Pilosocereus

It is fortunate when a hitherto poorly understood group of cacti is the object of a major research project. Such is the case with Pilosocereus, a genus of columnar cacti found from Mexico into South America. Daniela Zappi (1994) undertook such a study for her doctoral degree at the University of Sao Paulo, which involved fieldwork and studies of vegetative and seed morphology, and reproductive biology.

The name Pilosocereus is relatively recent, having been proposed by Ronald Byles and Gordon Rowley in 1957. Many species had been described long before but placed in Pilocereus, described by Charles Lemaire in 1839. Unfortunately, the type species of Lemaire's genus is the same as that of Cephalocereus, described a year earlier by Ludwig Pfeiffer: Cactus senilis = Cephalocereus senilis. Thus Pilocereus is nomenclaturally illegitimate. An attempt was made by Karl

Gordon Rowley

Pereskiopsis rotundifolia, photograph by Gordon Rowley

Schumann to redefine Pilocereus without Lemaire's type but that only resulted in another illegitimate name. A proposal was made in 1954 to the International Botanical Congress to retain Pilocereus by conserving that name, but it failed to pass. The solution was to coin a new name, which was done by Byles and Rowley (1957) in describing PHosocereus (type, P. leucocephalus) to include the 58 species formerly in Pilocereus. Many additional species have since been described but many appear to be frivolous and have been relegated to synonymy. Zappi suggests that the genus probably comprises 36 species and a naturally occurring hybrid.

The name PHosocereus is derived from the Latin pihsus, hairy, thus hairy cereus, similar to the Greek pilos, felted, from which the name Pilocereus was derived. PHosocereus is characterized as having abundantly woolly flowering areoles near the tips of the stems that produce night-flowering, bat-pollinated blossoms in summer. The flowers have fleshy, naked pericarpels and floral tubes.

PHosocereus Byles & G. D. Rowley 1957

Pilocereus K. Schumann 1894, illegitimate name, not Lemaire 1839

(see Cephalocereus) Pseudopilocereus Buxbaum 1968

Subfamily Cactoideae, tribe Cereeae. Plants shrubby or treelike, usually branched above the ground, to 10 m (33 ft) high. Stems green to gray to waxy blue, usually smooth, highly mucilaginous. Ribs 3-30, sinuses usually straight, often with cross-furrows. Areoles with felt, producing abundant woolly hairs to 5 cm (2 in) long near the stem tips and often coveringthe ribs, sometimes forming lateral cephalia. Spines variable, straight to rarely curved, sometimes translucent. Flower buds at acute or obtuse angles priorto expansion and opening of the flower. Flowers open in the evening, tubularto bell shaped, bat pollinated, white or pale pink, usually smooth and lacking areoles, 2.5-9 cm (1-3.5 in) long, 2-7 cm (0.8-2.8 in) in diameter; peri-carpelsandfloraltubesfleshyandlackingobvious bracts, often tinged with brown or purple; stamens numerous. Fruits globose to depressed globose, smooth, blackening, usually pendent, dehiscing by various means, pulp of varying colors; perianth parts persistent. Seeds dark brown or black, 1.2-2.5 mm long, snail shaped. Distribution: Mexico, Caribbean, and south throughout much of tropical South America in Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, and Paraguay.

PHosocereus albisummus P. J. Braun & Esteves 1987

Plants shrubby or treelike, to 2.8 m (9.2 ft) high, branching above ground level with well-defined trunks. Stems erect, gray-green, to 8.5 cm (3.3 in) in diameter. Ribs 6-12. Spines pale yellow or reddish, becoming grayish or black. Central spines 3-7, erect to bent downward, 5-10 mm (0.2-0.4 in)

long. Radial spines 11-13, flattened against the stem surface, 5-7 mm (0.2-0.3 in) long. Fertile portions of stems evident, with flower-bearing areoles on every rib near the stem tips, with grayish to brownish hairs to 22 mm (0.9 in) long and spines to 41 mm (1.6 in) long. Flower buds obtuse. Flowers silvery white to purplish, to 4.3 cm (1.7 in) long and 3.5 cm (1.4 in) in diameter. Fruits depressed globose, to 2.5 cm (1 in) long, dehiscing by lateral slits, pulp magenta. Distribution: western Minas Gerais, Brazil. PHosocereus albisummus is poorly known.

PHosocereus alensis (F. A. C. Weber ex Roland-Gosselin)

Byles & G.D.Rowley 1957 barba de viejo, pitahaya barbona

Pilocereus alensis F. A. C. Weber ex Roland-Gosselin 1905, Cephalocereus alensis (F. A. C. Weber ex Roland-Gosselin) Britton & Rose 1909 Pilocereus guerreronis Backeberg 1941, PHosocereus guerreronis (Backeberg) Byles & G. D. Rowley 1957

Plants treelike, branching from the base, 5-6 m (16-20 ft) high. Stems erect, hard, to 12 cm (4.7 in) in diameter. Ribs 14, tuberculate, minutely notched. Spines yellowish, becoming reddish, then gray. Central spine one. Radial spines 11-13, needle-like, to 20 mm (0.8 in) long, lowermost one deflexed and longest. Fertile portions of stems forming lateral cephalia with dense white hairs. Flowers appearing to be light purple. Fruits not known. Distribution: Chihuahua, Durango, Guerrero, Michoacán, México, Jalisco, and Sonora, Mexico. PHosocereus alensis has been studied too little.

PHosocereus albisummus

576 Pilosocereus alensis

576 Pilosocereus alensis

Pilocereus Cactus
Pilosocereus alensis

Pilosocereus arrabidae (Lemaire) Byles & G. D. Rowley 1957

facheiro da praia

Cactus heptagonus \lellozo 1829, not Linnaeus 1753 Cactus hexagonus Vellozo 1829, not Linnaeus 1753 (see Cereus hexa-gonus)

Pilocereus arrabidae Lemaire 1862, Cephalocereusarrabidae (Lemaire) Britton & Rose 1920, Pseudopilocereus arrabidae (Lemaire) Buxbaum 1968 Cereus warmingii K. Schumann 1890

Plants shrubby, 1-4 m (3.3-13 ft) high, often lacking trunks, branching near the base. Stems straight, slightly curved upward, yellow-green to dark green, 4.5-9.5 cm (1.8-3.7 in) in diameter, only slightly woody. Ribs 5-8, transverse folds oblique. Areoles produced on conspicuous podaria, hairs present or absent. Spines translucent at first, becoming brown to grayish. Central spines usually 2-4, often one ascending and two erect, 15-40 mm (0.6-1.6 in) long. Radial spines 7-10,2-20 mm (to 0.8 in) long, flattened against the stem surface. Fertile portions not differentiated. Flower-

bearing areoles along the stems. Flower buds acute. Flowers funnelform, 6-7 cm (2.4-2.8 in) long, 4-5 cm (1.6-2 in) in diameter. Fruits depressed globose, dehiscing by lateral slits,

3-5 cm (1.2-2 in) long, pulp magenta. Distribution: Rio de Janeiro, Espirito Santo, and southeastern Bahia, Brazil, from sea level to 100 m (330 ft).

Pilosocereus aureispinus (Burning & Brederoo) F. Ritter 1979

Coleocephalocereus aureispinus Burning & Brederoo 1974

Plants shrubby, to 2 m (6.6 ft) high, branching only basally. Stems erect, dark green, smooth, 5-9 cm (2-3.5 in) in diameter, weakly woody. Ribs 20-24, without transverse folds. Spines translucent to golden to rust colored, sometimes bristly. Central spines 8-16, ascending, 4-12 mm (to 0.5 in) long. Radial spines 14-16, flattened against the stem surface,

4-12 mm (to 0.5 in) long. Fertile portions of stems poorly differentiated. Flower-producing areoles lateral or forming rings around the stems, with sparse white hairs and golden bristles to 50 mm (2 in) long. Flower buds acute. Flowers narrow, externally dark, white, to 5 cm (2 in) long, 2-2.2 cm (0.8-0.9 in) in diameter. Fruits globose to depressed globose, 2-2.6cm (0.8-1 in) long, dehiscing by lateral slits, pulp white. Distribution: central Bahia, Brazil, at elevations of 450-550 m (1500-1800 ft).

