Passiflora eglandulosa J.M. MacDougal. Annals of the Missouri Botanical Garden 75: 1658-1662. figs 1, 2B, and 3. 1988.

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 : 88-90

7. Passiflora eglandulosa J.M. MacDougal. Annals of the Missouri Botanical Garden 75: 1658-1662. figs 1, 2B, and 3. 1988. Figs 36 -37

Type.

Guatemala. San Marcos: wet mountain forest at Aldea Fraternidad, W-facing slope of Sierra Madre between San Rafael Pie de La Cuesta and Palo Gordo (ca. 14°56'N, 91°52'W), 1800-2400 m, 10-18 Dec.1963, L. O. Williams, A. Molina & T. P. Williams 25997 (holotype: F [F0066764F, photograph seen]; isotypes: EAP [EAP110288. photograph seen], ENCB, C, G! [G00440998], NY! [NY00110403], S! [S04-205], US! [US00588642], W).

Description.

Slender, climbing, perennial vine 2-8 m long, sparsely to lightly pubescent with unicellular curved trichomes on petiole, stem, and stipule, (0.1)0.4-0.6(-0.8) mm long, 0.02 mm wide, also minutely antrorsely appressed-puberulent throughout (except ovary) with unicellular, curved trichomes, 0.05-0.10 mm long, 0.02-0.03 mm wide. Flowering stems 0.6-2.1 mm in diameter, terete or subterete, with little secondary growth (to 6 mm near base with corky, secondary growth). Stipules (3.5-)5.3-12.6(-20.0) mm long, 2.50-6.4(-9.0) mm wide, ovate, slightly oblique, acute to slightly attenuate, 5-9 veins departing from the base; petioles 0.7-4.6 cm long, eglandular. Laminas 2.5-12.0 cm long, 2.2-14.5(-17.0) cm wide, chartaceous, not variegated, ratio of leaf width to central vein length 0.28-1.88, 3-lobed 0.26-0.45 of the distance to the cordate leaf base, lateral lobes 1.6-8.7 cm long, 0.8-4.4 cm wide, ovate-triangular, acute to slightly attenuate, central lobes 2.5-11.1 cm long, 1.0-5.7 cm wide, ovate-triangular, acute to slightly attenuate, angle between the lateral lobes 127-170°, ratio of lateral to central lobe length 0.64-0.97, margins entire, primary veins 3, diverging and branching at base, laminar nectaries absent; tendril 0.3-1.1 mm wide, present at flowering node, absent in inflorescence. Flowers borne in leaf axils. Pedicels 5.6-20.0 mm long, 0.5-0.9 mm wide, (1-)2 per node; bract(s) absent; spur(s) absent (occasionally) or 5 retrorse spurs present between the bases of the sepals, 0.7-1.1 mm long. Flowers 16.9-21.5 mm in diameter with stipe 2.1-7.9 mm long, 0.5-0.9 mm wide; hypanthium 4.0-5.9 mm in diameter; sepals 5.5-8.5 mm long, 2.3-3.9 mm wide, ovate-triangular, acute to rounded, the 2-3 outermost with a (0.5-)0.8-1.2 mm blunt subapical horn, abaxially and adaxially greenish yellow, often with a flush of reddish purple (5PR 3/4-4/6) abaxially (rarely to fully dark reddish purple); coronal filaments in 2 series, the outer 24-31, 2.0-4.1 mm long, 0.1-0.3 mm wide, linear, reflexed above middle and the tips often slightly incurved, greenish yellow at base, yellow distally, ratio of outer coronal row to sepal length 0.30-0.70, the inner 18-34, 0.7-1.5 mm long, 0.1-0.2 mm wide, linear, often capitate, erect, greenish yellow, ratio of inner coronal row to outer coronal row length 0.28-0.66; operculum 1.4-2.9 mm long, plicate, greenish yellow, sometimes with a flush of reddish purple at center, whitish distally, the margin with narrow minutely fimbrillate teeth; nectary 0.06-0.88 mm high, 0.6-1.5 mm wide; limen recurved, 0.2-0.5 mm high, 0.2-0.3 mm wide, whitish, limen floor 1.6-2.1 mm in diameter, whitish; androgynophore 1.3-3.5 mm long, 0.8-1.3 mm wide; free portions of the staminal filaments 2.1-3.8 mm long, 0.3-0.5 mm wide, linear, greenish yellow; anthers 2.3-3.8 mm long, 0.5-1.7 mm wide, greenish yellow, long axis oriented perpendicular (or nearly so) to long axis of filaments at anthesis; styles 3.5-6.7 mm long including stigmas, 0.2-0.4 mm wide, greenish yellow; stigmas 0.5-0.9 mm in diameter; ovary 1.2-2.8 mm long, 0.8-2.2 mm wide, widely ellipsoid to globose, greenish yellow. Berry 8.0-14.4 mm long, (7-)9.0-15.3 mm in diameter, widely ellipsoid to globose, very dark purple with glaucous bloom. Seeds 4-10, 4.5-5.7 mm long, 3.1-3.5 mm wide, 2.0-2.7 mm thick, obovate in outline, acute at both ends, reticulate-foveate with each face marked with with ca. 15-19 foveae.

