Begonia rieckei Warb. [§ Petermannia

Ardi, W. H. & Thomas, D. C., 2022, SYNOPSIS OF BEGONIA (BEGONIACEAE) FROM THE NORTHERN ARM OF SULAWESI AND SANGIHE ISLAND, INDONESIA, INCLUDING THREE NEW SPECIES, Edinburgh Journal of Botany 79 (405), pp. 1-50 : 35-40

15. Begonia rieckei Warb. [§ Petermannia ], Bot. Jahrb. Syst. 13: 387 (1891).

– Type: Papua New Guinea, Sattelberg, F.R.R. Schlechter 20479 (holotype B [ B100217759 ]). Figure 12 .

Begonia axillipara Ridl., Trans. Linn. Soc. London, Bot. II (9): 60 (1916). – Type : Indonesia, New Guinea, Canoe Camp, 10 xi 1912, C.B. Kloss s.n (holotype BM [ BM000017329 ]), syn. nov.

Begonia brachybotrys Merr. & L.M.Perry, J. Arnold Arbor. 24: 56 (1943). – Type : Indonesia, New Guinea, Irian Jaya , Bernhard Camp, Idenburg river, iv 1939, L.J. Brass 14112 (holotype A [A00055445]; isotypes BM, L [ L0010281 ]), syn. nov.

Begonia brachyptera Merr. & L.M.Perry, J. Arnold Arbor. 29: 160 (1948). – Type: Papua New Guinea, Morobe Distr., Wantoat, 1940, J. & M.S. Clement 40896 (holotype A [A0005544], isotype MICH [MICH1115787]), syn. nov.

Begonia koordersii Warburg ex L.B.Sm. & Wassh., Phytologia 52(7): 444 (1983). – Type: Indonesia, Sulawesi, North Sulawesi, Minahassa , 1 iii 1895, S.H. Koorders 16246B (lectotype K [K000761117], designated by Smith & Wasshausen [1983: 444]; isolectotypes B [B100127763], BO), syn. nov.

Begonia peekelii Irmsch., Bot. Jahrb. Syst. 50: 360 (1913). – Type: Papua New Guinea, Bismarck Archipelago, New Ireland, New Mecklenburg , 21 ix 1910, G. Peekel 644 (holotype B [B100238530]), syn. nov.

Begonia pseudolateralis Warb. in Perkins, Fragm. Fl. Philipp. 1: 51 (1904). – Type: Philippines, Luzon, Isabela Prov., Malunu, O. Warburg 11793 (holotype B [B100238540]), syn. nov.

Mezierea salaziensis var. calleryana A.DC., Prodr. 15(1): 408 (1864). Begonia aptera var. calleryana (A.DC) Fern. - Vill., Fl. Filip. 4: 99 (1880). – Type: Philippines, Luzon, Laguna, Caluan, 1840, J.M.M. Callery s.n. (holotype P [P0190078]).

Begonia strictipetiolaris Irmsch., Bot. Jahrb. Syst. 50(4): 348 (1913). – Type: Indonesia, Sulawesi, North Sulawesi, Tomohon , 23 v 1894, K.F. & P.B. Sarasin 400a (holotype B [B100238028], isotype K [K000761115]), syn. nov.

Begonia lateralis Elmer ex Merr. nom. nud., Enum. Philipp. Fl. Pl. 3: 127 (1923).

