THE CACTACEAE

DESCRIPTIONS AND ILLUSTRATIONS OF PLANTS OF THE CACTUS FAMILY

BY

N. L. BRITTON anp J. N. ROSE

VotumE I

Tue CaRNEGIE INSTITUTION OF WASHINGTON

A Cactus Desert in Arizona. Photograph by D. T. McDougal.

THE CACTACEAE

DESCRIPTIONS AND ILLUSTRATIONS OF PLANTS OF THE CACTUS FAMILY

BY

N. L. BRITTON anp J. N. ROSE

a

Volume LI .

Tue CarneciE InstiruTIoN oF WASHINGTON tte Wasuincton, 1919

aN

CARNEGIE INSTITUTION OF WASHINGTON PUBLICATION No. 248, VoLumE I

Copies of this Boek were first issued JUN 21 i9fe

PRESS OF GIBSON BROTHERS WASHINGTON

Introduction...... . Order Cactales..... Family Cactaceae. .

CONTENTS.

CC Ce

ee

Key to Tribes........... 0... cece cee eee eee ee Tribe Pereskieae........... 0.0... cece cee eee

——~ Pereskia...... cee ee cee tenes

Key to Species... 20... .. 0c cece ee eee eee Tribe Opuntieae... 00.0... eee eee ee Key to Genera........ 2... cece cece ce eeees Pereskiopsis........... 00 e cece cee e cece eee Pterocactus....... 0.0... eee e ee eee eee

Opuntia..... ccc cece ee eee ees Key to Subgenera and Series.............. Subgenus Cylindropuntia.................

Series Ramosissimae.................-- Series Leptocaules..............-.--6-- Series Thurberianae.................... Series Echinocarpae..............000005 Series Bigelovianae.................--5 Series Imbricatae...............00.005- Series Fulgidae...............-0 00-0 eee Series Vestitae......... 0.0.0 e eee eee Series Clavarioides..............-2..00.- Series Salmianae...............00 0 ee eee Series Subulatae................0000 00 Series Miquelianae...................4. Series Clavatae......... 0... cece eee eee Subgenus Tephrocactus..............506- Series Weberianae................0.005. Series Floccosae..........-. eee eee 4s Series Glomeratae............2-0-2-000- Series Pentlandianae...................

PAGE. Family Cactaceae—continued. Tribe Opuntieae—continued. Opuntia—continued. Subgenus Platyopuntia................... 99 Series Pumilae... 0.0.0.0... cece ee eee 100 Series Curassavica€........... 0.000 c eee 102 Series Aurantiacae...............0.000 106 Series Tunae......... 0.0... cee cece eee 110 Series Basilares............. 0.000 ce eeee 118 Series Inamoenae................0.0005 125 Series Tortispinae.................0025 126 Series Sulphureae.......... 0.020.002 eee 133 Series Strigiles.......... 0... ...0 cee euee 136 Series Setispinae........... 00... cece eee 136 Series Phaeacanthae................... 139 Series Elatiores..............0.000ceuee 149 Series Elatae............. 0... cece eee 156 Series Scheerianae.................00.- 159 Series Dillenianae...................... 159 Series Macdougalianae ................. 169 Series Tomentosae.................000- 172 Series Leucotrichae.............0..0005 174 Series Orbiculatae.................008. 176 Series Ficus-indicae .................00. “4377 Series Streptacanthae.................. 181 Series Robustae............ 2.0. c cece 191 Series Polyacanthae.................... 193 Series Stenopetalae.................... 200 Series Palmadorae.................0005 201 Series Spinosissimae.................00- 202 Series Brasilienses.................0005 209 Series Ammophilae.................... 211 Series Chaffeyanae..................... 213 Grusonia... 0... cece eee eee tees 215 Appendix......... 0... c cee eee eee eee eee e eee 216 Index... 0... ccc cece cnet eeee . 227

PLATE I. PLATE 2.

PLATE 3.

PLATE 4.

PLATE

5 PLATE 6.

PLATE 7.

PLATE 8. PLATE 9.

PLATE 10. PLATE II.

PLATE 12. PLATE 13.

PLATE 14. PLATE 15. PLATE 16. PLATE 17.

PLATE 18. PLATE Ig.

PLATE 20. PLATE 21. PLATE 22. PLATE 23.

PLATE 24. PLATE 25.

PLATE 26. PLATE 27. PLATE 28. PLATE 29. PLATE 30. PLATE 31. PLATE 32. PLATE 33. PLATE 34.

PLATE 35.

PLATE 36.

ILLUSTRATIONS.

(1) Flowering joint of Opuntia spinosissima. Longitudinal section of flower of Op Opuntia spinosissima.

(2, 3) Single flower of Opuntia spinosissima. (4, 5)

untia spinosissima. (6) Cross-section of ovary of (7) Style of Opuntia spinosissima