Pilosocereus aurisetus (Werdermann) Byles 8c G. D. Rowley 1957

quiabo da lapa, quiab0 do inferno, rabo de raposa Pilocereus aurisetus Werdermann 1933, Pseudopilocereus aurisetus

(Werdermann) Buxbaum 1968 Pseudopilocereus werdermannianus Buining & Brederoo 1975, Pilosocereus werdermannianus (Buining & Brederoo) F. Ritter 1979, Pilosocereus aurisetus subsp. werdermannianus (Buining & Brederoo) P.J. Braun &Esteves 1995 Pilosocereus aurilanatus F. Ritter 1979, P. aurisetus subsp.aurilana-tus (F. Ritter) Zappi 1994, Pseudopilocereus aurilanatus (F. Ritter) P. V. Heath 1994

Pilosocereus werdermannianus var. densilanatus F. Ritter 1979, P. aurisetus subsp. densilanatus (F. Ritter) P. J. Braun & Esteves 1995 Pilosocereus supthutianus P. J. Braun 1985, P. aurisetus subsp. supthutianus (P. J. Braun) P. J. Braun & Esteves 1995

Plants shrubby, 1-3 m (3.3-9.8 ft) high, branching only basally. Stems erect, weakly woody, 2.8-7 cm (1.1-2.8 in) in diameter. Ribs 10-17, transverse folds absent. Spines 17-25, translucent, white to golden yellow. Fertile portions of stems strongly differentiated, lateral or subapical. Flower-bearing areoles with abundant long hairs and golden bristles. Flower buds acute. Flowers funnelform, reddish, 3.2-5 cm (1.3-2 in) long, 3-4 cm (1.2-1.6 in) in diameter. Fruits depressed globose, 2-3.5 cm (0.8-1.4 in) long, dehiscing by central slits, pulp white. Distribution: Minas Gerais, Brazil.

Two subspecies of Pilosocereus aurisetus are recognized by Zappi (1994). Subspecies aurisetus has white hairs on its flower-producing areoles, and stems to 2 m (6.6 ft) long and 2.8-5.5 cm (1.1-2.2 in) in diameter; it occurs in the Serra do Espinhac^o at elevations of650-1300 m (2100-4300 ft). Subspecies aurilanatus has golden hairs on its flower-producing areoles, and stems to 3 m (9.8 ft) high and 4.5-7 cm (1.8-2.8 in) in diameter; it occurs in the Serra do Cabral at 800-900 m (2600-2950 ft).

Pilosocereus azulensis N. P. Taylor & Zappi 1997

Plants treelike or shrubby, branching above the base, to 10 m (33 ft) high. Stems olive green to glaucous, smooth, 8-9.5 cm (3.1-3.7 in) in diameter. Ribs 6-12, sometimes with transverse folds. Areoles 5-7 mm (0.2-0.3 in) apart. Spines slender, yellowish brown, opaque. Central spines 1-11,5-40 mm (0.2-1.6 in), ascending to erect. Radial spines 8-12, flattened against the stem surface, 3-13 mm (to 0.5 in) long. Flower-producing portions only slightly modified, on apical and subapical areoles of one to three ribs. Flower-bearing areoles with white to gray hairs, to 3 cm (1.2 in) long. Flower buds acute. Flowers opening widely, pale green externally, 5.5-6.7 cm (2.2-2.6 in) long, 4.7-7 cm (1.9-2.8 in) in diameter. Fruits depressed globose, 4.2-6 cm (1.7-2.4 in) in diameter, dehiscing by lateral slits, pulp magenta. Distribution: municipality of Pedra Azul, Minas Gerais, Brazil.

Pilosocereus brasiliertsis (Britton & Rose) Backeberg 1960 Cephalocereus brasiliensis Britton & Rose 1920, Pilocereus brasilien-

sis (Britton & Rose) Werdermann 1933 ?Cereus sublanatus Salm-Dyck 1834, ? Pilocereus sublanatus (SalmDyck) Förster ex Backeberg & F. M. Knuth 1935, ?Pilosocereus sublanatus (Salm-Dyck) Byles & G. D. Rowley 1957 Pseudopilocereus ruschianus Buining & Brederoo 1980, Pilosocereus ruschianus (Buining & Brederoo) P. J. Braun 1988, Pilosocereus brasiliensis subsp. ruschianus (Buining & Brederoo) Zappi 1994

Plants erect or climbing shrubs, 2-4 m (6.6-13 ft) high with few branches, without well-defined trunks. Stems green to gray-green, 4.5-5.5 cm (1.8-2.2 in) in diameter, moderately woody. Ribs 4-6, transverse folds oblique. Areoles located on podaria, with long hairs. Spines reddish or yellowish brown, becoming gray. Central spine one, erect, 10-20 mm (0.4-0.8 in) long. Radial spines several, spreading, very short. Fertile portions of stems only slighdy differentiated, with flower-bearing areoles occurring randomly over the stems. Flower buds obtuse. Flowers narrowly funnelform, white, to 4.5 cm (1.8 in) long, 2.5-3 cm (1-1.2 in) in diame ter. Fruits depressed globose, to 2 cm (0.8 in) long, dehiscing by lateral slits, pulp magenta. Distribution: Espirito Santo, Rio de Janeiro, Bahia, and Minas Gerais, Brazil.

Two subspecies of Pilosocereus brasiliensis are recognized. Subspecies brasiliensis has dark green branches with 4-5 ribs, and the stem tips usually lack very woolly areoles; it occurs from sea level to 100 m (330 ft) along the coast of Espirito Santo and Rio de Janeiro. Subspecies ruschianus has gray-green stems with 5-6 ribs, and the stem tips have very woolly areoles; it occurs at 80-700 m (260-2300 ft) in southern Bahia, Espirito Santo, and eastern Minas Gerais.

Pilosocereus catingicola (Gürke) Byles & G. D. Rowley 1957

fach el ro

Cereus catingicola Gürke 1908, Cephalocereus catingicola (Gürke) Britton & Rose 1920, Pilocereus catingicola (Gürke) Werdermann 1933, Pseudopilocereus catingicola (Gürke) Buxbaum 1968 Pilocereus arenicola Werdermann 1933, Pilosocereus arenicola (Werdermann) Byles & G. D. Rowley 1957, Pilosocereus catingicola subsp. arenicola (Werdermann) P. J. Braun & Esteves 1995 Pilocereus hapaiacanthus Werdermann 1933, Pilosocereus hapala-canthus (Werdermann) Byles & G. D. Rowley 1957, Pseudopilocereus hapaiacanthus (Werdermann) Buxbaum 1968, Pilosocereus catingicola subsp. hapaiacanthus (Werdermann) P.J. Braun & Esteves 1995

Pilocereus rupicola Werdermann 1933, Pilosocereus rupicola (Werdermann) Byles & G. D. Rowley 1957, Pseudopilocereus rupicola (Werdermann) Buxbaum 1968 Pilocereus salvadorensis Werdermann 1933, Pilosocereus salvadoren-sis (Werdermann) Byles & G. D. Rowley 1957, Austrocephalocereus salvadorensis (Werdermann) Buxbaum 1966, Pseudopilocereus salvadorensis (Werdermann) Buxbaum 1968, Pilosocereus catingicola subsp. salvadorensis (Werdermann) Zappi 1994 Pilocereus sergipensis Werdermann 1933, Pilosocereus sergipensis (Werdermann) Byles & G. D. Rowley 1957, Pseudopilocereus sergipensis (Werdermann) Buxbaum 1968 Pilosocereus robustus F. Ritter 1979, P. catingicola subsp. robustus (F. Ritter) P. J. Braun & Esteves 1995, Pseudopilocereus robustus (F. Ritter) P. V. Heath 1994

