A foundation monograph of Ipomoea (Convolvulaceae) in the New World Author Wood, John R. I. Author Munoz-Rodriguez, Pablo Author Williams, Bethany R. M. Author Scotland, Robert W. text PhytoKeys 2020 143 1 823 http://dx.doi.org/10.3897/phytokeys.143.32821 journal article http://dx.doi.org/10.3897/phytokeys.143.32821 1314-2003-143-1 F6F11A6EE4FF5A1885CEA2B60AE965A5 220. Ipomoea batatas (L.) Lam. , Tabl. Encycl. 1 : 465. 1793. (Lamarck 1793: 465) Convolvulus batatas L. , Sp. Pl. 1 : 154. 1753. (Linnaeus 1753: 154). Type. INDIA. Herb. Linn. No.77.5 (S, lectotype designated by Biju, 2003 755). Convolvulus esculentus Salisb. , Prodr. Stirp. Chap. Allerton 123 . 1796. (Salisbury 1796: 123), nom. illeg. superfl. Type. Based on Convolvulus batatas L. Convolvulus edulis Thunb. ex Murray, Syst. Veg. , ed. 14: 203. 1784. (Murray 1784: 203). Type. JAPAN. Thunberg (holotype UPS). Batatas edulis (Thunb. ex Murray) Choisy , Mem. Soc. Phys. Geneve 6 : 435 [53]. 1834. (Choisy 1834: 435 [53]). Ipomoea edulis (Thunb. ex Murray) Niederl. , Bol. Mens. Mus. Prod. Argent. 3 (29): 190. 1890. (Niederlein 1890: 190). Ipomoea batatas var. edulis (Thunb. ex Murray) Makino , Fl. Japan 476 . 1925. (Makino 1925: 476). Convolvulus platanifolius Vahl , Symb. Bot. 3 : 26. 1794. (Vahl 1794: 26). Type. Illustration in L. Plukenet (1692: t.167, f. 3), lectotype designated here. Ipomoea platanifolia (Vahl) Roem. & Schult. , Syst. Veg. 4 : 220. 1819. (Roemer and Schultes 1819: 220). Ipomoea fastigiata var. platanifolia (Vahl) Griseb., Fl. Brit. W. I. 468. 1864 [pub. 1862]. (Grisebach 1862b: 468). Ipomoea villosa Ruiz & Pav. , Fl. Peruv. 2 : 12, t. 121. 1799. (Ruiz and Pavon 1799: 12). Type. ECUADOR. Guayaquil, Ruiz, Pavon & Dombey (lectotype MA814679, designated here; isolectotypes BM, OXF). Ipomoea catesbaei G. Mey. , Prim. Fl. Esseq. 103 . 1818. (Meyer 1818: 103). Type. Based on Catesby 2: 60, t. 60, lectotype designated here. Convolvulus fastigiatus Roxb., Fl. Indica , ed. 2, 2: 48. 1824. (Roxburgh 1824: 48) Type. INDIA. Bengal, (lectotype, icon Roxburgh 1355 (K), designated here). Ipomoea fastigiata (Roxb.) Sweet, Hort. Brit. , ed. 1: 188. 1826. (Sweet 1826: 188). Ipomoea batatas var. fastigiata (Sweet) Kuntze , Rev. Gen. Pl. 2 : 442. 1891. (Kuntze 1891: 442). Convolvulus edulis Vell. Fl. Flumin. 72. 1825 [pub. 1829]. (Vellozo 1829: 72), nom. illeg., superfl. based on Convolvulus batatas L. Convolvulus tuberosus Vell. Fl. Flumin. 72. 1825 [pub. 1829]. (Vellozo 1829: 72), nom. illeg., non Convolvulus tuberosus Spreng. (1824) . Type. BRAZIL. Not specified, (lectotype, original parchment plate of Flora Fluminensis in the manuscript section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651-057], designated here; later published in Vellozo , Fl. Flum. Icon. 2 : t. 57. 1827 [pub. 1831]). Convolvulus esculentus Vell. , Fl. Flumin. 73 . 