Grusonia

DNA sequence investigations of the opuntioid cacti by Steven Dickie and Robert Wallace (2001), seed studies by Wolfgang Stuppy (2001), and morphological and cytological analyses by Donald Pinkava (2001) have shed considerable light upon a confusing group. It has become clear that the large, inclusive genus Opuntia is polyphyletic, inviting subdivision into smaller, monophyletic groups. Unfortunately, the boundaries between some genera, including Grusonia, remain unclear. Stuppy suggests that Corynopuntia, Grusonia, Mareno-puntia, and Micropuntia should be placed in Cylindropuntia. Dickie and Wallace, on the other hand, believe that Cylindropuntia is distinct from the other groups, which should all be included in Grusonia. I have followed the International Cac-taceae Systematics Group's decision to keep Cylindropuntia separate from the others, thus Grusonia, as treated here, includes Corynopuntia, Marenopuntia, and Micropuntia, in large part following Harold Robinson (1973). Cylindropuntia can be distinguished from Grusonia because it has round spines covered completely by deciduous papery sheaths. The spines of Grusonia tend to be flattened, roughened or bulbous basally, with little or no sheath. Grusonia comprises 17 species.

Grusonia (type, Cereus bradtianus = G. bradtiana) was described by F. Reichenbach and published by Nathaniel Britton and Joseph Rose (1919-1923,1:215), though according to Index Kewensis the name was used horticulturally by Ernst Nicolai and published by Karl Schumann in 1894. The name honors Herman Gruson, a German plant collector from Magdeburg who sent his gardener, A. Mathesson, to Mexico in 1889 to collect specimens for him (Rowley 1997,367).

Grusonia F. Reichenbach exBritton & Rose 1919 Corynopuntia F. M. Knuth 1935 Micropuntia Daston 1947 Marenopuntia Backeberg 1950

Subfamily Opuntioideae. Plants either low growing and cushion forming or shrubby ortreelike, with indefinite growth and many erect branches. Roots usually tuberous but sometimes fibrous. Stems usually in distinct segments, cylindrical to club shaped, sometimes with tuberculate ribs. Leaves small, cylindrical, falling away early. Areoles with hairs, glochids, and spines. Spines without distinct papery sheaths or nearly so, somewhat flattened, roughened or bulbous basally. Flowers borne laterally oralmostapically, open duringthe day, pink, purple, yellow, or white. Fruits dry or fleshy, sometimes dehiscent, often sterile. Seeds whitish to yellowish, 2.5-5 mm longand in diameter; funicular envelopes glabrous or covered with thin trichomes; funicular girdle well developed; perisperm large. Distribution: southwestern United States, and Baja California Peninsula and several states of northern Mexico.

Grusonia aggeria (Ralston &Hilsenbeck) E. F. Anderson 1999

Opuntia aggeria Ralston & Hilsenbeck 1989

Plants low, forming clumps 3-9 cm (1.2-3.5 in) high. Stem segments short cylindrical to club shaped, 3.5-9 cm (1.4-3.5 in) long, 1.5-3 cm (0.6-1.2 in) in diameter. Glochids yellow, to 5 mm long. Spines mostly in distal areoles, 5-15, sometimes only one, 3-5 cm (1.2-2 in) long. Principal apical spines 0-5, ascending, diverging, reddish brown to gray, bulbous basally, round in cross section. Principal basal spines chalky white, bent downward, flattened, twisted or curved. Flowers bright yellow, to 2.5 cm (1 in) long. Fruits dry, cylindrical to ellipsoidal, 2-2.5 cm (0.8-1 in) or more long, without spines but with glochids. Distribution: Chihuahuan Desert of southeast Texas, into central Coahuila, Mexico.

