Passiflora pallida L. Sp. Pl. 955. 1753. non Passiflora pallida Lour., 1790. non Passiflora pallida Vell., 1827.

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 : 50-59

1. Passiflora pallida L. Sp. Pl. 955. 1753. non Passiflora pallida Lour., 1790. non Passiflora pallida Vell., 1827. Figs 23, 24

Passiflora hirsuta L. Sp. Pl. 958. 1753. non Passiflora hirsuta Loddiges, 1818. Type: Hispaniola, (no specimens extant; lectotype, designated here: Plumier Desc. Pl. Amer. pl. 88. 1693).

Passiflora minima L. Sp. Pl. 959. 1753. non Passiflora minima Blanco, 1837. Type: “Curassao,” [Curacao, Netherlands West Indies] 20 (lectotype, designated by Wijnands 1983 pg. 171: LINN 1070.20, microfiche seen).

Passiflora nigra Jacq., Observ. Bot. (Jacquin) 2: 27, pl. 46, fig. 3. 1767. Type: [Colombia, Cartagena, Boca Chica Inlet] (lectotype, designated here: Jacquin Observ. Bot. 2: 27 pl. 46, fig. 3. 1767).

Passiflora glabra Mill., Gard. Dict., eight ed., no. 4. 1768. non Passiflora glabra J.C. Wendl., 1805. Type: Based on Passiflora pallida L.

Passiflora parviflora Sw., Prodr. [O.P. Swartz] 97. 1788. Type: Jamaica, O. P. Swartz s.n. (holotype: S [S03-900][photograph seen]; isotype: MO! [MO-312541]).

Passiflora heterophylla Dryand., Hortus Kew. [W. Aiton] 3: 309. 1789. non Passiflora heterophylla Lam., 1789. non Passiflora heterophylla Jacq., 1797. Type: at Kew Gardens in ca. 1773, from the West Indies (no type material found).

Cieca minima (L.) Moench., Suppl. Meth. 102. 1802. Type: Based on Passiflora minima L.

Passiflora warei Nutt., Amer. J. Sci. Arts 5: 297. 1822. Type: United States of America. Florida: “Florida”, A. Ware s.n. (holotype: BM,! [BM000563877][photograph AAU!]).

Meioperis pallida (L.) Raf., Fl. Tellur. 4: 103. 1838. Type: Based on Passiflora pallida L.

Meioperis minima (L.) Raf., Fl. Tellur. 4: 103. 1838. Type: Based on Passiflora minima L.

Cieca pallida (L.) M.Roem., Fam. Nat. Syn. Monogr. 2: 142. 1846. Type: Based on Passiflora pallida L.

Cieca warei (Nutt.) M.Roem., Fam. Nat. Syn. Monogr. 2: 146. 1846. Type: Based on Passiflora warei Nutt.

Passiflora hirsuta var. parviflora (Sw.) M.Roem., Fam. Nat. Syn. Monogr. 2: 174. 1846. Type: Based on Passiflora parviflora Sw.

Passiflora suberosa var. minima (L.) Mast., Trans. Linn. Soc. London 27: 630. 1871. Type: Based on Passiflora minima L.

Passiflora suberosa var. hirsuta (L.) Mast., Trans. Linn. Soc. London 27: 630. 1871. Type: Based on Passiflora hirsuta L.

Type.

l’Isle S. Domingue (lectotype designated here: Plumier, Pl. Amer. pl. 89. 1693).

Description.