1825 [pub. 1829]. (Vellozo 1829: 73), nom. illeg., non Convolvulus esculentus Salisb. (1796) . Type. BRAZIL. Not specified, (lectotype, original parchment plate of Flora Fluminensis in the manuscript section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651-058], designated here; later published in Vellozo , Fl. Flum. Icon. 2 : t. 58 1827. [pub. 1831]). Convolvulus batata Vell. , Fl. Flumin. 73 . 1825 [pub. 1829]. (Vellozo 1829: 73). Type. BRAZIL. Not specified, (lectotype, original parchment plate of Flora Fluminensis in the manuscript section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651-059], designated here; later published in Vellozo , Fl. Flum. Icon. 2 : t. 59 1827. [pub. 1831]). Convolvulus cordatifolius Vell. , Fl. Flumin. 73 . 1825 [pub. 1829]. (Vellozo 1829: 73). Type. BRAZIL. Not specified, (lectotype, original parchment plate of Flora Fluminensis in the manuscript section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651-060], designated here; later published in Vellozo , Fl. Flum. Icon. 2 : t. 60 1827. [pub. 1831]). Convolvulus varius Vell. , Fl. Flumin. 73 . 1825 [pub. 1829]. (Vellozo 1829: 73). Type. BRAZIL. Not specified, (lectotype, original parchment plate of Flora Fluminensis in the manuscript section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651-061], designated here; later published in Vellozo , Fl. Flum. Icon. 2 : t. 61 1827. [pub. 1831]). Convolvulus variabilis Schltdl. & Cham. , Linnaea 5 : 116. 1830. (Schlechtendal and Chamisso 1830: 116). Type. MEXICO. Veracruz, Hacienda de la Laguna, Schiede & Deppe s.n. (holotype HAL0037741, isotype LE, n.v.). Ipomoea variabilis (Schltdl. & Cham.) Choisy in A.P. de Can. dolle , Prodr. 9 : 383. 1845. (Choisy 1845: 383). Ipomoea indica var. variabilis (Schltdl. & Cham.) L.O. Williams, Fieldiana , Bot. 32 (12): 191. 1970. (Williams 1970a: 191). Batatas xanthorhiza Bojer , Hort. Maurit. 225 . 1837. (Bojer 1837: 225). Type. MAURITIUS. "Cult. Danes les habitations". No specimen cited. Batatas edulis var. xanthorhiza (Bojer) Choisy in A.P. de Candolle , Prodr. 9 : 338. 1845. (Choisy 1845: 338). Batatas betacea Lindl., Bot. Reg. (Edwards) 25: 93. 1839. (Lindley 1839c: 93). Type. No specimen preserved, lectotype t. 56 in Bot. Reg. (Edwards) 26 (1839), designated here. Ipomoea apiculata M. Martens & Galeotti , Bull. Acad. Roy. Sci. Bruxelles 12 (2): 262. 1845. (Martens and Galeotti 1845: 262). Type. MEXICO. Veracruz, H. Galeotti 1381 (lectotype BR00006972851, designated here; isolectotypes BR, P). Ipomoea batatas var. apiculata (M. Martens & Galeotti) J.A. Mcdonald & D.F. Austin , Brittonia 42 (2): 118. 1990. (McDonald and Austin 1990: 118). Convolvulus attenuatus M. Martens & Galeotti , Bull. Acad. Roy. Sci. Bruxelles 22 : 265. 1845. (Martens and Galeotti 1845: 265). Type. MEXICO. Oaxaca, H.G. Galeotti 1399 (syntypes BR, P, G, MEXU). Batatas wallii Morren , Ann. Soc. Roy. Agric. Gand. 2 : 285-286, t. 74. 1846. (Morren 1846: 285). Type. GUATEMALA. Pere Wall de Poperingue s.n. (whereabouts uncertain). Ipomoea wallii (Morren) Hemsl., Biol. Cent.-Amer. , Bot. 2 (11): 396. 1882 (Hemsley 1882: 396). Ipomoea batatas var. leucorrhiza Griseb., Fl. Brit. W. Ind. 468. 