Passiflora sexocellata Schltdl., Linnaea 27: 521. 1854.

Porter-Utley, Kristen, 2014, A revision of Passiflora L. subgenus Decaloba (DC.) Rchb. supersection Cieca (Medik.) J. M. MacDougal & Feuillet (Passifloraceae), PhytoKeys 43, pp. 1-224 : 118-120

17. Passiflora sexocellata Schltdl., Linnaea 27: 521. 1854. Figs 52, 53

Type.

Mexico. Veracruz: along Hwy. 180 between Tampico and Pozarica, 12 mi N of Ozuluama, 38 km N of Naranjos, 110 m, 5 June 1987, T. B. Croat 66095 (neotype, designated here: MO! [MO-312537]).

Description.

Slender, climbing, perennial vine 2-6 m long or more, sparsely pubescent with unicellular curved trichomes on petiole, leaf, stem, sepal, and stipule, 0.20-0.64 mm long, 0.02-0.03 mm wide, also minutely antrorsely appressed-puberulent throughout with unicellular, curved trichomes, 0.03-0.12 mm long, 0.02-0.03 mm wide. Flowering stems 1.0-2.4 mm in diameter, terete to somewhat compressed, with the base woody and cork-covered. Stipules 2.5-6.0 mm long, 0.4-1.3 mm wide, narrowly ovate-triangular, acute; petioles 1.2-5.7 cm long, with 2 (rarely 3), opposite to subopposite, sessile, discoid nectaries with flat rims, 1.0-2.1 mm wide (on the widest axis), 0.3-1.3 mm high, borne in the proximal half of the petiole (0.34-0.54 of the distance from the base toward the apex of the petiole). Laminas 2.6-8.5 cm long, 6.5-23.5 cm wide, coriaceous, peltate (the distance from leaf base to point of petiole insertion 3.0-18.9 mm), transversely elliptic (widely divaricately bilobed) or sometimes 3-lobed, lateral lobes 3.7-12.9 cm long, 1.9-7.5 cm wide, elliptic, acute to attenuate, central lobe elliptic to obovate or present merely as a widely acute to obtuse tip (rarely retuse), central vein 1.8-7.0 cm long (measured from point of petiole insertion to the leaf apex), angle between the lateral lobes 132-188°, ratio of lateral lobe to central vein length 1.33-2.77, margins entire, hyaline, primary veins 3, diverging and branching above base, laminar nectaries present, 4-13, submarginal, associated with the minor veins of the abaxial surface, with 2-4 nectaries proximal to the lateral leaf veins, 0.5-1.4 mm in diameter, circular to widely elliptic, sessile; juvenile leaves bilobed and variegated, the variegation seen in some clones at maturity; tendril 0.3-1.0 mm wide, present at flowering node, absent in inflorescence. Flowers borne in leaf axils or inflorescences; inflorescences 2.0-18.5(-25.1) cm long, associated reduced laminas 2.0-4.3 mm long, 1.5-3.1 mm wide. Pedicels 1.9-15.8 mm long, 0.4-0.9 mm wide, 2 per node; bract(s) absent; spur(s) absent. Flowers 18.4-33.4 mm in diameter with stipe 3.1-8.6(-9.4) mm long, 0.5-1.3 mm wide; hypanthium 5.4-8.2 mm in diameter; sepals 6.5-13.3 mm long, 2.9-6.3 mm wide, ovate-triangular, acute to rounded, abaxially and adaxially greenish yellow; coronal filaments in 2 series, the outer 40-51, 5.5-8.4 mm long, 0.3-0.7(-0.8)mm wide, linear, more or less erect, very dark reddish purple (5RP 3/2) on proximal third, greenish yellow (5GY 8/4) on middle third, yellow on distal third (5Y 8/10), ratio of outer coronal row to sepal length 0.59-0.94, the inner 27-40, 2.3-3.8 mm long, 0.2-0.5(-0.6) mm wide, linear, capitate, erect, greenish yellow with a flush of very dark reddish purple at base, ratio of inner coronal row to outer coronal row length 0.35-0.52; operculum 1.2-2.0 mm long, plicate, greenish yellow with a flush of very dark reddish purple at base, the margin white with narrow minutely fimbrillate teeth; nectary 0.1-0.5(-0.6) mm high, 0.5-1.2(-2.9) mm wide; limen not recurved but inclined toward the operculum, 0.1-0.5(-0.7) mm high, 0.1-0.4(-0.5) mm wide, very dark red (5R 2.5/2), limen floor 2.5-5.1 mm in diameter, very dark red; androgynophore (2.1-)3.5-6.0 mm long, 0.4-1.3(-1.5) mm wide, the distal half dark red then greenish yellow with dark red spots and streaks; free portions of the staminal filaments 1.5-2.8 mm long, 0.5-0.9 mm wide, linear, greenish yellow; anthers 1.9-2.9 mm long, 0.6-1.5(-1.9) mm wide; styles 1.8-4.9(-5.4) mm long including stigmas, 0.2-0.5 mm wide, greenish yellow; stigmas 0.8-1.7 mm in diameter; ovary 1.6-2.7 mm long, 1.1-2.1 mm wide, widely ellipsoid to globose, greenish yellow. Berry (12.4-)18-23.1 mm long, (13.4-)18.4-33.0 mm in diameter, globose, very dark purple. Seeds ca. 40-50, 1.8-4.8 mm long, 2.1-2.9 mm wide, 1.5-2.1 mm thick, obovate in outline, acute at both ends, reticulate-foveate with each face marked with ca. 12-17(-19) foveae. Germination epigeal.