Slender, climbing, perennial vine 1-7 m long or more, sparsely to densely pubescent with unicellular curved trichomes on petiole, leaf, stem, and stipule, 0.20-0.30(-0.7) mm long, 0.02-0.03 mm wide, also minutely antrorsely appressed-puberulent throughout with unicellular, curved trichomes, 0.06-0.11 mm long, 0.02-0.03 mm wide. Flowering stems 0.6-1.6(-2.5) mm in diameter, terete or somewhat compressed, greenish yellow to very dark reddish purple, with the base woody and cork-covered. Stipules 2.1-6.9 mm long, 0.2-0.9 mm wide, narrowly ovate-triangular, sometimes slightly falcate, acute; petioles 0.3-1.8(-2.9) cm long, with 2 (rarely 1), opposite to alternate, stipitate or sometimes sessile, slightly obconical to capitate nectaries (very rarely crateriform), 0.3-0.8 mm wide (on the widest axis), 0.2-1.1 mm high, borne in the distal half of the petiole (0.49-0.92 of the distance from the base toward the apex of the petiole). Laminas 1.8-8.8(-12.0) cm long, (0.3-)1.4-8.2(-10.6) cm wide, membranous, unlobed to 3-lobed, lobed 0.20-0.50(-0.90) the distance to the leaf base, ovate to elliptic (rarely obovate), base cuneate to acute, lateral lobes 1.0-5.1(-6.8) cm long, 0.3-2.1(-3.0) cm wide, ovate to oblong, acute (rarely obtuse or rounded), central lobe ovate to elliptic (rarely obovate), central vein 1.8-8.8(-12.0) cm long, angle between the lateral lobes (33-)50-110(-152)°, ratio of lateral lobe to central vein length 0.46-0.78(-0.87), margins entire, hyaline, primary veins 1-3 (when more than one veins diverge and branch at base), laminar nectaries absent; tendril 0.2-0.7(-1.1) mm wide, present at flowering node. Flowers borne in leaf axils. Pedicels (2.0-) 3.3-9.4(-17.0) mm long, 0.3-0.6 mm wide, 2 per node; bract(s) absent or rarely with one narrowly ovate, acute, bract present on the distal half of the pedicel, 0.4-0.6 mm long, ca. 0.1 mm wide; spur(s) absent. Flowers (6.9-)11.7-20.4 mm in diameter with stipe 1.4-4.4(-6.3) mm long, 0.3-0.7 mm wide; hypanthium 2.8-4.1 mm in diameter; sepals (2.3-)4.0-7.0(-8.3) mm long, 1.2-3.3 mm wide, ovate-triangular, acute to rounded, reflexed at anthesis, abaxially and adaxially greenish yellow to very light greenish yellow (5GY 7/4, 8/4-8/2); coronal filaments in 2 series, the outer 20-30(-34), 1.2-4.0 mm long, (0.1-)0.2-0.6 mm wide, linear, slightly spreading, greenish yellow with yellow tips (5Y 8/10) or flushed with reddish purple (5RP 5/6-3/6) at base and greenish yellow at middle with yellow tips or very dark reddish purple (5RP 3/4-2.5/4) at base and yellow toward tips, ratio of outer coronal row to sepal length 0.20-0.69(-0.82), the inner (11-)20-34, 0.8-1.3 mm long, 0.04-0.16 mm wide, linear, capitate, erect, greenish yellow with yellow tips or greenish yellow flushed with reddish purple at base and yellow toward tips or very dark reddish purple with yellow tips, ratio of inner coronal row to outer coronal row length 0.36-0.66; operculum (0.6-)1.0-1.4 mm long, plicate, greenish yellow or greenish yellow with a flush of reddish purple at base or reddish purple or very dark reddish purple, margin white with minutely fimbrillate teeth; nectary (0.1-)0.2-0.4(-0.6) mm high, 0.2-0.6(-0.8) mm wide; limen recurved, erect or slightly inclined toward the operculum, 0.1-0.4 mm high, 0.1-0.3 mm wide, greenish yellow or greenish yellow flushed with red dish purple or reddish purple or very dark reddish purple, limen floor 1.6-2.6 mm in diameter, greenish yellow or greenish yellow flushed with reddish purple or reddish purple or very dark reddish purple; androgynophore (1.7-)2.2-3.5 mm long, 0.4-0.9 mm wide, greenish yellow or greenish yellow with a flush of reddish purple at base or greenish yellow with reddish purple spots and streaks or very dark reddish purple; free portions of the staminal filaments 1.4-3.0 mm long, 0.2-0.4 mm wide, linear, greenish yellow; anthers 1.1-1.9 mm long, 0.5-1.3 mm wide, pollen yellow; styles 1.6-4.3 mm long including stigmas, 0.1-0.4 mm wide, greenish yellow; stigmas 0.5-1.2 mm in diameter; ovary 1.1-1.8 mm long, (0.7-)1.0-1.5(-1.9) mm wide, ellipsoid to globose, greenish yellow. Berry 7.6-9.5 mm long, 6.9-8.8 mm in diamater, globose, or ellipsoid, very dark purple (5P 2.5/2). Seeds (4-)8-24(-33), 2.8-3.5 mm long, 1.9-2.2 mm wide, 1.1-1.4 mm thick, obovate in outline, acute at both ends, reticulate-foveate with each face marked with ca. 12-20 foveae; germination type epigeal.

Phenology.

Flowering and fruiting throughout the year.

Distribution.

In the New World tropics: Central America, Mexico, United States (Florida and Texas), Venezuela, and the West Indies. Introduced in the Old World tropics: Africa, Asia, and Australia. Growing in shrubs, trees or trailing on the ground in secondary successional areas and along the edges of dry tropical forests, both inland and near the seashore, primarily at low elevations but sometimes occurring at elevations as high as 800 m. Commonly associated with calcareous/alkaline substrate.

Ethnobotany.

In Réunion, the fruits may be used as a substitute for ink (Jean Jacques, pers. comm.).

Discussion.

Passiflora pallida as recognized here exhibits a substantial amount of morphological variation across its range. The various forms that the leaves may take have led to the proposal of many species and varietal names. For example, a plant of this species may possess only unlobed leaves, only trilobed leaves, or leaves that are unlobed, bilobed, and trilobed. This type of variation can be seen throughout the range of this species. However, the flowers of Passiflora pallida are diagnostically small, with a narrow hypanthium, short sepals, short coronal filaments, and narrow floral nectaries.