1864 [pub. 1862]. (Grisebach 1862b: 468). Type. ANTIGUA. Wullschlagel s.n. (whereabouts unknown). Ipomoea batatas var. porphyrorhiza Griseb., Fl. Brit. W. Ind. 468. 1864 [pub. 1862]. (Grisebach 1862b: 468). Type. JAMAICA. collector and whereabouts unspecified. Batatas edulis var. porphyrorhiza (Griseb.) Ram. Goyena , Fl. Nicarag. 2 : 649. 1911. ( Ramirez Goyena 1911: 649). Ipomoea batatas var. dissoluta Kuntze , Rev. Gen. Pl. 2 : 442. 1891. (Kuntze 1891: 442). Type. Not specified. Ipomoea batatas var. subscandens Kuntze , Rev. Gen. Pl. 2 : 442. 1891. (Kuntze 1891: 442). Type. INDIA. Deccan, not specified. Ipomoea fastigiata var. ciliata Huber , Bol. Mus. Paraense Hist. Nat. Ethnogr. 2 : 512. 1898. (Huber 1898: 512). Type. BRAZIL. Para, Rio Anauera-pucu , M. Guedes 582 (holotype MG). Ipomoea vulsa House , Muhlenbergia 3 (3): 45. 1907. (House 1907b: 45). Type. MEXICO. Veracruz, Orizaba, F. Mueller s.n. (holotype US00111491, isotypes NY, US). Ipomoea purpusii House , Ann. New York Acad. Sci. 18 : 248. 1908. (House 1908b: 248). Type. MEXICO. Veracruz, near Zacuapan, C.A. Purpus 2113 (holotype NY00319135, isotypes F, US). Ipomoea batatas var. lobata Gagnep. & Courchet , Fl. Indochine 4 : 241.1915. (Gagnepain and Courchet 1915: 241). Type. VIETNAM. Tonkin, Ninh-binh, Bon s.n. & Long-Tcheou, Beauvais (syntypes P?, n.v.). Ipomoea confertiflora Standl. , Publ. Carnegie Inst. Wash. 461 : 83. 1935. (Standley 1935: 83). Type. BELIZE. Rio Grande, W.A. Schipp 1236 (holotype F0054833, isotypes A, BM, GH, K, MICH, MO, NY, S). Ipomoea davidsoniae Standl., Publ. Field Nus. Nat. Hist. , Bot. Ser. 22 : 98. 1940. (Standley 1940c: 98). Type. PANAMA. Chiriqui, Bajo Mono, M.E. Davidson 595 (holotype F0O54838, isotype MO). Ipomoea mucronata Schery , Ann. Missouri Bot. Gard. 28 : 463. 1941. (Woodson and Schery 1941: 463). Type. PANAMA. Chiriqui, near Pena Blanca, R.E. Woodson & R.W. Schery 323 (holotype MO00340730). Ipomoea batatas forma trifida Moldenke , Phytologia 2 : 224. 1947. (Moldenke 1947: 224). Type. ECUADOR. Loja, La Toma, R. Espinosa 492 (holotype NY00319162). Ipomoea tiliacea var. merremioides Fosberg , Smithsonian Contrib. Bot. 21 : 15. 1975. (Fosberg and Sachet 1975: 14). Type. FRENCH POLYNESIA. Hiva Oa Island, M.H. Sachet 1300 (holotype US00111475, isotype P). Ipomoea tiliacea var. smithii Fosberg , Smithsonian Contrib. Bot. 21 : 15. 1975. (Fosberg and Sachet 1975: 15). Type. FIJI. Viti Levu, A.C. Smith 4468 (holotype US00111476, isotype BISH). Ipomoea tabascana J.A. McDonald & D.F. Austin , Brittonia 42 : 116. 1990. (McDonald and Austin 1990: 116). Type. MEXICO. Tabasco, S. limit of Ejido Lopez Zamora, D.F.Austin & F. de la Puente 7505 (holotype not at US, isotypes CIP [Lima], FTG, XAL, n.v.). Type. Based on Convolvulus batatas L. Description. Creeping (rarely climbing) perennial herb rooting from the stem and developing storage roots; stems extending to cover several metres, glabrous to coarsely pilose, often stiut in cultivated and feral forms. Leaves petiolate, very