27a. Syngonium podophyllum Schott, Bot. Zeitung (Berlin) 85:1851. TYPE: Mexico (Schott drawings 3223 and 3226; NYBG Photos 4330 and 4329 respectively).

Pothos auritus Willd. ex Schult., Mant. 3:301. 1827, non Syngonium auritum (L.) Schott. TYPE: Amer. Mend. Humboldt & Bonpland (not seen).
Arum auritum
Veil. non L., Fl. Flum. 9. tab. 113. 1827 (drawing only; no text). TYPE: Brazil (plate only).
Xanthosoma gracile Miq., Delect. Sem. Hort. Amstelodam. 1853. TYPE: Venezuela, Caracas, introduced into horticulture in Amsterdam.
Syngonium vellouanum
Schott, Oersterr. Bot. Wochenbl. 4:418. 1854. TYPE: Brazil (Schott drawings 3239 and 3240; NYBG Photos 4318 and 4319 respectively).
S. ruizii
Schott, Oesterr. Bot. Wochenbl. 4:148. 1854. TYPE: Peru (Schott drawing, NYBG Photo
S. gracile (Miq.) Schott, Syn. Aroid. 70.1856. S. riedelianum Schott, Syn. Aroid. 70. 1856. TYPE: Brazil (based on Philodendron riedelianum hort.
non Sehott, not seen). 5. decipiens Schott, Syn. Aroid. 69. 1856. TYPE: Brazil, Ilheus (based on Caladium arboreum hort., Schott drawing 3200; NYBG Photo 4320 of collection of Enherwg Maximillian 719).
S. poeppigii Schott, Syn. Aroid. 68. 1856. TYPE: Peru, Yurimaguas, Poeppig (Schott drawing 3226; NYBG Photo 4321) (see discussion).
S. affine Schott, Syn. Aroid. 67. 1856. TYPE: Surinam, Paramaribo, Wullschtagel 504 (Schott drawing
3192; NYBG Photo 4323).
S. willdenowii Schott, Syn. Aroid. 68. 1856. TYPE: Based on Pothos auritus Willd., Herb. No. 3104. fol. 1, Humboldt (P-, not seen).
S. xanthophilum Sehott, Prodr. Syst. Aroid. 214. 1860. TYPE: Mexico (specimen at W destroyed);
photo FM 29870; photo of Schott drawing, NYBG Photo 4335) (see discussion).
S. velloiianum Schott var. latilobum Engler, Fl. Bras. 3(2): 130. 1878. TYPE: Brazil, Rio de Janeiro,
S. vellownum var. riedelianum (Schott) Engler, Fl. Bras. 3(2): 130. 1878. S. vellownum var. decipiens (Schott) Engler, Fl. Bras. 3(2): 130. 1878. S. velloiianum var. poeppigii (Schott) Engler, Fl. Bras. 3(2): 130. 1878. S. amawnicum Engler, Pflanzenr. IV.23E (Heft 71): 128. 1920. TYPE: Brazil, Amazonas, Rio Jurua,
Vie 5614 (B, holotype). S. podophyllum var. multisectum Engler, Pflanzenr. IV.23E (Heft 71): 129. 1920. TYPE: Martinique,
Duss 519 (not seen). S. ternatum Gleason, Bull. Torrey Bot. Club 56:13. 1929. TYPE: Guyana, Waini River, De La Cruz.
1288 (NY, holotype).