The only species with which Passiflora pallida may be confused is Passiflora suberosa . Passiflora pallida is vegetatively similar to both Passiflora suberosa subsp. litoralis and Passiflora suberosa subsp. suberosa , and without flowering material these taxa can be difficult to distinguish. The position of the petiolar nectaries has often been used to separate species in closely related taxa in Passiflora . However, though the petiolar nectaries are generally located closer to the petiole apex in Passiflora pallida than in the South American populations of Passiflora suberosa subsp. litoralis and Passiflora suberosa subsp. suberosa , the upland Mexican/Central American populations of Passiflora suberosa subsp. litoralis also have petiolar nectaries positioned very near the petiole apex. The leaf base of Passiflora pallida is commonly not cordate. Passiflora suberosa possesses leaves that are frequently cordate, though this character is somewhat variable in the upland Mexican/Central American populations of Passiflora suberosa subsp. litoralis . Though foliage color is difficult to discern from herbarium specimens, my experience in the field and photos taken of Passiflora pallida and Passiflora suberosa in the field by others show that Passiflora pallida commonly possesses leaves that are paler in color and often less lustrous than Passiflora suberosa . Reproductive structures are more reliable in separating Passiflora pallida and Passiflora suberosa . The hypanthium of Passiflora pallida is commonly 2.8-4.1 mm in diameter and the inner coronal filaments are usually less than 1.5 mm long. In Passiflora suberosa s. l., the hypanthium is commonly 4.0-8.8 mm in diameter and the inner coronal filaments are frequently 1.5-3.9 mm long. The outer coronal filaments are also short, less than 4.0 mm in Passiflora pallida , and although that overlaps with the 2.5-8.1 mm range observed in Passiflora suberosa , the character is frequently observable in herbarium specimens. Where the distributions of Passiflora pallida and Passiflora suberosa overlap in the Antilles, Passiflora pallida is typically found in and along the edges of subtropical and tropical forests at or near sea level (rarely exceeding 200 m), whereas Passiflora suberosa commonly occurs in and along the edges of tropical forests above 500 m.

The most common variant of Passiflora pallida (as exemplified by E. Killip 41876, on Sugarloaf Key, Monroe Co., Florida, USA; E. Cabrera 1475, S of Akumal, Quintana Roo, Mexico; and J. Tillich 3558, in Black River, Mauritius) has ovate leaves that may be unlobed, bilobed or trilobed on the same plant. When unlobed, the leaves are commonly greater than 2.0 cm wide. When lobed, the leaves are usually shallowly lobed 0.20-0.41 the distance to the base, the lateral and central lobes are greater than 1.0 cm wide, and the angle between the lateral lobes is 45-100°. Another less common variant of Passiflora pallida (as exemplified by J. K. Small & C. Mosier 5511, from Cox Hammock, Miami-Dade Co., Florida, USA; and J. Small & J. Carter 194, between Perrine and Long Prairie, Miami-Dade Co., Florida, USA), has narrowly ovate leaves that may be unlobed, bilobed or trilobed on the same plant. When unlobed, the leaves are commonly less than 1.0 cm wide. When lobed, the leaves are usually deeply lobed 0.82-0.90 the distance to the base, the lateral and central lobes are commonly less than 0.7 cm wide, and the angle between the lateral lobes is greater than 100°. However, all the specimens brought together here as Passiflora pallida are all relatively small in stature in their native habitats in the New World, possess similar small flowers with short coronal filaments and occur in a similar range of elevations.

MacDougal has reported the appearance of an occasional, well-formed but small petal in other species within supersection Cieca (MacDougal, 1992). I have also seen this in Passiflora pallida in several of my greenhouse accessions and in the field in Quintana Roo, Mexico.

Stegmaier (1973) reported that Dasiops passifloris McAlpine ( Diptera : Family Lonchaeidae ) infests the fruits of Passiflora pallida in southern Florida. He found that the female fly oviposits on the fruits and the larvae feed on the arils and fruit flesh. In this study, in which he collected a total of 1040 wild passion fruits from Passiflora pallida occurring on a single farm in Hialeah, Florida, he also found that the mature fruit may contain from 4 to 17 seeds per fruit ( Stegmaier 1973).

Passiflora pallida is a pest plant where it occurs in many areas of the Old World. In New Guinea, Neville Kemp reports that the probable disperser of Passiflora pallida is the long-tailed macaque or crab-eating macaque ( Macaca fascicularis ) ( Kemp and Burnett 2003).

In Linnaeus’ 1753 edition of Species Plantarum, he describes three small-flowered entities, Passiflora pallida L. ("Habitat in Dominica, Brasilia"), Passiflora hirsuta L. ("Habitat in Dominica and Curassao") and Passiflora minima L. ("Habitat in Curassao"), for which the historical references include phrases such as "flore minore" ( Passiflora pallida L.), "flore & fructu minimis" ( Passiflora hirsuta ) and "flore flavescente omnium minimo" ( Passiflora minima L.). Charles Wright (1869), in an article discussing the genus Jussiaea L., chose the name Passiflora pallida over Passiflora minima . In the article he commented on the “embarrassing” status of the species of Passiflora and the unwise reliance upon vegetative morphology in species circumscription within the Cuban species of Passiflora (Wright, 1869:480). In the article he states, "I have lately carefully examined the Cuban species called Passiflora minima , hederacea , pallida , angustifolia , suberosa , &c., and come to this conclusion:- Passiflora pallida , L., is an old and appropriate name, to which belong Passiflora minima , L., and Passiflora angustifolia , Sw., certainly; Passiflora hederacea , Cav., Passiflora suberosa L., probably; and, from the description, I judge Passiflora lineariloba , Hook. f. to be only another form of it." It is possible that Passiflora hirsuta was not considered by Wright in his article because of the confusion surrounding its circumscription (see below) or because he had not encountered the taxon in Cuba.