variable in form but usually rather large, 3-15 x 5-12 cm, ovate or shallowly to deeply 3-5-lobed, cordate, shortly acuminate, both surfaces glabrous to coarsely pilose, abaxially somewhat glaucous and with prominent veins; petioles usually rather long, 4-15 cm. Inflorescence of long-pedunculate, axillary, dense umbellate cymes; peduncles 5-30 cm long, stout; bracteoles filiform, c. 2 mm long, caducous; secondary peduncles 5-15 mm; pedicels very short, 5-10 mm long; sepals 7-11 mm, unequal, margins often but not always ciliate, outer slightly shorter than inner, oblong-elliptic to oblong-oblanceolate, abruptly mucronate with a hair point c. 2 mm long, prominently 1-5-veined, the inner sepals broadly elliptic, rounded and mucronate; corolla 4-4.5 cm long, pink, often with a dark centre, glabrous; ovary pubescent, rarely fertile so capsules and seeds usually absent. Illustration. Figure 110E, F ; Acevedo-Rodriguez (2005: 165); Bosser and Heine (2000: 35); Deroin (2001: 173, 247) (as Ipomoea trifida ). Distribution. The sweet potato is of American origin but is now cultivated throughout tropical and subtropical regions of the world with greatest production reported from China. We have seen examples of cultivated plants from all parts of the Americas including Easter Island [ F. Fuentes 3 (K), 4 (K)] and Hawaii [ J. Stokes s.n. 1/1912 (K); Oahu, Christopherson et al. 1594 (K)] with the exception of the extreme south and Canada. Outside cultivation, plants are usually found in derelict fields and on roadsides near settlements Most cultivated plants are sterile but we have seen occasional specimens of apparently wild, fertile plants from various countries in tropical America including Colombia, Ecuador, Mexico, Panama and Venezuela. No apparently naturally occurring populations are reported from the Caribbean islands, Brazil or the Guianas. Obviously cultivated plants are not cited below but many of the records are of escapes from cultivation although some may be of wild populations. FRENCH GUIANA. Berthoud-Coulon 505 (BM). SURINAM. M. Berthoud-Coulon 507 (BM) BOLIVIA. (escapes from cultivation). La Paz : Murillo, Valle de Zongo, Cahua, 1300 m, 14 June 1980, S.G. Beck 3688 (CTES, CUSCO, FTG, LPB, MO, USZ). Santa Cruz : Nuflo de Chavez , Concepcion J.R.I. Wood & D. Soto 27939 (OXF, K, LPB, USZ); J.R.I. Wood et al. 28090 (LPB, OXF, USZ). PERU. Huanuco : J. Schunke 2013 (G). Lambayeque : T. Torres s.n. (USM). Loreto : Chanintia , A. Montalvo s.n. (USM); Boqueron , R. Ferreyra 1185 (USM); Iquitos, H. Murphy 301 (MO, OXF). Madre de Dios : Tambopata, Puerto Maldonado, I. Huamantupa & A. Montero 3671 (MO, OXF). Pasco : Oxapampa, Huancabamba, camino a Pozuzo, R. Rojas et al. 2513 (MO, OXF). Piura : E. Laure 5326 (P), 5370 (P). ECUADOR. Cotapaxi : B. Sparre 17329 (S). El Oro : Arenillas, E. Asplund 15676 (K, S). Guayas : San Ignacio, I. Holmgren 88 (S); Guayaquil, L. Fraser (BM). Los Rios : B. Sparre 17916 (S); C. Jativa & C. Epling 182 (S). Manabi : J. Brandbyge 42773 (AAU, ARIZ); Eggers 