FACING PLATES _PAGE Cactus Desert in Arizona......... 0. eee cece eee eee ees oe ee eee ete Frontispiece (1) Flowering branch of Pereskia pereskia. (2 and 3) F ruit of Pereskia pereskia. (4) Leafy branch of Pereskia sacharosa (5) Proliferous fruit of Pereskia sacharosa. . weet e ees beeen seen ees 10 (1) Flowering branch of Pereskia grandifolia. (2) Leafy branch of Pereskiopsis chapistle. (3) Leafy branch of Pereskiopsis pititache........ cece eee eee eee e cence eee eens settee 20 (1) Upper part of flowering joint of Nopalea cochenillifera. (2) Upper part of flowering joint of Nopalea auberi. (3) Fruit of Nopalea auberi. (4) Flowering joint of Nopalea dejecta..... 34 . Nopalea auberi as it grows near Mitla, Mexico.............-..00e0+s eee ees bette tenes 38 (1 and 2) Branch of Opuntia mortolensis. (3 and 4) Branch of Opuntia leptocaulis. (5) Flowering branch of Opuntia arbuscula. (6) Flowering branch of Opuntia kleiniae cece eee eee eens 48 (1) Leafy branch of Opuntia kleiniae. (2) Terminal branch of Opuntia vivipara. (3) Branch of Opuntia parryi. (4) Flowering branch of Opuntia echinocarpa. (5) Fruiting branch of Opuntia versicolor. ...... 0... ee ee teen eee e eee e nent eee e eens 50 (1) Type plant of Opuntia vivipara, near Tucson, Arizona. (2) A much branched plant of Opuntia OSs 6) (0) C0) 52 (1) Joint of Opuntia tetracantha. (2, 3, 4, 5) Flowering joint of Opuntia versicolor. (6) Proliferous fruits of Opuntia fulgida....... 0... ee eee eee ne ene e eee neenee 54 (1) Joint of Opuntia tunicata. (2, 3, 4, 5) Joint of Opuntia spinosior..... 0.0.5... 66 (1) Leafy branch of Opuntia imbricata. (2) Flowering branch of Opuntia prolifera. (3, 4) Form of Opuntia alcahes. (5, 6) Opuntia vestita... 0.0... cece cee eee eee teen e ene 68 (1) Clump of plants of Opuntia fulgida. (2) A very open plant of Opuntia spinosior.............. 70 (1) Opuntia exaltata as seen in the highlands of Peru. (2) Clump of Opuntia floccosa as it grows in the valleys of the Andes of eastern Perul....... 0... eee eee e eee enes 76 (1) Flowering branch of Opuntia burrageana. (2) Opuntia cylindrica. (3, 4) Joint of Opuntia stanlyi. (5) Flowering joint of Opuntia macrorhiza........ 0.0... cee ce eee nees 78 (1, 2) Part of joint of Opuntia exaltata. (3) Upper part of joint of Opuntia macrarthra. (4) Upper part of joint of Opuntia tortispina...... 0. ec ce cnc ee eee eee eee eens 80 (1) Top of Opuntia miquelii. (2) Old and young joints of Opuntia invicta. (3) Upper part of joint of Opuntia ignescens.... 2.0... cee eect e eee teen ete teen ee te teenies 98 (1) Joint of Opuntia pascoensis. (2) Joints of Opuntia taylori. (3, 4) Form of Opuntia repens. (5) Flower of Opuntia repens. (6) Flowering joint of Opuntia drummondii............... 102 (1) Two plants of Opuntia drummondii. (2) Joints of Opuntia retrorsa with flower. (3) Joints of Opuntia triacantha. (4) Joint of Opuntia jamaicensis. (5) Section of fruit of Opuntia _ JAMAICENSIS. ee ee ec cece e ence eee teen tent eee ee ne eeeeee 104 (1) Plant of Opuntia jamaicensis. (2, 3) Flower of Opuntia jamaicensis. (4) Longitudinal section of flower of Opuntia jamaicensis. (5, 6) Stamen of Opuntia jamaicensis. (7) Style of Opuntia jamaicensis........ 0... ccc cece cece cece ee eeeeeeneeeaunnteeneeees 112 (1) Flowering joint of Opuntia decumbens. (2) Fruiting joint of Opuntia decumbens. (3) Hybrid 116 Group of hardy Opuntia, mostly Opuntia tortispina, in grounds of New York Botanical Garden.... 126 (1) Joints of Opuntia microdasys. (2) Flowering joint of Opuntia macrarthra. (3) Fruit of Opuntia macrarthra. (4) Seed of Opuntia macrarthra. (5) Flowering joint of Opuntia opuntia..... 128 (1) Flowering joint of Opuntia fuscoatra. (2) Upper part of joint of Opuntia sulphurea. (3) Joint of Opuntia tenuispina......... 0000 ccc cece cece cece ceuececbeeetnvatennsnees 132 (1) Plant of Opuntia santa-rita. (2) Plant of Opuntia discata.......... 000 cc ceeccecccueeeuces 142 (1) Flowering joint of Opuntia atrispina. (2) Flowering joint of Opuntia phaeacantha. (3) Upper part of joint of Opuntia engelmannii.......... 0.00000 e eee eeeeeeens 144 (1) Flowering joint of Opuntia bergeriana. (2) Flowering joint of Opuntia elatior. (3) Flowering joint of Opuntia boldinghii. (4, 5) Joint of Opuntia clata..........00.0 00.0 cece eee eee ees 152 (1) Upper part of fruiting joint of Opuntia schumannii. (2) Flower of Opuntia schumannii. (3) Flowering joint of Opuntia vulgaris. (4) Flowering joint of Opuntia stricta.............. 156 (1) Flowering joint of Opuntia laevis. (2) Flowering joint of Opuntia dillenii. (3) Upper part of flowering joint of Opuntia aciculata.... 2... ccc cece ccc ucccueeueeuenaeees 160 (1) View of Opuntia keyensis. (2) View of Opuntia dillenii...............................0 162 Flowering joint of Opuntia linguiformis. 0.0... cece ccc cece ec ce cece ccceeeeeees 164 Flowering joints of Opuntia lindheimeri. (1) Orange-flowered race. (2) Red-flowered race........ 166 (t) Upper part of flowering joint of Opuntia leptocarpa. (2) Fruit of Opuntia leptocarpa. (3) Flower- a) U ing joint ! Opuntia velutina. (4) Upper part of joint of Opuntia megacantha............ 172 ae part o pont of Opuntia tomentosa. (2) Flowering joint of Opuntia brasiliensis. (3) (1) Part of soint 5 manch of Opuntia brasiliensis. (4) Joint of Grusonia bradtiana............... 174 Joint of Opuntia leucotricha, (2) Part of joint of Opuntia maxima. (3) Joint of Opuntia lasiz . 2 a) PI asiacantha. (4) Joint of Opuntia robusta... 00... 0. oe eee ccc c le cece ee cece eee. 180 I ant of Opuntia fragilis. (2) Flowering branch of Opuntia rhodantha. (3) Flowering joint of Opuntia polyacantha................................ 194

204

FIG.

THE CACTACEAE.

TEXT-FIGURES.

