Introduction
As part of an ongoing taxonomic study of the Andean Begonia species assigned to Begonia sect. Eupetalum (Lindl.) A.DC. and Begonia sect. Knesebeckia (Klotzsch) A.DC. by Doorenbos et al. (Reference Doorenbos, Sosef and De Wilde1998), four species, B. hydrophylloides L.B.Sm. & B.G.Schub., B. triramosa Irmsch., B. velata L.B.Sm. & B.G.Schub. and B. xerophyta L.B.Sm. & Wassh., each originally described only from its type collection, were studied.
Begonia hydrophylloides was originally described from a single fragmentary specimen, consisting of one leaf and a portion of inflorescence bearing a few male flowers and one immature female flower. The specimen had been collected in 1917 by Francis W. Pennell in Cundinimarca Department, Colombia. An extensive search of herbaria located three additional collections, all made in Meta Department, Colombia. Smith & Schubert (Reference Smith and Schubert1946, p. 8) tentatively classify Begonia hydrophylloides in Begonia sect. Huszia (Klotzsch) A.DC., currently recognised as a synonym of Begonia sect. Eupetalum by Doorenbos et al. (Reference Doorenbos, Sosef and De Wilde1998), who thought that the relatively large size of the petiole and peduncle make it “extremely probable” that this species is a “scapose plant with a tuberous base”. Based on Smith & Schubert's (Reference Smith and Schubert1946) original description, Doorenbos et al. (Reference Doorenbos, Sosef and De Wilde1998) reclassified the species in Begonia sect. Knesebeckia but offered no explanation for the sectional transfer. Additional material of Begonia hydrophylloides located during the current study has provided a better understanding of the morphology of this species and hence its sectional affiliation. Begonia hydrophylloides is confirmed to be scapose but is found to have a fleshy rhizome rather than a tuber. Among Andean Begonia, fleshy rhizomes in combination with a scapose habit are typically characteristic of the B. octopetala group (Begonia sect. Eupetalum). However, the many-flowered dichasial cymes of this species, with up to four branches, and the subequal fruit wings do not fit that group and instead suggest that it is most likely affiliated with the members of the polyphyletic Begonia section Knesebeckia.
Begonia triramosa was originally described from a single fragmentary specimen, consisting of a short apical portion of stem with a single leaf and two detached inflorescences bearing female flowers. The specimen had been collected in 1918 by Joseph N. Rose and George Rose in Chimborazo Province, Ecuador. Examination of the type specimen found that it lacks any features that separate it from Begonia acerifolia Kunth, and that it has the characteristic combination of thick stem, triangular lobed leaf blades, few-flowered inflorescences with a long peduncle and short branches, female flowers with five tepals, and markedly asymmetrical ovary wings found in that species. Fieldwork conducted in January 2014 and May 2016 at the type locality and surrounding area confirmed that Begonia acerifolia grows in this location (e.g. Tebbitt & Gutierrez 859).
Begonia xerophyta was originally described from a single fragmentary specimen, consisting of just the very top portions of two stems bearing two or three leaves, and a few mature male flowers and immature female flowers. The specimen had been collected in 1955 by Erik Asplund at 2100 m elevation between San Lucas and Loja in Loja Province, Ecuador. Fieldwork conducted in January 2014 and May 2016 found that the only Begonia species present in this area was B. acerifolia. Subsequent examination of the holotype of Begonia xerophyta showed that it was synonymous with B. acerifolia, as indicated by the combination of its unusually thick stems; shortly triangular lobed leaf margins; leaf blade undersurfaces with white tomentose hairs; few-flowered inflorescences; male flowers with four relatively large tepals, of which the outer pair are elliptic and red without and white within, and the inner pair broadly obovate and white to very pale pink; and female flowers with five tepals. The relatively high altitude (2100 m) of this collection site, coupled with the relatively dry slope on which the plants were growing, is also characteristic of Begonia acerifolia, which occupies an unusual habitat for a Begonia, the majority of species being found at lower elevation in more humid situations.
During the course of examining material of Begonia acerifolia for this study, it was also determined that B. erythrocarpa could not be distinguished from B. acerifolia. This discovery was surprising, because the type specimens of Begonia acerifolia and B. erythrocarpa differ substantially. Most notably, the type of Begonia acerifolia, which was collected in Loja Province, Ecuador, has cordate leaf bases and tepals with acute apices and serrate margins, whereas the type of B. erythrocarpa, which was collected in La Paz Department, Bolivia, has peltate leaf blades and tepals with obtuse apices and entire margins. Furthermore, fieldwork has shown that plants from southern Peru and Bolivia additionally differ in having thicker leaf blades with a tomentose covering of silvery white hairs on both surfaces and a fruit wing with a rounded apex. In contrast, the leaves of plants in Ecuador and northern Peru are often thinner and pubescent, and the hairs a paler white and therefore much less prominent, and the longest wing apex is subacute. However, fieldwork conducted throughout the range of these taxa, as well as examination of available herbarium material, shows that all these character combinations represent extremes in a single, highly variable species. When material in addition to the types is examined, every possible combination of these characters is observed. Furthermore, some populations in northern Peru contain individuals with either peltate or cordate leaf bases and/or either entire or serrate tepals. Similarly, greenhouse experiments have shown that seed collected from a population in Chota Province, Cajamarca Region, Peru, that closely matched the morphology of the Begonia erythrocarpa type collection, when grown, produced a mixture of progeny, some plants closely resembling the B. acerifolia type collection and others the B. erythrocarpa type collection (P. W. Moonlight, Royal Botanic Garden Edinburgh and University of Glasgow, personal communication). The variable nature of this species is further highlighted by the large number of synonyms previously recognised under Begonia erythrocarpa (Golding & Wasshausen, Reference Golding and Wasshausen2002), and which are transferred here to B. acerifolia. Begonia acerifolia as circumscribed here is therefore a common, widespread and highly variable species that occurs from the Andes of central Ecuador to northern Bolivia. Begonia acerifolia is currently classified in the polyphyletic Begonia sect. Knesebeckia (Doorenbos et al., Reference Doorenbos, Sosef and De Wilde1998; Tebbitt, Reference Tebbitt2016).
