Subtribe 3 Echinocereanae

Mostly low, simple or cespitose, terrestrial cacti, the stems 1-jointed or rarely few-jointed, ribbed; areoles borne on the ribs and spiniferous or rarely spineless; flowers always solitary at lateral* areoles, funnelform to campanulate; perianth-segments few to many; fruit smooth or spiny, with few exceptions fleshy and indehiscent or splitting on one side; seeds mostly black.

We recognize 6 genera, all South American except Echinocereus.

This subtribe, while somewhat uniform in its low, usually one-jointed stems, shows great variability in its flowers. Both Chamaecereus and Austrocactus are taken from Cereus of previous authors; Echinocereus has often been considered as a subgenus of Cereus. Echi-nopsis has usually been treated as a distinct genus related to Cereus, Bentham and Hooker, however, treating it as a subgenus of Cereus; in our opinion, it approaches Trichocereus in its flowers, but in habit resembles various genera in the Echinocactanae. Lobivia is segregated from Echinopsis. Rebutia has sometimes been recognized as a genus, but its species have usually been referred to Echinocactus or Echinopsis. The subtribe is nearest the Cereanae, but is also related to the Echinocactanae.

Key to Genera.

Ovary and fruit bearing clusters of spines at areoles.

Stigma-lobes always green; spines all straight 1. Echinocereus (p. 3)

Stigma-lobes red; some spines hooked 2. Austrocactus (p. 44)

Ovary and fruit not spiny.

Spines on tubercles as in Coryphantha; plants globular 3. Rebutia (p. 45)

Spines on ribs.

Plants very small, creeping, forming low clumps 4. Chamaecereus (p. 48)

Plants mostly large, solitary or cespitose.

Flower short-funnelform to campanulate; tube short 5. Lobivia (p. 49)

Flower long-funnelform; tube elongated 6. Echinopsis (p. 60)

1. ECHINOCEREUS Engelmann in Wislizenus, Mem. Tour North. Mex. 91. 1848.

Plants always low, perennial, erect or prostrate, sometimes pendent over rocks and cliffs, single or cespitose, globular to cylindric, prostrate or pendent if elongated; spines of flowering and sterile areoles similar; flowers usually large, but in some species small, diurnal, but in some species not closing at night; perianth campanulate to short-funnelform, scarlet, crimson, purple or rarely yellow, the tube and ovary always spiny; stigma-lobes always green; fruit more or less colored, thin-skinned, often edible, spiny, the spines easily detached when mature; seeds black, tuberculate.

Type species: Echinocereus viridiflorus Engelmann.

The first part of the generic name is from e^Lvog hedgehog, doubtless given on account of the spiny fruit in which Echinocereus is conspicuously different from the true Cereus.

We recognize 60 species. Professor Schumann admitted 39 species in his monograph of 1898, while more than 190 species and varieties have been proposed by other authors.

The genus is confined to the western United States and Mexico. It extends as far east as central Texas, Oklahoma, and Kansas, north to Wyoming and Utah, west to the deserts of southern California, the Pacific coast, and islands of lower California, and south to the City of Mexico.

Echinocereus has often been combined with Cereus. Engelmann, as well as Berger, treated it as a subgenus of Cereus, but Schumann gave it generic rank. As we understand the genus, it is not close to Cereus proper, but is much nearer to some of the other genera.

*Echinocereus baileyi is described as producing flowers from the young growth and appearing terminal; this habit has been observed in other species, but is inconstant.

In habit it simulates the South American genus, Echinopsis, while in flowers and fruits it comes near Erdisia, Bergerocactus, and Wilcoxia. While all the species are low in habit there is great variation in the manner and form of growth. Some are solitary; others grow in flat masses, and others in large rounded mounds. The flowers, while always having a spiny ovary and flower-tube and green stigma-lobes, have considerable variation in the shape and color of perianth-segments and in duration. The flower-buds as well as the young shoots are deep-seated in their origin and do not appear just at the areoles as in most cacti and hence must break through the epidermis when they develop. A somewhat similar result is produced in the flowering of some of the species of Rhipsalis. Echinocereus has been selected as the state flower of New Mexico.

Most plants of Echinocereus do not flower frequently in greenhouse cultivation.

The species are not readily grouped into series; our classification of them is largely artificial, taking flower-color as a more important character than it probably is in nature.

Key to Species

A. Flowers large, usually conspicuous, rarely only 2 to 3 cm. long.

B. Stems covered with long weak bristles or hairs, resembling a small plant of Ceph-

alocereus senilis 1. E. delaetii

BB. Stems variously covered with spines or rarely spineless, never like the above.

C. Flowers scarlet to salmon-colored, opening once, but lasting for several days.

Stems usually weak, often trailing, or at least becoming prostrate; ribs nearly continuous.

Flowers rosy red 2. E. scheeri

Flowers orange-red to salmon-colored.

Flowers 8 to 11 cm. long; wool from areoles on flower-tube long.

Flowers 8 to 10 cm. long; radial spines 9 or fewer 3. E. salm-dyckianus

Flowers is 11 cm. long; radial spines 10 to 12 4. E. huitcholensis

Flowers 6 cm. long or less; wool from areoles on flowers shorter than subtending scale 5. E. pensilis

Stems usually erect and stout; ribs more or less tubercled.

Plants forming large mounds, sometimes with 500 to 800 joints; spines white, long and flexuous 6. E. mojavensis

Plants in much smaller clusters; spines brownish or grayish, not long and flexuous.

Plant body with 12 to 14 ribs 7. E. leeanus

Plant body with 5 to 11 ribs (in one species 12).

Ribs 5 to 8 8. E. triglochidiatus

Axils of flower-scales filled with long cobwebby hairs.