PAGE 1. Hedge of Pereskia pereskia............ 7 2. Tree of Pereskia autumnalis............ II 3. Branches of Pereskia autumnalis........ 12 4. Branch of Pereskia lychnidiflora........ 12 5. Leafy branch of Pereskia nicoyana...... 13 6. Branch of Pereskia zehntneri........... 13 7. Cultivated plant of Pereskia zehntneri. . 14 8. Herbarium specimen of Pereskia moorei. 15 g. Tree of Pereskia guamacho............. 15 10. Flowering branch of Pereskia guamacho. 16 11. Leafy branch and flower of Pereskia colombiana ................0.0 0000s 17 12. Branch and fruit of Pereskia bleo....... 18 13. Fruit of Pereskia bahiensis............. 19 14. Leafy branch of Pereskia bahiensis...... 19 15. Tree of Pereskia bahiensis.............. 20 16. Hedge containing Pereskia grandifolia... 21 17. Branch of Pereskia zinniaeflora......... 21 18. Tree of Pereskia cubensis.............. 22 19. Leafy branch of Pereskia cubensis....... 22 20. Branch and fruit of Pereskia portulaci- 10) 0 23 21. Potted plant grown from a cutting of Pereskiopsis velutina............... 26 22. Branch of Pereskiopsis diguetii......... 27 23. Branch of Pereskiopsis opuntiaeflora .... 27 24. Branch of Pereskiopsis rotundifolia...... 27 25. Shows aclump of Pereskiopsis rotundi- folia... 2. eee ee nee 28 26. Branch of Pereskiopsis porteri.......... 28 27. Branch of Pereskiopsis aquosa.......... 29 28. Leaf of Pereskiopsis kellermanii......... 30 29. Leaf of Pereskiopsis kellermanii......... 30 30. Leaf of Pereskiopsis kellermanii......... 30 31. Seed of Pterocactus hickenii............ 31 32. Plant of Pterocactus hickenii........... 31 33. Branch of Pterocactus fischeri.......... 31 34. Seed of Pterocactus fischeri............. 31 35. Seed of Pterocactus pumilus............ 31 36. Seed of Pterocactus tuberosus.......... 31 37. Plant of Pterocactus tuberosus, showing a very large root.............000000+ 32 38. Potted plant of Pterocactus tuberosus... 33 39. Joint of Nopalea guatemalensis......... 35 40. Joint of Nopalea lutea................. 35 41. Large plant of Nopalea dejecta......... 36 42. Joints of Nopalea dejecta.............. 37 43. Joints of Nopalea karwinskiana......... 37 44. Joint of Nopalea inaperta.............. 37 45. Flower of Tacinga funalis.............. 38 46. Longitudinal section of flower of Tacinga funalis.... 2.0.0... ee eee ee eee 38 47. Section of stem of Tacinga funalis...... 38 48. Tip of young branch of Tacinga funalis. 38 49. Plant of Tacinga funalis, climbing over bushes... 0.2.2.2... 00 eee eee eee 39 50. Plant of Maihuenia valentinil.......... 40 51. Fruit of Maihuenia poeppigii........... 41 52. Joint and flower of Maihuenia brachydel- PHYS... oe eee 41 53. Plant of Maihuenia tehuelches.......... 4! 54. Branch of Opuntia ramossissima . 46 55. Section of stem of Opuntia ramosissima. 46 56. Plant of Opuntia leptocaulis. . 48

FIG.

V PAGE. . Section of stem of Opuntia leptocaulis’.. 48 . Joint of Opuntia caribaea............. 48 . Thicket formed of Opuntia caribaea..... 49 . Clump of Opuntia arbuscula............ 50 . Plant of Opuntia arbuscula............. 51 . Fruiting branch of Opuntia arbuscula... 51 . Flowering branch of Opuntia thurberi... 53 . Branch of Opuntia davisii.............. 55 . Branch of Opuntia viridiflora........... 55 . Branch of Opuntia whipplei............ 55 . Plant of Opuntia acanthocarpa......... 56 . Joint of Opuntia serpentina............ 58 . Plant of Opuntia bigelovil.............. 59 . Joint of Opuntia bigelovii.............. 59 . Potted plant of Opuntia ciribe.......... 60 . Joint of Opuntia ciribe..............0.. 60 . Potted plant of Opuntia cholla.......... 61 . Joint of Opuntia cholla................ 62 . Proliferous fruits of Opuntia cholla, devel- oping new joints..................... 62 . Proliferous fruits of Opuntia cholla, developing new joints................ 62 . Joints of Opuntia loydii................ 63 . Plant of Opuntia lloydii............... 63 . Plant of Opuntia imbricata............. 64 . Potted plant of Opuntia tunicata....... 65 . Plant of Opuntia pallida............... 65 . Potted plant of Opuntia molesta........ 67 . Joint of Opuntia prolifera.............. 69 . Potted plant of Opuntia prolifera....... 70 . Potted plant of Opuntia alcahes........ 70 . Joint of Opuntia verschaffeltii.......... 72 . Grafted plants of Opuntia clavarioides... 73 . Potted plant of Opuntia salmiana....... 74 . Joints of Opuntia salmiana............. 74 . Potted plant of Opuntia subulata....... 76 . Joint of Opuntia pachypus............. 77 . Joints of Opuntia schottii.............,. 81 . Joints of Opuntia clavata.............. 81 . Joints of Opuntia parishii.............. 82 . Joints of Opuntia pulchella............. 82 . Plants of Opuntia vilis...........0.0... 83 . Joints and cluster of spines of Opuntia bulbispina............0........0.00. 83 . Joints of Opuntia grahamii............. 84 . Plants of Opuntia weberi.............. 84 . Joints of Opuntia weberi............... 85 . Potted plant of Opuntia floccosa....... 86 . Mound of Opuntia lagopus............. 88 . Root, joints, and flower of O. australis. . 88 . Joints of Opuntia glomerata............ 89 . Joint of Opuntia aoracantha............ gt . Joint of Opuntia rauppiana............. 92 . Flowering plant and fruit of Opuntia sub- terranea....... ee eee 2 Joints of Opuntia hickenii.............. 92 Joint of Opuntia darwinii.............. 94 Joints of Opuntia atacamensis.......... 04 . Joints of Opuntia russellii. 94 . Joints of Opuntia ovata. .. Leas 95 . Potted plant of Opuntia sphacrica. Lees 96 . Joint of Opuntia skottsbergii..........., 97 . Joint of Opuntia nigrispina............. 97 . Joint of Opuntia pentlandii............. 97

VI

Fic. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139. 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. I5I. 152. 153. 154. 155. 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. 171. 172. 173. 174. 175. 176. 177. 178. 179. 180. 181. 182. 183. 184.