Grusonia agglomerata (A. Berger) E. F.Anderson 1999 Opuntia agglomerata A. Berger 1929, Corynopuntia agglomerata (A. Berger) F.M.Knuth 1935

Plants low, much branched and many-stemmed. Stem segments obovate, gray-green, 3-4 cm (1.2-1.6 in) long. Ribs 6, tuberculate. Areoles round, grayish white. Glochids white to yellowish. Principal spines 4-5, bulbous basally, yellowish, lower 2 curved and 10-15 mm (0.4-0.6 in) long, lateral 2 diverging and grayish white. Secondary or radial spines 6-7, erect, needle-like, short, whitish, 1-4 mm long. Flowers not known. Distribution: Coahuila, Mexico.

Grusonia bradtiana (J. M. Coulter) Britton &Rose 1919

ORGANILLO, VIEJO

Cereus bradtianus J. M. Coulter 1896, Opuntia bradtiana (J. M. Coulter)

K. Brandegee 1897 Grusonia cereiformis F. Reichenbach 1894, Opuntia cereiformis (F. Reichenbach) F. A. C. Weber 1898

Plants low, branching to form dense and impenetrable mats, to 1 m (3.3 ft) high. Stem segments green, 4-7 cm (1.6-2.8 in) in diameter. Ribs 8-10, low, tuberculate. Areoles white, 35 mm in diameter. Leaves linear, fleshy, green, early deciduous, to 8 mm (0.3 in) long. Spines 15-25, brownish yellow when young, later whitish, needle-like, rounded or slightly flattened, 1-3 cm (0.4-1.2 in) long, sometimes the largest ones directed downward. Flowers yellow, 3-4 cm (1.2-1.6 in) long. Fruits ellipsoidal, strongly umbilicate. Distribution: Coahuila, Mexico.

Grusonia bulbispina (Engelmann) H. Robinson 1973 Opuntia bulbispina Engelmann 1856, Corynopuntia bulbispina (Engelmann) F. M. Knuth 1935

Plants low, forming mats 0.6-1.2 m (24-47 in) wide. Stem segments ovoid, 2-2.5 cm (0.8-1 in) long, 1-1.2 cm (0.4-0.5 in) in diameter, with prominent tubercles 6-8 mm (0.2-0.3 in) high. Central spines 4, bulbous basally, 8-12 mm (0.30.5 in) long. Radial spines 8-12, needle-like, 3-6 mm (to 0.2 in) long. Flowers purple. Distribution: Coahuila, Durango, and Zacatecas, Mexico.

Grusonia clavata (Engelmann) H. Robinson 1973

CLUB CHOLLA, DAGGER CHOLLA

Opuntia clavata Engelmann 1848, Cylindropuntia clavata (Engelmann) F. M. Knuth 1930, Corynopuntia clavata (Engelmann) F. M. Knuth 1935

Plants low, spreading widely, forming mats 5-15 cm (2-5.9 in) high. Roots fibrous. Stem segments short club shaped, strongly narrowing basally, 2.5-7.5 cm (1-3 in) long, 1.5-3 cm (0.6-1.2 in) in diameter, with ovate tubercles. Areoles round, with white to gray wool. Glochids yellowish white, in upper parts of areoles, to 4 mm long. Spines 7-15, borne mostly on distal areoles. Principal apical spines 1-3, ascending, white to yellowish to tan, angular-flattened to nearly

round in cross section. Major lower spines 3-5, bent downward, white, flattened, the longer ones daggerlike, 12—35 mm (0.5-1.4 in) long. Flowers bright yellow, to 2.5 cm (1 in) long. Fruits barrel shaped to ellipsoidal, yellow, fleshy, 3-4.5 cm (1.2-1.8 in) long, 1.5-2.5 cm (0.6-1 in) in diameter, without spines but with glochids. Distribution: Great Plains grasslands of the United States into northern central Mexico. Stems of Grusonia clavata are used medicinally (Chapter 2, under Cacti as Medicine).

