My first glimpse of Browningia candelaris was on the outskirts of Arequipa, Peru, but the plants were inaccessible. Some days later we visited another site near Lima and I had the opportunity to examine closely this remarkable cactus. Of the species in the genus, B. candelaris is in many ways the most spectacular, developing into large candelabra-like trees with heavily spined trunks but completely spineless upper branches.

Browningia (type, B. candelaris) was named for a single species by Nathaniel Britton and Joseph Rose in 1920, honoring W. E. Browning, who was director of the Instituto Inglés in Santiago, Chile. There has been much dispute about the boundaries of the genus. Arthur Gibson (1992) believes that, based on anatomical data, the four species often included in Gymnanthocereus form a distinct group and should be recognized as a separate genus. However, the International Cactaceae Systematics Group and other researchers generally agree that Browningia contains 11 species, which at one time or another have been included in Azureocereus, Castel-lanosia, Gymnanthocereus, and Gymnocereus. Some also placed Rauhocereus riosaniensis in Browningia as well, but Rauhocereus is now recognized as a separate genus.

Browningia is characterized as having reproductive stems less spiny than the juvenile ones, and flowers open at night and with nearly naked pericarpels and floral tubes. The cacti flower in summer. Browningia is not common in cultivation.

Browningia amstutziae 133

Browningia Britton & Rose 1920 Gymnanthocereus Backeberg 1937 Azureocereus Akers & H. Johnson 1949 Castelianosia Cárdenas 1951 Gymnocereus Backeberg 1959

Subfamily Cactoideae, tribe Browningieae. Plants shrubby or treelike, usually columnar, branched, often with well-formed trunks, to 10 m (33 ft) high. Stems cylindrical, to 50 cm (20 in) in diameter. Ribs usually numerous, low. Areoleslarge;vegetativeareolesheavilyspined; reproductive areoles usually with few or no spines. Flowers open at night, tubular to funnelform with perianth parts spreading, white to purplish red, with overlapping and conspicuous scales; areoles of pericarpels and floral tubes naked or nearly so; floral tubes curving slightly. Fruits variable, usually small. Seeds diverse. Distribution: Bolivia, northern Chile, and Peru.

Browningia albiceps F. Ritter 1981

Plants treelike, 4-5 m (13-16 ft) high, often with trunks to 2 m (6.6 ft) high. Stems 8-10 cm (3.1-3.9 in) in diameter. Ribs about 17,12-15 mm (0.5-0.6 in) high, with notched margins. Areoles round, white, 7-10 mm (0.3-0.4 in) apart. Spines on young stems awl shaped to thickly needle-like. Spines of older stems 12-20, much reduced, bristle-like, yellowish, 5-20 mm (0.2-0.8 in) long. Flowers, fruits, seeds not known. Distribution: Cajamarca, Peru.

Browningia altissima (F. Ritter) Buxbaum 1965 Gymnanthocereus altissimus F. Ritter 1959, Gymnocereus altissimus (F. Ritter) Backeberg 1962

Plants treelike, 5-10 m (16-33 ft) high with well-developed trunks. Stems cylindrical, with 7-8 ribs. Central spine usually one, pointing downward, 2-6 cm (0.8-2.4 in) long. Radial spines 5-6, pointing outward, 0.5-1 cm (0.2-0.4 in) long. Flowers greenish white, 5-6 cm (2-2.4 in) long, with crowded, broad scales. Fruits green, elongated, juicy. Distribution: near Marañón, Peru.

Browningia amstutziae (Rauh & Backeberg) Hutchison ex Krainz 1965

Gymnocereus amstutziae Rauh and Backeberg 1957

Plants treelike, to 5 m (16 ft) high. Stems grayish to dirty green, 5-6 cm (2-2.4 in) in diameter. Ribs about 11. Areoles brown, close set. Spines as many as 15, bristle-like, brittle, brown, becoming gray or black with age. Central spines 6, barely differentiated as such, pointing downward, to 4.5 cm (1.8 in) long. Radial spines 8-10, bristly, flexible, brownish, becoming gray or black with age. Flowers opening widely, cream to white, to 4.5 cm (1.8 in) long and in diameter. Distribution: Rio Paucartambo, Peru.