THE CACTACEAE.

TEXT-FIGURES—continued.

PAGE. Joints of Opuntia pentlandii............ 98 Joint of Opuntia ignescens............. 98 Mound of Opuntia ignescens........... 98 Plant of Opuntia campestris............ 99 Joints of Opuntia ignota............... 99 Thicket of Opuntia pumila............. 100 Joints of Opuntia pumila............... 101 Joints of Opuntia pubescens............ 101 Joints of Opuntia curassavica........... 102 Joints of Opuntia borinquensis.......... 104 Joints of Opuntia militaris............. 104 Joints and flower of Opuntia tracyi..... 105 Joints and flowers of Opuntia pusilla.... 106 Joints of Opuntia aurantiaca........... 107 Potted plant of Opuntia schickendantzii. 107 Plant of Opuntia kiska-loro............ 108 Joints of Opuntia canina............... 108 Plant of Opuntia retrorsa.............. 109 Plant of Opuntia utkilio............... 110 Joints of Opuntia anacantha............ 110 Thicket of Opuntia bella............... II Joints of Opuntia bella................ 112 Joint of Opuntia bella... ..0......0.000. 112 Plant of Opuntia triacantha............ 113 Plant of Opuntia tuna................. 114 Joints of Opuntia tuna................. 114 Thicket of Opuntia antillana........... 115 Joints of Opuntia antillana............. 115 Plant of Opuntia decumbens........... 117 Plant of Opuntia depressa.............. 118 Joints of Opuntia lubrica............... 119 Landscape showing Opuntia treleasei.... 119 Joints of Opuntia basilaris............. 120 Plant of Opuntia microdasys........... 121 Potted plant of Opuntia, probable hybrid 121 Joint of Opuntia macrocalyx........... 122 Plant of Opuntia rufida................ 122 Plant of Opuntia pycnantha............ 123 Potted plant of Opuntia comonduensis.. 124 Plant of Opuntia inamoena............. 125 Joint of Opuntia inamoena............. 125 Joints of Opuntia allairei............... 126 Joints of Opuntia pollardii............. 126 Plant of Opuntia opuntia.............. 128 Fruit of Opuntia grandiflora............ 129 Flowering joint of Opuntia grandiflora. . 129 Flowering joints of Opuntia austrina.... 1 30 Joints, flower, and fruit of O. plumbea. . 131 Fruit of Opuntia stenochila............. 132 Fruit of Opuntia stenochila............. 132 Joint of Opuntia stenochila............. 132 Potted plant of Opuntia delicata........ 133 Joint of Opuntia soehrensii............. 135 Joint of Opuntia microdisca............ 135 Joints of Opuntia strigil................ 136 Joints of Opuntia ballii................ 137 Joints of Opuntia pottsii............... 138 Joint of Opuntia setispina.............. 138 Plant and fruit of Opuntia mackensensii 139 Joint of Opuntia macrocentra.......... 140 Joint of Opuntia tardospina............, 141 Cluster of spines of Opuntia gosseliniana 141 Joint of Opuntia gosseliniana..... 141 Joint of Opuntia angustata............. 142 Plant of Opuntia azurea............... 143 Joints of Opuntia azurea............._. 143 Joint of Opuntia covillei..........0.... 145 Joint of Opuntia covillei............ 2. 145

Fic. 185

186. 187. 188. 189. 190. IgI.

192. 193. 194. 195. 196. 197. 198. 199. 200. 201. 202. 203. 204. 205.

206. 207. 208. 209. 210. 211. 212. 213. 214. 215. 216. 217. 218. 219. 220. 221. 222. 223. 224. 225. 226. 227. 228. 229. 230. 231. 232. 233. 234. 235. 236. 237. 238. 239. 240. 241. 242. 243. 244. 245. 246. 247. 248. 249.

PAGE. Joint of Opuntia vaseyi................ 146 Potted plant of Opuntia occidentalis.... 147 Joint of Opuntia brunnescens........... 150 Fruit of Opuntia brunnescens........... 150 Joint of Opuntia galapageia............ 150 Flower of Opuntia galapageia........... 150 Joint and cluster of spines of Opuntia galapageia....... cc eee ee ce eee 151 Flowering joint of Opuntia delaetiana.. 152 Joints of Opuntia hanburyana.......... 154 Joint of Opuntia quitensis............. 154 Joint of Opuntia distans............... 155 Joint of Opuntia elata................. 157 Joints of Opuntia cardiosperma......... 157 Joint of Opuntia scheeri............... 159 Plant of Opuntia chlorotica............ 160 Joints of Opuntia chlorotica............ 160 Plant of Opuntia dillenii............... 162 Joint of Opuntia tapona............... 164 Potted plant of Opuntia littoralis....... 164 Joints of Opuntia cantabrigiensis........ 167