15105 (P). Pinchincha : B. Sparrre 14820 (S). COLOMBIA. Antioquia : Angelopolis , G. Gutierrez & F. Barklay 17C654 (BM). Boyaca : A.E. Lawrance 544 (BM). Cauca : La Paila, I.F. Holton s.n. [1853] (K). Cundinamarca : La Mesa, J. Triana 3807 (BM). Magdalena : Santa Marta, H.H. Smith 1912 (E). Meta : Villavicencio, J. Triana 3803 (BM). Putumayo : J. Ewan 16705 (BM). Valle : A. Gentry et al. 59527 (FTG). VENEZUELA. Dist. Fed. : Caracas-Guayra, A.H.G. Alston 5500 (BM). Zulia : A. Fernandez 20591 (MA). PANAMA. B.L. Seeman 488 (BM), 1604 (BM), 6453 (BM, MO); E.L. Tyson 6994 (BM, PMA); C. Whitefoord & A. Eddy 71 (BM); C. Hamilton et al. 1300 (FTG, MO); A. Ibanez et al. 1804 (MA). COSTA RICA. A.F. Skutch 2570 (K), 3672 (K); H. Pittier 13675 (K); Santa Elena-San Rafael, P. Wilkin 436 (BM); Puntarenas, Cordillera de Talamanca, F. Quesada et al. 1147 (BM); Limon, B. Hammel et al. 19673 (BM). NICARAGUA. M. Araquistain & J.C. Sandino 1384 (FTG); Zelaya, W.D. Stevens et al. 6453 (BM, MO). EL SALVADOR. G. Davidse et al . 37459 (MO). HONDURAS. Gracias a Dios, P. House 37 (BM); J. Saunders 709 (FTG). BELIZE. Georgeville-Augustine, G.R. Proctor 29630 (BM); D.R. Hunt 150 (BM). GUATEMALA. Alta Verapaz, H. von Tuerckheim 1437 (K); Bernoulli & Cario 1906 (K). MEXICO. Campeche : E. & H. Cabrera 13444 (BM, MEXU, MO). Chiapas : A. Reyes-Garcia & E. Martinez 132 (BM, MEXU); J.C. Soto et al. 13219 (BM, MEXU). Est. Mexico & Dist. Fed. : Temascaltepec: G.B. Hinton 2009 (K). Guerrero : G.B. Hinton 8501 (K), 9510 (K); Mina , 9699 (K). Oaxaca : D.F. Austin & F. de la Puente 7672 (FTG). Quintana Roo : Isla de Cozumel, E. & H. Cabrera 10541 (BM, MEXU). Veracruz : E. Kerber 37 (BM); M. Botteri 560 (BM, K, OXF); J. Linden 257 (K); H. Galeotti 1351 (K); Bandaril, Jalapa, E.K. Balls & W.B. Gourlay 5483 (E, BM); Gouin s.n. [1867] (P). Typification. Although often claimed to be an illegitimate name, Ipomoea fastigiata (Roxb.) Sweet appears to have been validly published. Sweet refers to Flora Indica, not Hortus Benghalensis but incorrectly gives the date as 1816, which is, in fact, incorrect for both these publications. Notes. Ipomoea batatas appears to have arisen naturally in pre-human times in Tropical America and is most closely related to I. trifida . Its origins are discussed by Munoz-Rodriguez et al. (2018). It is widely cultivated throughout the tropics and the orange-fleshed variety is of particular importance as it is rich in a precursor of Vitamin A. Ipomoea batatas is usually readily identified in the field because of its root tubers and perennial creeping habit, the stems rooting at the nodes. Herbarium specimens are distinguished by the strongly and usually abruptly mucronate sepals with a distinct mucro and a pronounced central vein with 2-4 less prominent lateral veins. The sepals are usually ciliate and the flowers characteristically clustered in a subumbelliform structure at the apex of a long peduncle. The leaves are commonly 3-lobed. Variation. Various apparently wild forms of Ipomoea