Flowers 5 to 6 cm. long; spines yellowish at first 9. E. polyacanthus

Flowers 3 cm. long; spines reddish at first 10. E. pacificus

Axils of flower-scales bearing short hairs.

Stems elongated and thinner than in E. octacanthus 11. E. acifer

Stems short and thicker than in E. acifer.

Stems pure green when old; central spine 1 12. E. octacanthus

Stems bluish green; central spines several.

Central spines 6; petals acutish 13. E. neo-mexicanus

Central spines mostly 4, sometimes 3 or 5.

Central spines more or less angled, somewhat curved 14. E. conoideus

Central spines terete, straight.

Central spines white or straw-colored 15. E. coccineus

Central spines gray to pinkish 16. E. rosei

CC. Flowers broad, rotate to campanulate, opening in sunlight, closing at night, usually purple, sometimes yellow or greenish yellow, rarely pink or nearly white, unknown in E. standleyi. D. Flowers yellow or greenish white. Ribs not strongly tubercled.

Plants densely cespitose 17. E. maritimus

P lants usual l y solita r y. Ribs very stout.

Ribs 5 to 8; spines on flower-tube and ovary short 18. E. subinermis

Ribs 8 or 9; spines on flower-tube and ovary acicular 19. E. luteus

Ribs low, usually hidden by the spines. Flowers small, 2.5 cm. long or less.

Areoles circular 20. E. chloranthus

Areoles elliptic 21. E. viridiflorus

Flowers large, 5 to 10 cm. long.

Flowers greenish white 22. E. grandis

Flowers yellow-red.

Key to Species—continued.

Central spines in more than 1 row 23. E. dasyacanthus

Central spines in a vertical row 24. E. ctenoides

Ribs strongly tubercled 25. E. papillosus

DD. Flowers purple.

E. Stems weak, slender, and creeping. Stems 2 cm. thick or less.

Areoles distant; spines not interlocking.

Perianth-segments narrowly oblong or linear-oblanceolate 26. E. blanckii

Perianth-segments oblong-erose 27. E. pentalophus

Areoles approximate; spines densely interlocking 28. E. sciurus

Stems 3 to 4 cm. thick 29. E. cinerascens

EE. Stems stout, usually erect or ascending.

F. Areoles elliptic to circular, closely set, often with pectinate spines. Areoles elliptic; spines pectinate. Central spine often very long.

Central spine dark 30. E. adustus

Central spine white 31. E. standleyi

Central spine, if present, short.

Spines of ovary and tube of flower slender and weak, the surrounding hairs long and cobwebby.

Spines variegated 32. E. perbellus

Spines of one color.

Spines strongly pectinate and appressed 33. E. reichenbachii

Spines not strongly pectinate, more or less porrect 34. E. baileyi

Spines of ovary and tube of flower short and stout, the surrounding hairs short. Central spine none.

Stems cylindric 35. E. rigidissimus

Stems globular 36. E. weinbergii

Central spines present 37. E. pectinatus

Areoles circular; spines not pectinate.

Central spines brown, much longer than white radials 38. E. fitchii'

Central spines not longer than radials.

Areoles about 5 mm. apart; spines densely interlocking 39. E. scopulorum

Areoles about 1 cm. apart; spines scarcely interlocking 40. E. roetteri

FF. Areoles nearly circular, not so closely set; spines never pectinate.

Ovary strongly tuberculate 41. E. chlorophthalmu:

Ovary not strongly tuberculate.

Plants strongly angled; flower pinkish 42. E. knippelianus

Plants not strongly angled; flower purple. Central spines none.

Spines 3 to 5; flower-tube and ovary without long wool from the areoles 43. E. pulchellus

Spines 6 to 8; flower-tube and ovary bearing long cobwebby wool from the areoles 44. E. amoenus

Central spines 1 or more.

Central spine 1 45. E. palmeri

Central spines several, much elongated, dagger-like 46. E. brandegeei

Flowers large, 6 to 12 cm. long.

Central spines none 47. E. hempelii

Central spines present.

Central spine solitary, rarely 2.

Spines red at base 48. E. merkeri

Spines not red at base.

Plants stout, erect 49. E. fendleri

Plants weak, becoming prostrate 50. E. enneacanthus

Central spines several. Spines never white.

Spines yellowish brown to red.

Spines short, usually stout, 10 mm. long 51. E. lloydii

Spines, at least some of them, very long 52. E. engelmannii

Spines bluish to blackish 53. E. sarissophorus

Spines usually white or straw-colored.

Ribs 11 to 13.

Flowers campanulate 55. E. conglomeratus

Flowers short-funnelform 56. E. stramineus

AA. Flowers minute, 1.2 cm. long or less 57. E. barthelowanus

1 58. E. mamillatus

AAA. Published species not grouped 59. E. ehrenbergii

3 60. E. longisetus

1. Echinocereus delaetii Gürke, Monatsschr. Kakteenk. 19: 131. 1909.

Cephalocereus delaetii Gürke, Monatsschr. Kakteenk. 19: 116. 1909.

Low, 1 to 2 dm. high, densely cespitose, completely hidden by the long, white, curled hairs; ribs indistinct; areoles closely set, bearing 15 or more white reflexed hairs 8 to 10 cm. long and a few stiff reddish bristles; flowers appearing near top of plant; perianth-segments pink, oblanceo-late, acute; stigma-lobes about 12; ovary covered with clusters of long, white, bristly spines; fruit not seen.

Type locality: Not cited.

Distribution: Known only from Sierra de la Paila, north of Parras, Mexico.

This is the most remarkable species in the genus; in aspect it resembles small plants of Cephalocereus senilis, and owing to this resemblance it was first described as a Cephalocereus. Its flowers, however, are so different from those of that genus that as soon as they were seen the plant was at once transferred to Echinocereus.