Part of joint and cluster of spines of

Opuntia procumbens................. 167 Joint of Opuntia cafiada............... 167 Joint of Opuntia pyriformis............ 168 Joint of Opuntia durangensis........... 169 Plant of Opuntia macdougaliana........ 170 Potted plant of Opuntia macdougaliana. 171 Joint of Opuntia wilcoxii............... 172 Plant of Opuntia tomentosa............ 173 Joint of Opuntia tomentella............ 174 Potted plant of Opuntia leucotricha..... 175 Joints of Opuntia orbiculata............ 176 Potted plant of Opuntia pilifera......... 177 Plants of Opuntia ficus-indica.......... 178 Fruit of Opuntia ficus-indica........... 178 Plant of Opuntia crassa................ 179 Potted plant of Opuntia maxima........ 180 Joint of Opuntia spinulifera............ 182 Joint of Opuntia lasiacantha............ 183 Joint of Opuntia zacuapanensis......... 183 Joint of Opuntia hyptiacantha.......... 183 Joint of Opuntia streptacantha......... 184 Potted plant of Opuntia megacantha.... 185 Plants of Opuntia megacantha.......... 186 Joint of Opuntia megacantha........... 186 Joint of Opuntia deamii................ 187 Joint of Opuntia eichlamii.............. 188 Joint of Opuntia inaequilateralis........ 188 Joint of Opuntia pittieri............... 189 Joint of Opuntia cordobensis........... 189 Fruit of Opuntia cordobensis........... 189 Joint of Opuntia quimilo............... 190 Fruit of Opuntia quimilo............... 190 Joint and flowers of Opuntia quimilo.... 191 Plant of Opuntia robusta.............. 192 Plant of Opuntia fragilis............... 194 Joints of Opuntia arenaria............. 195 Joint of Opuntia trichophora........... 195 Plant of Opuntia erinacea...... 196 Joint of Opuntia juniperina............ 197 Seed of Opuntia juniperina............. 197 Joint of Opuntia hystricina............. 197 Joint of Opuntia sphaerocarpa.......... 198 Joints of Opuntia polyacantha.......... 199 Joint of Opuntia stenopetala........... 200

Upper part of joint and flower of Opuntia stenopetala

i 2?

FIG. 250. 251. 252. 253. 254. 255- 256. 257. 258. 259. 260. 261. 262. 263. 264. 265. 266. 267. 268. 269. 270. 271. 272. 273. 274. 275. 276.

THE CACTACEAE.

TEX T-FIGURES—continued.

Plants of Opuntia palmadora.......... Joints of Opuntia palmadora........... Plants of Opuntia nashii............... Potted plant of Opuntia nashii.......... Joint of Opuntia bahamana............. Flower of Opuntia bahamana........... Plants of Opuntia macracantha......... Potted plant of Opuntia macracantha.. . Potted plant of Opuntia spinosissima.... Plants of Opuntia millspaughii.......... Plant of Opuntia moniliformis.......... Plant of Opuntia moniliformis.......... Plant of Opuntia moniliformis.......... Plants of Opuntia rubescens............ Plants of Opuntia rubescens............ Proliferous fruits of Opuntia rubescens. . Joint of Opuntia rubescens.............. Fruit of Opuntia brasiliensis............ Plant of Opuntia brasiliensis............ Branch of Opuntia bahiensis............ Joint and fruit of Opuntia bahiensis..... Plant of Opuntia bahiensis............. Plant of Opuntia ammophila........... Fruiting joint of Opuntia ammophila.... Flower of Opuntia argentina........... Potted plant of Opuntia chaffeyi........ Plant of Opuntia chaffeyi...............

PAGE.

F1G.277.

278. 279. 280. 281. 282. 283. 284. 285. 286. 287. 288. 289. 290. 291. 292. 293. 294. 295. 296. 297. 298. 299. 300. 301. 302.

Small joint of Nopalea gaumeri......... Elongated joint of Nopalea gaumeri.... Plant of Opuntia depauperata.......... Joint of Opuntia depauperata.......... Plant of Opuntia pestifer.............. Plant of Opuntia discolor.............. Joints of Opuntia pestifer.............. Joint of Opuntia discolor............... Joint of Opuntia guatemalensis......... Joint of Opuntia pennellii.............. Joints of Opuntia caracasana........... Plant of Opuntia aequatorialis.......... Joints of Opuntia aequatorialis......... Joints of Opuntia lata..........0.00.... Fruits of Opuntia lata.............00... Joint with flower of Opuntia macateci... Joint of Opuntia macateei.............. Plants of Opuntia soederstromiana...... Plants of Opuntia zebrina.............. Fruit of Opuntia zebrina............... Plants of Opuntia keyensis............. Section of flower of Opuntia keyensis.... Flower of Opuntia keyensis............. Joint of Opuntia bonplandii............ Plant of Opuntia dobbieana............ Plant of Opuntia dobbieana (without

legend).

a ey

VII

THE CACTACEAE

Descriptions and Illustrations of Plants of the Cactus Family

THE CACTACEAE.

INTRODUCTION.

The writers began field, greenhouse, and herbarium studies of the Cactaceae in 1904 and in the years following they made studies and collections over wide areas in the United States, Mexico, and the West Indies. It was first intended that these should be followed by a general description of the North American species only, but a plan for a more complete investigation of the family was proposed by Dr. D. ‘T. MacDougal in January 1911. This was approved by the trustees of the Carnegie Institution of Washington at its next regular meeting and a grant was made to cover the expenses of such an investigation. Dr. Rose was given temporary leave of absence from his position as Associate Curator in charge of the Division of Plants, United States National Museum, and became a Research Associate in the Carnegie Institution of Washington, with William R. Fitch and Paul G. Russell as assistants; Dr. Britton, Director-in-Chief of the New York Botanical Garden, was appointed an honorary Research Associate, while R. S. Williams, of the New York Botanical Garden, was detailed to select and preserve the specimens for illustration. Work under this new arrangement was begun January 15, 1912, and thus several lines of investigation were undertaken in a comprehensive way.

1. Reexamination of type specimens and of all original descriptions: This was necessary because descriptions had been incorrectly interpreted, plants had been wrongly identified, and the errors perpetuated; thus the published geographical distribution of many species was faulty and conclusions based on such data were unreliable. Not only had specific names been transferred to plants to which they did not belong, but generic names were interchanged and the laws of priority ignored. Many valid species, too, had dropped out of collections and out of current literature and had to be restored.

