Passiflora coriacea Juss., Ann. Mus. Natl. Hist. 6: 109. pl. 39, fig. 2. 1805.

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 : 109-112

15. Passiflora coriacea Juss., Ann. Mus. Natl. Hist. 6: 109. pl. 39, fig. 2. 1805. Figs 48, 49

Monactineirma coriacea (Juss.) Bory, Ann. Gén. Sci. Phys. 2: 138. 1819. Type: Based on Passiflora coriacea Juss.

Cieca coriacea (Juss.) M.Roemer, Prospect Fam. Nat. Syn. Monogr. 2: 148. 1846. Type: Based on Passiflora coriacea Juss.

Passiflora clypeata Sm., Cycl. [A. Rees] (London ed.) 26: Passiflora no. 20. 1814. Type: Colombia. Sin. loc., J. Mutis s.n. (lectotype, designated by Killip 1938, pg. 85: LINN 1070.16 [microfiche seen]).

Passiflora difformis Kunth in Humboldt, Bonpland and Kunth. Nov. Gen. Sp. 2: 136. 1817. Type: Colombia. Quindio: "in monte Quindiu juxta El Moral, alt. 1065 hex", A. Humboldt & A. Bonpland s.n. (holotype: P [P00307399, photograph seen], photograph AAU!, isotype: B, destroyed, P [P00307391, photograph seen], photograph AAU!).

Cieca difformis (Kunth) M.Roem., Prospect Fam. Nat. Syn. Monogr. 2: 140. 1846. Type: Based on Passiflora difformis Kunth

Passiflora cheiroptera Cortés, Fl. Colomb. ed. 2, fig between pages 112 and 113. 1919. Type: Colombia, (lectotype, designated here: Cortés, Fl. Colomb. ed. 2, fig between pages 112 and 113. 1919).

Type.

Colombia. Tolima: Santa Fé, near Honda, A. Humboldt & A. Bonpland s.n. (lectotype, designated here: P! [P00307401], photographs AAU!, DUKE!, isolectotype: P! [P00307391], photograph AAU!).

Description. Slender, climbing, perennial vine 2-8 m long or more, sparsely pubescent with unicellular curved trichomes on petiole, leaf, stem and stipule, 0.20-0.64 mm long, 0.02-0.03 mm wide, also minutely antrorsely appressed-puberulent throughout with unicellular, curved trichomes, 0.03-0.10 mm long, 0.02-0.03 mm wide. Flowering stems 1.0-2.9 mm in diameter, greenish yellow (5GY 8/4) to reddish purple (5RP 4/6), terete to somewhat compressed, with the base woody and cork-covered. Stipules 2.6-7.5 mm long, 0.4-1.0 mm wide, narrowly ovate-triangular, acute; petioles 1.1-4.3 cm long, with 2 (rarely 3), opposite to subopposite, sessile, discoid nectaries with flat rims, 1.1-2.1 mm wide (on the widest axis), 0.1-1.5 mm high, borne in the proximal two thirds of the petiole (0.21-0.64 of the distance from the base toward the apex of the petiole). Laminas 2.8-5.9 cm long, 6.2-18.8 cm wide, coriaceous, peltate (the distance from leaf base to point of petiole insertion 1.6-11.4 mm), transversely elliptic (widely divaricately bilobed) or sometimes 3-lobed, lateral lobes 3.3-9.5 cm long, 1.8-7.3 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 1.8-6.4 cm long (measured from point of petiole insertion to the leaf apex), angle between the lateral lobes (97-)110-160(-170)°, ratio of lateral lobe to central vein length 1.36-2.61, margins entire, hyaline, primary veins 3, diverging and branching above base, laminar nectaries present, 5-13, submarginal, associated with the minor veins of the abaxial surface, with 2-4 nectaries proximal to the lateral leaf veins, 0.7-1.3 mm in diameter, circular to widely elliptic, sessile; juvenile leaves bilobed and variegated; tendril 0.3-0.9 mm wide, present at flowering node, absent in inflorescence. Flowers borne in leaf axils or inflorescences; inflorescences 2.5-6.5(-12.0) cm long, associated reduced laminas 2.5-5.0 mm long, 1.5-2.8 mm wide. Pedicels 2.2-8.1 mm long, 0.4-1.1 mm wide, 2 per node; bract(s) absent; spur(s) absent. Flowers 18.0-30.0 mm in diameter with stipe 6.3-15.1 mm long, 0.7-1.0 mm wide; hypanthium 4.9-7.4(-8.1) mm in diameter; sepals 5.8-10.9 mm long, 3.3-6.4 mm wide, ovate-triangular, acute to rounded, abaxially and adaxially greenish yellow (5GY 8/4); coronal filaments in 2 series, the outer (36-)49-53, 3.1-5.3(-7.0) mm long, 0.2-0.5 mm wide, linear, spreading, dark reddish purple at base (5RP 3/6-4/6), medium reddish purple just below the middle (5RP 4/4-5/4), light reddish purple (5RP 6/6-6/8) just above middle and white on the distal third, ratio of outer coronal row to sepal length 0.43-0.76, the inner 33-50, 1.4-3.2 mm long, 0.2-0.5(-0.7) mm wide, linear, capitate, erect, dark reddish purple (5RP 3/6), lightening slightly towards tips, ratio of inner coronal row to outer coronal row length (0.29-)0.44-0.63(-0.72); operculum 1.3-2.0(-4.3) mm long, plicate, reddish purple (5RP 3/6-4/6), the margin with narrow minutely fimbrillate teeth; nectary 0.2-0.5(-1.5) mm high, 0.7-1.1 mm wide; limen recurved or sometimes erect, 0.2-0.5(-0.7) mm high, 0.1-0.4 mm wide, reddish purple (5RP 3/6-4/6), limen floor 2.0-3.5(-4.7) mm in diameter, pale greenish yellow with reddish purple (5RP 3/6-4/6) spots and streaks; androgynophore (3.3-)3.8-5.0 mm long, 1.0-1.5 mm wide, pale greenish yellow (5GY 8/2) with reddish purple (5RP 3/6-4/6) spots and streaks; free portions of the staminal filaments 2.4-3.2 mm long, 0.5-1.1 mm wide, linear, greenish yellow; anthers 1.9-3.9 mm long, (0.6-)0.9-2.3 mm wide; styles 3.2-4.6 mm long including stigmas, 0.2-0.5 mm wide, greenish yellow; stigmas 0.6-2.2 mm in diameter; ovary 1.7-2.9 mm long, 1.5-2.5(-4.0) mm wide, widely ellipsoid to globose, greenish yellow. Berry 17.1-21.0 mm long, 12.0-19 mm in diameter, globose, very dark purple (5P 2.5/2). Seeds ca. 44-61, 3.6-4.0(-5.0) mm long, 2.1-2.5 mm wide, 1.5-1.8 mm thick, obovate in outline, acute at both ends, reticulate-foveate with each side marked with ca. 15-17 foveae.