Begonia velata was originally described from a single collection composed of five sheets bearing male and female flowers. The collection was made in 1912 by Augusto Weberbauer in Píura Region, Peru, along the mountain pass leading from Palambla to Huancabamba. A combination of herbarium-based studies and fieldwork found that the species is relatively abundant at this site and also occurs much less frequently at Bosque Cachil in Contumazá Province, Cajamarca Region, Peru. Interestingly, this unusual disjunct distribution, which spans the Huancabamba deflection, is also mirrored by Begonia pseudopleiopetala Tebbitt (Tebbitt, Reference Tebbitt2015). Molecular data (P. W. Moonlight et al., unpublished) indicate that Begonia velata is nested within B. acerifolia, suggesting that the former may have evolved from within B. acerifolia. Given the morphology of Begonia velata, it is suggested that this species may have arisen as a stabilised hybrid between B. acerifolia and B. ludwigii Irmsch. Evidence for this comes from the observation that Begonia velata has a combination of the morphological characteristics found in B. acerifolia and B. ludwigii. In general, Begonia velata is morphologically very similar to B. acerifolia; however, it shares with B. ludwigii a symmetrical leaf blade, which is circular in outline (ovate in B. acerifolia), and also has a full or partial ring of hairs at the top of the petiole near the point of attachment to the leaf blade. The latter feature is unusual within Andean Begonia, being found within Peruvian Begonia sect. Knesebeckia only in B. ludwigii. Begonia ludwigii was originally described from Ecuador (Irmscher, Reference Irmscher1937) and has not previously been documented from Peru (Smith & Schubert, Reference Smith, Schubert and Macbride1941; Brako & Zarucchi, Reference Brako and Zarucchi1993). Recent fieldwork, however, found that Begonia ludwigii occurs both in Ecuador and in northern Peru, including in the vicinity of La Florida, Cajamarca Region, Peru (e.g. Tebbitt & Daza 846 (MOL), 6°52′S, 79°05′W). Furthermore, it hybridises with Begonia acerifolia, where the distributions of these two taxa overlap (e.g. Tebbitt & Daza 849 (MOL), 6°53′S, 79°03′W). Given that hybrid speciation is thought likely to have been an important mechanism of species formation in Begonia (Dewitte et al., Reference Dewitte, Twyford, Thomas, Kidner, van Huylenbroeck, Grillo and Venora2011), but that very little supporting data has been collected from natural populations of this genus, it would be of interest to investigate the potential hybrid origin of B. velata further.
Taxonomic Treatment
Begonia hydrophylloides L.B.Sm. & B.G.Schub., Caldasia, 4, no. 16: 7, 1946. – Type: Colombia, Cundinamarca, Moist bank, Guayabetal to “Monte Redondo” (= Monterredondo) south-east of Quetame, 1300–1500 m, 6 ix 1917, F.W. Pennell 1803 (holo NY!).
Acaulescent herb, with a fleshy rhizome. Rhizome probably horizontal but only known from a fragment, 2–4 cm in diameter. Stipules persistent, ovate 0.5–2 × 0.3–1.2 cm, apex acute, margin entire. Leaves few, alternate, basifixed; petiole ± continuing straight into main vein of blade, 20–50 cm long, glabrous; blade subsymmetrical, broadly ovate to almost orbicular in outline, 14–25 × 20–24 cm, base cordate, basal lobes spreading, sinus 2.5–5 cm deep, apex acute, margin triangular lobed, lobes 2–6.3 cm deep, lobes denticulate, teeth tipped with a short hair, membranous (when dried), green on both surfaces, upper surface glabrous, lower surface sparsely pubescent or glabrous, veins palmate, 6–8. Inflorescence solitary, erect, a dichasial cyme, with up to 4 branches, many-flowered, bisexual but strongly protandrous; peduncle 40–80 cm long, primary branches 0.6–3 cm long, secondary branches 0.35–1.5 cm long, glabrous; pedicels of male flowers 1.3–3.5 cm long, glabrous; pedicels of female flowers not observed; bracts deciduous, ovate or elliptic, 0.5–1.6 × 0.2–1.2 cm, apex acute, apiculate, margin entire, surfaces glabrous. Male flowers: tepals 4, spreading, white, white with a pink tinge, or pink, glabrous, outer two broadly ovate, 9–11 × 7–9 mm, apex acute, margin entire, glabrous, inner pair oblong, c.8 × 2 mm, apex obtuse, margin entire, glabrous; stamens c.50, attached along the length of a c.1.5 mm long torus, filaments 2–3 mm long, anthers symmetrically basifixed, obovoid, c.0.75 mm long, dehiscing by long unilateral slits, connectives not extended. Female flowers: (known only from immature specimens) bracteoles absent; tepals persisting in fruit, tepals 5, spreading, same colour as males, ovate, subequal, apex obtuse, margin entire; ovary 3-winged, wings subequal; styles 3, bifid. Fruiting pedicel to 4 cm. Fruit erect, body ellipsoid, to 1 × 0.7 cm, glabrous, longest wing marginiform, to 1 cm long, apex obtuse, shorter two wings marginiform, to 0.65 cm long, apex obtuse.