In the 1753 edition of Species Plantarum, Linnaeus indicated that he was well-acquainted with Passiflora pallida and refers to the diagnosis and drawing in his Dissertatio botanica de Passiflora (1745), that shows an unlobed, ovate leaf with two petiolar nectaries positioned near the apex of the petiole. Linnaeus cites an illustration by Plumier (pl. 89, in Description des plantes de l’Amérique 1693) that also exemplifies his Passiflora pallida . However, he also refers to a figure by Morison (1680) that shows a plant with a large flower that possesses sepals and petals (likely in the subgenus Passiflora ) with unlobed, ovate leaves. In the 1745 dissertation, Hallman specifically states that the flowers of Passiflora pallida L. are “pentapetala”, referring to the lack of petals; this decision was based upon the careful comparison of diagnoses from other petalous taxa in the treatment. An examination of the Linnaean herbarium (microfiche) did not reveal an herbarium specimen that could reasonably be attributed to the species described as Passiflora pallida by Linnaeus. There is one specimen in the Linnaean herbarium labeled Passiflora pallida , but it is a post 1753 accession that represents a large-flowered taxon from subgenus Passiflora . Though there is a small amount of confusion surrounding Passiflora pallida L., largely attributable to Linnaeus’ reference to Morison’s illustration and the post-Linnaean accession referred to above, it is clear from the diagnoses in the 1753 edition of Species Plantarum and the 1745 dissertation, that Linnaeus was referring to a plant that had unlobed, ovate leaves and small, pale, apetalous flowers ( Jarvis 2007). The lectotype of Passiflora pallida L. (designated here) is Plumier’s fig in Description des plantes de l’Amérique (1693) in which he illustrated several entities of both Passiflora pallida and Passiflora suberosa subsp. suberosa . Incidentally, Linnaeus chose the epithet, pallida , to refer to the pale-colored flower. Though the flowers are frequently pale in color, they may also be highly colored.

Linnaeus (1753) also describes Passiflora minima L. as a trilobed plant in which the central lobe is longer than the lateral lobes. He cites the diagnosis and drawing in the 1745 dissertation that shows a plant with narrowly trilobed leaves that lack petiolar glands. Linnaeus (1753) also refers to a figure by Plukenet (1696) that closely matches Linnaeus’ diagnoses and the drawing in the dissertation. Neither Linnaeus nor Hallman described the flowers of Passiflora minima L., but it can be inferred by the historical references in the dissertation that the flowers were small and lacked petals. In Killip’s treatment of Passiflora suberosa , he states that there are two sheets of Passiflora minima from the "West Indies" of uncertain origin in the Linnaean Herbarium and designated them "type of Passiflora minima " (1938:93) without specifying one of the sheets specifically. According to Jarvis (2007), the lectotype of Passiflora minima L. (designated by Wijnands 1983) is specimen 1070.20 (LINN). The lectotype closely matches Linnaeus’ diagnoses and the drawing in the dissertation and possesses small flowers apparently lacking petals (as observed on a microfiche of the herbarium). However, the lectotype of Passiflora minima is a very unusual example of the small-flowered entity, as the lack of petiolar nectaries in this taxon is very rare.

Passiflora hirsuta L. has been the source of confusion for several taxonomists of Passiflora , and under his treatment of Passiflora foetida var. moritziana (Planch.) Killip ex Pulle, Killip (1938) discussed the problem. Linnaeus (1753) cited several references in his treatment of Passiflora hirsuta , often with accompanying illustrations, that undoubtedly refer to Passiflora foetida . However, he also referred to an illustration by Plumier (pl. 88, in Description des plantes de l’Amérique 1693) that is clearly Passiflora pallida . As in his other species descriptions, he also cites the diagnosis and drawing in the 1745 dissertation by Hallman that shows a trilobed, densely pubescent leaf with rather large petiolar nectaries that are positioned on the distal half of the petiole and, thus, cannot be Passiflora foetida as this species lacks petiolar nectaries. The diagnosis in the 1753 edition of Species Plantarum is unclear. However, in the 1745 dissertation Hallman states that the flowers of this taxon are pale and small, the involucre is lanceolate, and the fruits are deep blue. Hallman goes on to say that the taxon that he is describing is somewhat similar to the next ( Passiflora foetida L.) but differs in that the flowers are opposite (paired) and the involucre consists of only a single bract. Hallman is clearly describing one of the entities in the Passiflora suberosa complex, as the flowers are commonly paired in the leaf axils, members of the species complex do sometimes possess one or two lanceolate bracts, and the fruits are very dark purple. In Passiflora foetida only one flower is present in the leaf axils, the involucre consists of three large bracts that are pinnatifid or pinnatisect, and the fruits are yellow to red. Though the leaf as illustrated in the dissertation is distinctly cordate and broadly ovate, which is a bit unusual for Passiflora pallida , Linnaeus’ reference to Plumier’s drawing leads me to conclude that Passiflora hirsuta L. is a synonym of Passiflora pallida . It is also the only original material that corresponds to the current concept of the species.

Selected specimens examined.