The plant is now largely imported into Europe and can be obtained from many dealers; it was named in honor of Frantz de Laet, a Belgian cactus dealer, who had imported many plants from Mexico through Dr. C. A. Purpus and other collectors.

Illustrations: Monatsschr. Kakteenk. 19: 119, as Cephalocereus delaetii; Monatsschr. Kakteenk. 22: 73; Rev. Hort. Belge 40: after 184.

Text-figure 1 is from a photograph of the plant received from M. de Laet.

2. Echinocereus scheeri (Salm-Dyck) Rümpler in Förster, Handb. Cact. ed. 2. 801. 1885.

Cereus scheeri Salm-Dyck, Cact. hort. Dyck.

Cespitose; stems procumbent, prostrate or ascending, decidedly narrowed towards the tip, 10 to 22 cm. long, yellowish green; ribs 8 to 10, rather low, not at all sinuate, somewhat spi-raled; spines 7 to 12, acicular, white with brown or blackish tips; flowers 12 cm. long, rose-red to crimson, with an elongated tube; perianth-segments oblanceolate, acute; fruit not known.

Type locality: Near Chihuahua.

Distribution: Chihuahua, Mexico.

The species was named for Frederick Scheer (1792-1868), who described the cacti for Seemann in the Botany of the Herald.

Lemaire used this name as early as 1868 (Les Cactées 57), but did not formally transfer or describe it, and it is not published or even mentioned by Lemaire in Manuel de l'Amateur de Cactus (1845) as stated in Blühende Kakteen under plate 14. It seems to be a distinct species, related to E. salm-dyckianus, but with differently colored flowers and shorter spines. The variety E. scheeri nigrispinus was used by Scheer in Botany of the Herald 291.

The type of this species seems to have been lost; it was collected by John Potts, a mining engineer, at one time stationed in Chihuahua, Mexico. We know the species only from description and illustrations.

Echinocereus scheeri vars. major and minor (Monatsschr. Kakteenk. 15: 175. 1905) and var. robustior (Monatsschr. Kakteenk. 15: 161. 1905) are only garden forms.

Illustrations: Curtis's Bot. Mag. 132: pl. 8096, as Cereus scheeri; Blühende Kakteen 1: Pl. 14; Schumann, Gesamtb. Kakteen f. 48.

Text-figure 2 is from a part of the second illustration above cited.

3. Echinocereus salm-dyckianus Scheer in Seemann, Bot. Herald 291. 1856.

Cereus salm-dyckianus Hemsley, Biol. Centr. Amer. Bot. 1: 545. 1880.

Echinocereus salmianus Rumpler in Forster, Handb. Cact. ed. 2. 809. 1885.

Cereus salmianus Weber, Dict. Hort. Bois 279. 1894.

Cespitose; stems more or less decumbent, 2 to 4 cm. in diameter, elongated, yellowish green; ribs 7 to 9, low, more or less sinuate; radial spines 8 or 9, acicular, yellowish, about 1 cm. long; central spine solitary, porrect, a little longer than the radials; flowers orange-colored, 8 to 10 cm. long, narrow, the tube elongated, the areoles of the flower-tube and ovary bearing white bristly spines and cobwebby hairs; perianth-segments oblanceolate to spatulate; filaments dark red; style longer than the stamens; fruit not seen.

Type locality: Near Chihuahua.

Distribution: States of Chihuahua and Durango, Mexico.

This species is in cultivation in Europe, and Dr. Rose saw it in flower at La Mortola in 1912; it was also collected by Dr. E. Palmer in Durango in 1906 (No. 205).

Fig. 2.—Echinocereus scheeri. Fig. 3.—Echinocereus salm-dyckianus.

We have not been able to see the type specimen and it is probably not in existence.

The specific name commemorates Joseph Franz Salm-Reifferscheid-Dyck (1773-1861), author of several important cactus treatises. He was the most distinguished cactologist of his time and possessed at his estate at Düsseldorf, Germany, one of the largest cactus collections ever brought together. Unfortunately, after his death the collection was permitted to disintegrate and most of his types were lost or thrown away.

Hybrids between E. salm-dyckianus and Heliocereus speciosus and with Epiphyllum species are reported.

Illustrations: Blühende Kakteen 1: pl. 29; Monatsschr. Kakteenk. 3: 129; Wildeman, Icon. Select. 6: pl. 202.

Text-figure 3 shows a part of the first illustration above cited.

4. Echinocereus huitcholensis (Weber) Gürke, Monatsschr. Kakteenk. 16: 23. 1906.

Cereus huitcholensis Weber, Bull. Mus. Hist. Nat. Paris 10: 383. 1904.

Plants 4 to 6 cm. long, 2 to cm. in diameter; radial spines 10 to 12; central spine usually solitary; flowers 11 cm. long or less, narrow, with a pronounced tube; color of perianth-segments uncertain but perhaps orange, as in E. salm-dyckianus; spines on ovary and tube weak, acicular; areoles of flower-tube bearing long cobwebby hairs.

Type locality: Sierra de Nayarit, Jalisco, Mexico.

Distribution: Known only from the type locality.

Weber described it as a Cereus, but without seeing flowers or fruit, basing it on the collection of L. Diguet of 1900. Three sheets of this collection are in the herbarium of the Museum of Paris, and with them are a flower, immature fruit, and two plants of another species, perhaps undescribed.

5. Echinocereus pensilis (K. Brandegee) J. A. Purpus, Monatsschr. Kakteenk. 18: 5. 1908.

Cereus pensilis K. Brandegee, Zoe 5: 192. 1904.