batatas are relatively distinct morphologically and have been recognized over the years. The plant treated as Ipomoea batatas var. apiculata ( I. apiculata , I. vulsa ) represents a form from coastal sand dunes near Veracruz but is also found in Campeche and Oaxaca. It is a slender plant, rooting at the nodes or twining, with deeply 3-5(-7)-lobed glabrous leaves, cymes of 1-3 flowers, very unequal sepals (the outer oblong, mucronate much narrower and shorter than the inner obovate sepals) and a distinctly campanulate corolla c. 3 cm long but with a tube c. 1.5 cm wide. We have seen the following additional specimens: MEXICO. Campeche. E. & H. Cabrera 12504 (IEB). Oaxaca : Ghiesbrecht s.n. (P03548796) Veracruz : D.F. Austin & F. de la Puente 7480 (FTG) , Vislet 1856 (P), G. Castillo-Campos et al. 1438 (IEB), E. Matuda 17095 (MEXU). The plants described as Ipomoea tabascana are very slender glabrous plants, rooting at the nodes and with strap-shaped, strongly sagittate leaves and few-flowered cymes. They are only known from marshy ground near the type locality in Tabasco (Austin et al. 1991). Molecular studies show I. tabascana to be nested within I. batatas , probably representing a hybrid between I. batatas and I. trifida . (Srisuwan et al. 2006; Munoz-Rodriguez et al. 2018). Ipomoea confertiflora also appears somewhat distinct and has sometimes been treated as belonging to I. trifida (Austin and Huaman 1996) because of the somewhat scarious sepals. It has oblong, somewhat twisted outer sepals. As well as the type, M. Araquistain & J.C. Sandino (FTG) from Nicaragua, N. Garwood et al . 819 (BM) from Costa Rica, I.F. Holton 540 (K) from Cauca, Colombia and C. H. Dodson 3811 (F) from Ecuador fit this form. Two distinct forms come from Ecuador. One of these is represented by the type of Ipomoea villosa Ruiz & Pav. from Guayaquil, which has been treated as I. leucantha . This has trilobed leaves and long, lanceolate, acuminate sepals 13-14 mm in length. Very similar is Asplund 15966 (S) from Manta, Manabi Province and U. Chavarria 1343 (BM, MO) from Costa Rica. Somewhat similar plants with entire leaves and slightly shorter sepals come from Pinchincha in Ecuador ( B. Sparre 14810 (S) and Piura in Peru ( E. Laure 5370 (P)). All of these plants have a large corolla 4.5-5 cm in length. They integrade with more typical forms of I. batatas in western Ecuador. Another distinct form comes from around Esmeraldas in Ecuador. This is a glabrous or sparsely pubescent twining herb with unequal, chartaceous, obovate to obrhomboid sepals with a single prominent central nerve extended as a mucro, the outer sepals 5-6 x 3 mm, the inner 7-8 x 4 mm. This was identified as a tetraploid form of Ipomoea batatas by Austin et al. (1992) although it had sometimes been previously identified as I. triloba or I. trifida . Examples include H. Balslev & W.C. Steere 3131 (GB), J. Hudson 730 (MO, US), L. Holm-Nielsen et al. 25318 (AAU, ARIZ) and B. Sparre 15286 (S), 15308 (S), 15341 (S) and 15517 (S).