1. Fruits dry at maturity, sometimes tardily so, tan to gray, usually bearing spines and apical flanges; plants low growing, less than 0.5 m (except O. aureispina) | → 2 |
1. Fruits fleshy or juicy (bleeding) at maturity, green, yellow, orange, pink, red, or purple; flowers and fruits usually spineless, rarely bearing spines and without apical flanges; plants low growing to treelike, to 8 m | → 10 |
2. Shrubs or trees with short trunks, erect, to 1(-1.5) m; fruits and stem segments glabrous; spines well distributed on stem segments, usually 4-12 per areole, bright yellow to orange, aging tan to blackish, not gray; inner tepals yellow with orange to red bases; Brewster County, Texas | O. aureispina |
2. Shrubs low, forming clumps or mats, only 1-3 stem segments tall, trunkless; fruits and stem segments glabrous or minutely papillose (use hand lens); spines variously distributed on stem segments, 0-20 per areole, of various colors including gray; inner tepals yellow, yellow with red bases, apricot, pink, to magenta; widespread | → 3 |
3. Seeds subspheric, 6.5-7 mm thick, girdle projecting 0.5-1 mm; inner tepals pink to magenta (rarely white); filaments red-magenta (rarely pale); stigma lobes white to cream; stem segments and fruits minutely puberulent (rarely glabrous), spineless except var. treleasei (Kern County, California), with areoles bearing 0-3(-8) spines, stout, to 10 mm, pale yellow | O. basilaris |
3. Seeds flattened, 1.5-3.5 mm thick; girdle projecting 1-3.5 mm; inner tepals yellow to red-magenta; filaments white, yellow, or red-magenta; stigma lobes green; stem segments and fruits glabrous or minutely puberulent, usually spiny | → 4 |
4. Areoles in distal 1/3 of each stem segment bearing 0-4 spines, spines absent from basal 50-100% of areoles of segment; flowers and fruits spineless or nearly so; stem segments often minutely papillose (use hand lens); Arizona-Utah boundary region | → 5 |
4. Areoles in distal 1/3 of each stem segment bearing 6-20 spines, spines absent from 0-40% of stem segment; flowers and fruits very spiny to nearly spineless; fruits and stems glabrous; widespread | → 7 |
5. Areoles in distal 1/3 of stem segment bearing 0(-1) spines, spines absent from basal 95-100% of stem segment; fruits spineless | O. aurea |
5. Areoles in distal 1/3 of stem segment bearing 1-4 spines (spines absent from basal 50-80% of stem segment); fruits spiny | → 6 |
6. Areoles (4-)7-8 per diagonal row across midstem segment; glochids conspicuous; longest spines per areole (35-)50-70 mm; fruit areoles 16-34. | O. pinkavae |
6. Areoles 3-5 per diagonal row across midstem segment; glochids inconspicuous; longest spines per areole 8-24 mm; fruit areoles 12-22. | O. fragilis |
7. Stem segments 6-27 × 5-18 cm, obovate to circular (or, if elliptic, then stem segments longer than 9 cm), firmly attached | O. polyacantha |
7. Stem segments (1.5-)2-9 × 5 cm, elliptic to narrowly obovate, flattened to sometimes nearly cylindric, firmly attached or readily detached | → 8 |
8. Depressed spines at base of areoles 0-3, 1-3 mm; longest spines 8-24 mm; stem segments easily detached; widespread | O. fragilis |
8. Depressed spines at base of areoles 5-7, 4-20 mm; longest spines 20-60 mm; stem segments sometimes easily detached (in O. ×columbiana) | → 9 |
9. Spines in most areoles; fruit top-shaped to barrel-shaped, usually less than 2 times longer than wide; lower Snake and Columbia River basins | O. ×columbiana |
9. Spines in all or only distal areoles of stem segment; fruit cylindric, often 2 times longer than wide; e New Mexico and w Texas | O. polyacantha |
10. Plants with stem segments spineless or bearing only vestigial spines (sometimes taxa have spineless individuals that cannot be safely identified without flowers and/or fruits—avoid sterile specimens or those from unhealthy or infested plants; see also spineless plants of dry-fruited species above in key) | → 11 |
10. Plants with at least some stem segments bearing spines | → 25 |
11. Fruits and stem segments puberulent (use hand lens) or velvety; trees or shrubs | → 12 |