134 Browning/a ca/ne ana

134 Browning/a ca/ne ana

Castellanosia Caineana
Browningia altissima

Browningia caineana (Cárdenas) D. R. Hunt 1991 Castellanosia caineana Cárdenas 1951

Plants treelike, branching only basally, to 6 m (20 ft) high. Steins long cylindrical, clearly jointed, somewhat flexible, gray-green, tapering slightly, 30-40 cm (12-16 in) long, 8-11 cm (3.1-4.3 in) in diameter. Ribs 9, divided by deep constrictions into sections. Areoles gray, round, to 3 cm (1.2 in) apart. Spines of vegetative stems brown. Central spines 3-4, 4-7 cm (1.6-2.8 in) long. Radial spines 15-16, 0.8-4 cm (0.3-1.6 in) long. Spines of flowering stems about 25, forming tufts of white to brown bristles, 1-4 cm (0.4-1.6 in) long. Flowers purplish red, 3-5 cm (1.2-2 in) long, with broad, overlapping scales. Fruits globose, greenish yellow, to 3 cm (1.2 in) long. Distribution: lowland Bolivia, possibly also in Paraguay.

Browningia candelaris Britton & Rose 1920 Browningia icaensis F. Ritter 1981

Plants treelike, producing crowns of several thick, nearly spineless branches, to 6 m (20 ft) high with large trunks, un-branched, heavily spined, to 50 cm (20 in) in diameter. Stems cylindrical, often curving downward, nearly spineless when mature. Ribs about 50, bearing close-set areoles. Spines 20-50 on trunks, straight, brown, 6-15 cm (2.4-5.9 in) long. Spines few or absent on juvenile stems, yellowish brown, very thin, brownish black, to 1.3 cm (0.5 in) long. Flowering branches spineless or with only a few bristles. Flowers tubular, white, 8-12 cm (3.1-4.7 in) long. Fruits fleshy, yellow, edible, to 7 cm (2.8 in) long. Distribution: northern Chile

Browningia Cactus

Browningia caineana, photograph by Jean-Marie Solichon

Browningia chlorocarpa, photograph by JeanMarie Solichon

Browningia hertlingiana

Browningia caineana, photograph by Jean-Marie Solichon

Browningia chlorocarpa, photograph by JeanMarie Solichon

Browningia hertlingiana and southern Peru. There is concern regarding the conservation status of Browningia candelaris in some localities. Heavy grazing apparently has eliminated all seedlings, seriously threatening long-term viability of the populations.

Browningia chlorocarpa (Kunth) W. T. Marshall 1945 Cactus chlorocarpus Kunth 1823, Cereus chlorocarpus (Kunth) A. P. de Candolle 1828, Gymnanthocereus chlorocarpus (Kunth) Backeberg 1937, Seticereus chlorocarpus (Kunth) Backeberg 1959

Plants treelike, branching to form fairly dense crowns, to 1.5 m (4.9 ft) or more high with short trunks. Stems dull green, branching freely. Ribs 9-10, forming low tubercles. Areoles brownish. Spines brownish to blackish, becoming gray with age. Central spines 1-4, unequal, awl shaped, straight, stiff, to 5 cm (2 in) long. Radial spines 8-10, radiating, to 1 cm (0.4 in) long. Flowers reddish orange. Distribution: near Huanca-bamba, northern Peru. Browningia chlorocarpa is poorly known.