Phenology
Flowering August to December.
Distribution
Foothills of western Cordillera Oriental in central Colombia (Cundinamarca and Meta).
Habitat
Found at 500–1500 m, on shady, moist slopes in montane forest, often by rivers.
IUCN conservation category
The area of Colombia where this species occurs has seen limited botanical activity in recent years, so it is likely that the species is both more widespread and more common than is suggested by the low number of current collections. The lack of information related to both the distribution and the abundance of this species necessitates assigning an IUCN (2016) category of Data Deficient (DD).
Additional specimens examined. Colombia. Meta: Sabanas de San Juan de Arama, margen Izquierda del río Güejar, alrededores del alerrizaje “Los Micos”, 14 ii 2007, J.M. Idrobo 560 (COL); San Juan de Arama, margen Izquierda del río Güejar, paso de “Los Puercos”, 24 viii 2007, J.M. Idrobo 471 (COL [2]); Parque Nacional Natural Tinigua, Río Duda, Serranía Chamusa, Centro de Investigaciones Ecológicas La Macarena, Trocha Baño Chamusa, ca mirador W, D.W. Stevenson 1884 (NY).
A photograph of the holotype (Pennell 1803) of Begonia hydrophylloides is available via the Begonia Resource Centre (Hughes et al., Reference Hughes, Moonlight, Jara, Tebbitt, Wilson and Pullan2015–).
Begonia acerifolia Kunth, Nov. Gen. Sp. (quarto ed.) 7: 186, t. 644, 1825. – Type: Ecuador, [Loja Province], Loxam, 1060 m, A.J.A. Bonpland 3333 (holo P! [barcode: P00679517], photo E! [barcode: E00157074], photo G! [barcode: 358682]).
Begonia erythrocarpa A.DC., Ann. Sci. Nat. Bot., IV, 11: 121, 1859. – Type: Bolivia, Dep. La Paz, Prov. Larecaja, v 1847, H.A. Weddell 4729 (lecto P! [barcode: P01900755], designated here; klepto G-DC, isolecto P! [barcode: P01900754]), syn. nov.
Begonia dolabrifera C.DC., Bull. Herb. Boissier, sér. 2, 8: 324, 1908. – Type: Ecuador, Pallatanga, [2°1′S, 78°58′W], ix 1891, L. Sodiro 594 (holo G! [barcode: G00237358] [photo K!, F!, MO!], iso F [barcode: v0052627F], iso G, iso P! [barcode: P01900768], iso QPLS!).
Begonia griseocaulis Irmsch., Biblioth. Bot., 116: 112, 1937. – Type: Ecuador, Prov. Chimborazo, Tal des Rio Chanchan, oberhalb Huigra, um 1600 m, 23 ix 1933, L. Diels 1173 (holo B), syn. nov.
Begonia pennellii L.B.Sm. & B.G.Schub., Publ. Field Mus. Nat. Hist, Bot. Ser. 13: 196, 1941. – Type: Peru, Cuzco Region, Pillahuata, Cerro de Cusilluyoc, thickets above Pillahuata, 13°10′30′′S, 71°35′53′′W, 2800–3100 m, 3–6 v 1925, F.W. Pennell 14126 (holo GH!, iso B), syn. nov.
Begonia triramosa Irmsch., Bot. Jahrb. Syst. 74: 613, 1949. – Type: Ecuador, vicinity of Huigra, mostly on the Hacienda de Licay, 3 ix 1918, J. N. Rose & G. Rose 22493b (holo US!, photo NY!), syn. nov.
Begonia pennellii subsp. lobato-ovata Irmsch., Bot. Jahrb. Syst., 76: 84, 1953. – Type: Peru, Cuzco Region, Prov. Calca, Distr. Lares, Lares valley above Mantoc, 2400–2500 m, 8 iii 1929, A. Weberbauer 7906 (syn B, BM!, F, GH!), syn. nov.
Begonia pennellii var. longiloba Irmsch., Bot. Jahrb. Syst., 76: 85, 1953. – Type: Peru, Prov. Huanta, Dep. Ayacucho, Weg von Tambo über Osno zum Flusse Apurimac, 2700–2800 m, 1 vi 1910, A. Weberbauer 5622 (syn B, F; photo F!, K!, P [barcode: P06841739]) syn. nov.
Begonia pennellii forma macrantha Irmsch., Bot. Jahrb. Syst., 76: 86, 1953. – Type: Peru, Píura, Prov. Ayabaca, Westabhänge der Anden über Frias, Geogr. Br. 4°50′–5°S, 2900–3000 m, 20 v 1912, A. Weberbauer 6402 (holo B!), syn. nov.
Begonia lobato-peltata Irmsch., Bot. Jahrb. Syst., 76: 86, 1953. – Type: Peru, Puno Region, Prov. Sandia, 10 vii 1903, A. Weberbauer 503 (lecto B! [barcode: B100243008] designated here; isolecto B! [barcode: B100243009], G!), syn. nov.
Begonia macbrideana Irmsch., Bot. Jahrb. Syst., 76: 87, 1953. – Type: Peru, Junín Region, Tarma, entre Huacapistana y Palca, [11°14′S, 75°32′W], 2200 m, A. Weberbauer 2011 (holo B, iso MOL!, iso herbarium of Universidad Nacional Agraria La Molina!), syn. nov.
