Cassiinae pt 1 NY-Botanical_gardens_Vol. 35_1 - Copy.pdf - Antbase

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8 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. that of some sennas; and in several members of Senna ser. Coluteoideae the pod is at once indehiscent and multUocular Uke that of Cassia. The long woody pipe- Uke pod of Senna spectabilis superficially resembles that of some true cassias, but when ripe it sphts open along the ventral suture and it arises from a flower whoUy unhke that of any Cassia in its androecium and asymmetric corolla, fea tures that remain in close harmony with related sennas characterized by piano- compressed pods. Cassia lacks petiolar glands, but whUe the gland characteristic of the majority of sennas is absent in some series of the genus it is infaUibly present wherever the pod approaches that of Cassia in form or other attributes. An internaUy pulpy pod, common but not ubiquitous in Cassia, occurs also in some sennas (e.g. S. pendula, S. bacillaris); it is a speciaUzation doubtless related to seed dispersal which we suppose to have developed independently in the two genera, as it has in some neotropical acacias. By itself, therefore, the pod does not provide a sure means of separaring Cassia from Senna, although the few exceptions are not genuine obstacles but merely distracrions due to convergent evolution. The fallibility of the pod deserves emphasis, nonetheless, because of its influence on the historical taxonomy of Cassunae. Cassia and Senna became first known to European botanists through the pods of C. fistula and of S. italica and S. alexandrina, articles of herbal medicine imported from the Orient by way of Egypt and Asia Minor. The fistular woody muUilocular pulp-laden pod of Cassia appeared (and really is) fundamentally distinct from the bifacial papery umlocular dry pod of Senna, and it was in these terms that Tournefort (Inst. Rei Herb. 618, t. 390, 392. 1700) defined the genera. And it was to Tournefort's durably influential work that the origin of confusion between Cassia and Senna can be traced. While Tournefort illustrated Cassia by pods of C. fistula and C. grandis and by a flower probably of the latter, he also listed as species of Cassia seven American sennas, mostly known to him only from antecedent accounts by Piso and Plumier. All of these were deliberately absorbed by Linnaeus (1753, 1754, U. cc), along with several sennas discovered after 1700 and the five then-known species of Chamaecrista, into the heterogeneous genus that survived into modern times. In this historically momentous decision Linnaeus adapted, in the high-handed manner that shocked many contemporary botanists, a famihar name to an unfamihar usage. For the genus Cassia described in the fifth edition of Genera Plantarum is no longer Cassia of the ancients but explicitly Senna. The carefully described androecium, which is pro foundly at odds with that of the genuine cassias or of the chamaecristas attributed to Cassia in Species Plantarum, is that of Senna as observed alive in the conservatories at Hartekamp and Upsala; for the androecium of Cassia sens, str., a genus of tropical trees that had not yet been brought to flower in cultivation, was evidently unknown to him. The world monographs of Cassia by DeCandolle in CoUadon (1816), Vogel (1837, incorporating a wealth of BrazUian sennas discovered by Sellow) and Bentham (1871), while attaining progressively sophisti cated understanding of relationships between natural groups of Cassiinae, reaffirmed the generic concept of Linnaeus, and stifled the voices of protest. In fact the intratribal generic hmits, as we understand them today, had been worked out intuitively as early as 1739 by Jacob Breyne (Prod. fasc. rar. pl. secund. 50), who first described the genera Chamaecrista and Chamaecassia, the latter equivalent to Senna P. MiU. Later on, piecemeal partitions of Linnaean Cassia were attempted by Persoon and WiUdenow, who abstracted from it the true cassias as Cathartocarpus (1805) and Bactyrilobium (1809) respectively, and also by Link (1831) and Roxburgh (1832), who contradictorily expelled the sennas and chamaecristas and reserved a purified Cassia for the immediate kindred of
