Passiflora megacoriacea K. Porter-Utley., 2014

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 : 112-118

16. Passiflora megacoriacea K. Porter-Utley. sp. nov. Figs 50, 51

Diagnosis.

Passiflora scandens; stipulae 0.4-0.7 mm latae; petioli in parte proximali biglandulosi; folia peltata glandulosa bilobata vel obscure ad distincte trilobata, lobis centralibus obtusis ad acutis, lobis lateralibus acutis vel raro acuminatis, marginibus integris; pedunculi ebracteatis vel raro unibracteatis, stipites florum 1.7-5.7 mm longi; petala nulla; sepala 10.0-20.5 mm longa, viridiflava; filamenta coronae 2-seriata, filamentis exterioribus linearibus, 6.8-14.0 mm longis, pro parte maxima viridiflavis, ad apicem flavidis, interdum ad basim purpureis, filamentis interioribus capitatis, 2.3-5.6 mm longis, pro parte maxima viridiflavis, ad apicem flavidis, interdum ad basim purpureis; operculum plicatum; androgynophorum 4.1-10.0 mm longum; ovarium glabrum; fructus ellipsoidei; semina 4.9-5.1 mm longi, 3.0-3.1 mm lata, retifoveata.

Type: Costa Rica. Limón: bluff above mouth of river at Moín, about 7 km N of Limon, sunny clay bank along road, 9 Aug. 1980, J. M. MacDougal 1204 (holotype: DUKE! [DUKE00274532]; isotypes: C!, CAS! [CAS00767084]).

Description.

Slender, climbing, perennial vine 2-4 m long or more, sometimes trailing on ground, sparsely pubescent with unicellular curved trichomes on petiole, leaf and stem, 0.2-0.4 mm long, 0.02-0.03 mm wide, also minutely antrorsely appressed-puberulent throughout with unicellular, curved trichomes, ca. 0.1 mm long, 0.02-0.03 mm wide. Flowering stems 1.0-2.4 mm in diameter, terete or somewhat compressed, with the base woody and cork-covered. Stipules 1.7-5.7 mm long, 0.4-0.7 mm wide, narrowly ovate-triangular, acute, longitudinally striate-nerved; petioles 1.1-3.8 cm long, inserted 1.4-9.6 mm from the basal margins of the peltate blades, with two, round or elliptic, opposite to subopposite, sessile or shortly stipitate (rare), saucer-shaped nectaries with flat rims, 1.3-2.1 mm wide (on the widest axis), 0.4-1.6 mm high, commonly borne in the distal half of the petiole (0.30-)0.53-0.77 of the distance from the base toward the apex of the petiole. Laminas (2.7-)3.3-7.1(-8.2) cm long, 6.6-17.3 cm wide, sometimes glaucous beneath, coriaceous, peltate, transversely elliptic (widely divaricately bilobed) or 3-lobed, lateral lobes 3.5-9.1 cm long, 3.7-25 cm wide, elliptic, acute to slightly attenuate, central lobe elliptic to ovate or present merely as an acute to obtuse tip, central vein 2.4-7.7 cm long (measured from point of petiole insertion), angle between the lateral lobes 104-176°, ratio