Begonia xerophyta L.B.Sm. & Wassh., Phytologia, 44: 245, pl. 7, 1979. – Type: ECUADOR, Prov. Loja, between Loja and San Lucas, very steep and dry slope, c.2100 m, [3°45′S, 79°15′W], 8 x 1955, E. Asplund 18036 (holo S!), syn. nov.
Caulescent, rhizomatous herb. Rhizome horizontal, 7–20 cm long, 0.75–1.25 cm in diameter, unbranched. Stem erect to sprawling, becoming somewhat woody at maturity, usually 0.5–2 m tall, 0.75–1.25 cm in diameter, unbranched, internodes 1–20 cm long, usually glabrous, occasionally tomentose. Stipules persistent, ovate, 1.5–1.8 × 0.8–1.4 cm, apex subacute, margin entire. Leaves alternate, basifixed or peltate; petiole usually joining blade at a transverse angle but occasionally continuing straight into main vein of blade, 5.5–13 cm long, glabrous; blade juicy to subcoriaceous, asymmetrical, usually ovate or occasionally suborbicular in outline, 5.5–18 × 4.5–20 cm, base peltate or cordate, basal lobes when present overlapping to spreading, sinus to 4.5 cm deep, apex acute, margin usually deeply 6- to 8-lobed, rarely with 6–8 shallow angular lobes, lobes usually narrowly triangular, 1–5.5 cm deep, serrulate to serrate, teeth to 1.5 mm long, teeth sometimes tipped with a short hair, upper surface usually pale green, occasionally olive green with silvery white stripes along part of or the entire length of the main veins, glabrous or moderately pubescent, hairs white, simple, lower surface green or red, moderately pubescent or tomentose, hairs white to silvery white, veins palmate, 6–9. Inflorescences few, axillary from the upper portion of the stem, erect, symmetrical cymes, 2- or 3-branched, branches 0.9–6.5 cm, usually bearing 12 flowers, bisexual, male flowers beginning to open before the first female flowers but both sexes eventually open concurrently; peduncle 15–40 cm long, glabrous; pedicels of male flowers 1.5–6 cm long, glabrous to tomentose; pedicels of female flowers 0.5–1.65 cm long, glabrous to tomentose; bracts caducous, ovate, 1.1–1.6 × 0.9–1 cm, apex obtuse, margin entire. Male flowers: tepals 4, spreading, white, sometimes reddish pink flushed on the outer surfaces or rarely both surfaces, outer two transversely ovate to broadly elliptic, 2.1–2.4 × 1.85–2.4 cm, apex subacute to rounded, margin entire to serrate, glabrous, inner two elliptic or obovate to rhomboid-obovate, 0.7–2.5 × 0.85–1.8 cm, apex subacute to rounded, margin entire to serrate, glabrous; stamens 50–100, attached along the length of a 2–3 mm long torus, filaments 1–2.5 mm long, anthers symmetrically basifixed, obovoid to obovoid-cuboid, 1–2 mm long, dehiscing by unilateral slits, connectives not extended. Female flowers: bracteoles absent; tepals not persisting in fruit, 5, spreading, same colour as males, usually elliptic to rhomboid, inner one or two sometimes obovate, subequal, 1.5–1.9 × 0.9–1.7 cm, glabrous, apex subacute to rounded, margin entire to serrate; ovary body ellipsoid to broadly ellipsoid, 5.25–15 × 5–9 mm, pale green to pink, glabrous, with one long wing and two rib-like wings, longest wing ligulate, ligulate-triangular or triangular, apex rounded to subacute, to 2.4 × 2.1 cm, 3-locular; placentae bifid, bearing ovules on both surfaces of placental branches; styles 3, shortly fused at base, 5–7 mm long, bifid from about one-third to almost half their height, branches erect, stigmatic papillae in a once-spiralled band. Fruiting pedicel to 1.2 cm long. Fruit subnutant, body broadly ellipsoid, to 1.5 × 1 cm, glabrous, wings as in ovary.
Phenology
Typically flowering March to July, with a peak during April and May, but producing a few flowers throughout the year.
Distribution
Andes of Ecuador (Azuay, Bolivar, Cañar, Chimborazo, El Oro and Loja), Peru (Amazonas, Cajamarca, Cuzco, Junín, Píura and Puno) and Bolivia (La Paz).
Habitat
Locally common and often forming large populations in open disturbed sites, such as roadsides, pastureland or along trails in disturbed montane forest. Typically growing at altitudes between 2000 and 3100 m but down to 1500 m and up to 3250 m. Usually terrestrial and growing in soil or moss but occasionally epiphytic on the trunks of trees and shrubs. Frequently growing among tall herbs or shrubs and using these for support.
IUCN conservation category
Begonia acerifolia is a widespread and common species that prefers disturbed areas and is therefore assessed as Least Concern according to IUCN criteria (2016).