More or less cespitose; the stems often erect, 30 cm. high or, when growing on cliffs, hanging, and then nearly 2 meters long, 3 to 4 cm. in diameter; ribs 8 to 10, low; areoles about 10 mm. apart; spines needle-like, at first yellow, becoming reddish gray, the longest not over 2 cm. long; radial spines about 8; central spine 1; flowers orange-red, narrow, 5 to 6 cm. long; areoles on ovary and tube bearing short, yellow or white wool and chestnut-colored bristly spines; fruit globular, 1.5 to 2 cm. in diameter; seeds black, rugose, very oblique at base.

Type locality: Sierra de la Laguna, Lower California.

Distribution: High mountains of the Cape region of Lower California.

This species is unlike most of the known Lower Californian species in that it grows in the high mountains of the Cape region and is in fact more closely related to the species of the mountains of the United States and Mexico than to any of its near neighbors. It is a beautiful plant; Dr. Rose saw it in flower in Darmstadt in June 1912, where it was grown from a hanging basket.

The type specimen is in the Brandegee Herbarium at the University of California, but a duplicate and a photograph of the type are preserved in the United States National Herbarium.

Illustration: Monatsschr. Kakteenk. 18: 3.

6. Echinocereus mojavensis* (Engelmann and Big-

803. 1885.

Cereus mojavensis Engelmann and Bigelow, Proc.

Cereus bigelovii Engelmann, Pac. R. Rep. 4: pl. 4, f.

Cespitose, growing in massive clumps, often forming mounds, with hundreds of stems (500 to 800 have been recorded); stems globose to oblong, 5 to 20 cm. long, pale green; ribs 8 to 13, 5 to 6 mm. high, but becoming indistinct on old parts of stem, somewhat undulate; areoles circular, about 1 cm. apart; spines all white, or in age gray; radial spines about 10, acicular, spreading, curved, I to 2.5 cm. long; central spine subulate, porrect or somewhat spreading, often weak, 3 to 5 cm. long; flowers rather narrow, 5 to 7 cm. long, crimson; perianth-segments broad, obtuse or even retuse; areoles on ovary with white felt and short acicular spines; fruit oblong, 2.5 to 3 cm. long.

Type locality: On the Mojave River in California.

Distribution: Southeastern California to Nevada and Utah, western Arizona, and reported from northwestern Mexico.

* This species was named for the Mojave Desert, California, where it was first found. The specific name is sometimes incorrectly spelled mohavensis. Munz and Johnston report that the flowers are "pale scarlet tinged with nopal-red."

Fig. 4.—Echinocereus mojavensis.

Engelmann compares this species with E. fendleri and in that relationship most writers have since treated it, but in its habit and shape of flower it suggests a closer relationship to the scarlet-flowered species such as E. polyacanthus.

Cereus bigelovii was doubtless the first name applied to this plant, but for some reason it was afterward changed, although not in the case of the legend for the first illustration cited below.

Illustrations: Pac. R. Rep. 4: pl. 4, f. 8, as Cereus bigelovii; Curtis's Bot. Mag. 126: pl. 7705, as Cereus mojavensis.

Text-figure 4 is from a photograph obtained through S. B. Parish, taken by Dr. P. A. Munz near Pinos Wells, southern edge of the Mojave Desert, altitude 1,335 meters.

7. Echinocereus leeanus (Hooker) Lemaire in Förster, Handb. Cact. ed. 2. 828. 1885.

Cereus leeanus Hooker in Curtis's Bot. Mag. 75: pl. 4417. 1849. Echinocereus multicostatus Cels in Förster, Handb. Cact. ed. 2. 834. 1885. Echinocereus leeanus multicostatus Schumann, Gesamtb. Kakteen 289. 1898.

Plant erect, about 3 dm. high, 1 dm. thick at base, tapering gradually toward the top, simple so far as known; ribs 12 to 14, acute, bearing rather closely set areoles; spines about 12, acicular, very unequal in length, the central and longest about 2.5 cm. long; flowers brick-red, 5 to 6 cm. long; inner perianth-segments somewhat spreading, spatulate to obovate, 3 cm. long, acute; filaments elongated, quite as long as the style.

Type locality: Northern Mexico.

Distribution: Mexico, but range undetermined.

The only herbarium specimens so-named which we have seen are two sheets in the herbarium of the Berlin Botanical Garden, representing two flowers which are scarlet, slender, 9 cm. long, with pale brownish spine-clusters intermixed with cobwebby white hairs. We have studied a plant sent from the Berlin Botanical Garden to the New York Botanical Garden which has not flowered.

This species differs from its relatives in having more numerous ribs.

The type specimen was presented to the Royal Botanic Gardens at Kew about 1842 by Mr. James Lee, owner of the Commercial Gardens at Hammersmith, near London, for whom it was named, and it is said to have come to him from France. Echinocereus pleiogonus and E. multicostatus, which Schumann refers here as synonyms, were described about the same time from specimens introduced into France by Cels. It is not at all unlikely that all three had a common origin. Cereus multicostatus Cels (Schumann, Gesamtb. Kakteen 288. 1898) is only a catalogue name.

A small specimen of this species obtained from Berlin in 1914 has been grown at the New York Botanical Garden, presumably correctly identified. The young areoles are white-woolly; the spines are acicular, the outer ones white, the central ones with brown tips.

Illustrations: Curtis's Bot. Mag. 75: pl. 4417; Gard. Mag. Bot. 2: pl. facing 81, as Cereus leeanus; Schumann, Gesamtb. Kakteen f. 49.

Text-figure 5 is from a photograph of the first illustration above cited.

8. Echinocereus triglochidiatus Engelmann in Wislizenus, Mem. Tour North. Mex. 93. 1848.

Cereus triglochidiatus Engelmann in Gray, Pl. Fendl. 50. 1849.

Cereus gonacanthus Engelmann and Bigelow, Proc. Amer. Acad. 3: 283. 1856.

Cereuspaucispinus Engelmann, Proc. Amer. Acad. 3: 285. 1856.