2. Assembling of large collections for greenhouse and herbarium use: Extensive greenhouse facilities were furnished by the New York Botanical Garden and the United States Department of Agriculture, while the herbaria and libraries of the United States National Museum and of the New York Botanical Garden furnished the bases for the researches. The New York Botanical Garden has also cooperated in contributing funds in aid of the field operations, in clerical work, and a large number of the illustrations used have been made there, the paintings and line drawings mostly by Miss Mary E. Eaton.

3. Extensive field operations in the arid parts of both Americas: Many of these deserts are almost inaccessible, while the plants are bulky and if not handled care- fully are easily destroyed. Many plants require several years to mature, in some cases many years to flower in cultivation. Through these explorations were obtained the living material for the greenhouse collections and for exchange purposes, as well as herbarium material for permanent preservation. Of much importance, also, were field observations upon the plants as individuals, their form, habit, habitat, and their relations to other species.

3

4 THE CACTACEAE.

Early in 1912 Dr. Rose went to Europe to study the collections there and to arrange for exchanges with various botanical institutions having collections of these plants. He spent considerable time at London, mainly at the Royal Botanic Gardens, Kew, where through the courtesy of the Director, Sir David Prain, he was able to examine the greenhouse, illustrative, and herbarium material for which this institution has long been famed. ‘The collection at the British Museum of Natural History and that of the Linnaean Society of London were examined. At Paris he studied the collections at the Natural History Museum, many of which have historic interest; one of his interesting discoveries there was that the Pereskia bleo, collected by Baron Friedrich Alexander von Humboldt in Colombia, is a very different species from the plant which for nearly a century has been passing in our collections and literature under that name. He also visited the famous botanical garden of the late Sir Thomas Hanbury, at La Mortola, Italy, and through the courtesy of Lady Hanbury was given every possible facility for the study of this collection; Mr. Alwin Berger, who was then curator in charge, had brought together one of the most extensive representations of this family to be found growing in the open in any place in the world. Here in the delightful climate of the Riviera were grown many species which were apparently just as much at home as they would have been in their desert habitats. Dr. Rose also visited Rome, Naples, Venice, and Florence, where he saw smaller collections in parks and private gardens. At Munich he examined certain types in the Royal Botanical Museum, then under the charge of Dr. L. Radlkofer, and saw some interesting species in the Royal Botanical Garden then being organized by Dr. K. Goebel. At Berlin he examined the herbarium and living specimens in the Berlin Botanical Garden, through the courtesy of Dr. A. Engler, and the West Indian collection through the courtesy of Dr. I. Urban. He then went to Halle and saw L. Quehl’s collection of mammillarias; to Erfurt, where he saw the Haage and Schmidt, and Haage Jr. collections; to Darmstadt to see the Botanical Garden under Dr. J. A. Purpus; and to Antwerp to see DeLaet’s private collection.

In 1913 Dr. Britton and Dr. Rose visited the West Indies. Dr. Britton, who was accompanied by Mrs. E. G. Britton, Miss D. W. Marble, and Dr. J. A. Shafer, collected on St. Thomas and the other Virgin Islands, Porto Rico, and Curacao. At the latter island he rediscovered the very rare Cactus mammillaris, which had not been im cuyation cr many years. Dr. Rose, who was accompanied by William Coie St Chien , ; Russell, also stopped at St. Thomas, and collected on St.

ot. pher, Antigua, and Santo Domingo.

John F Can, ana oe eritton again visited Porto Rico and, assisted by Mr. vnct and the sat ee ewardson Brown, explored the entire southwestern arid s Desecheo, Mona, and Muertos.

In 1914 Dr. Rose went to the west coast of at Jamaica and Panama. Peru, central Bolivi

South America, making short stops a made extensive collections in central and southern a, and northern and central Chile. At Santiaso. Chil

, ba qe *) e : 5 , Cs he examined a number of Philippi’s types in the National Museum and obtained

some rare specimens from the B i Sohrens otanical Garden through the courtesy of Johannes

INTRODUCTION. 5

In 1915 Dr. Rose, accompanied by Paul G. Russell, visited Brazil and Argentina on the east coast of South America, collecting extensively in the semiarid parts of Bahia, Brazil, and in the region about Rio de Janeiro, so rich in epiphytic cacti. In the deserts about Mendoza and Cérdoba, in Argentina, collections were also made. Here he also arranged for exchanges with the leading botanists and collec- tors. The following persons have made valuable contributions from the regions visited: Dr. Leo Zehntner, Joazeiro, Brazil; Dr. Alberto Lofgren, Rio de Janeiro, Brazil; Dr. Carlos Spegazzini, La Plata, Argentina; Dr. Cristébal M. Hicken, Buenos Aires, Argentina; and Dr. Carlos S. Reed, Mendoza, Argentina.

In October and November 1916, Dr. Rose, accompanied by Mrs. Rose, visited Curacao and Venezuela, studying especially the cactus deserts about La Guaira and Puerto Cabello. A number of photographs were taken by Mrs. Rose.

While en route for Venezuela, arrangements were made with Mr. Harold G. Foss to make a collection of cacti at Coro, Venezuela. Among the specimens obtained were species not found farther east in Venezuela, so far as known.

In 1916 Dr. Britton, assisted by Mr. Percy Wilson, studied the cacti of Havana and Matanzas Provinces and those of the Isle of Pines, Cuba.

In 1918 Dr. Rose, assisted by George Rose, visited Ecuador on behalf of the United States Department of Agriculture, aided by the Gray Herbarium of Har- vard University and the New York Botanical Garden; about thirty rare or little- known species were obtained.

Through the expenditure of about $2,400, contributed by Dr. Britton, a very important collection of cacti was made by Dr. J. A. Shafer during a six months’ exploration from November 1916 to April 1917 of the desert regions of northwestern Argentina, southeastern Bolivia, northeastern Argentina, and adjacent Uruguay and Paraguay. Fortunately, for the purposes of this work, this collection was brought back to New York by Dr. Shafer in time for the information yielded by it and by his field observations to be used in the manuscript. It has given us first-hand information concerning over 120 species of cacti as to which we have previously known little.