Perennial, monoecious herb, erect, up to c. 1 m tall; stems glabrous except for microscopic glandular hairs. Stem branched, internodes 2–15 cm long, swollen at the nodes, green or red or brownish red with white spots. Leaves basifixed, alternate; stipules caducous, 2.5–3 × 1.2–1.8 cm, elliptic, boat-shaped, thick, midrib abaxially prominent, glabrous, apex acute, margin entire, reddish to reddish-greenish; petioles 8–23 cm long, adaxially shallowly channelled, concolorous with the stem, glabrous, except for the microscopic glandular hairs; lamina basifixed, 10–26.5 × 7–14 cm, asymmetrical, broadly ovate to broadly elliptic, margin entire to dentate to scalloped and denticulate between the larger teeth, base cordate and lobes sometimes overlapping, apex acute to acuminate, adaxial surface green or reddish-greenish, with white spots between the veins in juvenile stage, glabrous, smooth velvety or glossy, abaxially pale green or maroon red, sparsely hairy on the veins, veins reddish green; venation palmate-pinnate, primary veins 7–8, actinodromous, secondary veins craspedodromous. Inflorescences: bisexual; compressed cymose with up to 12 female flowers and 25 male flowers in a mixed inflorescence, peduncles c.1(–10) mm long, green-reddish, glabrous, bracts broadly elliptic to suborbicular, 2–2.5 × 1.8–2.4 cm, thick, pale green or reddish-greenish, glabrous. Male flowers: pedicels 11–15 mm long, white or white tinged with pink, glabrous; tepals 2, white or white tinged with pink, 5–12 × 2–12 mm, elliptic to suborbicular, apex rounded, outer surface glabrous; androecium yellow, symmetrical, on a c. 1 mm long column; stamens c.60, free filaments 0.5–1 mm long, fused at the base, anthers up to c. 0.5–1.5 mm long, narrowly obovate, dehiscing through unilaterally positioned slits that are c.1/2 as long as the anthers. Female flowers: pedicels c. 6 mm long, green-reddish, glabrous; tepals 5, white or white tinged with pink, subequal, 6–12 × 2–7 mm, ovate to elliptic, abaxially glabrous; ovary (excluding wings) 6–12 × 6–12 mm, ellipsoid, ovoid or globoid, glabrous, red or greenish or white, locules 3, placentation axile, placentae bilamellate, wingless or with 3 poorly developed wings, very narrow, green, widest point up to 1 mm (apically to subapically); style c. 3 mm long, basally fused, 3-branched, each stylodium bifurcate in the stigmatic region, stigmatic surface a spirally twisted papillose band, orange. Fruits: peduncles 5–15 mm long; pedicels 12–25 mm long; seed-bearing part ellipsoid, ovoid, or globoid, 15–20 × 10–15 mm (excluding the wings), glabrous, fleshy, indehiscent, wing shape as for ovary, up to 2 mm at the widest point (apically to subapically). Seeds barrel-shaped, c. 0.3 mm long.

Distribution. Widespread in east Malesia (Sulawesi, Philippines, Moluccas, New Guinea, Pacific Islands). In Sulawesi from North, Gorontalo and Central Provinces. See Figure 2 .

Habitat. Primary to secondary lowland to upland forest, in half to full shade, at 10–1500 m elevation.

Proposed IUCN conservation category. Least Concern (LC). Begonia rieckei has a wide distribution in central and eastern Malesia (EOO, 5,791,908 km 2) and occurs in various habitats including primary to strongly disturbed lowland to upland forests, and even oil palm plantations. Some of the populations are found in protected areas in Central Sulawesi Province, such as Pangi Binangga Forest Reserve and Lore Lindu National Park. Consequently, we assess this species as Least Concern ( IUCN Standards and Petitions Subcommittee, 2019).

Additional specimens examined. INDONESIA. Sulawesi. Northern arm of Sulawesi. Western North Sulawesi: Sojol foothill: 7 viii 2018, W.H. Ardi WI304 ( BO) ; 8 viii 2018, W.H. Ardi WI309 ( BO) . Central North Sulawesi: Gorontalo: Gunung Boliohutu , 22 iv 2002, M. Mendum et al. 141 (E) . Eastern North Sulawesi: Manado : J.E. Tiejsmann ( BO) ; anonym 41 (B); Goeroepahi , 18 iii 1917, Kauderns 18 ( L, S) ; Bolaang Mongondow , 15 iii 1985, E.F. de Vogel & J.J. Vermeulen 6556 ( L) ; Sulawesi Utara: Motomboto, 23 xii 1994, J.J. Afriastini et al. 2872 (L). Minahasa: Tomohon 11 vii 1894, K.F. & P.B. Sarasin 400 (B); 23 v 1894, K.F. & P.B. Sarasin 400a (BAS); Bojong, s.d., O. Warburg 15189 (B); Wiau Complex, 29 vi 1956, L.L. Forman 319 (K).

Central Sulawesi Province: Mount Lumut , on river bank, 24 iv 2008, D.C. Thomas & W.H. Ardi DCT08-62 ( BO, CEB, E) ; Kulawi road divide, Lore Lindu National Park , 29 vii 2018, W.H. Ardi et al. WI244 ( BO, SING) ; Mount Sojol, Balungkang II villane, Donggala , 7 viii 2018, W.H. Ardi et al. WI304 ( BO, SING) ; Sigi, Sidaoenta , 12 vii 1939, S. Bloembergen 4200 ( BO, L) ; Sungai Pehoeia, 25 iv 1975, W. Meijer 9603 ( L). Pangi Binangga Forest Reserve: Parigi , 20 vii 2018, W.H. Ardi et al. WI207 ( BO, SING, FIPIA) ; 20 vii 2018, W.H. Ardi et al. WI208 ( BO, FIPIA, SING). Luwuk : Batui River, 15 x 1989. M. Coode 5945 ( BO, L) ; Hanga-hanga waterfall, 8 iv 2008, D.C. Thomas & W.H. Ardi DCT08-23 (E) . Province unknown: A.B. Meyer s.n (B); J.G.F. Riedegel s.n (K).