Cereus hexaedrus Engelmann and Bigelow, Proc. Amer. Acad. 3: 285. 1856.

Echinocereus paucispinus Rümpler in Förster, Handb. Cact. ed. 2. 794. 1885.

Echinocereus gonacanthus Rümpler in Förster, Handb. Cact. ed. 2. 806. 1885.

Echinocereus hexaedrus Rümpler in Förster, Handb. Cact. ed. 2. 807. 1885.

Echinocereus paucispinus triglochidiatus Schumann, Gesamtb. Kakteen 281. 1898.

Echinocereus paucispinus gonacanthus Schumann, Gesamtb. Kakteen 281. 1898.

Echinocereus paucispinus hexaedrus Schumann, Gesamtb. Kakteen 281. 1898.

Always cespitose, with few or many simple stems, these 2 to 6 dm. long, 5 to 8 cm. in diameter, deep green, erect or spreading, 5 to 8-ribbed; spines 3 to 8, various, nearly terete to strongly angled, when young reddish to yellow, but gray in age, usually spreading, often all radial, 3 cm. long or less; flowers scarlet, 5 to 7 cm. long; perianth-segments oblong, obtuse, 3 cm. long; areoles on the flower-tube and ovary few, white-felted, the subtending scales small and red; spines on ovary and flower-tube few, red and white; fruit at first spiny, but in age smooth, bright red, 3 cm. in diameter; seeds 1.6 mm. in diameter or less.

Fig. 6 and 7.—Echinocereus triglochidiatus.

Type locality: Wolf Creek, New Mexico.

Distribution: Western Texas, New Mexico, and Colorado.

The species here described is very variable as to habit and number and kind of spines, and has generally been separated into three or four species; Schumann treated it as a single species with three varieties; we have recognized only a single species, but it is possible that Echinocereus paucispinus which has nearly terete spines should be restored for certain plants in Texas.

Echinocereus monacanthus Heese (Gartenflora 53: 215. f. 32, with wrong legend) may belong here. It is a small one-jointed plant, 10 cm. long, with 7 ribs, and bears but a single spine at an areole. The flowers and fruit are unknown. The plant is said to be a native of Mexico and Texas.

Illustrations: Pac. R. Rep. 4: pl. 5, f. 2, 3, as Cereus gonacanthus; Pac. R. Rep. 4: pl. 5, f. 1, as C. hexaedrus; Cact. Mex. Bound. pl. 56, as C. paucispinus; Schelle, Handb. Kakteenk. 137. f. 66; Blühende Kakteen 3: pl. 124; Rümpler, Sukkulenten 139. f. 74; Förster, Handb. Cact. ed. 2. 793. f. 102, as Echinocereus paucispinus; N. M. Agr. Exp. Sta. Bull. 78: pl. 16, as E. gonacanthus; Pac. R. Rep. 4: pl. 4, f. 6, 7, as Cereus triglochidiatus; Cact. Journ. 2: 18, as Echinocereus paucispinus flavispinus.

Text-figure 6 is from a photograph of a plant sent to the New York Botanical Garden by Dr. Rose in 1913, from near Las Vegas, New Mexico; text-figure 7 shows a flowering plant sent to Washington from near Kerrville, Texas, by Mr. B. Mackensen in 1912.

Fig. 8.—Flower of Fig. 9.—Flowering Fig. 10.—Flower of Fig. 11.—Top of joint of Echinocereus

Echinocereus polya- branch of Echinocereus Echinocereus neo- conoideus.

canthus. X 0.6. polyacanthus X 0.6. mexicanus. X0.6.

9. Echinocereus polyacanthus Engelmann in Wislizenus, Mem. Tour North. Mex. 104. 1848.

Cereuspolyacanthus Engelmann in Gray, Pl. Fendl. 50. 1849.

Cespitose, forming clumps of 20 to 50 stems, pale green but often tinged with red; ribs usually 10, low; areoles approximate; spines gray when old, at first pale yellow, becoming more or less purplish; radial spines about 12; centrals 4, straight, elongated; flowers crimson, 6 cm. long; spines on ovary and flower-tube yellow, intermixed with cobwebby wool; fruit and seeds unknown.

Fig. 8.—Flower of Fig. 9.—Flowering Fig. 10.—Flower of Fig. 11.—Top of joint of Echinocereus

Echinocereus polya- branch of Echinocereus Echinocereus neo- conoideus.

canthus. X 0.6. polyacanthus X 0.6. mexicanus. X0.6.

9. Echinocereus polyacanthus Engelmann in Wislizenus, Mem. Tour North. Mex. 104. 1848.

Cereuspolyacanthus Engelmann in Gray, Pl. Fendl. 50. 1849.

Cespitose, forming clumps of 20 to 50 stems, pale green but often tinged with red; ribs usually 10, low; areoles approximate; spines gray when old, at first pale yellow, becoming more or less purplish; radial spines about 12; centrals 4, straight, elongated; flowers crimson, 6 cm. long; spines on ovary and flower-tube yellow, intermixed with cobwebby wool; fruit and seeds unknown.

Type locality: Cosihuiriachi, Chihuahua.

Distribution: Chihuahua and Durango, Mexico, to western New Mexico and southeastern Arizona.