Plants treelike or shrubby, branching above the base, 3-10 m (9.8-33 ft) high. Stems olive green to glaucous, smooth, 3.512 cm (1.4-4.7 in) in diameter. Ribs 4-12, sometimes with transverse folds. Spines yellowish brown, opaque. Central spines 1-11, ascending to erect, 2-40 mm (to 1.6) in. Radial spines 8-12, flattened against the stem surface, 3-13 mm (to 0.5 in) long. Flower-producing portions only slightly modified, on apical and subapical areoles of one to three ribs. Flower-bearing areoles with white to gray hairs, to 3 cm (1.2 in) long. Flower buds acute. Flowers opening widely, pale green externally, 5.5-6.7 cm (2.2-2.6 in) long, 4.7-7 cm (1.9-2.8 in) in diameter. Fruits depressed globose, 4.2-6 cm

(1.7-2.4 in) in diameter, dehiscing by lateral slits, pulp magenta. Distribution: caatinga vegetation of Bahia north into Sergipe, Alagoas, Pernambuco, Rio Grande do Norte, and Ceara, Brazil.

Two subspecies of Pilosocereus catingicola are recognized by Zappi (1994). Subspecies catingicola has stems 8-12 cm (3.1-4.7 in) in diameter with 4-6 ribs, and stout central spines 10-40 mm (0.4-1.6 in) long; it occurs at elevations

Pilocereus
Pilosocereus chrysacanthus

of 200-850 m (660-2800 ft) throughout much of Bahia. Subspecies salvadorensis has stems 3.5-6 cm (1.4-2.4 in) in diameter with 6-12 ribs, and slender central spines 2-10 mm (to 0.4 in) long; it occurs in Sergipe, Alagoas, Pernambuco, Rio Grande do Norte, Ceara, and Bahia.

Pilosocereus chrysacanthus (F. A. C. Weber) Byles 8c G. D.

Rowley 1957 GOLDEN OLD MAN, G0LDEN-SPINED CACTUS Pilocereus chrysacanthus F. A. C. Weber 1897, Cephalocereus chrysacanthus (F. A. C. Weber) Britton & Rose 1909

Plants treelike, 3-4 m (9.8-13 ft) high, branching mostly from the base. Stems more or less erect, glaucous green. Ribs 9-12. Spines 12-15, yellowish, becoming darker with age, the longest 3—4 cm (1.2-1.6 in). Flower-producing portions well defined, located apically and on one side of each stem. Flower-bearing areoles with long white hairs and yellow spines. Flowers with a rose tint, 7-8 cm (2.8-3.1 in) long. Fruits globose, 3-4 cm (1.2-1.6 in) in diameter, pulp purple. Distribution: Puebla and Oaxaca, Mexico.

Pilosocereus chrysostele (Vaupel) Byles 8c G. D. Rowley 1957

FACHEIRO, RABO DE RAPOSA

Cereus chrysostele Vaupel 1923, Pilocereus chrysostele (Vaupel) Werdermann 1933, Pseudopilocereus chrysostele (Vaupel) Buxbaum 1968

Plants shrubby, 1.5-3 m (4.9-9.8 ft) high, branching only basally. Stems erect, pale green, smooth, 4-7 cm (1.6-2.8 in) in diameter. Ribs 22-23, without transverse folds. Spines translucent pale yellow. Central spines 6-8, ascending, 5-12 mm (0.2-0.5 in) long. Radial spines 9-12, flattened against the stem surface, 4-10 mm (to 0.4 in) long. Flowering portions strongly differentiated. Flower-bearing areoles on lateral and subapical parts of the stems, with white hairs to 20 mm (0.8 in) long and golden brisdes 40-60 mm (1.6-2.4 in) long. Flower buds obtuse. Flowers stout, short, pinkish or brownish, 4.5-5 cm (1.8-2 in) long, 3.7 cm (1.5 in) in diameter. Fruits depressed globose, 3-3.5 cm (1.2-1.4 in) in diameter, dehiscing by lateral or adaxial slits, pulp magenta. Distribution: northeastern Brazil near the borders of Ceara, Paraiba, and Pernambuco at elevations of 430-600 m (1400-2000 ft).

Pilosocereus densiareolatus F. Ritter 1979

CABEQA DE VELHO, FACHEIRO DA LAPA

Pseudopilocereus densiareolatus (F. Ritter) P. V. Heath 1994

Plants treelike, 2-6 m (6.6-20 ft) high with well-defined trunks. Stems erect, olive green to bluish, glaucous, weakly

Pilosocereus catingicola subsp. catingicola, photograph by Urs Eggli

PHosocereus floccosus 579

woody, smooth, 3-7 cm (1.2-2.8 in) in diameter. Ribs 13-19, without transverse folds. Spines translucent, brown to golden, becoming gray with age. Central spines 6-15, ascending,

9-35 mm (0.4-1.4 in) long. Radial spines 14-16, flattened against the stem surface, 4-7 mm (to 0.3 in) long. Flowering portions of the stems slightly to well differentiated. Flower-bearing areoles forming lateral cephalia with white to gray to brown hairs 30-80 mm (1.2-3.1 in) long and golden bristles. Flower buds obtuse. Flowers short, tubular, appearing olive green on the outside, 5-6 cm (2-2.4 in) long, to 4 cm (1.6 in) in diameter. Fruits depressed globose, 2-3 cm (0.81.2 in) long, dehiscing by lateral or adaxial slits, pulp magenta. Distribution: northern Minas Gerais and western Bahia, Brazil, at elevations of 450-800 m (1500-2600 ft).

Pilosocereus diersianus (Esteves) P. J. Braun 1988 Pseudopilocereus diersianus Esteves 1981

Plants shrubby, to 2 m (6.6 ft) high, branching above the base. Stems erect, green to gray-green, 5-9 cm (2-3.5 in) in diameter. Ribs 19-22, transverse folds not visible. Spines golden brown to gray-brown. Central spines 10, ascending, 5-60 mm (0.2-2.4 in) long. Radial spines 10-20, flattened against the stem surface. Flower-producing portion strongly differentiated, on lateral and subapical parts. Flower-bearing areoles with white hairs to 30 mm (1.2 in) long and yellow bristles to 10 cm (3.9 in) long. Flowers stout, flared, 3.5-5 cm (1.4-2 in) long, 3-3.5 cm (1.2-1.4 in) in diameter. Fruits depressed globose, 2-3 cm (0.8-1.2 in) long, dehiscing by lateral slits, pulp magenta. Distribution: limestone rocks in northeastern Goias, Brazil, at elevations of600-700 m (20002300 ft).

Pilosocereus estevesii p. j. Braun 1999

Plants treelike, often candelabra-like, to 3.2 m (10.5 ft) high, with erect columnar branches. Stems erect, green to greenish blue to greenish gray, 2.5-6.5 cm (1-2.6 in) in diameter. Ribs

10-13, with straight grooves between, slightly depressed between areoles. Areoles round to oval, to 1.3 cm (0.5 in) apart. Spines needle-like, straight, radiating, light yellow to brownish to grayish. Central spines 1-7 but variable in number. Radial spines 13-17, to 16 mm (0.6 in) long. Flower-producing portion well differentiated as an intermittent, compact, circular, ring-shaped structure. Flower-bearing areoles with brownish to grayish bristles to 47 mm (1.9 in) long. Flowers open at night, more or less funnelform, to 4.7 cm (1.9 in) long, whitish to pale cream. Fruits depressed globose to globose, rosy cream with dark rose pulp, to 3.6 cm (1.4 in) in diameter, dehiscing by lateral slits. Distribution: Serra do Muquem, Bahia, Brazil.