Representative specimens examined. Ecuador. Bolívar: road Balsapampa–San Miguel, La Guardia, [1°48′S, 79°6′W], 2500 m, 16–17 v 1968, G.W. Harling, G. Storm & B. Ström 9571 (GB, MO, NY). Chimborazo: Juan de Velasco – Pallatanga, km 9, 1°52′S, 78°54′W, 2610 m, 22 v 1990, P.M. Jørgensen, R. King, P. Peterson & E. Judziewicz 91827 (AAU, MO, QCA, QCNE); Pallatanga, Comunidad Jesús del Gran Poder, from Panza Redonda (South Pan American Highway), 2 km S on cobble/dirt road, c.4 km NE of Pallatanga, 1°58′S, 78°56′W, 2800–3200 m, 10–14 vii 1995, J.L. Clark, Z. Aguirre, B. Greene, T. Harris & C. Watt 1263 (MO, QCNE); Pallatanga, x 1858, R. Spruce 5539 (K [2] = paratype: B. griseocaulis Irmsch.); Cañon of the río Chanchan, about 5 km N of Huigra, 5000–6500 ft, 19–28 v 1945, W.H. Camp E-3435 (BM, NY); Cantón Alausi, roadside bank between Alausi and Huigra, 02°14′S, 78°55′W, 2108 m, 20 v 2016, M.C. Tebbitt & D. Gutierrez 859 (QCA); Alausi Canton, parroquia Huigra, sector Chasmay, 2°16′28′′S, 78°59′21′′W, 1500 m, 27 v 2006, C.E. Ceron & C.I. Reyes 57504 (MO [2]); Prov. Chimborazo, Chimborazo–Cañar border, between Sta. Rosa (8300 ft) and Joyagshi (9000 ft), 6 vii 1945, W.H. Camp E-4023 (G [2], GH [2], K [2], MO [2], NY, P [2], US). Cañar: between Suscal and Chontamarca, north rim of the valley of the Río de Cañar, [2°26′S, 79°4′W], 23 iv 1943, W.H. Camp E-2853 (GH); route de Zhud à la Côte, 1 km après l'embranchement de Zhud, 2°29′S, 79°1′W, 2700 m, 19 v 1988, C. Huttel 1339 (QCA, QCNE). Azuay: between Río Molleturo at Huigra and Cruz Pamba crossing Río Mehuir, north of Molleturo, 1645–2315 m, 2 vi 1943, J.A. Steyermark 52926 (GH); west of Patul 3 kms between Huahualcay and Río Patul below Pasas de Pinglion, 2670–3275 m, 19 v 1943, J.A. Steyermark 52609 (GH, NY); Portete del Tarqui, Cuenca-Girón, 3°6′10′′S, 79°7′41′′W, 2700–2950 m, 5 iv 1974, G. Harling & L. Andersson 13223 (GB, MO); Vallée du rio Jubones, route de Pasaje a Cuenca, 8 km après Giron, 3°17′S, 79°20′W, 2550 m, 9 v 1988, C. Huttel 1110 (QCA, QCNE). El Oro: along Quebrada de Pampa de Los Cedros, tributary to Río Palma, NE of San Pablo, 2285 m, 12 viii 1943, J.A. Steyermark 53801 (GH); Cantón Atahualpa, Piñas to Machala road, c.2 km S of Mirador, 03°32′S, 79°40′W, 26 v 2016, 2119 m, M.C. Tebbitt & D. Gutierrez 865 (QCA). Loja: Cerro de Chilla, Saraguro-Manú, Km 45.4, 3°33′37′′S, 79°22′12′′W, 2760 m, 2 v 1994, P.M. Jørgensen, C. Ulloa, H. Vargas & G. Abendaño 703 (AAU, MO, NOLS, QCA, QCNE); road Loja – Saraguro, km 30 La Contradia, 3°51′S, 79°14′W, 2600 m, 7 ix 2000, J.E. Madsen & P. Lozano 7225 (AAU, QCNE); road from Celica to Alamor, northern and shorter route, passing through Mercadillo, km 8.75 from Plaza Central in Celica, c.3°58′S, 80°00′W, 1987 m, 5 v 1989, J. Smith 1968 (QCA, QCNE); summit and upper slopes of Cerro Villonaco, [3°58′8′′S, 79°16′1′′W], 2600–2750 m, 12 iv 1974, G.W. Harling & L. Andersson 13446 (GB, NY); old road Loja–Catamayo, c. km 10, 2700 m, 24 v 1988, J.E. Madsen, B. León, K. Young & S.S. Renner 74414 (AAU, MO, NY, QCA, QCNE); Loma Santiago, 3 km S of Santiago and 14 km S of San Lucas, 2900 m, 4°0′S, 79°58′W, 2900 m, 18 ii 1993, G. Harling & B. Ståhl 26489 (GB, QCA); Loja – Las Palmas, Cerro El Tambo, just S of Cerro Villonaco, 4°4′S, 79°14′W, 2750–3020 m, 23 vii 1990, P.M. Jørgensen, C. Ulloa, M. Gavilanes, P. Mena & L. Suarez 92094 (QCA, QCNE); Parque Nacional Podocarpus, 4°5′S, 79°11′W, 2750–2850 m, 15 v 1988, B. Øllgaard, J.E. Madsen & L. Christenson 74206 (AAU, E, MO, NY, QCA, QCNE); Nudo de Cajanuma, 7 km S of Loja, [4°5′36′′S, 79°12′W], 8000–8400 ft, W.H. Camp E-121 (GH, NY); Env. 10 km au sud de Loja sur la route de Malacatos, 4°14′S, 79°15′W, 2250 m, 12 v 1988, C. Huttel 1181 (QCA, QCNE); Cía. Agrominera Curishiro, cerca a Parque Nacional Podocarpus, 4 horas a pie desde Masanamaca, 4°16′S, 79°5′W, 2400 m, 2–4 iv 1990, P. Mena 2745 (QCA); Cerro Sozoranga, Colaisaca-Utuana, Km 0.5, 4°19′14′′S, 79°41′16′′W, 2340 m, 24 iv 1994, P.M. Jørgensen et al. 569 (AAU, MO, QCA, QCNE); Sur la route de Lauro Guerrero à Orianga, versant occidental très humide et souvent couvert de brouillard du Cerro Negro, 4°20′S, 79°5′W, 2450 m, 15 v 1988, C. Huttel 1267 (MO, QCA, QCNE); Huacabamba, 1 xi 1876, É-F. André 4396 (NY [photo MO]).