Phenology.

Flowering and fruiting January-May, July-September and December.

Distribution.

El Salvador, Guatemala, and Honduras. Growing in shrubs and small trees in shady ravines and at the edges of premontane to montane broad-leaved forests on volcanic cones; 1500-2650 m.

Discussion.

For many years after the publication of Killip’s 1938 monograph, the name Passiflora trinifolia Mast. was applied to two distinct taxa: Passiflora eglandulosa and Passiflora trinifolia . In fact, Standley and Williams (1961), in their description of Passiflora trinifolia , combined information from Killip’s description of Passiflora trinifolia , which strictly applied to Passiflora trinifolia in the sense of Masters, and their own personal observations of Passiflora eglandulosa ( MacDougal 1988). It is true that the two species both possess wide foliose stipules, similarly trilobed leaves at fertile nodes and seeds with the micropylar end and chalazal beak erect and not inclined toward the raphe. However, Passiflora eglandulosa is distinguished by flowers with longer flower pedicels, spurs that occur between each of the sepals, narrower sepals, narrower outer coronal filaments, shorter inner coronal filaments that are not broadly capitate, narrow limen floors, short staminal filaments, and anthers that present pollen laterally as opposed to subproximally. The seeds are longer and wider than those of Passiflora trinifolia , and as its name implies, Passiflora eglandulosa lacks both laminar and petiolar nectaries; petiolar nectaries have been seen on only one specimen, M. Veliz 16059. Passiflora eglandulosa possesses flower buds that are slightly horned at the apex and flowers that are oriented above rather than near or below the horizontal plane. In comparing the habitats of the two species, MacDougal found that Passiflora eglandulosa is found in shady ravines and at the edges of wet premontane to montane broad-leaved forests on volcanic cones, whereas Passiflora trinifolia is found in open, seasonally dry pine/oak forests on rock outcrops. In addition, Passiflora eglandulosa is a larger plant that may climb to 4 m or more, but Passiflora trinifolia rarely exceeds a height of 1 m. The chartaceous leaves of Passiflora eglandulosa are bright green adaxially and possess drip tips, but the leaves of Passiflora trinifolia are dark green, lack long drip tips and are very stiff and rigid ( MacDougal 1988).

Passiflora eglandulosa is also similar vegetatively to Passiflora tacanensis , a species found in montane forests on Volcán Tacaná of Chiapas, Mexico. Both species possess wide, foliose stipules. However, the two species are easily separated because Passiflora tacanensis possesses petiolar nectaries. The fruits of Passiflora eglandulosa also possess fewer than 10 seeds, whereas Passiflora tacanensis possesses ca. 20 seeds per fruit.

The development and physiology of the floral nectary of Passiflora eglandulosa , misidentified as Passiflora trinifolia , was examined by Durkee et al. ( Durkee et al. 1981). She found that the floral nectary development and nectar secretion in this species is similar to that in the two other species of Passiflora that she studied. She concluded that the activity of an intercalary meristem increased starch deposition in the amyloplasts of the secretory cells parallels the maturation of the nectary phloem, and granulocrine secretion in the “starchy” nectaries does not occur. She also observed large membrane-bound protein bodies in the phloem parenchyma cells ( Durkee et al. 1981).

Benson et al. (1975), in a study of the coevolution of plants and herbivores, reported that Heliconius hortense is an herbivore of Passiflora eglandulosa (misidentified as Passiflora trinifolia ). This report was confirmed by MacDougal ( MacDougal 1988).