Moluccas. Sula Islands , Mangoli, 26 ix 1939, S. Bloembergen 4689 ( BO, L) ; Moluccas, 1888, Jaheri 231 ( BO) ; Halmahera : Ternate Island, 10 i 1921, Anonymous 1295 ( BO) ; Baccan Islands : Gunung Sibela, 27 x 1974, E.F. de Vogel 3701 ( L). Buru : M.M.J.v Balgooy 5048 ( K, L) ; Fakal , 30 viii 1921, L.J. Toxopeus 460 ( BO) ; Wae Duna River , 25 xi 1984, M.M. J.v Balgooy 4911 ( L) ; 26 xi 1984, M.M.J.v Balgooy 4917 (L) ; 11 x 1984, Mogea & Ismail JPM 5237 ( BO) ; 22 xi 1984, H.P. Nooteboom 5222 ( L) ; 25 xi 1984, H.P. Nooteboom 5277 ( L) ; Wae Langa , 9 xi 1984, H.P. Nooteboom 5087 ( BO, L) ; 9 xi 1984, H.P. Nooteboom 5089 ( BO) ; Waekosi , 4 xi 1984, M.M.J.v Balgooy 4596 ( K, L). Seram: Roemoga, 4 ix 1938, P. Buwalda 5945 ( L) ; Mon Valley , 7 iii 1996, R.F. Ellen 1203 (K) ; Sikeu Walala , 20 xii 1996, M. Kato et al. 1114 ( L) ; Buria , 3 ii 1985, M. Kato et al. 6001 ( L) ; Lelesiru , 24 xii 1996, M. Kato 1167 ( L) ; 3 vii 1918, Kornasi 1507 ( BO) ; 4 vii 1918, Kornasi 1532 ( BO), 1507 ( BO) ; 20 xi 1917, Kornasi 471 ( BO), 472 ( BO) ; 23 xi 1917, Kornasi 515 ( BO) ; 18 i 1918, Kornasi 837 ( BO) ; 10 xii 1983, Ramlanto 357 ( BO) ; 1 x 1918, L.M.R. Rutten 1764 ( BO) ; Beka , 7 x 1918, L.M.R. Rutten 1772 ( BO) ; 10 iii 1919, L.M.R. Rutten 2106 ( BO) ; 10 x 1917, L.M.R. Rutten 242 ( BO) ; Manusela National Park , 15 ii 1985, M. Kato 6673 (A, L) ; Hatumete , 20 ii 1985, M. Kato et al. 6947 ( L) ; Maraina , 2 i 1985, Ueda K., Okamoto, M. & Mahjar U.W. 2834 ( L) .

New Guinea. West Papua: 9 v 1973, J. Raynal 17645 (P); vi–viii 1928, E. Mayr 546 (B); Yapen Island, 18 ii 2009, G.C.G. Argent et al. 147 (E); Aru Islands, Wokam Islands, Dosinamaloe, 29 v 1938, P. Buwalda 5069 (L); Upper Ramu, ix 1939, J. & M.S. Clemens 40750 (MICH); Sorong, 19 viii 1948, D.R. Pleyte 617 (L); Sentani, 9 iii 1973, J. Raynal 16685 (P); Cycloop Mts: Faika River, 8 viii 1961, P.v. Royen & H. Sleumer 6536 (L); Jappen-Biak: G. Wawah, 22 vii 1939, L.J.V. Dijk 44 (L); G. Wawah, 22 vii 1939, L.J.V. Dijk 215 (L) ; 16 ix 1939, L.J.V. Dijk 802 ( L) ; Arijom, 29 ix 1939, L.J.V. Dijk 923 ( L). Vogelkopf: Ije River Valley , 2 xi 1961, P.v. Royen & H. Sleumer 7650 ( L) ; Arfak Mts : Mupi Dessa, Trail from Mupi village to G. Humibou , near Sungai Mupi , between Kali Umera (stream) and K. Ngwes, 11 iv 1995, M.J.S. Sands 6741 (E) ; Arfak Mts : Minjambau, 22 v 1962, C. Versteegh 12685 ( L). Mamberamo: 3 iv 1914, A.C.T. Thompson 643 ( L) ; 28 viii 1953, C. Versteegh 47 ( L). Rouffaer River : viii 1926, Doctors v. Leeuwen-Reynvaan 9744 ( L) ; viii 1926, Doctors v. Leeuwen-Reynvaan 9905 ( L), 9743 ( L) ; Koode River , 16 viii 1938, E. Meijer Dress 653 ( L). Mt Jaya: 7 xii 1998, M.J.S. Sands 7210 ( L) ; 7 v 2000, T. Utteridge et al. 10615 ( L) ; 18 xi 2000, S. Atkins et al. 640 ( L) .

