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 27. Begonia humilis Dryand. in Aiton (1789) Figs 42A , 43 Hortus Kewensis 3: 353 ( Dryander 1789 ) . – Type : TRINIDAD AND TOBAGO • Cultivated in Saint Vincent from seed collected in Trinidad ; Anderson s.n. ; lectotype: LINN [ LINN-HS1474-3-1 ], designated here. – COUNTRY UNKNOWN • Cultivated in Mr Lee’s Garden ; 1788; coll. unknown s.n. ; syntype: LINN [ LINN-HS1474-3-2 ] . Dryander (1791: 166) ; Link (1822: 396) ; Otto & Dietrich (1836: 359) ; Fisch. et al. (1842: 51); Klotzsch (1855: 157) ; Walpers (1858: 883) ; de Candolle (1861: 343 , 1864: 297 ); Schulz (1911: 26) ; Smith & Schubert (1941a: 193 , 1952: 38 ); Smith (1973: 215) ; Smith & Wasshausen (1979: 238 , 1986: 9 , 1989: 52 ); Brako & Zarucchi (1993: 193) ; Vásquez et al. (2005: 112–125) . Begonia lucida Haw. Saxifragearum Enumeratio : 197 ( Haworth 1821a ). – Type : TRINIDAD AND TOBAGO • Cultivated in Saint Vincent from seed collected in Trinidad ; Anderson s.n. ; neotype: LINN [ LINN-HS1474-3-1 ], designated here . de Candolle (1861: 343) ; Schulz (1911: 27) . Begonia subhumilis A.DC., Annales des Sciences Naturelles Botanique, Série 4 11: 124 ( de Candolle 1859 ). – Type : ECUADOR – [ Prov. Guayas ] • Guayaquil , Las Bodegas ; coll. unknown 8 ; holotype: K [ K000536756 ]; isotype: B [ B10036204 ] . de Candolle (1864: 298) ; Smith & Wasshausen (1979: 238) . Begonia pavoniana A.DC., Annales des Sciences Naturelles Botanique, Série 4 11: 142 ( de Candolle 1859 ). – Type : PERU • 1778; H. Ruiz & J.A. Pavón s.n. ; lectotype: B [ B100507473 , F neg. 20808], designated by Moonlight & Jara-Muñoz (2017: 21) ; isolectotype: OXF . de Candolle (1864: 381) ; Schulz (1911: 27) . Begonia guyanensis var. cearensis C.DC. , Bulletin de L’Herbier Boissier II 1: 315 ( de Candolle 1901 ). – Type : BRAZIL – Ceará State • C. Huber 206 ; holotype: G . Smith & Wasshausen (1984: 467) . Begonia lokobeensis Humb. , Bulletin du Muséum National d’Histoire Naturelle Sér. 3, Botanique 1 (45): 80 ( Humbert 1972 ). – Type : MADAGASCAR • Nossi-Bé : Sambirano , Lokobe ; Nov. 1932 ; Perrier de la Bâthie 18708 ; holotype: P [ P00137255 ]; isotype: P [ P00137256 ] . Begonia alemanii Brade , Rodriguésia 9: 30 ( Brade 1945 ) . – Type : BRAZIL – Ceará State • F.Allemão & M. de Cysneiros 1352 ; holotype: R [ R000037072 , R000037072 a]; isotype: RB [ RB00536658 ] . Jacques & Mamede (2005: 582) . Begonia hirsuta Ruiz ex Klotzsch (nom. inval.; nom. rej. pro syn. Pilderia hirsuta Klotzsch (nom. inval.; nom. nud.) Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 127 ( Klotzsch 1854 ). de Candolle (1864: 381) ; Schulz (1911: 27) . Fig. 42. Distribution of Begonia sect. Doratometra (Klotzsch) A.DC. in Peru and surrounding countries. A . B. humilis Dryand. (red). B . B. semiovata Liebm. (red). Black lines indicate country borders, grey lines indicate major administrative divisions, blue lines indicate rivers, and shading indicates elevation. Pilderia hirsuta Klotzsch (nom. inval.; nom. nud.), Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 127 ( Klotzsch 1854 ) . Begonia haematotricha hort ex A.DC. (nom. inval.; nom. rej. pro syn. Begonia humilis var. porteriana (Fisch, C.A.Mey & Avé-Lall.) A.DC. ), Flora Brasiliensis 4 (1): 344 ( de Candolle 1861 ). Pilderia erythrotricha Klotzsch ex A.DC. (nom. inval.; nom. rej. pro syn. Begonia humilis var. porteriana (Fisch, C.A.Mey & Avé-Lall.) A.DC. ), Flora Brasiliensis 4 (1): 344 ( de Candolle 1861 ). Begonia hirsuta hort. ex A.DC. (nom. inval.; nom. rej. pro syn. Begonia humilis Dryand. ), Flora