Browningia columnaris f. Ritter 1981

Plants solitary, columnar, branching little, blue-green, 3-5 m (9.8-16 ft) high, 8-10 cm (3.1-3.9 in) in diameter. Ribs 13-19, notched, 12-18 mm (0.5-0.7 in) high. Areoles oval to round, 10-25 mm (0.4-1 in) apart. Spines brown to grayish brown. Central spines 4, forming a cross, strongly awl shaped, 3-6 cm (1.2-2.4 in) long. Radial spines about 12, awl shaped, to 4 cm (1.6 in) long. Flowers not known. Fruits globose, indehiscent. Distribution: Rio Pampas, Ayacucho, Peru.

Browningia hertlingiana (Backeberg) F. Buxbaum 1965 Clistanthocereus hertlingianus Backeberg 1937, Azureocereus hertiin-

gianus (Backeberg) Backeberg 1956 Azureocereus nobiiis Akers 1949

Plants treelike, columnar, 5-8 m (16-26 ft) high with trunks about 1 m (3.3 ft) high from which several stems arise. Stems pale blue-green, unbranched, erect, to 30 cm (12 in) in diameter. Ribs 18 or more, tuberculate. Areoles somewhat sunken. Spines of juvenile stems yellowish gray; central spines 1-3, stout, to 8 cm (3.1 in) long; radial spines as many as 4-6. Spines of flowering stems as many as 30, flexible, yellowish, bristle-like. Flowers white, to 5 cm (2 in) in diameter. Fruits dry, 2.5 cm (1 in) in diameter. Distribution: Mantara Valley, southern Peru.

Browningia microsperma (Werdermann & Backeberg)

W. T. Marshall 1947 Cereus microspermus Werdermann & Backeberg 1931, Gymnanthocereus microspermus (Werdermann & Backeberg) Backeberg 1937,

Trichocereus microspermus (Werdermann & Backeberg) Borg 1937, Gymnocereus microspermus (Werdermann & Backeberg) Backeberg 1959

Plants treelike, 4-7 m (13-23 ft) high, with upright branches arising from trunks to 30 cm (12 in) in diameter. Stems not jointed, erect, gray-green. Ribs 12-20, rounded. Spines 30 or more, yellowish or reddish brown, pointing downward, to 1.2 cm (0.5 in) long; some areoles bear single long spines. Flowers white, to 6 cm (2.4 in) long and in diameter. Fruits green, 5-6 cm (2-2.4 in) in diameter. Distribution: northern Peru.

Browningia pilleifera (F. Ritter) Hutchison 1968 Gymnanthocereus pilleifera F. Ritter 1966 Gymnanthocereus macracanthus F. Ritter 1981

Plants treelike, much branched, 3-4 m (9.8-13 ft) high. Ribs 7-9, triangular or rounded, with areoles becoming confluent

Azureocereus Viridis
Browningia candelaris, also illustrated on page 76; photograph by James Mauseth

on older stems. Spines present on juvenile stems, becoming gray. Central spines 0-3, to 50 mm (2 in) long. Radial spines 0-3,1-5 mm long. Spines absent on mature stems. Flowers white, covered with dry scales, 3.2-3.5 cm (1.3-1.4 in) long. Fruits club shaped, green, to 2 cm (0.8 in) in diameter. Distribution: Balsas, along the Maranon, Peru.

Browningia viridis (Rauh & Backeberg) F. Buxbaum 1965 Azureocereus viridis Rauh & Backeberg 1957

Plants treelike, candelabra-like with several erect branches, to 10 m (33 ft) high with distinct trunks. Stems green, to 8 cm (3.1 in) in diameter. Ribs 18 or more, thickened around the areoles, sometimes becoming tuberculate. Spines 10-20, yellow with dark tips, to 7 cm (2.8 in) long. Flowers white, to 5 cm (2 in) in diameter. Distribution: Apurimac Valley, Peru. Browningia viridis appears to be closely related to B. hertlin-giana.

Cacti Cultivation

Browningia viridis, photograph by Jean-Marie Solichon

Browningia pilleifera

Browningia viridis, photograph by Jean-Marie Solichon

Browningia pilleifera

Carnegiea 137

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