Ipomoea L.

Wood, John R. I., Munoz-Rodriguez, Pablo, Williams, Bethany R. M. & Scotland, Robert W., 2020, A foundation monograph of Ipomoea (Convolvulaceae) in the New World, PhytoKeys 143, pp. 1-823 : 1

Ipomoea L. Sp. Pl. 1: 159. 1753, nom. cons.

urn:lsid:ipni.org:names:

Acmostemon Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 13: 106. 1936. (Pilger 1936: 106). Type. Acmostemon angolensis Pilg. (= Ipomoea viscoidea Choisy ).

Adamboe Raf. , Fl. Tellur. 4: 79. 1836 [pub. 1838]. (Rafinesque 1838a: 79). Type. Adamboe bicolor Raf. (= Ipomoea campanulata L.).

Amphione Raf. , Fl. Tellur. 4: 79. 1836 [pub. 1838]. (Rafinesque 1838a: 79). Type. Amphione lobata Raf. (= Ipomoea triloba L.).

Apopleumon Raf. , Fl. Tellur. 4: 72. 1836 [pub. 1838]. (Rafinesque 1838a: 72). Type. Apopleumon bignonioides (Sims) Raf. (= Ipomoea bignoniodes Sims ).

Argyreia Lour. , Fl. Cochinch. 1: 95, 134. 1790. (Loureiro 1790: 134). Type. Argyreia obtusifolia Lour. (= Ipomoea obtusifolia (Lour.) J.R.I. Wood & Scotland).

Argyryon St.-Lag., Ann. Soc. Bot. Lyon. 7: 120. 1880. (Saint-Lager 1880: 120). Orth. Variant of Argyreia .

Astripomoea A. Meeuse , Bothalia 6: 709. 1958. (Meeuse 1958: 709). Type. Based on Astrochlaena Hallier f.

Astrochlaena Hallier f., Bot. Jahrb. Syst. 18: 120. 1894 [pub. 1893]. (Hallier 1893b: 120), nom. illeg., non Astrochlaena Corda (1845) . Type. Astrochlaena lachnosperma (Choisy) Hallier f., lectotype designated by Meeuse, Taxon 4: 199. 1955. (= Ipomoea lachnosperma Choisy ).

Batatas Choisy , Mém. Soc. Phys. Genève 6: 434 [52]. 1833 [pub. 1834]. (Choisy 1834: 434). Type. Batatas edulis (Thunb.) Choisy , lectotype designated by Roberty, Boissiera 10: 147 (1964). (= Ipomoea batatas (L.) Lam.).

Blinkworthia Choisy , Mém. Soc. Phys. Genève 6: 430 [48]. 1833 [pub. 1834]. (Choisy 1834: 430). Type. Blinkworthia lycioides Choisy (= Ipomoea lycioides (Choisy) J.R.I. Wood & Scotland).

Bombycospermum J. Presl , Reliq. Haenk. 2: 137.1835. (Presl 1831-5: 137). Type. Bombycospermum mexicanum J. Presl (= Ipomoea bombycina (Choisy) Benth. & Hook f.).

Bonanox Raf. , Ann. Gén. Sci. Phys. 8: 272. 1821. (Rafinesque 1821: 272). Type. Ipomoea bonanox (= Ipomoea alba L.).

Calboa Cav. , Icon. 5: 51. 1799. (Cavanilles 1799: 51). Type. Calboa vitifolia Cav. (= Ipomoea neei (Spreng.) O'Donell ).

Calonyction Choisy , Mém. Soc. Phys. Genève 6: 441. 1833. [pub. 1834]. (Choisy 1834: 441) Type. Based on Bonanox Raf.

Calycanthemum Klotzsch in W.C.H. Peters , Naturw. Reise Mossambique 6 (Bot. 1): 243. 1861. (Peters 1861: 243). Type. Calycanthemum leucanthemum Klotzsch (= Ipomoea leucanthemum (Klotzsch) Hallier f.).

Cleiemera Raf. , Fl. Tellur. 4: 77. 1836 [pub. 1838]. (Rafinesque 1838a: 77). Type. Several syntypes cited including Cleimera hederacea (Jacq.) Raf. (= Ipomoea hederacea Jacq. ).