There are still a few cactus regions which ought to be explored, but the following summary will show the wide field from which we have obtained information.

Our field investigations have covered practically all the cactus deserts of Mexico. The most important of these are the vicinities of Tehuacan and Tomellin, the plains of San Luis Potosi, the chalky hills surrounding Ixmiquilpan, the lava fields in the Valley of Mexico and above Cuernavaca, the deserts of Querétaro, the west coast of Mexico extending from the United States border to Acaponeta, and the seacoasts and islands of Lower California. Other regions in Mexico containing cacti, but not in such great abundance as the foregoing, are those about Pachuca, Oaxaca City, Mitla, Jalapa, Iguala, Chihuahua City, and Guadalajara. All the work in Mexico, however, was done prior to 1912, for, owing to political disturb- ances, no field work there has been feasible since that time.

In the United States our work has extended over the cactus regions of Florida, Texas, New Mexico, Arizona, southern California, western Kansas, and south- eastern Colorado.

6 THE CACTACEAE.

In the West Indies we have explored all of the Greater Antilles, the Bahamas, the Virgin Islands, St. Christopher, Antigua, Barbados, and Curacao.

In South America our field study included the most important deserts of Peru, Bolivia, and Chile, and parts of Brazil, Venezuela, Ecuador, and Argentina. The cactus deserts of South America are so extensive and so remote from one another that it was possible to visit only a part of them in the four seasons allowed for their exploration.

Among many enthusiastic volunteers whose contributions of specimens and data have greatly supplemented our own collections and field studies, the following deserve especial mention:

Mr. Henry Pittier has made valuable sendings from Colombia, Venezuela, Panama, Costa Rica, and Mexico; Mr. O. F. Cook, from Guatemala and Peru; Mr. G. N. Collins, the late Federico Eichlam, Mr. R. H. Peters, Mr. C. C. Deam, Mrs. T. D. A. Cockerell, Baron H. von Tiirckheim, and the late Professor W. A. Kellerman have sent important collections from Guatemala; Mr. A. Tonduz, Mr. Otén Jiménez, Dr. A. Alfaro, Mr. C. Wercklé, and Mr. Alfred Brade, local collectors and naturalists in Costa Rica, have sent much good material from their country; Mr. William R. Maxon has sent new and rare material from Costa Rica, Guatemala, and Cuba; Professor C. Conzatti and his son, Professor Hugo Conzatti, Dr. C. A. Purpus, Dr. Elswood Chaffey, Mrs. Irene Vera, M. Albert de Lautreppe, and the late Mr. E. A. H. Tays have sent us many interesting specimens from Mexico; Mr. W.E. Safford made a valuable collection in Mexico in 1907; E. W. Nelson and E. A. Goldman, who have collected so extensively in Mexico and the Southwest, have obtained many herbarium and living specimens for our use; Mrs. Gaillard, who lived at Panama several years while the late Colonel D. D. Gaillard was a member of the Isthmian Canal Commission, collected interesting cacti, including Epiphyllum gaillardae; the late Dr. H. EK. Hasse sent specimens from southern California and Arizona; C. R. Orcutt, the well-known cactus fancier, has aided us In many ways besides sending us specimens from his collections; Dr. R. E. Kunze has frequently sent specimens, especially from Arizona: General T imothy FE. Wilcox, for whom Wilcoxia was named, has sent us specimens from the Southwest, while his son, Dr. G. B. Wilcox, contributed several sendings from the west coast of Mexico and Guatemala; Dr. D. T. MacDougal has sent many specimens from all over the Southwest, especially from Mexico, Arizona, and southern California; ne nas Mas contin se remote deserts, which have yielded interesting

) ; y excellent photographs, quite a number of which are reproduced in this report (Plate 1, etc.). Professor F. E. Lloyd, while located nd fa and neta ses Nex, mace saree collections of living, herbarium, photographs. Dr. Forrest Shreve has ‘sent vain acid alk Sketches, and Arizona and Mr. W. H. Long from New M ico: My am Beals yom none Schaller have furnished interesting speci pre ene E euler enaes be fornia species; Professor J.J Thoraber has my fe ‘laabe tain “ti Souter ne and notes from Arirong : a 7 a as made valuable contributions of material

; - M. E. Jones, Mr. I. Tidestrom, Mr. Thomas H.

INTRODUCTION. 7

Kearney, and Professor A. O. Garrett have all sent specimens from Utah; Profes- sor T. D. A. Cockerell and Mr. Merritt Cary have sent specimens from Colorado; Dr. P. A. Rydberg has brought many specimens from the Rocky Mountain region; Messrs. Paul C. Standley, EF. O. Wooton, Vernon Bailey, and H. L. Shantz have sent specimens from the southwestern United States; Brother Leén, of the Colegio de la Salle, Havana, and Dr. Juan T. Roig, of the Estacién Agrondémica, Santiago de las Vegas, Cuba, have contributed Cuban specimens, and Dr. J. A. Shafer has collected widely in Cuba; Mr. William Harris, of Hope Gardens, Jamaica, has col- lected for us in Jamaica; Dr. John K. Small has obtained collections from nearly all over the southeastern United States, aided by Mr. Charles Deering. Dr. Henry H. Rusby and Dr. Francis W. Pennell have contributed plants and specimens from Colombia, collected in 1917 and 1918. Mr. Frederick V. Coville, of the United States Department of Agriculture, has made many valuable suggestions during the progress of the investigation.

In our studies we have also had use of the cacti of the following American collections: Herbarium of the Missouri Botanical Garden at St. Louis; the Gray Herbarium of Harvard University; the Rocky Mountain Herbarium at Laramie, Wyoming; the collection of the United States Department of Agriculture; the herbarium of the University of California, especially the Brandegee collection; and the herbarium of the Field Museum of Natural History.

The types of the new species described in this work are deposited in the herbaria of the New York Botanical Garden and the United States National Museum, unless otherwise indicated.