of lateral lobe to central vein length 0.85-2.47, margins entire, hyaline, primary veins 3, diverging and branching above base, laminar nectaries present, 6-10, submarginal, associated with the minor veins of the abaxial surface, 0.3-1.5 mm in diameter, circular to widely elliptic, sessile; tendril 0.3-1.1 mm wide, present at flowering node, absent in inflorescence. Flowers borne in leaf axils or inflorescences; inflorescences 5.6-11.7 cm long, associated reduced laminas 2.6-9.0 mm long, 1.0-2.8 mm wide. Pedicels 4.4-17.5 mm long, 0.4-1.1 mm wide, 2 per node; bract(s) 1 (rare) or absent; spur(s) absent. Flowers 29.5-56.7 mm in diameter with stipe 2.6-6.1 mm long, 0.9-1.4 mm wide; hypanthium (7.8-)8.1-16.1 mm in diameter; sepals 10.0-20.5 mm long, 4.3-12.1 mm wide, ovate-triangular, acute to rounded, abaxially and adaxially very pale greenish yellow; coronal filaments in 2 series, the outer 31-40, 6.8-14.0 mm long, 0.4-1.1 mm wide, linear, dilated toward tips, erect, greenish yellow (5GY 8/4) with very light yellow tips (5Y 8/6), ratio of outer coronal row to sepal length 0.48-0.85, the inner (12-)30-45, 2.3-5.6 mm long, 0.1-0.4 mm wide, linear, capitate, erect, greenish yellow with whitish apices or greenish yellow with a mere flush of reddish purple (5RP 4/8-4/10) at the very base and whitish tips, ratio of inner coronal row to outer coronal row length 0.30-0.52; operculum 2.1-4.2 mm long, plicate, flushed with reddish purple toward the base and whitish toward the tips, the margin with narrow minutely fimbrillate teeth; nectary 0.4-0.8(-2.3) mm high, 0.8-2.5 mm wide; limen recurved or rarely inclined slightly away from androgynophore, 0.4-1.3 mm high, 0.2-0.9 mm wide, whitish, limen floor 3.3-8.4 mm in diameter, whitish with reddish purple spots and streaks toward base; androgynophore 4.1-10.0 mm long, 0.9-1.9 mm wide, whitish at base with reddish purple spots and streaks becoming light greenish yellow toward apex; free portions of the staminal filaments 2.8-3.9 mm long, 0.5-1.3 mm wide, linear, greenish yellow; anthers 2.8-4.4 mm long, 0.8-2.6(-5.1) mm wide; styles 3.2-6.7 mm long including stigmas, 0.3-0.7 mm wide, greenish yellow; stigmas 1.2-2.5 mm in diameter; ovary 2.1-4.0 mm long, 1.4-3.6 mm wide, widely ellipsoid to globose, greenish yellow. Berry 24.0-27.0 mm long, 19.0-25.0 mm in diameter, ellipsoid, very dark purple (5P 2.5/2) with a glaucous bloom at maturity, immature fruit greenish yellow, sometimes mottled with white or yellow. Seeds (27-)45-50(-60), 4.9-5.1 mm long, 3.0-3.1 mm wide, 1.9-2.0 mm thick, obovate in outline, acute at both ends, reticulate-foveate with each face marked with ca. 15-20 foveae.