Pilosocereus flavipulvinatus (Burning & Brederoo) F. Rit ter 1980

Pseudopilocereus flavipulvinatus Buining & Brederoo 1979 Pilosocereus carolinensis F. Ritter 1979, Pseudopilocereus carolinen-sis (F. Ritter) P. V. Heath 1994, Pilosocereus flavipulvinatus subsp. carolinensis (F. Ritter) P. J. Braun & Esteves 1995

Plants treelike, slender, sparsely branched, 2-8 m (6.6-26 ft) high, often leaning on other vegetation, with well-defined trunks. Stems erect to inclined, olive green, glaucous at the tips, 3-9 cm (1.2-3.5 in) in diameter. Ribs 6-9, transverse folds oblique. Areoles located on rounded stem projections, with yellowish felt. Spines translucent, pale yellow, becoming gray, ascending. Central spines 3-5, erect, 15-30 mm (0.61.2 in) long,lowermost one longest. Radial spines 12-15, flattened against the stem surface, 5-14 mm (0.2-0.6 in) long. Fertile portions not differentiated. Flower-bearing areoles subapical, on two to four ribs. Flower buds acute. Flowers funnelform, 5.8-6 cm (2.3-2.4 in) long, 3.5-3.7 cm (1.4-1.5 in) in diameter. Fruits depressed globose, 3-3.5 cm (1.2-1.4 in) in diameter, dehiscing by lateral slits, pulp magenta. Distribution: dense caatinga vegetation at elevations of 120-200 m (390-650 ft) along the border of Piaui and Maranhao, Brazil.

Pilosocereus flexibilispinus P. J. Braun & Esteves 1990

Plants treelike, to 8 m (26 ft) high, branched above the ground, with well-defined trunks. Stems erect, gray-green to bluish, 5-12 cm (2-4.7 in) in diameter. Ribs 9-11, high and narrow, transverse folds oblique. Spines yellow to brownish red. Central spines 5-11, erect to bent away, 17-31 mm (0.7-1.2 in) long. Radial spines 14-20, flattened against the stem surface, 11 mm (0.4 in) long. Fertile portions strongly differentiated. Flower-bearing areoles near the stem tips, on most ribs, with gray hairs to 23 mm (0.9 in) long and flexible spines to 75 mm (3 in) long. Flower buds obtuse. Flowers flared, to 4.8 cm (1.9 in) long and 2.8 cm (1.1 in) in diameter. Fruits globose to slightly depressed globose, 3.6 cm (1.4 in) in diameter, dehiscing by lateral slits, pulp red. Distribution: limestone at an elevation of about 600 m (2000 ft) in Tocantins, Brazil.

Pilosocereus floccosus Byles & G. D. Rowley 1957 Pseudopilocereus floccosus (Byles & G. D. Rowley) Buxbaum 1968 Pilosocereus quadricostatus F. Ritter 1979, P. floccosus subsp. quadricostatus (F. Ritter) Zappi 1994, Pseudopilocereus quadricostatus (F. Ritter) P. V. Heath 1994

Plants variable in habit, shrubby or treelike, sometimes branching above the ground with few or many branches, 1-5 m (3.3-16 ft) high with well-defined trunks. Stems erect, rough and warty, gray-green, 5-11 cm (2-4.3 in) in diameter.

580 Pilosocereus floccosus

Ribs 4-8, transverse folds not visible. Spines opaque, yellowish brown or reddish, becoming gray, not differentiated as centrals and radials. Fertile portions strongly differentiated. Flower-bearing areoles forming subapical crowns or rings around the stems, with pale brown or gray woolly hairs 15-20 mm (0.6-0.8 in) long and dark bristly spines to 35 mm (1.4 in) long. Flower buds obtuse. Flowers flared, 4-5 cm (1.6-2 in) long, to 3 cm (1.2 in) in diameter. Fruits depressed globose or depressed laterally, 3-5 cm (1.2-2 in) in diameter, dehiscing by lateral, abaxial, or adaxial slits, pulp bright red. Distribution: limestone and gneiss outcrops in Minas Gerais, Brazil.

Two subspecies of Pilosocereus floccosus are recognized. Subspecies floccosus has stems 5-9 cm (2-3.5 in) in diameter with 5-8 ribs; it occurs on limestone in the Serra do Espin-ha^o at elevations of 600-850 m (2000-2800 ft). Subspecies quadricostatus has stems 8-11 cm (3.1-4.3 in) in diameter with 4-5 ribs; it occurs on gneiss at 250-800 m (820-2600 ft).

Pilosocereusfulvilanatus (Buining & Brederoo) F. Ritter 1979

Pseudopilocereus fulvilanatus Buining & Brederoo 1973 Pilosocereus rosae P. J. Braun 1984, P. fulvilanatus subsp. rosae (P. J. Braun) Zappi 1994

Plants shrubby to treelike, branching above ground, 2-4 m (6.6-13 ft) high. Stems erect, gray-green, 5-12 cm (2-4.7 in) in diameter, with intense wax coating near the tips. Ribs 4-8, transverse folds absent. Areoles nearly contiguous. Spines opaque, yellow-brown to black, thickened basally. Central spines 1-7,10-30 mm (0.4-1.2 in) long. Radial spines 8-10,

Pilosocereus fulvilanatus subsp. fulvilanatus flattened against the stem surface, 8-15 mm (0.3-0.6 in) long. Fertile portions strongly differentiated. Flower-bearing areoles subapical or lower, on one to several ribs, with abundant golden brown hairs 20-40 mm (0.8-1.6 in) long. Flowers funnelform, 3.6-6 cm (1.4-2.4 in) long, 3-4 cm (1.2-1.6 in) in diameter. Fruits depressed globose, 3.5-6.5 cm (1.4-2.6 in) in diameter, dehiscing by lateral slits, pulp magenta. Distribution: Minas Gerais, Brazil.

Two subspecies of Pilosocereus fulvilanatus are recognized. Subspecies fulvilanatus has stems 8-12 cm (3.1-4.7 in) in diameter, 4-7 ribs, and 1-7 central spines; it is found at elevations of 720-1000 m (2400-3300 ft) in the drainage of the Rio Jequitinhonha of the Serra do Espinhaco. Subspecies rosae has stems to only 5.5 cm (2.2 in) in diameter, 6-8 ribs, and 3-4 central spines; it is found at 800 m (2600 ft) in the drainage of the Rio das Velhas of the Serra do Espinhaco.

Pilosocereus glaucochrous (Werdermann) Byles & G. D. Rowley 1957

Pilocereus glaucochrous Werdermann 1933, Pseudopilocereus glaucochrous (Werdermann) Buxbaum 1968

Plants treelike, sparsely branched, 3-5 m (9.8-16 ft) high with well-defined trunks. Stems tending to lean, blue-green to gray-green, glaucous, appearing silverish, 3-7 cm (1.2-2.8 in) in diameter. Ribs 5-10, transverse folds present. Areoles on low, rounded podaria, with blackish felt and white hairs. Spines translucent when young, golden yellow to gray. Central spines 2-5, ascending, 20-40 mm (0.8-1.6 in) long. Radial spines 7-14, mostly spreading, flattened against the stem surface, 8-15 mm (0.3-0.6 in) long. Fertile portions not or only slightly differentiated, apical to subapical. Flower buds obtuse. Flowers flared, pinkish to reddish in appearance, 4— 5.2 cm (1.6-2 in) long, to 2.5 cm (1 in) in diameter. Fruits depressed globose, dehiscing by lateral slits, pulp purplish. Distribution: highland caatinga vegetation, central Bahia, Brazil.