Cleiostoma Raf. , Fl. Tellur. 4: 80. 1836 [pub. 1838]. (Rafinesque 1838a: 80). Type. Cleiostoma villosum Raf. (= Ipomoea pes-tigridis L.).

Clitocyamos St. -Lag., Ann. Soc. Bot. Lyon 7: 128. 1880 (Saint-Lager 1880: 128). Type. Clitocyamos pinnatifidus Raf. (= Ipomoea quamoclit L.)

Coiladena Raf. , Fl. Tellur. 2: 12 1836 [pub.1837]. (Rafinesque 1837: 12). Type. Coiladena hyemalis Raf. = Ipomoea sp. incert.

Cryptanthela Gagnep., Notul. Syst. (Paris) 14: 24. 1950. (Gagnepain 1950: 24). Type. Cryptanthela sericea Gagnep. (= Ipomoea thorelii Gagnep. )

Decaloba Raf. , Fl. Tellur. 4: 76 1836 [pub. 1838]. (Rafinesque 1838a: 79). Type. Several syntypes cited, including Decaloba mutabilis (Ker-Gawl) Raf. = Ipomoea indica (Burm.) Merrill

Diatremis Raf. , Ann. Gén. Sci. Phys. 8: 271. 1821. (Rafinesque 1821: 271). Type. Convolvulus nil L. (= Ipomoea nil (L.) Roth).

Dimerodiscus Gagnep., Notul. Syst. (Paris) 14: 25. 1950. (Gagnepain 1950: 25). Type. Dimerodiscus fallax Gagnep. (= Ipomoea polymorpha Roem. & Schult. ).

Doxema Raf. , Fl. Tellur. 4: 75. 1836 [pub. 1838]. (Rafinesque 1838a: 75). Type. Doxena sanguinea (Vahl) Raf. (= Ipomoea hederifolia L.).

Elythrostamna Bojer ex Desjardins, Rapp. Annuel Trav. Soc. Hist. Nat. île Maurice 1: 31. 1836. (Desjardins 1836: 31). Type. Elythrostamna convolvulacea Bojer ex Desjardins (= Ipomoea pileata Roxb. ).

Euryloma Raf. , Fl. Tellur. 4: 75. 1836 [pub. 1838]. (Rafinesque 1838a: 75). Type. Convolvulus grandiflorus L. (= Ipomoea alba L.).

Exallosis Raf. , Fl. Tellur. 4: 83 1836 [pub.1838]. (Rafinesque 1838a: 83). Type. Exallosis biflora (L.) Raf. (= Ipomoea biflora (L.) Pers.).

Exocroa Raf. , Fl. Tellur. 4: 80. 1836 [pub. 1838]. (Rafinesque 1838a: 80). Type. Exocroa egyptiaca Raf. (= Ipomoea cairica (L.) Sweet).

Exogonium Choisy , Mém. Soc. Phys. Genève 6: 443. 1834. (Choisy 1834: 443). Type. Exogonium bracteatum (Cav.) Choisy ex G. Don , lectotype designated by House, Bull. Torrey Bot. Club 35: 98. (1908). (= Ipomoea bracteata Cav. ).

Fraxima Raf. , Fl. Tellur. 4: 83. 1836 [pub. 1838]. (Rafinesque 1838a: 83). Type. Fraxima ebractea (Desr.) Raf. (= Ipomoea ebracteata (Desr.) Choisy ).

Gynoisa Raf. , Fl. Tellur. 4: 75 1836 [pub. 1838]. (Rafinesque 1838a: 75). Type. Gynoisia carolina (L.) Raf. (= Ipomoea cordatotriloba Dennst. ).