United States. Florida: Brevard Co.: near Cape Malabar, Curtiss 974 (BM, G, GH, M, MIN, MO, NY, US). Broward Co.: along U.S. 27, 6 mi. N of Andytown, Beckner 769 (MO); along canal N of Rt. 84 near Florida State Forestry Station, W of Florida Turnpike, Correll et al. 40198 (MO); along US 27, 1 mi. S of intersection with Fla. 84 at Andytown ("Twenty Mile Bend"), ca. 18 mi. W of Ft. Lauderdale, Ward & Burch 3334 (FLAS). Charlotte Co.: Bull Key, opposite Lemon City, Small & Carter s.n., 6 November 1903 (NY). Collier Co.: on tram, lower Fahkahatchee strand, Atwater 684 (FLAS); Goodland Point, Marco Island, Brass 18081 (FLAS, US); lagoon embankment W of Everglades City, Lakela 31716 (CONN, FLAS, GH, MIN). Duval Co.: Mouth of the St. John’s River, Curtiss 973 (FLAS, G, GH, M, MO, NY, US). Hendry Co.: E side on dirt road, 5 mi. S of Fla. 846, ca. 10 mi. due ESE of Immokalee, Ward et al. 5396 (FLAS). Highlands Co.: The Archbold Biological Station, Cooley et al. 9434 (GH). Hillsborough Co.: 2511 LaSalle St., Almeda 367 (FLAS). Indian River Co.: near Roseland; near corner of 110th St. along road bordering Indian River, 0.3 mi. S of junction with US 1., Wunderlin & Beckner 6490 (NO). Lake Co.: in city park, Leesburg, Baltzell 6619 (FLAS). Lee Co.: Pine Island, S of jct. Rt. 767 & Rt. 78, Lakela, Long & Broome 305897 (FLAS). Levy Co.: Waccasassa Bay State Preserve, N of Turtle Creek, Abbott & Williams 8461 (FLAS); Seahorse Key, West et al. s.n., 1 August 1958 (GH). Manatee Co.: Perico Island, Tracy 7655 (BM, G, GH, MIN, MO, NY, US). Martin Co.: S.R. 714, 16 mi. W of Palm City, McCart 11226 (FLAS). Miami-Dade Co.: 0.2 mi. N of SW 288th St., to the W side of SW 170th Ave., Homestead, Goldman & Hammer 1654 (MO); between Perrine & Long Prairie, Small & Carter 194 (US); Goodburn Hammock, Small & Mosier 5923 (G, NY, UC); Miami, Tracy 9168 (BM, G, GH, MIN, MO, PH, NY, TEX, US); along US 41, 2.2 mi. W of int. with Fla 27 W of Miami, Ward & Burch 3983 (FLAS). Monroe Co.: along road 4A about 0.5 mi. S of Islamorada, Deam 61084 (DUKE); Sugerloaf Key, Killip 41876 (B, NO, TEX, US); Key West, Hammock between Flagler Ave. and airport, Killip 44478 (US); N Key Largo, 6.5 mi. from US 1, on Fla 905, Long et al. 2800 (MIN); Cox Hammock, J. K. Small & C. Mosier 5511 (NY); Pinecrest, S of Fla. 94 ("Loop Road"), 4 mi. W of Miami-Dade-Monroe county line, NE corner of Ward & Burch 3309 (FLAS, GH). Palm Beach Co.: Singer Island, 9 mi. S of Lost Tree Village, McCart 11155 (FLAS). Pinellas Co.: N end of Long Key, town of St. Petersburg Beach, near Ciega Bay, Ward & Ward 2333 (FLAS). Polk Co.: vicinity of Crooked Lake, McFarlin 3958 (TEX). St. Johns Co.: S end near South Point loop, Harrison & Harrison 722 (FLAS). St. Lucie Co.: along edge of Indian River, Fla. 707, 1.3 mi. S of jct. with Fla. 712, ca. 4 mi. S of Ft. Pierce city limit, Ward & Crosby 4839 (FLAS, NY). Sarasota Co.: 2482 Linwood Dr., north of Bee Ridge and Webber, east of McIntosh, south of Bahia Vista, west of I-75, Abbott, J. R. 14284 (FLAS); Manasota Key, 6070 Manasota Key Road (S.R. 776), 3.2 mi. from S.R. 774, Lott & Lott DT1085 (FLAS). Volusia Co.: N end of Meritt Island, Apollo Beach between Turtle Mound and the House of Refuge Site, Cape Canaveral National Seashore, Judd et al. 3245 (FLAS, NY, U). Texas: Cameron Co.: La Palmas Plantation, about 4 mi. SW of Brownsville, Correll 14852 (GH, TEX, US). Hidalgo Co.: ca. 0.5 mi. ESE of Anzaldua Dam, ca. 4.5-4.6 airmi. S of jct. of US Rt. 83 and F.M. 1016 at Mission, Río Grand Valley National Wildlife Refuge, Gabrielson Tract, Mission Quadrangle, 105-110 ft., Carr & Hernández 14374 (LL).

Antigua and Barbuda. Antigua, Bodkin Estate, Box 1258 (BM, US).

Bahamas. Acklins and Crooked Islands: Acklins Island, Gold Rock, Brace 4414 (NY, US); Acklins Island, about 4 mi. N of Pinefield, Correll 44459 (NY). Bimini: S Bimini, Howard & Howard 10072 (A, GH, NY, S, US). Cat Island: Atlantic shore at Bird Point, Byrne 242 (A). Exuma: Hummingbird Cay, at top of Mt. Earlham, Nickerson & Gross 3017 (A, MO). Freeport: Grand Bahama, ca. 0.5 mi. W of Hawksbill Creek channel (vicinity of Eight Mile Rock), Webster 10914 (US). Fresh Creek: Andros, across road from Andros Town Airport, Fehling 17 (NY). Governor’s Harbour: Great Abaco, Abaco, along Queen’s Hwy. about 8 mi. S of Marsh Harbour, Correll & Wassjausen 52077 (US). Harbour Island: N Eleuthera, in coppice near turn-off to road to ferry slip to Harbour Island, about 1.5 mi. N of Lower Bogue, Correll 40996 (NY). Kemps Bay: Andros Island, Mangrove Cay, Bryant 4 (GH). Long Island: upper beach strand, Clarence Town, Harbor Area, Hill 823 (A). Marsh Harbour: Abaco, along Forest Drive, about 1.5 mi. NW of Marsh Harbour, Correll & Meyer 44600 (NY). New Providence: near Harrold & Wilson Ponds, Degener 18964 (NY). Nichollstown and Berry Islands: Anderson Cay, Great Harbour Cay, Correll & Correll 43669 (NY). San Salvador: off Jake Jone’s Road (to Little Lake S from Queen’s Hwy.) near Barker’s Point, NW part of island, Romansky et al. 13 (FLAS).