Phenology.

Flowering and fruiting throughout the year.

Distribution.

Bolivia, Colombia, Ecuador, Perú, and Venezuela; reported once from Guyana (Lejos 43, B, destroyed). Growing in shrubs and small trees in secondary successional areas, along the edges of moist tropical forests near rivers and streams, and along the seashore, 0-1500 m.

Ethnobotany.

Timothy Plowman in a note on a specimen collected by him in 1976 (T. Plowman 6029), noted that in Perú a medicine for the liver is prepared from Passiflora coriacea by boiling the whole plant and then drinking the syrup.

Discussion.

Passiflora coriacea is extremely similar to Passiflora sexocellata and Passiflora megacoriacea in its vegetative morphology, but is easily distinguished by its flowers. The flowers of Passiflora coriacea possess long floral stipes as compared to their pedicels (the stipes are usually two to three times the length of the pedicels) and an operculum that is dark reddish purple. Passiflora sexocellata has floral stipes that are commonly shorter than or equal in length to the pedicels and an operculum that is greenish yellow with a flush of dark reddish purple at the base and a white margin. Passiflora megacoriacea possesses floral stipes that are commonly less than half the length of the pedicels and an operculum that is greenish yellow with a white margin or greenish yellow with a mere flush of reddish purple at the base and a white margin. Passiflora coriacea is also distinguished by outer coronal filaments that may appear banded with light to dark reddish purple. In addition, the outer coronal filaments are more dilated distally, much like Passiflora megacoriacea but in contrast to Passiflora sexocellata . The limen floor in Passiflora coriacea is very light greenish yellow with dark reddish purple spots and streaks, again much like Passiflora megacoriacea . Passiflora sexocellata usually possesses a very dark red limen floor.

Schlechtendal (1854) attempted to use mostly vegetative characters to distinguish Passiflora coriacea from Passiflora sexocellata ; incidentally, he was the first to notice differences in the stipe and pedicel lengths of the two species. He used the following characters to differentiate them: position of the petiolar nectaries, the number of laminar nectaries, the shape of the stem, leaf venation, the leaf margin, and the leaf texture. However, in my analysis of these species, I did not find any of these vegetative characters to be wholly reliable in distinguishing between these two species. Both have petiolar nectaries that occur in various positions below the middle of the petiole, stems that are terete to somewhat compressed, five distinct leaf veins, thick leaf margins and coriaceous leaves. Passiflora sexocellata does tend to have fewer nectaries than Passiflora coriacea on average, but there is a significant amount of overlap in the range of variation.

A clone of Passiflora coriacea (MacDougal 3029) did not produce fruits by autogamy in years in cultivation. This greenhouse accession was given to me by MacDougal, who originally received it as seedlings from J. Zarucchi (Zarucchi et al. 6102).