Grusonia dumetorum (A. Berger) E. F. Anderson 1999 Opuntia dumetorum A. Berger 1929, Corynopuntia dumetorum (A. Berger) F. M. Knuth 1935, Platyopuntia dumetorum (A. Berger) F. Ritter 1979

Plants forming clumps to 50 cm (20 in) high. Stem segments more or less cylindrical, grayish green, somewhat pubescent, slightly tuberculate. Areoles round, with white wool and a few white hairs. Glochids whitish. Spines several, unequal, diverging, yellowish brown, 1.2-1.5 cm (0.5—0.6 in) long. Flowers, fruits not known. Distribution: Tamaulipas, Mexico. Grusonia dumetorum is poorly understood.

Grusonia emoryi (Engelmann) Pinkava 1999 CURSED CHOLLA, DEVIL CHOLLA, PEEBLE'S CHOLLA, STANLY'S CLUB CH0LLA, WRIGHT'S CLUB CHOLLA Opuntia emoryi Engelmann 1856

Opuntia stanlyi Engelmann 1848, not validly published; Corynopuntia stanlyi (Engelmann) F. M. Knuth 1935, Grusonia stanlyi (Engelmann) H. Robinson 1973

Plants forming low, much branched mats, 15-30 cm (5.9-12 in) high. Roots fibrous. Stem segments curved club shaped, narrowing gradually or abruptly basally, 7-19 cm (2.8-7.5 in) long, 2.5-5 cm (1-2 in) in diameter, with prominent tubercles 2-5 cm (0.8-2 in) high. Areoles round, with white to gray wool. Glochids yellow to brown, 5-7 mm (0.2-0.3 in) long. Spines evenly distributed or only on distal areoles, yellow, reddish brown, to tan, spreading. Principal apical spines 3-5, sometimes only one, ascending, diverging, angular to flattened to nearly round in cross section. Principal basal spines 3-5, diverging, sharp edged, angular to flattened. Flowers yellow, 2-3 cm (0.8-1.2 in) long. Fruits cylindrical to ellipsoidal, yellow, fleshy, 4-9 cm (1.6-3.5 in) long, 1.4-4 cm (0.6-1.6 in) in diameter, without spines but with dense yellow glochids. Distribution: Sonoran and Chihuahuan Deserts of southeastern Arizona, southwestern New Mexico, and southwest Texas, and neighboring Chihuahua, Mexico.

Grusonia grahamii (Engelmann) H. Robinson 1973 GRAHAM CLUB CACTUS, GRAHAM'S DOG CACTUS, MOUNDED DWARF CHOLLA

Opuntia grahamii Engelmann 1856, Corynopuntia grahamii (Engelmann) F. M. Knuth 1935

Plants forming low mats 8-20 cm (3.1-7.9 in) high. Roots tuberous. Stem segments cylindrical, curved, narrow basally, 3-7 cm (1.2-2.8 in) long, 1.5-3.5 cm (0.6-1.4 in) in diameter, with narrow tubercles 10-15 mm (0.4-0.6 in) high. Areoles round, with white to yellowish wool. Glochids white to yellow, abundant in basal areoles, to 6 mm (0.2 in) long. Spines 6-15, mostly in distal areoles, white to reddish brown, 30-55 mm (1.2-2.2 in) long. Principal apical spines 2-5, ascending, diverging, round in cross section. Principal basal spines erect, the marginal ones bent downward, white, angular to flattened to somewhat round in cross section. Flowers bright yellow, 2-2.5 cm (0.8-1 in) long. Fruits cylindrical

Grusonia clavata

Grusonia emoryi

Grusonia clavata

Grusonia emoryi to ellipsoidal, fleshy, yellowish, 2-3.5 cm (0.8-1.4 in) long, without spines but with yellow glochids. Distribution: Chi-huahuan Desert of southern New Mexico and southwestern Texas, and Chihuahua and Durango, Mexico.

