; 230 m a.s.l., 17 Jul. 1974 ; B. Berlin 1717 ; MO [ MO-286153 ]. – Prov. Bagua • Dist. Imaza , Comunidad Nativa Yamayakat ; 300 m a.s.l.; 4°55′ S , 78°19′ W ; Jan. 1995 , V . Hodges & J. Gorham 123 ; HUT , MO [ MO-286158 ], US [ US00672906 ] • 10 minutes drive S of Chiriaco on main road from Bagua to Santa María de Nieva ; 5°12’51′00″ S , 78°20′02″ W ; 367 m a.s.l.; 30 Jan. 2016 ; P.W. Moonlight & A. Daza 134 ; E [ E00885572 ], G , MO , MOL • Dist. Jumbilla , along road Jumbilla-Rioja ; 5°39′58″ S , 77°46′19″ W ; 1900–1940 m a.s.l.; H. van der Werff , L. Valenzuela , G. Shareva & A. Reyes Barrantes 25374 ; HOXA . – San Martín Region : Prov. Rioja • Aguas Verdes ; 5°41′17″ S , 77°37′53″ W ; 1100 m a.s.l.; 8 Jun. 2012 ; H. van der Werff , L. Valenzuela , G. Shareva & A. Reyes B. 25374 ; E [ E01007289 ], MO [2: MO-2991759 , MO-2991760 ]. – Prov. Moyobamba • Zepelacio ; [ Jepelacio ], near Moyabamba ; [ 6°06′ S , 76°55′ W ]; ca 1100 m a.s.l.; Jul. 1934 ; G. Klug 3716 ; BM , K , MO [ MO-1642421 ], NY , US [ US00222172 ] . – Prov. San Martín • An Felsen des Pongo de Chilcayo , Tarapoto ; [ 6°26′ S , 76°16′ W ]; Oct. 1902 ; E. Ule 6467 ; G , K , L • Trail above cataratas de Ahuashiguas on road from Tarapoto to Yurimaguas ; 6°27′21″ S , 76°18′30″ W ; 761 m a.s.l.; 4 Feb. 2016 ; P.W. Moonlight & A. Daza 176 ; E [ E00885457 ], G , MOL • Near Tarapoto ; [ 6°28′ S , 76°19′ W ]; Jun. 1855 ; R.E. Spruce 4859 ; BM [ BM000777903 ], E [ E00157075 ], G [ G00237559 ], G-BOIS , G-DC [ G00676066 ], K [ K000006032 ]; MG [ MG019634 ], NY [ NY00118630 ], OXF , TCD [ TCD0005563 ]. – Prov. Mariscal Cáceres • Dist. Tocache Nuevo , río Huallaga , margen derecha del, Balsa Probana ; Apr. 1982 ; S. Barrier s.n. ; US [2: US00222173 , US00222174 ] • Dist. Campanilla , route from Tarapoto to Tocache ; 7°35′10″ S , 76°41′30″ W ; 467 m a.s.l.; 6 Feb. 2016 ; P.W. Moonlight & A. Daza 185 ; E [ E00885471 ], MO , MOL . – Ucayali Region : Prov. Purús • Cuenca del río Purús , río Curanja , cerca la communidad nativa de Colombiana ; 10°04′ S , 71°06′ W ; 300– 350 m a.s.l.; 15 Feb. 2000 ; J.G. Graham & J. Schunke V. 900 ; F [ V0088042F ], G , US [ US01008607 ]. – Huánuco Region : Prov. Pachitea • Puerta Inca , forest along small river just N of town ; 9°18′ S , 74°58′ W ; 250–300 m a.s.l.; 15 Sep. 1982 ; R.B. Foster 8830 ( F , MO [ MO-1642433 ], NY , USM . – Pasco Region : Prov. Oxapampa • Comunidad Nativa Alto Lagarto , Reserva Comunal Yanesha ; 10°08′04″ S , 75°22′06″ W ; 500 m a.s.l.; 28 Mar. 2012 ; R. Rojas & G. Ortiz 8195 ; E [ E01007276 ], HOXA . – Junín Region : Dist. Satipo • La Molina University Field Station , ca 10 km N of Satipo on 5S , ca 2 km E of field station ; 11°10′08″ S , 74°38′53″ W ; 782 m a.s.l.; 8 Jul. 2014 ; P.W. Moonlight 65 ; E [ E00724452 ], MOL , USM . – Madre de Dios Region : Prov. Manu • Cocha Cashu uplands ; 11°45′ S , 71°10′ W ; 400 m a.s.l.; P. Núñez 6127 ; MO [ MO-2228061 ], US [ US00222194 ] • Parque Nacional del Manu , río Manu , Cocha Cashu Station ; 11°50′ S , 71°25′ W ; 350 m a.s.l.; 21 Jul. 1984 ; R.B. Foster 9668 ; F , MO [ MO-1642416 ], NY , USM • ibid.; 11°50′ S , 71°25′ W ; ca 400 m a.s.l.; 17 Aug. 1983 ; A.H. Gentry 43642 ; MO [ MO-1642410 ]. – Cusco Region : Prov. La Convención • Dist. Echarati , Pagoreni well site ; 11°41′ S , 73°00′ W ; 350 m a.s.l.; P. Nuñez , H. Beltrán , W. Nauray , L. Acurio , R. de la Colina , N. Llerena & A. Salas 21778 ; US [ US00625232 ], USM • Campamento San Martín-C , Camimisaca Production Unit , S and SW of camp ; 11°47′08″ S , 72°41′57″ W ; 24 Jan. 1997 ; P. Acevedo R. , D. Bell , K. Rankin & S.F. Smith 9050 ; G , K , MO [ MO-1642432 ], NY , US [ US00594260 ], USM • Dist. Echarati , Armihuari well site ; 11°51.88′ S , 72°46.69′ W ; 535 m a.s.l. ; 14 May 1997 ; P. Nuñez , H. Beltrán , F. Ramirez , J. Tenteyo P. 20081 ; US [ US00625231 ]. Fig. 74. Begonia maynensis A.DC. A . Habit. B . Staminate flower, front view. C . Pistillate flower, front-side view. D . Branch of infructescence. Illustration by Peter Moonlight from photographs of P.W. Moonlight & A. Daza 185 (E); all scale bars are estimated. Description Caulescent herb, to 40 cm high. Stem erect, unbranched or rarely branching at the base; internodes to 3 cm long, to 6 mm thick, flexuous, brown to pale green, glabrous. Stipules late deciduous, lanceolate, 10–22 × 4–6 mm , apex acuminate, opaque to translucent, light green to brown, glabrous, margin entire, aciliate. Leaves > 3, alternate, basifixed; petiole 0.5–5 cm long, pale green, glabrous; blade subsymmetric, elliptic to oblanceolate, to 22 × 10 cm , succulent, apex acuminate, base obliquely cuneate, margin irregularly dentate, aciliate to sparsely ciliate, upper surface pale to dark green, sometimes with silver blotches between the major veins, glabrous, lower surface pale green to purple, glabrous, veins pinnate, appearing 3 veined from the base, with 6–8 secondary on each side of the lamina. Inflorescences 1–5, bisexual, axillary, erect, cymose, with 6 branches, bearing up to 32 staminate flowers and 32 pistillate flowers, protandrous; peduncle to 4.5 cm long, white, glabrous, bracts late deciduous, lanceolate, 3–5 × 1–2 mm , opaque, white, glabrous, apex acute, sometimes mucronate, margin entire, aciliate. Staminate flowers : pedicels to 15 mm long, glabrous; tepals 4 spreading, outer 2 lanceolate to ovate, 4–8 × 4–7 mm , apex acute, white, sometimes tinged pink or yellow, glabrous, margin entire, aciliate, inner 2 narrowly lanceolate, 4–6 × 2 mm , apex acute, white, glabrous, margin entire, aciliate; stamens 20–25, spreading, yellow, filaments 1.5–2 mm long, attached along the length of a 2 mm long column, anthers broadly obovoid, ca 0.5 × 0.5 mm , dehiscing via lateral slits, connectives not extended, symmetrically basifixed. Pistillate flowers : pedicels to 10 mm long; bracteoles 1, directly beneath the ovary, lanceolate, 3–5 × 1–2 mm , opaque, white, glabrous, apex acute, sometimes mucronate, margin entire, aciliate; tepals 5, subequal, deciduous in fruit, spreading, lanceolate to ovate, 5–11 × 3–5 mm , apex obtuse to acute, white, sometimes tinged yellow or pink, glabrous, margin