Juvenile plant with stems slightly glaucous; petioles sheathed l/'^-2/3 their length; blades simple, cordate, 7-14 cm long, becoming sagittate or hastate, acuminate at the apex, the anterior lobe somewhat constricted at the base, the posterior lobes usually more or less triangular, directed downward or prominently outward; blades on climbing plants 12-27 cm long. Adult plants with stems sometimes glaucous; sap milky; internodes 2.8-14.5 cm long, 0.5-3.5 cm wide (dry);
petioles 15-60 cm long, sometimes glaucous, sheathed 2/3 their length (the sheath free-ending), rounded to obtusely angular above the sheath; blades pedatisect, the surface dark green above, pale below (sometimes glaucescent in South America); leaflets 3-11, united to free; lowermost leaflet variously auriculate at the base, the auricles oblong to oblong-elliptic to broadly elliptic; median leaflet ob-ovate to broadly elliptic, abruptly acuminate at the apex, broadly or narrowly decurrent at the base, 16-38 cm long, 6-17 cm wide; rachis usually angular on blades with more than 3 segments; primary lateral veins 3-4(-7) pairs on the median leaflet, sunken above, prominently raised beneath; collective veins 2 or 3; tertiary veins all distinct.
Inflorescences 4-11 per axil; peduncle sometimes glaucous somewhat compressed laterally, erect and usually less than 9 cm long at anthesis, pendent, to 13 cm long in fruit; spathe 9-11 cm long; spathe tube sometimes glaucous, narrowly ovoid to ellipsoid, 3-4 cm long, 1.8-2 cm diam., green inside and out; spathe blade greenish white to creamy white or sometimes yellow (in South America) (sometimes green outside and creamy white inside), 6-7.5 cm long, 3-5 cm wide, long-cuspidate at the apex; pistillate portion of the spadix 1-2 cm long, 6-9 mm diam., greenish cream, the flowers irregularly (5-6(-7)-sided, adherent, the stigma discoid-capitate, sometimes 2- or 3-lobed; sta-minate portion of the spadix 4-7 cm long, 7-15 mm diam., cream, the synandrium with anthers usually 4, cross-shaped, 3.5-4 mm wide, partially or completely fused.
Infructescences red to reddish orange or yellow (rarely brown) at maturity; syncarp ovoid, brownish, scruffy with darker brown flecks, 3-5.5(-7) cm long, 1.5-3.5 cm wide; seeds many, ovoid, 7-11 mm long, 5-7 mm wide, black or brown, enveloped in a soft, grayish, sweet, pulpy mesocarp. Figs. 64, 65, 69.

DISTRIBUTION: Syngonium podophyllum is the most widespread and variable species in the genus, ranging from Mexico (states of San Luis Potosf and Vera-cruz) to the Guianas, Brazil, and Bolivia. It is found principally on the Atlantic slopes in Mexico but commonly extends to the Pacific slope in Guatemala, Costa Rica, and Panama. It ranges from sea level to usually less than 1000 m and is more abundant below 750 m, especially between 100 and 500 m. Ellenberg 3857 (Peru, Tingo Maria-Pucalpa) at 1510 m and Lehmann 5316 (Colombia, Rio Negro in Andes of Popayan) at 1400-1600 m are both well out of the normal range of the species. They are sterile collections and doubtfully placed with S. podophyllum. In Central America it is most frequent in regions of tropical moist forest but also occurs in premontane wet forest. It is replaced by S. macrophyllum in tropical wet forest life zones.
Syngonium podophyllum and S. macrophyllum are often difficult to separate where they occur together in wetter parts of tropical moist forest and in premontane wet forest, but S. macrophyllum is usually distinguishable by its much larger vegetative and fertile parts, its smoother, thicker blades, which tend to dry brown rather than greenish as in S. podophyllum, and by its ovate-cordate juvenile blades.
The morphological variation in the species is great. Mexican plants may have up to 11 leaflets (usually 9), have leaflets often more widely separated and have stems, peduncles and spathe tubes glaucous, whereas Panamanian populations generally have mostly 3-5 (sometimes 7) leaflets with the lowermost leaflet often conspicuously auriculate.
Unusual variation is also represented in collections from Darien Province, Panama. Here the auricles of the lateral leaflets are frequently obovate and scarcely pinched-off rather than more or less oblong and usually conspicuously pinched-off as is typical of most Central American populations. These Darien populations most closely resemble typical material of what was previously considered S. vellozianum.
Infructescences in Mexico are commonly bright red whereas those in Panama and in South America are generally yellow to orange or red orange.