Brasiliensis 4 (1): 346 ( de Candolle 1861 ). Begonia hirsuta Pav. ex A.DC. (nom. inval.; nom. rej. pro syn. Begonia pavoniana A.DC. ), Prodromus Systematis Naturalis Regni Vegetabilis 15 (1): 381 ( de Candolle 1864 ). de Candolle (1864: 381) ; Schulz (1911: 27) . Begonia hitchcockii auct. non Irmsch: R.Vásquez et al. , Arnaldoa 12 (1–2): 112–125 ( Vásquez et al. 2005 ). Begonia humilis var. porteriana (Fisch., C.A.Mey. & Avé-Lall.) A.DC. pro parte in A.DC. Flora Brasiliensis 4 (1): 344 ( de Candolle 1861 ) ; pro parte in A.DC. Prodromus Systematis Naturalis Regni Vegetabilis 15 (1): 298 ( de Candolle 1864 ). Begonia semiovata Liebm. pro parte in R.Vásquez et al. , Arnaldoa 12 (1–2): 112–125 ( Vásquez et al. 2005 ). Etymology The epithet is formed from the Latin word ‘ humilis ’ meaning ‘lowly’ or ‘humble’ and refers to the small stature of the species. Selected specimens examined PERU – Loreto Region : Prov. Loreto • río Yanayacu , campamento Palizada ; 5°03′03″ S , 74°12′4″ W ; 90 m a.s.l.; 3 Nov. 1992 ; C. del Carpio & J. Ruiz 1633 ; USM . – San Martín Region : Prov. San Martín • Dist. Tarapoto , Prope Tarapoto ; [ 6°27′ S , 76°20′ W ]; Jun. 1855 ; R.E. Spruce 3982ª ; BM , K [2], NY , TCD • Dist. Tarapoto , 10.6 km from junction in road that goes to río Huallaga ; 6°30′ S , 76°21′ W ; 530 m a.s.l.; 10 Apr. 1984 ; T. B. Croat 58093 ; MO [ MO-2180398 ], USM • Route from Tarapoto to Tocache ; 7°44′36″ S , 76°40′03″ W ; 894 m a.s.l.; 6 Feb. 2016 ; P.W. Moonlight & A. Daza 188 ; E [ E00885890 ], MOL . – [ Prov. Tocache ] • Dist. Tocache Nuevo , Vicinity of Tocache ; 400–700 m a.s.l.; 1979; J. Schunke V. 11028 ; MO [ MO-1642628 ], US [ US00672844 ] • Dist. Tocache , Balsa probana (margen derecha del río Huallaga) ; [ 8°08′ S , 76°32′ W ]; 24 Sep. 1970 ; J. Schunke V. 4447 ; COL , G , NY [2], US [ US00222168 ] • Dist. Tocache , Quebrada de Pólvora , 10 km abajo de Puerto Pizana (margen derecha del río Huallaga) ; [ 8°07′ S , 76°32′ W ]; 1 Jun. 1970 ; J. Schunke V. 4950 ; NY [2], US [ US00222161 ]. – Ucayali Region : Prov. Padre Abad • Boquerón km 228 ; [ 9°04′ S , 75°42′ W ]; 460 m a.s.l.; 10 Aug. 1943 ; C.A. Ridoutt 12986 ; MO [ MO-1642528 ], USM [3] • Dist. Padre Abad , Boquerón de Padre Abad ; 9°04′47″ S , 75°42′10″ W ; 450 m a.s.l.; 8 Feb. 2016 ; P.W. Moonlight & A. Daza 202 ; E [ E00885888 ], MO , MOL . – Prov. Atalaya • road from Puerto Ocopa to Atalaya , ca km 107; 10°53′43″ S , 73°54′08″ W ; 623 m a.s.l.; 15 Feb. 2016 ; P.W. Moonlight & A. Daza 248 ; E [ E00885895 ], MO , MOL , P • 105 km from Las Cavernas on trail from Paujil ; 10°54′06″ S , 74°01′22″ W ; 633 m a.s.l.; 14 Feb. 2016 ; P.W. Moonlight & A. Daza 247ª ; E [ E00885887 ], G , MO , MOL . – Huánuco Region : Prov. Leoncio Prado • Dist. Rupu Rupu , Tingo María , limestone hills opposite airport ; 9°18′ S , 75°59′ W ; 9 Dec. 1981 ; T. Plowman , M. Ramírez R. & T.M. Perry 11243 ( USM • Dist. Mariano Dámaso Berau , Bello • [ 9°23′ S , 75°59′ W ]; 750 m a.s.l.; 31 May 1967 ; M. Fukushima 6530 ; HUT , US [ US00222159 ]. – Prov. Puerto Inca • Panguana ; 9°37′ S , 74°56′ W ; 21 Jan. 1983 ; F.G. Seidenschwarz 205/1 ; US [ US00424974 ]. – Huánuco - Ucayali Region • La Divisoria ; [ 9°10′ S , 75°47′ W ]; 5 Aug. 1965 ; A. Aldave & M. Fernández 5616 ; HUT . – Pasco Region : Prov. Oxapampa • Dist. Pozuzo , route from Pozuzo to Codo de Pozuzo ; 10°00′22″ S , 75°30′31″ W ; 655 m a.s.l.; 20 Feb. 2016 ; P.W. Moonlight & A. Daza 286 ; E [ E00885569 ], MO , MOL • Dist. Pozuzo ; [ 10°04′ S , 75°33′ W ]; 1778–1788; H. Ruiz & J.A. Pavón s.n. ; MA [ MA813500 ] • Dist. Palcazú , Atrás de la Escuela de nivel secundario San Pedro de Pichanaz , camino a la montaña ; 10°30′17″ S , 75°04′08″ W ; 633 m a.s.l.; 1 Apr. 2017 ; A. Orejuela & J. Castillo 2933 ; E [ E01053425 ], USM . – Junín Region : Prov. Chanchamayo • río Paucartambo Valley , near Perene Bridge ; [ 10°57′ S , 75°14′ W ]; 700 m a.s.l.; 19 Jun. 1929 ; E.P. Killip & A.C. Smith 25370 ; NY , US [ US0222309 ] • La Merced ; [ 11°04′ S , 75°20′ W ]; ca 700 m a.s.l., 29 May–5 Jul. 1929 ; E.P. Killip & A.C. Smith 23713 ; NY , US [ US00222312 ] • Hacienda Genoa ; [ 11°05′ S , 73°25′ W ] ca 1600 m a.s.l.; 2 Jul. 1962 ; F. Woytkowski 7361 ; K , MO [ MO-1642501 ], US [ US00222167 ]. – Prov. Tarma • San Luis ; 13 Aug. 1944 ; C.A. Ridoutt 14525 ; USM • Camino Ramazuy a San Luis ; 16 Aug. 1944 , C.A. Ridoutt s.n. ; USM [2]. – Prov. Concepción • San José ; 18 Aug. 1944 ; C.A. Ridoutt 14587 ; USM . – Prov. Satipo • La Molina University Field Station , ca 10 km N of Satipo on 5S , ca 1 km E of field station ; 11°10′12″ S , 74°58′54″ W ; 782 m a.s.l.; 20 Jun. 2014 ; P.W. Moonlight & A. Daza 21 ; E [ E00724455 ], MOL • 5 km SE of Satipo on road to Mazamari ; [ 11°16′ S , 74°36′ W ]; 620 m a.s.l.; 24 May 1979 ; D.C. Wasshausen & F. Encarnación 1093 ; K , MO [ MO-1642522 ], US [ US00222155 ], USM . – Ayacucho Region : Prov. La Mar • Along new road between Hacienda Luisiana and Santa Rosa Bridge ; 12°23′ S , 73°47′ W ; ca 585 m a.s.l.; 7 Jun. 1968 ; T.R. Dudley 9089 ; MO [ MO-1642527 ] • Along río Marantari , a tributary of río Apurimac, below Santa Rosa Bridge ; [ 12°41′ S , 73°43′ W ]; 580 m a.s.l.; 28 May 1975 ; D.C. Wasshausen & F. Encarnación 483 ; K , US [ US00222160 ] • Between Tambo , San Miguel , Ayna and the Hacienda Luisiana ; ca 12°52′ S , 73°47′ W ; 15 Aug. 1968 ; T.R. Dudley 11719 ; F, US [ US00222163 ]. – Madre de Dios Region : Prov. Madre de Dios • Manu , Parque Nacional del Manu , alrededores de la estación Cocha Cashu ; 11°52.8′ S , 71°22.9′ W ; 350 m a.s.l.; 12 Oct. 1983 ; Y. Mäkinen & R. Kalliola 1792 ; USM . Description Caulescent herb, to 50 cm high. Stem erect, branching; internodes to 5 cm long, to 5 mm thick, succulent, green to red, glabrous. Stipules persistent, lanceolate, 4–10 × 2–4 mm , apex acute, translucent, white, glabrous, margin entire, ciliate. Leaves > 3, alternate, basifixed; petiole 1–5.5 cm long, green to red, glabrous to villous; blade asymmetric, lanceolate, to 11.5 × 4.5 cm , succulent, acuminate, base obliquely truncate to obliquely cordate, basal lobes not overlapping, sinus to 6 mm deep, margin double serrate, ciliate, upper surface green, glabrous to sparsely pilose, lower surface pale green to red, glabrous, veins palmate but with 1 primary vein, 5–9 veined from the base, with 1–4 secondary veins on the larger side, 1–2 on the smaller side. Inflorescences 1–3 per stem, bisexual, axillary, erect, cymose, with 2 branches, bearing up to 4 staminate flowers and 4 pistillate flowers, protandrous; peduncle to 4 cm long, red, glabrous, bracts persistent, lanceolate to ovate, 1.5–5 × 0.5–2 mm , translucent, white, glabrous, apex rounded to acute, margin entire, ciliate. Staminate flowers : pedicels to 11 mm long, glabrous; tepals 2, spreading, circular, 2–4 × 2–4 mm , apex obtuse rounded, white, glabrous, margin entire, aciliate; stamens 8–10, projecting, yellow, filaments 0.5–1 mm long, free, anthers linear, 0.5–1.5 × 0.2 mm , dehiscing via lateral slits, connectives not extended, symmetrically basifixed. Pistillate flowers : pedicels to 7 mm long; bracteoles 2, directly beneath the ovary, lanceolate, 1.5–2 × 0.75–1.5 mm , translucent, white, glabrous, apex acute, margin entire, ciliate; tepals 5, subequal, persistent in fruit, spreading, elliptic to ovate, 1–2.5 × 0.5–2 mm , apex obtuse, white, glabrous, margin entire, aciliate; ovary body ovoid, 2–4 × 1–2 mm , white, glabrous, unequally 3-winged, wings triangular, largest 4–8 × 1–4 mm , smallest 2–7 × 4– 2 mm ; 3-locular, placentae branches entire, bearing ovules on both surfaces; styles 3, yellow, free, 1–2 mm long, once-divided, stigmatic papillae n a spirally twisted band. Fruiting pedicel to 10 mm long. Fruit body ovoid, to 9 × 6 mm , drying brown, wings same shape as in ovary, the largest expanding to 10 × 10 mm , the smallest expanding to 10 × 6 mm . Fig. 43. Begonia humilis Dryand. A . Habitat. B . Habit. C . Staminate flower. D . Pistillate flower flowers. All photographs taken by P.W. Moonlight from P.W. Moonlight & A. Daza 247a in Atalaya Province, Ucayali Region. Proposed conservation assessment Widespread and common in Peru and across its range, which includes much of the Caribbean and Amazonia. Frequent in disturbed areas, including next to roads, and found in numerous protected areas. We assess B. humilis as Least Concern (LC). Notes Smith & Schubert (1941a) described the placentae of B. humilis as varying from entire to divided. We have dissected several fruits of Peruvian individuals B. humilis and have only observed entire placentae. This may be due to confusion with the highly similar species B. hirtella , which has divided placentae, or simply because we have dissected a biased selection of ovaries. Synonymy notes Begonia haematotricha hort. Boiss and Pilderia erythrotricha Klotzsch were both cited by Alphonse Pyramus de Candolle as synonyms of his B. humilis var. porteriana (Fisch., C.A.Mey. & Avé-Lall.) A.DC. based upon annotations by Klotzsch in Berlin herbarium ( de Candolle 1861: 344 ). As such, they are both rejected names. We cite them in the synonymy of B. humilis because all specimens we have seen annotated with either name are clearly this taxon. De Candolle’s variety is excluded from this treatment (see Excluded names) but included as a pro parte synonym of B. humilis because of these specimens. The only known material of B. humilis collected by H. Ruiz and J.A. Pavón are three sheets housed in Madrid, Berlin, and Oxford herbaria. The sheets in Berlin and Oxford are labelled as B. hirsuta , a name they also used for our concept of B. hirta . Ruíz’s diaries state they wrote a description of B. hirsuta in Muña, but this probably applies to B. hirta , which they collected there. Ruiz’s diaries state that they wrote a description of B. purpurea earlier in Pozuzo. This likely refers to B. humilis because the habitat here is suitable and the name matches that on an illustration housed in Madrid. In 1788, the same year Ruiz and Pavón returned from their travels, O. Swartz published the name B. purpurea Sw. for a species found in Jamaica ( Swartz 1788: 86 ). Perhaps Ruiz and Pavón saw this publication and were aware this name was now unavailable because the herbarium sheet they prepared for Madrid herbarium is labelled as B. glabra . Unknown to most of the botanical world, this name was unavailable too due to the earlier publication of B. glabra Aubl. ( Aublet 1775: 916 ) , a name that remained overlooked throughout much of the 1800s. Ruiz and Pavon’s names B. glabr a, B. hirsuta and B. purpurea all remain unpublished names. Typification notes The protologue of B. humilis states the species is found in the West Indies ( Dryander 1789: 353 ), citing Scottish botanist Alex Anderson, who was at that time the superindendent of the government botanic garden in Saint Vincent . Anderson sent most of his herbarium specimens to Joseph Banks, and they are now in the herbarium of the British Museum. Despite a thorough search, we have not been able to find any specimens collected by Anderson in BM herbarium. The protologue of B. humilis also cited living material grown by James Lee and Lewis Kennedy in 1788, who had a nursery in Hammersmith in London. In their preparatory work for the Flora of Venezuela , Smith & Wasshausen (1979) stated that the type of B. humilis was “probably in BM” and in 1984 the same authors cited a specimen in the BM that was grown in Kew Gardens as the holotype . If this sheet can be located, then this citation is an effective lectotypification. Smith & Wasshausen (1984) were probably referring to a sheet with three specimens and three separate barcodes (BM001008469, BM001008470, BM001008471). The only text written on this sheet is written in pencil and says “Dryand! Begonia humilis Hort. Kew. iii 353”. This suggests that Dryander saw the sheet and used it to describe B. humilis ; however, this sheet is not cited in the protologue, so it is original material rather than a syntype . The herbarium of the Linnean society contains a sheet with two collections, the first grown in the “Garden at St Vincent” by “Mr Anderson” from seeds from Trinidad (LINN-HS 1474-3-1 ) and a second from “Mr Lee’s Garden, 1788” (LINN-HS 1474-3-2 ). These specimens match the citation in the protologue so are syntypes of B. humilis . Syntypes take priority over uncited original material in lectotype designation ( Turland et al. 2018 :Article 9.12) so it is appropriate to designate a lectotype from among the syntypes that supersedes Smith & Wasshausen’s effective lectotypfication ( Turland et al. 2018 : Article 9.19). We select A. Anderson s.n. (LINN-HS 1474-3-1 ) as the lectotype because its wild origins are known. Adrian Hardy Haworth described B. lucida Haw. as resembling B. humilis but with slightly different flowers, including a longer peduncle ( Haworth 1821a: 197 ). This description was based upon living material grown in Chelsea, London, and originating in Trinidad . Haworth’s specimens were distributed between Kew and Oxford herbaria, but most of the specimens in Oxford were discarded by Fielding. We have not seen any specimens resembling B. humilis in either Kew or Oxford that were cultivated in Chelsea or collected in Trinidad before 1821. It is therefore appropriate to designate a neotype of this name. We follow de Candolle (1861) and Schulz (1911) , who both included B. lucida as a synonym of B. humilis so select A. Anderson s.n. ; (LINN [LINN-HS 1474-3-1 ]) as a neotype . This is the lectotype of B. humilis so it ensures B. lucida will remain a synonym of B. humilis . Identification notes Recognised in Peru as a small annual or short-lived perennial with palmate-pinnate venation and stipules with long-ciliate margins. Distribution and ecology Known from Dominca, Jamaica , Saint Vincent , Trinidad and Tobago , French Guiana , Suriname , Guyana , Venezuela , Colombia , Ecuador , Peru , Bolivia , and Brazil . Within Peru , known from Loreto , San Martín , Ucayali , Huánuco , Pasco , Junín , Ayacucho , and Madre de Dios Regions ( Fig. 42A ). Found in Amazonia and lower montane forests at an elevation of 90–1500 m a.s.l. Begonia humilis appears to be an annual species and is typically found in the shade of dense forest along watercourses or on wet banks.