Phenology.

Flowering and fruiting throughout the year.

Distribution.

Colombia, Costa Rica, and Panama. Growing in shrubs or trees in secondary successional areas, along the edges of tropical moist to premontane wet forests, and near the seashore, 0-1100 m altitude.

Discussion.

Passiflora megacoriacea is relatively common in Costa Rica and Panama. John MacDougal brought my attention to the variation of vegetative and floral characters of some of the Costa Rican and Panamanian specimens then identified as Passiflora coriacea .

Passiflora megacoriacea , as noted above in the discussion of Passiflora coriacea , is very similar to Passiflora coriacea and Passiflora sexocellata , and although not sympatric, without reproductive material it can be difficult to separate them. Passiflora megacoriacea may be recognized by commonly having petiolar nectaries found on the distal half of the petiole, (0.30-) 0.50-0.77 of the distance from the base toward the apex of the petiole, and although that overlaps the 0.21-0.54(-0.64) range of Passiflora coriacea and Passiflora sexocellata , the character is easily seen in herbarium specimens. Passiflora megacoriacea can also possess deeply trilobed leaves (commonly 0.11-0.61 the distance to the base), especially in populations along the Pacific coast of Costa Rica and in the Panamá Canal Zone, whereas Passiflora coriacea and Passiflora sexocellata do not possess deeply trilobed leaves (commonly less than 0.11 the distance to the base). The reproductive structures of these three species provide a number of distinguishing characters. Passiflora megacoriacea possesses floral stipes that are commonly less than half the length of the pedicels, whereas Passiflora coriacea possess stipes that are usually two to three times the length of the pedicels and Passiflora sexocellata has floral stipes that are commonly just shorter than or rarely up to two times the length of the pedicels. The overall size of the flower of Passiflora megacoriacea exceeds that of both Passiflora coriacea or Passiflora sexocellata , with Passiflora megacoriacea commonly having a wider hypanthium, longer sepals, larger and fewer outer coronal filaments, a longer androgynophore, longer staminal filaments, longer anthers, and a longer operculum. The most informative of these is the length of the androgynophore, with Passiflora megacoriacea having an androgynophore that is 6.9-8.8 mm long and the androgynophores of both Passiflora coriacea and Passiflora sexocellata not exceeding a length of 5.9 mm. In addition, the nectary floor is raised in Passiflora megacoriacea , never raised in Passiflora sexocellata , and only rarely raised in Passiflora coriacea . The outer coronal filaments of both Passiflora megacoriacea and Passiflora sexocellata are erect, while those of Passiflora coriacea spread to ca. 140-160°. The flowers of Passiflora megacoriacea are commonly referred to as white, greenish white, or cream on herbarium labels and this is due to it having no (or relatively little) reddish purple coloration in the mature flowers; the flowers of Passiflora coriacea and Passiflora sexocellata both commonly have a significant amount of reddish purple coloration.

According to Benson et al. (1975), Passiflora megacoriacea (based on geography) has a different passionflower butterfly herbivore than Passiflora coriacea and Passiflora sexocellata . Heliconius cydno has been reported to be the primary herbivore of Passiflora megacoriacea in Panamá and southeastern Costa Rica, though Heliconius erato is also known to utilize this species. Heliconius erato is the primary herbivore of Passiflora coriacea and Passiflora sexocellata . Dryas julia is also an herbivore of Passiflora sexocellata ( Benson et al. 1975).

In an unpublished manuscript, MacDougal determined the total sugar concentration measured as sucrose equivalents in percent weight per total weight to be 29-44% in Passiflora megacoriacea (MacDougal 409). He found the flower odor to be sweet, waxy, and strong. These data indicate that the flowers are likely utilized by bees.

Specimens examined (paratypes).

COSTA RICA. Cartago: Pasture beside Río Pejibaye, 2 km SW of Taus, 750 m, Lent 2960 (F); Las Vueltas (de Tucurrique), 635 m, Tonduz 12808 (US). Guanacaste: Parque Nacional Guanacaste Estación Biología Volcán Cacao, 1100 m, Alvarado 28 (CR, MO); Parque Nacional Rincón de la Vieja Liberia, Cordillera de Guanacaste, Estación Las Pailas, 800 m, Espinoza 708 (CR, MO). Herédia: Los pastizales de la Finca de Napoleon Murillo, Chacon 778 (DUKE); Finca La Selva, the OTS Field Station on the Río Puerto Viejo, just E of its junction with the Río Sarapiquí, 100 m, Grayum 2782 (DUKE); N base of hills to the S of the Río Sarapiquí, opposite Chilamate, 60-100 m, Grayum et al. 5316 (MO). Limón: Between Siquerres and the Río Pacuare, and remnant forest on steep hills S of the railroad bridge over Río Pacuare, 50-100 m, Burger & Liesner 6868 (F, MO); Canton de Siquirres, llanura de Santa Clara, puente sobre Río Barbilla, 50 m, Chavarria & Solis 955 (MO); Talamanca, Sixaola, en la fila entre Gandoca & Manzanillo frente a Punta Mona, 50-100 m, Herrera & Bloemen 7632 (F, MO, US); along beach between Port Limón and Moin, Pittier 3630 (BM, US); Parque Puerto, Vargas, Poveda & de Ramury 3270 (CR, F). Puntarenas: Carara National Park, near Río Carara, near guard post, 120 m, Gentry et al. 79273 (CR, MO); Canton de Buenos Aires, cañon del Río Grande de Terraba, cerca del Proyecto Boruca-ICE, 100 m, Hammel et al. 17870 (CR, MO); Parque Nacional Corcovado, Sirena, Río Claro Trail-Río Claro, 0-150 m, Kernan 131 (MO); Canton de Osa, R.B. Isla del Caño, Península de Osa, 1 m, Lepiz 462 (MO); Burica Península, unnamed quebrada opposite Quebrada Macho of Panamá, 11 mi. S of Puerto Armuelles, 20-200 m, Liesner 184 (MO); Reserva Biología Carara, 200 m, Morales 1267 (MO); Bords du Río Platanar, Hacum, pres Buenos Aires, 250 m, H. Pittier 6584 (MO); Canton de Buenos Aires Reserva Indígena Boruca, 200 m, Rojas & Zuniga 158 (CR, MO).