Phenology.

Flowering and fruiting throughout the year.

Distribution.

Mexico and Central America (except Costa Rica and Panama). Growing in shrubs, trees or trailing on the ground in secondary successional areas, along the edges of semideciduous to deciduous, dry to wet tropical forests, both inland and near the seashore, 0-1171 m.

Ethnobotany.

The vine is sold in Guatemalan herb markets and is sold dried where the plant does not grow naturally ( Morton 1981). A decoction of the leaves is commonly taken as a diuretic, especially in the treatment of kidney infections ( Morton 1981). In El Salvador and Honduras the leaves are combined with lard and used as a poultice on wounds and swellings ( Morton 1981).

Discussion.

Passiflora sexocellata is very similar to Passiflora coriacea and Passiflora megacoriacea , and some of their similarities and differences are discussed under their respective descriptions. According to Jan Meerman (pers. comm.), Passiflora sexocellata and Passiflora xiikzodz grow side by side in Belize, with Passiflora sexocellata growing in the sun and Passiflora xiikzodz growing in the shade. Where these two species are found in the Yucatán Peninsula of Mexico, I found that Passiflora sexocellata occurs in wetter forests along rivers and lakes to the west and Passiflora xiikzodz and the related Passiflora itzensis are found in drier forests to the east. However, MacDougal concluded that these plants grow together at some sites in the Yucatán ( MacDougal 1992). These two species are easily separated because Passiflora xiikzodz and Passiflora itzensis possess petiolar nectaries at or near the apex of the petiole whereas Passiflora sexocellata has petiolar nectaries on the proximal half of the petiole. In addition, numerous floral characters can be used to distinguish between them. The most obvious difference is the number of coronal rows, with Passiflora xiikzodz and Passiflora itzensis possessing seven series and Passiflora sexocellata possessing only two.

Klucking (1992) classified the leaf venation of Passiflora sexocellata , identified as Passiflora coriacea , as actinodromous and pinnate secondary venation with irregular to regular intercostal venation consisting of lineate to transverse veins. The peltate, trilobed leaf illustrated is the typical form for Passiflora sexocellata . There are three primary veins and two acrodromal veins which extend two-thirds the length of the lateral lobes, the lateral lobes are acute, the angle between the lateral veins is 150°, and there are six laminar nectaries apparent on the abaxial surface ( Klucking 1992).

In Belize, Meerman (2001) found that Heliconius erato is an herbivore of Passiflora sexocellata (which he identified as Passiflora coriacea ). Benson et al. (1975) found that Dryas julia and Heliconius erato were herbivores of Passiflora sexocellata (again, identified as Passiflora coriacea ).