11. Fruits and stem segments glabrous or glaucous; trees, shrubs, or forming low clumps | → 13 |
12. Stem segments bright green; areoles (9-)11-16 per diagonal row across midstem segment; usually low shrubs to l m, with many small stem segments; escapes from cultivation | O. microdasys |
12. Stem segments grayish green; areoles 8-13 per diagonal row across stem segments; low to medium-sized shrubs to 1.5 m, with fewer, larger stem segments; Texas | O. rufida |
13. Glochids inconspicuous on midstem segment areoles (use hand lens), short in crescent at adaxial edge of areole or covered by white cottony hairs in subapical region; plants known from cultivation or occasionally as escapes | → 14 |
13. Glochids usually conspicuous on midstem segment areoles, 3-5+ mm, densely to sparsely arranged; native taxa | → 15 |
14. Plants trees, 3-6 m, with well-developed trunks, to 30-45 cm diam.; stem segments broadly oblong to ovate to narrowly elliptic, usually (20-)40-60 × 2-3+ cm; glochids along adaxial margin of areoles | O. ficus-indica |
14. Plants low, spreading shrubs, to 2 m, trunkless; stem segments obovate to ovate or circular, 15-25 × 10-20 cm; glochids scattered, usually covered by conspicuous white cottony hairs | O. ellisiana |
15. Plants forming low clumps (in Florida specimens may be erect shrubs, sometimes with short trunks); fruits narrowly cylindric with only slightly tapering bases, tardily ripening, fleshy, surfaces red with greenish to reddish pulp | → 16 |
15. Plants small trees or shrubs; fruits barrel-shaped, sometimes stipitate, fleshy or juicy, surfaces reddish to purple with pulp of various colors | → 18 |
16. Stem segments lustrous; stigma lobes white; inner tepals yellow through- out; e, c United States | O. humifusa |
16. Stem segments dull; stigma lobes cream to yellowish; inner tepals yellow with red bases or red throughout; c United States (some overlapping ranges with above) | → 17 |
17. Inner tepals yellow, with red basal portions; spines ± stout (0.5 mm diam. near base); stem segments commonly subcircular to obovate, flabby and cross wrinkled when stressed | O. macrorhiza |
17. Inner tepals red throughout; spines slender (± 0.2 mm diam. near base); stem segments cuneate-obovate to commonly rhombic, ± firm | O. pottsii |
18. Stem segments ± tuberculate, margins scalloped between marginal areoles; fruits juicy, barrel-shaped, tapering below, purplish throughout; primarily se United States coast | O. stricta |
18. Stem segments with margins entire, not scalloped; fruits juicy or fleshy, various in shape and color; widespread | → 19 |
19. Glochids widely and evenly spaced, radiating like a star or pin cushion; Webb County, s Texas, and cultivated | O. aciculata |
19. Glochids not evenly spaced radially, in crescent at adaxial margin and also sometimes in subapical tuft; Texas to California, native | → 20 |
20. Areoles with subapical tuft of glochids poorly developed or absent, in addition to crescent at adaxial margin | → 21 |
20. Areoles with subapical tuft of glochids well developed, in addition to crescent at adaxial margin | → 22 |
21. Glochids moderately dense in adaxial crescent, of ± even lengths along margins of areole; style pink to red | O. ×vaseyi |
21. Glochids crowded in adaxial crescent, increasing in length abaxially along margin of areole; style white | O. chlorotica |
22. Glochids sparse, of irregular lengths, encircling (or nearly so) areole margins, also in subapical region, these often embedded in wool; large shrubs (sometimes with short trunk); fruits juicy | O. engelmannii |
22. Glochids dense, subequal or increasing in length toward adaxial margin of crescent, also dense in subapical tuft; small trees or shrubs; fruits fleshy or juicy | → 23 |
23. Inner tepals yellow throughout; erect shrubs or small trees to 2 m; stem segments lavender to red-purple when under stress, at least around areoles | O. santa-rita |
23. Inner tepals yellow with red bases; erect to trailing shrubs to 1.5 m; stem segments green, reddish, or purplish, particularly when under stress | → 24 |
24. Shrubs erect, erect to decumbent; stem segments purplish when under stress | O. macrocentra |