1982] CASSIINAE—CASSIA C. fistula. When Britton and colleagues, in 1930-1936, revived Cassia sensu Link and Roxburgh, its just claim to generic status was devaluated by association with the false claims of a horde of Ul-considered segregates from Senna. Nevertheless, Bentham's often quoted remark (1871, p. 503) to the effect that "no tolerably fair botanist" had ever mistaken a cassia for anything else is true only provided that the concept of a subtribe Cassunae be substituted for a comprehensive genus Cassia. In his work on Caesalpiniaceae and Mimosaceae the generic criteria empha sized by Britton were primarily carpological and his taxonomy became as a con sequence highly formahzed, like that of Rydberg in the papihonate legumes. WhUe the elastically dehiscent pod of Chamaecrista effectively and consistently, even by itself, separates that great genus from Cassia and Senna collectively, there is no one single feature of the pods of these two genera that will consistently separate one from the other. The terete indehiscent pulpy pod of Senna bicapsularis is morphologically very close to that of some genuine Cassiae, even in its biseriate seeds, and the chain of character states that has been found to lead from an ideahzed piano-compressed dehiscent pod in Senna to a terete indehiscent one in Cassia sens, restr. has made the case for the latter fatally vulnerable and given decisive support to Bentham's inclusive concept. Our definition of Cassia is, however, no longer based'on particulars ofthe fruit, but on an infallible syndrome of characters expressed in form of inflorescence, bracteoles and floral morphology that rigorously defines a genus much further removed from any one set of Sennae than any one group of these is from another. In his masterly discussion of Cassia sect. Fistula, Bentham (1871, p. 507) found insuperable difficulty in arranging the species into subsidiary groups, owing to lack of correlation between spatial dispersal and phenetic characters, particularly size and number of leaflets, persistence of floral bracts, detaUs of staminal and carpological structure, and substance (pithy or pulpy) enveloping the seeds. With the advantages of more complete materials and knowledge of several species discovered since 1871, we find on the contrary that it is possible to distinguish between the capricious effects of parallel or convergent evolution and clues to presumed genetic relationships that are confirmed by patterns of distribution on a continental scale. We have accordingly arranged the American cassias and those exotic ones of which we have detaUed knowledge into named series, our purpose being to bring out relative degrees of relationship between groups and stress the isolation within the genus of several unique species. FoUowing a key to these series, which encapsulates our basic classification of Cassia, will be found (p. 10) a practical key to species native and commonly cultivated in the Americas which is adapted for use both with flowering and fruiting specimens. Key to Series of genus Cassia 1. Cavity of pod divided by transverse septa into one rank, or into 2 parallel ranks, of 1-seeded locules; seeds transverse and broadside to the septa, obcompressed, their hilum situated near base of one broad face. 2. Seeds stacked in one vertical column within the pod, the intervening septa traversing the whole cavity. 3. Both sutures of the pod thickened and externally prominent and each consisting of 2 ribs, the pod consequently bluntly compressed-quadrangular. Bracts persistent into anthesis; anthers of 4 fertile antepetalous stamens erect; spp. 3 of S. America (Amazon Basin and the Guianas to s.-e. Brazil). Petals yellow. ser. Tetrapleurae (p. 36). 3. Sutures of pod either thickened or not, if prominent the semiferous one 2-ribbed and the opposed one 1-ribbed. 4. Pod massively woody 3.5-5 cm diam, coarsely 3-ribbed by the sutures and the valves coarsely transverse-venose; anthers of 3 long abaxial stamens pilose and
Page 1: •'•mmmmmmfii^'''- '->mmmmmmmm^'
Page 4 and 5: ADVISORY BOARD MEMOIRS OF THE N E W
Page 6 and 7: This material is based upon researc
Page 8 and 9: IV FOREWORD In looking back over th
Page 11 and 12: DECEMBER 1982 MEMOIRS OF THE NEW YO
Page 13 and 14: 1982] CASSIINAE—INTRODUCTION 1. F
Page 15 and 16: 1982] CASSIINAE—CASSIA Fig. 1. An
Page 17: 1982] CASSIINAE—CASSIA commonly m
Page 21 and 22: 1982] CASSIINAE—CASSIA 11 sulcate
Page 23 and 24: 1982] CASSIINAE—CASSIA 13 i. ser.