In 1990, Joanna Turner collected Plaster bees, Colletes sp., that regularly visited flowers of Passiflora sexocellata in Belize. The bees are approximately 10 mm long, 3-4 mm high, including some off-the-ground leg clearance, and have a thorax that is 2.0-2.5 mm high, and were identified by Rick Clinebell at MO (pers. comm.).

Passiflora sexocellata was originally described by Schlechtendal in 1854. He cited " Passiflora marmorea hort.", as a synonym, but this horticultural name was not validly published. It is interesting that the specific epithet “marmorea” means marbled, as the leaves of this species are often variegated. Holm-Nielson et al. (1988), in the synonomy of Passiflora coriacea , stated that Passiflora sexocellata is an illegitimate name that was based upon material of Passiflora coriacea Juss. and Passiflora difformis Kunth. However, I do not see any reason why Schlechtendal’s species has to be considered illegitimate. He carefully describes the plant from cultivated material that he had at hand in the Botanical Gardens in Halle, Germany and spends a paragraph differentiating his species from both Passiflora coriacea and Passiflora difformis . I was unable to locate the type of Passiflora sexocellata , and U. Braun (curator of the herbarium at the Herbarium at the Institut fur Geobotanik und Botanischer Garten, Halle) was unable to find any material under the name Passiflora sexocellata . Braun was also unable to locate appropriate material under Passiflora coriacea or Passiflora difformis . Other species of Passiflora from Mexico and Central America were in cultivation in Europe by 1830 ( Loudon 1830), and it is plausible that Schlechtendal had such material at hand when he described Passiflora sexocellata . Schlechtendal’s Passiflora sexocellata seems to fit the description of the Mesoamerican entity that I am recognizing as a species distinct from Passiflora coriacea and other similar taxa from supersection Cieca . However, some of the vegetative characters that he uses to distinguish Passiflora sexocellata are actually quite variable, but he only had one live specimen available to him when he described the species. He describes the flower as having five green sepals, outer coronal filaments that are “lilac” at the base but "greenish yellow" otherwise, inner coronal filaments that are dilated at the apex and “lilac” in color at the tips and lighter toward the base, an operculum that is dull “lilac” at the base and becoming "greenish yellow" toward the apex, and a "greenish yellow" androgynophore. The use of the term “lilac” is somewhat misleading, but the description of how the colors vary on the various parts of the flower is diagnostic. For example, Passiflora coriacea possesses outer coronal filaments that are reddish purple at the base but obviously white toward the tips with a band of reddish purple and not "greenish yellow." In addition, the operculum of Passiflora coriacea is wholly reddish purple. However, Passiflora sexocellata possesses outer coronal filaments that are reddish purple at their bases, greenish yellow at their middles and yellow at their apices and an operculum that is dark reddish purple at the base and greenish yellow otherwise (often with a white margin). Based upon Schlechtendal’s detailed description, I apply the name Passiflora sexocellata to this species and have designated a neotype that perfectly illustrates the diagnostic characters of the taxon, with the colors of the corona and limen floor still very vibrant.

Selected specimens examined.