Kolofonia Raf. , Fl. Tellur. 4: 73 1836 [pub. 1838]. (Rafinesque 1838a: 73). Type. Kolofonia albivenia (Lindl.) Raf. (= Ipomoea albivenia Lindl. )

Latrienda Raf. Fl. Tellur. 4: 81. 1836 [pub. 1838]. (Rafinesque 1838a: 81). Type. Various syntypes cited including Latrienda soldanella (L.) Raf. (= Calystegia soldanella (L.) R.Br. and Latrienda imperati (Vahl) Raf. (= Ipomoea imperati (Vahl) Griseb. )

Legendrea Webb & Berth, Hist. Nat. Iles. Canar., Bot. 3, 2: 26. 1844. (Webb and Berthelot 1844-50: 26). Type. Legendrea mollissima Webb. & Berthel. (= Ipomoea corymbosa (L.) Roem. & Schult.).

Lepistemon Blume , Bijdr. Fl. Ned. Ind. 722. 1826. (Blume 1826: 722). Type. Lepistemon flavescens Blume (= Ipomoea binectarifera (Wall.) J.R.I. Wood & Scotland).

Lepistemonopsis Dammer, Pflanzenw. Ost-Afrikas C : 331. 1895. (Engler 1895). Type. Lepistemonopsis volkensii Dammer (= Ipomoea volkensii (Dammer) J.R.I. Wood & Scotland).

Lettsomia Roxb. , Fl. Ind. 2: 75. 1824. (Roxburgh 1824: 75). Type. Lettsomia cuneata (Willd.) Roxb. , designated by Pfeiffer (1874: 95). (= Ipomoea cuneata (Willd.) J.R.I. Wood & Scotland).

Marcellia Choisy, Mém. Soc. Phys. Genève 10: 443. 1844. (Choisy: 1844: 443). Type. Marcellia villosa Mart. ex Choisy (= Ipomoea marcellia Meisn. ).

Melascus Raf. , Fl. Tellur. 4: 81 1836 [pub. 1838]. (Rafinesque 1838a: 81). Type. Melascus latifolius Raf. (= Ipomoea alba L.).

Mina Cerv. in La Llave & Lexarza , Nov. Gen. Descr. 1: 3. 1824. (La Llave and Lexarza 1824: 3). Type. Mina lobata Cerv. (= Ipomoea lobata (Cerv.) Thell. ).

Modesta Raf. , Fl. Tellur. 4: 76. 1836 [1838]. (Rafinesque 1838a: 76). Type. Modesta paniculata (L.) Raf. (= Ipomoea mauritiana Jacq. ).

Moorcroftia Choisy , Mém. Soc. Phys. Genève 6: 431. 1833 [pub.1834]. (Choisy 1834: 431). Type. Moorcroftia pinangiana Choisy , lectotype designated by Roberty (1964: 143) = Ipomoea pinangiana (Choisy) J.R.I. Wood & Scotland.

Navipomoea Roberty , Boissiera 10: 147. 1964.. (Roberty 1964: 147). Type. Navipomoea involucrata (P. Beauv.) Roberty (= Ipomoea involucrata P. Beauv. ).

Neorthosis Raf. , Fl. Tellur. 4: 125 1836 [pub. 1838]. (Rafinesque 1838a: 125). Type. Not clearly cited, possibly Neorthosis coccinea (L.) Raf. (= Ipomoea coccinea L.).

Ornithosperma Raf., Fl. Ludov. : 149. 1817. (Rafinesque 1817: 149). Type. Ornithosperma serotina (DC.) Raf. (= Ipomoea orizabensis (G. Pelletan) Ledeb. ex Steud. ).

Paralepistemon Lejoly & Lisowski , Bull. Jard. Bot. Belg. 56: 196. 1986. (Lejoly and Lisowski 1986: 196). Type. Paralepistemon shirensis (Oliv.) Lejoly & Lisowski (= Ipomoea shirensis Oliv. ).

Pentacrostigma K. Afzel. , Svensk. Bot. Tidskr. 23: 181. 1929. (Afzelius 1929: 181). Type. Pentacrostigma nyctanthum K. Afzel. (= Ipomoea longituba Hallier f.).

Pharbitis Choisy , Mém. Soc. Phys. Genève 6: 438. 1833 [pub.1834]. (Choisy 1834: 438). Type. Pharbitis hispida (Zuccagni) Choisy (= Ipomoea purpurea (L.) Roth).