Page 25 and 26: 1982] CASSIINAE—CASSIA 15 1.3-2.8
Page 27 and 28: 1982] CASSIINAE—CASSIA 17 modific
Page 29 and 30: 1982] CASSIINAE—CASSIA 19 and is
Page 31 and 32: 1982] CASSIINAE—CASSIA 21 Adult l
Page 33 and 34: 1982] CASSIINAE—CASSIA 23 Fig. 3.
Page 35 and 36: 1982] CASSIINAE—CASSIA 25 cm, the
Page 41 and 42: 1982] CASSIINAE—CASSIA 31 exactly
Page 43 and 44: 1982] CASSIINAE—CASSIA 33 bowl-sh
Page 45 and 46: 1982] CASSIINAE—CASSIA 35 s.-e. V
Page 49 and 50: 1982] CASSIINAE—CASSIA 39 like) a
Page 51 and 52: 1982] CASSIINAE—CASSIA 41 sometim
Page 53 and 54: 1982] CASSIINAE—CASSIA 43 sylvis:
Page 55 and 56: 1982] CASSIINAE—CASSIA 45 Lvs 13-
Page 59 and 60: 1982] CASSIINAE—CASSIA 49 f produ
Page 61 and 62: 1982] CASSIINAE—CASSIA 51 garden
Page 63 and 64: 1982] CASSIINAE—CASSIA 53 Klug, G
Page 65 and 66: 1982] CASSIINAE—CASSIA 55 Cassia
Page 67 and 68: 1982] CASSIINAE—CASSIA 57 Not see
Page 69 and 70:
1982] CASSIINAE—CASSIA 59 Cassia
Page 71 and 72:
1982] CASSIINAE—CASSIA 61 Opposit
Page 73 and 74:
1982] CASSIINAE—CASSIA 63 of leaf
Page 75 and 76:
1982] CASSIINAE—SENNA 65 neglecta
Page 77 and 78:
1982] CASSIINAE—SENNA 67 Pubescen
Page 79 and 80:
1982] CASSIINAE—SENNA 69 tionary
Page 81 and 82:
1982] CASSIINAE—SENNA 71 zygomorp
Page 83 and 84:
1982] CASSIINAE—SENNA 73 apical s
Page 85 and 86:
1982] CASSIINAE—SENNA 75 alternat
Page 87 and 88:
1982] CASSIINAE—SENNA 77 our clas
Page 89 and 90:
1982] CASSIINAE—SENNA 79 pressed
Page 91 and 92:
1982] CASSIINAE—SENNA 81 technica
Page 93 and 94:
1982] CASSIINAE—SENNA 83 parallel
Page 95 and 96:
1982] CASSIINAE—SENNA 85 18. Pod
Page 97 and 98:
1982] CASSIINAE—SENNA 87 3. Senna
Page 99 and 100:
1982] CASSIINAE—SENNA 89 tertiary
Page 101 and 102:
1982] CASSIINAE—SENNA 91 never so
Page 103 and 104:
1982] CASSIINAE—SENNA 93 3d. Senn
Page 105 and 106:
1982] CASSIINAE—SENNA 95 broad ma
Page 107 and 108:
1982] CASSIINAE—SENNA 97 with ran
Page 109 and 110:
1982] CASSIINAE—SENNA 99 ed. 2, 1
Page 111 and 112:
1982] CASSIINAE—SENNA 101 duction
Page 113 and 114:
1982] CASSIINAE—SENNA 103 broadsi
Page 115 and 116:
1982] CASSIINAE—SENNA 105 4. Blad
Page 117 and 118:
1982] CASSIINAE—SENNA 107 many-fl
Page 119 and 120:
1982] CASSIINAE—SENNA 109 neighbo
Page 121 and 122:
1982] CASSIINAE—SENNA 111 through
Page 123 and 124:
1982] CASSIINAE—SENNA 113 a multi
Page 125 and 126:
1982] CASSIINAE—SENNA 115 Vesture
Page 127 and 128:
1982] CASSIINAE—SENNA 117 those o
Page 129 and 130:
1982] CASSIINAE—SENNA 119 obhquel
Page 131 and 132:
1982] CASSIINAE—SENNA 121 Chamaef
Page 133 and 134:
1982] CASSIINAE—SENNA 123 3 abaxi
Page 135 and 136:
1982] CASSIINAE—SENNA 125 little
Page 137 and 138:
1982] CASSIINAE—SENNA 127 biseria
Page 139 and 140:
1982] CASSIINAE—SENNA 129 Trees o
Page 141 and 142:
1982] CASSIINAE—SENNA 131 Lf-rach
Page 143 and 144:
1982] CASSIINAE—SENNA 133 Strongl
Page 145 and 146:
1982] CASSIINAE—SENNA 135 interca
Page 147 and 148:
1982] CASSIINAE—SENNA 137 cium it
Page 149 and 150:
1982] CASSIINAE—SENNA 139 atively
Page 151 and 152:
1982] CASSIINAE—SENNA 141 pus, da
Page 153 and 154:
1982] CASSIINAE—SENNA 143 affinit
Page 155 and 156:
1982] CASSIINAE—SENNA 145 quangul
Page 157 and 158:
1982] CASSIINAE—SENNA 147 . . ."
