606 John R.I. Wood et al. / PhytoKeys 143: 1–823 (2020)

3501 (NY). Dominica: Wilbur et al. 7984 (BM). Martinique: Hahn 1200 (BM). St Lu- cia: G.R. Proctor 18117 (BM). St Vincent: H.H. & G.W. Smith 490 (K). Grenada: G.R. Proctor 17206 (BM); P. Beard 1266 (K, NY, S). Barbados: E.G.B. Gooding 189 (BM). TRINIDAD. W.E. Broadway 9401 (K). Tobago: H.F.A. von Eggers 5624 (K). NETHERLANDS ANTILLES. Aruba, Bonaire, Curaçao fide Proosdij (2012). HAWAII. Phillips & Johnson 716 (MO); A.A. Heller 2097 (BM); G.W. Barklay 1328 (BM). Notes. Ipomoea pes-caprae is commonly divided into two subspecies or varieties. Only subsp. brasiliensis (or var. emarginata, if recognised at varietal level) occurs in the New World. It is recognised by its emarginate leaves, whereas the type from the north- ern Indian Ocean area has deeply bilobed leaves, the lobes somewhat divergent. Opin- ions about the status of these two forms have varied over the years. Recent molecular studies (Miryeganeh et al. 2014) suggest the two forms are genetically separate and rarely hybridise but some intermediates occur and the issue is not yet fully resolved. Ipomoea pes-caprae is sometimes confused with I. asarifolia but the latter has sub- reniform leaves and very unequal, often muricate sepals. Molecular data suggests this species is most closely related to a small clade of Aus- tralian species. It is widespread on tropical sea shores. Its total world distribution is given in detail by St John (1970). We have been unable to trace any publication data for the combination. Ipomoea pes-caprae var. brasiliensis (L.) A. St.-Hil.

340. Ipomoea amnicola Morong in Morong & Britton, Ann. New York Acad. Sc. 7: 170. 1892. (Morong and Britton 1892: 170)

Type. PARAGUAY. Banks of the Pilcomayo, T. Morong 974 (lectotype NY00319140, designated here, isolectotypes MO, NY, R). Description. Somewhat succulent twining or trailing perennial, completely gla- brous in all parts. Leaves petiolate, 2–8(–12) × 2–8-(10) cm, ovate, sometimes broadly so, usually constricted in the middle to form a tapering acuminate apical portion, base cordate with rounded auricles, abaxially slightly glaucous; petioles 1–10 cm. Inflores- cence of lax to rather dense, many-flowered, pedunculate, simple or compound cymes; peduncles 1–5 cm; bracteoles 1–3 mm, lanceolate to ovate, caducous; secondary pe- duncles 5–20 mm; pedicels 0.8–2.5 cm; sepals slightly unequal, coriaceous, glabrous, outer 4–6 × 3–4 mm long, ovate-elliptic, convex, obtuse and shortly mucronate, inner 5–7 × 4–5 mm long, broadly oblong-elliptic to obovate, rounded, with broad scarious margins; corolla 2–5.5 cm long, pale lilac to pink with dark centre, glabrous, funnel- shaped, the limb 2.5–3.5 cm diam., unlobed. Capsules 7–12 × 6 mm, conical, shortly rostrate, glabrous; seeds 5–7 × 2.5–4 mm, reddish brown, the surface minutely tomen- tellous, the angles densely pilose. We recognise two subspecies, which intergrade in the region around the Pantanal and perhaps elsewhere. A foundation monograph of Ipomoea in the New World 607

340a. Ipomoea amnicola subsp. amnicola

Ipomoea nuda N.E. Br. Trans. & Proc. Bot. Soc. Edinb. 20: 63. 1894, nom. illeg., non Ipomoea nuda Peter 1891. (Brown, NE: 1894: 63). Type. PARAGUAY. RVo Pilco- mayo. J.G. Kerr 12 (not found at K).