Peru. Píura: Prov. Ayabaca, Distr. Ayabaca, Las Lomas to Ayabaca road, above Chinchinpampa, 4°40′S, 79°45′W, c.2710 m, 28 v 2015, M.C. Tebbitt & A. Daza 843 (MOL); Prov. Ayabaca, Yacupampa-Cuyas (Ayabaca), 2500 m, 26 v 1971, A.M. Lopez, H. Fabris, A. Sagástegui & H. Aguado 7777 (HUT); Prov. Huancabamba, above Canchaque on the Huancabamba Pass, 5°22′39′′S, 79°33′32′′W, 2371 m, 27 i 2016, P.W. Moonlight & A. Daza 120 (E); Cerro Porculla 3 km to 11 km W of Continental Divide down W coast of Cerro, 5°51′41′′S, 79°31′14′′W, 12 vi 1966, G. Edwin & J.V. Schunke 3763 (USM). Amazonas: Prov. Chachapoyas, ladera del Cerro Tambo Viejo, road between Balsas and Leymebamba, 6°47′S, 77°56′W, 2810 m, 24 v 2015, M.C. Tebbitt & A. Daza 831 (E, MOL). Cajamarca: Prov. Chota, near Las Palmas, c.24 km NE of Chota, 6°9′28′′S, 78°35′47′′W, 2024 m, 25 vii 2014, P.W. Moonlight & A. Daza 68 (E); Prov. Santa Cruz, Distr. Pulan, El Cedro, 6°47′35′′S, 79°5′16′′W, 2600 m, 31 i 2008, L. Santa Cruz 2113 (USM); Santa Cruz Province, c.3 km (por aire) ENE Monteseco, [6°50′36′′S, 79°6′18′′W], 1800 m, 9 v 1987, J.C. Santisteban C. & J.B. Guevara B. 34 (F, HUT, MO, NY); Prov. San Miquel, Distr. La Florida, above La Florida, 6°53′S, 79°03′W, 1787 m, 30 v 2015, M.C. Tebbitt & A. Daza 847 (MOL); Prov. San Miguel, Bosque de Quellahoreo, de noreste de la localidad de Tongod, 2700 m, 14 ix 1991, J. Sánchez Vega & A. Brónes 5800 (CPUN). Cuzco: Prov. La Convención, Distr. Santa Ana, Madre Selva, 12°53′49′′S, 72°45′2′′W, 1500 m, 21 iii 2004, L.G. Valenzuela, E. Suclli & G. Calatayud 3090 (MO); Prov. La Convención, Distr. Maranura, Mesa Pelada, 12°32′S, 72°22′W, 2547 m, 20 iv 2005, L. Valenzuela, E. Suclli, I. Huamantupa, J. Farfán, L. Cardenas, V. Chama & J. Latorre 5546 (AMAZ, CUZ, HUT, MO, MOL, US, USM); Prov. La Convención, Distr. Vilcabamba, Vilcabamba, Pajonal, 13°7′27′′S, 72°58′55′′W, 3470 m, 31 v 2002, W.L. Galiano, L.G. Valenzuela, E. Suclli, I.C. Huamantupa & A. Carazas 4108 (MO); Prov. La Convención, Distr. Huayopata, Balconpata, 12°52′1′′S, 72°32′46′′W, 2200 m, 15 iv 2004, G. Calatayud, I. Huamantupa, L. Cardenas, H. Coasaca & E. Apaza 2178 (CUZ, MO, US); Prov. Calca, Distr. Yanatile, above intersection of roads coming from Amparaes, Quebrada Honda, and Lares, along road to Amparaes, 12°58′S, 72°03′W, 3660 m, 5 i 2015, M.C. Tebbitt & A. Daza 797 (MOL); Prov. Urubamba, Distr. Ollytaytambo, Road from Quillabamba to Ollytaytambo, 13°5′7′′S, 72°22′52′′W, 3173 m, 7 viii 2014, P.W. Moonlight & A. Daza 96 (E); Prov. Urubamba, Huiñaihuaina, [13°11′S, 72°32′W], 3000 m, 22 vi 1948, C. Vargas C. 7243 (CUZ, US); Prov. Urubamba, on trail between Machupichu–Lucmayoc, 2000 m, vii 1941, C. Vargas Calderón 2075 (CUZ [photo MO]); Prov. Urubamba, Distr. Machu Pichu, Alcamayo, nacientes del río, 13°9′S, 72°30′W, 2900 m, 19 v 2003, I. Huamantupa, J. Farfán & G. Huallparimachi 3227 (CUZ, MO, US); Prov. Paucartambo, Hacienda Pillco, 2800 m, 12 iv 1967, C. Vargas C. 019239 (CUZ); Prov. Paucartambo, Distr. Kosnipata, Pillahuata, Parque Nacional Manu, 13°9′42′′S, 71°35′39′′W, 2700 m, 24 iii 1992, A.E. Cano 5296 (USM); Prov. Quispicanchi, Marcapata, En los alrededores de la mina cerca de la ciudad de Marcapata, en la carretera Cuzco-Maldonado, 13°25′25′′S, 70°54′15′′W, 1200 m, 20 vii 1987, P. Núñez & N. Núñez 8200 (MO); Prov. Quispicanchi, Distr. Marcapata, Limacpunko, Community of Union Arasa, Cullebrayoc Trail, 13°29′30′′S, 70°52′50′′W, 2674 m, 12 x 2011, J.D. Wells 941 (USM), J.D. Wells 963 (BRIT); Prov. Quispicanchi, 33 km past the peak (4750 m), on road from Ocongate to Marcapata, just before Opispata, 3110 m, 21 vii 1978, J. Aronson & P.E. Berry 525 (MO). Puno: Prov. Carabaya, Distr. Ollachea, road directly below Ollachea, 13°47′S, 70°28′W, 1949 m, 9 i 2015, M.C. Tebbitt & A. Daza 810 (MOL); Prov. Sandia, 10 vii 1903, 2100–2300 m, A. Weberbauer 503 (B, G); Prov. Sandia, 8 km N of Limbani, on trail, [14°4′40′′S, 69°41′17′′W], 3000 m, 19 v 1942, R.D. Metcalf 30543 (G, MO, US); Prov. Sandia, 2–6 km from Oconeque, 1800–2100 m, 22–25 v 1942, R.D. Metcalf 30607 (MO, US); Prov. Sandia, c.15 km on road below Sandia, 14°14′38′′S, 69°24′78′′W, 1600 m, 16 ii 2002, R.T. Pennington, T.D. Pennington & A. Daza 1112 (K, MOL); Prov. Sandia, entre Sandia y Cuyocuyo, 14°25′43′′S, 69°30′58′′W, 3100 m, 16 v 1966, R.A. Ferreyra 16771 (MO, USM [2]).
