Many think of Epiphyllum and orchid cactus as synonymous, but orchid cacti are now placed in Schlumbergera or Diso-cactus, closely related genera in the same tribe, Hylocereeae. One encounters numerous so-called epiphyllum hybrids, which in fact rarely involve species of Epiphyllum as a parent. Rather, these magnificent plants, bearing shimmering, iridescent flowers, are mostly hybrids of related genera such as Disocactus, Pseudorhipsalis, and Selenicereus.

Epiphyllum (type, E. phyllanthus) is generally recognized as comprising epiphytic cacti with flattened leaflike shoots and large white flowers open at night. The flowers also tend to have very long floral tubes. The genus was described in 1812 by Adrian Haworth, the meaning of the name a bit misleading for it is from the Greek, meaning upon a leaf. Haworth was referring to the flowers, borne on what he thought were leaves but, in fact, are stems. Epiphyllum comprises 19 species. Myron Kimnach (1964,1965a,b, 1967,1990a,b) has published many excellent articles on the genus.

Epiphyllum Haworth 1812 Phyllocactus Link 1831 Marniera Backeberg 1950

Subfamily Cactoideae, tribe Hylocereeae. Plants mostly epiphytic but some lithophytic, freely branching, erect, climbing, or pendent shrubs, often with aerial roots. Older stems round in cross section basally, usually spineless, often woody. Younger stems flattened, leaflike, having margins with broadly rounded or sharp teeth, sometimes lobed or pinnatifid between the areoles. Spines usually absent. Flowers borne laterally, solitary, usually open at night, saberform or funnelform, 10-30 cm (3.9-12 in) long; pericarpels with small scales, rarely with hairs or bristles; floral tubes long, with naked scales, abruptly dilated at the throats; outer perianth parts whitish, yellowish, or pinkish, inner pale yellow or white; stamens inserted in the throats. Fruits ovoid or oblong, bearing small scales and areoles, somewhat ridged, spineless, 4-9 cm (1.6-3.5 in) long, 2-5 cm (0.8-2 in) in diameter. Seeds kidney shaped, black. Distribution: primarily Central America and Mexico but with a few species extending into the Caribbean and northern South America.

Epiphyllum anguliger (Lemaire) D. Don 1855 moon cactus, queen ofthe night Phyllocactus anguliger Lemaire 1851 Phyllocactus darrahii K. Schumann 1903, Epiphyllum darrahii (K. Schumann) Britton & Rose 1913

Primary stems cylindrical or three-angled basally. Lateral stems lance shaped, flattened except basally somewhat fleshy, green, with pronounced midribs, rather stiff, margins wide toothed and deeply lobed, to 1 m (3.3 ft) long, 4-8 cm (1.6-3.1 in) wide. Areoles sometimes with one or two small white bristles. Flowers scented, 15-18 cm (5.9-7.1 in) long, 10-13 cm (3.9-5.1 in) in diameter; outer perianth segments lemon or golden yellow, inner pure white. Distribution: central and southern Mexico. Fruits of Epiphyllum anguliger are gooseberry-like (Chapter 2, under Cacti as Food).

Epiphyllum cartagense (F. A. C. Weber) Britton 8c Rose 1913 Phyllocactus cartagensis F. A. C. Weber 1902

Plants forming thickets with erect, arched, or pendent stems. Primary stems three-angled or round in cross section basally, spineless, to 2 m (6.6 ft) long. Roots often aerial, numerous. Secondary stems arising at nearly right angles from upper portions of primary stems, arranged in three rows, basally diverse, apically flattened, lobed and toothed, 2-7 cm (0.82.8 in) wide. Areoles spineless. Flowers usually borne on the flattened portion of secondary stems, tubular funnelform, 15-18 cm (5.9-7.1 in) long, floral tubes, very slender, nearly straight on erect stems or strongly upcurved on pendent stems; outer perianth parts yellow, inner white. Fruits ellip-

Epiphyllum columbtense 287

soidal, 7-8 cm (2.8-3.1 in) long, slightly scented, red. Distribution: Costa Rica.