24. Shrubs low, with decumbent to trailing branches; stem segments green, sometimes reddish when under stress | O. phaeacantha |
25. Fruits and stem segment surfaces puberulent (use hand lens); inner perianth all yellow to orange-red, sometimes apically tinged red; spiny to spineless trees, bearing trunks; escapes from cultivation | tomentosa, O. leucotricha; see discussion under genus |
25. Fruits and stem segments glabrous or glaucous, not puberulent (except for pubescence of areoles); inner perianth yellow, orange, pink red, magenta (rarely white), sometimes red basally; natives and escapes from cultivation | → 26 |
26. Spines (0-)1-3(-5) per areole, strongly barbed; stem segments 2.5-18 cm, easily detached; plants creeping, forming mats or clumps, or dense shrubs to 1 m; inner tepals yellow throughout; se United States, coastal (or slightly inland) and island plants, West Indies | → 27 |
26. Spines 1-18+, not strongly barbed; stem segments of various lengths, not easily detached; habit various; inner tepals yellow throughout or with bases of a different color; widespread | → 28 |
27. Plants creeping or forming mats; stem segments narrowly elliptic to linear, sometimes somewhat cylindric, 2.5-5(-8) × 1.2-2.5 cm; spines 1-2 in distal 3/4 of stem segment areoles; Texas to North Carolina | O. pusilla |
27. Plants forming clumps or low shrubs; stem segments narrowly elliptic, 5-18 × 3-7 cm; spines 1-3 in most areoles; Florida, West Indies. | O. triacantha |
28. Large trees, 3-6 m; trunks well developed, to 45 cm diam.; stem segments 25-60 cm; spines, when present, subulate, white, sometimes basally yellow to brown, erect to spreading; glochids inconspicuous, less than 2 mm; stem segments very thick, obovate, elliptic to oblong; known only from cultivation and escapes | O. ficus-indica |
28. Small trees or shrubs, erect to prostrate, to 2.5 m, usually much shorter; trunk, if present, short, to 25 cm diam.; stem segments rarely to 20-30 cm; spines, if present, subulate or acicular; glochids conspicuous, more than 2 mm; stem segments variable in thickness and shape | → 29 |
29. Shrubs prostrate, trailing or forming clumps (sometimes erect, even rarely short trunked in O. humifusa in Florida) sometimes with enlarged rootstocks; stem segments to 5-11(-17) cm | → 30 |
29. Shrubs or trees, erect to sprawling, several segments tall; stem segments 10+ cm | → 33 |
30. Fruits subspheric or short ovoid; stigma lobes yellow-green to green; flowers broadly spatulate; stem segments fleshy, green; spines usually distributed over distal 30-85% of stem segments; major spines erect to reflexed; minor spines at base of areole 2-8, deflexed | O. tortispina |
30. Fruits elongate, subterete, becoming stipitate-obconic with seed development; stigma lobes white to yellow-cream (rarely tinted green); flowers broadly cupped to rotate; stem segments fleshy to flabby, shiny to dull green; spines absent or only in distal 10-30(-50)% of stem segment; major spines erect to spreading; minor spines 0-2(-3), spreading to deflexed | → 31 |
31. Inner tepals yellow throughout; stigma lobes white; stem segments lustrous, green; spines, if present, 1 or 2(-3) on marginal areoles, dark; major spine porrect, stout, usually terete | O. humifusa |
31. Inner tepals yellow with red bases (occasionally only blushed) or red throughout; stigma lobes cream (or faint tinge of green); stem segments dull dark green; major spines (0-)1-3(-6) in distal areoles, rather stout to slender and flexible, terete or 1 central flattened, white to gray | → 32 |
32. Inner tepals yellow, with red basal portions; spines ± stout (± 0.5 mm diam. near base); stem segments commonly subcircular to obovate, of- ten flabby and cross wrinkled when stressed | O. macrorhiza |
32. Inner tepals yellow, with red basal portions; spines slender (± 0.2 mm diam. near base); stem segments cureate-obovate to commonly rhom- bic, ± firm | O. pottsii |
33. Stem segments somewhat tuberculate, margins scalloped between marginal areoles; fruits juicy, barrel-shaped, tapering below, red-purple to purplish through- out; se United States, primarily coastal | O. stricta |