Bolivia. La Paz: Prov. Franz Tamayo, Madidi, Pelechuco-Apolo, Coranara Parque Nacional Madidi, sector Coranara, al lado del río Pelechuco, 14°46′21′′S, 68°59′9′′W, 2300 m, 15 vi 2008, A.F. Fuentes & H. Huaylla 13066 (MO); Prov. Bautista Saavedra, ANMI Apolobamba, Carpa Primeros restos de casas al lado de carretera bajando de Charazani, 15°11′24′′S, 68°53′8′′W, 2530 m, 17 iv 2005, A.F. Fuentes, R. Cuevas, E. Cuevas & H. Pariamo 6882 (MO); Prov. Sud Yungas, 16°19′37′′S, 67°49′55′′W, 2000 m, 24 iii 2007, T. Särkinen, C.E. Hughes, A. Wortley & P. Duchan 2058 (FHO).
In the original publication of Begonia dolabrifera (de Candolle, Reference De Candolle1908), there is a typographic error regarding the author name. It should read ‘C. DC’ instead of ‘D. DC’.
In the original publication of Begonia triramosa (Irmscher, Reference Irmscher1949), there is a typographic error regarding the collection number of the holotype. It should read ‘22493b‘ instead of ‘224936’.
Photographs of representative specimens of both living and dried plants of Begonia acerifolia are available via the Begonia Resource Centre (Hughes et al., (Reference Hughes, Moonlight, Jara, Tebbitt, Wilson and Pullan2015–).
Begonia velata L.B.Sm. & B.G.Schub., Publ. Field Mus. Nat. Hist., Bot Ser. 13(4/1): 201. 1941. – Type: Peru, Píura Region, Prov. Huancabamba, above Palambla, 2700 m, iv 1912, A. Weberbauer 6021 (holo GH!; iso F [2]!, NY!, US!).
Caulescent, rhizomatous herb. Rhizome horizontal, 5–20 cm long, 0.75–2 cm in diameter, unbranched. Stem erect to sprawling, usually 0.5–2 m tall, 0.75–2 cm in diameter, unbranched, internodes 1–19 cm long, glabrous. Stipules persistent, ovate to ovate-triangular, 1–1.5 × 0.6–0.9 cm, apex acute, margin entire. Leaves alternate, basifixed; petiole running straight into main vein of blade, 7.5–23 cm long, glabrous, bearing a dense ring of or a few scattered simple, translucent hairs around the top of the petiole at the position where it joins the blade; blade juicy, symmetrical, orbicular, 7.5–17 × 7.5–15 cm, base cordate, basal lobes spreading, sinus to 12 cm deep, apex acute, margin deeply 6-lobed, lobes ovate-triangular, 3.5–13 cm deep, serrulate, teeth sometimes tipped with a short hair, upper surface green, moderately pubescent, hairs pink, simple, lower surface silvery green, moderately pubescent, hairs white, veins palmate, 6–8. Inflorescence axillary from the upper portion of the stem, erect, a symmetrical cyme, 3-branched, branches 0.5–5 cm, bearing up to 12 flowers, bisexual, male flowers beginning to open before the first female flowers but both sexes eventually open concurrently; peduncle 15–40 cm long, glabrous; pedicels of male flowers 1.1–3.5 cm long, glabrous; pedicels of female flowers c.1 cm long, glabrous; bracts deciduous, ovate, 0.7–1.4 × 0.5–0.7 cm, apex acute, margin entire. Male flowers: tepals 4, spreading, white, sometimes reddish pink flushed on the outer surfaces or rarely on both surfaces and sometimes additionally with a green tinge on the outer surfaces of the outer two tepals, outer two elliptic to broadly elliptic, 1.4–1.6 × 1.2–1.4 cm, apex rounded, margin entire, glabrous, inner two obovate, c.1.5 × c.1.1 cm, apex rounded, margin entire, glabrous; stamens 45–75, attached along the length of a 1–2 mm long torus, filaments 1–2.5 mm long, anthers symmetrically basifixed, obovoid, c.1 mm long, dehiscing by unilateral slits, connectives not extended. Female flowers: bracteoles absent; tepals not persisting in fruit, 5, spreading, white, sometimes reddish pink flushed on the outer surfaces, obovate, subequal, 0.7–0.9 × 0.8–1 cm, glabrous, apex subacute, margin entire; ovary body broadly ellipsoid, 5.25–10 × c.6 mm, pale green to pink, glabrous, with one long wing and two rib-like wings, longest wing usually ligulate to occasionally triangular-ligulate, apex rounded to almost truncate, to 1.6 × 1 cm, 3-locular; placentae bifid, bearing ovules on both surfaces of placental branches; styles 3, shortly fused at base, c.6 mm long, bifid from about half their height, branches upright to flattened and somewhat spreading, stigmatic papillae in a once-spiralled band. Fruiting pedicel c.1 cm long. Fruit subnutant, body broadly ellipsoid, to 1.5 × 1 cm, glabrous, wings as in ovary.