British Overseas Territory. Turks and Caicos Islands: E Caicos, Jacksonville and vicinity, Millspaugh & Millspaugh 9073 (NY).

British Virgin Islands. Anegada, near settlement, Britton & Fishlock 1005 (NY); Virgin Gorda, Fishlock 136 (NY).

Cuba. Camagüey: Silla de Cayo, Cayo Romano, Shafer 2529 (BM, NY, US). Cienfuegos: Farallones de Guajímico, on the coast E of Cienfuegos, Morton 10492 (US). Ciudad de la Habana: Near Havana, Curtis 552 (BM, C, G, GH, M, MINN, MO, NY, PH, PR, US). Guantánamo: Beside the Río Jauco, S coast of Oriente, 50 m, Morton & Alain 9159 (US). Holguin: Holguin, Cerro de Fraile, Ekman 7551 (S). Isla de Juventud: along road from Nueva Gerona to Santa Bárbara, Killip 45109 (US). La Habana: between Madruga and Robles, Shafer 36 (NY). Matanzas: vicinity of Matanzas, Empalme, Britton et al. 584 (NY). Pinar del Río: trail from Buenaventura to San Juan de Guacamaya, Wilson 9348 (NY, U). Santiago de Cuba: between Sardinero and Siboney, Santiago, Clemente 6396 (GH). Villa Clara: 5 km W of Santa Clara, Howard et al. 441 (A, MIN, UC).

Dominica. Saint Patrick: SE coast, path between Delices & Belvedere Estate, 250 m, Whitefoord 3758 (BM).

Dominican Republic. Baoruco: Dos Brazos, 8.5 km N of Neiba, 400 m, Maas et al. 8391 (U). Barahona: N end of Beata Island, Howard 12449 (A, US). Dist. Nacional: 8 km from La Batatas (via Laguna La Jagüita) on road to Mata de Piedra and La Catalina, 20 m, Mejia & Zanoni 9744 (MO, NY). El Seibo: vicinity of Higuey, near Canada Honda, Howard & Howard 9754 (BM, GH, NY, S, US). Elías Piña: Vieja Estrelleta, 9.1 km SW de El Cercado en la carretera a Hondo Valle, entre Sonador & Juan Santiago, 740 m, Zanoni 27976 (JBSD). Independencia: aprox. 12 km S de Duvergé, en el lugar llamado Monte Palma, 860 m, Garcia et al. 4451 (B). La Altagracia: half-way between Boca del Yuma town and El Caracol (old bridge over Río Duey, N of town), 20-60 m, Zanoni et al. 10740 (JBSD, MO, NY). La Romana: Río Cumayasa river valley, just N of town of boca de Cumayasa, on SW side of river mouth, 0-20 m, Mejia & Ramírez 14786 (JBSD, MO, NY). La Vega: vicinity of Piedra Blanca, knob S of Piedra Blanca, 200-500 m, Allard 17783a (US). María Trinidad Sánchez: Cabo Frances Viejo, Smith 10452 (JBSD, NY). Monte Cristi: near Puerto Libertador, Manzanilla Bay, Howard & Howard 9633 (BM, GH, NY, S, US). Pedernales: 22 km N del puerto de Cabo Rojo en la carretera de Alcoa Exploration Company a las Mercedes & Aceitillar, 400 m, Zanoni & Pimentel 25943 (JBSD). Puerto Plata: en manigua costera, Cabarete, Puerto Plata, Alain & Liogier 26347 (JBSD). Samaná: Samaná, en la zona Costera entre el km 4 & el km 5 E del pueblo de Las Terrenas, en el área de “protillo,” 3 m, Zanoni & Mejia 17728 (NY). San Cristóbal: Nigua, Faris 442 (US). San Pedro de Macoris: Town S of Boca de Soco, at SW bank of Río Soco at its mouth, small village along river and sea coast, 5 m, Mejia & Zanoni 8595 (JBSD). Santiago Rodríguez: 20 km desde Sabaneta en la carretera a Monción, 250 m, Zanoni & Pimentel 25441 (JBSD, MO, US).

French Overseas Department. Guadeloupe. Lamentin, Apres Duportail, en prenant le sentier du Chemin des Contrebandiers, 300 m, Jeremie 295 (A, US). Martinique: Saint Martin, near Mullet Pond, Boldingh 2764 (U).

Grenada. St. George: near Mount Parnassus, Broadway 1720 (GH, NY).