33. Stem segments not or only slightly tuberculate, margins entire, not noticeably scalloped; fruits juicy or fleshy, various in shape and color; primarily w United States | → 35 |
34. Shrubs, densely branched, to 1 m; stem segments (4-)10-17.5 × (3-)5-7 cm; spines 2-3(-5) per areole, straight, commonly spirally twisted; fruits reddish and fleshy at maturity, fruit areoles 17-21, bearing few spines to ±9 mm; Florida Keys. | O. cubensis |
34. Shrubs, sparsely branched, to 2 m; stem segments 10-25(-40) × 7.5-16 cm; spines absent or 0-11 per areole, straight or curved (not spirally twisted); fruits purplish and juicy at maturity, fruit areoles 6-10, spineless; coastal se United States | O. stricta |
35. Major spines acicular, flexible, 1-8+ per areole, all strongly deflexed (sometimes some spines divergent along edges of stem segments); midstem segment areoles bearing marginal glochids that increase in length toward base of areole; fruits subspheric or barrel-shaped | → 36 |
35. Major spines acicular or subulate, 1-15+ per areole, not all deflexed; midstem segment areoles bearing various other glochid patterns; fruits ellipsoid to spheric, sometimes long tapering at base | → 37 |
36. Erect shrubs to small trees, to 2.5 m; spines all yellow, aging blackish; fruits large, barrel-shaped, 30-60 × 18-40 mm | O. chlorotica |
36. Erect to sprawling shrubs, to 1 m; spines reddish brown with yellow tips, aging blackish; fruits small, subspheric, 15-28 × 12-23 mm | O. strigil |
37. Glochids widely spaced (not dense), at least in most areoles, usually irregular in length; inner tepals of uniform color | → 38 |
37. Glochids densely clustered, in brushy apical tuft of unequal lengths or, more commonly, in adaxial crescent, partially encircling areoles, of equal length or decreasing in length from apex; inner tepals of uniform color or with bases of different color | → 40 |
38. Glochids prominent, radiating in all directions from areoles (like star or pincushion), very widely and evenly spaced; spines 1-3(-8), brown, tipped yellow or mottled brown, basal in areole, usually deflexed; flowers yellow, orange, or red throughout | O. aciculata |
38. Glochids obvious, in adaxial crescent, partially encircling areole margins and usually also in subapical clusters above major spines; spines 1-6(-10) per areole; major spines yellow, darkening with age, sometimes with chalky white coating (yellow spines show through when wetted), and sometimes with red-brown bases; tepals yellow throughout (sometimes pink, buff, or red, rarely white) | → 39 |
39. Shrubs (occasionally with short trunk) spreading to decumbent, to 3 m; major spines subulate, commonly flattened to angular at base, in some varieties chalky white-coated; tepals yellow (sometimes orange, pink, or red, rarely white); fruits juicy, purple, barrel-shaped, often with long tapering bases; widespread | O. engelmannii |
39. Shrubs erect, to 1 m; major spines terete to flattened near base, often curved, yellow, aging red or red-brown, never chalky; tepals yellow to buff; fruits juicy, reddish purple, barrel-shaped, about as wide as long; local in Chisos Moun- tains, w Texas | O. chisosensis |
40. Midstem segment areoles strongly obovate, each bearing many large glochids in brushy adaxial crescent; inner tepals yellow to orange throughout without red centers; divergent spines, dark red to black with strongly differentiated yellow tips; deflexed spines ± chalky white | O. atrispina |
40. Midstem segment areoles subcircular to elliptic to narrowly obovate, each bearing tidy (not brushy) glochids of equal lengths or of decreasing length from apex; inner tepals of 1 color or with differently colored bases; spines divergent and or deflexed, if bearing yellow tips, not in sharp contrast to basal color | → 41 |
41. Areoles with subapical tuft of glochids poorly developed or absent, in addition to adaxial crescent; inner tepals yellow, orange, or red, often bearing red to salmon bases; filaments yellow or orange to pink; style usually pink to red (rarely whitish); stigma lobes green to yellow-green; sw California | → 42 |