MEXICO. Campeche: road between Ulumal and Canosayab, Porter-Utley & Mondragón 311 (CICY); road (MEX15) between El Estado de Mexico and Monclova, close to El Estado de Mexico, Porter-Utley & Mondragón 314 (FLAS, CICY); Champoton, Steere 1888 (US). Chiapas: Mpio. Ocosingo, el ejido Chajul a la orilla del Río Lacantun, 150 m, Martínez et al. 26047 (XAL). Oaxaca: 5 mi. E of Temascal (10 mi. W of Veracruz border), 45 ft., Janzen s.n., 13 November 1963 (UC). PUEBLA: Mpio. Tenampulco, Tenampulco, Chavez & Kerbel 327 (CICY). San Luis Potosi: Mpio. Ciudad Valles, ca. 1 km upstream from Rancho Pago Pago on Río Mesillas, 120 m, Fryxell & Anderson 3449 (CHAPA, MO). Tabasco: cerca de la parcela de Don Justo Hernández, Ejido Fernández Manero, km 12.1 del camino hacia cacaos de la desviación KM 32 de la carretera Villahermosa hacia Escarcega, Cowan 2815 (CAS, NY); San Isidro, near Balancan, Matuda 6045 (LL); road (MEX 180) between Minatitlan and Villahermosa, Porter-Utley & Mondragón 384 (CICY, FLAS). Veracruz: Mpio. Coatzacoalcos, Coatzacoalcos, entro las dos lenguas de la laguna del Ostion, Castillo-Campos & Acosta 16155 (XAL); Mpio. Tlacotalpan, along the hwy. following the Río Papaloapan towards the coast, 2 km NE of Tlacotalpan, 2 m, Nee & Taylor 26567 (F, MO); Playa Escondida, Porter-Utley & Mondragón 326 (CICY, FLAS); La Palmilla, Mpio. de Tlapacoyan, Ventura 1270 (CHAPA, MEXU); El Encanto, Mpio. de Tlapacoyan, Ventura 19595 (CAS, XAL).

BELIZE. Belize: along Belize River near Burrel Boom, near sea level, Gentry 8046 (MO); Caves Branch Base Camp, Whitefoord 1327 (BM). COROZAL: Alfonsoville, Gentle 821 (MO, NY, US). Orange Walk: Mi. 54, N Hwy., Dwyer & Liesner 12214 (MO). Stann Creek: Swasey Branch, Monkey River, Gentle 3931 (GH, NY).

EL SALVADOR. Cabañas: Ilobasco, Villacorta & Rivas 2117 (MO). La Libertad: Santa Tecla, Garcia 151 (UC). San Salvador: Cerro de San Jacinto, near San Salvador, 800-1171 m, Standley 20602 (GH, NY, US). San Vicente: vicinity of San Vicente, 400-500 m, Standley & Padilla 3444 (F). Sonsonate: vicinity of Izalco, 400-600 m, Pittier 1949 (US).

GUATEMALA. Alta Verapaz: 1 km N de finca Mercedes, Telemán, Panzós, 32 m, Martínez et al. 22859 (MEXU); Pantín, below Tamahú, 600 m, Standley 70882 (F). Chiquimula: Chocón Plantation, Watson s.n., 20 March 1885 (GH). Esquintla: San Luis, N of Escuintla, 450 m, Standley 60135 (F). Izabal: 27 km from junction of Atlantic Rt. with road to Tikal, McDade 210 (DUKE). Petén: SE part of Cerro Cauhui, Walker 1172 (MO). Retalhuleu: above Asintal, on road toward Colomba, 750-800 m, Standley 87879 (F). San Marcos: vicinity of Pajapita, 120 m, Molina & Molina 27104 (F). Santa Rosa: Region of La Morenita, NE of Chiquimulilla, 400 m, Standley 78869 (F). Suchitepéquez: Las Ánimas, 650 m, Shannon 274 (US). Unknown State: Tejada 248 (US).

HONDURAS. Atlántida: Valle Río Lean near El Mazapán N of Mezapa, 20 m, MacDougal et al. 3298 (BM, CHAPA, MO, TEFH). Comayagua: 1 km SW Palmitia, 840 m, Lentz 996 (TEFH); Pitosolo Yojoa, 500 m, Valerio & Rodriguez 2895 (F). Cortes: Mountains E of Lake Yojoa, 600-800 m, Morton 7760 (US). Gracias A Dios: Leymus, orilla del Río Segovia o Wanki, 100 km SO de Puerto Lempira, 30 m, Nelson & Cruz 8707 (TEFH). Santa Barbara: San Pedro Sula, 1200 m, Thieme 5242 (US).

NICARAGUA. Chontales: Cerro Oluma, 750 m, Gentry et al. 43989 (MO). Matagalpa: Carretera al Tuma 6 km NW de Cuatro Esquinas, 700-800 m, Guzman et al. 812 (MO). Region Autonomista Atlántico Norte: matorrales de la Playa S de Puerto Cabezas, 0 m, Molina 14759 (F).