Epiphyllum caudatum (Vaupel) Britton 8c Rose 1913 Phyllocactus caudatus Vaupel 1913

Plants bushy, erect, semiepiphytic, to 1 m (3.3 ft) high. Primary stems cylindrical, thin. Secondary stems elongate lance shaped, narrowed basally, green, margins low and undulate, 15-20 cm (5.9-7.9 in) long, 3-4 cm (1.2-1.6 in) wide. Flowers open at night, white. Distribution: Oaxaca, Mexico. Epiphyllum caudatum is poorly known and may be simply a population of E. pumilum.

Epiphyllum columbiense (F. A. C. Weber) Dodson 8c A. H. Gentry 1977

Phyllocactus phyllanthus var. columbiensis F. A. C. Weber 1898, Epiphyllum phyllanthus var. columbiensis (F. A. C. Weber) Backeberg 1959

Plants much branched, epiphytic, with pendent branches, 1-2 m (3.3-6.6 ft) long. Primary stems round in cross section and petiole-like for 1-3 cm (0.4-1.2 in). Secondary stems delicate, linear to oblong, flattened, 7-10 cm (2.8-3.9 in) long, 1.5-4.5 cm (0.6-1.8 in) wide, marginally lobed and toothed. Areoles 2-4 cm (0.8-1.6 in) apart. Flowers salver-form, scented, white, 7-10 cm (2.8-3.9 in) long, 3.5-5.5 cm (1.4-2.2 in) in diameter; floral tubes slender. Fruits ellipsoidal, red, ridged, to 4 cm (1.6 in) long and 2 cm (0.8 in) in diameter. Distribution: Costa Rica, Panama, Colombia, Venezuela, and Ecuador.

Epiphyllum anguliger, photograph by Werner Rauh

288 Epiphyllum costar/cense

Epiphyllum costaricense (F. A. C. Weber) Britton & Rose 1913

Phyllocactus costaricensis F. A. C. Weber 1902, Epiphyllum thoma-

sianum var. costaricensis (F. A. C. Weber) Kimnach 1965 Phyllocactus macrocarpum F. A. C. Weber 1902, Epiphyllum macro-carpum (F. A. C. Weber) Backeberg 1959

Plants bushy, to 2 m (6.6 ft) long. Stems thin, green, to 30 cm (12 in) long and 7 cm (2.8 in) wide, margins slightly indented. Areoles 4-6 cm (1.6-2.4 in) apart. Flowers very large, funnelform, curved; outer perianth parts salmon colored with yellowish tips, inner white. Fruits red, slightly ribbed, to 9 cm (3.5 in) long and 7 cm (2.8 in) in diameter. Distribution: Costa Rica and Panama.

Epiphyllum crenatum (Lindley) D. Don 1855 Cereus crenatus Lindley 1844, Phyllocactus crenatus (Lindley) Lemaire 1845

Phyllocactus caulorrhizus Lemaire 1851, Epiphyllum caulorrhizum

(Lemaire) D. Don 1855 Phyllocactus cooperi Regel 1884, Epiphyllum cooperi (Regel) Clover

1941, xSeleniphyllum cooperi (Regel) G. D. Rowley 1962 Epiphyllum crenatum var. kimnachii Bravo 1964, Marniera macroptera var. kimnachii (Bravo) Backeberg 1966

Plants erect, semiepiphytic, to 1 m (3.3 ft) high. Primary stems cylindrical or three-angled, becoming woody. Secondary stems thick, leaflike, gray-green, margins scalloped and notched, to 60 cm (24 in) long and 4-10 cm (1.6-3.9 in) wide. Areoles spineless. Flowers open at night but lasting into the next day, scented, 20-29 cm (7.9-11 in) long, 10-20 cm (3.9-7.9 in) in diameter; outer perianth parts greenish or pinkish yellow, inner creamy white. Distribution: Mexico, Guatemala, and Honduras.

Two distinct varieties of Epiphyllum crenatum are recognized. Variety crenatum is distributed throughout the range

Epiphyllum crenatum var. crenatum, photograph by Roy Mottram of the species; it has stems 6-10 cm (2.4-3.9 in) wide with oblique lobes and has been used extensively as a parent in intergeneric and interspecific hybrids. Variety kimnachii has stems only 4-6 cm (1.6-2.4 in) wide and semicircular lobes; it probably comes from Oaxaca, Mexico. Variety kimnachii has a confusing history. It was long presumed to be a hybrid between Epiphyllum crenatum and Selenicereus grandiflorus and was called xSeleniphyllum cooperi, but Kimnach (1967) has shown that it is not a hybrid.

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