Diagnosis. Inflorescence of usually rather dense axillary cymes; peduncles 1–5 cm; outer sepals 4–5 mm long, inner sepals 5–5.5 mm long; corolla 2–3 cm long, pale lilac with dark centre, the limb 2.5–3 cm diam; seeds 5 × 2.5 mm. Illustration. Figures 2G, 141E, 162. Distribution. This subspecies has an amphitropical distribution being found in the southern United States and South America. In South America it is most common as a species of dry Chaco scrub near the Andes in western Argentina, western Paraguay and southern Bolivia but penetrates the Andean cordillera along dry river valleys. It also occurs in dry areas of NW Peru and neighbouring parts of Ecuador and in the upper Magdalena valley in Colombia. In the United States it is perhaps introduced and is most common in the Rio Grande region of Texas. No records from Mexico have been traced. ARGENTINA. Catamarca: Brizuela 626 (LIL); Pomán, P.D. Cantino 807 (CORD, GH). Chaco: C. O’Donell 5563 (LIL). Córdoba: Cuezzo 903 (LIL); Pocho, A.T. Hunziker & J.A, Caro 13477 (CORD). Corrientes: T.M. Pedersen 3866 (C, P, S); A. Schinini 4470 (ASU, CTES). Formosa: S. Pierotti 4175 (LIL, P). Jujuy: A.L. Cabrera 34061 (MO). La Rioja: Stucker 17135 (LIL); General Ángel Peñalosa, A.T. Hunziker et al. 15117 (CORD, MO). Salta: L.J. Novara et al. 8901 (S). Santa Fe: S. Venturi 297 (LIL). Santiago del Estero: T. Meyer 17076 (LIL). PARAGUAY. Chaco región. Alto Paraguay: F. Mereles 6728 (FCQ). Boquerón: F. Mereles & R. Degen 5150 (FCQ), 5680 (FCQ), 5948 (CTES, FCQ). Central: E. Zardini 2674 (FCQ, MO). Paraguarí: Carpegua, T. Rojas 3371 (S). Presidente Hayes: Maroma, M. Peña-Chocarro et al. 1918 (BM, 2556 (BM); F. Mereles & R. Degen 6425 (FCQ). BRAZIL. Mato Grosso do Sul: Faz. Uberaba, J. Almeida de Jesus 1735 (RB); Es- trada Pantaneira, E.P. Heringer 831 (NY). BOLIVIA. Inter-andean dry valleys and chaco. Chuquisaca: 100 km E of Boy- uibe, B. Mostacedo & T.J. Killeen 354 (NY, LPB, USZ); Zudañez, Puente Inca, J.R.I. Wood et al. 2724 (K, LPB, USZ). Cochabamba: Campero, Puente Arce, J.R.I. Wood 28119 (K, OXF, USZ). La Paz: Sud Yungas: S.G. Beck 22444 (K, LPB); Tamayo, ANMI Madidi, A. Araujo-M et al. 2869 (LPB, MO). Potosí: Charcas, Río Caine bridge, J.R.I. Wood et al. 23244 (K, LPB). Santa Cruz: Ángel Sandoval, Candelaria, J.R.I. Wood et al. 24870 (K, LPB, UB, USZ). Chiquitos, Taperas: J.R.I. Wood et al. 27873 (K, LPB, USZ). Caballero: La Palisada, J.R.I. Wood & A. Haigh 21839 (K, LPB, P); Cordillera, Abapó, J.R.I. Wood & F. Mamani 27484 (K, LPB, USZ). Ibañez, M. Nee 49480 (LPB, MO, NY, USZ); Ñuflo de Chávez, San Julián,J.R.I. Wood & D. Soto 27947 (K, LPB, OXF, USZ); Vallegrande, Río Grande, G.A. Parada et al. 4387 (MO, USZ). Tarija: Gran Chaco, Palos Blancos, J.R.I. Wood et al. 28028 (LPB, OXF, USZ). 608 John R.I. Wood et al. / PhytoKeys 143: 1–823 (2020)

Figure 162. Ipomoea amnicola subsp. amnicola. A habit B outer sepal C inner sepal D corolla opened out to show stamens E ovary and style F habit with capsules G seed. Drawn by Rosemary Wise A–E from Wood & Mamani 27484; F–G from Wood & D. Soto 27947.

PERU. Amazonas: Río Chamaya, Bagua-Olmos, T. Croat 58302 (MO). Ca- jamarca: T. Croat 58367A (MO); P.C. Hutchison & J.K. Wright 6734 (F, UC). Lam- bayeque: Llatas Quiroz 2402 (F). ECUADOR. Loja: La Toma-El Tambo, J.E. Madsen et al. 7772 (AAU). COLOMBIA. Upper Magdalena Valley. Huila: F.R. Fosberg 19610 (US). Tolima: Honda, E. André 561 (K). UNITED STATES. Georgia: Spalding County, W. Hardcastle s.n. (GA); Mis- souri: Jackson, B.F. Bush 9691 (BM, MO). Texas: Cameron County, R. Runyon 2916 A foundation monograph of Ipomoea in the New World 609

(BM), 2904 (S); Hidalgo County, E.U. Clover 301 (MEXU); Kleberg County, W.R. Carr 25097 (MEXU). Typification. There are two sheets of Morong 974 at NY. We have selected the best of these as lectotype, rather than the sheet labelled as holotype in an unknown hand as this lacks most diagnostic details. Note. In the field Ipomoea amnicola (especially subsp. amnicola) is usually easily recognised by the relatively small corolla which is pale pink with a dark centre. It often blankets shrubs and small trees where it occurs. The leaves are quite glabrous, usually somewhat glaucous and slightly fleshy. It is not a very easy plant to dry successfully so leaves are often deciduous on herbarium specimens. It can be confused rather easily with species from the Batatas Clade.