Pilosocereusgounellei (F. A. C. Weber) Byles & G. D. Rowley 1957

alastrado, chique-chique, xiquixique, xique-xique, xique-xique das pedras

Pilocereus gounellei F. A. C. Weber 1897, Cephalocereus gounellei (F. A. C. Weber) Britton & Rose 1920, Cereusgounellei (F. A. C. Weber) Lutzelburg 1926, Pseudopilocereus gounellei (F. A. C. Weber) Buxbaum 1968 Pilocereus setosus Gurke 1908

Cephalocereuszehntneri Britton & Rose 1920, Cereus zehntneri (Britton & Rose) Lutzelburg 1926, Pilosocereus zehntneri (Britton & Rose) F. Ritter 1979, P. gounellei subsp .zehntneri (Britton & Rose) Zappi 1994, Pseudopilocereus zehntneri (Britton & Rose) P. V. Heath 1994

Cacto Chique Chique

Pitosocereus lanuginosus 581

Pseudopilocereussuperfloccosus Buining & Brederoo 1974, Pitosocereus superfloccosus (Buining & Brederoo) F. Ritter 1979 Pilosocereus braunii Esteves 1989

Plants shrubby, rarely treelike, 0.5-4 m (1.6-13 ft) high with numerous branches arising from the main stem. Stems erect, becoming inclined or even horizontal with age, 3.7-9 cm (1.5-3.5 in) in diameter, new stems arising subapically, olive green, sometimes glaucous. Ribs 8-15, with sinuate sinuses, transverse folds present. Spines opaque to translucent, golden to reddish to brown to gray. Central spines 1-10. Radial spines 10-20, flattened against the stem surface. Fertile portions subapical, weakly to strongly differentiated. Flower buds obtuse, completely encircled by long silky hairs. Flowers funnelform, 4-9 cm (1.6-3.5 in) long, 2.5-6 cm (1-2.4 in) in diameter. Fruits globose to depressed, 4.5-6 cm (1.82.4 in) in diameter, dehiscing by adaxial or abaxial slits. Distribution: northeastern Brazil.

Two subspecies of Pilosocereus gounellei are recognized. Subspecies gounellei has strong, stout, brownish to grayish, opaque spines with the centrals distinctly longer than the ra-dials; it occurs throughout northeastern Brazil. Subspecies zehntneri has slender, fragile, translucent, golden to reddish spines with the centrals about the same length as the radials; it is found only in northern central Minas Gerais to northern Bahia.

Pilosocereus lanuginosus (Linnaeus) Byles & G. D. Rowley 1957

Cactus lanuginosus Linnaeus 1753, Cephalocereus lanuginosus (Linnaeus) Britton & Rose 1909 Cereus moritzianus Otto ex Pfeiffer 1837, Cephalocereus moritzianus (Otto ex Pfeiffer) Britton & Rose 1920, Pilosocereus moritzianus (Otto ex Pfeiffer) Byles & G. D. Rowley 1957 Cephalocereus colombianus Rose 1909, Pilosocereus colombianus

(Rose) Byles & G. D. Rowley 1957 Cephalocereus tweedyanus Britton & Rose 1920, Pilosocereus tweedyanus (Britton & Rose) Byles & G. D. Rowley 1957 Cereus backebergii Weingart 1930, Pilosocereus backebergii (Weingart) Byles & G. D. Rowley 1957 Cereus claroviridis Backeberg 1931, Pilosocereus claroviridis (Backeberg) Byles & G. D. Rowley 1957 Pilosocereus gironensis Rauh & Backeberg ex Byles & G. D. Rowley 1957

?Pilosocereus tuberculosus Rauh & Backeberg 1957 Pilosocereus tillianus R. Gruber & Schatzl 1982

Plants treelike, ascending or spreading with flexible branches, to 10 m (33 ft) high with well-defined trunks. Stems erect, strongly blue at first, becoming blue-green and glaucous, 6-10 cm (2.4-3.9 in) in diameter. Ribs 9-13, transverse folds present. Spines black with red, later gray. Central

Pilosocereus glaucochrous, photograph by Urs Eggli

Pitosocereus lanuginosus 581

Pilosocereus glaucochrous, photograph by Urs Eggli

582 Pilosocereus lanuginosus spines 1-4, 2-7 cm (0.8-2.8 in) long, one distinctly longer and pointing downward. Radial spines 9-15,1-2 cm (0.40.8 in) long. Fertile portions distinct, involving two or three ribs, interrupted, with spines and white or light brown wool 2-3 cm (0.8-1.2 in) long. Flowers bell shaped, 6-7.5 cm (2.4-3 in) long, to 4 cm ( 1.6 in) in diameter. Fruits depressed globose, pulp purplish. Distribution: Curaçao and throughout northern South America in Colombia, Trinidad, Venezuela, Ecuador, and Peru.

Pilosocereus leucocephalus (Poselger) Byles & G. D. Rowley 1957

cabeza de viejo, old man cactus, old man of mexico, ôrgan0, pitayôn, tuno

Pilocereus leucocephalus Poselger 1853, Cephalocereus leucocephalus (Poselger) Britton & Rose 1909 ?Cereus cometes Scheidweiler 1840, ? Cephalocereus cometes (Scheidweiler) Britton & Rose 1909, ?Pilosocereus cometes (Scheidweiler) Byles & G. D. Rowley 1957 Cephalocereus maxonii Rose 1909, Pilosocereus maxonii (Rose) Byles

& G.D.Rowley 1957 Cephalocereus palmeri Rose 1909, Pilosocereus palmeri (Rose) Byles

& G.D.Rowley 1957 Cephalocereus sartorianus Rose 1909, Pilosocereus sartorianus

(Rose) Byles & G. D. Rowley 1957 Pilocereus tehuacanus Weingart 1927, Pilosocereus tehuacanus (Weingart) Byles & G. D. Rowley 1957

Plants treelike, 2-5 m (6.6-16 ft) high, branching from below with well-defined short trunks. Stems erect or ascending,

Pilocereus Leucocephalus
Pilosocereus lanuginosus, photograph by Keith Grantham

green to blue-green, 6-10 cm (2.4-3.9 in) in diameter. Ribs 7-12. Spines brownish, becoming gray with age. Central spine one, 2-3 cm (0.8-1.2 in) long. Radial spines 8-12, slender, 1-2 cm (0.4-0.8 in) long. Fertile portions usually well defined. Flower-bearing areoles apical to lateral, involving three or four ribs, with abundant silky white hairs 4-10 cm (1.6-3.9 in) long. Flowers tubular to bell shaped, pinkish to whitish, to 6 cm (2.4 in) long. Fruits globose, to 4 cm (1.6 in) in diameter. Distribution: Tamaulipas, Mexico, south to Guatemala and Honduras.

Pilosocereus machrisii (E. Y. Dawson) Backeberg 1960 Cephalocereus machrisii E. Y. Dawson 1957, Pseudopilocereus machrisii (E. Y. Dawson) Buxbaum 1968 Pilocereus cuyabensis Backeberg 1933, illegitimate name; Pseudopilocereus cuyabensis (Backeberg) Buxbaum 1968, illegitimate name Pseudopilocereus jauruensis Buining& Brederoo 1978, Pilosocereus jauruensis (Buining & Brederoo) P. J. Braun 1984 Pilosocereus paraguayensls F. Ritter 1979 Pilosocereus pusillibaccatus P. J. Braun & Esteves 1979 Pilosocereus saudadensis F. Ritter 1979, Pseudopilocereus saudaden-

sis (F. Ritter) P. V. Heath 1994 Pseudopilocereus parvus Diers & Esteves 1982, Pilosocereus parvus

(Diers & Esteves) P. J. Braun 1988 Pilosocereus cristalinensis P. J. Braun & Esteves 1987 Pilosocereus densivillosus P. J. Braun & Esteves 1987 Pilosocereus lindanus P. J. Braun & Esteves 1987

Plants shrubby, branching only at ground level, 0.4-3.5 m (1.3-11.5 ft) high. Stems erect, weakly woody, dark green to

Pilosocereus me jg n ific us
i'ilos ocereus lei tcocephalus

blue-green, somewhat gray or glaucous, smooth, 3.2-9 cm (1.3-3.5 in) in diameter. Ribs 8-15, transverse folds absent. Spines translucent when young, reddish, brownish, or golden. Central spines 3-8, ascending, the largest bent backward, 15-40 mm (0.6-1.6 in) long. Radial spines 9-24, flattened against the stem surface, 3-25 mm (to 1 in) long. Fertile portions strongly differentiated on three or more ribs, with yellow to gray hairs 20-30 mm (0.8-1.2 in) long. Flower buds acute. Flowers funnelform, white with reddish tips, 3-8 cm (1.2-3.1 in) long, 3-6 cm (1.2-2.4 in) in diameter. Fruits depressed globose, 2-3.5 cm (0.8-1.4 in) in diameter, dehiscing by lateral slits, pulp white. Distribution: elevations of 150-1200 m (490-3900 ft) in Goias, Mato Grosso, Mato Grosso do Sul, western Bahia, and western Minas Gerais, Brazil, and eastern Paraguay.