Plesiagopus Raf. , Fl. Tellur. 4: 78. 1836 [1838]. (Rafinesque 1838a: 78). Type. Convolvulus pes-caprae L. (= Ipomoea pes-caprae (L.) R.Br.).

Pseudipomoea Roberty , Boissiera 10: 147. 1964. (Roberty 1964: 147). Type. Pseudipomoea repens (L.) Roberty (= Ipomoea repens (L.) Lam.).

Quamoclit Mill., Gard. Dict. Abr. ed. 4(3). 1754. (Miller 1754). Type. Based on Ipomoea quamoclit L.

Quamoclita Raf. , Fl. Tellur. 4: 74. 1836 [pub. 1838]. (Rafinesque 1838a: 74). Type. Various heterogeneous species cited.

Rivea Choisy , Mém. Soc. Phys. Genève 6: 407. 1833 [pub.1834]. (Choisy 1834: 407). Type. Rivea hypocrateriformis (Desr.) Choisy. Lectotype designated by Manitz (1976: 313). (= Ipomoea hypocrateriformis (Desr.) J.R.I. Wood & Scotland)

Samudra Raf. , Fl. Tellur. 4: 72 1836 [pub. 1838]. (Rafinesque 1838a: 72). Type. Samudra speciosa (L.f.) Raf. (= Ipomoea nervosa (Burm. f.) J.R.I. Wood & Scotland)

Stictocardia Hallier f., Bot. Jahrb. Syst. 18: 159. 1894 [pub. 1893]. (Hallier 1893b: 159). Type. Stictocardia tiliifolia (Desr.) Hallier f. (= Ipomoea tiliifolia (Desr.) Roem. & Schult. )

Stomadena Raf. , Fl. Tellur. 2: 12 1836 [pub.1837]. (Rafinesque 1837: 12). Type. Stomadena violacea Raf. (= Ipomoea sp. incert.)

Tereietra Raf. , Fl. Tellur. 4: 124 1836 [pub.1838]. (Rafinesque 1838a: 124). Type. Tereietra violacea (L.) Raf. (= Ipomoea violacea L.)

Tirtalia Raf. , Fl. Tellur. 4: 71. 1836 [pub. 1838]. (Rafinesque 1838a: 71). Type. Various syntypes cited.

Turbina Raf. , Fl. Tellur. 4: 81. 1836 [pub. 1838]. (Rafinesque 1838a: 81). Type. Turbina corymbosa (L.) Raf. (= Ipomoea corymbosa (L.) Roth ex Roem. & Schult.).

Type.

Ipomoea pes-tigridis L.

Description.

Annual or perennial herbs, subshrubs, lianas, shrubs or small trees, very varied in habit but, most commonly, twining, less commonly decumbent or erect; vegetative parts glabrous or variously hirsute. Leaves without true stipules, alternate, usually petiolate, entire, lobed or compound with separate leaflets; pseudostipules sometimes present. Inflorescence characteristically of axillary cymes, but sometimes very dense and subcapitate or reduced to single flowers or corymbose to foliose paniculate in form, or subterminal and racemose to spicate in erect species; peduncles variable in length, rarely absent; bracts usually indistinguishable from leaves except in species with a terminal inflorescence; bracteoles very small to large, persistent or caducous, scarious, chartaceous or foliaceous, occasionally forming an involucre; pedicels short or long, rarely absent; calyx of five equal or unequal sepals, very variable in texture, coriaceous, herbaceous, scarious, persistent, often enlarging in fruit; corolla ± often showy, small or (usually) large, commonly funnelform, sometimes hypocrateriform, campanulate or suburceolate, pink or white with 5 distinct darker midpetaline bands, the limb distinct from the tube; stamens 5, usually included, equal or unequal in length, dilated and glandular-pilose at base, inserted near base of corolla tube; anthers usually narrowly oblong; pollen spheroidal, pantoporate, echinulate, the grains relatively large; disc annular, ovary 2(-5) locular, 4 (-10)-ovulate, glabrous or pubescent; style simple, filiform; stigma subglobose, 2(-3)-lobed, rarely ( Astripomoea and some species in the Arborescens Clade) lobes somewhat elongate. Fruit a globose, 4 (-10) valved capsule or indehiscent; seeds (1-)4-6(-10), triquetous, ovoid or subglobose, glabrous or variously hirsute.