340b. Ipomoea amnicola subsp. chiliantha (Hallier f.) J.R.I. Wood & Scotland, comb. & stat. nov urn:lsid:ipni.org:names:77208082-1

Ipomoea chiliantha Hallier f., Bull. Herb. Boiss. 7 (5), append. 1: 50. 1899. (Hallier 1899c: 50). Type. PARAGUAY. “Villa occidental”, Lorentz s.n. (holotype B†, lec- totype GOET, designated by Wood et al. 2015: 29).

Type. Based on Ipomoea chiliantha Hallier f. Diagnosis. Inflorescence of usually long pedunculate, axillary cymes, sometimes compounded; peduncles 5–13 cm; outer sepals c. 6 × 4 mm long, inner c. 7 × 5 mm; co- rolla 4–5.5 cm long, pink, darker in the centre, limb 3–3.5 cm diam.; seeds 7 × 4 mm. Illustration. Figure 163; O’Donell (1959b: 147) as Ipomoea chiliantha. Distribution. This subspecies seems to prefer seasonally flooded swampy ground both in Bolivia, Paraguay and the Brazilian Pantanal. ARGENTINA. Misiones: T.M. Pedersen 5497 (C, S). Chaco: Isla Anequera, A. Krapovickas & C. Cristóbal 12733 (CTES), A. G. Schulz 2059 (CTES, LIL). Formosa: Dept. Pilcomayo, C. Cristóbal et al. 2146 (CTES), Santa Fe: Pensiero & Tivano 3212 (CTES). Corrientes: Dept. Capital, S.G. Tressens et al. 769 (CTES, MO). PARAGUAY. Alto Paraguay: Est. Cerrito, F. Mereles 7006 (FCQ). Central: Ypacaraí, E. Hassler 11582 (BM, K), 12532 (BM). Concepción: San Luis: K. Fiebrig 4485A (BM, K, MO). Cordillera: E. Hassler 1856 (K); E. Zardini & U. Velázquez (MO). Presidente Hayes: Est. Santa Maria del 12, M. Peña-Chocarro et al. 2565 (BM, FCQ); km 130, Ruta Transchaco, F. Mereles 2244 (CTES), Puente Remanso, F. Mereles 4460 (FCQ); km 58, Ruta Transchaco, A. Krapovickas & C. Cristóbal 43220 (CTES, F, FCQ, K). San Pedro: abundant in plain west of Com. 25 de Diciembre, J.R.I. Wood & G. González 28473 (FCQ); between Río Apa y Río Aquidaban, K. Fiebrig 4483 (BM). BRAZIL. Mato Grosso do Sul: Puerto Murtinho, Robert 885 (K). BOLIVIA. Beni: Cercado, Puerto Varodor, Maldonado et al. 58 (LPB). Santa Cruz: Chiquitos, El Tinto–Quimome, J.R.I. Wood & B. Williams (OXF, K, LPB, USZ); 610 John R.I. Wood et al. / PhytoKeys 143: 1–823 (2020)

Figure 163. Ipomoea amnicola subsp. chiliantha. A habit B outer sepal C inner sepal D corolla opened out to show stamens E ovary and style F fruiting cyme G fruiting calyx and capsule. Drawn by Rosemary Wise A E from Mereles 162; B–D from Petersen 5497; F, G from Mereles 2550.

Germán Busch: Yacuses, J.R.I. Wood & D. Villarroel 25541 (K, LPB, UB, USZ); Ñuflo de Chávez J.R. Abbott 16966 (BOLV, HSB, LPB, MO, USZ); Puente San Miguelito, J.R.I. Wood et al. 27743 (OXF, K, LPB, USZ). Note. Subsp. chiliantha has the appearance of a large-flowered subsp. amnicola and intermediates occur particularly in the Corumbá region. Parada et al. 947 (USZ, MO, OXF) from Carmen Rivero Tórrez and Wood & Pozo 26078 (K, LPB, UB, USZ) from San José de Chiquitos are examples from Bolivia but intermediates appear more com- monly in the Brazilian Pantanal.

• Species 341–343 comprise a small clade with distinctive ribbed sepals