Pilosocereus magnificus (Buining 8c Brederoo) F. Ritter 1979 facheir0

Pseudopilocereus magnificus Buining & Brederoo 1972

Plants shrubby to treelike, 1.5-5 m (4.9-16 ft) high with or without well-defined trunks. Stems weakly woody, erect, pale blue, waxy, smooth, 4-7.5 cm (1.6-3 in) in diameter. Ribs 5-12, transverse folds visible only at stem tips. Spines bristly, translucent, golden yellow to brown. Central spines 8, erect, to 15 mm (0.6 in) long. Radial spines about 16, ascending, to 10 mm (0.4 in) long. Fertile portions only slightly differentiated. Flower-bearing areoles in clusters of three to six, occurring randomly along the stems, especially on middle to lower parts, with white hairs. Flower buds obtuse. Flowers narrow, flared, to 6 cm (2.4 in) long and 2.3 cm (0.9 in) in diameter. Fruits depressed globose, 2.5-3 cm (1-1.2 in) in diameter, dehiscing by various slits, pulp magenta. Distribution: northeastern Minas Gerais, Brazil, at elevations of 250-800 m (820-2600 ft).

Pilosocereus multicostatus F. Ritter 1979 Pseudopilocereus multicostatus (F. Ritter) P. V. Heath 1994

Plants shrubby, branching at ground level, 1.5-3.5 m (4.911.5 ft) high, usually without well-defined trunks. Stems weakly woody, erect, shiny dark green, smooth, 3.8-7.5 cm (1.5-3 in) in diameter. Ribs 18-25, transverse folds absent. Spines translucent, golden yellow to brown. Central spines 3-7, ascending, 10-20 mm (0.4-0.8 in) long. Radial spines 15-18, flattened against the stem surface, 5-10 mm (0.2-0.4 in) long. Fertile portions only slightly or not differentiated. Flower-bearing areoles randomly distributed along the stems, mostly subapical, with sparse gray or white long hairs, and with flexible golden bristles to 40 mm (1.6 in) long. Flower buds acute. Flowers flared, to 4.7 cm (1.9 in) long, 2.9-3 cm

(1.1-1.2 in) in diameter. Fruits depressed globose, 2.5-3 cm (1-1.2 in) in diameter, dehiscing by adaxial or abaxial slits, pulp magenta. Distribution: northeastern Minas Gerais, Brazil, at elevations of 670-900 m (2200-2950 ft).

Pilosocereus occultiflorus P. J. Braun 8c Esteves 1999

Plants treelike, candelabra-like, branching above 1 m (3.3 ft), to 7 m (23 ft) high with woody trunks to 23 cm (9.1 in) in diameter. Stems erect, robust, slightly curved basally, matte gray-green, to 3 m (9.8 ft) long, 6-9.5 cm (2.4-3.7 in) in diameter. Ribs 8-9, obtuse, triangular in cross section, with an undulating furrow between the ribs. Areoles 5-9 mm (0.20.4 in) apart. Spines straight to slightly curved, translucent, light yellow with brownish tips, becoming gray with dark tips with age. Central spines 6-13, pointing outward or downward, 2.5-9 mm (to 0.4 in) long. Radial spines about 20,911 mm (0.4 in) long. Fertile portions facing west. Flower-bearing areoles lateral, with chestnut-colored to brownish gray wool, on one to three or more ribs. Flowers 3-4 cm (1.2-1.6 in) long, 2-2.5 cm (0.8-1 in) in diameter. Fruits flattened globose, pale rose to greenish, pulp white, dehiscing by lateral slits. Distribution: western Minas Gerais, Brazil.

Pilosocereus oligolepis (Vaupel) Byles 8c G. D. Rowley 1957 Cereus oligolepis Vaupel 1913, Pilocereus oligolepis (Vaupel) Werdermann 1933, Pseudopilocereus oligolepis (Vaupel) Buxbaum 1968 Cephalocereus kanukuensis Alexander 1939, Pilosocereus kanukuensis (Alexander) Leuenberger 1987, P. oligolepis subsp. kanukuensis (Alexander) P.J. Braun & Esteves 1995

Plants shrubby, branched or unbranched basally, 1-2 m (3.36.6 ft) high. Stems erect or somewhat sprawling, gray-green to dark green, smooth, 3-5 cm (1.2-2 in) in diameter. Ribs 45. Spines opaque, dark brown. Central spines 1-6, erect, 1030 mm (0.4-1.2 in) long. Radial spines 8-16, flattened against the stem surface, 4-7 mm (to 0.3 in) long. Fertile portions slightly or not differentiated. Flower-bearing areoles with few to many white or brown hairs. Flowers slightly curved, flared, to 7 cm (2.8 in) long. Fruits depressed globose, 2-2.5 cm (0.8-1 in) in diameter. Distribution: northern Brazil and adjacent southwestern Guyana at elevations of 150-550 m (490-1800 ft). Pilosocereus oligolepis is poorly known.

Pilosocereus pachycladus F. Ritter 1979

facheiro, facheiro azul, mandacaru de facho Pseudopilocereus pachycladus (F. Ritter) P. V. Heath 1994 Pseudopilocereus azureus Buining & Brederoo 1975 Pilosocereus atroflavispinus F. Ritter 1979, Pseudopilocereus atroflav-

ispinus (F. Ritter) P. V. Heath 1994 Pilosocereus azureus F. Ritter 1979

Pilosocereus oreus F. Ritter 1979, Pseudopilocereus oreus (F. Ritter) P. V. Heath 1994

Pilosocereus pernambucoertsis F. Ritter 1979, P. pachycladus subsp. pernambucoensis (F. Ritter) Zappi 1994, Pseudopilocereus pernambucoertsis (F. Ritter) P. V. Heath 1994 Pilosocereus splendidus F. Ritter 1979, Pseudopilocereus splendidus

(F. Ritter) P. V. Heath 1994 Pilosocereus superbus F. Ritter 1979, Pseudopilocereus superbus

(F. Ritter) P. V. Heath 1994 Pilosocereus cyaneus F. Ritter 1981 Pilosocereus cenepegive/Rizzini &A. Mattos 1986 Pilosocereus schoebelii P. J. Braun 1987

Plants usually treelike, 2-10 m (6.6-33 ft) or more high. Stems erect, gray-green to blue-green, glaucous, smooth, 5.5-11 cm (2.2-4.3 in) in diameter. Ribs 5-19, transverse folds visible at the stem tips. Spines translucent, golden yellow, becoming gray. Central spines 1-12, ascending to erect, 0.5-30 mm (to 1.2 in) long. Radial spines 8-18, flattened against the stem surface, 5-15 mm (0.2-0.6 in) long. Fertile portions slightly to well differentiated. Flower-bearing areoles subapical on one or more ribs. Flower buds acute or obtuse. Flowers extremely variable, 4-7 cm (1.6-2.8 in) long, 2.2-4.5 cm (0.9-1.8 in) in diameter. Fruits depressed globose, dehiscing by lateral slits, pulp magenta. Distribution: common and widely distributed in northeastern Brazil.