Specimens examined.

EL SALVADOR. Ahuachapán: Cerro Grande de Apaneca, 1700 m, Weberling 2610 (M). Santa Ana: Mountain Cerro Verde, 1800 m, Molina & Montalvo 21514 (F, NY). Sonsonate: near top of Cerro Verde, 1860 m, Croat 42222 (MO); Laguna de las Niñas, 1829 m, 13°53'N, 89°47'W, Villacorta 750 (MO); Laguna Verde, 1650 m, 13°54'N, 89°48'W, Villacorta & Gonzalez 683 (MO).

GUATEMALA. El Progreso: Montaña Canahui, between Finca San Miguel and summit of mountain, near upper limits of Finca Caieta, 1600-2300 m, Steyermark 43787 (F). Guatemala: Choacorral, km 20 aprox. llendo a San Juan Sacatepéquez, 2000 m, Castillo et al. 82347 (F); Santa Catarín Pinula, cerca la Cuidad Guatemala, barranca de Paraje Solar, Km 15.8 carr. de Cd. al Salvador, 1860 m, 14°32 N, 90°27 W, MacDougal & MacVean 6210 (MO); vicinity of San Andrecillo, 1700 m, Molina & Molina 27543 (F, U, US); near Canales, 1900 m, Williams & Molina 11822 (F). Huehuetenango: Mpio. Jacaltenango, Montaña Aqo’ma, 2278 m, 15°40'N, 91°39'W, Véliz et al. 16059 (BIGU). Jalapa: Volcán Jumay, N of Jalapa, 1300-2200 m, Steyermark 32352 (F). Quetzaltenango: 2.5 mi. below tunnel at Santa María de Jesus between km post 202-203 on Hwy. 97, 14°42'N, 91°32'W, MacDougal 316 (FLAS, MO); slopes of Volcán de Zunil, at and above Aguas Amargas, 2430-2850 m, Standley 65404 (F, US); along road above Santa María de Jesús, 1680 m, Standley 84846 (F, US); El Pocito, S of San Martín Chile Verde, on road to Colomba, 2200 m, Standley 84997 (F, G); slopes and ridges between Quebrada Chicharro and Montaña Chicharro, on SE-facing slopes of Volcán Santa María, 1300-1400 m, Steyermark 34360 (F, US). San Marcos: road between San Rafael Pie de La Cuesta and Palo Gordo, 3 km from Aldea Fraternidad toward San Marcos, parcelamiento "La Lucha," between Km posts 264265, 2150 m, 14°56 N, 091°51 W, MacDougal et al. 6234 (MO); road between San Rafael Pie de La Cuesta and Palo Gordo, 1 km above Aldea Fraternidad, between Km posts 266267, 1900 m, 14°56 N, 091°52 W, MacDougal et al. 6237 (MO); road between San Rafael Pie de La Cuesta and Palo Gordo, 3 km from Aldea Fraternidad toward San Marcos, parcelamiento "La Lucha," between Km posts 264265, 2150 m, 14°56 N, 091°51 W, MacDougal et al. 6249 (MO); Barranco Eminencia, road between San Marcos and San Rafael Pie de la Cuesta, in upper part of the barranco between Finca La Lucha and Buena Vista, 2500-2700 m, Standley 86379 (F); Barrancos 6 mi. S and W of Tajumulco, NW slopes of Volcán Tajumulco, below cliffs along Río Malacate, 2300-2800 m, Steyermark 36663 (F, US); on outer slopes of Tajumulco Volcano, Sierra Madre mountains about 8-10 km W of San Marcos, 2300 m, Williams et al. 26864 (F, GH, NY, US). Suchitepéquez: Volcán Santa Clara, between Finca El Naranjo and upper slopes, 1250-2650 m, Steyermark 46628 (F, US). Zacapa: Ravine bordering Quebrada Alejandria, summit of Sierra de las Minas, vincinity of Finca Alejandria, 2500 m, Steyermark 9859 (F).

HONDURAS. Santa Bárbara: Cuestas de piedra caliza, Dep. de Santa Bárbara, 10 km W de Lago Yojoa, 1500-2000 m, 14°55'N, 88°5'W, Clewell & Hazlett 3858 (MO, TEFH).