Papuasia. Bismarck Archipelago. New Britain: Hoskins Subdistr., Dakamanu , 21 xii 1967, M. Coode 32645 ( L) ; Kandrian Subdistr., Alimbit River , 8 x 1965, A. Gillison 22466 ( L) ; Ulamona Mission , 6 vi 1973, R.S. Isles & A.N. Vinas 32353 ( L) ; 20 ii 1989, P. Katik 64088 ( L) ; Pomio Subdistr., Mt Lululua , 4 v 1973, P.F. Stevens & Y. Leiean 58247 (E, L) ; Kokopo Subdistr., Valilie River, 14 viii 1969, H. Streimann 44374 ( L). Manus: Mt Dremsel , 26 x 1974, D. Foreman & P. Katik 59195 (E, L) ; Buyang , 11 iii 1981, K. Kerenga & J. Croft 77357 ( L) ; Rambutyo Island, 16 iii 1981, K. Kerenga & J. Croft 77376 ( L). New Ireland: Kavieng Subdistr., Schieinitz Range , Logagon Village , 24 x 1974, J. Croft & Y. Leiean 65630 ( L) ; Konos Province , NE of Lelet Farm, 26 x 1984, O. Gideon 77137 ( L) ; Logging concession N of Danfu river, 25 v 2013, A.D. Poulsen et al. 10001 (E) .

PAPUA NEW GUINEA. Baridi , 18 xi 1935, C.E. Carr 14995 ( BM, L), 14975 (B) ; Bewepi Creek , 3 ix 1962, Henry 14820 (E, L) ; Mt Bosavi , 27 viii 1986, O. Gideon 57420 ( L) ; Aitape Subdistr. Pes Village , 8 vi 1961, P.J. Darbyshire & R.D. Hoogland 7896 ( L) ; Nassau Mts , x 1926, Doctors v. Leeuwen-Reynvaan 10545 ( L) ; Yodda Valley , 19 ix 1953, R.D. Hoogland 3941 ( L) ; Hozeke , 29 xi 1984, K. Kerenga & C. Baker 56917 ( L) ; Kokoda , 25 vii 1964, A.N. Millar 23496 ( L) ; Western Highland Province: Hagen Subdistrict: Trauna Creek , 7 vii 1968, A.N. Millar 37627 ( L) ; Kar Kar Island, 16 vii 1968, A.N. Millar ( L). Sepik District: Maprik Subdistr: Prince Alexander Range: Mt Turu , 24 viii 1959, R. Pullen 1582 ( L) ; Madang Prov.: Saidor Subdistr., Wumundi , 11 viii 1984, C.D. Sayers 19778 ( L) ; Chimbu Province: Crater Mt Wildlife Management area , 12 ix 1996, W.N. Takeuchi 11125 (A, E, L) ; Sangwep Logging Area , 13 iii 1975 , J.F. Veldkamp 6168 ( L); Baiyer River , 28 vi 1981, A.N. Vinas 3286 ( L) ; East Sepik Province: Maprik subprovince, 24 vii 1980, J. Wiakabu 73419 ( L) ; 18 xi 1935, C.E. Carr 14975 (B); Isuarava , 15 ii 1936, C.E. Carr 15598 (B, L) ; Kokoda, 15 ii 1936, C.E. Carr 16124 ( B, L). Morobe District: Sattelberg , 28 iii 1936, J. & M.S. Clemens 2244 (B) ; Yunzaing , 11 vi 1936, J. & M.S. Clemens 3281 (B) ; Sattelberg , J. & M.S. Clemens 379a (B) ; Wantoat , 6 iv 1940, J. & M.S. Clemens 41205 ( MICH) ; Sattelberg , 31 x 1935, J . & M.S. Clemens 655 (B); Wafi River , 4 iii 1985, B.J. Conn 1800 ( L) ; Mt Kaindi , 29 iv 1992, R. Hoft 2186 ( L) ; Bulolo : Crooked Logging Area, 5 ix 1966, H. Streinmann & A. Kairo 27900 ( L) ; Bulolo: Dengalu Village , Floyd 5245 ( L) ; Kipu , 13 xi 1968, A. Gillison & A. Kairo 25692 ( L) ; Sopa , 19 vii 1962, T.G. Hartley 10354 ( L) ; Bumbu Loggin Area , ix 1961, Henry 14321 ( L) ; Lae Subprovince : Musom Village, 19 vi 1978, P. Katik 70827 ( L) ; Wantoat , 9 viii 1968, A.N. Millar 12139 ( L) ; Garaina , 10 v 1971, B.C. Stone 10169 ( L) ; Atzera Range , 10 vii 2001, W.N. Takeuchi et al. 15424 ( L) ; Bubia , xii 2001, W.N. Takeuchi 15806 ( L) ; Bubia , 17 vii 2001, W.N. Takeuchi 16398 ( L) ; Sankwep logging area, 23 iv 1968, J.S. Wormsley 37119 ( L). Northeast New Guinea. West Sepik Province: Bewani, 31 viii 1982, K. Kerenga 55469 ( L) ; Wutung Patrol Post , 8 ix 1982, K. Kerenga 56405 ( L) ; Vanimo , 30 xi 1971, H. Streimann 52967 ( L) ; Bewani : Mt Yungat, 20 ix 1982, J. Wiakabu 50578 ( L). Kaiser-Wilhelmsland: 10 iv 1889, F.C. Hellwig 601 (B) ; Simmpliqe Stellen au Mimjeiu lei Kelel, 24 vi 1907, F.R.R. Schlechter 16175 (E). Josephstaal: 22 vii 1999, W.N. Takeuchi 13422 (A, L) ; 28 vii 1999, W.N. Takeuchi 13420 (A, L) .