In Central America S. podophyllum is most closely related to and confused with S. angustatum. The two species occur together in similar habitats throughout much of their range in Central America, and according to Birdsey (1955) they may hybridize. (See the Key for characters used in separating the two species.) Syngonium podophyllum may also be confused with S. neglectum Schott in its vegetative condition. In the field S. podophyllum can be distinguished from S. neglectum by its glaucous stems. Although the species tends not to have glaucous
stems in the southern part of its range, there is no confusion with S. neglectum, which is restricted to Mexico.
The species is equally variable in South America. I have been unable to separate Central American material from S. velloyanum of South America. The widespread S. vellovanum was treated by Engler & Krause (1920) as a distinct species based on its having leaflets with the leaf tissue united at the base. However, this is not always the case and the degree of separation of the leaf segments is in fact quite variable, ranging from leaves which have the segments closely united to leaves which have the segments relatively remote from one another. The same is true for the species in Central America, as I have already noted. Other variations in South American material of S. podophyllum involve the shape and size of the auricles on the lateral lobes. South American material generally has fewer leaflets (most commonly 3 distinct leaflets with the lateral leaflets conspicuously auriculate) and corresponds to the species in Panama and the lower part of Central America; however, it may have up to 5 distinct leaflets with a prominent "pinched-off auricle, such as in material Engler and Krause called S. amawnicum. All of these characters are extremely variable and are not consistent in material from a single geographical area.
Another synonym of S. podophyllum is S. ternatum. Type material with this name, like that of S. yurimaguense, has poorly developed auricles on the lateral lobes. Considering the immense variability of S. podophyllum in a single population (or even on a single plant) there is no justification for maintaining these different leaf forms.
I believe that S. xanthophilum is a synonym of S. podophyllum. The description was of a sterile collection cultivated in Vienna. The collection as illustrated by Schott's painting (NYBG Photo 4335) represents an apparently juvenile plant. The leaves are remarkably similar to juvenile leaves of S. podophyllum. Further evidence that the Schott illustration represents a juvenile of S. podophyllum is that a photograph of an adult specimen of S. podophyllum was also later made at Vienna. This specimen seen in Field Museum Type Photo 29870 was perhaps collected from the same plant after it was more mature.
Mention should be made of the name S. auritum Poeppig which occurs in the literature. Poeppig (1845) prepared a description of S. auritum Schott with apparently no intention of considering it a new species. Schott (1856) described S. poeppigii saying that it was based on S. auritum Poeppig, a nonexistent name. Engler & Krause (1920) in their revision of Syngonium included S. poeppigii Schott and S. auritum Poeppig in synonymy with S. vellozianum var. oblongi-sectum Engler thus perpetuating an error by Schott.
Although South American material of S. podophyllum may ultimately prove to consist of other subspecies or varieties with additional field work, enough is known at this time to recognize S. peliocladum as a variety of S. podophyllum. Variety peliocladum differs from the typical variety only in having tuberculate projections on the stems. Tom Ray (pers. comm.), who has studied populations in the field in Costa Rica, suspects that these projections may have no taxonomic significance whatever.
Syngonium podophyllum has been collected in flower and fruit throughout the year.
Valley, Pffeifer 2069 (US), Standley 52751, 53330, 54537, 56603, 56648 (F, US), 53486 (US). Yuncker 5091 (F), 5090 (MICH). COPAN: Below Copan Antigua, Barkley & Herndndez 40278 (GH); E of Copan, Croat 42512, 42524, 42527 (MO); Vicinity of ruins of Copan, Carbon 533 (F, UC). CORTES:
N ofLago de Yojoa, Croat 42719 (MO). EL CAYO: Vaca, Gentle 2599 (MICH). YORO: Quebrada Sica, Standley 53931 (US). SANTA BARBARA: San Pedro Sula, Thieme 5529 (US).