Page 159 and 160:
1982] CASSIINAE—SENNA 149 pair of
Page 161 and 162:
1982] CASSIINAE—SENNA 151 Amply l
Page 163 and 164:
1982] CASSIINAE—SENNA 153 of the
Page 165 and 166:
1982] CASSIINAE—SENNA 155 Hithert
Page 167 and 168:
1982] CASSIINAE—SENNA 157 27. Sen
Page 169 and 170:
1982] CASSIINAE—SENNA 159 persal
Page 171 and 172:
1982] CASSIINAE—SENNA 161 Weak di
Page 173 and 174:
1982] CASSIINAE—SENNA 163 ovary d
Page 175 and 176:
1982] CASSIINAE—SENNA 165 Lvs 4-5
Page 177 and 178:
1982] CASSIINAE—SENNA 167 Pod pen
Page 179 and 180:
1982] CASSIINAE—SENNA 169 Fig. 16
Page 181 and 182:
1982] CASSIINAE—SENNA 171 coriace
Page 183 and 184:
1982] CASSIINAE—SENNA 173 petiole
Page 185 and 186:
1982] CASSIINAE—SENNA 175 ±3.5-8
Page 187 and 188:
1982] CASSIINAE—SENNA 177 a membe
Page 189 and 190:
Page 191 and 192:
1982] CASSIINAE—SENNA 181 hkewise
Page 193 and 194:
1982] CASSIINAE—SENNA 183 leaflet
Page 195 and 196:
1982] CASSIINAE—SENNA 185 present
Page 197 and 198:
1982] CASSIINAE—SENNA 187 42f. Se
Page 199 and 200:
1982] CASSIINAE—SENNA 189 Lvs bel
Page 201 and 202:
1982] CASSIINAE—SENNA 191 or acut
Page 203 and 204:
1982] CASSIINAE—SENNA 193 Stipule
Page 205 and 206:
1982] CASSIINAE—SENNA 195 leafles
Page 207 and 208:
1^^^] CASSIINAE—SENNA 197 In and
Page 209 and 210:
1982] CASSIINAE—SENNA 199 parenri
Page 211 and 212:
1982] CASSIINAE—SENNA 201 r^jCico
Page 213 and 214:
Page 215 and 216:
Page 217 and 218:
1982] CASSIINAE—SENNA 207 Chamaef
Page 219 and 220:
1982] CASSIINAE—SENNA 209 dackU i
Page 221 and 222:
Page 223 and 224:
Page 225 and 226:
1982] CASSIINAE—SENNA 215 Araguai
Page 227 and 228:
1982] CASSIINAE—SENNA 217 Racemes
Page 229 and 230:
1982] CASSIINAE—SENNA 219 broader
Page 231 and 232:
Page 233 and 234:
1982] CASSIINAE—SENNA 223 Pohl."