Haiti. Artibonite: NE of Gros Morne, 235 m, Leonard 9788 (US). Nord: vicinity of St. Michel del Atalaye, 350 m, Leonard 7140 (NY, US). Nord-Ouest: vicinity of Port de Paix, Leonard & Leonard 11181 (NY, S, UC, US). Ouest: herbarium at the Faculte D’Agronomie at Medicine Veterinaire at Damien, 310 m, Paul & Porter-Utley AP516 (FLAS).

Jamaica. Clarendon: Peckham Woods, near Aenon Town, 2300 ft., Crosby & Anderson 1233 (DUKE). Hanover: Lucea, Hitchcock s.n., 3 January 1891 (MO). Kingston: vicinity of Kingston, Maxon & Killip 338 (BM, GH, MO, US). Manchester: vicinity of Mandeville, Crawford 742 (NY, PH). Portland: Navy Island, Fredholm 3076 (NY, US). St. Andrew: 1.5 mi. SSW of Lucky Valley, along the road to Bull Bay, 403 ft., Porter-Utley et al. P-57 (FLAS). St. Ann: Union Hill and vicinity, N slopes of Mount Diablo, 400-750 m, Maxon 10398 (US). St. Catherine: Pigeon Island, 10 mi. off Old Harbour Bay, Maxon & Killip 1580a (US). St. Elizabeth: near pit 101, Kaiser mine area S of Gutters, Howard & Proctor 13763 (A). St. James: near the mouth of Great River, W of Montego Bay, sea level, Maxon & Killip 1426a (US). St. Thomas: 14 mi. SE of Kingston toward Morant Point, Wunderlin 5135 (MO). Trelawny: Quickstep Forestry Road, Kay SQFR1 (FLAS). Westmorland: about 2 mi. W of White House, 0-100 m, Yuncker 18026 (NY). Unknown Parish: Ehb 32 (B).

Netherlands Antilles. Bonaire: Mont Kr, Boldingh 7396 (U); Stoffers 545 (U). Curaçao: near Sint Christoffelberg, Curran & Haman 205 (CAS, GH, PH, US). Sint Eustatius: N rim of the Quill, 580 m, Howard 18112 (A).

Netherlands Autonomous Country. Aruba: Arnoldo 187 (U); Boldingh 6515 (NY, U); Stoffers 2036 (U).

Puerto Rico. Bayamón: Bo. Hato Tejas, series of mogotes W of Rt. 871 (only central pair collected), 25-100 m, Axelrod & Axelrod 2346 (MO). Cabo Rojo: Salinas de Cabo, Rojo ad Punta de Aguila, Urban 644 (BM, G, GH, M, PR, S). Caja de Muertos: Cayo Muertos, N.L. Britton, Cowell & Brown 5030 (NY). Ciales: Bo. Hato Viejo, Rt. 6685, 1 km N jct Rt. 632, 50 m, Axelrod & Axelrod 4430 (NY). Dorado: Mpio. de Dorado, Rte 693 at the freeway extension and on mogotes just E, Taylor et al. 10050 (MO). Guánica: prope Guánica in litoralibus ad salinas, Urban 3488 (BM, G, GH, M, MO, NY). Isla Desecheo: Desecheo Island, Warshall 106 (GH). Mayagüez: Isla de Mona; Sardinera, lado E, Acevedo & Siaca 4302 (NY). Ponce: Ponce to Peñuelas, Britton et al. 1764 (NY). Quebradillas: Rt. 437 ca 1-2 km S of rte 113, 100-200 m, Taylor & Gereau 10487 (MO). Rincón: Rincón, Urban 5667 (G). Toa Alta: Rt. 677 ca. 3 km S of Rt. 2, 100 m, Taylor & Miller 10409 (FLAS, MO). Vega Alta: Rte 620 km, 4.0 m, Taylor & Molano 8681 (MO).

Saint Lucia. Vieux-Fort, Maria Island, Pierre et al. 261 (A).

Trinidad and Tobago. Trinidad: Bird of Paradise Island, Yellowtail Walk, 60 m, Webster 24186 (TRIN). Tobago: Banaan, Broadway 4236 (M).

United Kindom Overseas Territory. Anguilla: Bottom district, N of The Valley, Proctor 18538 (A, BM, US). Bermuda: Spittle Pond, Brown 718 (GH, NY, PH, US). Cayman Islands: Grand Cayman, near larva survey site 80, trans island road, Brunt 2164 (BM). Paget: Paget, Harshberger s.n., 19 June 1905 (MO).

United States Virgin Islands. St. Croix: A. Benzon 199-5098 (C); Reugen 199 (C). St. John: Cruz Bay, Maria Bluff, 90 m, Acevedo et al. 2330 (US). St. Thomas: Orsted s.n. (C).

Mexico. Campeche: Mpio. Hopelchén, 11 km S de la frontera Yucatán-Campeche, ca. de San Antonio Yax-che, Carnevali et al. 5675 (CICY). Quintana Roo: Hwy. 307 between Chetumal and Cancún, 30 m, Porter-Utley & Mondragón 393 (FLAS, CICY); road between Chetumal and Cancún, Porter-Utley & Mondragón 398 (CICY). Tamaulipas: Gómez Farias, 3 km below city plaza off main road, MacDougal 259 (DUKE, US). Veracruz: Mpio. Emiliano Zapata, 0.5 km de la desviación a Carrizal por la carretera Xalapa-Veracruz, Calzada 1838 (F, XAL). Yucatán: off of Mexico 180 between X-can and X-Uilub, 60 m, Porter-Utley & Mondragón 403 (CICY); road between X-can andX-Uilub, small path off main highway, Porter-Utley & Mondragón 405 (CICY); on small dirt road off of the road between Vallodolid and Tulum, Porter-Utley & Mondragón 407 (CICY); Mérida, CICY, Jardín Botánico, 20 m, Porter-Utley & Mondragón 412 (CICY); Oxkutzcab, Labná, S de la entrada, 10 m, Puch & Narvaez 488 (CICY, XAL); Oxkutzcab, Xul, camino antiguo a Benito Juárez 4 km, 60 m, Sanabrio & Sima 194 (CICY).