PHILIPPINES. Catanduanes Island: Catanduanes , 7 xi 1996, Reynuso & R.S. Majaducon 24877 ( L) ; 9 xi 1996, Reynuso & R.S. Majaducon 24958 ( L). Cebu Island : Cantipla , 26 iii 1971, Anon. 10 ( L) ; i 1994, D. Bicknel 804 ( K). Leyte Island : 30 viii 1913, C.A. Wenzel 517 ( BM). Luzon Island : 1861, A.F. Jagor 814 ( B) ; Aurora, Brgy. Bianoan , 19 iii 1993, Barbon et al. 9299 ( PNH) ; Cagayan Province: Claveria , 3 viii 1995, Garcia et al. ( L) ; Leyte, 10 iii 1914, C.A. Wenzel 634 ( BM) ; Mt Dimaxinggay , 13 iii 1993, Barbon et al. 9202 ( L, PNH) ; Ilocos Norte Province: Bangui , viii 1918, M. Ramos 33037 ( L) ; Apayao, v 1917, E. Fenix 28141 ( BO, K, P) ; Ifugao: Lake Ambuaya , 29 iii 1991, Reynoso et al. 7283 ( K) ; Camarines: Mt Isaro (g), xi–xii 1913, M. Ramos 22017 ( BM, L) ; Cavite Province: Mt Palay Palay Nat. Park. , 22 iii 1995, Reynoso et al. 14957 ( L) ; Quezon: Polillo Island , C.B. Robinson 6903 ( BO, L, P) ; Labra Prov.: Poblacion Gangal , 14 xi 1996, A.F. Fuentes 38594 ( L) ; San Jose Village: San Mariano: Bo. Disulap , 26 iv 1961, H. Gutierrez 78078 ( L) ; Laguna, vi–viii 1915, R.C. McGregor 22801 ( P) ; Bataan: Mt Mariveles: Lamao River , 13 i 1904, R.S. Williams 523 ( K) ; Tayabas : iii 1888, O. Warburg 13086 ( B) ; Atimonan , iii 1905, E.D. Merrill 4409 ( B, K, P) ; Casiguran , v–vi 1925, M. Ramos & G. Edano 45733 ( BM) ; Mauban , i 1913, M. Ramos 19481 ( BM, L) ; Albay Province, Mt Malinao , 29 i 1956, G. Edano 34447 ( L) ; Mayon Volcano , 2 vi 1953, D.R. Mendoza 18312 ( L) ; Taqui River , 27 x 1995, Reynoso et al. 21272 ( L) ; Mt Malinao , 29 x 1995, Reynoso et al. 21354 ( L) ; Mt Mayon, 17 ix 1991, Reynoso et al. 3506 ( K, L). Isabela Province: San Jose Village , 6 iii 1997, Reynoso et al. 2008 ( L) ; Malunu , O. Warburg 11792 ( B) ; 6 iii 1997, P. Wilkie et al. 29008 ( E, K). Sorsogon: vii–viii 1915, M. Ramos 23437 ( B, P) ; Irosin , v 1957, G. Edano & H. Guiterrez 38546 ( BM, K, L) ; Mt Bulusan, vii 1915, A.D.E. Elmer 16661 ( BM, BO, K, L). Mindanao Island: Agusan : Mt Urdaneta , viii 1912, A.D.E. Elmer 13494 ( BM, BO, E, K, L, U) ; Butuan: Ojot River , 21 vi 1961, D.R. Mendoza 42468 ( L) ; Zamboanga: Mt Tubuan , x 1919, M. Ramos & G. Edano 36654 ( BM, BO) ; Basilan, viii–ix 1912, J. Reillo 16142 ( BM, L, P). Mindoro Island: Sibuang River , 12 ii 1985, C.E. Ridsdale 831 ( L) ; Mt Halcon , i–ii 1948, G. Edano 2492 ( BO) ; i–ii 1948, G. Edano 3492 ( BO) ; iii 1922, M. Ramos & G. Edano 40679 ( K) ; Lantuyan , 29 iii 1991, Anon. 440 ( K, L) ; Baco River , E.D. Merrill 991 ( B, K, P). Negros: Negros Occidental: Mt Katugasan , iii 1954, G. Edano 21819 ( K, L). Panay Island : Capiz: Libacao, v–vi 1919, A. Martelino 35397 ( B, BM, K, P) ; Aklan Province: Libacao : Poblacion, C.I. Peng & R.R. Rubite 23801 ( HAST). Samar Island: Laquilacon , vi 1924, McGregor 437559 ( BM) .

