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 31. Begonia geraniifolia Hook. Figs 48A , 49 Curtis’s Botanical Magazine 62: t. 3387 ( Hooker 1835 ). – Eupetalum geraniifolium (Hook.) Klotzsch, Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 121 ( Klotzsch 1854 ). – Type : PERU – [ Lima Region : Prov. Lima ] • Amancaes ; [ 11°50′ S , 77°01′ W ]; A. Matthews 134 ; holotype: K [ K000252038 ]. Klotzsch (1855: 141) ; Walpers (1858: 875) ; de Candolle (1864: 281) ; Smith & Schubert (1941a: 188) ; Brako & Zarucchi (1993: 192) . Begonia petalodes Lindl. , The Botanical Register 21: t. 1757 ( Lindley 1835 ). – Eupetalum petalodes (Lindl.) Lindl., A natural system of botany, 2 nd ed. : 440 ( Lindley 1836 ). – Type : lectotype : t. 1757 in Lindl., The Botanical Register 21: t. 1757 ( Lindley 1835 ), designated here . Klotzsch (1855: 140) ; Walpers (1858: 875) ; de Candolle (1864: 281) ; Smith & Schubert (1941a: 189) . Eupetalum lindleyanum Gaudich., Voyage Autour du Monde, Exécuté Pendant les Années 1836 et 1837 sur la Corvette La Bonite, vol. 3(Atlas) : pl. 50 ( Gaudichaud-Beaupré 1842 ). – Begonia gaudichaudii Walp. (nom. illeg.; nom. superfl.), Repertorium Botanices Systematicae 5: 768 ( Walpers 1846 ) . – Type : PERU – Lima Region : Prov. Callao • San Lorenzo ; [ 12°05′ S , 77°13′ W ]; Jul. 1836 ; C. Gaudichaud-Beaupré s.n. ; lectotype: G , designated here ; isolectotypes: G , G-DC [2: F neg. 7328], K [2], MO [2: MO-2264392 , MO-2264405 ], P [3: P05586571 , P05586572 , P05586573 ], RB [ RB00053825 ], US [2: US00222150 , US00222151 ]. Walpers (1858: 875) ; Gaudichaud-Beaupré (1866: 18) ; de Candolle (1864: 281) ; Smith & Schubert (1941a: 189) . Eupetalum kunthianum Klotzsch, Gattungen und Arten 1854: 141 ( Klotzsch 1855 ) . – Type : PERU – Lima Region : Prov. Callao • San Lorenzo ; [ 12°05′ S , 77°13′ W ]; Jul. 1836 ; C. Gaudichaud-Beaupré s.n. ; lectotype: G , designated here ; isolectotypes: G , G-DC [2: F neg. 7328], K [2], MO [2: MO-2264392 , MO-2264405 ], P [3: P05586571 , P05586572 , P05586573 ], RB [ RB00053825 ], US [2: US00222150 , US00222151 ] . Walpers (1858: 875) ; de Candolle (1864: 281) ; Smith & Schubert (1941a: 189) . Eupetalum tuberosum Klotzsch, Gattungen und Arten 1854: 142 ( Klotzsch 1855 ) . – Type : PERU – [ Lima Region ] • ad Amancaes ; [ 11°50′ S , 77°01′ W ]; H. Ruiz & J.A. Pavón s.n. ; lectotype: B [ B100243038 , F neg. 20862], designated here . Walpers (1858: 875) ; de Candolle (1864: 281) . Begonia tuberosa Ruiz ex Klotzsch (nom. inval.; nom. rej. pro syn. Eupetalum tuberosum Klotzsch (nom. inval.; nom. nud.)), Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 121 ( Klotzsch 1854 ) . Eupetalum gaudichaudii Klotzsch (nom. inval.; nom. nud.), Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 121 ( Klotzsch 1854 ) . Eupetalum kunthianum Klotzsch (nom. inval.; nom. nud.), Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 121 ( Klotzsch 1854 ) . Eupetalum tuberosum Klotzsch (nom. inval.; nom. nud.), Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 121 ( Klotzsch 1854 ) . Eupetalum lindleyanum herb. ex Klotzsch (nom.inval.;nom. nud.), Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1854: 141 ( Klotzsch 1854 ) . Etymology Named for the superficial resemblance of the species’ leaves to those of geraniums and cranesbills ( Geranium L., Geraniaceae ). Selected specimens examined PERU – Cajamarca Region : Prov. Contumazá • Arriba del Rupe , siguiendo la carretera ChileteContumaza ; [ 7°17′ S , 78°49′ W ]; 1500 m a.s.l. ; 10 Mar. 1986 ; I. Sánchez Vega 4009 ; CPUN • Road between Cascas and Contumazá , 7°23′34″ S , 78°47′47″ W ; 2892 m a.s.l. ; 23 May 2015 ; M.C. Tebbitt & A. Daza 830 ; E , MOL • Yetón ; [ 7°23′ S , 78°48′ W ]; 1840 m a.s.l. ; 22 Jan. 1972 ; A.M. Lopez & A. Sagástegui 7897 ; HUT . – La Libertad Region : Prov. Trujillo • Cerro Prieto ; [ 7°55′ S , 78°48′ W ]; 675 m a.s.l. ; 14 Jul. 1949 ; N. Angulo 1017 ; F [ V0078822F ], HUT • Cerro Campana ; [ 8°07′ S , 79°05′ W ]; 600 m a.s.l. ; 10 Aug. 1985 ; J. Mostacero L. , L. Leiva G. , F. Mejía C. , F. Peláez P. 1076 ; HUT , MO [ MO-1642688 ], NY [2: NY01085843 , NY01085844 ] • Cerro Chuputur ; [ 8°11′ S , 78°56′ W ]; 400– 500 m a.s.l. ; 25 Jul. 1948 ; N. Angulo & A. López 0830 ; HUT . – Ancash Region : Prov. Santa • Lomas de Lupín , entre Barranca y Huarmey ; [ 10°21′ S , 77°56′ W ]; 400–500 m a.s.l. ; 29 Jul. 1959 ; R. Ferreyra 13791 ; MO [ MO-1642691 ], USM . – Lima Region : Prov. Chancay • Lomas de Lachay ; 11°20′ S , 77°30′ W ; 400 m a.s.l. ; 1 Sep. 1991 ; A.H. Gentry , C. Díaz , R. Ortiz & C. de Maestria 74500 ; MO [ MO-1643519 ] • ibid.; [ 11°22′ S , 77°21′ W ]; 300–500 m a.s.l. ; 1 May 1996 ; A. Cano & Y. Roque 7584 ; USM . – Prov. Huaral • On the Pan American Highway , km 78 N of Lima , 2 km east of the km stone , Lomas de San Juan San Cayetano de Chancayllo ; [ 11°28′ S , 77°18′ W ]; ca 100 m a.s.l. ; 27 Aug. 1964 ; P.C. Hutchinson 6301 ; F [ V0078820 F ], K , MO [ MO-2180399 ], NY , US [ US00222135 ], USM . – Prov. Lima • Lomas de Carabayllo entre los cerros San Diego y Parades ; [ 11°47′ S , 77°04′ W ]; 8 Jul. 2000 ; J. Roque , A. Cano , C. Cáceres & A. Ramirez 1626 ; USM • in monte Amancaes prope Lima ; [ 11°50′ S , 77°01′ W ]; 21 Jul. 1876 ; E. André 4088 ; F [ V0078819F ], K , NY • Quebrada de Manzano , trail to Cerro Menzano , E of Pachacamac ; 12°15′ S , 76°55′ W ; 300–600 m a.s.l. ; 14 Sep. 1986 ; S. Knapp 8328 ; MO [ MO-1642692 ]. – Prov. Huarochiri • Lomas El Manzano , Cucuya , Pachacamac ; [ 12°12′ S , 79°50′ W ]; 16 Jul. 1994 ; S. Llatas Q. 3508 ; HUT , UMS • San Bartolomia ; [ 11°55′ S , 76°32′ W ]; Jul. 1876 ; P.A. Savatier 1396 ; K . Fig. 48. Distribution of Begonia sect. Eupetalum (Lindl.) A.DC. in Peru and surrounding countries. A . B. geraniifolia Hook. (red) and B. joshii Moonlight (blue). B . B. neoharlingii L.B.Sm. & Wassh. (red) and B. weberbaueri Irmsch. (blue). Black lines indicate country borders, grey lines indicate major administrative divisions, blue lines indicate rivers, and shading indicates elevation. Fig. 49. Begonia geraniifolia Hook. Reproduction of the type illustration of Begonia petalodes Lindl. by Miss Sarah Ann Drake from Lindley (1835 : t. 1757) from the copy at the Royal Botanic Garden Edinburgh. Description Caulescent, rhizomatous herb, to 35 cm high. Rhizome ellipsoid, 2–5 × 2–4 cm , with 1–3 growing points. Stem erect, rarely branching; internodes to 15 cm long, to 10 mm thick, succulent, red, glabrous. Stipules persistent, broadly ovate, 6–10 × 4–12 mm , apex obtuse, opaque, red, glabrous, margin irregularly-dentate, aciliate. Leaves 4–6, alternate, basifixed; petiole 1–4 cm long, pale green to red, glabrous; blade subsymmetric, ovate, to 6 × 9.5 cm , succulent, apex indistinct, base truncate to cordate, basal lobes not overlapping, sinus to 10 mm deep, margin serrate, with up to 5 irregular, triangular lobes, aciliate, upper surface green, glabrous, lower surface pale green flushed red, glabrous, veins palmate, 6–8 veined from the base. Inflorescences 1–4 per stem, bisexual, axillary, erect, cymose, with 6 branches, bearing up to 4 staminate flowers and 2 pistillate flowers, protandrous; peduncle to 15 cm long, red, glabrous, bracts late-deciduous, elliptic to ovate, 2–8 × 1–4 mm , opaque, white to red, glabrous, apex acute, margin entire to serrate, aciliate. Staminate flowers : pedicels to 2 mm long, glabrous; tepals 4, spreading, outer 2 ovate to ovate, 8–10 × 6–9 mm , apex acute to obtuse, white to pink, glabrous, margin entire, aciliate, inner 2 obovate, 6–8 × 5–12 mm , apex truncate, often with two rounded apical lobes, pink, glabrous, margin entire, aciliate; stamens ca 20, spreading, yellow, filaments 1–2 mm long, free, anthers obovoid, 1–1.5 × 0.8 mm long, dehiscing via lateral slits, connectives extending to 0.2 mm , symmetrically basifixed. Pistillate flowers : pedicels to 20 mm long; bracteoles lacking; tepals 5 or rarely 6, subequal, deciduous in fruit, spreading, the largest ovate, 4–9 × 3–5 mm , apex truncate, white to pink, glabrous, margin entire, aciliate, the smallest elliptic, 3–7 × 2–4 mm , apex acute, white to pink, glabrous, margin entire, aciliate; ovary body ovoid, 2.5–3 × 3–4 mm , green to red, glabrous, sub-equally 3-winged, wings narrow-triangular, 2–3 × 5–10 mm ; 3-locular, placentae branches divided, bearing ovules on both surfaces; styles 3, yellow, free, 2–4 mm long, 4–6 times divided, stigmatic papillae in a spirally twisted band. Fruiting pedicel to 55 mm long. Fruit body obovoid, to 6 × 8 mm , drying brown, wings same shape as in ovary, expanding to 7 × 35 mm . Proposed conservation assessment Common in the Lomas surrounding Lima and Trujillo and likely also in less well collected Lomas between. It seems scarcer in its Andean range, but this probably reflects collecting biases; few botanists collect in the Andes during the wet season. The species’ EOO is> 30 000 km 2 and, while there has been some urban encroachment onto Lomas formations near Lima , we know of no specific threats to this species. We assess B. geraniifolia as Least Concern (LC). Notes To assess the sectional relationships of B. joshii , Moonlight et al. (2020) included a phylogenetic tree with two accessions labelled as B. geraniifolia . The resultant tree resolved B. geraniifolia as paraphyletic with B. joshii nested within, and with significant genetic difference between the two accessions of B. geraniifolia . They therefore recommended future authors give particular attention to the circumscription of this species. We now recognise many of the specimens previously recognised as B. geraniifolia as B. neoharlingii L.B.Sm. & Wassh. , including the accession “ B. geraniifolia 1” in Moonlight et al. (2020 : P.W. Moonlight & A. Daza 116 ). Our new circumscription of B. geraniifolia and B. neoharlingii are mostly geographically and environmentally distinct as well as morphologically distinct (see Identification notes). Begonia geraniifolia is mostly a species of coastal Lomas hills from La Libertad to Lima Regions at elevations < 500 m . In the north of its range, in Cajamarca and La Libertad Regions, it has infrequently been collected on the western slopes of the Andes at elevations up to 1840 m a.s.l. in valleys that trap fog in the wet season. In contrast, B. neoharlingii has been exclusively collected in the Andes at elevations above 1750 m a.s.l. and usually from 2000–3500 m a.s.l. Its habitat includes relict northwest Peruvian montane forest and areas of montane forest on the eastern slope of the Andes, but also includes areas that trap fog on the western slopes of the Andes. The specimen É.F. André 2376 (K) is B. geraniifolia but the specimen label says it was collected in Cauca , Colombia . We consider it extremely unlikely that this species occurs in Colombia and suggest this is likely an incorrect label. Sheets of B. geraniifolia collected by Ruiz and Pavón housed in B (B100243038, F neg. 20862, the lectotype of E. tuberosum Klotzsch. ex Klotzsch ), G-BOIS, and MA (MA813499) are labelled as “ Begonia tuberosa ”. The same name is used as part of a detailed description on the label of a sheet from Dombey’s herbarium now