entire, aciliate; ovary body ellipsoid, 7–15 × 2–5 mm , white, glabrous, sub-equally 3-winged, wings triangular, 12–20 × 5–10 mm ; 3-locular, placentae branches divided, bearing ovules on both surfaces; styles 3, yellow, free, ca 4 mm long, once-divided, stigmatic papillae in a spirally twisted band. Fruiting pedicel to 25 mm long. Fruit body ellipsoid to ovoid, to 20 × 8 mm , drying brown, wings same shape as in ovary, expanding to 25 × 15 mm . Proposed conservation assessment Assessed by Moonlight et al. (2017b) as Least Concern (LC). Typification notes The protologue of B. maynensis cited the collection R.E. Spruce 4859 as the type ( de Candolle 1859: 126 ). Moonlight et al. (2017b) attempted to lectotypify the name based upon a collection in G-DC (G00676969). Smith & Wasshausen (1979) had however already cited this sheet as the type of the name. This was the first time the name was lectotypified so should be followed ( Turland et al. 2018 : Article 9.19). It could be argued that Smith & Wasshausen only achieved the first stage of lectotypification as there are two sheets of R.E. Spruce 4859 at G-DC, but they also cited a photograph of the type at F. There is no photograph of the second G-DC sheet (G00676066) in F so this represents an unambiguous lectotypification. Furthermore, the sheet cited by Smith & Wasshausen (1979) has the following locality description on its label: ‘In sylvis humidis andium Maynensibus’ while the second sheet was collected ‘prope Tarapoto’. The dates on these two collections are also different ( Aug. 1856 vs Jun. 1855 ), so they are clearly two separate collections. There is only one other sheet of R.E. Spruce 4859 that matches the lectotype locality and date, which is held in Kew (K000006031). We consider this sheet an isolectotype. All other known sheets with this number were collected ‘prope Tarapoto’ in June 1855 , so cannot be considered isolectotypes. De Candolle clearly believed these sheets were also B. maynensis because he determined most of them as this species and filed one duplicate as B. maynensis in the herbarium of the Prodromus (G-DC, G00676066). We do not consider these sheets collected in ‘prope Tarapoto’ as syntypes or as isolectotypes. Identification notes Begonia maynensis is the most common terrestrial Begonia found in Amazonian and lower montane Peru with cuneate leaf bases and leaves clustered towards the apex of its stems. It is distinguished from B. buddleiifolia , which shares these characters, by its smooth and glabrous (vs bullate and glandular haired) leaf laminae. Distribution and ecology Known from Colombia , Ecuador , Peru , and Acre State in Brazil . Within Peru , known from Loreto , Amazonas, San Martín , Ucayali , Huánuco , Pasco , Junín , Madre de Dios , and Cusco Regions ( Fig. 71B ). Found in Amazonian Forest, lower and rarely middle montane forest at an elevation of 100–1940 m a.s.l. Begonia maynensis is typically collected from riverbanks or rocks in and around small streams but has also been found growing as an epiphyte in very humid forests.