MEXICO: No other location, Kenoyer 428 (F). CAMPECHE: Santa Leonor, Barlow 14-3 (MICH, WIS). CHIAPAS: Between Chiapilla and San Lucas, Laughlin 278 (F, MEXU); Escuintia, Matuda 16723 (GH), s.n. (7-12-47, 8-30-47) (MEXU), 17939, 20946 (MEXU); S of Isthuatan, Croat 47835 (CAS, MEXU, MO); N of Isthuatan, Croat 47870 (MO); SE of Mapastepec, Breedlove & Thorne 30680 (MO); Munc. of Las Margaritas, on Guatemala Border, Breedlove & McClintock 34088 (DS);
Ocosingo, Breedlove 33950, 34463 (DS); N of Ocozocoautia, Breedlove 38214 (DS), Croat 40629, 40652 (MO); SW of Palenque, Croat 40318, 40338 (MO); N of Pichucaico, Croat 40083 (MO); NW of Pueblo Nuevo Solistahuacan, Croat 47799 (MO); WNW ofSoyalo, Breedlove 37216 (DS); Between Tapachula and Union Juarez, Croat 47204 (MO). GUERRERO: Acapuico and vicinity. Palmar 346 (F, GH, MEXU, MO, UC, US); E ofAgua de Obispo, Crisman & Willis 292 (MICH); NAYARIT: NW of Jalcacatlan, Croat 45306 (MO). OAXACA: Cahuapa River, Williams 8905 (F); Island of Malsaya, Quintero 657 (MEXU); Near Junction of Ismo Highway and Road to Matfas Romero, Carlson 2193 a (F, MO); Tuxtepec, Chavelas & Perry 237 (MEXU); Chavelas 278 (MEXU); Martinez & Calderon 556 (GH, MEXU, UC, US); Ubrero, Williams 9290, 9525 (F); S of Valle Nacional, Croat 39721, 39737 (MO); NE of Valle Nacional, Moore & Bunting 8909 (BH). PUEBLA: No other location, Bravo 520 (MEXU); Near Maria Andrea, Rzedowski 23379 (MICH). SAN LUIS POTOSI: Palmer s.n. (US);
NW of Huichihuayan, Croat 39247 (MO); NW of Tamazunchale, Croat 39282 (MO), Dunn & Dunn 19152 (MO); Xilitia, Villar & Herndndez 7145 (MEXU). TABASCO: W of Cardenas, Conrad et al. 2767 (MO); Road to del Golfo, Contu 176B (MEXU); Between Macuspana and El Carmen, Gilly & Herndndez 367 (MICH); Near Teapa, Matuda 37496 (MEXU). TAMAULIPAS: NNW of Gomez Farias, Martin 69 (MICH). VERACRUZ: Lot 637 (Jalapa); Near Boca del Rio, Paxson et al. 17M634 (F, MEXU); Campo Experimental de Hule, El Palmar Zongolica, Santos 2317, 3634 (MICH); Cana de las Perdidas, Chavelas et al. 2372 (MEXU); W ofCatemaco, Quintero 1495 (MEXU); Cerro Gordo, Dorantes et al. 4261 (Jalapa); E ofCoatzacoalcos, Croat 40061 (MO); Coatzacoalcos River, Williams 8420 (MICH); Between Conejo and Huatusco, Moore & Bunting 8865 (BH, MO); Along Highway 185, N of border of Veracruz, Croat 40010 (MO); Hidalgotitlan, Vdzquez 782 (F, Jalapa, MEXU);
S of Highway between Cordoba and Veracruz, Croat 39616 (MO); S side of Laguna de Sontecomapau, Lot 1285 (F, GH, MEXU); Lake Catemaco, Calzada 54 (F); Road to La Palma and Balzapote, Calzada 2447 (F, Jalapa, MO); Las Choapas, Soto & Horowitz 49 (Jalapa); N of Martinez de laTorre, Nevling & Gomez-Pompa, 532 (GH, MEXU); NE of Minatitlan, King 1042 (MICH); District ofPapantIa, Kelly 156, 269 (BH); Between Papantia and Tajin, Moore & Bunting 8955 (BH); Road to Playa Escondida, Breckon & Breckon 2043 (MO); Rancho Tepellaw, Herndndez s.n. (Jalapa);
San Andres Tuxtia, Dressier & Jones 25 (GH), Kennedy & Horvitz 3687 (F), Soto & Horvitz 30 (F), Sousa 2107 (MEXU); San Francisco, Smith 1307 (F); E of San Pablo, Gutierrez 9 (MEXU); Vicinity Santiago Tuxtia, Leija & Garza 5418 (MEXU); On Road to Santecomapan, Moore & Bunting 8934 (BH); W of Sayula de Aleman, Croat 40023 (MO); S of Tampico, Palmer 386 (GH, MO, US);
YUCATAN: Chichen Itza, Lundell & Lundell 7506 (MEXU, MICH).