housed in P (P00679108). The annotations on the label of the sheet in MA suggest Ruíz, Pavón, and Dombey believed this was possibly B. tuberosa Lam. (or B. capensis L.f.). This is a rare case where the name used by Ruiz and Pavón matches the name on an illustration housed in Madrid (MA-AJB04-D-1389) by Isidro Gálvez, demonstrating a clear and unambiguous link between specimens and the illustration. Typification notes Begonia geraniifolia was described by W.J. Hooker in 1835 ( Hooker 1835 ), who cited material received from Mr Matthews and living plants grown in Glasgow and collected by Mr McLean in 1833. In 1835, Hooker was the director of Glasgow Botanic Gardens, and was appointed as director of the Royal Botanic Gardens Kew in 1841. He took much of his herbarium collection to London, which formed the basis of the original Kew herbarium (K). Specimens remaining in Glasgow were incorporated into the herbarium of the Royal Botanic Garden Edinburgh (E) in 1965. We have been unable to locate any material collected from McLean’s material in either Edinburgh or Kew and even if we did, we would not consider this syntype material as the protologue referred to living rather than herbarium material. There is a single sheet containing material of B. geraniifolia collected by Matthews in the herbarium at Kew ( Matthews 134 [K000252038]). This sheet includes five full plants of B. geraniifolia and two labels: the Matthews label and a label for plants collected by Mr W. Nation in 1862, after the description of B. geraniifolia . It is not absolutely clear which plants were collected by Nation and which by Matthews, so there is an argument for treating the Matthews material as lost and designating the only remaining original material (the illustration in the protologue) as a lectotype . We however believe the plant mounted closest to the Matthews label was almost certainly collected by Matthews. We therefore treat this specimen as the holotype of B. geraniifolia and the homotypic name Eupetalum geraniifolium (Hook.) Klotzsch. J. Lindley described B. petalodes Lindl. based upon material sent to him by Mr Richard Harrison in April 1833 ( Lindley 1835 ). He presumed the species was from Brazil , but the illustration provided within ( Fig. 49 ) clearly shows B. geraniifolia . A peculiarity of this plate is the unclear number of tepals on the pistillate flowers. Begonia geraniifolia always has five tepals on the pistillate flowers, but Lindley remarked in the protologue “two or four being the number of floral envelopes”. We believe the artist saw five tepals but, to make five tepals look like either two or four, drew an indistinct number of flowers. In all other respects, the plate clearly shows B. geraniifolia . The protologue of this species did not cite any herbarium material and we have been unable to locate any sheets that could reasonably be considered as original material. We therefore designate the plate in the protologue as the lectotype of B. petalodes . The first appearance of the name E. lindleyanum Gaudich. in the literature is in an 1842 article by C. Gaudichaud-Beaupré, where it is accompanied by an illustration that includes several important characters for identification ( Gaudichaud-Beaupré 1842 : pl. 50). This name is validly published because, prior to the 1 st of January 1908 , an illustration with analysis was acceptable in the place of a written description or diagnosis ( Turland et al. 2018 : Article 38.7) and figures separate from the main illustration of the organism showing details aiding identification constitutes analysis ( Turland et al. 2018 : Article 38.9). In 1866, Gaudichaud published text to accompany his 1842 publication, but for E. lindleyanum , this only included a caption for the plate ( Gaudichaud-Beaupré 1866: 18 ). A collection made by Gaudichaud in 1836 is deposited in various European herbarium, of which there are two duplicates in the main herbarium in Geneva (G). One of these is labelled as “ Eupetalum lindleyanum ”. We designate this as the lectotype of the name. In 1846, Walpers transferred Gaudichaud’s E. lindleyanum into the genus Begonia as B. gaudichaudii Walp. ( Walpers 1846: 768 ) . This is in contravention of the nomenclatural code as Walpers cited Eupetalum lindleyanum in synonymy when the name B. lindleyanum should have been adopted ( Turland et al. 2018 : Article 51.1). Walpers’ B. gaudichaudii is therefore an illegitimate superfluous name. The protologue of E. kunthianum cites material collected material collected in San Lorenzo and Lima and held in “herb. Kunthii” that is labelled as “ Eupetalum lindleyanum ” ( Klotzsch 1855: 141 ) . Kunth’s herbarium no longer exists, and its specimens have been distributed to various herbaria. The locality description matches a collection made by Gaudichaud on the voyage of the Bonite in July 1836 . The only duplicate of this collection that is labelled as “ Eupetalum lindleyanum ” is the same sheet held in G that we designate as the lectotype of E. lindleyanum . We therefore designate this specimen as the lectotype of E. kunthianum . In the protologue of E. tuberosum , Klotzsch cited material collected in Lima and San Juan by Ruiz and Pavón ( Klotzsch 1855: 142 ). There are no known herbarium sheets that match the description. Instead, it seems likely that Klotzsch misinterpreted the labels of material collected by Ruiz and Pavón in June 1776 in Lomas de Amancaes near Lima . These collections are labelled as “Flor de San Juan ”, which is the local name for B. geraniifolia ( Steele 1964: 80 ) . It seems Klotzsch misinterpreted “Flor de San Juan ” name as a locality. Material collected in Amancaes by Ruiz and Pavón is present in B, G, and G-BOIS but only the sheet in B (B100243038, F neg. 20862) has Klotzsch’s handwriting upon it, so we designate it as the lectotype of E. tuberosum . We do not consider the sheets in G and G-BOIS as isolectotypes because Ruiz and Pavón did not use collection numbers, so it is impossible to know whether these collections were made on the same day or at the same locality. Identification notes Begonia geraniifolia and B. neoharlingii are superficially similar to members of B. sect. Australes , which are also geophytic, caulescent herbs. They differ in their multifid rather than bifid styles. The two species are relatively simple to distinguish when flowering. Begonia neoharlingii has a pair of bracteoles directly beneath the ovary on the pistillate flower, whereas B. geraniifolia lacks bracteoles. The inner tepals of the staminate flowers of B. neoharlingii are rounded at the apex, while those of B. neoharlingii have two rounded lobes at the apex, and the staminate tepals of B. neoharlingii are much larger (8–30 × 9–12 mm vs 6–10 × 9 mm ). When sterile, B. neoharlingii can be distinguished as a generally larger plant with leaves with a crenate or rarely dentate margin (vs serrate) and only rarely with lobed leaves (vs usually with 3–5 triangular lobes). Distribution and ecology Endemic to Peru and known from Cajamarca , La Libertad , Ancash , and Lima Regions ( Fig. 48A ). Found in coastal Lomas formations at an elevation of 0–500 m a.s.l. and in dry forest and scrubland at elevations up to 1500 m a.s.l. Begonia geraniifolia is a geophytic species and dies back to its rootstock in the dry season. In the coastal Lomas, the wet season is from July to November and in the Andean Dry Shrublands it is usually March to May.