Grusonia invicta (T. Brandegee) E. F. Anderson 1999

DAGGER CHOLLA

Opuntia invictaT. Brandegee 1889, Corynopuntia invicta (T. Brandegee) F. M. Knuth 1935

Plants small, low, many-stemmed, forming dense masses 2045 cm (7.9-18 in) high and to 2 m (6.6 ft) wide with interweaving, erect and clambering stems. Stem segments widely obovate to club shaped, 8-15 cm (3.1-5.9 in) long, 4-6 cm (1.6-2.4 in) in diameter, with well-defined tubercles 2-5 cm (0.8-2 in) long, 1-2 cm (0.4-0.8 in) wide, 1 cm (0.4 in) high. Areoles nearly round, with gray wool, to 1.5 cm (0.6 in) in diameter. Glochids only rarely present. Spines 10-25, very rigid, gray with darker tips, diverging to bent downward, very flattened but four-angled, 1-5 cm (0.4-2 in) long, to 3 mm wide basally. Flowers yellow, 4-6 cm (1.6-2.4 in) in diameter; pericarpels very spiny. Fruits ovoid, moderately fleshy, 4-5 cm (1.6-2 in) long, 2.5-3 cm (1-1.2 in) in diameter, very spiny. Distribution: central region of Baja California Peninsula, Mexico.

Grusonia kunzei (Rose) Pinkava 1999

DEVIL CHOLLA, KUNZE CLUB CHOLLA, WRIGHT'S CLUB CHOLLA Opuntia kunzei Rose 1908

Grusonia wrightiana E. M. Baxter 1935, Opuntia wrightiana (E. M. Baxter) Peebles 1937

Plants much branched, forming mats to 50 cm (20 in) high. Stem segments club shaped, curved, narrowing basally,

Grusonia invicta

10-15 cm (3.9-5.9 in) long, 2.5-4 cm (1-1.6 in) in diameter, with prominent tubercles 2-3 cm (0.8-1.2 in) high. Areoles round, with gray to white wool. Glochids yellow, few, 5-6 mm (0.2 in) long. Spines 17-27, abundantly distributed along the stems and obscuring them. Principal apical spines tan to reddish tan, diverging, angled basally, the longest to 50 mm (2 in). Principal basal spines tan with white margins, strongly flattened, bent downward, the longest to 45 mm (1.8 in). Flowers yellow to pale yellowish green, 1.5-2 cm (0.60.8 in) long. Fruits obovate to oblong, lemon yellow to yellow, fleshy, sometimes proliferating, 4-7.5 cm (1.6-3 in) long, 1.5-4 cm (0.6-1.6 in) in diameter, with glochids and spines. Distribution: Sonoran Desert of southwestern Arizona, and northern Baja California and Sonora, Mexico.

Grusonia marenae (S. H. Parsons) E. F. Anderson 1999 Opuntia marenae S. H. Parsons 1936, Marenopuntia marenae (S. H.

Parsons) Backeberg 1950

Plants shrubby, erect, much branched, 15-60 cm (5.9-24 in) high, the older parts woody. Roots tuberous. Stems not segmented, slender cylindrical, branching, shiny green, to 20 cm (7.9 in) long, 0.8-1.5 cm (0.3-0.6 in) in diameter, with tubercles. Areoles round, 3 mm in diameter, with very small glochids and some needle-like spines. Leaves awl shaped, green or reddish purple, to 1 cm (0.4 in) long. Principal apical spines 2-6, whitish, to 2 cm (0.8 in) long, bulbous basally. Principal basal spines dark, directed downward, bulbous basally. Secondary spines 7-12, white, needle-like, next to the surface, 0.3-1 cm (to 0.4 in) long. Flowers borne terminally, sunken into the stem, often open for several days, whitish, without receptacular tubes, 6-8 cm (2.4-3.1 in) in diameter. Fruits dry, ripening in the swelling stem tips, dehiscing laterally. Distribution: Sonora, Mexico.