Heliconius erato ( Lepidoptera : Nymphalidae , Heliconiinae) has been reported to be an herbivore of Passiflora coriacea in the central Colombian valleys (Cauca and Magdalena) ( Benson et al. 1975).

Fajardo et al. (1998) in a study on the genetic variation analysis of the genus Passiflora using RAPD markers, used Passiflora coriacea and Passiflora adenopoda DC. as representatives of taxa from subgenus Decaloba . They found Passiflora coriacea to be genetically distant from the other taxa in his study, including Passiflora adenopoda , but due to insufficient data, they were not able to discuss the significance of this result ( Fajardo et al. 1998).

In Antoine Laurent de Jussieu’s original description of Passiflora coriacea (1805) he included a detailed diagnosis and drawing of the species. The lectotype of Passiflora coriacea (at P), closely resembles the drawing in Jussieu, but there are no locality data on the specimen. The isolectotype of Passiflora coriacea consists of two leaves and a small portion of the stem and does not resemble the type drawing of the species, but written on the specimen are locality and descriptive data in Jussieu’s hand.

Selected specimens.

BOLIVIA. La Paz: Prov. Alto Beni, Chaco, cerca de Santa Ana de los Mozetenes, 450 m, Seidel & Schulte 2525 (TEX).

COLOMBIA. Antioquia: Mpio. Salgar, along road to Salgar, 4 km from Bolombolo, Bolívar Road, 900 m, Zarucchi et al. 6102 (HUA, MO). Bolívar: vicinity of Cartagena, Heriberto 392 (US). Caldas: Entre Aranca & Manizales 35 km de Manizales, 1500 m, Escobar & Uribe 483 (HUA, LL). Chocó: Mpio. Ríosucio, Parque Nal. Nat. Los Catios, camino Tilupo Peye, Quebrada Peye, 40 m, Forero et al. 1770 (MO). Cundinamarca: Población de Nariño, bosque donde finaliza la carretera de los Mangos, 350-450 m, Fernández et al. 5480 (MA). Huila: about 5 km N of Villavieja; upper basin of Río Magdalena, 500 m, Mason 13808 (UC). Magdalena: about 8 km N of Codazzi, 250 m, Haught 3752 (S, US); Flanco N de la Sierra Nevada de Santa Marta, Romero 761 (US). Santander: River Suratá valley, between Bucaramanga and El Jaboncillo, 800-1500 m, Killip & Smith 19062 (GH, US). Tolima: Río Cuello, New Quindío Trial, Cordillera Central, 1000-1500 m, Hazen 9652 (GH, US). Valle: Mpio. Tuluá, Corr. Mateguada, Jardín Botánico, 1100 m, Escobar 1045 (HUA); Mpio. Yotoco, ingenio La Carmelita, sección San Martín, zona A, dentro de un guadual que esta en medio de la cana, mas o menos una hora de Mediacanoa, 950 m, Ramos et al. 2811 (MO).

ECUADOR. Esmeraldas: El Timbre, towards Esmeraldas, Asplund 16532 (AAU, S).

PERÚ. Cuzco: La Convencion, N bank of Río Alto Urubamba, across from village of Kiteni, 500 m, Knapp & Mallet 6356 (US). Madre De Dios: Prov. Tambopata, Cuzco Amazónico, across Río Madre de Dios on road to Lago Sandoval, 200 m, Gentry & Curso de Posgrado de la Universidad de San Marcos 68962 (F, MO). San Martín: Prov. Mariscal Caceres, Dtto. Tocache Nuevo, Schunke 3823 (F, GH, MO, US).

VENEZUELA. Distrito Federal. Cerro Naiguatá, slopes near the sea to the N, above the town of Naiguatá, Lomas de Las Delicias, between Quebrada Basenilla and Quebrada Guayoyo, 9-12 km SE of Hacienda Cocuizal, 1000-1300 m, Steyermark 92078 (US). Falcón: Parque Nacional Quebrada de la Cueva El Toro, trail going to La Piedra, 600-900 m, Liesner et al. 7831 (MO, VEN). Lara. E border near state of Yaracuy, Guaremal River, NE of Barquisimeto, Meijer & Smith 56 (MO). Yaracuy. Dist. Urachiche, Quebrada Higueronal, afluente del Río Urachiche, W de Urachiche, cerca de la Sabana de Mendez, 50 m, Steyermark et al. 124671 (VEN). Zulia. Dist. Mara, alrededores del Puesto "El Bosque" de la Guardia Nacional, 1450-1600 m, Bunting et al. 12264 (MO).

CULTIVATED MATERIAL. United States. Missouri, cultivated at the Missouri Botanical Garden, from material collected by J.L. Zarucchi (Zarucchi 6102) in Antioquia, Colombia, MacDougal 3029 (FLAS, MO); Florida, cultivated at the University of Florida from material collected by J.L. Zarucci (Zarucchi 6102) in Antioquia, Colombia, Porter-Utley P-66 (FLAS).