Belize. Belize: Caye Caulker, N Island, Whitefoord 8223 (BM, F, MA, MO). Stann Creek: Northeast Cay, Glover’s Reef, 0-3 m, Fosberg & Sachet 53819 (B, BM, F, GH, MO, NY, US). Toledo: NE Sapodilla Cay, Spellman & Stoddart 2322 (MO, US).

Guatemala: Petén: Dos Lagunas, Ixcanrío, on Aguas Turbias Road, Contreras 8687 (F, LL, MO).

Honduras. Islas de la Bahía: Cayo Grande de Cayos Vivorillo, Valerio 270 (MO, TEFH).

Nicaragua. Zelaya: Cayo Palmeta, 0-10 m, Stevens & Krukoff 20764 (MO).

Panamá. Panamá: Bella Vista, at sea level, Killip 12039 (US). San Blas: Soskatupu, island ca 1.5 mi. long, 0.5-0.7 mi. broad, 0-150 ft., Elias 1692 (MO, UC).

Colombia. Atlántico: Usiacurí, Arroyo del Higuerón, 100 m, Dugand & Garcia 2277 (US). Bolívar: vicinity of Turbaco, 200-300 m, Killip & Smith 14329 (GH, US). Magdalena: Buritaca, 50 mi. E of Santa Marta, Smith 1531 (NY). San Andrés and Providencia: San Andrés Island, along beach near Sound Bay Cemetary, Weston & Weston 5542 (UC). Sucre: Mpio. Cartagena, Archipelago San Bernardo, Isla de Tintipán, Mar Caribe, 2 horas por bote NW de Tolú, Callejas & Bornstein 11031 (HUA).

Venezuela. Bolívar: Carapo, unknown collector 27 (PR). Dependencias Federales: Archipielago Los Testigos, Isla Testigo Grande, Playa Guzmán, Fernández José, Flores-Javier & Fernández 607 (NY). Falcón: Dist. Silva, al pie de los penascos calcareos, S de la Punta Faustino, SE de Chichiriviche, 1-3 m, Steyermark & Manara 110380 (MO, US). Sucre: vicinity of Cristóbal Colón, La Planisa, Broadway 340 (US).

Comoros. Anjouan: Schlieben 11161 (B, M, MO). Gran Comore: S edge of Moroni, D’Arcy 17538 (MO). Moheli: Schlieben 11248 (B, M, MO).

French Overseas Possession. Glorioso Islands, NW corner of Gloriosa Island, sea level, Frazier 107b (CONN, US).

Madagascar. Lemberano, Hildebrandt 3264 (G, M).

Republic of Seychelles. Aldabra Islands: Mahé Island, Mahé, Pointe La Rue hill, 1200 ft., Osborne-Day 124 (BM).

British Indian Ocean Territory. Chagos Archipelago, Diego Garcia, Hutson 27 (BM, US).

India. West Bengal: Calcutta, Kuntze 6385 (NY).

Maldives. Seenu: Addu Atoll, Gan Island, S side of airstrip, Sigee 55 (US).

Mauritius. Agalega Islands: S Island, Stoddart 7263 (US). Unknown District and Dependency: Black River, Tillich 3558 (MSB).

Singapore. Telok Paku, Sinclair 6467 (US).

Sri Lanka. Central: campus of Univ. of Ceylon, Peradeniya, 500 m, Comanor 324 (MO, US). Sabaragamuwa: 12th mi. post on the road between Panamure and Kilanne, Ratnapura Dist., Balakrishnan & Jayasuriaya NBK911 (US). Western: Induruwa, Jacobsen 13-6 (C).

Australia. Queensland: Rockhampton, Boorman s.n., August 1912 (B).

Commonwealth of the Northern Mariana Islands. Saipan, E of Ogso Tafotchau just N of Kannat Tadung Laulau, 170-190 m, Fosberg 50550 (US).

Federated States of Micronesia. Pohnpei: Ascension Island, 500 ft., Saltis 385/5 (BM).

French Overseas Territory. New Caledonia: Sous-bois S sol calcaire, Îlot Maître pres Nouméa, Guillaumin & Hurlimann 727 (NY, US).

Palau. Koror: R. Bishop P-10192 (US).

Solomon Islands. Guadalcanal: Lunga, sea level, Brown 1448 (BM).

United States Territory. Guam: Trust Territory Compound, NAS, Agana, 70 m, Fosberg 46212 (BM, UC, US).

Cultivated Material. United States: North Carolina, cultivated at Duke University 1980-1984 from seeds sent by Jack Longino in 1979 from Florida, MacDougal 662 (FLAS, MO); Florida, cultivated at the University of Florida from material collected at Passiflora Society International meeting, Porter-Utley P-65 (FLAS).