Grusonia kunzei

Grusonia marenae

Grusonia moelleri (A. Berger ex Y. Wright) E. F. Anderson 1999

Opuntia moelleri A, Berger ex Y, Wright 1929 Corynopuntia moellerina F. M. Knuth 1935

Plants many-stemmed with small branches. Stem segments more or less club shaped, green, 4-7 cm (1.6-2.8 in) long, 34 cm (1.2-1.6 in) in diameter, with large, elongated tubercles. Principal spines usually 6, bulbous basally, upper ones radiating and erect, lower ones directed downward, flattened, whitish, slightly pubescent, to 1.6 cm (0.6 in) long. Secondary or radial spines numerous, white, fine, upper ones grouped like glochids. Flowers transparent greenish yellow, 5-6 cm (2-2.4 in) long, to 5 cm (2 in) in diameter; peri-carpels strongly tuberculate and spiny. Distribution: Coahu-ila, Mexico.

Grusonia parishii (Orcutt) Pinkava 1999 GROUND MAT CHOLLA, PARISH CHOLLA Opuntia parishii Orcutt 1896, Corynopuntia parishii (Orcutt) F. M. Knuth 1935

Plants low, much branched, forming mats 10-20 cm (3.97.9 in) high. Stem segments club shaped, 5-9 cm (2-3.5 in) long, 2-3 cm (0.8-1.2 in) in diameter, with tubercles. Areoles round, with gray to white wool. Glochids yellow, 5-8 mm (0.2-0.3 in) long. Spines 14-22, borne on distal areoles or over entire segment, white to brown, with yellowish tips. Principal apical spines about 5, brownish, diverging, somewhat round in cross section, angular to flattened basally, to 42 mm (1.7 in) long. Principal basal spines 5-6, whitish to pinkish to tan, strongly bent backward, flattened, 25-45 mm

(1-1.8 in) long. Flowers pale yellow with reddish midribs, . 1.5-2.2 cm (0.6-0.9 in) long. Fruits yellow, fleshy, 3.5-5.5 cm (1.4-2.2 in) long, 1.5-2 cm (0.6-0.8 in) in diameter, without spines but with dense yellow glochids. Distribution: Mojave and Sonoran Deserts of southeastern California, southern Nevada, and northwestern and southern Arizona.

Grusoniapulchella (Engelmann) H.Robinson 1973 DWARF CHOLLA

Opuntia pulchella Engelmann 1863, Corynopuntia pulchella (Engelmann) F. M. Knuth 1935; Micropuntia pulchella (Engelmann) C. Holland 1997, not validly published Micropuntia barkleyana Daston 1947, Opuntia barkleyana (Daston)

G. D. Rowley 1958 Micropuntia brachyrhopalica Daston 1947, Opuntia brachyrhopalica

(Daston) G. D. Rowley 1958 Micropuntia spectatissima Daston 1947, Opuntia spectatissima (Daston) G. D. Rowley 1958 Micropuntia gracilicylindrica Wiegand & Backeberg 1956, Opuntia gra-

cilicylindrica (Wiegand & Backeberg) G. D. Rowley 1958 Micropuntiapygmaea Wiegand & Backeberg 1956, Opuntia pygmaea

(Wiegand & Backeberg) G. D. Rowley 1958 Micropuntia tuberculosirhipalica Wiegand & Backeberg 1956, Opuntia tuberculosirhipalica (Wiegand & Backeberg) G. D. Rowley 1958 Micropuntia wiegandil Backeberg 1957, Opuntia wiegandii (Backeberg) G. D. Rowley 1958

Plants low, much branched, forming mats 10-20 cm (3.97.9 in) high. Roots tuberous. Stem segments cylindrical to ellipsoidal to club shaped, 1-4 cm (0.4-1.6 in) long, scfme-times as long as 10 cm (3.9 in), and 0.5-2.5 cm (0.2-1 in) in diameter, with rather prominent tubercles. Areoles round, with white to gray wool. Glochids yellow to reddish yellow, barbed, to 0.8 cm (0.3 in) long. Spines 8-15, mostly in distal areoles, white to reddish brown, becoming gray with age, divergent to bent downward, flattened, longest to 6 cm (2.4

Grusonia parishii

Gymnocalycium 347

in). Flowers rose to purple, 1.5-2.5 cm (0.6-1 in) long. Fruits obconical, smooth, reddish, fleshy, 1.5-3 cm (0.6-1.2 in) long, 0.8-1.2 cm (0.3-0.5 in) in diameter. Distribution: Mo-jave Desert of central Nevada and neighboring border areas of California and Utah.