Phenology
Flowering from April to June.
Distribution
Andes of Peru (Cajamarca and Píura).
Habitat
Typically growing as an epiphyte on trunks of trees and shrubs, occasionally terrestrial. Found at altitudes between 1500 and 3100 m.
IUCN conservation category
Begonia velata is known from two locations within a total area measuring 150 km2. Within these two locations, ongoing conversion of the local forest to agricultural land has caused a decline in the quality of available habitat. The current population size of the species is estimated to consist of no more than 10,000 individuals. Begonia velata is according assessed using IUCN criteria as Vulnerable (VU B2) (2016).
Additional specimens examined. Peru. Píura: Region, Prov. Morropon, Dist. Chalaco, Cima del cerro Mijal, Pajonal de jalca, 5°4′56′′S, 79°44′47′′W, 3100 m, 10 v 2003, I. Sánchez Vega, R. Cruz Córdova & E. Peña Cruz 11952 (CPUN); Prov. Huancabamba, El Tambo, [5°21′34′′S, 79°33′3′′W], 3000 m, 2 vi 1961, C. Acleto 228 (USM); Prov. Huancabamba, Distr. Canchaque, Canchaque – Minas Turmalina, [5°22′21′′S, 79°34′32′′W], 2200 m, 23 vii 1975, A. Sagástegui A., J. Cabanillas S. & O.C. Dios 8279 (HUT, MO); Huancabamba Province, above Canchaque on the Huancabamba Pass, 5°22′35′′S, 79°34′59′′W, 1758 m, 26 i 2016, P.W. Moonlight & A. Daza 107 (E); Prov. Huancabamba, Distr. Canchaque, above Palambla, 5°22′S, 79°35′W, 1710 m, 27 v 2015, M.C. Tebbitt & A. Daza 838 (E); Prov. Huancabamba, Distr. Canchaque, “Chorro blanco”, [5°23′S, 79°35′W], 1500–1900 m, 18 iv 1987, C. Díaz & S. Baldeón 2457 (USM); Prov. Huancabamba, Distr. Canchaque, Chorro Blanco, 5°23′S, 79°35′22′′W, 1500 m, 18 iv 1987, C. Díaz & S. Baldeon 2457 (MO). Cajamarca: Prov. Contumazá, Bosque Cachil, 6°23′18′′S, 79°17′28′′W, 2500 m, 16 vi 1994, A. Sagástegui, S. Leiva G. & P. Lezama 15307 (MO).
On the type sheets and in the original description (Smith & Schubert, Reference Smith, Schubert and Macbride1941) of Begonia velata, the species is described as having “fiery red” tepals. Fieldwork found that the species never has red tepals but instead has tepals that are white, although these are sometimes reddish pink flushed on the outer surfaces or rarely on both surfaces and sometimes additionally have a green tinge on the outer surfaces of the outer two tepals. Presumably this inconsistency resulted from a mix-up with collection notes. It is easy to see where this mistake could have originated, because at its type locality Begonia velata grows close to B. polypetala, which does have vivid scarlet tepals.
A photograph of the collection Tebbitt & Daza 838 in the living state is available via the Begonia Resource Centre (Hughes et al., Reference Hughes, Moonlight, Jara, Tebbitt, Wilson and Pullan2015–).
Acknowledgements
I thank the curators of AAU, B, BKL, BM, BR, BRIT, C, CAS, CGE, COL, CORD, CPUN, CUZ, E, E-GL, F, G, GB, G-BOISS, G-DC, G-DEL, GH, GOET, GUAY, HUT, K, LIL, LP, LPB, MEDEL, MO, MOL, MOL-WEB, NY, OXF, P, QCA, QCNE, QPLS, S, SEL, SI, TEX, UC, US, USM, USZ, W and Z for loaning material or allowing me to work in their herbaria. Fieldwork was funded by a series of grants from the American Begonia Society. I thank the Ecuadorian Ministerio del Ambiente and the Ministario del Ambiente del Perú for granting me permission to conduct the fieldwork and collect specimens. Álvaro J. Pérez (QCA), Carlos Augusto Reynel Rodriguez (MOL) and Tiina Särkinen (E) assisted with obtaining permits and gave advice regarding fieldwork, and Aniceto Daza Yomona (MOL), Adolfo Jara Muñoz (ANDES), Cristina Toapanta (QCA) and David Gutierrez (Ecuador) are gratefully acknowledged for their assistance in the field.