Distribution.

A mainly tropical genus, which is almost absent from temperate regions. In its widest circumscription (that is including Argyreia and Stictocardia ), it is about equally common in all three tropical regions although the greatest numbers are found in the Americas. A feature of the genus is the existence of a group of around 30 species which are pantropical in distribution, many as the result of early or prehistoric dispersal. There are significant numbers of endemic species on some large islands including Cuba, Hispaniola, Madagascar and Australia but endemics on small islands or island groups are uncommon.

••• Clade A. (Species 1-233). This enormous clade includes over half the species found in the Americas. There is no obvious morphological character that unites the clade but it divides into three smaller clades. Species in the first two of these, Clade A1 (species 1-127) and Clade A2 (Species 128-215), appear always to have pollen with relatively few echinulae (Figure 9A-C ) while that of Clade A3 (Species 218-232) has more numerous echinulae (Figure 10A, B ). In addition, there are 3 species within clade A that lie outside Clades A1-3. These are Ipomoea setosa and I. peruviana (species 216-217) that are sister to each other and together sister to Clades A1-2. Lastly, Ipomoea cryptica (Species 233) is sister to Clade A3. Figure 1 .

•• Clade A1 (Species 1-127) is very heterogenous morphologically although notable for the absence of annual species and of species with a hypocrateriform corolla and exserted stamens. It includes a number of smaller clades, which are indicated in the text, as well as the following major, principally South American, radiation, which we refer to as the Jalapa radiation after its most widespread species.

• The Jalapa radiation (Species 1-83) is centred on Paraguay, Bolivia, southern Brazil and the extreme north of Argentina. It is very poorly represented in North America. The exact boundaries of the radiation are unclear but evidence suggests that Ipomoea carnea (Species 84) and subsequent species should be excluded ( Muñoz Rodríguez et al. 2019). Species in this radiation are very varied in habit but the corolla is always pubescent on the exterior. The sepals are usually ± flat, somewhat soft in texture and pubescent.

The radiation appears to be actively evolving and there are several clusters of species, which are difficult to delimit or are bridged by intermediates. To date molecular studies have not shed much light on these relationships or on species monophyly. Most species are unresolved with samples of some species, notably Ipomoea malvaeoides and I. hirsutissima appearing in several places, although in other cases samples from multiple accessions indicate monophyletic species. Results from the few species for which we have extensive sequence data confirm some species relationships suggested by morphology such as Ipomoea malvaeoides with I. paludosa , or I. argentinica with I. longibarbis but raise serious questions over others that are suggested by morphology, such as I. megapotamica with both I. hieronymi and I. opulifolia or I. nitida with I. psammophila .

Figures

Figure 1. Cladogram showing the principal clades into which Ipomoea can be divided, the two main clades indicated as OWC (Old World Clade) and NWC (New World Clade). The placement of traditionally recognised genera and groups is shown in the corresponding clade.

Figure 9. Pollen of Ipomoea species. A I. hieronymi (Wood et al. 28055) B I. wolcottiana (Hughes et al. 1911) C I. bonariensis (Wood et al. 27871) D I. bahiensis (Queiroz 15975) E I. maurandioides (Krapovickas & Cristobal 1573) F I. corymbosa (Jurgensen 612) G I. sericosepala (Wood 28122) H I. tiliifolia (Beddome 5581). Photos by Robert Scotland.

Figure 10. Pollen of Ipomoea species. A I. triloba (D'Arcy 317) B I. cryptica (Steinbach 6311) C I. purpurea (Parada & Rojas 2664) D I. alba (Wood et al. 27828) E I. dumosa (Hinton et al. 9479) F I. hederifolia (Queiroz 15975) G I. stans (Y. Mexia 275112) H I. suffulta (Pringle 4755). Photos by Robert Scotland.