Two subspecies of Pilosocereus pachycladus are recognized. Subspecies pachycladus has stems with 5-12 high, broad ribs, long central spines easily distinguished from the radials, and flower-bearing areoles that are densely hairy; it occurs in Bahia and Minas Gerais south of 10° south latitude. Subspecies pernambucoensis has stems with 13-19 low, narrow ribs, central spines the same length as the radials and not easily distinguished from them, and flower-bearing areoles that are not obviously hairy; it occurs in northeastern Brazil north of 10° south latitude.

Pilosocereuspentaedrophorus (Cels) Byles & G. D. Rowley 1957

facheiro, facheiro fino, mandacaru deveado

Cereus pentaedrophorus Cels 1858, Pilocereus pentaedrophorus (Cels) Console ex K. Schumann 1897, Cephalocereus pentaedrophorus (Cels) Britton & Rose 1920, Pseudopilocereus pentaedrophorus (Cels) Buxbaum 1968 Pilosocereus pentaedrophorus subsp. robustus Zappi 1994

Plants shrubby or treelike, sparsely to much branched, to 6 m (20 ft) high. Stems woody, erect or leaning, glaucous blue-green, 3-7.5 cm (1.2-3 in) in diameter. Ribs 4-10, transverse folds horizontal. Spines translucent yellowish brown. Central spines 0-3, ascending to erect to bent backward, 10-26 mm (0.4-1 in) long. Radial spines 3-12, flattened against the stem surface, 4-20 mm (to 0.8 in) long. Fertile portions not differentiated. Flower-bearing areoles subapical, present on several ribs. Flower buds curved and obtuse. Flowers flared, 3.55.5 cm (1.4-2.2 in) long, to 2.8 cm (1.1 in) in diameter. Fruits depressed globose, 2-3 cm (0.8-1.2 in) in diameter, dehiscing by lateral slits, pulp purplish or magenta. Distribution: Pernambuco, Bahia, and Minas Gerais, Brazil.

drophorus Pilosocereus piauhyensis
photograph by Urs Eggli

Two subspecies of Pilosocereus pentaedrophorus are recognized. Subspecies pentaedrophorus is usually treelike and sparsely branched with long, slender, leaning stems to 4.5 cm (1.8 in) in diameter with 4-6 obtuse ribs; it occurs from Bahia to Pernambuco. Subspecies robustus is shrubby or treelike with stout branches that never lean, to 7.5 cm (3 in) in diameter with 6-10 acute ribs; it occurs in southern Bahia and northeastern Minas Gerais.

Pilosocereus piauhyensis (Gürke) Byles 8c G. D. Rowley 1957

facheiro, rabo de raposa

Cereus piauhyensis Gürke 1908, Cephalocereus piauhyensis (Gürke) Britton & Rose 1920, Pilocereus piauhyensis (Gürke) Werdermann 1933, Pseudopilocereus piauhyensis (Gürke) Buxbaum 1968 Pseudopilocereus mucosiflorus Buining & Brederoo 1977, Pilosocereus mucosiflorus (Buining & Brederoo) F. Ritter 1979, Pilosocereus piauhyensis subsp. mucosiflorus (Buining & Brederoo) P. J. Braun & Esteves 1995

Pilosocereus gaturlanensis F. Ritter 1979, Pseudopilocereus gaturia-nensis (F. Ritter) P. V. Heath 1994, Pilosocereus piauhyensis subsp. gaturianensis (F. Ritter) P. J. Braun & Esteves 1995

Plants shrubby, branching only at the base, 1.5-2.5 m (4.98.2 ft) high. Stems erect, weakly woody, shiny dark green, smooth, 5-7.5 cm (2-3 in) in diameter. Ribs 14-21, transverse folds absent. Spines translucent golden. Central spines 5-9, ascending to erect, 5-15 mm (0.2-0.6 in) long. Radial spines 11-16, flattened against the stem surface, 3-8 mm (to 0.3 in) long. Fertile portions slightly or not differentiated. Flower-bearing areoles subapical, with long white to gray hairs, and flexible golden bristles. Flower buds obtuse, Flowers flared, 5.5-7.5 cm (2.2-3 in) long, 3-4 cm (1.2-1.6 in) in diameter. Fruits depressed globose, 3.8 cm (1.5 in) in diameter, dehiscing by lateral slits, pulp magenta. Distribution: Piaui, Rio Grande do Norte, and probably Cearä, Brazil.

Pilosocereuspolygonus (Lamarck) Byles 8c G. D. Rowley 1957

bahama dildo, key tree cactus, robin tree cactus

Cactus polygonus Lamarck 1783, Cephalocereus polygonus (Lamarck)

Britton & Rose 1909 Pilocereus robinii Lemaire 1864, Cephalocereus robinii (Lemaire) Britton & Rose 1920, Pilosocereus robinii (Lemaire) Byles & G. D. Rowley 1957, Cereus robinii (Lemaire) L. D. Benson 1969 Cephalocereus bahamensis Britton & Rose 1909, Pilosocereus baha-

mensis (Britton & Rose) Byles & G. D. Rowley 1957 Cephalocereus keyensis Britton & Rose 1909, Pilosocereus keyensis

(Britton & Rose) Byles & G. D. Rowley 1957 Cereus brooksianus Vaupel 1912, Pilosocereus brooksianus (Vaupel)

Byles & G. D. Rowley 1957 Cephalocereus deeringii Small 1917, Pilosocereus deeringii (Small) Byles &G.D. Rowley 1957

Plants shrubby or treelike, 3-10 m (9.8-33 ft) high. Stems erect or ascending, bluish to blue-green, especially when young, 5-10 cm (2-3.9 in) in diameter. Ribs 5-13, narrow, transverse folds evident. Spines dense, yellowish, becoming gray, spreading, 10-25 mm (0.4-1 in) long. Fertile portions not differentiated. Flower-bearing areoles with dense white wool. Flowers 5-6 cm (2-2.4 in) long, 2.5-5 cm (1-2 in) in diameter. Fruits depressed globose. Distribution: Florida, Bahamas, Cuba, and Hispaniola. Pilosocereus polygonus is listed (as P. robinii) as endangered in the U.S. Endangered Species Act.

Pilosocereus purpusii (Britton 8c Rose) Byles 8c G. D. Rowley

1957 viejo

Cephalocereus purpusii Britton & Rose 1920 ?Cephalocereus collinsii Britton & Rose 1923, ?Pilosocereus collinsii (Britton & Rose) Byles & G. D. Rowley 1957

Pilosocereus purpusii, photograph by Charles Glass

Plants shrubby, sometimes solitary but usually branched basally, sometimes laterally, 2-3 m (6.6-9.8 ft) high. Stems erect, olive green, sometimes bluish near the tips, 4-5 cm (1.6-2 in) in diameter. Ribs 7-12, broad, transverse folds visible. Spines opaque, yellowish, becoming gray or almost black. Central spine usually one, sometimes as many as 4, 15-25 mm (0.6-1 in) long. Radial spines 8-10, flattened against the stem surface, to 10 mm (0.4 in) long. Fertile por

Pilosocereus quadricentralis, photograph by Charles Glass tions well defined. Flower-bearing areoles apical and sub-apical, with numerous silky hairs. Flowers to 7 cm (2.8 in) long, pale red with white. Fruits globose. Distribution: Sina-loa, Nayarit, Jalisco, Michoacán, and Oaxaca, Mexico.