Page 235 and 236:
1^^2] CASSIINAE—SENNA 225 between
Page 237 and 238:
1982] CASSIINAE—SENNA 227 sometim
Page 239 and 240:
1982] CASSIINAE—SENNA 229 sometim
Page 241 and 242:
1982] CASSIINAE—SENNA 231 the hor
Page 243 and 244:
1982] CASSIINAE—SENNA 233 paratyp
Page 245 and 246:
Page 247 and 248:
1982] CASSIINAE—SENNA 237 tiary c
Page 249 and 250:
1982] CASSIINAE—SENNA 239 2.5-9(-
Page 251 and 252:
Page 253 and 254:
1982] CASSIINAE—SENNA 243 Cuba; d
Page 255 and 256:
1982] CASSIINAE—SENNA 245 the ste
Page 257 and 258:
1982] CASSIINAE—SENNA 247 in outh
Page 259 and 260:
1982] CASSIINAE—SENNA 249 "In Bra
Page 261 and 262:
1982] CASSIINAE—SENNA 251 the lon
Page 263 and 264:
1982] CASSIINAE—SENNA 253 fainriy
Page 265 and 266:
1982] CASSIINAE—SENNA 255 is not
Page 267 and 268:
1982] CASSIINAE—SENNA 257 70b. Se
Page 269 and 270:
1982] CASSIINAE—SENNA 259 racemes
Page 271 and 272:
1982] CASSUNAE—senna 261 other co
Page 273 and 274:
1982] CASSIINAE—SENNA 263 largely
Page 275 and 276:
Page 277 and 278:
1982] CASSIINAE—SENNA 267 sides t
Page 279 and 280:
1982] CASSIINAE—SENNA 269 found,
Page 281 and 282:
1982] CASSIINAE—SENNA 271 cordate
Page 283 and 284:
1982] CASSIINAE—SENNA 273 and hya
Page 285 and 286:
1982] CASSIINAE—SENNA 275 Key to
Page 287 and 288:
1982] CASSIINAE—SENNA 277 Peduncl
Page 289 and 290:
1982] CASSIINAE—SENNA 279 margine
Page 291 and 292:
1982] CASSIINAE—SENNA 281 Sandy b
Page 293 and 294:
1982] CASSIINAE—SENNA 283 cm, at
Page 295 and 296:
1982] CASSIINAE—SENNA 285 like th
Page 297 and 298:
Page 299 and 300:
1982] CASSIINAE—SENNA 289 not con
Page 301 and 302:
1982] CASSIINAE—SENNA 291 almost
Page 303 and 304:
1982] CASSIINAE—SENNA 293 filamen
Page 305 and 306:
1982] CASSIINAE—SENNA 295 2. Lfts
Page 307 and 308:
1982] CASSIINAE—SENNA 297 2. Fls
Page 309 and 310:
1982] CASSIINAE—SENNA 299 Main ca
Page 311 and 312:
Page 313 and 314:
1982] CASSIINAE—SENNA 303 a trifl
Page 315 and 316:
1982] CASSIINAE—SENNA 305 sentiaU
Page 317 and 318:
1982] CASSIINAE—SENNA 307 PUosulo
Page 319 and 320:
1982] CASSUNAE—senna 309 stamens,
Page 321 and 322:
1982] CASSIINAE—SENNA 311 of 3 ab
Page 323 and 324:
Page 325 and 326:
1982] CASSIINAE—SENNA 315 Pubesce
Page 327 and 328:
Page 329 and 330:
1982] CASSIINAE—SENNA 319 ovate,
Page 331 and 332:
Page 333 and 334:
1982] CASSIINAE—SENNA 323 buds su
Page 335 and 336:
1982] CASSIINAE—SENNA 325 orbata
Page 337 and 338:
1982] CASSIINAE—SENNA 327 glabrou
Page 339 and 340:
1982] CASSUNAE—senna 329 nation o
Page 341 and 342:
1982] CASSIINAE—SENNA 331 varying
Page 343 and 344:
1982] CASSIINAE—SENNA 333 seldom
Page 345 and 346:
1982] CASSIINAE—SENNA 335 to and
Page 347 and 348:
1982] CASSIINAE—SENNA 337 Charact
Page 349 and 350:
Page 351 and 352:
Page 353 and 354:
1982] CASSIINAE—SENNA 343 Dry roc
Page 355 and 356:
1982] CASSIINAE—SENNA 345 llOg. S
Page 357 and 358:
1982] CASSIINAE—SENNA 347 exempli
Page 359 and 360:
1982] CASSIINAE—SENNA 349 17. Lft
Page 361 and 362:
1982] CASSIINAE—SENNA 351 (3-)4-6
Page 363 and 364:
1982] CASSUNAE—senna 353 a photog
Page 365 and 366:
1982] CASSUNAE—senna 355 the pedu
Page 367 and 368:
1982] CASSIINAE—SENNA 357 filifor
Page 369 and 370:
1982] CASSIINAE—SENNA 359 Stamens
Page 371 and 372:
1982] CASSIINAE—SENNA 361 Pods "o
Page 373 and 374:
1982] CASSIINAE—SENNA 363 obtuse
Page 375 and 376:
1982] CASSIINAE—SENNA 365 the vex
Page 377 and 378:
1982] CASSIINAE—SENNA 367 Racemes
Page 379 and 380:
Page 381 and 382:
1982] CASSIINAE—SENNA 371 cm, the
Page 383 and 384:
1982] CASSIINAE—SENNA 373 Pod pen
Page 385 and 386:
Page 387 and 388:
1982] CASSIINAE—SENNA 377 chilens
Page 389 and 390:
1982] CASSIINAE—SENNA 379 mm, the
Page 391 and 392:
1982] CASSIINAE—SENNA 381 found t
Page 393 and 394:
1982] CASSIINAE—SENNA 383 76 of c
Page 395 and 396:
1982] CASSIINAE—SENNA 385 mostly
Page 397 and 398:
1982] CASSIINAE—SENNA 387 replace
Page 399 and 400:
1982] CASSIINAE—SENNA 389 eastern
Page 401 and 402:
1982] CASSIINAE—SENNA 391 h is ju
Page 403 and 404:
1982] CASSIINAE—SENNA 393 in sylv
Page 405 and 406:
1982] CASSIINAE—SENNA 395 Araguai
Page 407 and 408:
1982] CASSIINAE—SENNA 397 or almo
Page 409 and 410:
1982] CASSIINAE—SENNA 399 Pod ess
Page 411 and 412:
1982] CASSIINAE—SENNA 401 only ±
Page 413 and 414:
1982] CASSIINAE—SENNA 403 locaUy
Page 415 and 416:
1982] CASSIINAE—SENNA 405 COLUTEO
Page 417 and 418:
1982] CASSIINAE—SENNA 407 within
Page 419 and 420:
Page 421 and 422:
1982] CASSIINAE—SENNA 411 but the
Page 423 and 424:
1982] CASSIINAE—SENNA 413 interfo
Page 425 and 426:
1982] CASSIINAE—SENNA 415 cm; bra
Page 427 and 428:
1982] CASSIINAE—SENNA 417 -35) mm
Page 429 and 430:
1982] CASSUNAE—senna 419 mexicana
Page 431 and 432:
1982] CASSIINAE—SENNA 421 places,
Page 433 and 434:
Page 435 and 436:
1982] CASSIINAE—SENNA 425 137d. S
Page 437 and 438:
1982] CASSIINAE—SENNA 427 across
Page 439 and 440:
1982] CASSIINAE—SENNA 429 6. Vena
Page 441 and 442:
1982] CASSIINAE—SENNA 431 on the
Page 443 and 444:
1982] CASSIINAE—SENNA 433 var. gl
Page 445 and 446:
1982] CASSIINAE—SENNA 435 Caledon
Page 447 and 448:
1982] CASSIINAE—SENNA 437 occiden
Page 449 and 450:
1982] CASSIINAE—SENNA 439 piano-c
Page 451 and 452:
1982] CASSIINAE—SENNA 441 Bot. 24
Page 453 and 454:
1982] CASSIINAE—SENNA 443 valves
Page 455 and 456:
1982] CASSUNAE—senna 445 scarcely
Page 457 and 458:
1982] CASSIINAE—SENNA 447 Cassia
Page 459 and 460:
1982] CASSIINAE—SENNA 449 Sesse &
Page 461 and 462:
1982] CASSIINAE—SENNA 451 Neg. 24
Page 463 and 464:
1982] CASSUNAE—senna 453 tierra t
Page 465:
^m\m'^Y%^ws:m.yitmxmfsmsiiimmammKmm
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