NICARAGUA: CHONTALES: Above Cuapa, Stevens 3694 (MO). JINOTEGA: SE ofYali, Croat 42934 (MO). ZELAYA: Alamicamba above Rio Prinzapoica, Neill 3901 (MO); E of Siuna, Neill 4511 (MO).
PANAMA: BOCAS DEL TORO: Chiriquicito, Lewis et al. 2049 (GH, MO); Vicinity of Chiriqui Lagoon, Von Wedel 2616 (GH); Station Milla 7.5, Croat 38128 (MO). CANAL ZONE: Barro Colorado Island, Aviles35 (F), Croat 5815, 7027, 7429, 8913, 10197, 11694, 16218, 20888 (MO), 10260, 11426, 11921 (F, MO), 10876 (F, MO, PMA, UC), 12558 (F, MO, SCZ), Foster 1939 (F, GH, MICH, PMA), Gentry 428 (MO), Hutchison & Wright 2864 (UC), Shattuck 54 (F, GH, MO), Standley 40914 (US), Woodworth & Vestal 585 (F, GH, MO); SE of Achiote, Gentry & Nee 8663 (MO); Near Alhajuela, Dodge 16596 (MO); N of Frijoles, Standley 27451 (US); Vicinity Gamboa, Croat 32975 (MO), 32977 (PMA), Nee 7745 (US), Standley 28419 (US); Quebrada Bonita, Steyermark & Alien 17211 (MO); S of Rio Providencia, Tyson & Blum 4003 (MO, SCZ); Near Summit Gardens, Dodge & Hunter 8661 (MO). CHIRIQUI: Burica Peninsula, vicinity of Puerto Armuelles, Busey 466 (MO, NY), Croat 21936, 26704, 35055 (MO), Liesner 67 (F, MO), 504 (MO, US), Woodson & Schery 930 (F, NY); Vicinity of Gualaca, Croat 49822 (MO); Vicinity of Palmas Bellas, Mori & Kallunki 1971 (MO). COLON: SW ofPortobelo, Liesner 1080 (US); E ofPortobelo, Liesner 1109 (MO, US); Vicinity of Portobelo and Nombre de Dios, Croat 33553, 33636, 49784 (MO); Vicinity Portobelo, Croat 49746 (MO). DARIEN:
Vicinity of gold mine at Cana, Croat 37599, 37659, 37676, 38034 (MO); Cerro Sapo, Hammel 1257 (MO); Vicinity El Real, Duke 5013 (MO, GH), Gentry & Clewell 6917 (MO), Pittier 6568 (GH, US);
Vicinity of Pay a. Stern et al. 199 (GH, MO, US); Rio Tuquesa, Le Clew 188 (MO), PANAMA:
Vicinity of Cerro Azul, Croat 11508 (MO); El Llano-Carti Road, Croat 33776 (MO), Nee 10520 (US);
Vicinity Rio Maje, Croat 14524, 34438, 34658 (MO).