PANAMÁ. Canal Zone (currently separated into the provinces of Colón and Panama): Shoreline of E side of Peña Blanca Point across from front no. 8 light, Barro Colorado Island, Croat 6732 (MO); Río Majé, along river from waterfalls near Bayano Lake to Finca of Chocó Indian Eduardo Maycha, ca. 2 mi. upstream, 30-60 m, Croat 34557 (MO); vicinity of Panamá Railroad crossing at Guillard Hwy., across road from former Summit Hills golf course, Croat & Zhu 76290 (MO); between Chilibre & Madden Dam on Transisthmian Hwy., Dwyer & Correa 9397 (MO); Forest preserve, near Green Park, Folsom 228 (MO); Barro Colorado Island, SE of Gross Point, Foster 2285 (DUKE); Barro Colorado Island, tower clearing, Foster 769 (DUKE); Gaillard hwy., mi. 12-13, Garwood 1861A (F); Gatún Locks, Gilbert 409 (FLAS); on brush along railroad, Summit Gardens, Hammel 1787 (MO); ca. de Represa Madden, Campo de Exploradores, Kant 21 (DUKE); junction of Chiva-Chiva and Gaillard Hwy., 50 m, Knapp & Schmalzel 4870 (MO); Pipeline Road, ca. 5 km from beginning, just NE of crossing of Río Siristes, 128 m, 9°10N, 79°45 W, MacDougal et al. 6315 (MO); Around Alahajuela, Chagres Valley, 30-100 m, Pittier 3456 (US); Boy Scout Road, Madden Dam area, Porter et al. 4014 (MO, UC); Las Cascadas Plantation, near Summit, Standley 29594 (US); Darien Station, Standley 31617 (US); near Survival School, Curundú, Tyson 1054 (MO); Boy Scout Camp on Madden Lake, Tyson 5454 (MO); Fort Clayton, no. 519, the old hospital building, Tyson & Blum 3901 (MO, US); Shore N of end of Chapman Trail, Woodworth & Vestal 501 (A, F, MO); Río Vigue Beach, Zetek 5564 (MO). Chiriquí: near San Juan, Seemann, s. n., 1844 (K). Coclé: N rim of El Valle de Antón, 600-1000 m, Allen 1667 (MO); Forest behind Club Campestre, 700 m, Duke 13270 (MO); Behind Hotel Turístico, El Valle, 2200 ft., Hammel 1778 (MO); NE of El Valle de Antón, 2000 ft., Lewis et al. 1703 (MO); 2.4 km (air) N of the church at El Valle, 725 m, 08°37 N, 80°08 W, MacDougal & Lezcano 6274 (MO); Above and N of El Copé, road to the old saw mill that used to be called “Whiskey” near the continental divide, now a national park reserve, 484 m, 08°39 N, 80°35 W, MacDougal et al. 6299 (MO). Darién: 3 km S of Jaqué, 0-100 ft., D’Arcy & Sytsma 14553 (MO); Hill ca. 1 mi. NE of Nura, 200 m, Duke 10084 (3) (ECON, MO). Panamá: 1 km E of Chorrera City limits, Folsom 3466 (MO); SE slope of Cerro Campana, Lewis et al. 3130 (MO). San Blas: on mainland in front of Ustupo, D’Arcy 9527 (MO).

COLOMBIA. Bolívar: Torrecilla, near Turbaco, 150-300 m, Killip & Smith 14415 (GH, US).