Begonia rieckei is a common species in eastern Malesia and is morphologically highly variable. Several species have been described in the Begonia rieckei complex, but the distinguishing characters are often minor and therefore we advocate a wider concept (see Hughes, 2008).

Previously, some of the taxa in the Begonia rieckei complex, including B. axillipara , B. brachybotrys and B. pseudolateralis , were incorrectly classified under Begonia sect. Sphenanthera (the section is now synonymised under Begonia sect. Platycentrum , see Moonlight et al., 2018), primarily based on the presence of fleshy fruits. Tebbitt (2000) revised and transferred them into Begonia sect. Petermannia because they have some typical characters of the section, such as female flowers with five tepals, male flowers with two tepals, and anthers dehiscing through unilaterally positioned slits, and the anther connectives are not extended as is typical for species in Begonia sect. Platycentrum . This placement was confirmed by chloroplast DNA data, which indicated that the Begonia rieckei complex is nested within a clade of Sulawesi endemics ( Thomas et al., 2012).

Tebbitt (2000) and Tebbitt & Dickson (2000) kept some taxa in the species complex separate, including Begonia brachybotrys (New Guinea) and B. pseudolateralis (the Philippines), based on differences in the leaf margin and venation characters. However, all the species have very characteristic generative structures in common, including compressed, cymose, bisexual inflorescences with up to 12 female flowers and 25 male flowers, and fleshy indehiscent fruit that are wingless or have poorly developed wings.

Figures

Figure 2. Distribution maps of Begonia species occurring on the northern arm of Sulawesi. Distribution points are from georeferenced collections from major herbarium collections (B, BO, E, K, KRB, L, SING), the Begonia Resource Centre (Hughes et al., 2015–) and the Sulawesi Begonia Data Portal (Thomas et al., 2013, continuously updated).

Figure 12. Begonia rieckei. A, Habit, reddish leaves with white dots along the margin and between the veins; B, inflorescence; C, male flowers; D, female flower; E and F, infructescence; G, ovary (crosssection of middle part). A from W.H. Ardi WI410; B, D and F from W.H. Ardi WI207; C, E and G from W.H. Ardi WI391. Photographs: W. H. Ardi.