Pilosocereus quadricentralis (E. Y. Dawson) Backeberg 1960

Cephalocereus quadricentralis E. Y. Dawson 1948

Plants treelike, to 5 m (16 ft) high, branching well above the ground with well-defined trunks. Stems erect, glaucous green, 7-8 cm (2.8-3.1 in) in diameter. Ribs 9, prominent. Spines brownish red, becoming gray, bulbous basally. Central spines 4, diverging, 20-35 mm (0.8-1.4 in) long. Radial spines 11-13, diverging, 10-20 mm (0.4-0.8 in) long. Fertile portions well defined. Flower-bearing areoles lateral, apical, with abundant white wool. Flowers large. Fruits globose. Distribution: Oaxaca and Chiapas, Mexico.

Pilosocereus royenii (Linnaeus) Byles & G. D. Rowley 1957 Cactus royenii Linnaeus 1753, Cephalocereus royenii (Linnaeus) Britton & Rose 1909 Cereus nobilis Haworth 1812, Pilocereus nobilis (Haworth) K. Schumann 1894, Cephalocereus nobilis (Haworth) Britton & Rose 1909, Pseudopilocereus nobilis (Haworth) Buxbaum 1968 Cereus monoclono s A. P. de Candolle 1828, Cephalocereus mono-clonos (A. P. de Candolle) Britton & Rose 1909, Pilosocereus mono-clonos (A. P. de Candolle) Byles & G. D. Rowley 1957 Cereus swartzii Grisebach 1860, Cephalocereus swartzii (Grisebach) Britton & Rose 1909, Pilosocereus swartzii (Grisebach) Byles & G. D. Rowley 1957 Pilocereus urbanianus K. Schumann 1897, Pilosocereus urbanianus

(K. Schumann) Byles & G. D. Rowley 1957 Cephalocereus millspaughii Britton 1909, Pilosocereus miilspaughii

(Britton) Byles & G. D. Rowley 1957 Cephalocereus brooksianus Britton & Rose 1912 Cephalocereus barbadensis Britton & Rose 1920, Pilosocereus barba-

densis (Britton & Rose) Byles & G. D. Rowley 1957 Cephalocereus gaumeri Britton & Rose 1920, Pilosocereus gaumeri (Britton & Rose) Backeberg 1960

Plants treelike, branching basally or above the ground, 2-8 m (6.6-26 ft) high, often with well-defined trunks. Stems erect or sometimes ascending, stout, blue-green, 7-9 cm (2.8-3.5 in) in diameter. Ribs 6-11. Spines variable, yellowish to reddish. Central spines 1-6,32-60 mm (1.3-2.4 in) long. Radial spines about 9,19-26 mm (0.7-1 in) long. Fertile portions not well differentiated. Flower-bearing areoles subapical and lateral, on one to three ribs, with long white hairs. Flowers rose pink, to 5 cm (2 in) long, 3-4 cm (1.2-1.6 in) in diameter. Fruits depressed globose, pulp white. Distribution: Yucatán Peninsula, Mexico, and Bahamas, Dominican Repub-

Pilosocereus quadricentralis, photograph by Charles Glass

Pilosocereus tuberculatus 587

lie, Jamaica, Puerto Rico, Tobago, U.S. Virgin Islands, and Lesser Antilles. Pilosocereus royenii is one of the most common cacti in the Caribbean.

Pilosocereus Xsubsimilis Rizzini & A. Mattos 1986

Pilosocereus xsubsimilis is almost certainly a naturally occurring hybrid, P. floccosus subsp. quadricostatus x P. magnifi-cus. Plants shrubby or treelike, sparsely branched above the ground, 2-5 m (6.6-16 ft) high with well-defined trunks. Stems weakly woody, erect, dark green to olive green, somewhat glaucous, slightly rough, 7-10 cm (2.8-3.9 in) in diameter. Ribs 4-7, transverse folds absent. Spines translucent when young, yellowish to brown to red. Central spines 4-5, ascending to erect, 7-15 mm (0.3-0.6 in) long. Radial spines 10-14, ascending to erect, sometimes deflexed, 6-10 mm (0.2-0.4 in) long. Fertile portions somewhat to well differentiated. Flower-bearing areoles on all ribs near the stem tips, forming rings, with black bristly spines to 30 mm (1.2 in) long and white hairs. Flowers to 5 cm (2 in) long and 2.5 cm (1 in) in diameter. Fruits depressed globose, pulp magenta. Distribution: northeastern Minas Gerais, Brazil.

Pilosocereus tuberculatus (Werdermann) Byles & G. D.

Rowley 1957 CAXACUBRI

Pilocereus tuberculatus Werdermann 1933, Pseudopilocereus tuberculatus (Werdermann) Buxbaum 1968

Plants treelike, 2-6 m (6.6-20 ft) high, branching freely sub-apically, spreading to 6 m (20 ft) wide, usually with well-defined trunks. Stems strongly woody, erect to inclined, olive green, waxy when young, 3-6 cm (1.2-2.4 in) in diameter. Ribs 4-7, with sinuate sinuses, transverse folds pronounced and oblique. Areoles borne on podaria. Spines light brown, becoming gray, sharp, brittle, erect. Central spines 3-5,3042 mm (1.2—1.7 in) long, lowermost one longest and often curved downward. Radial spines 10-12, flattened against the stem surface, curved downward, 4-10 mm (to 0.4 in) long. Fertile portions not differentiated. Flower-bearing areoles subapical. Flower buds acute. Flowers 6-6.7 cm (2.4-2.6 in) long, to 3 cm (1.2 in) in diameter. Fruits globose, to 4 cm (1.6 in) in diameter, dehiscing by lateral slits, pulp magenta. Distribution: northwestern to northeastern Bahia and adjacent Pernambuco, Brazil.

588 Pilosocereus tuberculatus

Pilosocereus tuberculatus, photograph by Urs Eggli

Pilosocereus ulei (K. Schumann) Byles & G. D. Rowley 1957

cabeqadevelho

Pilocereus ulei K. Schumann 1903, Cereus ulei (K. Schumann) A.

Berger 1905, Pseudopilocereus ulei (K. Schumann) Buxbaum 1968 Cephalocereus robustus Britton & Rose 1920

Plants shrubby to treelike, branching above the bases, to 5 m (16 ft) high with well-defined trunks. Stems erect, gray-green, waxy, 9-12 cm (3.5-4.7 in) in diameter. Ribs 6-8, transverse folds absent. Spines translucent only when young, reddish brown, becoming blackish gray. Central spine one, 12-18 mm (0.5-0.7 in) long, erect, sometimes pointing downward. Radial spines 8-10, flattened against the stem surface, 3-10 mm (to 0.4 in) long. Fertile portions well differentiated. Flower-bearing areoles subapical, on most ribs, with stiff white or gray hairs to 11 cm (4.3 in) long. Flower buds obtuse. Flowers flared, 4.2-4.5 cm (1.7-1.8 in) long, 2.7-3 cm (1.1-1.2 in) in diameter. Fruits depressed globose, 2.5-3 cm (1-1.2 in) in diameter, dehiscing by lateral or adax-ial slits, pulp magenta. Distribution: southeastern Rio de Janeiro, Brazil.

Pilosocereus vilaboensis (Diers 8c Esteves) P. J. Braun 1988 Pseudopilocereus vilaboensis Diers & Esteves 1983 Pilosocereus rizzoanus P. J. Braun & Esteves 1992

Plants shrubby, branched basally, to 1.4 m (4.6 ft) high. Stems erect, pale green, smooth, 3.5-5.5 cm (1.4-2.2 in) in diameter. Ribs 12-16, transverse folds absent. Spines translucent, golden to rust colored, sometimes bristle-like. Central spines 2-8, erect, 4-30 mm (to 1.2 in) long. Radial spines 10-20, flattened against the stem surface, 4-30 mm (to 1.2 in) long. Fertile portions poorly differentiated. Flower-bear -ing areoles produced randomly, subapical, with golden bris tles and long white hairs. Flowers flared, funnelform, to 5 cm (2 in) long, 3-5 cm (1.2-2 in) in diameter. Fruits globose, 3-4 cm (1.2-1.6 in) in diameter, pulp white. Distribution: western central Goias, Brazil.

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