Echinocereus polyacanthus was described by Dr. Engelmann in 1848, based upon specimens collected by Dr. A. Wislizenus at Cosihuiriachi, a small mining town west of the city of Chihuahua. The next year Dr. Engelmann transferred it along with the other species of his genus, Echinocer-eus, to Cereus, and in 1859, in his report on the Cactaceae of the Mexican Boundary, redescribed and illustrated the species; the specimens used by him for this report, however, were largely from Texas and New Mexico, and this additional material represents a quite distinct species. In order to prove this point Dr. Rose in 1908 visited Cosihuiriachi, the type locality, and collected living and herbarium specimens which were found to be specifically distinct from the so-called E. polyacanthus from the El Paso region which now bears the name E. rosei Wooton and Standley. The habit of the two species is similar, but the armament is somewhat different and the flowers of the true Echinocereus polyacanthus produce an abundance of wool in the axils of the scales which is

Fig. 12.—Echinocereus polyacanthus.

lacking in the other species. The distribution of this species is much more restricted than has usually been given for it; it has been reported from Texas to California and as far south as La Paz, Lower California. The plant illustrated on plate 66 of Blühende Kakteen as this species must be referred elsewhere.

Illustrations: Tribune Hort. 4: pl. 139, as Echinocereus polyacanthus var.; Förster, Handb. Cact. ed. 2. f. 101; (?) Monatsschr. Kakteenk. 15: 41; (?) 17: 169; Schelle, Handb. Kakteenk. 139. f. 67; Shreve, Veg. Des. Mt. Range pl. 24; Bull. Torr. Club 35: 83. f. 1; Cact. Journ. 1: 89.

Figure 8 shows a flower of an herbarium specimen collected by Dr. Palmer near Madera, Chihuahua, in 1908; figure 12 is from a photograph of a plant collected by Professor Lloyd in the Santa Catalina Mountains, Arizona.

10. Echinocereus pacificus (Engelmann).

Cereusphoeniceuspacificus Engelmann, West Amer. Sci. 2: 46. 1886.

Cereuspacificus Coulter, Contr. U. S. Nat. Herb. 3: 397. 1896.

Cespitose, growing in clumps 30 to 60 cm. in diameter, sometimes containing 100 stems, these 15 to 25 cm. long, 5 to 6 cm. in diameter; ribs 10 to 12, obtuse; spines gray, with a reddish tinge; radial spines 10 to 12, 5 to 10 mm. long; central spines 4 or 5, the longest sometimes 25 mm. long; flowers deep red, rather small, about 3 cm. long; areoles on ovary and flower-tube bearing long tawny wool and reddish-brown bristly spines; fruit spiny.

Type locality: Todos Santos Bay, Lower California.

Distribution: Northern Lower California, recorded, apparently erroneously, from farther south.

Although the type is from the coastal hills we are inclined to refer here Mr. Brande-gee's plant from the San Pedro Martir, collected May 5, 1893; the specimen shows flowers and a spine-cluster.

Mr. Brandegee's plant from Comondu Cliffs, also referred hereby Coulter, may belong elsewhere; it is without flowers, however, and we are uncertain of its relationship. The spines are long and acicular and Mr. Brandegee's notes state that the stems are not dense but sometimes hang from the rocks.

Figure 9 shows a small flowering branch of an herbarium specimen collected by C. R. Orcutt in northern Lower California in 1883.

11. Echinocereus acifer (Otto) Lemaire in Förster, Handb. Cact. ed. 2. 798. 1885.

Cereus acifer Otto in Salm-Dyck, Cact. Hort. Dyck. 1849. 189. 1850.

? Echinopsis valida densa Regel, Gartenflora 1: 295. 1852.

Echinocereus acifer tenuispinus Jacobi in Förster, Handb. Cact. ed. 2. 798. 1885.

Echinocereus acifer brevispinulus Jacobi in Förster, Handb. Cact. ed. 2. 798. 1885.

Echinocereus durangensis Rümpler in Förster, Handb. Cact. ed. 2. 799. 1885.

Echinocereus durangensis nigrispinus Rümpler in Förster, Handb. Cact. ed. 2. 800. 1885.

Echinocereus durangensis rufispinus Rümpler in Förster, Handb. Cact. ed. 2. 800. 1885.

Echinocereus acifer trichacanthus Hildmann, Monatsschr. Kakteenk. 1: 44. 1891.

Echinocereus acifer durangensis Schumann, Gesamtb. Kakteen 287. 1898.

Echinocereus acifer diversispinus Schumann, Gesamtb. Kakteen 287. 1898.

Cespitose, glossy green, erect; ribs 10, strongly tubercled; radial spines 5 to 10, 10 to 16 mm. long, pale brownish, bulbose and purplish at base; centrals 4 (Schumann says 1), stout, purplish brown, the three upper erect, the lower and stouter one subdeflexed; flowers scarlet.

Type locality: Not cited.

Distribution: Durango and Coahuila, according to Professor Schumann.

Professor Schumann recognized three varieties, based chiefly on the differences in the spines.

We have studied a small plant secured from the Berlin Botanical Garden.

The illustration in Blühende Kakteen cited below shows a plant with almost continuous ribs and one stout central spine. It presumably represents a different species.

Echinocereus acifer was mentioned by Lemaire in 1868 (Les Cactées 57), but the name was not published at that place.

Illustrations: Monatsschr. Kakteenk. 1: 44, as Echinocereus acifer trichacanthus; Förster, Handb. Cact. ed. 2. 637. f. 85; (?) Gartenflora 1: pl. 29, as Echinopsis valida densa; (?) Blühende Kakteen 2: pl. 106; Gartenwelt 9: 410; Gard. Chron. III. 36: 245. f.

12. Echinocereus octacanthus (Mühlenpfordt).

Echinopsis octacantha Mühlenpfordt, Allg. Gartenz. 16: 19. 1848.

Cereus roemeri Engelmann in Gray, Pl. Fendl. 50. 1849. Not Mühlenpfordt, 1848.

Echinocereus roemeri Rümpler in Förster, Handb. Cact. ed. 2. 792. 1885.

Cereus octacanthus Coulter, Contr. U. S. Nat. Herb. 3: 395. 1896.