Grusonia reflexispina (Wiggins & Rollins) E. F.Anderson 1999

Opuntia reflexispina Wiggins & Rollins 1943, Corynopuntia reflexispina (Wiggins & Rollins) Backeberg 1958

Plants low, moderately branched, forming clumps 20-40 cm (7.9-16 in) high, 30-100 cm (12-39 in) wide, with erect candelabra-like trunks. Stem segments cylindrical to broadly club shaped, pale yellowish green, glabrous, 5-10 cm (2-3.9 in) long, 0.8-1.5 cm (0.3-0.6 in) in diameter, with broad tubercles. Areoles round to elliptical, with white wool and twisted hairs soon falling away. Glochids numerous in older areoles, yellow, 1-7 mm (to 0.3 in) long. Principal spines 2-5, broadly awl shaped, flattened, rough, bulbous basally, with small papillae. Principal apical spine one, erect, gray, 6-10 mm (0.2-0.4 in) long. Principal basal spines gray to almost white, bent backward, fine, 8-15 mm (0.3-0.6 in) long. Secondary or radial spines about 10, needle-like to flattened, 4-6 mm (0.2 in) long. Flowers borne in clusters at segment tips, yellow. Fruits dry, green. Distribution: Sonora, Mexico.

Grusonia schottii (Engelmann) H. Robinson 1973

CLAVELLINA, DEVIL CACTUS, DOG CH0LLA, SCH0TT CLUB CACTUS

Opuntia schottii Engelmann 1856, Corynopuntia schottii (Engelmann)

F.M.Knuth 1935 Opuntia densispina Ralston & Hilsenbeck 1992

Plants low, forming extensive mats 7.5-9 cm (3-3.5 in) high. Roots fibrous. Stem segments club shaped, 2-6.5 cm (0.82.6 in) long, 1.5-3 cm (0.6-1.2 in) in diameter, with broad

Grusonia schottii

tubercles to 2 cm (0.8 in) high. Areoles round, with white or yellow wool. Glochids yellowish white to yellow, to 0.7 cm (0.3 in) long. Spines 11-17, borne mostly on distal areoles, spreading to bent downward, white to reddish brown, 3-7 cm (1.2-2.8 in) long. Principal apical spines 2-3, ascending, diverging, somewhat round to angular-flattened in cross section, reddish brown. Principal basal spines divergent or bent downward, tan to brown, sharp edged, flattened to angular-flattened. Flowers bright yellow, to 2 cm (0.8 in) long. Fruits cylindrical to ellipsoidal, yellow, fleshy, without spines but with yellow glochids. Distribution: Chihuahuan Desert from southern central Texas into central Coahuila, Mexico.

Grusonia vilis (Rose) H. Robinson 1973

Opuntia vilis Rose 1909, Corynopuntia vilis (Rose) F. M. Knuth 1935

Plants very small, creeping, forming mats 10-15 cm (3.9-5.9 in) high, to several meters wide. Stem segments prostrate, becoming ascending or erect, the terminal segments vertical and club shaped, turgid, pale green, to 5 cm (2 in) long, with low tubercles. Areoles with white wool when young. Principal spines on the prostrate portions reddish with white tips, round in cross section, somewhat rough, on the club-shaped segments reddish with yellowish white tips, 1-4 cm (0.4-1.6 in) long. Secondary or radial spines more than 12. Flowers brilliant purple, to 4 cm (1.6 in) long. Fruits pale green, darkening as they dry, tuberculate, spiny, 2.5-3 cm (1-1.2 in) long. Distribution: Zacatecas and Coahuila, Mexico.

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