41. Areoles with dense subapical tuft (above major spines) of glochids, in addition to adaxial crescent; inner tepals yellow throughout or yellow with red centers (or rarely all reddish); filaments white, cream, or yellowish; style white; stigma lobes whitish or yellowish, sometimes tinged pale green; widespread | → 45 |
42. Shrubs or small trees; fruits red-purple with paler interiors; areoles 8-10 per diagonal row across midstem segment; spines 5-13 per areole, well distributed on stem segments, reflexed, usually curved, yellow; inner tepals yellow throughout | O. oricola |
42. Shrubs (sometimes with short trunks); fruits red-purple throughout; areoles 5-7(-8) per diagonal row across midstem segment; spines 0-11 per areole, well distributed on stem segments to spineless, spreading and reflexed, terete or flattened, straight to slightly curved, yellow ± white coated, often yellow to brown based; inner tepals of various colors, some with different colored bases | → 43 |
43. Spines 4-11 per areole, well distributed on stem segments (rarely spines restricted to apical and basal areoles only); longest spines spreading, generally terete, others reflexed; inner tepals yellow to dull red | O. littoralis |
43. Spines 0-6 per areole, well distributed to absent on stem segments; longest spines spreading to reflexed, generally flattened; others reflexed or very short; inner tepals yellow, orange, deep pink, or dull red, often darker basally | → 44 |
44. Spines 0 or 1-6, distributed over distal 0-70% of stem segments; shrubs sprawling to spreading; stem segments 7-22 cm; inner tepals yellow, orange, or dull red; filaments orange-yellow; style usually pink | O. ×vaseyi |
44. Spines 3-6, distributed over distal 90-100% of stem segment; shrubs sprawling to erect; stem segments 18-35 cm; inner tepals yellow to deep pink; filaments yellow or white; style pink or white | O. ×occidentalis |
45. Inner tepals yellow throughout or yellow with faint reddish blush at bases; spines, if present, yellow, horn colored, or red-brown; spreading, deflexed, or erect, to 60 mm; small trees with erect, short trunks | → 46 |
45. Inner tepals yellow with red bases or reddish throughout; spines, if present, of various colors, placements, and sizes; shrubs erect to sprawling | → 47 |
46. Spines, if present, 1-2 in few distal or marginal areoles per stem segment, deflexed to erect, yellow to horn colored, aging reddish brown, terete, to 40 mm; stem segments thin, often purplish (particularly under stress), at least around areoles; s Ari- zona | O. santa-rita |
46. Spines in most areoles, 4-8+ per areole, spreading to deflexed, reddish yellow to red-brown, subterete to flattened or channeled adaxially, sometimes twisted and curved, to 40-60 mm; stem segments thick, gray-green to yellow-green; nw Ari- zona, California, Nevada | O. ×curvispina |
47. Fruits sparingly spiny in distal areoles, yellow to red, slightly tuberculate, often sterile, shriveling and drying tan; spines 1-5(-8) per areole, well distributed on stem segment, white with red base, becoming dark red-brown with pale tips, often curved and twisted | O. ×spinosibacca |
47. Fruits not spiny, red to purplish, not tuberculate, fleshy to juicy; spines on stem segments, if present, 0-10(-20+) per areole, variously colored and distributed on stem segment | → 48 |
48. Spines 0-15+ per areole, few and at distal areoles or many and evenly distributed on stem segments; major spines usually reddish brown to blackish (or rarely yellow in w Texas), often partly to wholly chalky white, usually terete except at base, 30-170 mm, erect to upward-spreading (appearing to "need a haircut"), rarely also with small, deflexed spines; stem segments often purplish (particularly under stress), at least around areoles | O. macrocentra |
48. Spines (0-)2-8 per areole, in distal 1/3-2/3 areoles of stem segments; major spines gray to tan with red-brown bases or usually red-brown, sometimes blackish, usually flattened, 30-80 mm, spreading or deflexed, also with 1-3+ small, deflexed gray to tan spines; stem segments sometimes reddish under stress | O. phaeacantha |