Cespitose, with many simple joints; joints ovoid, yellowish green, 7 to 10 cm. long, 5 to 7 cm. in diameter; ribs 7 to 9, obtuse, somewhat tubercled; areoles when young white-woolly, in age naked, 8 to 16 mm. apart; spines rigid, grayish brown; radial spines 7 or 8, 10 to 24 mm. long; central spine solitary, stouter than the radials, porrect, 2 to 3 cm. long; flowers red, 5 cm. long, remaining open for several days; fruit unknown.

Type locality: Northern Texas.

Distribution: Known to us definitely only from northwestern Texas, but reported by Coulter from New Mexico and Utah.

M. Cary in 1907 collected at Dolores, Colorado, a plant which comes nearer this species than anything which we have seen, except a plant from Marathon, Texas, which has the armament and flowers called for by the original description. The plant referred to in the following illustration in the Garden is said to have come from northern California but this is undoubtedly an error.

Illustrations: Hort. Franc. II. 7: pl. 22; Garden 13: 291, as Cereus roemeri.

13. Echinocereus neo-mexicanus Standley, Bull. Torr. Club 35: 87. 1908.

Cespitose, but with only a few stout simple joints, 18 to 25 cm. long, 7 cm. in diameter, obtuse, glaucous-green; ribs 11 or 12, obtuse, low, somewhat tuberculate; areoles 1 to 1.5 cm. apart; spines slender, subulate, somewhat spreading; radial spines 13 to 16, the longest only 1.5 cm. long, white or nearly so; central spines 6, the lowest one yellowish to almost white, the others reddish, sometimes 4 cm. long; flowers abundant, appearing near the top or along the sides of the plant, 5 cm. long, narrow and not spreading at the mouth, bright scarlet; perianth-segments acute, firm in texture, 2 cm. long, 6 mm. broad; stamens about half as long as the style; stigmalobes 7; ovary and flower-tube bearing clusters of bristly spines; fruit not known.

Type locality: Mesa west of the Organ Mountains, New Mexico.

Distribution: Known only from the type locality.

Figure 10 is copied from the illustration above cited.

14. Echinocereus conoideus (Engelmann and Bigelow) Rümpler in Förster, Handb. Cact. ed. 2.

807. 1885.

Cereus conoideus Engelmann and Bigelow, Proc. Amer. Acad. 3: 284. 1856.

Echinocereusphoeniceus conoideus Schumann, Gesamtb. Kakteen 283. 1898.

Plants cespitose; joints somewhat conic at apex; ribs 9 to 11; radial spines 10 to 12, slender, rigid; central spines 2.5 to 8 cm. long, generally 5. cm. long; flowers 6 cm. long, scarlet, slender; ovary and flower-tube spiny.

Type locality: On the Upper Pecos, New Mexico.

Distribution: Southeastern New Mexico and western Texas.

The species is closely related to Echinocereus coccineus, perhaps not specifically distinct from it.

Coulter takes for this species the name Cereus roemeri Mühlenpfordt (Allg. Gartenz. 16: 19. 1848), which we refer to E. coccineus.

Illustrations: Pac. R. Rep. 4: pl. 4, f. 4, 5, as Cereus conoideus; N. Mex. Agr. Exp. Sta. Bull. 78: pl. [17]; Bull. Torr. Club 35: 8. f. 2.

Figure 11 is copied from the first illustration above cited.

15. Echinocereus coccineus Engelmann in Wislizenus, Mem. Tour North. Mex. 94. 1848.

Cereus roemeri Mühlenpfordt, Allg. Gartenz. 16: 19. 1848.

Cereus coccineus Engelmann in Gray, Pl. Fendl. 50, 51. 1849. Not Salm-Dyck, 1828. ? Echinopsis valida densa Regel, Gartenflora 1: 295. 1852. Cereus mojavensis zuniensis Engelmann, Proc. Amer. Acad. 3: 281. 1856. Cereusphoeniceus Engelmann, Proc. Amer. Acad. 3: 284. 1856. Echinocereus phoeniceus Rümpler in Förster, Handb. Cact. ed. 2. 788. 1885. Echinocereus phoeniceus albispinus Rümpler in Förster, Handb. Cact. ed. 2. 789. 1885. Echinocereus phoeniceus longispinus Rümpler in Förster, Handb. Cact. ed. 2. 789. 1885. Echinocereus phoeniceus rufispinus Rümpler in Förster, Handb. Cact. ed. 2. 789. 1885. Echinocereus krausei De Smet in Förster, Handb. Cact. ed. 2. 789. 1885. Echinocereus mojavensis zuniensis Rümpler in Förster, Handb. Cact. ed. 2. 803. 1885. Echinocereus phoeniceus inermis Schumann, Monatsschr. Kakteenk. 6: 150. 1896. Echinocereus roemeri Rydberg, Bull. Torr. Club 33: 146. 1906. Not Rümpler, 1885.

Usually densely cespitose, often forming large mounds a meter in diameter, containing sometimes 200 simple stems, these 2 dm. high or less, 3 to 5 cm. in diameter; ribs 8 to 11, somewhat tubercled; radial spines acicular, 8 to 12, 1 to 2 cm. long, usually white; central spines several, longer and stouter than the radials, usually yellowish or whitish but in some specimens reddish or blackish; flowers crimson, 5 to 7 cm. long; perianth-segments broad, obtuse or retuse; areoles on flower and ovary felted and bearing short white bristly spines.

Type locality: About Santa Fe, New Mexico.

Distribution: New Mexico and Arizona to Utah and Colorado.

Schumann describes and figures a plant entirely without spines, but whether it is common or not we do not know. This was published as a variety, E. phoeniceus inermis (Monatsschr. Kakteenk. 6: 150. 1896), but it often passes as E. inermis, although never described as such. Some years ago such a plant was sent to Washington from Utah by Mr. M. E. Jones and is still growing in the Cactus House, but it has not since flowered.