In greenhouse collections many kinds of cacti grow very slowly, and flower only after many years’ cultivation. We have a number of plants of this kind from various parts of America. It is hoped that some of them may bloom during the period of publication of this book and thus enable us to include them in an appendix.

Fic. 1.—Pereskia pereskia. Grown as a hedge.

8 THE CACTACEAE.

Order CACTALES.

Perennial, succulent plants, various in habit, mostly very spiny, characterized by specialized organs termed areoles. Leaves usually none, except in Pereskia and Pereskiopsis, where they are large and flat but fleshy, and in Opuntia and its relatives, where they are usually much reduced and mostly caducous, terete, or subulate. Spines very various in size, form, arrangement, and color, sometimes with definite sheaths. The areoles are peculiar and complex organs, situated in the axils of leaves when leaves are present, and bearing the branches, flowers, spines, glochids, hairs, or glands; in some genera two kinds of areoles occur, either distinct or united by a groove. Flowers usually perfect, either regular or irregular, usually solitary but sometimes clustered, sometimes borne in a specialized terminal dense inflorescence called a cephalium; perianth-tube none, or large and long, the limb spreading or erect, short or elongated, the lobes few or numerous, often inter- grading in shape and color, but sometimes sharply differentiated into sepals and petals; stamens commonly numerous, elongated or short, sometimes clustered in series, the filaments usually borne on the throat of the perianth, the small oblong anthers 2-celled; style one, terminal, short or elongated; stigma-lobes 2 to many, usually slender; ovary 1-celled, distinct, or immersed in a branch or forming a part of a branch; ovules numerous. Fruit a berry, often juicy and sometimes edible, sometimes dry, in one species described as capsular and dehiscing by an operculum, in others opening by a basal pore. Seeds various; cotyledons two, accumbent, sometimes minute knobs, often broad or elongated; endosperm little or copious; radicle terete.

The order consists of the following family only:

Family CACTACEAE Lindley, Nat. Syst. ed. 2. 53. 1836.

Characters of the order as given above. The family is composed of three tribes.

KEY To TRIBES.

Leaves broad, flat; glochids wanting; flowers stalked (sometimes short-stalked), often clustered Leaves (except in Pereskiopsis) terete

flowers sessile. Areoles with glochids (except in Mathuenia); vegetative parts bearin small and fugacious; flowers rotate (petals erect in Nopalea) Areoles without glochids; usually no leaves on the vegetative par with definite tubes (except Rhipsalis)

g leaves, which are usually

occ c ec eee eee cece eee eee 2. Opuntieae ts (except cotyledonary); flowers

eee eee eee cece e teen een eeeeee 3. Cereeae

Tribe 1. PERESKIEAE.

Stems and foliage as in other dicotyledonous plants; flowers more or less stalked, their parts all distinct; gloc testa of seed thin, brittle.

inflorescence in some species compound; hids wanting; ovule with short funicle;

The genus Pereskia, the only representative of this tribe, is, on account of its similarity to other woody flowering plants, considered the nearest cactus relative to the other families, but this relationship is in all cases remote.

The nearest generic relatives of Pereskia in the cactus family are doubtless the following:

Pereskiopsis, some of whose species were first assigned to the genus Pereskia, but they have different foliage and the areoles often bear glochids.

Opuntia, whose species have leaves, though much reduced and usually caducous, other- wise very different ; but some of the species of Opuntia were first referred to Pereskia.

Mathuenta (two of whose species have only recently been taken out of Pereskia), whose seeds are similar but the areoles lack glochids, otherwise very different.

This tribe has a wide geographic distribution, but is found wild only in the tropics.

[7)-

¢ ils of the leaves, neither sheathed nor barbed: glochids (found only in the Opuntieae) wanting; leaves alternate, broad, flat, deciduous, or somewhat fleshy ; wheel-shaped; stamens numerous: shell, the embryo strongly curved ;

Lees 1. Pereskieae

PERESKIA. 9

Type species: Cactus pereskia Linnaeus. In 1808 about 25 names had been proposed in Pereskia, but, in his monograph published

that year, Karl Schumann accepted only 11 species. Several new ones have been proposed

since the publication of Schumann’s monograph.

The species are native in Mexico, the West Indies, Central America, and South America. Some of the species are much used as stocks for growing the various forms of Zygocactus, Epiphyllum, and other cacti requiring this treatment; P. pereskia is most used and P. grandifolia next. Several species are widely cultivated as ornamentals in tropical regions; they do not flower freely under glass in northern latitudes. All species studied by us in the living state grow readily from cuttings.

The typical species seems to have been first introduced into Europe from the West Indies in the latter part of the sixteenth century. A straight-spined species was first described and figured by L. Plukenet in 1696, who called it a portulaca, and the next year by Commerson as an apple (Malus). In 1703 C. Plumier described the genus Pereskzia, basing it upon a single species. The genus was repeatedly recognized by Linnaeus in his earlier publications, and by some pre-Linnaean botanists, but in 1753 Linnaeus merged it into Cactus along with a number of other old and well-established genera; but it was retained by Philip Miller in 1754 in the fourth edition (abridged) of his Gardeners’ Dic- tionary and has since been generally recognized as a genus by botanical and horticultural authors.

The name is variously spelled Pezrescia, Peireskia, Perescia, and Pereskia.

Named for Nicolas Claude Fabry de Peirese (1580-1637).

KEy TO SPECIES.

Climbing vines, the twigs with a short pair of reflexed spines from each areole, the stem with

acicular spines (Series 1. Typicae)........-5 6.6.0 e eee eee 1. P. pereskia Shrub or trees with slender straight spines (Series 2. Grandifoliae). Petals toothed or fimbriate. Petals somewhat toothed..........060 cece eee ett ene eee e eee 2. P. autumnalis Petals fimbriate. Species from Mexico; ovary turbinate.. 0... ett ee eee 3. P.lvchnidiflora Species from Costa Rica; ovary pyriform. 2.66 eee 4. P. nicoyana Petals entire, at least