PERU: AMAZONAS: Rio Cenepa, Ancuash 122, 1157, 1193 (MO), Berlin 371 (MO); S of Chavez Valdivia, Berlin 838 (MO). AYACUCHO: Near Kimpitirique, Killip & Smith 23007 (NY, US). HUA-NUCO: Honoria, Schunke 1592 (F); Tingo Maria, Ellenberg 3826, 3857 (MO). JUNI'N: N of La Merced, Killip & Smith 23626 (NY, US). LORETO: Iquitos, Killip & Smith 27398, 27448 (NY, US), Martin et at. 1636 (F); La Victoria, Williams 2631 (F), 2782 (F, US); Letieia, Williams 3054 (F); Province Maynas, Diaz & Jaramillo 9 (MO), Revilla 1087 (MO); N of Iquitos, Revilla 855 (MO); Rio Huallaga, Williams 4986 (F, US), Croat 17855 (MO); Soledad on Rio Itaya, Killip & Smith 29680 (NY, US);
NE of Yurimaguas, Kiltip & Smith 27901 (US). MADRE DE DIOS: Parque Nacional de Manu, Foster & Janson 5216 (F), Gentry et at. 26771 (MO), Terborgh 6624 (F); Tambopata, Gentry & Revilla 16299 (F, MO, NY). SAN MARTIN: Mariscal Caceres, Gentry et al. 26733 (MO), Schunke 3862 4932 (F, GH, MO, NY, US), 7711 (MO, US).
PUERTO Rico: At Km 28.1 on Route 191 near Florida, Wagner 1003 (M1CH).
SINGAPORE: No other location, Clemens 22553 (UC).
SURINAM: No other location, Samuels 175 (GH, K); Corantijn River, Boer & Wessels 548 (A);
Slopes of Hendriktop, Daniels 978 (US); Jodensavanne-Mapane creek area, Lindeman 4782 (F, MO, NY); Lucie River, Maguire et al. 53991 (NY), Prance et al. 54619 (F, K, NY, US), Prance et al. 55668 (NY); N of Paramaribo, Maguire & Stahel 22800 (F, NY, US); Tibiti, Lanjouw & Lindeman 1616 (K, NY, US).
TRINIDAD: Siparia Quarry, Britton & Broadway 2795 (NY, US).
UNITED STATES: FLORIDA: Coral Gables, Kaplan 478 (UC); Madiera Beach, Anderson & Mason 19328 (CM).
VENEZUELA: AMAZONAS: Tencua, Colchester 2052 (K); Vicinity Mision de Santa Maria de los Guaicas, Steyermark 106164 (MO, VEN). APURE: San Camilo Forest Reserve, Steyermark et al. 101396 (MO, NY, VEN). BARINAS: Vicinity of Barinas, Bunting 2271 (MY); SW of Barinas, Smith 3292 (MY, VEN); Cano Barragan, Bunting 2257 (MY); Caparo Forest Reserve, Steyermark et al. 102051 (MO, VEN); Rio Caparo, Liesner & Con-idle! 9413 (MO). BOLIVAR: S of Plateau of Nuria, Bunting & Holmquist 4317 (MY); Vicinity of Salto Para, Rio Caura, Steyermark et al. 113104 (F, MO, US). DELTA AMACURO: Sierra Imataca, Steyermark 87381 (F, K); Tucupita, Davidse & Gomalez 16548 (MO). MERIDA: Between San Cristobal and El Vigia, Bunting 2403 (MY). TACHIRA: Vicinity of San Cristobal, Broadway 578 (US). Vicinity La Grita, Bunting 2483, 2491, 2497, 2500 (MY); E of Hacienda El Palmar, Steyermark et al. 120463 (MO). TRUJILLO: La Ceiba, Williams 12661 (US). ZULIA: S of Concha, Bunting 2774, 2788 (MY); Hacienda Las Chorros, Farrari & Trujillo 1569 (MY);
Sierra de Perija, Steyermark & Fernanda 99633A (NY, US); SE of Mission of Los Angeles de Tocucu, Steyermark 99878 (US).