Coulter has combined this species with Mammillaria aggregata Engelmann (Emory, Mil. Reconn. 157. f. 1. 1848) taking both up as Cereus aggregatus Coulter (Contr. U. S. Nat. Herb. 3: 396. 1896), but we do not believe that they are the same; Rydberg has used the name Echinocereus aggregatus (Bull. Torr. Club 33:146. 1906) for this plant.

Echinopsis valida densa Regel (Gartenflora 1: pl. 29. 1852; also Förster, Handb. Cact. ed. 2. f. 85) is referred by Schumann to both Echinocereus acifer and E. phoeniceus (Gesamtb. Kakteen 239, 283). To us it suggests E. fendleri, although it has differently colored flowers.

Illustrations: ? Gartenflora 1: pl. 29, as Echinopsis valida densa; Gartenwelt 1: 85, 89; 4: 159, as Cereus phoeniceus; Gartenwelt 1: 89, as Cereus phoeniceus inermis; Gartenwelt 4: 159, as Echinocereus phoeniceus inermis; Curtis's Bot. Mag. 110: pl. 6774, as Cereus paucispinus (fide Schumann); Pac. R. Rep. 4: pl. 4, f. 9, as Cereus bigelovii zuniensis; Gartenwelt 4: 15 7, as Echinocereus phoeniceus; Monatsschr. Kakteenk. 6: 151, as Echinocactus phoeniceus inermis (through typographical error); N. Mex. Agr. Exp. Sta. Bull. 78: pl. [18].

Plate 11, figure 1, shows a flowering plant collected by E. A. Goldman in Arizona; figure 2 shows an open flower.

16. Echinocereus rosei Wooton and Standley, Contr. U. S. Nat. Herb. 19: 457. 1915.

Cespitose, forming small compact clumps, the stems 1 to 2 dm. long, 5 to 8 cm. in diameter, sometimes as many as 40; ribs 8 to 11, obtuse; areoles rather closely set; spines pinkish to brownish gray; radial spines about 10, spreading; centrals 4, 4 to 6 cm. long; flowers 4 to 6 cm. long, scarlet; inner perianth-segments broad, obtuse; spines on ovary and flower-tube brownish or yellowish, intermixed with short hairs; fruit spiny.

Type locality: Agricultural College, New Mexico.

Distribution: In mountains and dry hills and sometimes on the mesas of southern New Mexico, western Texas, and adjacent parts of northern Mexico.

BRITTON AND ROSE, VOL. III

PLATE II

BRITTON AND ROSE, VOL. III

PLATE II

Cespitose Cactus Illustration
M. E. Eaton del.

1. Top of flowering plant of Echinocereus coccineus.

2. Flower of the same.

3. Flowering plant of Echinocereus chloranthus.

4. Flowering plant of Echinocereus viridiflorus.

5. Flowering plant of Echinocereus maritimus.

(All natural size.)

THE CACTACEAE.

This species passes generally as E. polyacanthus Engelmann, a Mexican species with which it was confused in the Report on the Cactaceae of the Mexican Boundary, but as stated by Mr. Standley that species "is amply separated by the presence of long, white wool in the areoles of the ovary and fruit."

Illustrations: Cact. Mex. Bound. pl. 54, 55, as Cereus polyacanthus.

Figure 13 is from a photograph of a plant obtained on the Sierra Blanca, Texas, by Rose, Standley, and Russell in 1910, which afterward flowered in the cactus house of the U. S. Department of Agriculture.

Fig. 13.—Echinocereus rosei. Fig. 14.—Echinocereus maritimus.

17. Echinocereus maritimus (Jones) Schumann, Gesamtb. Kakteen 273. 1898.

Cereus maritimus Jones, Amer. Nat. 17: 973. 1883.

Cereus flaviflorus Engelmann in Coulter, Contr. U. S. Nat. Herb. 3: 391. 1896.

Echinocereus flaviflorus Schumann, Gesamtb. Kakteen 274. 1898.

Decidedly cespitose, often forming clumps 60 to 90 cm. broad and 30 cm. high, sometimes containing 200 joints; individual joints globose to short-cylindric, 5 to 16 cm. long; ribs 8 to 10; areoles 10 to 12 mm. apart; radial spines about 10, spreading; central spines 4, stout and angled, 2.5 to 3.5 cm. long; flowers small, including the ovary 3 to 4 cm. long, arising from near the top of the plant, light yellow; inner perianth-segments oblanceolate, rounded at apex; ovary not very spiny; fruit not seen.

Type locality: Ensenada, Lower California.

Distribution: West coast of Lower California.

This is a low, coastal species, perhaps extending all along the west coast of central Lower California. It was first found by Marcus E. Jones at Ensenada and was recently collected at the same locality by Ivan M. Johnston, April 7, 1921 (No. 3007). Dr. Rose found it in abundance about San Bartolomé Bay and introduced a great quantity into cultivation. It has frequently flowered both in the New York Botanical Garden and at Washington. A specimen of the original collection is preserved in the U. S. National Museum.

The name Cereus glomeratus Engelmann, unpublished, is cited by Orcutt (West Amer. Sci. 13: 28. 1902) as a synonym of Echinocereus maritimus. The name is also used by Schumann (Gesamtb. Kakteen 274. 1898), but it is not found in any of Engelmann's works.

Plants collected by Mr. C. R. Orcutt in Lower California have been referred as Echinocereus orcuttii (Kew Bull. Misc. Inf. 1921: 36), without description.

Illustration: Cact. Journ. 2: 123.

Plate 11, figure 5, shows a plant in flower. Figure 14 is from a photograph of a plant of the same collection brought to Washington from San Bartolomé Bay, Lower California, in 1911 by Dr. Rose (No. 16189).

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